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Library

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QPP BBVA PUPYVPAVPAWPAPAPAUAUAYPAPVYAAP

THE

GARDENS BULLETIN

SINGAPORE

BYOVPV PV PVPVPUPUPVPVUPUPYVPYUP VPA VPVPUAUA>VA™IV AV

Vol. XVII, Part I Ist November, 1958

YY YVPVPYBPYUPV POY. BAPUAYAAAUAAAAP-PQO’VDAV

CONTENTS

PAGE

KOSTERMANS, A. F. C. H.: New and Critical Malaysian Plants, V_. 1

Korisa, K.: On the Periodicity of tree-growth in the tropics . iota

. GILLILAND, H. B.: Plant Communities on Singapore Island . oh: canoe

KERN, J. H.: Juncaceae, a new family record for Malaya. Pera 3

Gheisid, J.: Ararocarpus—A Monstrosity . 93

SINCLAR, J.: Florae Malesianae Precursores—XX. The Gene Gym-

nacranthera (Myristicaceae) in Malaysia . - 96

Book REVIEW: Common Malayan Plants—(Selected Drawing with

notes) by H. B. GILLILAND . : 121

To be purchased at the Botanic Gardens, Singapore

Price: $12.50

Published by Authority

Prey AT THE GOVERNMENT FRINTING OFFICE, SINGAPORE,

y R. D. GILLESPIE, ACTING GOVERNMENT PRINTER

1958

ys = oh ye gaat et Si SA wrk ant ec are htt Z

i AR

“RECEIVED x

Ving

”

PYLE

CONTENTS

. Page

Part 1—Ist November, 1958

KOSTERMANS, A. J. G. H.: New and Critical Malaysian Plants, V Be +, 1

KoriBa, K.: On the Periodicity of tree growth in the tropics pies ae ii

GILLILAND, H. B.: Plant Communities on Singapore Island = oy $2

Kern, H. J.: Juncaceae, a new family record for Malaya ... Se Pe OF

SINCLAIR, J.: Ararccarpus—A Monstrosity ie esis - 4, SS

SINCLAIR, J.: Florae Malesianae Precursores—XX. The Genus Gymnacranthera

(Myristicaceae) in Malaysia fle es ae ae ct 96

Book REVIEW: Common Malayan Plants—(Selected Drawings with notes) by H. B.

Gilliland . rye sx ate Py * ely

xe = * *

Part 2—Centenary Issue—Sth December, 1959

DEDICATION pi ts a ed ae ea £23

PURSEGLOVE, J. W.: History and Functions of Botanic Gardens with special reference

to Singapore a = a ss: cs 2 AES

RUSSELL, T. A.: Kew and Singapore iy aie bi APS oD

VAN STEENIS, C. G. G. J.: Singapore and Flora Malesiana a oe pL

Lam, H. J.: A tale of two cities: Singapore and Leiden .. aa SLOG

BurRKILL, I. H.: A note on the Gardens’ Jungle o< ay vee ah

WYATT-SMiTH, J.: The Singapore Botanic Gardens and Forestry in Malaya wee ke

WYCHERLEY, P. R.: The Singapore Botanic Gardens and Rubber in Malaya za 175

ANoNyMous (4 Malayan Agriculturist): The Contribution to Agriculture in Malaya

by the Singapore Botanic Gardens — ae om e. hOY

Ho.ttum, R. E.: Orchids, Gingers and Bamboos; Pioneer work at the Singapore

Botanic Gardens and its significance for Botany and Horticulture .. ae 490

Furtapo, C. X.: Singapore’s Contribution to the Study of Palms Be x 195

QUISUMBING, E.: Manila and the Singapore Gardens A ts sa, 199

BuRKILL, H. M.: The Botanic Gardens and Conservation in Malaya .. se 204

KusNotTo SETyopIwiryo: The Singapore Botanic Gardens and the Central Institute

for Nature Research in Indonesia at Bogor. Comments on Past and Present

Co-operation and the Need to continue it .. 4a os es 206

Corner, E. J. H.: The Importance of Tropical Taxonomy to modern Botany .. 209

Gi_mour, J. S. L.: The International Code of Nomenclature for Cultivated Plants,

with Special Reference to Tropical Botanic Gardens... ie Sere ls

FosBerG, F. R.: The Importance of Biological Research in the Pacific Region

Mattsson, L.: Role of Botanic Gardens in the Humid Tropics and U.N.E.S.C.O’s

Programme related to them

GILLILAND, H. B. and WANTMAN, M. J.: Regenerating High Forest on Singapore

Island .

ROBINSON, R. A.: The pH of Rain Water from the Rotanic Gardens ..

MOLESWORTH-ALLEN, M.: Malayan Fern Notes

MOLESWORTH-ALLEN, B.: Malayan Fern Notes, II

KERN, J. H. and VAN STEENIS, C. G. G. J.: An Interesting new Record from the

Malayan Beach: Spilanthes urens Jacq., its synonymy and distribution

Furtapo, C. X.: A New Aroid from Sarawak

FurtTapbo, C. X.: Some New or Noteworthy species of Malaysia

JOHNSON, A.: The Genus Sphagnum in Malaysia

JOHNSON, A. and TAN KrAP SENG: Cleome ciliata Schum. et Thonn. in Singapore

OBITUARIES:

Haji Mohamed Nur bin Mohamed Ghous, B.E.M.

Ngadiman bin Haji Ismail

Kwan Koriba

Part 3—18th May, 1960

SANTAPAU, H.: I. H. Burkill in India a

FurtTabo, C. X. and R. E. Hotttum: I. H. Burkill in Malaya

VAN STEENIS, C. G. G. J.: Phellodendron, a genus of trees new to the Malayan Flora

(Rutaceae)

HOLTTuM, R. E.: Vegetative Characters distinguishing the various groups of ferns

included in Dryopteris of Christensen’s Index Filicium, and other ferns of similar

habit and sori

Corner, E. J. H.: Taxonomic Notes on Ficus Linn., Asia and Australasia. Sections

1-4 .

(Editor): Posthumous Publication of New Dipterocarp Species from North Borneo

B.E.S.: G. H. S. Woop, M.A., F.L.S. (A tribute)

INDEX

THE

GARDENS’ IBULLETIN

SINGAPORE

BV_YVPVOVPY(PYUPUPUPYW(P UIP AUM AU AAU ™OV

Vol. XVII, Part | Ist November, 1958

VV VPYWV VV PYU(PVYV(PYUPUPYUPYU(PYU( UV ™° AY VV

New and Critical Malaysian Plants V*

By A. F. G. H. KOSTERMANS

Forest Service of Indonesia, Bogor

1. Brownlowia purseglovei Kostermans—Fig. 1.

(Tiliaceae ).

Arbuscula, foliis subobovato-oblongis, paginam inferiorem squa-

mulis rotundatis minutis obtectam. Inflorescentia pauciflora brac-

teis bracteolisque magnis persistentibus. Petala rosea, oblonga,

basin versus subcontracta. Calyx campanulatus, lobis magnis

acutis, squamulis dense obtectis.

Typus.—Purseglove 4662 (SING).

Shrub, 3 m. Branches grey-brown, lenticellate. Stipules subulate,

up to 12 mm. long, densely scaly. Leaves chartaceous, subobovate-

oblong, up to 32 cm. long, 12 cm. wide; top conspicuously, broadly

acuminate; base acutish; upper surface glabrous, glossy, nerves

impressed; lower surface golden, densely covered with minute,

round scales (scales with numerous radial lines); midrib strongly

prominent; lateral nerves about 20 pairs, arcuately anastomosing

near margin; the basal pair (or pairs) slightly ascendant; veins

reticulate, prominulous. Petiole 1 cm. Inflorescence few-flowered,

not or hardly branched, densely covered with scales. Bracts and

bracteoles large, persistent. Pedicel 6-8 mm. Calyx 6-8 mm.,

cup-shaped, 3—4 lobed, (lobes slightly more than half the cup,

acutish, triangular). Petals 3-5, pink, in bud imbricate, narrowly

oblong, up to 3 cm., glabrous, narrowed towards base, apex acut-

ish. Stamens numerous, filaments up to 1 cm. long, slender; an-

thers yellow, subpeltate (opened). The 5 staminodes glabrous, 6

* The series I-IV appeared separately, issued by the Planning Bureau,

Forest Service of Indonesia, Bogor, 1954-1956.

Gardens Bulletin, S.

mm. long, narrowly lanceolate, acute. Ovary 1-2 mm., obovoid,

densely covered with scales; style 9 mm.; stigma inconspicuous.

Distribution—Only known from type locality.

Fig. 1. Brownlowia purseglovei Kostermans.

Bud x 5; flower x 5.

Vol. XVII. (1958).

Specimens examined: Sarawak, G. Pueh, alt. 70 m., Sept., f1.,

Purseglove 4662 (SING, BO).

The species is outstanding by its non-peltate, large leaves and

large flowers. In the sheet at hand, one opened flower is present,

which has 3 calyx lobes, 3 petals and 5 staminodes; there is also

one bud, which has 5 petals.

__ The species is called after Mr. Purseglove, former Director of the

Botanic Gardens, Singapore, the discoverer of the specimen.

2. Alseodaphne magnifica Kostermans, spec. nov.—Fig. 2, 3.

(Lauraceae).

Arbor ramulis teretibus nitidis viridis (in sicco nigris) glabris;

foliis chartaceo-coriaceis oblanceolatis ad obovato-ellipticis, acu-

minatis, basi in petiolum latum decurrente, discoloribus (in sicco).

Inflorescentia axillaria paniculata paulo et breviter ramosa, flori-

bus flavibus (post anthesin rubescentibus). Fructus ellipsoideus,

niger, sub-obliquus, pedicello nudo impositus.

Tree, up to 32 m. tall; bole 21 m. long, 70 cm. in diameter.

Buttresses thick, inconspicuous. Bark yellowish brown, rather

flaky. Branchlets smooth, green (black when dried); apical bud

acute, minutely sericeous. Leaves alternate, glabrous, chartaceous-

coriaceous, glossy green (when dried dark brown beneath), ob-

lanceolate to obovate-elliptical; top acuminate; base decurrent in-

to the broad, up to 1:5 cm. long petiole; midrib, lateral nerves

(about 7-9 pairs) and the lax reticulation prominulous (lateral

nerves rather erect) on both surfaces. Inflorescences axillary,

developing immediately below the new flush, sparsely, minutely

sericeous, glabrescent, consisting of panicles of 2—5 cm. long with

few and short branchlets; peduncle rather stout. Lateral branch-

lets bearing 2—4 flowers in axils of minute, triangular bracts.

Pedicel rather stout, 2-4 mm. long, sparsely, minutely pilose.

Flowers yellow, turning dark purplish red after anthesis, about

3—4 mm. in diameter; outer sepals shorter than inner ones, trian-

gular-ovate, acute 1-5 mm. long; inner tepals ovate, acutish 2—2-5

mm. long. Stamens sessile, densely, minutely pilose; outer ones

ovate, acutish, 1 mm. long with introrse, slanting cells; inner ones

smaller, rectangular, obtuse, cells lateral; basal glands sessile.

Ovary glabrous, ovoid-ellipsoid, 1-1-5 mm. long, style distinct,

1 mm., stigma conspicuous, peltate, laterally incised. Fruit ellipsoid,

often a little oblique and club-shaped, glossy blue-black; endo-

carp 1—2 mm. thick, juicy; cotyledons large, flat-convex.

Typus.—Kostermans 11070 (BO).

Distribution.—Sumatra.

Gardens Bulletin, S.

Specimens examined.—Sumatra. Lampong Districts, Gedang

Harta, ster., Teijsmann H.B. 4429 (kaju djoos); West Coast

Sumatra, Priaman, alt. 400 m., Apr., ster., bb. 6721 (medang

djambu; fancy name) (BO, L); Hort. Bogor., culta sub V A 24,

Oct., fl., fr., Kostermans 11070 (A, B, BO, BRI, BZF, UC, CAL,

LU, CANB, BISH, K, KEP. L, LAE, MEL, NY, P, PNH, SAN,

SING, NWS).

os II EME GE GDR ELLE DE A

Fig. 2. Alseodaphne magnifica Kostermans.

Photo: Bot. Gardens, Bogor, Sth Nov., 1955.

Vol. XVII. (1958).

It is possible, that the tree in the Bogor Gardens was grown

from seeds of the specimen H.B. 4429. The vernacular names—

as is usual in Lauraceae—have no value at all and should be con-

sidered as pure fantasy.

sit

b ZN

Be:

ATG)

Fig. 3. Alseodaphne magnifica Kostermans no. 11070 29 Oct., 1955 Keb.

Raya photo.

a & b, flowers < 5; c, outer stamen x 10; d, ovary x 10;

e, inner stamen x 15; f, flower seen from above x 10.

3. Sympetalandra borneensis Stapf

(Caesalpiniaceae-Dimorphandrae).

Sympetalandra borneensis Stapf in Hook. Icon. 4 (8): t. 2721.

1901; Merrill, Bibl. Enum. Born. pl. (J. Str. Br. Roy. Asiat. Soc.

Spec. Numb.) 295. 1921; in Univ. Calif. Publ. Bot. 15: 98. 1929.

—Haviland 1628 (K).

Erythrophloeum densiflorum (Elm.) Merr., l.c. (non aliis).-—

Villamil 52.

Erythrophloeum unijugum Airy-Shaw in Kew Bull. 1939: 180.

—Agullana For. Dept. 1780 (K).

Serianthes gigalobium Kostermans in Reinwardtia 2: 357.

1953.—bb. 20030 (BO).

The reason why this species was described under four different

names and three different genera originates from the fact, that the

Gardens Bulletin, S.

leaves vary between simply pinnate to bipinnate (unijugate and

bijugate) and that I described the species from a fruiting speci-

men, whereas Merrill and Airy-Shaw examined flowering speci-

mens.

With all type specimens at hand now, I have been able to clear

up all controversies. I collected several times fruit of this species in

the island of Nunukan, where it is common locally on rather

loamy soils. The simply pinnate leaves are not only found near the

inflorescences, but occur also on sterile branchlets, which are less

vigorously developed. Unijugate leaves are quite common among

bijugate ones on the same tree.

Consequently Merrill was right in referring the specimen Elmer

20923 to Sympetalandra borneensis. Airy-Shaw’s contention, that

the specimens certainly belong to Erythrophloeum, is wrong, as

the pods are different: they are non-dehiscent and very thick.

Sympetalandra is a distinct genus, allied to the African genus

Burkea Hook. f., which has equally non-dehiscent (but very thin)

pods. In leaf characters it does not differ from Burkea. The identi-

fication of the sterile Sumatra specimens, enumerated in Rein-

wardtia, is not sure. The seeds have a pink seedcoat. The pods are

yellowish green.

Additional specimens (other specimens are mentioned in Rein-

wardtia, l.c.).—British North Borneo. Sandakan, Kabili For. Re-

serve, alt. 7 m., Nov., ster., Agama B.N.B. For. Dept. 9953 (BO,

K); Kandasan—Tenompok, 12 miles W. of Ranau, Kota Belud, alt.

1500 m., Dec. fr., San 16398 (BO, SAN); Sepilok For. Reserve,

15 miles W. of Sandakan, alt. 50 m., April, in bud, Wood &

Charington, San. 16313 (A, BO, BRI, K, KEPONG, L, SING);

ibid., June, fl., N.B.F.D. A808 (BO, K); Tawau, along bank of

Marutai R., Agullana 1780 (BO, K). Sarawak, Kuching, Sept.,

in bud, Haviland & Hose 1628 (BO, K). Indonesian Borneo.

East Borneo, Nunukan Isl., Nov., ster., Kostermans 8790 (A, BO,

K, L, SING, PNH); ibid, Nov., fr., Kostermans 8727 (A, BO, K,

L, SING, PNH); ibid., Nov., fr. Meyer 2176 (A, BO, K, L);

ibid., young tree, Kostermans 8727A (BO, L).

Living material was brought by me to the Bogor Botanical

Gardens, where it is cultivated.

Vernac. names.—Tadarun (Dusun—Kandasan).

4. Teijsmanniodendron sinclairii Kostermans, spec. nov.—Fig. 4.

Arbor, ramuli apice et petioli et foliorum pagina inferior et in-

florescentia minute asperi. Folia elliptica rigide coriacea, petiolo

apice incrassato. Inflorescentia racemosa, dense bracteosa, haud

vel vix ramosa.

Vol. XVII. (1958).

Tree 5 m. tall. Branches and branchlets compressed, stout, grey,

the latter, minutely asperate. Living leaves medium green and

glossy above with a yellowish tinge, yellowish green beneath.

Dried leaves rigid coriaceous, bullate, glossy, glabrous above, less

Fig. 4. Teijsmanniodendron sinclairii Kostermans.

After Kiah & Sinclair S.F.N. 40877. Flowering branch (not

sure), fruit x 5, part of inflorescence x 3.

Gardens Bulletin, S.

glossy and asperate beneath, elliptical, up to 13 & 25 cm., top

acuminate, base acute, midrib prominent in a grove and nerves

sunken above; lower surface: midrib prominent, lateral nerves (8—

10 pairs) prominent, arcuate, near margin arcuately anastomos-

ing, secondary nerves lax, prominent, reticulation obscure. Petiole

stout, 3—4 cm. long, at apex with a conspicuous, globose swelling

at the lower surface. Inflorescences axillary, up to 30 cm. long,

raceme-like; the inflorescence consists sometimes of a short main

peduncle with 2 or 3 long branches. Lower half or one third of

inflorescence bare, peduncle stout, compressed, sparsely asperate.

Flowers 2—3 together on a very short peduncle in the axils of up

to 8 mm. long, narrowly ovate or lanceolate (narrowed towards

base) acute, stiff, persistent bracts, purple, with a yellow brown

patch on lower lip. Calyx campanulate, 3 mm. long, lobes reflexed,

ovate, acutish, 1-5-2 mm. long. Corolla not seen. Fruit cup

shallow, 5-7 mm. deep, 10-12 mm. in diam., margin irregularly

incised, fruit subglobose to obovoid-globose, glabrous, depressed

at apex, 10-12 mm. in diam. with an apical and often a lateral)

median suture, 2-celled with 2 seeds.

Type specimen.—Sinclair and Kiah bin Saleh S.F.N. 40877

(SING).

Distribution.—Malay Peninsula, Borneo.

In my paper on Teijsmanniodendron (in Reinwardtia 1: 97.

1951), I stated, that I had no access to the type specimen of T.

holophyllum Kosterm., in which I included arbitrarily specimens

deviating by their tomentum. Moldenke (in schedae) considered

the pilose specimens (bb. 12144) a variety (var. pubescens

Mold.) of T. holophyllum.

The material, collected by Sinclair and Kiah makes it clear,

that the pilose (scabrous) specimens represent a new species, dif-

ferent from T. holophyllum not only by its scabrosity but also by

the stout inflorescence and larger flowers and fruit. In sterile con-

dition the species is easily recognised by the scabrous (touch) of

the lower leaf surface.

Specimens examined.—Malay Peninsula. Trengganu, Sg. Nerus

at 16th mile Kuala Trengganu right bank, Sept., fl., fr., Sinclair and

Kiah bin Saleh, S.F.N. 40877 (A, BO, K, L, SING). Borneo.

Berau, Inaran, ster., bb. 12144 (BO, L).

5. Abarema nediana Kostermans, spec. nov.—Fig. 5.

Arbor parva ramulis cylindricis laevibus perparce perminuteque

tomentellis, foliis bipinnatis bijugatis foliolis chartaceis, late ellip-

ticis obscure acuminatis, basi in petiolulum contracta supra nitida

ray

Varailis »

Stipes tain reine eam ag CROSS

| Conus:

EA RERS ADORT. BOT. REG. Kew,

i Ete Species: ms

“FLORA or NORTH BORNEO.

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bac

bak : *

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AON A atin, oun

Bi Yee ere pigs

Fig. 5. Abarema nediana Kostermans.

Gardens Bulletin, S.

glabra reticulata subtus terna perminute sparseque adpresse pilosa

petioluli parvi; glandulis pedunculorum ovalibus, glandulis inter

petiolulorum subrotundatis, rachillae basin versus glandulis mini-

mis bigeminis.

Paniculae supra—axillaribus dense minuteque tomentellis, flori-

bus sessilibus dense minuteque adpresse-tomentellis; ovario glabro,

stipitato.

Typus.—N.B.F.D. A142 (BO).

Tree 13 m., 10 cm. in diameter; branchlets cylindrical, smooth,

sparsely and very minutely pilose, with a few scattered pale tiny

lenticels. Leaves bipinnate, bijugate; rachis pilose about 8 cm. long,

near its base with an elliptical, about 2 mm. long, slightly raised

gland with a central depression; base with 2 short (1 cm.) decur-

rent spurs; petiolar part 4-5 cm.; rachillae pilose, lower ones 1—2

cm. long, with 1 or 2 pairs of folioles, near their bases with a pair

of minute, raised glands; interpetiolular glands raised, almost orbi-

cular, 1 mm. in diameter; petiolules 2-3 mm. long; upper rachillae

up to 6 cm. long, with 3 pairs of folioles; folioles chartaceous,

broadly elliptical, the apical ones up to 5 & 7 cm., the basal ones

2 < 3 cm., apex obscurely acuminate, base contracted into petiole;

above glossy glabrous with prominulous reticulation; lower surface

dull with sparse, minute, apressed tomentum; lateral nerves 4—6

pairs.

Panicles slightly above the axils of the leaves, up to 30 cm.

long, sometimes with leaves; densely, minutely tomentellous; flow-

ers white, sessile, densely, minutely sericeous-tomentellous; calyx

infundibuliform, 1-5-2 mm. long with 0-5 mm. long teeth; corolla

4—5 mm., with 2 mm. long, lanceolate, acutish lobes; staminal tube

as long as corolla tube; ovary glabrous, stipitate.

Specimens examined.—Colony of North Borneo. Lumanggas

Is]., Simporna, Lahad Datu Forest Distr., seashore. alt. 1 m.,

Oct., fl., B.N.B. For. Dept., A.142 (BO, K).

The species is named in honour of Mr. Nedi of the Staff of the

Herbarium Bogoriense, who during 20 years has accumulated in

astonishing knowledge of Indonesian plants and who is a much

appreciated assistant in routine identifying.

The species is characterised by the twin glands near the bases of

the rachillae.

According to a note on the field label the plant is used for clean-

ing teeth and seems to have medicinal properties for dental

diseases.

7 he

S06

g 8

S279

§ 10

$11

$12

On the periodicity of tree-growth in the

tropics, with reference to the mode of

branching, the leaf-fall, and the

formation of the resting bud.

By KWAN KORIBA

Hirasaki University, Japan

(with 3 text-figures)

CONTENTS

1. Introduction

. The climate of Singapore

3. Periodicity of tree-growth in its relation to the mode of

branching

4. Materials and methods

5. Terminal growth . :

. Lateral growth

1. Substituting growth

2. Apposing growth

. Evergrowing trees .

1. The terminal evergrowing -

2. The evergrowing tree with lateral growth

. Trees with branches of manifold growth

1. Terminal manifold growth

2. Lateral manifold growth

. Intermittent growth

The transition from evergrowing to intermittent

Deciduous trees , : ; i ;

1. Transition from intermittent-evergreen to deciduous .

2. The leaf-period and the date of leaf-fall and leafing .

The flowering season ; :

1. The flowering of evergrowing trees .

2. Seasonal flowering

3. Contemporary flowering

4. Non-seasonal flowering

14.

16.

17.

18.

20.

oi.

25.

24.

23.

26.

2a.

Gardens Bulletin, S.

The fruiting season

The frequency of species of the four types of periodicity .

Comparison of the leaf-fall and leafing periods of sles ea

pore with those of Ceylon and Java. ;

The grade of leaf-fall of trees in various districts .

The deciduous habit and the height of crown

Deciduous trees and their habitats .

The systematic occurrence of deciduous trees

The geographical distribution of deciduous trees

The geological change of the Malay Peninsula in the past

The transition of shoot-growth from evergrowing to deci-

duous (Fig. 1) . i : :

The morphological development of the resting bud with

special reference to the formation of scales. The anes

buds (Fig. 2) ,

The retrogression of the bud-forming ability

The intermediation of hormonal activity on the process of

periodicity (Fig. 3) : :

Conclusion

Summary

Appendix

Literature cited

Page

S 1. Introduction

DURING My STAY at Singapore, from 1942 to 1945, it was quite

unexpected to find that there were such various growth-forms of

trees compared with the temperate region. In temperate coun-

tries deciduous trees are clearly distinguishable from the evergreen,

and leaf-fall is in accordance with the winter. Even a slight fluctua-

tion of chilly weather affects the date of leaf-fall, so that the ex-

acting influence of the climate is conspicuous. This holds true,

also, in the monsoon region with a dry season.

In Singapore, most trees are naturally evergreen, and some of

them are evergrowing in accordance with the favourable climate,

but most of the evergreens are intermittent in their growth and some

are even deciduous in spite of so uniform a climate, though the

bare spell is very short and the flowering or fruiting may go on

during that time. Besides, there are some trees, in which the leaf-

ing, flowering, etc. are different according to individual branches

or stocks. So we can see among trees every possible transition of

growth-form from the evergrowing to the deciduous.

Another peculiarity to be pointed out in Singapore is that the

leaf-fall does not coincide with the calendar year. Beside those

which shed leaves once, twice, or thrice seasonally every year,

there are those which shed leaves from every several months to

more than one year non-seasonally. Yet there is no exacting change

of climate to enforce this leaf-fall, since other trees or even other

stocks of the same species remain clad with a green crown.

To elucidate why such diverse behaviour of the tree-growth is

displayed was the chief aim of the present investigation.

The periodicity of tree-growth is a natural consequence of the

activity of the growing point. But before discussing the detail, it is

necessary first to consider the climatic character of the region. As

to general features, one may refer to manuals of climatology and

related literature.

§ 2. The climate of Singapore

As is well known, south-eastern Asia and Malaysia undergo, as

monsoon regions, a distinct dry season alternatively. But in Singa-

pore, lying just between those two regions, the monsoon effect is

extremely moderated and there is no truly dry season. The climate

is warm and wet, often cloudy, with insolation neither very long

nor intense.

Gardens Bulletin, S.

The most definite seasonal feature is the rainy season from Octo-

ber to January. The monthly rainfall-amounts to 20 cm. occa-

sionally more than 30 cm. (HOLTTUM 740). The excess of

precipitation, however, does not affect the tree-growth particularly,

as it is drained away by the surface run-off and seepage, though it

is beneficial to the seasonal growth of fungi (CORNER ’35).

On the other hand a long spell of rainless days, together with

intense radiation, affects the periodicity of trees not a little. The

seasonal colour of tree-growth may be attributed mainly to such

occurrences. The so-called dry spells occur in Singapore twice a

year, February to March and July to August, and correspond

nearly to the dry season of India and Malaysia respectively. On the

mainland of the Malay Peninsula, however, the distribution of the

dry season is not the same; it is different on each side of the coast,

as well as inland (STEWART 730).

In Singapore, even in the dry months, the rain-fall amounts

usually to 13-16 cm., rarely 6 cm. According to HOLTTUM

(40) the number of times in which the monthly rainfall was less

than 2:5 inches (= 6-3 cm.) during 50 years (1890-1939) was

twenty-two, or once every two years (see Tab. 1a), and the long-

est drought during 10 years (1927-1937) was 22 days extending

from January to February. The frequency of rainless days, conti-

nued for 14-20 (Tab. 1b) and 7—13 days (Tab. 1c) respectively

during that time, was as follows:—

TABLE I.—DISTRIBUTION OF THE RAINLESS MONTHS IN SINGAPORE

Month I If WI IV V_ VI VILVIILIX X XI XII

a a ea AGS: Yana anya ae Se Gr Sie Des eka during 50 years

b ty ae BT esl inn Voteane ists 4 Sok ere ees 10 years

Coe be) 2 Se a a5 ars ore ee a OP eee oot

From these data it is discernible that the rainless month can

shift one month or more according to the year, indicating the non-

seasonal character of drought in Singapore.

As long as the dry spell continues, the temperature rises day by

day, and the decrease of humidity is more acute. Intense radiation

and decrease of water-content by transpiration seem to induce

anthogenesis and accelerate leaf-fall as well. A shower after the

dry spell causes, then, new leafing often conspicuously. At any

rate, as the climate is fairly uniform, even a short dry spell can

give rise to the periodic feature of tree-growth.

Naturally the trees which respond in this way are not very abun-

dant, or are restricted to their susceptible phase only, while others

show no special reaction to such a triflng change of climate. Even

Vol. XVII. (1958).

among deciduous trees there are those which maintain their proper

periodicity regardless of the dry spell.

If one compares the periodicity of tree-growth in Singapore with

that of the monsoon regions, north and south, or of the tropics of

other continents, one finds that there are some kinds of trees that

are evergreen in Singapore, while deciduous in other regions. In

this respect, Singapore is one of the most uniform climates in the

tropics, and moreover with abundance of tree-species. But, then,

why in the district of such a uniform climate such various growth-

forms occur, is a question to be considered further.

§ 3. The periodicity of tree-growth in its relation

to the mode of branching

The periodicity of tree-growth is derived from the behaviour of

the growing point. If all twigs grow continually, producing leaves

successively, the tree is “evergrowing”. If however the growing

point ceases to grow or slows down after forming a certain num-

ber of leaves, or tends to form an inflorescence, the growth of the

twig becomes intermittent.

But, even if some twigs cease to grow or become flowering

twigs, the tree is still evergrowing as a whole, provided that some

axillary buds are growing into new twigs. The periodicity is, there-

fore, not only related with the activity of the single shoot-apex,

but also with the mode of branching of trees. Such individual dif-

ferences in branch-growth. in the tropics was noted by SCHIM-

PER (798) and various subsequent authors. We may designate it

the manifold growth of tree-branches.

This behaviour is seen not merely in individual branches or

limbs of one tree, but also among trees of the same species. That

is, some stocks are just in new leaf, while others are clad with

dark green old leaves or in flower—the ‘“stockwise manifold

growth”, in contrast to the “branchwise manifold growth” on the

same tree. Among young avenue trees of Spathodea campanulata

both forms of manifold growth can be seen, while in old trees all

stages of the branchwise manifoldness are mingled together in each

tree, so that the stockwise difference ultimately disappears.

Now, trees with intermittent growth are very common in the tro-

pics. New leaf-sprays all over the twigs are often seen. If leaves

of the previous season still remain at that time, the tree is ever-

green just as in the temperate zone, but if they have fallen off

beforehand, it is deciduous, though in the tropics there are various

transitions.

Gardens Bulletin, S.

The periodicity of the tree-growth is determined therefore in two

ways:—

(1) whether the activity of growing points is continuous, mani-

fold or intermittent, and

(2) how the growing points are distributed in each branch,

terminal or lateral.

The mode of branching in general is well known morphologic-

ally, but the periodicity of tree-growth is characterized also by

the behaviour of the growing point, which is rather diverse and

not always intermittent in the tropics and qualifies the periodical

character. ee

By the combination of these two processes, we get six types of

tree-growths as mentioned below:—

THE MODE OF GROWTH AND BRANCHING, AND THEIR COMBINATIONS

( evergrowing \

(1) Activity of growing point < manifold evergreen or deciduous

(intermittent J

terminal

lateral

{ terminal-evergrowing

J lateral-evergrowing ,

terminal-manifold

{3) Mode of growth of tree < itcrak aasssfobd evergreen

| terminal-intermittent deciduous

| lateral-intermittent

§ 4. Materials and methods

Trees examined were those:—

(2) Site of srowing posts ia declan {

(1) Planted in the Botanic Gardens of Singapore, including

exotic trees,

(2) of the primary forest of the Gardens,

(3) of Bukit Timah Forest Reserve,

(4) of the primary forests of the Water Catchment Area,

(5) of various other places on the Island and the Peninsula.

The identification of wild trees was entirely dependent on the

instruction of Mr. E. J. H. Corner, without whose help the present

research could hardly have been accomplished. Dr. R. E. Holttum

gave me also advice concerning the leaf-fall. I express here my

sincerest thanks to them on this occasion.

To classify the growth-types of trees, as mentioned above, is

however not always easy. Especially the distinction of the ever-

growing from the intermittent is difficult, as there is no sharp dis-

tinction between them (§ 10). If one can see the fresh foliage all

Vol. XVII. (1958).

over the twigs, as often is the case in jungles, avenues and gardens,

it is certain that the tree is intermittent in its growth, but if one

can not catch such a chance, it is necessary to investigate the fol-

lowing accessory criteria: —

(1) In evergrowing twigs, the leaves are nearly uniform in

size, similar in colour except a few young ones, and

often with 1—2 ageing leaves at the base. Besides, the

internodes are nearly equidistant.

(2) In trees with intermittent growth, the leaf-size and the

length of internodes are not uniform. At the basal part

of the shoot of each season, there are some scars of

scale-leaves that protected the young shoot initially.

(3) In trees with intermittent growth, there is usually a sudden

change of bark colour, wearing off of epidermis, ap-

pearance of cork-layer, lenticels, etc., whereas in conti-

nually growing twigs all changes are gradual.

Trees with manifold growth are easily discernible by comparing

the branches here and there. Their growth is never continuous,

though usually evergreen, and individual branches may be deci-

duous.

Deciduous trees are very conspicuous at times of leaf-fall. We

can find a full account of them in the literature (CORNER °40

and others). The mode of branching has no direct relation to

periodic deciduousness.

We shall first examine the mode of branching morphologically

and then, combined with the activity of the growing point, the

mode of tree-growth in detail.

S 5. Terminal growth

The terminal growth of trees in the tropics may be grouped into

three types, the tuft-trees, the trunked and the ramiferous.

Tuft-trees produce leaves only from their thick stem, as in tree-

ferns, cycads, palms, papaya-trees, and they rarely branch. Trunk-

ed trees, designated as the monopodial tree, have a main trunk

with relatively slender limbs. This form is very abundant in the

tropics, not only in gymnosperms, but also in dicotyledonous trees.

The ramiferous tree has a short trunk, from which thick limbs

subdivide and ramify gradually into slender branches and twigs.

Naturally there is no sharp distinction between the latter two. All

young dicotyledonous trees grow at first monopodially, but some

trees cease sooner or later their terminal growth as their inherent

character, though this may be varied more or less by environmental

conditions.

Gardens Bulletin, S.

The trunked tree sometimes tends to the many-stemmed form

near the base, and conversely the ramiferous tree in dense forest

may tend to the trunked form up to the high branches.

That there are many monopodial trees in the tropics is one of

their characteristics, indicating the favourable conditions of growth.

Even in trees with sympodial growth, the pseudo-monopodial

form is occasionally seen, as in Alstonia. They seem to be suited

to dense forest. All these forms are recorded systematically in

Corner’s book (1940).

Among trees with terminal growth, there are still some distinc-

tions according to whether the organs produced are uniform or not,

and how long the apical growth can last. In tree-ferns and most of

the palms the kind of organ formed is uniform, while in cycads a

set of scales and foliar leaves are produced in each season, so that

the growth is intermittent. In the flowering season, then, the female

trunk of cycads forms scales, sporophylls and foliar leaves as one

set, while on the male trunk, on account of the terminal position of

the male cone, the stem-apex, withers and growth proceeds by an

axillary bud sympodially. The Corypha-palm on the contrary,

having formed a terminal inflorescence after about thirty years

growth, dies down, while the Sago-palm (Metroxylon) succeeds

this growth with axillary saplings.

Terminal growth is therefore not necessarily endless. It is a mor-

phological alternative to lateral growth. If the main axis terminates

with an inflorescence after some seasons, it is a limited terminal

growth, but if the inflorescence is axillary and growth has no mor-

phological limitation, it is an indefinite terminal growth, even if its

duration be not very long.

§ 6. Lateral growth

When the growing point dies out or becomes an inflorescence,

then one or more lateral buds grow instead and sympodial growth

ensues in each season. In some trees, too, chiefly on their hori-

zontal branches, one or more axillary buds of the underside grow

more vigorously than the main shoot, which then grows slowly as

a short shoot, while the lateral twigs take its place in the continua-

tion of the branch. The same process is repeated in every season,

adding new twigs successively. The former two cases may be

named “substituting growth” and the latter “apposing growth”.

If the growth of lateral buds precedes flowering or dying out of

the terminal part, the twig remains evergrowing, but if it falls be-

hind, growth becomes intermittent, though there is no sharp boun-

dary between the two extremes.

Vol. XVII. (1958).

1. Substituting growth

Two cases may be mentioned, namely the dying of the terminal

bud or twig and the fruiting or decaying of the terminal inflores-

cence. The inflorescence may be represented either by a single

flower, an inflorescence, or the anthocladium (GOEBEL ’31), e.g.

Delonix, but as the morphological details are not important, we

may call them all inflorescences.

As a typical case of the dying of the terminal bud, Xanthophyl-

lum Curtisii may be mentioned. It is an endemic tree of Malaya,

growing twice a year intermittently. Though the shoot is provided

with green coriaceous leaves, its terminal bud dies every season

and an adjoining axillary bud substitutes the growth of the twig

(§ 22, Fig. 1). In Crataeva, Sapium, and several genera of Legu-

minosae, a few upper internodes die away and the remaining lower

buds develop. The young tree of Sapium discolor is especially in-

teresting in respect to its mode of branching. At the end of each

season, the apical part of the axis dies away and one of the appa-

rently whorled branches grows upward and takes the place of the

main stem. The trunk thus assumes a pseudomonopodial form.

Then, after a height of several metres is attained, two branches

grow upwards simultaneously, so that the trunk appears pseudo-

dichotomous, and these limbs repeat this mode of branching; so

in a fairly tall tree a ‘dichotomously branched’, flat-topped crown

develops. In Alstonia scholaris a tall pseudo-monopodial trunk

grows up and a handsome pagoda form is assumed, as illustrated

in Corner’s book, Fig. 15.

2. Apposing growth

This is the so-called “Terminalia-type branching” (RACIBOR-

SKI ’01, p. 50; CORNER ’40, p. 30), and involves horizontally

spreading branches, such as Terminalia, Fagraea, Achras, and

Palaquium, whether evergreen or deciduous, and occurs even in

temperate trees such as Cornus and Idecia. It is homologous with

the rhizomes of Polygonatum and many others. In Tectona and

Eriobotrya, when the branches are horizontally directed, the same

mode of branching occurs, but growth is usually more or less up-

wards, and the more upward it is, the more vigorously the main

twig grows, tending ultimately to normal ramification. This type

of branching is, therefore, a case of geomorphosis caused by hor-

monal activity in which the branches of the upper side are res-

tricted in their elongation, while those of the underside are

accelerated.

So much for the morphology of branching. Let us now consider

the growth-forms of trees in general.

Gardens Bulletin, S.

§ 7. Evergrowing trees

There are three types of evergrowing trees: —

(1) Shoots are all evergrowing by terminal growth—“‘the ter-

minal evergrowing”’.

(2) The shoots show a limited growth by

(a) dying of apical parts,

(b) development of the inflorescence,

but the growth is maintained by the development of

substituting or apposing twigs without resting.

(3) The shoot-growth is intermittent and sometimes decidu-

ous, but the phase of development is manifold accord-

ing to branches, so that the tree as a whole is ever-

growing (§ 8).

Among the evergrowing trees, we may distinguish further two

types: the inherent and the environmental. There are many ende-

mic “evergrowing trees of inherent type”. But some trees, though

evergrowing in Singapore, are intermittent or deciduous in other

regions, so that they may be named “the facultative evergrowing”’..

For instance, Duabanga sonneratioides, Thespesia populnea, and

Trema orientalis are deciduous in India (TROUP ’21) and Trema

is even bare for 22 days (WRIGHT ’05). Mimosa sepiaria and

Psidium guajava are semideciduous in San Paulo and the former

is deciduous in Rio Grande do Sul (30° S) (IHERING ’23).

Hibiscus tiliaceus is intermittent in East Java (FABER 735). All

these trees are evergrowing in Singapore.

Naturally it is not certain whether the deciduous habit of such

trees is environmental or inherent, as there may exist racial dif-

ferences in each region among widely distributed species (§ 22).

1. The terminal evergrowing

This is the simplest case of the twig-growth. The terminal part.

of each twig has a constantly growing bud, forming equal-sized

leaves one after another, leaving a few ageing leaves behind. The

growing point is protected by outer developing leaves, petioles or

stipules. Young leaves may have hairy coverings or secrete gum-

resin and are more or less sheathing. The hairs are usually simple,

occasionally long, branched, tufted or slimy according to species.

Very often the young bud is covered by an outer leaf or a pair

of opposite leaves, the margins of which fit together by entangling

hairs so as to make a tube or cone. As the inner bud grows, the

Vol. XVII. (1958).

connection of the margin is gradually separated and the next leaf,

or leaves, restore the arrangement, as in Duabanga sonneratioides,

Horsfieldia brachiata, and Campnosperma auriculata.

The leaves of Archytaea and Gordonia are glabrous and the

young bud is surrounded by the involute leaf-base, or outer leaf

respectively, so as to recall the leaf-sheath of Tradescantia. When

the younger part is protected by the petiole, this is more or less

grooved, and in cases of opposite leaves as in Gnetaceae, Apocy-

naceae, Loganiaceae, etc., the young bud is enclosed between their

bases, which often make a tube above the bud filled with an exu-

dation of gum-resin or slime. Wormia suffruticosa shows a peculiar

form of bud-covering, in which the equitant petiole encloses the

bud completely (FABER ’35, Fig. 136).

Stipules are often useful for the protection of the bud, too, espe-

cially when they are large, as in Jackia ornata (CORNER ’40,

Fig. 190) and Leea (LUBBOCK ’99, Fig. 188). Trees of Rubia-

ceae, Polygonaceae, Magnoliaceae, and Moraceae have as well-

developed a stipular covering, hood- or cone-shaped, as those of

temperate regions. They are described in detail by POTTER

(91), GROOM (’92), LUBBOCK (’99) and others.

a. Tuft-trees

The tree-ferns, most palms and the papaya-tree are the typical

monocaulous trees. Cycas, Pandanus, Dracaena, etc. are often

polycaulous or branched. The growing point develops steadily,

deeply protected by a tolerable number of young leaves. Most

palms produce inflorescences at the same rate, one by one after

the development of the subtending leaf. In the papaya the surface

of the growing point is slightly convex and the leaf-primordia are

arranged in 2:3 parastichy. In each primordium, there deve-

lops at first the columnar petiole with narrow brim-like lamina on

the top. The growing point is therefore in direct communication

with outer air, though the laminas of older leaves extend hori-

zontally and make a shelter over the young parts.

b. Evergrowing trunk-tree

a. Trunk and branches both evergrowing

Casuarina equisetifolia, Canangium odoratum, Macaranga tri-

loba, Knema malayana, and Adinandra dumosa belong to this

category. Adinandra represents a typical case of the broad-leafed

evergrowing tree. The leaves are formed in 2/5-phyllotaxis and

both the main axis and lateral branches grow continually, bearing

flowers afterwards in their leaf-axils. In Canangium most lateral

pat

Gardens Bulletin, S.

branches are drooping and, after growing a few metres without

thickening and branching, fall away from the abcission layer at the

base: they are, so to speak, a kind of short shoot functionally.

Macaranga is a well known ant-plant (CORNER 740, p. 263).

The ants remove with the growth of twigs to the upper internodes

successively. It seems difficult for ants, if the branching be sym-

podial. The South American ant-trees, Cecropia and Triplaris, are

also terminal in their growth.

B. Trees with evergrowing trunk

Glochidion and Taraktogenus may be cited. G. superbum and

others bear on their main axis narrow scale-leaves only, and this

main axis grows slowly. In contrast, the lateral branches bear

leaves in two rows (CORNER, Lc. Pl. 58). The leaf, or leaves,

at the basal part of the twig are small; the next are bigger; and,

then, they become smaller again. With further growth of the twig

the undulation of the leaf-size is repeated. I observed three such

undulations on one twig. This does not however depend on the

seasonal change of the climate, as the phase of fluctuation is not

the same for all branches. It indicates therefore the very beginning

of the rhythmical growth of the twig (§ 10).

y. Trees with evergrowing lateral branches

Anisophyllea disticha and Horsfieldia sylvestris may be men-

tioned as interesting cases. In A. disticha the main axis produces

only small leaves, and from this axis 5 lateral branches develop in

pseudo-whorled arrangement periodically. The leaves on each

branch are arranged spirally in 4 rows, but distinctly anisophyllous

with two small\ upper rows and two large lower rows (CORNER

lic. Pl. 15), just like the shoot of Selaginella. That the dicotyledon-

ous tree and Selaginella can assume an analogous form is very

remarkable. Similar instances are also recorded in Hepaticae,

Musci and Podostemonaceae (GOEBEL 28, p. 317).

In contrast another species of Anisophyllea, A. Griffithii, bears

the usual coriaceous leaves on the twigs. They are apparently alter-

nate, but, on close examination of the young twig, it is seen that

the leaves are opposite, but the upper leaf of each node, on ac-

count of a strong hypotropous anisophylly, dies away so early in

development that one can hardly find its remains.

Horsfieldia is many-stemmed when grown in the open. The main

axis of each stem produces in every season (perhaps half a year)

ten leaves in 2/5 phyllotaxis, the lower five of which have long

internodes but no axillary branches, while the upper five with short

internodes produce axillary branches, which bear big leaves of the

Vol. XVI. (1958).

same size (35 < 65 cm.) in two alternate rows equidistantly

(+ 2-4 cm.). These branches grow continually without noticeable

secondary thickening and branching, so that they hang down gra-

dually, attaining finally 3-5 m., producing small flower-clusters in

the leaf-axils or on the branch behind the leaves. These branches

are therefore likewise short shoots as in Canangium (mentioned

above). The branches, though without abscission layer, fall off

just like a long compound leaf, lying then sinuously on the ground

like snakes.

c. Evergrowing ramiferous trees

This is the most usual type of tree, in which from the trunk large

limbs ramify gradually to slender branches and twigs. Such trees

are generally monopodial in young stages, but sooner or later the

main axis ceases to grow or lateral branches grow vigorously and

represent main limbs and branches. Of the evergrowing, ramifying

type, there are many, for instance Campnosperma, Elaeocarpus,

Commersonia bartramia, etc.

2. The evergrowing tree with lateral growth

There are two types of growth, the substituting and the appos-

ing, as already mentioned.

a. The substituting growth behind the inflorescence

The flowering of the terminal inflorescence is either seasonal,

contemporary or non-seasonal according to species, but new shoots

can be seen throughout the year. |

In the seasonally flowering trees of evergrowing type, the season

is once or twice every year according to species. As once-flowering

trees may be named Parkia speciosa, Cassia nodosa, and C. sia-

mea. In the last named, the growth of branches is tolerably irre-

gular and the flowering continues from May to October according

to branches, so that it seems rather manifold or represents a transi-

tional case to it. Perhaps the species originated in the monsoon

region and is tending to everflowering in Singapore.

As twice-flowering trees, there are Ervatamia dichotoma, AI-

bizza falcata, Enterolobium saman, and Cassia multijuga, among

which the last named Cassia is also irregular in the branch-growth

and its flowering continues for three months every season.

Among the contemporary flowering trees, there are shrubby

Coffea spp. and Stenolobium stans. The latter flowers several times

every year gregariously, stimulated probably by the temperature-

fall, as in Coffea. The irregularly branched raceme contains flower-

buds of various ages (CORNER l.c. Fig. 44) and only the older

Gardens Bulletin, S.

buds react to the stimulus, while younger ones wait for the next

chance. New lateral shoots grow all the time irregularly.

As trees of non-seasonal flowering, there are Cerbera, Ochrosia,

Kopsia, Plumeria, Ixora, Vitex and others. The flowering is more

conspicuous twice every year, but the flowering can be seen at any

time, so that it seems to be transitional to manifold growth and

flowering. This is especially the case in Kopsia and Plumeria.

Wormia suffruticosa has continual substituting growth. The infio-

rescence is terminal at every node, but the axillary vegetative bud

grows early and assumes the terminal position, so that the inflo-

rescence seems opposite to the subtending leaf. Jatropha multijuga

has the same mode of growth.

b. Apposing growth

Fagrea fragrans grows luxuriously in Singapore with a hand-

some crown and is well adapted to the soil-conditions. It is

originally monopodial, but in open ground develops often “sub-

trunks” with big limbs. They branch in Terminalia-fashion and are

evergrowing. Each short shoot produces one pair of opposite leaves

every two months, and counting from the number of leaf-scars, each

short shoot dies away after about 6-7 years, but new shoots are

added one by one laterally, extending the branches outwards.

Flowers in axillary clusters are conspicuous in May and October

(§ 12). F. gigantea also shows apposing growth in the same fashion.

Apposing growth is not confined to evergrowing trees, but occurs

also in trees of manifold and intermittent growth, some of which

are deciduous, as mentioned below:—

Manifold: Achras Zapota, Palaquium gutta;

Intermittent: Pterocymbium tinctorium, Terminalia Muelleri;

Deciduous: Sterculia macrophylla, Firmiana fulgens.

§ 8. Trees with branches of manifold growth

So long as a tree shows continuous growth, there is no manifold

growing. In lateral-growing trees, if the development of axillary

buds occurs in every branch non-seasonally, it may be regarded as

a manifold growth in short steps. In typical manifold-growing

trees, however, whether the growth is terminal or lateral, one can

discern the growing and resting branches, or the vegetative and

flowering branches at a glance. This is seen very often in sympo-

dially growing trees, because even a slight difference of the deve-

lopment phase of new shoots behind the dying twig or inflorescence

exerts a further shift of growth and this affects the growing phase

and emphasizes the individual differences more and more.

Vol. XVII. (1958).

This is the usual case of manifold growth accompanied by lateral

branching. But there are still two other cases of the manifold

growth. The one is the big tree in general. As a tree grows tailer,

the light-condition, the ascent of sap, etc., become uneven accord-

ing to the position of the branches, and, so far as the climate is

uniform, exerting no seasonal restriction, each branch may submit

to a position effect and the result tends to be manifold.

As the second case, various exotic trees may be named. Most of

them are adapted inherently to the dry or cold season with inter-

mittent growth, and even under the uniform climate the tendency

does not disappear so easily in each twig. By a combination of

twig-individuality and position-effect, the manifold growth is em-

phasized. This is not only the case in the warm region, but ob-

served also in the cool alpine region with uniform climate. For

instance, temperate trees as Magnolia, Pyrus and others, trans-

planted to the Alpine Garden of Tyjibodas (1425 m) of Java,

show similar manifold growth (HABERLANDT ’93, SCHIMPER

"98, FABER ’35). I have observed also these features in that

Garden, as well as on the summit of Mt. Tankoeban Prahoe

(2076 m), where Vaccinium varingiifolium shows typical mani-

foldness remarkably.

So far concerns the branchwise manifold growth. As to the

stockwise manifold growth, it is very common in transplanted

trees, e.g. Lagerstremia flos-regine, Cassia fistula, Delonix regia,

Spathodea campanulata, Hevea brasiliensis and various other

avenue trees. Perhaps at the time of transplanting of nursery trees,

a slight difference in treatment, as the cutting away of branches

and roots, affects the further growth of the young trees, and, as

there is no climatic restriction, each tree or branch take its own

way and manifold growth follows gradually.

1. Terminal manifold growth

This is observed in intermittently growing trees and there are

transitions from nearly evergrowing to deciduous, e.g. Myristica

fragrans, Eugenia aromatica, Michelia alba, Carapa guianensis

and some genera of Leguminosae.

Myristica seems at a glance to be evergrowing, but really inter-

mittent in a slight degree. In the Moluccas, its native district, the

fruit-season is once every year, so the tree in Singapore seems to

be converted to manifold growth of nearly evergrowing habit. The

axillary flowers develop in ascending order one by one and the

fruits of various size are seen in various positions on twigs here

and there. For Eugenia aromatica the condition seems almost the

same, though no flower is seen.

Gardens Bulletin, S.

Michelia alba is of continental origin and in the north and mid-

dle part of Malaya it is more or less deciduous (CORNER, Lc.

p. 434), but in Singapore it is evergreen and manifold. Both the

resting and growing bud are covered by the characteristic scale-

like stipules of the family. It grows twice every year vigorously,

but new shoots can be seen throughout the year with fragrant

white flowers, more abundant in some stocks than others.

Carapa guianensis produces brown or light green new shoots

here and there over the dark green crown. The resting buds are

covered densely by ten or more scale-like petioles even in Singa-

pore. In Ceara (4° S), Brazil, the tree is deciduous (IHERING

23)

Ambherstia, Brownea and Saraca show the most conspicuous

manifold growth represented by new, white or brown, hanging

shoots mingled with the green crown. Besides, some conifers,

such as Pinus Massoniana, P. Merkusii and others, also grow

manifoldly in Singapore.

2. Lateral manifold growth

a. Substitution of the withered apical part

In some trees one or more apical buds of twigs wither after pro-

ducing a certain number of leaves. New shoots then arise from the

buds of the proximal part and substitute with their new growth.

The withering has no intimate relation with the season, so that the

new growth becomes manifold. Crataeva Roxburghii, Semecarpus

sinensis, Pistacia formosana, and Sapium sebiferum (deciduous in

China) are instances.

b. Substitution of the inflorescence

Tectona grandis grows well in Burma, East Java and other

monsoon regions and is deciduous, while in Singapore it tends to

be evergreen, though the growth is poor. The development of the

shoot is extremely manifold, forming the terminal inflorescence

non-seasonally. Crataeva (mentioned above) also forms a terminal

inflorescence sporadically.

c. Apposing growth

Achras zapota is, though evergreen, distinctly intermittent in the

growth of its successive short shoots. The resting bud is covered by

petiolar scales and the growth of twigs is fairly manifold, so that

the ripening of axillary fruit is non-seasonal. Palaquium gutta of

the same family shows a similar mode of growth, though nearly

evergrowing.

Vol. XVII. (1958).

d. Stockwise manifold growth

The leafing, flowering or leaf-fall of each stock proceeds in this

case with different phase, indicating that the climatic influence is

rather ineffective for periodicity. This is especially noticeable in

avenue trees with attractive flowers or young foliage. Lagerstroe-

mia flos-reginae, Cassia fistula, Peltophorum pterocarpum, Delonix

regia, and young trees of Spathodea campanulata give to travellers

an impression, that the tropics is the land of everflowering. The

bright-red foliage of Cinnamomum iners is likewise striking.

The flowering season of Cycas Rumphii is also stockwise mani-

fold and, as it is dioecious, the pollination may sometimes be

missed, when the trees are not abundant.

§ 9. Intermittent growth

In this case the unfolding of new leaves occurs periodically every

year once or more, and full-grown shoots show a fluctuation in the

size of leaves as well as in the length of internode, with resting

buds on the apex and in the axils. The mode of branching, ter-

minal or lateral, has no direct connection with the periodicity. As

trees of terminally intermittent growth, there are Kurrimia pani-

culata, Quercus conocarpa, Durio zibethinus, and as the substitut-

ingly intermittent ones Xanthophyllum Curtisii and Cleistanthus

heterophyllus. Those with intermittently apposing growth have

been stated already (§ 7—2-b).

For the covering of resting buds in the tropics, there are various

modes and grades. Some are nearly naked as in the growing bud,

but others are covered tightly by scales derived from the petiole,

stipules or modified leaf-base or blade.

The axillary buds are usually in the resting condition. They are

sometimes naked, sometimes covered by a pair of prophylls. This

is also the case in the evergrowing trees (§ 22).

New foliage is usually light green, sometimes brownish red or

even blue (CORNER, lL.c. p. 85), and often more attractive than

the flower. When strolling in the forest, one will notice very often,

that many trees are in new leaf gregariously. This appearance is

very common after a dry spell of several days, but one can see at

any time new leafing here and there, denoting that new leafing is

induced either by the weather condition or by the internal activity,

facultatively or by a combination of both.

In trees with intermittent growth, therefore, there are three types

of the leafing: the seasonal, the contemporary and the non-sea-

sonal. In the seasonal leafing the frequency is 1—3 times every

year, twice being the most usual in Singapore. Contemporary leaf-

ing is not regular in calendar months, and occurs in accordance

Gardens Bulletin, S. :

with a slight change of the weather. Non-seasonal leafing is much

more irregular. Some instances of leafing are as follows (Roman

number denotes the month) :—

(a) Seasonal leafing: once every year: Emblica officinalis

(III), Parkia speciosa (III), Erythrina lithosperma

(X); Twice a year, chiefly in February to March and

August to September after the dry spell: Kurrimia

paniculata, Castanopsis and Quercus spp., Calophyllum

inophyllum, Garcinia mangostana, Parinarium corym-

bosum, Nephelium lappaceum, Gordonia spp.; thrice

a year: Eugenia grandis (III, VII, XII).

(b) Contemporary leafing: Flacourtia rukam (3 times), Cin-

namomum iners (3—4 times), Rhodamnia trinervia

(4—5 times). In Cinnamomum one can see, besides the

usual contemporary leafing, sporadic leafing according

to stocks at any time of year, or the non-seasonal leaf-

ing is here more or less combined. In Rhodamnia the

leafing as well as the flowering in the same district is

simultaneous, but the leafing and flowering do not

necessarily coincide. Probably flowering is caused by a

sudden fall of temperature, while the leafing is induced

after a dry spell of weather.

year) and many trees with non-seasonal leaf-fall.

§ 10. The transition from the evergrowing to the intermittent

In the tree-growth of the tropics, there are various transitions

from the evergrowing to the intermittent and, further, to the deci-

duous (§ 11 and 22). The first indication of intermittent growth

developing from evergrowing is fluctuation of leaf-size. Bridelia

tomentosa grows continually in its main stem, and the lateral

branches bear leaves in two rows alternately. But the leaves are

not of uniform size, the spray showing a more or less waved out-

line as a whole. In long branches one can see three such waves re-

presenting three periods of growth. The leaf-size is larger in the

middle part and smaller (ca. 4 in size) in the proximal and distal

parts of each period with shorter internodes. Glochidion of the

same family shows likewise a fluctuation of the leaf-size, though

it is more or less disturbed when secondary branches appear.

Some conifers, such as Araucaria Bidwillii, Dacrydium elatum

and one variety of Cryptomeria japonica (var. araucarioides), also

show this fluctuation.

Vol. XVII. (1958).

With further advance of the periodic tendency, a few proximal

leaves of each season wrap up the inner part of the shoot, and fall

off after development of the latter. For instance in Rheedia mad-

runo, one or two leaves, though green, are merely 1/5 of the

normal length. In Gordonia singaporeana the first leaf of each

season is small, light green and falls early. The second leaf is also

a little smaller than usual, brownish-green in colour containing

anthocyan, falling likewise early, and from the third leaf onwards

the normal form is assumed. In Myristica fragrans the resting bud

is naked and the proximal two leaves embrace each other, pro-

tecting the inner leaves, and fall early after elongation of the shcot.

With still further development of the intermittent tendency, the

growing point becomes a resting bud, formed after the production

of a number of leaves, initiated by primordia as incomplete cover-

ings, or well-developed scales, wrapping a number of younger

leaves. The bud-envelope is represented sometimes by incomplete

leaves, at other times only by petioles or stipules without blade.

When the petiole serves as the covering, the lamina does not deve-

lop well (Achras zapota). When the leaf is sessile, the covering

may change gradually from scales to the normal leaf (Lucuma),

and, in the) cases of the stipulate leaf, the entire leaf-element may

form at first a scale (Aporosa nigricans), or at first only stipules

are formed and then, after forming some prophylls, the foliar

leaves develop abruptly (Kurrimia paniculata) or with gradual

transition (Brassaia actinophylla).

How shall we then define the intermittent growth in contrast to

the evergrowing, as they are only of gradual change in nature? It

seems, however, adequate to define the shoot as evergrowing, when

leaves at the onset of the new shoot, even if small in size, remain

as assimilating leaves, as in Bridelia and Rheedia. But, if the

leaves, even if green, are distinctly small and fall off early (Myris-

tica) or show a difference in size and colour (Gordonia), one

may define the shoot-growth as intermittent. Such a rhythmic

character of the shoot is quite independent of climatic changes and

is undoubtly caused by inherent hormonal activity on the growing

point, by which the formation, growth and senescence proceed

periodically, often with a pre-determined number of leaves (§ 23).

KLEBS (715) once claimed that the periodicity of tree-growth

could be controlled artificially. But the continual growth of trees

is seen only in evergrowing trees or saplings, which latter tend then

to rhythmic growth. The opinion that the supply of minerals may

cause the evergrowing or intermittent growth is also inconceivable,

as in so uniform a climate as Singapore both types of tree, the

Gardens Bulletin, S.

evergrowing and intermittent, flourish side by side, and also

stockwise manifold growth occurs. At any rate, the basic cause is

the inherent factor, by which the trend to periodicity is deter-

mined.

§ 11. Deciduous trees

In evergreen trees with intermittent growth, the older leaves fall

off after new leafing. But there are trees in which leaf-shedding

precedes new leafing, exposing thus naked branches, and thereby

characterising the deciduous tree. In them three phases of deve-

lopment, leaf-fall, naked branches and new leafing, follow succes-

sively. The mode of branching is not intimately related, whether

it is the terminal (Ficus, Hevea), the substituting (Cassia fistula,

Lagerstromia spp., Sapium discolor) or the apposing growth (Ter-

minalia catappa, Sterculia macrophylla).

The naked phase is usually not very long in the tropics, though

flowering or fruiting may even proceed during this spell, e.g. Pari-

shia insignis, Cratoxylon formosum, Firmiana fulgens, Parkia

javanica, Cochlospermum religiosum, and Oroxylum indicum. In

regions with a long dry spell, the number of such flowering trees

increases much more (TROUP ’21).

1. Transition from the intermittent evergreen to the deciduous

In Singapore the resting period of trees is rather short and some-

times indistinct. Usually the spell is one week or so, rarely a few

days only. Moreover there are some trees, in which new leafing

begins before leaf-fall has finished. Various irregularities occur too.

As an instance, a para-rubber tree in the Botanic Gardens

showed in March, 1944, from the top downwards, a transition of

new foliage, naked branches, tinted leaves and old green leaves,

and in April of the next year the same nuance of phases. The tree

took for the leaf-change of the entire crown nearly three months.

Swietenia macrophylla from Central America is, so to speak,

a deciduous tree, but most avenue trees of it have emitted new

brown shoots by the time the old leaves are falling off. Dyera cos-

tulata is deciduous, but I saw once on a big tree in the Gardens

that the old leaves remained on a few of the lowest limbs until

the next period of leaf-change. Trees tend in general to be more

irregular in their periodicity, the bigger they grow.

Thus the transition from the deciduous to the intermittent-ever-

green may be classified in four stages: —

(1) Holodeciduous: entire crown becomes bare at least for a

few days: Peltophorum pterocarpum.

Vol. XVII. (1958).

(2) Semideciduous: new and old foliage, as well as naked

branches, appear irregularly according to the height,

direction, etc. of branches: Hevea brasiliensis.

(3) Vice-deciduous: leaf-fall is finished during one week

promptly, but new shoots appear before old leaves fall

off completely: Swietenia macrophylla.

(4) Semi-evergreen: some branches do not shed leaves till

next season, tending to be intermittent-evergreen:

Dyera costulata.

Naturally these grades of the leaf-fall are not absolutely constant,

and may be shifted according to year or district. Deciduous trees

of N. Malaya can easily become semi-deciduous in Singapore, if

the season is not sufficiently dry.

Both the leaf-fall and new leafing occur in most trees nearly

simultaneously, but in some the leaf-fall proceeds gradually for

2—4 months, while the new leafing occurs at once, as in Sapium

discolor and Terminalia subspathulata. These cases indicate that

the new leafing does not necessarily follow on the formation of the

abscission layer of old leaves. Both of them are independent pro-

cesses (§ 18) and, if the former occurs earlier than the latter, the

tree is evergreen, while it is deciduous when the former is delayed.

2. The leaf-period and the date of leaf-fall and leafing

The duration of the leaf-bearing period of deciduous trees in

Singapore is three months to more than two years. They may

be seasonal or non-seasonal, and the latter may be contemporary.

Seasonal leafing or leaf-fall is in most cases once or twice in a

year, chiefly after a dry spell, rarely thrice. Non-seasonal leafing

is sometimes shorter than one year and sometimes longer accord-

ing to species. We may designate them as the “shorter” and

“longer” leaf-period of deciduous trees respectively.

(a) The average date of leafing of seasonally deciduous trees

(The names in brackets indicate exotic trees: e: early, m: middle,

and |: later part of the month given in Roman numerals) (HOLT-

TUM ’°40, CORNER ’40).

(1) Once every year (2) Twice every year

(Canarium rufum) Maier Pe (Couroupita guianensis) 15/TII, 20/IX

(Cariniana sp.) .. VIII,m | Elateriospermum tapos .. 1, VII-VIII

(Hymenaea courbaril) .. II Peltophorum pterocarpum 12/TI, 10/VIIl

(Kigelia pinnata) bes LA (Pentaspadon officinale) I-—-V, X-XI

Parkia javanica eae | ae Terminalia catappa .. 2/TT, 11/VHI

Terminalia subspathulata ... IV,e (3) Thrice every year

Ficus caulocarpa fen b> Vi, oe;

XI, ¢

Gardens Bulletin, S.

(b) The mean leaf-period of the non-seasonal deciduous trees

(HOLTTUM, lL.c.).

(1) Shorter Period Months (2) Longer Period Months

Ficus variegata .. .. | 6.1-6.5 | (Hevea brasiliensis) FA 13.3

Sterculia macrophylla .. | 6.9-7.0 | (Homalium graniflorum) .. 13.7

Adenanthera pavonina ae 7.25 | Anisoptera megistocarpa .. 13.8

(Delonix regia) .. Ls 8.8 | Cedrela glaziovi sis 14.0

Cratoxylon formosum Ee 9.1 | Caesalpinia ferrea a 14.2

Lagerstroemia flos-reginae .. | 8.1-9.4 | Parishia Maingayi it 15.6

Cassia fistula .. .. |9.1-10.5 | Heritiera elata ats 20.5

(Salmalia malabarica) .. |9.3-10.5 | (H. macrophylla). sia | LOST

Cassia nodosa .. a 1 |

§ 12. The flowering season

The flowering season may similarly be arranged into four

groups: continuous, seasonal, contemporary and non-seasonal.

Continuous flowering follows as a natural sequence in some

evergrowing trees, in the leaf-axils of which the flower buds are

formed, though evergrowing trees are not necessarily everflowering.

Seasonal flowering takes place 1-3 times every year according to

species, and in definite season. Contemporary flowering occurs,

though not at a definite season, in all trees of a species in the same

district simultaneously. Non-seasonal flowering is indefinite in the

calendar year and can vary by branches and stocks according to

species, so that branchwise and stockwise manifoldness appear.

Flowering occurs sometimes in consequence of the development

of new shoots, on which the flower is afterwards formed. This is

especially the case with the terminal inflorescence of the shoot of

each season. But in trees with axillary, ramiflorus, or cauliflorous

flowers no such relation exists or climatic conditions are the chief

cause.

Among trees with evergrowing shoots, about half flower conti-

nuously, while the rest are either seasonal, contemporary or non-

seasonal. Most manifoldly growing trees are also non-seasonal in

flowering, but some are seasonal in relation to the climate. Among

intermittently growing trees, the seasonal leafing shoot is also

seasonal in flowering, and on the non-seasonal leafing shoot fol-

lows generally the non-seasonal flowering, too.

Vol. XVII. (1958).

1. The flowering of evergrowing trees

Evergrowing trees are not all everflowering. If the flowers are

formed in the axils of twigs and open successively, they are ever-

flowering. But if the flowering is controlled by the weather, it may

tend to seasonal or contemporary flowering. If the shoot growth is

substituting and the inflorescence is terminal, the flowering may

become seasonless according to the shoot growth. And when the

formation of flower buds is sporadic, as in the lateral branches of

Anisophyllea disticha, so the flowering is optional or non-seasonal,

even if the branches grow continuously. The following instances

are the chief cases of flowering types of the evergrowing trees:—

(a) Everflowering: Carica papaya, Adinandra dumosa, Archy-

taea Vahlii, Commersonia bartramia, Glochidion spp.,

Macaranga spp., Hibiscus spp., Thespesia populnea,

Scaevola frutescens, Sesbania grandiflora, Leucaena

glauca, Mimosa sepiaria, Solanum Wrightii, Symplocos

fasciculata, Wormia_ suffruticosa, Psidium guajava,

Melastoma malabathricum.

(b) Seasonal flowering: Twice a year: Casuarina spp., Fag-

raea fragrans, Artocarpus, Morinda, Gardenia, Hors-

fieldia. Once a year: Hydnocarpus, Elaeocarpus spp.

chids.

(d) Non-seasonal flowering: Arthrophyllum, Bridelia, Canan-

gium, Leea, Mallotus, Rhizophora.

2. Seasonal flowering

This is intimately correlated with the climatic course, but two

cases must be distinguished. The one is that in which the leafing is

associated with the season, so that the flowering occurs as a conse-

quence also seasonally. The other is that the flowering is induced

directly by the seasonal change, independent of the leafing. To the

former belong the evergreen, the intermittent and the deciduous

trees either with the terminal or axillary inflorescence, and to the

latter the evergrowing and intermittent-growing trees with either

terminal, axillary or cauline flowers. Some instances are tabulated

below. The number in brackets indicates the frequency of flowering

in every year.

a. Seasonally leafing tree

Flower terminal: deciduous—Peltophorum pterocarpum (2).

Jf Eugenia grandis (3).

( evergreen \ Garcinia mangostana (2).

Flower axillary

deciduous—Terminalia catappa (2).

Gardens Bulletin, S.

b. Seasonally flowering tree

Flower terminal—Fagraea auriculata (1).

Flower axillary—Fagraea fragrans (2), Casuarina spp. (2),

Erythrina subumbrans (1).

Flower ramiflorous—Horsfieldia sylvestris (2).

Flower cauliflorus—Durio zibethinus (2).

Casuarina equisetifolia is distributed from Australia to the

eastern coast of India, adapted to typical monsoon regions.

So, even under evergrowing conditions in Singapore, it shows

two flowering seasons, probably as a_ specific character.

The flowering of Fagrawa fragrans is however a little different.

The flower-buds are formed in the dry spells of February

and July and open in May and October. The flowering

is especially conspicuous in May (HOLTTUM ’35), but it

fluctuates more or less according to the year. For instance, in

February, 1945, it was rainy and a short spell of rainless days

came in April, so that most trees were seen in flower July to

August. In big trees, however, some branches flowered in May

tending to be slightly manifold. Besides, some young trees flowered

in March or in December of the previous year; thus even a stock-

wise difference was noticed. For the anthogenesis of Fagraea there

is necessary, therefore, the timely co-operation of the internal ten-

dency with the external climatic course.

Generally, if the formation of flower-buds is controlled by

weather and in one season the weather is not effective enough, so

the trees can not react uniformly all together. In some stocks or

branches the bud-formation can diminish or not occur. Then this

irregularity can exert a compensatory after-effect at the next season,

so that even seasonal flowering tends to be seasonless or manifold.

3. Contemporary flowering

In this case the young buds formed can not flower right away;

they rest in a certain stage of development, waiting for the sudden

fall of temperature by a shower, by which they are stimulated to

grow anew, and after a certain number of days all flowers of the

district open gregariously. They are called the temperature-flowers.

(CORNER, l.c.p. 38).

The most conspicuous gregarious flowering was studied with

Dendrobium crumenatum, an epiphytic orchid, in detail (COSTER

°26, KUIJPER ’33). Among trees and shrubs, Pterocarpus indicus,

Murraya paniculata, Randia macrantha, Stenolobium stans, and

Coffea spp. may be named (HOLTTUM ’40, CORNER 740).

The inflorescence of Pterocarpus follows on the new leaf-growth;

Vol. XVII. (1958).

the buds are formed in successive stages of development, and the

old buds react to the temperature fall, while the younger ones wait

for the coming chance, so that the flowering of an inflorescence is

divided into several stages with pauses of a few days or more. The

flowering of Pterocarpus tinctorius of Africa is also divided into

four periods (v. NOLDE ’40). Stenolobium stans forms the inflo-

rescence throughout the year and one can see gregarious flowering

about ten times every year. Besides, Flacourtia rukam shows 2-3

times and Rhodamnia trinervia 4—5 times, flowering and fruiting.

4. Non-seasonal flowering

This is seen in shoots and stocks of manifold growth with the

inflorescence either terminal, axillary or ramiflorous.

a. Branch-growth manifold

( terminal—Cassia splendens, Vitex, Kopsia,

Inflorescence Txora.

| axillary—Michelia.

b. Stock-growth manifold

Inflorescence terminal—Lagerstroemia flos-reginae.

Anthocladium terminal—Delonix regia.

Inflorescence ramiflorous—Cassia fistula.

Branch and stockgrowth manifold—inflorescence terminal—

Spathodea.

The shoot of Kopsia produces a terminal inflorescence after

some pairs of leaves are formed, and, while the flower-buds are

still young, the lateral branches grow out to a tolerable size. This

mode of growth is repeated irrespective of the season, so that both

the flowering and the shoot-growth go on non-seasonally together.

In K. singapurensis, though the flowering season predominates in

two seasons every year, yet one can see some flowers at any time,

and in K. fruticosa the flowers are seen throughout the year.

Thevetia, Ochrosia and Cerbera of the same family, and Ixora and

Vitex show similar behaviour.

Spathodea produces a terminal inflorescence on each substituting

new shoot, and each inflorescence bears scores of buds opening

_ Successively, so that old trees are decorated with big flame-red

flowers all over the crown the year round.

Gardens Bulletin, S-

Among the four groups just mentioned, the everflowering and

the non-seasonal ones are controlled inherently, while the contem-

porary and seasonal ones, though inherent originally, are actually

induced at the last step by some trifling change of the weather.

In the humid region of the temperate zone, the formation of

flower-buds is controlled generally by the temperature and photo-

period, and in trees it is induced chiefly by the seasonal change of

temperature, while a dry spell, even if it may affect the abundance

of flowers, does not control anthogenesis. In Singapore, on the

contrary, there is no seasonal change of temperature and photo-

period, but the length of the dry spell, accompanied by intensified

insolation and thermoperiod, and the sudden fall of temperature

during a shower are the chief meteorological conditions. Some

species of trees are sensitive to such slight weather changes for

their anthogenesis.

§ 13. The fruiting season

The ripening of fruit after flowering takes usually 2-5 months

according to the species. In some cases flowering is the morpholo-

gical sequence of the leafing, but in others not. In the former case,

leafing, flowering and fruiting are three consecutive processes,

while in the latter the season of fruiting is subject only to the

flowering. But the ripening of fruits is sometimes affected by their

position on the branches or the inflorescence. For instance, the

raceme of Adenanthera pavonina develops soon after the leafing

of every seven months period (§ 11-2), and it flowers gradually

during two months, but the ripening of pods continues for several

months, so that the glossy red seeds are dispersed nearly through-

out the year.

In Singapore anthogenesis is induced chiefly in the dry spells of

February and July; the flowering follows two months later, and the

fruiting is most abundant in June to July and January. This is re-

markable in durian, mangosteen and rambutan, though one and

the same tree does not produce every season regularly. According

to OCHSE (731), these fruits appear in Java only once every year

with longer duration than in Singapore.

The fruiting season of the everflowering Averrhoa and Psidium

and of the manifoldly flowering Achras is extremely long, so to

speak nearly non-seasonal, though with fluctuation. Flacourtia

rukam fruits 2—3 times contemporarily. Papaya, Musa and Ananas

show stockwise manifoldness of fruiting, so that they serve as

good table-fruits throughout the year.

Vol. XVII. (1958).

§ 14. The frequency of species in the four types of periodicity

The number of species I could examine personally or extract

from the literature (RIDLEY ’22—24, CORNER 740) amount to

543, belonging to 298 (211 + 87 exotic) genera and 73 families

(after ENGLER’s system). There are still 13 other families in

Malaya containing trees, which I could not investigate.* Achras,

Aleurites, Anacardium, Anona, Carica, Ceiba, Delonix, Hevea,

Jatropha, Leucaena, Muntingia, Plumiera, Psidium, Sesbania,

Spathodea, Thevetia and others are included as exotic genera.

Under the exotic species observed, those that are not common in

Malaya are all those planted in the Botanic Gardens, Singapore.

Among exotic trees enumerated in the Appendix, there are four

families not indigenous to Malaya, namely Salicaceae, Bixaceae,

Papayaceae and Cactaceae. Besides, there are some families or

genera in which only exotic species are treated, though the families

or genera themselves are indigenous to Malaya, as Pittosporaceae,

Polygonaceae, Proteaceae, Solanaceae, Thymeliaceae, as well as

various genera.

Naturally there is no boundary between shrub and tree, and

there are many shrubs that grow continually just like young trees,

but shrubs as Lantana and Ardisia are omitted from the list, as

there are many trees in these families, while big evergrowing

shrubs of Goodeniaceae, Solanaceae and Asclepiadaceae are

added.

Among the Malayan families, those containing evergrowing trees

amount to 35, or nearly the half of the total families examined.

Families containing deciduous trees amount to 32, or about the

same as those of the evergrowing ones. Compared with them, the

families containing intermittently growing trees are the most abun-

dant, namely 48, or including the exotic families 52. The number

of species in each group was as follows (exotic species added in

brackets), though sharp boundaries between the growth-forms do

not exist: —

evergrowing — manifold intermittent deciduous

103 (46) 14 (26) 154 (60) 108 (31)

It may be remarked, however, that the deciduous trees are all

easily noticeable in contrast with the evergrowing trees, while the

latter are not distinguishable unless actually examined, so that the

number of the latter given above seems distinctly less than the

actual rate. Concerning the intermittently growing trees, as the

writer observed only a few species of the genera which contain

* Gonystylaceae, Staphyleaceae, Sabiaceae, Samydaceae, Styraceae, Acan-

thaceae, Caprifoliaceae, Crypteroniaceae, Cunoniaceae, Epacridaceae, Monj-

miaceae, Hermandiaceae, Opiliaceae.

Gardens Bulletin, S.

many similar species, the number given above is also distinctly

less than the actual rate. The manifoldly growing trees are much

more numerous in exotic species than in the Malayan (26:14).

At any rate it may be asserted that among all tree-species of

Malaya the evergrowing trees, even with the addition of Palma-

ceae, Coniferae and tree-ferns, amount to less than 20 per cent,

probably 15 per cent. The deciduous trees total about 5 per cent

over the whole Malay Peninsula, though some of them do not

shed leaves in Singapore. The remaining 75 per cent are the usual

evergreen trees with intermittent growth.

In Java, according to KOORDERS (798) the deciduous trees

amount to about 5 per cent among indigenous ones, distributed

chiefly in the eastern and middle Java (FABER °35, p. 374). In

Ceylon according to WRIGHT (05, p. 463), deciduous trees

make about 14 per cent among 650 indigenous species and 6 per

cent (17 species) among endemic species.

The occurrence of deciduous trees in dicotyledonous families is

fairly even, just like the evergrowing trees, being more abundant

in big families (see Appendix).

How such a circumstance has been derived, is a thesis to be

discussed further. If one observes the deciduous trees more closely,

they are not all mesophytic; there are even hygrophilous-decidu-

ous trees living in swampy ground, and contrarily there are some

hygrophobic trees unable to grow in as wet a district as Singapore.

From these facts, the interrelation of the deciduous trees and the

environment seems much more indirect.

§ 15. Comparison of the leaf-fall and leafing periods of Singapore

with those of Ceylon and Java

So far we have dealt with the behaviour of tree-growth in Singa-

pore and Malaya. Now, if we compare its features with those of

Ceylon and Java, it is very instructive to discover what characters

of periodicity are easily affected by external influences. The most

remarkable difference among the deciduous trees in these three

regions is the calendar data of the leafing related to the monsoon.

From India to Indo-China the rainy season extends from May

to October, especially rainy from June to September, and the dry

season in winter from November to April. Temperature is relatively

low in the first part of the dry season from December to January,

while it is very hot from March to May on account of fine weather,

but then, despite the sun’s higher position, it gets cooler on ac-

count of the cloudy sky (WRIGHT ’94, TROUP ’21, CHAM-

PION °36). These changes of seasonal climate deviate naturally

Vol. XVII. (1958).

with the distance and direction of the ocean as well as the moun-

tain range. For instance in Ceylon, the south-west monsoon pre-

dominates from February to July and the north-east monsoon

from August to January, so that the seasonal change is three

months earlier, and moreover it differs according to the south-

western side and north-eastern side of the island (WRIGHT lL.c.).

But as a whole, the general feature of the seasonal change in

S.E. Asia is as mentioned above.

In India the- deciduous trees of northern origin shed their leaves.

usually during the winter, those of southern origin mainly in the

second half of the dry season, indicating the hereditary habit of

their birth-place. In Peradeniya (473 m.) February is the most

rainless and dryest season, and about half of the deciduous trees

(47 per cent) shed their leaves in this month; before and after

that month the proportion decreases gradually until in the rainy

season of September it attains the minimum (2 per cent)

(WRIGHT, l.c.).

In Buitenzorg the monthly mean maximum temperature is 29°—

31° and the mean minimum 21°—22:5°, so that there is no verit-

able seasonal effect on the growth of the plant (SCHWEIZER

*32). Of the rain-fall, however, there is a notable fluctuation,

though it is sufficient all over the year. From July to September

the mean monthly rain-fall amounts to 135 mm., while in October

it is 706 mm. On the other hand, in Soemaran, Middle Java,

while in February the rain-fall amounts to 300 mm., in July to

August it is less than 10 mm., and eastwards it diminishes more.

In Pasoeruan, East Java, from August to September the mean

rain-fall amounts to a few millimetres only (COSTER ’23). East-

wards in Lesser Sunda Islands it diminishes more and more, with

a longer dry spell, so that it tends to be the typical monsoon

climate (RENSCH ’30).

Thus the dry and rainy seasons are in Ceylon and Java diamet-

rically opposite, while in Singapore, situated in the intermediate

region, the climate is mild and non-seasonal. It is instructive there-

fore to compare the leaf-period of these three regions.

The leaf-period of various trees has been reported in India,

Ceylon and Burma by WRIGHT (794), TROUP (’21), STAMP

(25), CHAMPION (’36), in Java by VOLKENS (712), SIMON

(14), COSTER (’23), and in Singapore by HOLTTUM (’31,

°40), CORNER (’40) and others. If we compare the leafing

period or the leaf-fall season of some deciduous trees reported

commonly in these regions, we can find a striking contrast be-

tween them, as has been stated by many authors already.

Gardens Bulletin, S.

TABLE 5

The leafing or leaf-fall period in Singapore, Ceylon and Java. after

HOLTTUM, COSTER and WRIGHT, I.c. (Month in Roman number).

SINGAPORE CEYLON EAST JAVA

Trees + te

leaf-period leaf-fall bare leaf-fall

(months) period days period

Adenanthera pavonina .. | 7.25 (6.0-8.5) X-XII 30 -

Delonix regia se 8.8 (7.7-10.6) IV-V wae VIII-X

Lagerstroemia flos-reginae 9.2 (8.9-9.4) XII-I fz | eae

Salmalia malabarica .. 99: 93-105) XII-I ca. 3 VII-X

months

Cassia fistula .. .. | 10.6 (9.1-10.5) II-Ill 18 VIII-X

3, nodosa . | 17 Ute) III —— V-VII

Tamarindus indica a» | 129 (400-17.7) III-IV —— | VIII-XI

These trees in Singapore are, except Tamarindus, in their leaf-

period all shorter than one year, while in other districts the leaf-

fall is one year regularly with a shift of nearly half a year in the

leaf-fall season. According to these data, the difference of climate

has two effects on the periodicity—the shifting of the period as

well as the leaf-duration. In other words, the active period of the

tree-growth is transformed rather flexibly by environmental con-

ditions. This is also the case with the grade of leaf-fall (§ 16),

while the bud-forming ability, as we will see later (§ 23), is rather

constant.

The vegetative period seems, therefore, the most easily movable

character of trees in accordance with the change of climatic cycle.

According to WRIGHT (’05, p. 7) the flowering season of Acacia

dealbata, indigenous to south-eastern Australia, has been entirely

shifted following the climatic cycle in the Nilgiris in India. It

flowered from 1845 to 1850 in October just as in Australia, but

then the flowering season was gradually advanced, in 1860 to

September, in 1870 to August, in 1878 to July and lastly in 1882

to June, just the spring season of the new habitat. It took about

forty years for the seasonal acclimatization. Whether a tree is

able to acclimatize by lengthening the active period, the writer

could not find a suitable example, though it seems not impossible

according to the circumstances. In the trees mentioned in Table 5

Vol. XVII. (1958).

(above), it is not known in what sequence they have had their

active period transformed.

In non-seasonally leafing trees in Singapore, some are shorter

and others longer ‘than one year in their leaf-period ($ 11- im

Cratoxylon formosum in Singapore has a leaf-period of 9:1

months, while in Penang (HOLTTUM ’40) and even in East

Johore (CORNER ’40) it shows a one year cycle normally,

though the bare spell is not known. Probably the real vegetative

period is longer than in Singapore. In contrast, the Para-rubber

tree has in Singapore 13-3 months leaf-period, while it shows even

in North Malaya 12 months regularly and in Ceara (4° S), Brazil,

it stands two months in the resting condition. Singapore seems in

this respect to be a favourable habitat for rubber production. For

trees with a period of more than 13-7 months, we have no data

from other districts to compare.

If one assumes that the leaf-period including the rest period* of

deciduous trees is one year normally, except those with a half-year

period, then this habit is maintained in those districts with dis-

tinctly dry or cold seasons irrespective of the length of the rest-

period, and the periodicity in Singapore is shortened or lengthened

according to the species as stated above. But in those regions with

a long dry season, the vegetative period becomes far shorter than

one year, so that the same trees with a short leaf-period at Singa-

pore might display their proper length of activity, while those with

a longer period might have lengthened their activity under the

favourable conditions of Singapore. But if the active period of

trees in Singapore be their natural length, then the length in dryer

districts should be a forced one destined to become one year.

Very interesting in this respect is the fact that Terminalia

catappa, which has a half-year period in the tropics and sheds

leaves in Singapore about January to February and July or August,

tends in Rio de Janeiro (23° S), according to a letter from Mr.

Corner, to be of one year cycle with a long rest period in winter

(May to October) when the leaves are gradually shed: it has,

therefore, six months vegetative period grading into six months

rest. The length of rest period seems, therefore, to be a flexible

character, by adjustment of which the shifting of seasonal change

is attained. Besides, there may be some trees in which the length

of their activity has already become inherent at each habitat.

Trees of northern origin, as Fagus, Carpinus, Ulmus, etc., are

provided probably with a one-year period, but those of southern

origin as Ficus, Mallotus, Albizzia, of which some allied species

* The flowering and fruiting may occur during either the leaf- period

or the resting period according to the species.

Gardens Bulletin, S.

in the tropics grow continually, produce in the temperate zone

their leaves until summer and close the shoot into a dormant bud

in autumn. In these trees it is not clear whether the annual perio-

dicity is inherent or induced. Leaves have their own life-time, but

the shoot can continue its growth so far as the bud does not close.

§ 16. The grade of leaf-fall of trees in various districts

In Singapore, not only the leaf-fall and the length of active

period, but also the grade of leaf-fall, are different from those of

the other districts. Many species of trees occur on the Continent,

in Malaysia and in other regions, aboriginally or by introduction.

If one compares the behaviour of leaf-fall in these regions, the

Telative habit of trees against the dry season can be seen.

(1) Evergrowing in Singapore, but deciduous in other dis-

tricts :—

Trema orientalis (Himalaya to Polynesia).

Ficus elastica (Himalaya to Java. Leaf-fall in Ceylon

II-IV irregularly).

Duabanga sonneratioides (India to N. Malaya. Leaf-

fall in India in IV, the hot season).

(2) Exotic trees evergrowing or evergreen in Singapore, but

deciduous in original districts: —

Melia azedarach (N. India).

Antidesma bunius (Himalaya to Australia).

Sapindus mukorosii (China).

Mimusops elengi (India, Burma, Ceylon).

Tectona grandis (S.E. Asia. Leaf-fall in India XJ-HI,

in Java VII-IX).

Tectona hamiltoniana (Burma, deciduous in _ hot

season).

(3) In Singapore semi-deciduous, in other districts deciduous :—

Ficus religiosa (India).

Tamarindus indica (Africa to Asia).

Albizzia odoratissima (Himalaya to Siam).

Dillenia indica (India, Siam, Malaya).

(4) Deciduous in N. Malaya, evergreen in S. Malaya: —

Elateriospermum tapos (W. Malaysia).

Dillenia ovata (China, Siam, W. Malaysia).

The effect of the dry season on the leaf-fall is clearly to be seen.

Vol. XVII. (1958).

§ 17. The deciduous habit and the height of crown

Most shrubs and young trees in the tropics are evergrowing,

while tall trees are often deciduous. It is stated that in the highest

stratum of the primary forest in the tropics many deciduous trees

are to be seen (RICHARDS ’52). Indeed, with the raising of the

crown, not only the ascent of sap gets difficult, but also the insola-

tion and wind are far more intense than in the lower strata, so

that the lifting high over the forest canopy may induce the decidu-

ous habit or the habit may relate with the height of crown. Dyera

and Irvingia are instances of deciduous trees more than 200

feet high. But there are also tall evergreen trees of the same

height, as Balanocarpus and Dryobalanops. Trees projecting higher

than 100 feet are more numerous among evergrowing trees than

deciduous ones in Malaya and India (CORNER *40, TROUP ’21),

though this may be due chiefly to the abundance of evergreen

species in the tropics. Tall trees in the same family are sometimes

evergreen and sometimes deciduous as shown below. (Deciduous

trees and the height of trees in feet in brackets).

Juglandaceae: Engelhardia spicata (100), (E. nudiflora 100);

Moraceae: Artocarpus elasticus (150), (Antiaris toxicaria

a TS) *

Leguminosae: Parkia speciosa (150), (Koompassia excelsa

150-260);

Anacardiaceae: Melanorrhoea Wallichiana (150), (Melanorr-

hoea sp. 150, Pentaspadon 2 spp. 150);

Sterculiaceae: Pterospermum javanicum (150), (Sterculia

macrophylla 150).

It may be understood that tall trees are not all deciduous.

Moreover there are many deciduous trees of relatively low height,

as for instance: Excoecaria agallocha (30), Cassia fistula (30),

Sterculia rubiginosa (30), Flacourtia rukam (40), Lannea gran-

dis (50) and many others.

Even if the deciduous habit is fitted for the highest story of the

forest, the bare period is usually short, and in wet regions or in the

wet season it seems rather a waste of time for photosynthesis. If

one examines the deciduous trees more closely, it becomes clear

that the deciduous habit is not merely related with dryness alone.

Viewed from their distribution in the tropics, the deciduous trees.

are not all xerophobic, on the contrary some are even hygrophobic

as stated below.

Gardens Bulletin, S.

§ 18. Deciduous trees and their habitats

In the temperate zone the texture of the leaves of deciduous

trees and of evergreens are clearly distinguishable, namely mem-

branous versus coriaceous, while in the tropics of uniform climate

both kinds of leaf-texture are found as a generic character either

in deciduous or evergreen trees, whether their habitat be wet or

dry. One can not decide, therefore, from the leaf-texture aione

either habit or habitat. This may be attributed to the circumstance

that, under uniform climate, trees are allowed to grow continually,

irrespective of their leaf-texture. On the other hand, even if the

leaf is coriaceous, it tends to be deciduous, if the abscission layer

is formed earlier than the new leafing, as in Ficus caulocarpa and

others.

Most deciduous trees are mesophytic, but there are also some

hygrophilous and hygrophobic trees, though the water-condition of

the soil and of the air is not necessarily the same.

Hygrophilous deciduous trees indigenous to Malaya, growing in

lowland forest or swampy ground are Anisoptera megistocarpa,

Beilschmiedea malaccensis, Canarium rufum, Excoecaria agallo-

cha, Mallotus sp., Terminalia phellocarpa, and T. subspathulata.

‘The Mallotus sp. is an undergrowth tree deciduous in the swampy

forest and shows that the deciduous habit can develop even in

wet forest. Excoecaria grows in the mangrove zone. Among trees

of wide distribution, some are hygrophilous and deciduous. For

instance, Cedrela toona grows on wet soil of valleys and river-

sides; Bischofia javanica is a deciduous tree, which however tends

to be evergreen in wet ground (TROUP ’21). The teak-tree of

monsoon regions tends to be semi-deciduous on riversides in the

flood-zone (SIMON 714, COSTER ’23), but in Singapore its

growth is inferior and manifold (§ 8-2). Probably the shoot is

hygrophobic, but the root is slightly hygrophilous. Trees which are

semi-deciduous or evergrowing in S. Malaya, but deciduous in

other districts, seem generally to be mesophytic or slightly xero-

phobic.

Among deciduous trees there are some which seem rather hygro-

phobic. SCHIMPER (98, p. 262) stated that some trees shed

leaves in the wet season. In Ceylon there are also about 2 per cent

of deciduous trees, which shed leaves in the September rainy

season (WRIGHT, ’05). In Java Spondias mangifera sheds leaves

in the wet season of January to April, and Tetramelis nudiflora in

December (SIMON 714). In Trengganu (Malaya), an unidentified

species of Melanorrhoea sheds its leaves in October and Novem-

ber at the beginning of the rainy season (CORNER, ’40).

Vol. XVII. (1958).

Leaf-fall occurs, not only after senescence, but also after a few

days in the dark or under a wet bell-jar, in high temperature or

other abnormal conditions. In the tropics the rainy season is often

_ accompanied by damp and gloomy weather, so SIMON imagined

that the leaf-fall may be attributed partly to these conditions.

Naturally such a habit seems rather rare among deciduous trees in

general, but there are still some that, considered from their distri-

bution, seem to be hygrophobic. They are trees, which, in spite of

their distribution widely in S.E. Asia or Malaysia or further to

Australia or Africa, do not occur in Malaya or S. Malaya, as for

instance :—

Ailanthus malabarica: \ndia to N. Malaya.

Albizzia lebbec: Africa to India and Australia.

Alstonia scholaris: Ceylon to N. Malaya, Java and Philippines.

Anthocephalus cadamba: India to north of Johore, Java to

New Guinea.

Butea frondosa: India to China and Java, but not in Malaya.

Dillenia aurea: India to N. Malaya and Java to Philippines.

Diospyros montana: India, Australia.

Feronia limonia: India, Siam to N. Malaya.

Flacourtia indica: India, Malaysia.

Garuga pinnata: Himalaya to Siam, Java and eastwards.

Hymenodictyon excelsum: India to N. Malaya, Java to Philip-

pines. ;

Neesia altissima: Siam to Penang, West Malaysia.

Oroxylum indicum: India, China to N. Malaya, Malaysia to

Philippines.

Tetrameles nudiflora: India to N. Malaya, Java to Timor.

Ailanthus flowers and fruits in Penang, but it does not thrive in

S. Malaya even under cultivation (BURKILL °35). Tetrameles

Sheds its leaves in the rainy season, as mentioned above.

Roughly speaking, the deciduous habit is an internal process, in

which the abscission layer is formed in a quick tempo. Its final

step may be precipitated by some physiological disturbance as

dryness, wetness or other condition according to the species. In a

non-seasonal warm climate the habit does not matter very much,

but in the temperate or monsoon regions it may have a survival

value.

Gardens Bulletin, S.

§ 19. The systematic occurrence of deciduous trees

Among 73 dicotyledonous families that the writer could investi-

gate, there are naturally many families which contain no decidu-

ous tree, such as Casuarinaceae, Elaeocarpaceae, Fagaceae, Loga-

niaceae. Magnoliacede. Malvaceae, Myricaceae, Myristicaceae,

Myrsinaceae, Myrtaceae, Rhamnaceae, Rhizophoraceae, Symplo-

caceae, Theaceae and some other small families (see Appendix),

though most of the families, if they occur in temperate regions,

contain deciduous trees.

Now, the deciduous trees in the tropics are distributed in various.

families fairly evenly, just as are the evergrowing ones. Families

rich in genera and species are generally rich also in deciduous

trees, though not proportionally.

Some genera contain several deciduous trees. For instance, Arto-

carpus (contains 2 deciduous species), Ficus (5), Parishia (4),

Spondias (4), Dillenia (3), Cratoxylon (3), Terminalia (5), and

Alstonia (6), that is, 8 genera, in 6 families. But these families

contain also evergrowing species, except Guttiferae, indicating how

easily the periodic habit can vary even in one and the same family.

Among the 73 families investigated, 17 Malayan families con-

tain both evergrowing and deciduous trees, namely: Anacardia-

ceae, Apocynaceae, Bombacaceae, Combretaceae, Dilleniaceae,

Dipterocarpaceae, Euphorbiaceae, Hamamelidaceae, Lauraceae,

Leguminosae, Lythraceae, Meliaceae, Moraceae, Rubiaceae, Sapo-

taceae, Sterculiaceae and Ulmaceae. If one takes exotic trees into

account, the following 8 families may be added: Anonaceae,

Bignoniaceae, Flacourtiaceae, Guttiferae, Lecythidaceae, Ruta-

ceae, Sapindaceae and Tiliaceae.

Of these families, the following genera contain trees of both

forms of growth: Gironniera, Ficus, Bridelia and Mallotus. The

genera which contain a few deciduous trees among intermittently

growing ones are many (see Appendix).

Besides, there are a few tropical genera which contain among

many evergreen trees only one deciduous species:—

TABLE 6

NUMBER OF SPECIES

Species -————__——_———__| Dediduous

In the world | In Malaya

Beilschmiedia (malaccensis) al | 20 | 15 | 1

Dalbergia (loiveri) > <n 120 15 1

Canarium (rufum) BK Ja4 100 12 1

Mallotus sp. ig 22 100 ) 25 1

Lucuma (Maingayi) .. om, 50 8 1

Vol. XVII. (1958).

These trees are therefore, so far as the Malayan species are con-

cerned, the single deciduous species of each genus.

Moreover, even in the tropical families Anonaceae and Diptero-

carpaceae, some deciduous trees are seen, and in the genus Jlex,

most species of which are evergreen even in the temperate zone,

I. macrocarpa in Malaya is semideciduous. There are also a few

monotypic genera with deciduous species, namely: Elateriosper-

mum tapos of W. Malaysia, which is in §. Malaya semideciduous,

and Feronia limonia, which is distributed from India to the north-

ern boundary of Malaya, is deciduous. Further in some genera of

the tropics, only the single Malayan species is deciduous, though

it is not known whether the other species of each genus are also

deciduous or not:

Altingia excelsa: 2 spp. in the tropics.

Irvingia malayana: 4 spp.

Trigonachras acuta: 9 spp.

Tetrameles nudiflora: 4 spp.

As these cases indicate, deciduous trees occur even in a wet

and warm climate, so the origin of the inherent deciduous factor

seems not very complicated and can be developed in various

families and genera rather optionally. The formation of the abscis-

sion layer before new leafing is its fundamental condition, and the

habit can be continued so long as it is not a handicap in the

struggle for existence.

§ 20. The geographical distribution of deciduous trees

In Malaya there are endemic deciduous trees, some of which

‘spread all over Malaya, but others are confined to the northern or

southern part of the peninsula. Besides, there are indigenous deci-

duous trees common to S.E. Asia or Malaysia. With Sumatra the

affinity is the most intimate, but there are also trees common to

the Philippines, New Guinea, and the Pacific Islands to Australia.

On the other hand, there are also trees common to islands of the

Indian Ocean and Africa, and a few show the most wide distribu-

tion over all these regions.

If most Malayan deciduous trees are the same as those of the

Continent, we may assume that most of them have migrated from

S.E. Asia, where a distinct dry season predominates, whereas if

the trees in common are few, the relation of the deciduous habit

with dryness is not essential. On the other hand, if the Malayan

Gardens Bulletin, S.

elements have more in common with those of Malaysia and re-

moter districts, it seems more probable that at least some Malayan

elements may be hygrophilous deciduous trees common to the

Oceanic Islands, though even in Malaysia there are wide regions

with a long dry spell. If, again, endemic deciduous trees are abun-

dant, the habit seems to have developed of itself by mutation.

Naturally such a test tells nothing about the actual course, yet it

is not without interest to see the trend of the deciduous habit.

For this purpose the writer divided the deciduous trees of

Malaya into three Groups: (1) those endemic to Malaya, (2)

those common with the Continent, and (3) those common with

the Pacific region, each with subdivisions.

No. of

1) Endemic species of Malaya species Total

a: throughout the Malay Peninsula ite i

b: the northern half .. ™. vi = a

c: the southern half .. S¢ ot. aoe 6...)

2) Common to the Continent

a: S.E. Asia to Philippines, New Guinea or Pacific Islands 12- }

b: S.E. Asia or Africa to Australia 7 ae

c: S.E. Asia to Malaysia a va eee (

d: S.E. Asia to Malay Peninsula .. es . ae. §

3) Common to the Pacific Islands

a: Indian Ocean to the Pacific .. a2 _ ae

b: Malaysia to the Pacific MS sat df 1 2%

c: Malaysia ‘z - AF 3

d: West Malaysia ug ae Re PRLS ef

The number of species in these groups is in ratio of 32: 49: 26.

In Malaya therefore the deciduous species common with those of

the Continent are much more than the endemic ones (49: 32),

but the relation with the Malaysia and Oceanic Islands is not

much less, or the difference is not very significant. The groups,

here classified, are however based only on a territorial distinction.

But in E. Malaysia there are some islands with distinctly dry

season (RENSCH ’30) and Oceanic Islands have few species in

common with Malaya. The most similar region, both in the climate

as well as in the species in common, is W. Malaysia. If one divides

all the named regions roughly into two, the wet and the dry, so

the number of species of the former, including Malaya and W.

Malaysia amounts to 32 + 20 = 52, and that of the rest distri-

buted in more or less dry regions amounts to 55, so that the both

groups are nearly equal. In this respect the deciduous trees of

Malaya tend in their affinities neither to the dry, nor to the wet

regions.

Vol. XVII. (1958).

But if a species continues its distribution after having acquired

the deciduous habit, it is not enough to relate this to the present cii-

mate alone. It is necessary to consider conditions in the past. Let

us examine, therefore, the past geological history of the Malay

Peninsula and see whether there were some changes that might

have induced the deciduous habit.

§ 21. The geological history of the Malay Peninsula

Malaya is not an ancient peninsula. It originated, together with

W. Malaysia in the Cretaceous. The foundation of the Peninsula

consists of conglomerate, the materials of which flowed down from

the Gondwanaland and spread over both the northern and southern

Hemispheres in the Mesozoic (SCRIVENOR ’28). Since then,

until the mid-Tertiary, the climate of the eastern Hemisphere was

mild and tropical plants ranged widely from Asia to Europe.

Fossil plants embedded in London clay of the Old Tertiary are

common to families and genera of South China and Malaya to-

day and even several closely related species were found (REID and

CHANDLER ’33). The climatic gradient from south to north of

these regions was gentle and probably warm and cloudy, so that

even the tropical zone was not very hot, as at present, and the

temperate zone was not very cold in winter.

When the winter is not very cold, the tropical flora can flourish,

as the summer is warm enough for growth. The tropical rain forest

in Asia spreads at present over 28° north latitude from north-

eastern Burma to the Himalayan district. At Sibsagar (27° N.),

where the tropical rain-forest still predominates, the mean mini-

mum in January is 49° F., and in the extreme attains 42° F. The

rain-fall amounts to 95-5 inches (— 242-5 cm.), but chiefly in

~ summer, while in December only 0-5 inches (= 1:3 cm.) falls,

and yet there exists the rain-forest (CHAMPION ’36). According

to BROOKS (’22) the mild climate in the arctic regions of the

Tertiary would have been attained by increasing inflow of warm

surface-water from the tropics to the pole by topographical chan-

ges, even without the change of the earth’s axis or continental

drift.

In the Miocene Malaya made up, together with the West Malay-

sian Islands, the so-called Sundaland, faced eastwards to the Sahul

Shelf including New Guinea and Australia, with which it was con-

nected sometimes by a land, sometimes by a chain of archipelagoes

(MERRILL °23, LAM °’27, ARLDT ’38). In the Pleistocene,

Java separated first, then Borneo and Sumatra and lastly the

Malay Islands which later were connected with the Continent.

Gardens Bulletin, S.

Such a chronological sequence of W. Malaysia is shown well in

the floristic affinity of its parts. So far as the Malay Peninsula is

concerned, the difference of the floristic composition north and

south of 77° N. makes the estimation probable (SYMINGTON

43). In the Pleistocene the Glacial Periods were repeated, and

migration and changes of growth-form were also striking.

During such bewildering changes, what kind of trees gained the

ability to form the resting bud is not clear. Considering the north-

ern hemisphere as a whole, trees with the ramifying growth, as

Ginkgo, seemed easily to endure the unfavourable season by form-

ing resting buds. Most conifers continued their monopodial growth

by forming buds in winter. Dadoxylon indicum, a Permian conifer

of Gondwanaland in India, shows distinct growth-rings in the

trunk (SEWARD °33). Also some fossil-woods with growth-rings

have been found in Europe and Alaska (BROOKS ’22), though

most of the ancient fossil-woods show ringless wood. Naturally,

even in trees with the intermittent growth, so far as the shoot-

growth is manifold, the formation of growth-rings tends to be in-

distinct (COSTER ’27, ’28), so that the presence or absence of

growth-rings alone can not give the evidence of intermittent

growth and bud-formation.

In the era of Sundaland, the climate of Malaya was probably

like present-day Burma (compare the maps in MERRILL °23 and

LAM °27), where from the tropical wet forest along the western

coast to the thorn-forest of the midland several transitional vege-

tation are seen (STAMP ’25, CHAMPION 736) (§ 18). At least

there would have existed some districts with a long dry spell in

the Sunda-midland of that era. Therefore, if some trees originated

in Sundaland, like the Dipterocarpaceae, the birth place of which

is inferred to lie between Borneo and Malaya (SYMINGTON

l.c.), they may have acquired the deciduous habit in spite of the

tropical origin. In this respect, the comparative study of resting

buds and growth behaviour in general of tropical trees may tell us

the previous history of their ancestors.

But, since the formation of the Malay Peninsula, there have

elapsed ten thousand years or more, and, considered from the

viewpoint of adaptation, it seems rather strange to have perpe-

tuated for so long a time the deciduous habit in a uniform climate.

Perhaps among the genetic characters some are easily formed but

not easily reversed, while others behave inversely, though we do

not know how the deciduous and bud-forming characters are rea-

lized as internal processes. The transformation of the bud-forming

Vol. XVIT. (1958).

character within a certain systematic group in respect to the

range of distribution tells us however something about the evo-

lutionary change of the growth habit of trees indirectly (§ 24).

§ 22. The transition of shoot-growth from the evergrowing to the

deciduous

The growth process, whether periodic or not, is essentially an

inherent character. Even if it is automatic, it is realized only when

the external conditions allow, while under abnormal conditions it

can be realized only with difficulty. On the other hand, the para-

tonic reaction goes on subject to environmental change, sometimes

freely and sometimes enforced by it, so that the reaction proceeds

either actively or passively, and varies widely in its behaviour.

Or, in other words, the internal and external factors work together

intricately.

Trees of ancient types show mostly non-periodical gradual

growth. Perhaps the ancestors of all phyla of trees were non-

periodic in their growth and so far as they remain in wet and

warm regions, they have continued their habit unaltered, as for

instance, tree-ferns, cycads and palms with single thick stem, in

which substituting growth is not generally permitted. Most conifers

show the monopodial growth as much as possible, even if provided

with the means of substitution. In dicotyledonous trees, the mode

of branching is represented in various ways. The tropical trees of

primitive families show mostly monopodial growth, but even in

trees of advanced families, so far as they grow in a favourable en-

vironment, monopodial growth ‘still predominates, while in the

temperate zone sympodially ramifying trees are usual, especially

those with slender twigs. This is partly due to the dying out of the

apical parts through physiological dryness, as may be estimated by

the number of wilting buds, which are more abundant after a

severe winter. But in some trees terminal buds die in summer as an

inherent character, as in Tilia, Ulmus, Syringa and others. In either

case the more easily the substitution occurs, the more adaptable is

the tree to severe and variable conditions.

But, even in Singapore, there are a few trees in which the ter-

minal bud dies out (§ 9) as Xanthophyllum Curtisii and Crataeva

Roxburghii. The resting buds of the former (Fig. 1), both ter-

minal and axillary, are peculiar in construction. They are protected

by a pair of prophylls ca. 0:5—1-5 cm. long, forming an acute

quadrangular white pyramid, very conspicuous amidst the dark

green leaves. The inside of the scales consists chiefly of an air

chamber with a tiny bud at the base. What teleological meaning

)

Gardens Bulletin, S.

such a construction has is not clear, but absorption and conduc-

tion of heat of insolation will be relieved at least for a while. In

spite of such a structure, the terminal bud dies away early every

season as an inherent process. The ancestor of the species seems

therefore to have been -well adapted to dry spells, though in

Malaya, especially in Singapore at present, no such crisis may

happen. |

Fig. 1. Xanthophyllum Curtisii.

t, terminal bud withered; a, axillary buds.

On the other hand, among some families and genera of the in-

termittently growing habit, one can find a few species of the ever-

growing habit in Singapore. For instance, Gironniera subaequalis,

Elaeocarpus sphaericus, E. stipularis, Campnosperma auriculata,

C. minor, Adinandra dumosa, Archytaea Vahlii, and Symplocos

fasciculata. Perhaps there may be still more. As most trees of these

families and genera are distributed on the Asiatic Continent and

are intermittent or deciduous in their growth, so these evergrowing

species seem to have acquired the habit in Malaya.

In temperate regions, all trees, whether evergreen or deciduous,

are intermittent in their growth, forming the resting buds and

shedding leaves in a definite season. But when they are trans-

planted to warmer regions, their behaviour differs specifically. For

instance, the beech-tree transplanted to southern regions sheds

leaves though with delay, ultimately, subject to the autonomic

periodicity, while fig and mulberry-trees transplanted to the tro-

pics grow continually, becoming positive in activity on account of

the favourable conditions.

Conversely, trees of southern origin as Robinia, Sterculia and

others, do not close the shoot in the temperate regions, but form

new leaves continually and indefinitely until the arrival of the cocl

season, when the bud-forming process is at length induced. The

Vol. XVII. (1958).

palms in the temperate zone, for instance Trachycarpus excelsa of

Japan, show intermittent growth, flowering in late spring by send-

ing out 5—7 inflorescences at once from the axils. The morpholo-

gical sequence is just the same as in tropical palms with axillary

inflorescences, which appear however in the tropics after develop-

ment of successive leaves with equal intervals, one by one, non-

seasonally. The flowering of palms in the temperate regions is,

therefore, externally controlled intermittent growth, though it is

not known whether this periodic flowering has become inherent

or is still reversible in warmer regions.

There are besides a few widely distributed trees which are ever-

growing in Singapore, but deciduous under seasonal climate, as

Trema orientalis, distributed from India to Polynesia. We can not

say which of the two regions is the normal and which is the pheno-

copy (GOLDSCHMIDT ’35), or whether each represents an

ecotype (TURESSON ’30).

Physiological reactions are controlled naturally by prevailing

conditions, but so far as a species shows a local difference in its

activity, even if of slight character, it is not easily modified by

mere transplantation to other localities. In the temperate regions

the same species of tree from the north is generally earlier in leaf-

ing as well as in leaf-fall than that from the south; the former is

more frost-resistant than the latter. The variation is fixed as an

autonomic character. As a rare case, it was found, however, in

Carya of temperate America, that a southern species was more

frost-resistant than a northern species (WHITE ’35); it indicates

the existence of a genetic character not apparent in the original

southern district. TURESSON (7°30) has found in ca. 30 perennial

plants distributed widely in Europe and Asia, north and south as

well as inland and coasta!, that the species in each locality forms

an ecospecies in flowering, height of stem and so forth, adapted to

the local environment. Even in the photoperiodic response,

FURTAUER (’40) found in Epilobium hirsutum, which spreads

from Abo (61° N.), Finland to Kirstenbosch (34° S.), S. Africa,

that the photoperiodic response is long day in the high latitude,

short day in the low latitude and neutral in intermediate regions,

representing hereditary local races.

Similarly the periodicity of the tree-growth in each locality seems

mainly of inherent character (see also § 25). Recently the beha-

viour of polygenes in various quantitative characters has been taken

up in the experimental field and analyzed in detail (see for ins-

tance: MATHER et al ’43, WOODWORTH et al ’52). The local

difference of the periodic behaviour of trees seems in this respect

to be an instructive theme for further study.

Gardens Bulletin, S.

§ 23. The morphological development of the resting bud with

special reference to the formation of scales

For the formation of the resting bud three procedures are in-

volved: (1) the growth in length is retarded by the development

of a rhythmic tendency, (2) the leaf-primordia formed on the

growing point cease to grow further till to the next season, and

(3) some primordia metamorphose into the protecting members,

though in the tropics these processes are gradually simplified.

In primitive tropical families, as Anonaceae and Myristicaceae,

most of the trees are evergrowing without closing the shoot. The

formation of buds is in this respect a change of growth reaction

for protection of the growing point against unfavourable condi-

tions. But the restriction of the shoot growth does not necessarily

cause the formation of bud-scales. The latter is an independent

process, and trees destitute of that ability, when restricted in their

growth by drought, can not form a resting bud, e.g. Carica, Aca-

lypha, Jatropha, Hibiscus, Lantana and others (COSTER ’23).

In tropical trees with intermittent growth, the terminal portion of

the resting shoot is usually not covered by scales, but loosely by a

few young leaves, provided often with dense hairs or gum-resin.

Morphologically, these procedures are far simpler than the meta-

morphosis of foliar organs.

But some deciduous trees, the genera of which are of continental

origin, as Kurrimia and Pygeum, are provided with well-formed

scales, and those from central and tropical America, as Achras

and Couroupita, with metamorphosed succulent petioles. Even in

evergreen species of Quercus and Castanopsis, which originated

in calid regions, the buds are covered with dense corky scales,

like those of the temperate species. The same holds true with trees

transplanted from the calid regions such as Eryobotrya, Cinna-

momum, and Pittosporum. Scaly leaves are here not only the re-

duced phyllome, but with suberification added.

In trees of Magnoliaceae, Moraceae, and Rubiaceae, in which

the terminal bud is protected, as a family character, by a pair of

stipules or petiolar bases, the growing shoot can become a resting

bud without special morphological contrivance so that the trees of

these families are found in various growth-forms from the ever-

growing to the intermittent or even to the deciduous, and distri-

bute themselves widely in calid regions.

The development of foliage leaves of intermittently growing

shoots is sometimes gradual with transition from scales or protec-

tive leaves to normal ones in each season, but usually the change

is as abrupt as in temperate trees of the northern origin, in which

even a definite number of foliage leaves is laid down in the bud

Vol. XVII. (1958).

(LUBBOCK ’99). For instance in Fagus, Carpinus, etc., after

development of new shoots during one month or so, the twig ter-

minates with apical and axillary buds of the next spring without

growing further, notwithstanding that the season is just getting

warmer. Bud-formation here is, therefore, not induced by low tem-

perature, but only by an inherent process.

In Carya of N. America, at first about seven scales are found

during spring and early summer as the covering of the bud for

the next spring; 3-4 pairs of foliage leaves follow during the

summer and, then, 2—3 heteromorphic scales, which represent the

outermost scales of two years later and shed themselves soon

after the growth of the inner bud (FOSTER ’31). Similar develop- _

ment of foliar organs is seen also in Aesculus (FOSTER ’29). In

Pinus sylvestris, a tiny bud of two years hence is already formed

on the tip of the bud of the next year (LUBBOCK, l.c.), and in

Salix reticulata of the arctic region, the new shoot is formed two

years and a half in advance, in accordance with the short vegetative

season (RESVOLL ’25). Even in Shorea Hemsleyana of Diptero-

carpaceae, on the top of each bud is formed a young tiny bud of

two seasons later (Fig. 2).

The tree is endemic to S. Malaya and Sumatra and ecologically

has no need to form such a young bud with hurried tempo. At

Fig. 2. Shorea Hemsleyana x ca 4/5. Right figure seen from the opposite

side.

A bud just elongating, with leaves 1-5, and with young bud

of next season on the top covered by stipules a and b. (Compare

SYMINGTON ’43, p. 71) (ca. 4/5).

Gardens Bulletin, S.

any rate, such a process is hardly conceivable without the assump-

tion of a specific hormonal activity.

All these cases show that the formation of buds is not a direct

reaction to unfavourable climatic change. On the contrary, it oc-

curs as an inner process rhythmically after development of foliage

leaves, either in warmer or in colder regions, and even in the

case where low temperature or drought induce bud-formation, it

is only an indirect condition and not able to cause bud-formation

by itself, unless the plant is endowed with bud-forming ability

genetically.

If we summarise the development of the resting bud from the

ever-growing to the deciduous, it may be written as follows:—

(1) Evergrowing bud.

(2) Periodic fluctuation of leaf-size on the shoot.

(3) Change of proximal leaves of each period to protecting

leaves, small in size, often more hairy or producing.

anthocyan, etc. and falling early.

(4) Protecting leaves reduced to scales, often represented by

petioles or stipules, embracing or surrounding the

young shoot tightly and often corky.

(5) Young bud ceases to grow after forming a definite num-

ber of foliar organs.

(6) Young bud hastens further to form still younger bud of

two seasons later.

Each species of trees has its own rank of bud-development, as

mentioned above.

In the evergrowing shoot leaf-fall keeps pace with the opening

of new leaves, one after another. After acquiring the periodic

tendency, however, the basal leaves fall early, while the laminar

leaves remain longer as assimilating organs and the more the

periodic tendency is intensified, the more simultaneous become

leafing or leaf-fall.

In conclusion, the formation of the bud is realized by the intri-

cate co-operation of the hereditary characters and the environ-

ment. In the tropics the formation of buds is often autonomic,

while in the temperate zone the seasonal change may induce it.

AXILLARY BUDS: Now, concerning the axillary buds, they

are in some shoots just bare like the terminal one, as in Alangium

Ridleyi, Adenanthera pavonia, Delonix regia and many species

of Anonaceae and Myristicaceae. In Spathodea campanulata the

prophylls are also green, though different in shape. But in most

Vol. XVII. (1958).

cases they are protected by a pair of prophylls, rather tightly even

in evergrowing shoots. In the evergrowing Averrhoa and Caesal-

pinia sappan, although the terminal bud is bare, the axillary buds

are protected by prophylis, and in Sterculia ferruginea they are

covered with many scaly stipules besides the prophylls. Naturally

most of the axillary buds remain dormant without further growth,

except when they develop as cauline inflorescences after a long

spell.

It may be seen, therefore, that in the wet tropics the grade of

protection of axillary buds is manifold and not intimately related

with the mode of the shoot-growth.

§ 24. The retrogression of the bud-forming ability

Bud-forming ability does not always proceed unidirectionally

to the complication of scale-members. As already stated (§ 22),

some tropical trees, viewed from the systematic and comparative

standpoints, seem to be transformed from the intermittent to the

evergrowing, together with retardation of the bud-forming ability.

For instance the genus Adinandra, which is distributed mainly in

mountainous regions of S.E. Asia to Malaysia, produces well-

protected buds, whereas Adinandra dumosa spreads over the low-

land of W. Malaysia growing continually, forming no resting bud.

Gordonia of the same family spreads over the highlands of tro-

pical regions widely, while G. singaporeana grows in the lowlands

of Malaya, and, though the growth is slightly intermittent (§ 10),

never forms a resting bud, and suggests equally the transition to

the evergrowing. The genus Alangium, distributed from Africa to

the Pacific Islands and Formosa, seems distinctly intermittent in

its growth, but A. Ridleyi, an endemic species of Malaya, has no

scales in the resting bud, which is covered only by two young

leaves as if nearly evergrowing. Peltophorum pterocarpum is deci-

duous with leaf-fall twice every year, well adapted to the mon-

soon-regions ranging from Indo-China to Malaya and further to

Australia; in Singapore, however, it does not close the bud, which

is covered only by a few basal leaves densely clad with hairs.

There may be many more cases of such a retrogression of bud-

formation.

Thus bud-formation goes on, either in evergreen or deciduous

twigs, not always parallel with the cessation of shoot-growth.

Under a favourable climate, the bud can be much simplified. The

formation of bud-scales is therefore a different process that may or

may not accompany the retardation of shoot growth. But, as one

Gardens Bulletin, S-

looks at trees as a whole, including those of the temperate zone,

the two processes are seen to be correlated with each other, and

the more tightly the bud is closed, the more intermittent is its mode

of growth. Though originally different, these two processes are

combined together by natural selection, except under uniformly

favourable conditions. The internal: processes of bud-formation

are not clear, but hormonal activities derived from inherent cha-

racters undoubtedly control this behaviour. Periodicity of tree-

growth as a whole is, indeed, sufficiently comprehensible only when

one assumes hormonal activity as an inherent process.

§ 25. The intermediation of hormonal activity on the process of

periodicity

As stated above, the formation of scales, foliar leaves, etc. in a

definite order around the growing point of trees rhythmically is

not caused by the direct effect of climatic changes, but as a con-

sequence of internal specific processes that ought to be attributed

to the quantity and distribution of respective hormones. An in-

teresting example of abnormal scale formation caused evidently

by the disturbance of hormonal distribution, was observed by the

writer in a growing shoot of Lucuma Maingayi (Fig. 3).

10 f

| SOE

ae =

Biel

Fig. 3. A young shoot of Lucuma Maingayi grown horizontally.

Vol. XVII. (1958).

In a new shoot with scales and foliage leaves in 2/5 phyllotaxis,

the Ist to 5th phyllomes were scales and the 6th, 8th and 10th

were foliage leaves, while the 7th and 9th between the above

mentioned leaves should have been foliage leaves but were typical

scales. The new shoot was in horizontal position and the reverted

scales were on the underside. It is therefore conceivable that some

hormones, destined for green leaves or an antihormone to such,

received disturbance on both sides, upper and lower, at the begin-

ning of differentiation, and as a result the disorder ensued, just

like the behaviour of auxine in geotropic response (SCHRANK

°50). This fact shows that the final form of the leaf is not predis-

tined by the primordium. It may also be inferred that in Kurrimia

and others mentioned previously (§ 23), at the beginning of for-

mation of the resting bud, some hormones appear that prevent the

normal development of foliage leaves, and after exhaustion of the

same, the primordia tend to be normal. But in Carya (FOSTER,

l.c.) the leaf-forming hormone seems to be exhausted after the ap-

pearance of a definite number of leaves, and then an inversed

course of hormonai activity takes place forming the scales again.

The shoot of Bridelia shows the first indication of the rhythmic

growth of leaf-size, as already mentioned (§ 10). If the leaf-size

is determined absolutely by the quantity of nutrient materials, it

seems more probable that the basal or apical portion of the shoot

in each season would produce the larger leaves. In most tropical

trees with intermittent growth, the leaves at about one-third from

the base of the new shoot tend to be the largest, but in Cassia

and others the leaf-size increases in the terminal direction. In other

words, there is no rule concerning the distribution of leaf-size in

shoots of each season. Similarly in temperate regions it is manifold

according to species (DENNERT ’26). As the leaf-primordia

around the growing point are uniform in size, so the subsequent

difference in size should result from the mode of cell-division in-

duced by the activity of the respective hormones, while each cell

attracts its own nutritive materials from below.

In the shoot of each season, the internodes belonging to the

scales and incomplete leaves are generally very short, but in some

Species they grow much longer than those of the foliar leaves, for

instance in Actinodaphne malaccensis, Phoebe macrophylla,

Knema laurina, Diospyros lanceifolia, Ptycopyxis costata and the

basal portion of apposed twigs of the Terminalia-type. It is clear

therefore, that there is no causal connection between leaf-size and

length of internode, even if in most cases they are more or less

Gardens Bulletin, S.

proportional. Were they to be connected, it might be attributed to

nutrition, but, as it is not really so, it ought to be attributed at

least to two groups of hormones, which effect the leaf-size and the

length of the internode respectively. They can behave indepen-

dently according to species.

Now, concerning the leaf-fall, it is well known that this is affect-

ed by the formation of the abscission layer and the separation of

cells along it. The layer is formed generally at the base of the

petiole, for the formation of which some hormonal activity should

be concerned.

For instance, the tomato does not form the abscission layer on

the leaf, but the fruit is provided with a junction, from which it

separates at maturity. But there was found a mutant which failed

to form the junction, so that the ripe fruit was detached from the

receptacle (BUTLER ’36). This shows that a genetic factor con-

trols the hormonal activity. The abscission layer is not formed,

however, always in a definite position. The leaf-cutting of the

sweet potato sometimes forms the abscission layer abnormally in

the middle of petiole (YASUI °46). This fact indicates that, dur-

ing the formation of the layer, some substance, probably a pre-

cursor of a hormone of cell division, descends from the lamina

through the parenchyma downwards and meets on the way with an

ascending substance from the petiole and is activated by it. That

the abscission layer has a smooth surface, may be attributed to

the flatness of the diffusion front, even though it is often curved

according to species (PFEIFFER ’28).

The formation of an abscission layer is relatively rare in tree-

ferns, monocotyledonous trees, and herbs. Even in conifers some

do not form it, as Cunninghamia, Araucaria, Cryptomeria, while

Taxodium, Metasequoia, etc. form it at the short-base, just like

Canangium odoratum, Sonneratia acida, and Populus nigra. In

dicotyledonous trees, whether evergreen or deciduous, and whether

in the tropics or in the temperate region, this process goes on re-

gularly. Especially in the temperate region it occurs not only be-

fore the new leafing, but also before the severe winter, well under-

stood as the adaptation. In trees of one and the same species, a

delicate difference of various characters is often found, especially

in cultivated forms. For example, in Para-rubber trees grown in

the same plantation, some shed the leaves earlier, but some later,

and this is characteristic of each clone. As to the quality and yield

of the latex a similar difference is seen (SCHWEIZER 732). In

avenue trees of Aesculus, Fagus, and Ulmus there are seen also a

Vol. XVII. (1958).

difference of leafing and leaf-fall according to trees, which show

the same difference every year (LUBBOCK l.c., WELLENSIEK

°34). This is also seen in Ginkgo-trees, the so-called living fossil.

Whether these characters may be transmitted through seeds or not

wants experimental proof, but at all events the last step of the

physiological process is reaiized by the activity of cells induced by

hormones, by which the kind of materials, the amount and time

of secretion, their co-operation or antagonism, etc. are determined

inherently, even if their actions are affected at the same time by

external conditions.

In herbs, there are many cases in which the seasonal behaviour

has been genetically investigated. For instance in Hyoscyamus

niger, there are annual and perennial races, but the habit can be

altered by low temperature or by grafting (MELCHERS ’39).

Concerning the periodicity of the nyctinastic movement of Phaseo-

lus multiflorus, there was found an inherent variation of 15-35

hours at 22° C. according to individuals, and the statistical mode

for one cycle was 25 hours (BUNNING ’33). Yet under natural

conditions their proper cycle is controlled to 24 hours. Moreover,

there is a difference of the cyclic course of movement. In some

individuals the leaves sink quickly and lift slowly, while others

behave in the inverse way (BUNNING ’36).

The effect of synthetic hormones on the flower-formation and

the ripening or abscission of fruits is well known. The fruiting of

papaya-trees is noteworthy in this respect. Axillary female flowers

open successively and continually, but the further development of

them ceases after 10—20 fruits are formed, though all flowers are

evenly pollinated, and after a few months the next crop begins

forming a second group of fruits, and so on; it seems nothing but

a hormonal control.

In Singapore there are some trees which do not flower or bear

fruit, as for instance Ravenala madagascariensis, which flowers

and fruits in Burma. Borassus flabellifera, Nephelium litchi, and

Bouea macrophyila can ripen in Penang. It is said that the sterility

may be attributed to the wet climate. But it is not clear, whether

a wet climate alone is responsible, or whether intense insolation

or a definite thermoperiod (WENT ’48, ’49) or photoperiod is

needed. After all, it ought to be related with the production of the

hormone-complex.

In effect, the periodicity of tree-growth is controlled by various

hormonal activities as well as by the environment. Some hormones

are formed as an inherent character almost invariably, but others

Gardens Bulletin, S.

are more or less controlled by the external conditions. The hor-

mones that partake in bud-formation and other autonomic pro-

cesses are formed nearly independently of seasonal conditions,

especially under the uniformly favourable climate. The formation

of hormone in the case of leaf-fall or of abscission layer becomes

the more intense, the more susceptible the tree is or the more

severe the climatic changes. Conversely, the more favourable the

conditions or the more insensitive the tree, the more autonomic is

the production of hormones.

§ 26. Conclusion

If we compare trees in tropical and temperate regions, each

group shows many adaptive characters. In the tropics there are

many monopodial trees of primitive type, while in the temperate

regions most trees, except some conifers, are ramiferous with

slender twigs provided with numerous growing points in the form

of axillary buds, which are able to substitute injured parts. Re-

generation is easier in conifers than tree-ferns, and in dicotyledon-

ous trees than monocotyledonous ones, as may be seen from the

fact that monocaulous thick-stemmed trees scarcely extend further

north than the calid region.

The existence of deciduous trees in the tropics of uniform cli-

mate is, however, not adaptive in its nature. The habit is not

directly concerned with water or temperature relations. It is an

inherent character in its origin, just like the evergrowing and

intermittent growth-habit.

In the temperate region, from the systematic point of view, most

deciduous trees are of northern origin, but in the tropics the deci-

duous trees are distributed in various families fairly evenly, being

generally more abundant in larger families. There are various

genera which contain both evergrowing and deciduous species,

and even in some genera of tropical origin one can find a few

deciduous species. The periodic habit of trees seems, therefore, to

arise in any region in any family, so long as certain changes of

hormonal activity can occur.

Geographically, the deciduous species of Malaya are related

with those of both Continental Asia and Malaysia, though the

former has slightly more species in common than the latter. If one

divides all the related regions roughly into two, the wet W. Malay-

sia and other drier districts, so the number of deciduous species is

also nearly the same. Thus the deciduous habit has no preference

for the dry or wet habitat.

Vol. XVI. (1958).

Considered from the geological conditions in the past, Malaya

constituted in the early Pleistocene the western part of the Asiatic

continent, the so-called Sundaland, and there might have been

many chances for origination of deciduous trees in the midland

of that era, which were distributed then to the coastal region, now

representing the Malay peninsula, where they have retained their

habit unaltered. But there are, on the other hand, some genera,

most species of which are intermittently evergreen, that occur on

the mountain region inland with some species distributed in the

lowland with evergrowing habit. Growth-habit seems, therefore,

not generally changeable, yet possibly so in some genera.

Periodicity of tree-growth is the rhythmical advance of the

growth-process realized by various consecutive reactions. Partial

aspects of it, like leafing, leaf-period, leaf-fall, etc. show sometimes

even clonal variations with respect to the calendar date, which

occur fairly precisely according to stocks. Such trivial characters

have originated internally and are transmitted to the offspring. It

has been called the stockwise variation in this paper.

The real process is not known, but, so far as it appears as an ex-

ternal character, it stands in direct bearing with the environment,

and the more favourable the condition of the district, the more

freely the inherent ability exhibits itself; conversely, the more un-

favourable the local condition, the more restricted is the play-

ground of expression, whether autonomic or paratonic. And the

more restricted the existing conditions, the more adaptive seems

the character to one’s eye, and the more favourable the condition,

the more untidy becomes the habit.

After all, the periodic habit, or the change of it, is not neces-

sarily the adaptive, but, so far as the environment permits the

habit, the tree can exist competently mingled with trees of other

habit. That in Singapore such various growth-habits are displayed

may be attributed to these circumstances.

§ 27. Summary

(1) In the tropics of uniform climate, as at Singapore, there are

in tree-growth various transitions from the evergrowing to the

deciduous, so that it is not enough to classify the trees simply as

evergreen and deciduous as in temperate regions.

(2) Periodicity of tree-growth is characterized by the activity

of the growing point as well as by the mode of branching. The

former includes the continuous, the intermittent and the manifold

growth, and the latter the terminal and the lateral branching. By

Gardens Bulletin, S.

combination of these two categories, one can distinguish six types

of tree-growth morpho-physiologically. They may be classified

further in detail.

(3) Evergrowing trees are everleafing with the growing bud at

the apex of the twig, and the leaf-size and the length of internode

are uniform.

(4) Leafing of intermittently growing trees is seasonal, con-

temporary or non-seasonal, and at the end of each period a resting

bud forms on the shoot-apex. Leaf-size and the length of inter-

node are variable, usually with a sudden change in appearance of

the cork-layer between each period.

(5) Manifoldly growing trees show a branchwise difference in

the developmental phase caused (a) by dying out of the terminal

bud or the apical portion of the twig or by forming the inflores-

cence; (b) by trees growing to a fairly big size; and (c) by the

exotic nature of trees with intermittant growth, their branches hav-

ing a strong inherent tendency to rhythmical growth, which is

shifted, however, according to individual branches under the uni-

form climate.

(6) The deciduous trees are rather conspicuous in their appear-

ance, but they represent only a special case of intermittent growth,

the distinction being only whether leaf-shedding proceeds or lags

behind the leafing, whence arise all sorts of transitions.

(7) The leaf-period of deciduous trees in Singapore varies from

three months to more than two years according to species, usually

with a short bare spell, but this has nothing to do with the hiber-

nation of temperate trees. The frequency of seasonal leaf-fall is

1—3 times per annum according to species, that of the contem-

porary ones 3—4 times, and among the non-seasonal ones some

are shorter and others are longer than one year.

(8) Four categories of flowering can be distinguished, as in the

case of leafing. Everflowering is limited to a certain number of

evergrowing trees, while the rest are seasonal, contemporary or

non-seasonal, just as in the intermittent and manifoldly growing

trees. Ever- and non-seasonal flowerings are autonomic in nature,

while seasonal and contemporary flowerings are chiefly determined

by trifling changes of the weather.

(9) In Malaya, evergrowing trees together with the manifold

ones compose at most about one-fifth of the total indigenous tree-

species, and the deciduous about one-twentieth, the rest being the

intermittent evergreen trees.

Vol. XVII. (1958).

(10) Deciduous species occur there fairly evenly in various

dicotyledonous families. Most of these families include not only

the usual evergreen trees with periodical growth, but also ever-

growing as well as deciduous ones, indicating that the deciduous

habit may occur in various families rather optionally.

(11) Some trees, though deciduous in other regions as India

and Malaysia, may be evergreen in Singapore, while others are

not seasonal in their leaf-shedding.

(12) In the Malay Peninsula there are, besides endemic deci-

duous trees, those common with Continental Asia or the Oceanic

Islands—chiefly W. Malaysia—or those distributed in both re-

gions, so the deciduous trees as a whole show no preference as to

the habitat, whether dry or wet.

(13) There is also no intimate relation between the deciduous

habit and the height of crown, tall evergreen trees being more

abundant than deciduous ones. Low deciduous trees are also met

with not infrequently.

(14) Deciduous trees are, moreover, not all the same in water

relations. There are, besides many mesophytic trees, some hygro-

philous and others hygrophobic, the former growing in wet

swampy forest and yet deciduous, while the latter do not occur in

S. Malaya.

(15) The Malay Peninsula, as a land on the Sunda Shelf, was

during the Tertiary period a part of the Asiatic Continent together

with W. Malaysia, so that under the then predominating climate

there might have originated a number of deciduous species in the

midland of that area, which retained the same habit unaltered,

while others seem to have changed their habit from the periodic

to the evergrowing after the formation of the present peninsula

since the Pleistocene.

(16) Generally, intermittent growth, whether evergreen or de-

ciduous, is accompanied by the formation of a resting bud. But

these two processes are not always carried on in the same degree.

Under a favourable climate the latter is more easily simplified than

the former.

(17) The formation of leaf-scales may be attributed to hor-

monal activities. Similarly the formation of the abscission layer,

the length of periodic activity and many other characteristics of

trees are all controlled internally and show even clonal differences.

This is especially evident in the uniform climate like that of

Singapore, where various growth-habits are displayed without res-

triction, whereas in the temperate or monsoon regions the external

control predominates.

Appendix

Gardens Bulletin, S.

The periodicity of tree-growth in four types, together with loca-

lities. Exotic genera and species in parenthesis.

Abbreviations

Periodicity types: e: evergrowing, m: manifold, i: intermittent,

d: deciduous.

Ad: Andaman Islands

Af: Africa

Am: America

As: Asia

Ass: Assam

Au: Australia

B: Borneo

Br: Brazil

Bu: Burma

CAm: Central America

Cd: Calid Regions

Ce: Celebes

Ch: China

Cosm: Cosmopolitan

Cy: Ceylon

EAs: Eastern Asia

EMs: East Malaysia

Eu: Europe

F: Formosa

Hm: Himalaya

Hond: Honduras

I: India

IC: Indo-China

IO: Indian Ocean

Localities

J: Java

Jam: Jamaica

Jp: Japan

M: Malaya

Mc: Mascarene Islands

Med: Mediterranean

Mg: Madagascar

Mo: Molucca

Ms: Malaysia

Mx: Mexico

NAu: North Australia

NB: North Borneo

NC: New Caledonia

NEM: North-East

Malaya

NG: New Guinea

Nic: Nicobar Islands

NM: North Malaya

NT: North Temperate

P: Pacific Islands

Per: Peru

Ph: Philippine Islands

Po: Polynesia

Q: Queensland

S: Sumatra

SAf: South Africa

SCh: South China

SEA: South-East Asia

Si: Siam

SM: South Malaya

Sol: Solomon Islands

T: Tenasserim

Ti: Timor

Tr: Tropics

TrAf: Tropical Africa

TrAm: Tropical America

TrAs: Tropical Asia

TrM: Tropical moun-

tains

Ven: Venezuela

WI: West Indies

WMA: West Malayan

Archipelago

WMs: West Malaysia

WP: Western Pacific

Islands

Vol. XVII.

Genera

Alangium

(Anacardium)

Bouea

Campnosperma

bed

Dracontomelum

(Lannea)

Mangifera

Melanorrhoea

Parishia

Pentaspadon

(Pistacea)

(Pleiogynium)

Semecarpus

Sorindaea

(Spondias)

3°

Anaxagorea

{Anona)

Canangium

Drepananthus

Polyalthia

Xylopia

Alstonia

Cerbera

Dyera

Ervatamia

Kopsia

Ochrosia

(Plumeria)

(Thevetia)

(1958).

Distribu- | No. of | Do of :

| tion _| species jal Species

Alangiaceae

| Am-P | 22. | 7 | Ridleyi

Anacardiaceae

Am 8 | (occidentale)

I-Mo 5 2 | microphylla

As,Af,Am| 10 3 | auriculata

| macrophylla

| minor

| I, Ms, P 5 1 | mangiferum

Af, I i a | (grandis)

SEA-Ms 40 20 | oe

oetida

| (indica)

\WMs 15 | 10 | malayana

| Bu, Ad, 10 5 | insignis

| WMs, Ph

Maingayi

paucijuga

pubescens

T, M,B few | 3 officinale

Med, As, 5 | (formosana)

Q 1 (Solandrii)

TrAs, Au, 40 10 vernicifera

| Po

| Af, Mg 20 madagascariensis

Y FrAs: 6 | (cytkerea)

| TrAm

| (lutea)

| (monbin)

(pinnata)

Anonaceae

| Am, As 8 ) 2 javanica

Am,Af,As| 60 | > Reticulata

Bu-Au 2 | odoratum

S-NG, Ph 8 | 4 pruniferus

| ramiflorus

As: AE. 60 | 40 Beccarii

Au :

| stenopetala

Tr 100 | 16 ferruginea

Apocynaceae

At... AS 30 6 | angustifolia

Au, Po

| angustiloba

| latifolia

macrophylla

| scholaris

spatulata

I-WP 5 2 manghas

odollam

M,S,B 2-3 1 | costulata

Tr 170 10 corymbosa

dichotoma

SEA 12 | fruticosa

| singapurensis

Mc,As,Po . mary rescence

acuminata

| (obtusa)

Am 10 (peruviana)

vse TO NESSES

ooo a

Bp BR B

Anan AO

AaAQ &

a & A228 «

Gardens Bulletin, S.

istribu- | No. of ; : ae ’

Genera | es u Sacces Meee Species Localities |emi d

Aquifoliaceae

Ilex Cd 200 2 cymosa WMs i

’” | | | macrophylla | WMs | d

Asclepiadaceae

(Calotropis) | Af, As | 3 | 1 | (gigantea) | I | e

Araliaceae

Arthrophyllum Ad-NG 10 8 | diversifolium WMs e

(Brassaia) Au 1 (actinophylla) Au i

Schefflera ETr 80 30 | heterophylla M,S i

Bignoniaceae

(Catalpa) Am, EAs 10 (longissima) WI m

(Jaccaranda) SAm 30 (filicifolia) SAm i

(Kigelia) Af, Mg 3 (pinnata) Af d

Oroxylum La iC 1 indicum I-Ph, NM| m

(Spathodea) Af 2 (campanulata) Af e

(Stenolobium) Mx-SAm 4 (stans) SAm e

(Stereospermum) ETr 20 fimbriatum Bu-NM d

(Tabebuia) Cd (rosea) Mx i

” (triphylla) WI m

Bixaceae

(Bixa) Am 1 (orellana) Am e

(Cochlospermum) | Tr | 12 | (religiosum) I d

Bombacaceae

(Adansonia) ETr 10 (digitata) Af d

(Ceiba) TrAm 10 (pentandra) 4009 d

Durio As 12 zibethinus Ms i

Neesia WMs 6 altissima M, B,S, J

(Ochroma) SAm, WI 1 lagopus Br e

(Pachira) m (aquatica) Am

»» (cyathophora) Am

Salmalia Af, As, 11 malabarica Hm, Cy,

Au Bu, Ss, d

wa Valetoni Ms d

Borraginaceae

Cordia Neel 300 dichotoma ee Ms,

u, Po

»” (sebastina) TrAM,

“ subcordata I-WP m

%» (umbraculaeformis) TrAm

Burseraceae

Canarium At, I, Ms, 109 (album) IC

” nitidum M i

9 rufum M _d

Santiria Ms 30 laxa M

% (rostrata) B

9 rubiginosa Bu, i d

" tomentosa i d

Vol. XVII. (1958).

ae |

Genera | ae Dad Meiaya Species Localities |emi d

Cactaceae

(Cereus) | Am | 300 (hexagonus) Am e

(Peireskia) ed tie aoe 1 (grandifolia) | Am i

| |

Capparidaceae

Crataeva af 20 | 2 | religiosa ETr m

- | | (Roxburghii) | I, Cy, IC, m

| 1

Caricaceae

Casuarinaceae

Casuarina | Au, Ms, I 23 1 equisetifolia I-P e

” | | | (sumatrana) | Bu, WMA| e

} | |

Celastraceae

Elaeodendron Cd 30 1 (quadrangulatum) Br e

Kurrimia I, Ms 3-4 2 paniculata I, WMs | m_

» robusta I, WMs | i

Combretaceae

Lumnitzera IO-WP a | Zz coccinea | Af-P e

a | | | racemosa | Mc-P e

Compositae

Vernonia | Cd | 500 | 8 | javanica | I, Ms | i

Datiscaceae

Tetrameles | Ms, Ph | 1 | 1 | nudifiora | 2 J, | d

| i

Dilleniaceae

Dillenia | As, Au 10 6 | aurea L a NM,

3 | | grandifolia M d

» indica I, Si, M i

Yr) | ovata a, IC, i

2 reticulata M d

Wormia Pe Au, 20 4 albiflos M, NB e

”» excelsa M,S, B i

” pulchella M, S, B i

a | suffruticosa WMs e

Dipterocarpaceae

Anisoptera Bu-NG 17 7 | megistocarpa SM,S d

Dipterocarpus | I-Ce 69 34 grandiflorus Ad, Si, Ph d

% Kunstleri S, NM d

% | Lowii M, S, B d

Gardens Bulletin, S.

Distribu- | No. of | Do. of : sf :

Genera den species | Malaya Species Localities |em i d

Dipterocarpaceae—contd.

Dryobalanops B, M,S 6 2 aromatica |B, M.S je

Hopea | I-NG 56 9 mengarawan S, SM i

Pentacme Bu-Ph 6 1 siamensis Bu; Si, IC, d

Shorea | I-Mo,Ph | 95 58 Curtisii Si, M i

” Hemsleyana NM

> leprosula Sime S, d

a (robusta) I, Bu i

os sumatrana A, M i

Ebenaceae

Diospyros } I-Bu-J, Au] 300 ; 70 cordifolia | I, Si, Ms, d

99 (ebenum) B.Cy i

9 lanceifolia WMs i

% (mollis) Ss i

ys oblonga M i

Elaeocarpaceae

Elaeocarpus | I, Ms, Au 100 30 ferrugineus M,B i

9» littoralis Bus) SS i

WMs

99 Mastersii WMs i

» obtusus Bu, M,S, i

% petiolatus IC, WMs i

99 polystachyus M,B i

ay | sphaericus I, WMs le

” stipularis IC, WMs |e

| |

Erythroxylaceae

Txonanthes | SEA-MG 11 2 | icosandra M,S i

| | | reticulata | M, S, B d

Euphorbiaceae

Acalypha Tr, SAf 400 3 siamensis Si, NEM i

Agrostistachys I, Ms, Ph 11 3 sessilifolia M i

{Aleurites) As, P 5 (moluccana) Mol e

Antidesma At, As: 140 21 (bunius) J-Au i

» , (diandrum) I-Bu i

Aporosa As-NG 60 20 | Benthamiana M-S i

> bracteosa M, S, B i

Es nigricans M i

Baccaurea As, Au 60 22 Kunstleri M i

2 latifolia M i

99 macrophylla M e ,

> malayana M i

99 parviflora M,S i

ap pyriformis M i

(Bischofia) I, Ms, Po 2 1 (javanica) I, Ms, Po d

Bridelia | Eftr 50 8 retusa I, Si, M d

” tomentosa I, Ms e

‘Claoxylon Mg,As,P| 57 4 | indicum I, Ms ‘

Cleistanthus Af, I, Ms 110 30 heterophyllus M i

Croton Cd | 500 10 (eleuteria) Am ee

” (laevifolium) IO i

Elateriospermum WMs 1 1 tapos M,S, J ie

Emblica As, Am 11 1 officinalis SEA i

Euphorbia Cd 750 10 neriifolia I, Ms e

Excoecaria ETr 26 agalloca Af-Au d

Glochidion As-Au 130 24 laevigatum M e

0 microbotrys M e

- sericeum M, B, J e

“i singaporense SM e

- superbum SM-Ph ¢

Vol.. XVII. (1958).

Genera

( Hevea)

(Hura) .

(Jatropha)

Macaranga

Mallotus

Microdesmis

Pimelaeodendron

Ptycopyxis

Sapium

”

Castanopsis

9°

Quercus

Flacourtia

Homalium

Hydnocarpus

”

Pangium

Scaevola

Calophyllum

Cratoxylon

Kayea

Mesua

(Rheedia)

Distribu-

tion

CdAs

Eu, As,

NG, Am

I, Ms

I-Ms-Ph

WMs

Au, ETr |

ETr

No. of

Species

Do. of

Malaya

Species

Euphorbiaceae—contd.

12 (brasilliensis)

2 (crepitans)

150 (multifida)

200 24 javanica

triloba

100 25 sp.

paniculatus

2 1 (casearifolia)

4 1 Griffithii

1 1 costata

100 3 discolor

indicum

sebiferum

Fagaceae

40 15 Andersonii

inermis

400 40 (bambusaefolia)

conocarpa

craterophora

hysterix

lucida

Flacourtiaceae

10 3 indica

jangomas

rukam

50 8 grandiflorum

30 7 (anthelmintica)

Kurzii

2 1 edule

Goodeniaceae

2 | 1 | frutescens

Guttiferae

50 25 inophyllum

12 5-6 arborescens

formosum

ligustrinum

Maingayi

100 40 atroviridis

(dulcis)

(ferruginea)

Forbesii

Hombroniana

(Livingstonii)

mangostana

xanthochymus

12 11 ferruginea

2-3 2 ferrea

17 madruno

Localities

emi

I-Mc (not

Sm)

ETr

I, Ms

Si, M

Si, IC

Ass, Bu, |e

I-Po e

I-WP

IC, Ms,Ph

I, WM, Ph

fmt pete bmi pede

feats eho eek peahe prate pete beaks fee

AA,

Gardens Bulletin, S.

Genera | apenas oe ae bya Species Localities |em i d

Hamamelidaceae

Altingia I, Ms, Ch 2 1 excelsa I,M,J d

Bucklandia | i Bu, 1 1 populnea M e

Rhodoleia M, Ch 3 2 ovalifolia M i

m Teysmannia | M,S i

Juglandaceae

Engelhardtia I-Ch, M 10 | 4 nudifiora M d

ae | | | Wallichiana | M | i

Lauraceae

Actinodapvhne | EAs, Ms 50 15 bancana M m

oe | malaccensis | SM m

Beilschmiedia Tr 20 15 malaccensis SM d

Cinnamomum As, Au, 150 17 (camphora) Ch, F, Jp m

xs iners I, WMs m

io (zeylanicum) I, Cy m

‘Cryptocarya Cd 50 17 Griffithii | M, B i

Lindera As, NAm 60 14 lucida SM, WMs| e

Litsea | As, Au 200 56 | gracilis M i

“ { megacarpa M i

2 singaporensis | M i

Phoebe As, Am 40 6 grandis | WMs i

e macrophylla i

99 opaca M,J,S,B i

Lecythidaceae

Barringtonia | Af, As, Au 30 15 asiatica ETr i

at (acutangula) 5, Si, i

5 racemosa IT, Ms, Po i

(Bertholetia) | Br 1 (excelsa) r d

(Couroupita) Am 4 (guianensis) Guiana d

(Lecythis) Am 45 (ollaria) TrAm e

{(Napoleona) WAf 5 (imperialis) TrAf i

Leguminosae

(Acacia) Au, Af 450 (auriculaeformis) NAu m

+ (catechu) I, Bu, Si m

5 (confusa) Pho i

i | (glomerosa) SAm m

99 (interjecta) WI e

Adenanthera | ETr 3 2 bicolor Cy, M d

a pavonina I, Ch, Ms, d

| Mo

Albizzia ETr etc 100 4 falcata EMs e

45 (odoratissima) Hm, Bu, d

| Cy, Si

{Amherstia) | Bu 1 (nobilis) Bu m

{Andira) | Am, Af 25 (inermis) re WI, i

(Baphia) Af, Mg 30 (nitida) TrAf i

(Brownea) SAm, WI 15 (ariza) New m

Granada

ue | (capitella) Ven m

- | (coccinea) Jam m

es | (grandiceps) Ven m

- (macrophvlla) m

(Butea) ee Ch: 3 (frondosa) I, Bu, Ch, d

Vol. XVII. (1958).

Genera | ieee a ote ee Species Localities |emi d

Leguminosae—contd.

Caesalpinia Tr 100 9 (coriaria) | Am e

. (ferrea) Br d

ia sappan I, Ms i

Cassia Tr 400 17 (excelsa) Br i

¥ (fistula) I, Cy d

Pe (javanica) J, S, Ph d

a (leptophylla) Br d

a (multijuga) TrAm e

ae nodosa Bu, Ch, d

WMs

+ siamea I, Ms e

os (splendens) Am e

Cynometra Tr 20 3 inaequifolia ot WMs, i

wy ramiflora I, Cy, Ms i

Dalbergia - Au, 120 15 | (Oliveri) Bu, Si d

fe)

(Delonix) Af, Mc 3 (regia) Mg _d

Dialium SEA ils} 8 Wallichii M i

(Enterolobium) Am 6 (saman) SAm e

Erythrina Tr 30 5 fusca Me I, d

Ms, P

- indica I-P d

lithosperma Bu, M i

es , subumbrans or WMs, i

(Gliricidia) Am 4 (sepium) CAm _d

(Haematoxylon) Am, WI et (campechianum) Am i

(Hymenaea) Am 8 (courbaril) WI _d

Inocarpus am, Ms, 1 1 edulis Ms-P i

Intsia ETr 12 2 Bakeri Si. WMs, d

a bijuga Mc-Po d

Koompassia IC, Ms 4 2 excelsa M,S,B d

s malaccensis M,S d

(Leucaena) Am, P 9 (glauca) SAm e

(Lonchocarpus) png Af, 120 (cyanensis) WAf i

Millettia Af,I, Ms,| 150 IS atropurpurea Bus. Si i

Au WMs

sa (Manni) TrAf i

(Mimosa) ae As, 300 - | (Ssepiaria) SAm e

(Myroxylon) | Am 8 (toluiferum) Am d

Parkia Tr 10 3 | javanica Ms d

= speciosa I-J i

Peltophorum Tr 7 3 dasyrachis Si, S, M d

a pterocarpum IC-M-Au d

(Pentaclethra) Am, Af 3 filamentosa Am i

A (macrophylla) Af i

(Phyllocarpus) Br 1 (septentrionalis) Br e

(Piptadenia) TrAm 45 (colubrina) Br i

Pithecellobium TrAs, 60 10 ellipticum W Ms i

TrAm

és jiringa WMs

i. microcarpum S,M,B i

Pongamia I-Fiji few 1 pinnata SEA-P d

(Prosopis) Cd 25 (juliflora) Am e

Pterocarpus Tr 15 1 indicus Ms d

Saraca I-Ms 20 ¢ bijuga Si, M i

= declinata ot M, S, i

3 (indica) I

a thaipingensis M i

(Sesbania) Tr 40 Ps (grandiflora) EAs €

Sindora I-Ms, Af 9 5 coriacea M d

99 Wallichii M d

Sophora Cd 25 1 tomentosa I-P i

Tamarindus Af, As 1 1 indica TrAs d

(Tetrapleura) WAf Sa (Thonningii) TrAf i

(Trachylobium) Af, As 3 | (verrucosum) EAf, Mc i

Gardens Bulletin, S.

Distribu- | No. : e ae 5

Genera | ; fi see ” Seta Moke Species | Localities |emi d

Loganiaceae

Fagraea RrAs: 60 20 auriculata M,. J, IC, i

WP * Ph

a fragrans T, Si, Ms je

35 gigantea Ss e

o | Ridleyi SM i

Lythraceae

Duabanga j|Hm-Ph 2 1 sonneratioides I-NM e

Lagerstroemia CdAs, 20 3 floribunda Buy Se d

Mg, NG, M

es flos-reginae J-Au, NM d

2 ovalifolia WMs d

Sonneratia As-P 4-5 3 acida I-P e

% alba I-P e

Magnoliaceae

(Magnolia) | As,NAm; 21 (fusca) Ch e

Michelia I-Ms 12 5 (alba) i e

% | champaca I, M e

Malvaceae

Hibiscus Tr ) -1S0 12 macrophyllus IC, “Si je

W Ms

As (rosa-sinensis) Ch? e

29 (schizopetalus) EAf e

an tiliaceus Tr e

‘Thespesia Af, As 6 1 populnea ETr e

Au |

Melastomaceae

Memecylon ETr 150 35 heteropleurum M,S,B i

Pternandra Ms 18 5 capitellata T-M i

ns coerulescencs var.

Jackiana W Ms-Mol i

a echinata M, B e

Meliaceae

Aglaia I, Ms, Au ! 90 40 trichostemon M,B e

Carapa Tr i 6 3 (guianensis) SAm m

‘Cedrela As, Am 16 3 (glaziovii) Am ‘

‘Chisocheton I, Ms 30 11 spicatus M i

(Chloroxylon) I, Cy 1 (swietenia) I, Cy ’

Melia ETr 12 4 (azedarach) Hm i

(Swietenia) Am 5 (macrophylla) Hond i

op (mahogani) WI i

Moraceae

Antiaris | Af, I, Ms 3-4 1 toxicaria iE Ch,

Artocarpus | As-P 50 20 bracteatus NM

is | (communis) P e

es dadah WMs d

=f Gomezianus WMs i

e rigida , WMs i

(Castilloa) Am, Cuba 3 (panamensis) Mx, Per je

(Cecropia) SAm 30-40 (peltata) | SAm e

Vol. XVII. (1958).

Genes | Distribu-

Streblus

Myrica

Horsfieldia

”

Aegiceras

(Eucalyptus)

”>

Eugenia

Melaleuca

(Psidium)

Rhodamnia

Pisonia

»”»

Ouratea

tion

En M,

SCh, Ph

Au, EMs

Au, Ms,

No. of

species

Do. of

Malaya

Species

Moraceae—contd.

600

1,000

130

120

100

alba

(calophyllus)

caulocarpa

elastica

glomerata

indica

(irregularis)

(Kurzii)

lepicarpa

religiosa

superba

variegata

asper

Myricaceae

| 1 | Farquhariana

Myristicaceae

globularia

(grandis)

(sylvestris)

Myrsinaceae

| 1 | corniculatum

Myrtaceae

150

(corymbosa)

(longifolia)

(Naudiniana)

(robustus)

(aromatica)

densiflora

javanica

(liliiformis)

longiflora

malaccensis

myrtifolia

polyantha

Ridleyi

spicata

leucodendron

(guajava)

trinervia

Nyctaginaceae

(alba)

excelsa

Ochnaceae

| 4

corymbosa

crocea

Localities | emid

I-NG

Ms (not

ooa

i te pte ete

Pte poate pete phe ty fake fem pete beak ep y

Gardens Bulletin, S.

Genera | Demin hee Lette’ Species | Localities |emi d

Olacaceae

Gonocaryum Bu, M 4 2 longe-racemosum (iM e

Ochnostachys B, M 1-2 1-2 amentacea B, M

Scorodocarpus M, B 1 1 borneensis B

Oxalidaseae

Averrhoa Ms 2 2 bilimbi , Ms e

ss carambola | J, EMs e

Connaropsis M,B 8 8 Griffithii | M

38 | macrophylla | M

Pittosporaceae

Pittosporum | ECd | 60 | 3 | (tobira) | Ch |

Polyglaceae

Xanthophyllum | Ms, I, Au | 30 | 21 | Curtisii | M | i

Polygonaceae

(Triplaris) SAm 10 (americana) SAm e

2» | | | (surinamensis) SAm e

Proteaceae

Buckinghamia Q 1 (celsissima) | Au

(Grevillea) Au 170 (robusta) Au

(Stenocarpus) EAu, NC 15 (sinuatus) | Au

Rhamnaceae

Zizyphus Cd 100 10; calophylla | M,B,S

% (Jujuba) I-Ch

% (mauritiana) | IO m

Rhizophoraceae

Anisophyllea f trAs At 10 7 disticha WMs e

” Griffithii M

Bruguiera As-EAf 5 4 gymnorhiza ETr e

Ceriops ETr 1 1 Candolleana ETr e

<Gymnotroches Si, Ms 2 1 axillaris M c

Kandelia I, Ms 1 1 Rheedii Ms e

Rhizophora Cd 3 2. conjugata ECd e

% mucronata ECd e

Rosaceae

Parinarium cy 50 12 | corymbosum Ms

Pygeum WAf 30 15 parviflorum WMs

99 polystachyum NM

Pyrus NTemp- 70 1 granulosum I-Ms

TrMt

Rubiaceae

Adina ECd 10 2-3 j{ rubescens M

Anthocephalus I, Ms 3 1 cadamba I-NM

Canthium ETr 100 10 dicoccum SEA-Ms |e

ss glabrum Si, M e

Vol. XVII. (1958).

Genera : amma od ated Species Localities |e mi d

Rubiaceae—contd.

Gardenia ETr-Temp 70 10 carinata M je

Py (coronaria) Bu e

Ixora ETr 120 20 congesta T, M e

x multibraceteata SSi, M e

Jackia M, S, B 1 1 ornata e

Morinda tr 50 7 citrifolia I, Ms, P je

99 > elliptica e

Randia Tr 100 25 anisophylla S, M,B e

a3 macrophylla S,M i

Scyphiphora As, Au 1 1 hydrophyllacea As, Au e

‘Timonius As, Po 30 10 Wallichianus M e

Urophyllum As, Af 40 13 hirsutum M i

i streptopodium M e

Rutaceae

¢(Aegle) I 1 (marmelos) e

Atalantia Hm, Au 10 6 spinosa I, Bu, Si, i

Feronia i, Si 1 1 limonia I, Si d

Glycosmis SEA, Ms 20 8 chlorosperma W Ms i

Salicaceae

(Salix) Cosm 160 | | (tetrasperma) I-Ch d

Sapindaceae

Erioglossum As-Af 4 1 rubiginosum M i

(Filicium) As-Af 3 (decipiens) I, Cy e

Guioa I, Ms, Au 60 4 pubescens WMs, Ph i

Lepisanthes As 20 a cuneata M i

{Litchi) Ch 1 (chinensis) Ch i

Nephelium As, Au 70 16 lappaceum Ms i

% ophioides M i

Paranephelium I-Ms 8 3 macrophyllum Si, NM i

Pometia I, Ms, Au 7 4 pinnata I, Ms, Po i

Trigonachras Ph, M 9 1 acuta SM d

Xerospermum I-Ms 20 . 6 muricatum M i

Sapotaceae

Achras) Mx, CAm 1 (zapota) e

Lucuma Am, As 50 8 Maingayi M, S, B d

Madhuca I-Ms 40 21 Kingiana M i

Mimusops) ETr 20 2 (elengi) I, Bu, Cy i

Palaquium I, Ms, Po 100 14 Clarkianum M

2” gutta S, M, B €

” Maingayi M d

% obovatum ir. MM, 'S d

Payena Ms 12 12 dasyphylla M,J,S e

: glabra ;

{Pouteria) Am 30 (suavis) Uruguay i

Simarubaceae

Eurycoma IC, Ph, B 2 2 longifolia IC-M, S, i

Irvingia Af-Ms 4 1 malayana — S; d

Solanaceae

Solanum Cd | 1,225 16 | (Wrighti) Bolivia e

id)

Genera |

(Cola)

Commersonia

(Dombeya)

Firmiana

Heritiera

Kleinhovia

Pterocymbium

(Rheevesia)

Scaphium

3°

Sterculia

Tarrietia

Symplocos

o”

Adinandra

Archytaea

Gordonia

Pyrenaria

Ternstroemia

(Thea)

(Phaleria)

Gardens Bulletin, S.

Distribu- | No. of | Do. of

tion species | Malaya Species

Sterculiaceae

Af 50 (acuminata)

Aue: 8 1 Bartramia

Ms, Ch

Af 100 spectabilis

I-Ms 2 1 fulgens

‘As, At. 7 2 elata

littoralis

(macrophylla)

Mc-Po 1 1 hospita

Bu, WMA 2 Z javanicum

tubulatum

Hm-Ch 3 (thyrsoidea)

Bu; IC; 4 3 affine

; linearicarpum

Tr 200 20 (carthaginensis)

ferruginea

foetida

macrophylla

parviflora

rubiginosa

I, Ms, Au 10 6 | javanica

Symplocaceae

Cd 300 30 fasciculata

| | rubiginosa

Theaceae

SEA 20 10 | acuminata

angulata

dumosa

Hulletti

integerrima

macrantha

maculosa

parvifolia

villosa

TrAm, 4 2 Vahli

CdAs, 20 10 penangensis

singaporeana

I-M af 2 acuminata

As, Am 300 6 bancana

SEA, Ch, 40 (cochin-chinensis)

Jp |

Thymeliaceae

(Blumei)

Localities

Len |

M, S; Beale

WMs

(highland)

bs J 4-0

Si, Ms e

Pte tte pete pete

emid

rom

a ARAAAR A A AAA

Vol. XVII. (1958).

Genera

‘Gironniera

”

Trema

””

”

Avicennia

.°

Callicarpa

(Citharexylum)

Gmelina

Premna

‘(Tectona)

Vitex

”

Alscdeia

Distribu- | No. of | Do. of

tion species | Malaya

Ulmaceae

SEA 12 4

Tr 30

Verbenaceae

Tr 4 4

Tr 50 7

Am 20

As, Au 12 1

ECd 50 13

I, Bu, Si, 3

Cd 140 16

Violaceae

Tr | 50 | 19

Vitaceae

As,Af, Au 70 12

Species

nervosa

subaequalis

angustifolia

orientalis

variegata

alba

intermedia

lanata

officinalis

(Rheevesii)

(quadrangulare)

(arborea)

tomentosa

(grandis)

| (Hamiltoniana)

(capitata)

coriacea

pubescens

Maingayi

aequata

angulata

indica

rubra

Localities

WMs (not

Java)

SEA

WMs

Hm-Po

SEA-Ms

emi d

oaoodon

ooo

Literature cited

ARLDT, T.: Die Entstehung der Kontinente und ihrer Lebewelt. 2

Aufl. Berlin. 1936-8.

Brooks, C. E. P.: The evolution of climate. London. 1922.

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(36):

BuRKILL, I. H.: Dictionary of the economic products of the Malay

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BUTLER, L.: Journ. of Her. 27 (36).

CHAMPION, H. G.: Indian forest records. (New Series). Silvi-

culture. Vol. I. New Delhi. 1936.

CorRNER, E. J. H.: Garden’s Bull. 9 (35); Wayside trees of

Malaya. 2 vols. Singapore. 1940.

CosTER, C.: Ann. Jard. Bot. Buitenz. 33 (23), 35 (26), 37 (27),

38 (28).

DENNERT, E.: Die individuelle fluktuierende Variabilitat. Jena.

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FABER, F. C. v.: Pflanzengeographie auf physiologischer Grund-

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GOEBEL, K.: Organographie der Pflanzen. 3 Aufl. I. 1928. Jena:;

Bliitenbildung und Sprossgestaltung. 1931. Jena.

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Groom, P.: Trans. Linn:‘Soc, 2 Ser. 3:(92),

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Symposia Exp. Biology VII (53).

IHERING, H. v.: Bot. Jahrb. 58 (23).

KLess, G.: Jahrb. f. wiss. Bot. 56 (15).

Koorpers, S. H.: Forstl.-naturw. Zeitschr. (1898).

KoriBa, K.: Journ. Jap. Bot. 7 (31).

KUIJPER, J.: Rec. trav. bot. néerl. 30 (33).

LAM, H. J.: Ann. Jard. Bot. Buitenz. 37 (27).

Lussock, J.: Buds and stipules. London. 1899.

MATHER, K.: Biol. Reviews 18 (43).

MELCHERS, G.: Biol. Zbl]. 57 (39), Ber. d. d. bot. Ges. 57 (39).

MERRILL, E. D.: Proc. Pan-Pacific Sci. Congr. Austr. 1923.

Mo.iscuH, H.: Die Lebensdauer der Pflanze. 1929. Jena.

Gardens Bulletin, S.

No_LpE, B. I. v.: Bot. Jahrb. 71 (40).

OcusE, J. J.: Fruits and Fruitculture in the Dutch East Indies.

1931. Batavia.

PFEIFFER, H.: Die pflanzliche Trennungsgewebe. 1928. Berlin.

PoTTER, M. C.: Journ. Linn. Soc., Bot. 28 (91).

Raciporski, M.: Ann. Jard. Bot. Buitenz. 17 (01).

REID, E. M. AND CHANDLER, M. E. J.: The London Clay flora.

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RENSCH, B.: Eine biologische Reise nach den kleinen Sunda-In-

seln. 1930. Berlin.

RESVOLL, T. R.: Flora, 118/9 (25).

RicHArpDs, P. W.: The tropical rain forest. 1952. Cambridge.

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e293, TIE“(24)..

SCHIMPER, A. F. W.: Pflanzengeographie auf physiologischer

Grundlage. 1898. Jena.

SCHRANK, A. R.: Ann. Rev. of Plant Physiol. 1 (50).

SCHWEIZER, J.: Mitteil. naturf. Ges. Bern. 1932.

SCRIVENOR, J. B.: The geography of Malayan ore deposits. 1928.

SEWARD, A. C.: Plant life through the ages. 1933. Cambridge.

Simon, S. V.: Jahrb. f. wiss. Bot. 54 (14).

STAMP, L. D.: The vegetation of Burma. 1925. Calcutta.

STEWART, C. D.: Malayan Meteorological Service, Summary of

Observations, 1930. 1931. Kuala Lumpur.

SYMINGTON, C. F.: Foresters’ Manual of Dipterocarpaceae.

Malayan Forester Records 16 (43). Syonan.

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Waite, O. E., Zeitschr. f. ind. Abst. u. Vererb. 69 (35).

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Plant Communities on Singapore Island

By H. B. GILLILAND

Professor of Botany,

University of Malaya, Singapore

DESCRIPTIONS OF plant communities on Singapore Island are infre-

quent in the literature. This is perhaps natural because attention

has been directed in the first place to the identification of the

members of its rich flora. General papers have been published by

Corner (1941) and others, and one or two papers concerning the

specific make-up and dynamic behaviour of the vegetation also

exist. The first of these was a paper published by I. H. Burkill in

1919, and constituted the record of his observations upon a piece

of then 30-year-old secondary forest (belukar) which he had cut

down for the purpose of developing the Botanic Gardens. Burkill’s

data are very full, and can be represented as shown in Table 1, in

such a way as to indicate the reproductive status of the principal

species. In selecting from Burkill’s data for presentation in this

‘table, I have chosen the species present in reasonable amount, or

sometimes have been guided by the importance of the species con-

cerned in other communities. Those which are lumped together at

the end of the Table under “Others” amount to 25 per cent of the

total number of individuals recorded and comprise some 95 further

species.

It is clear that the Common Ivy-Palm (Arthrophyllum diversi-

folium), whose population is present in all size classes, has been

growing and reproducing successfully for a considerable period on

this site. It is in this case the nearest equivalent to what is meant

by a dominant in other climates. The Leechwood (Anisophyllea

disticha) is here its principal shrubby partner. The Broad-leaved

Oil Fruit (Elaeocarpus petiolatus) of which two large specimens are

recorded, has evidently not been reproducing itself successfully

until recently.

By analogy this might well be called an Arthrophyllum/Aniso-

phyllea Mixed Forest indicating that, while in the conventional

sense no dominant species was present, the Common Ivy Palm

and the Leechwood appear to have been the most successful and

characteristic species.

Table 1 is prepared in such a way as to render a comparison

with the results obtained by the present author elsewhere (Gilli-

land, 1952) readily possible.

Vol. XVII. (1958).

TABLE 1

THIRTY- YEAR-OLD BELUKAR: BOTANIC GARDENS: SINGAPORE

DATA OBTAINED FROM BURKILL, 1919*

SAMPLE AREA 13,680 SQUARE FEET

Species 0-1+1-3 3-5 5-7 Over7 _ Total 48

Arthrophylum diversifolium 125 10 16 13 164 5.5

Rhodamnia trinervia 229 — 2 11 242 8.2

Cinnamonum iners 315 2 4 — 32] 10.8

Timonius wallichianus 107 2 1 — 110 3.7

Macaranga triloba 106 _ 1 — 107 3.6

Adinandra dumosa i el — 1 4 96 52

Elaeocarpus petiolatus uy geet — — 2 36 i2.

Gordonia singaporeana .. 25 1 1 1 28 9

Gynotroches axillaris i ee 1 — —_ 86 2.9

Pygeum polystachyum ee 1 — 1 — 2 1

Ixonanthes reticulata — — — | 1 —

Anisophyllea disticha 345 — — — 345 11.6

Elaeocarpus mastersii 199 — — — 199 6.7

Gironniera nervosa 190 — — —_ 190 6.4

Palaquium bancanum 144 — — — 144 4.8

Eugenia grandis 139 — — — 139 4.7

Others — — — — 754 25.5

¥otals) .°2,135 16 27 32 2,964 99.8

* Gard. Bull. $.S8.2:5:145, 1919.

Holttum, in a paper published in 1954, recorded those plants

which constitute what he called “Adinandra belukar” and gave

his observations upon the species which behaved as pioneers, ini-

tiating succession on this island. Table 2 summarises his findings.

There are trees and shrubs, perennial herbs and climbers. My own

observations fully confirm Professor Holttum’s conclusions with

regard to these species.

TABLE 2

PIONEER ‘ADINANDRA BELUKAR’; SINGAPORE ISLAND

DATA FROM HOLTTUM, 19547

TREES AND Melastoma malabathricum Macaranga javanica Archytea vahlii

SHRUBS

Adinandra dumosa

Wormia suffruticosa

‘PERENNIAL Lycopodium cernuum

HERBS

Gleichenia linearis

Eriachne pallescens

CLIMBERS Nepenthes rafflesiana

} Mal. Jour. Trop. Geogr. 3;27, 1954.

Ficus alba

Rhodamnia trinervia

Gahnia tristis

Scleria laevis

Bromheadia palustris

Nepenthes gracilis

Fagraea fragrans

Myrica farquhariana

Spathoglottis plicata

Arundina graminifolia

Imperata cylindrica

Nepenthes ampullaria

Gardens Bulletin, S.

At the corner of the “catchment area” which forms part of the

Nature Reserves on Singapore fronting Lornie Road and adjoin-

ing the Royal Singapore Golf Course, occurs a large specimen of

the tree /xonanthes reticulata prominent amongst the species of a

manifest secondary woodland. An examination of the ground at

this site disclosed evidence of military occupation in the shape of

slit trenches, latrine pits, foundations of temporary buildings, etc.

from which it is logical to deduce considerable occupation of the

area, with its concommitant destruction of the vegetation, during

the 1941-6 period.

By a fortunate chance Mr. Corner, in discussing the Ten Men

Tree in his well-known ‘Wayside Trees of Malaya’ mentions this

specific locality for the tree. In correspondence he notes that he

can confirm the existence of this species at this site in 1934. So we

are not without some history of this particular area. The Ten Men

Tree has stood for at least twenty years—a fact confirmed by the

size of the bole of a large specimen, which has a girth of more than

ten feet—and has survived severe disturbance which occurred

-around it some ten to fifteen years ago.

This area was transected by chain using the technique described

in the paper referred to (Gilliland, 1952). However, since so many

plants and species were present only 1,000 feet of transect were

enumerated. Furthermore, in this case the records of each chain

length (600 sq. ft.) of the transect were recorded separately. In

this way it became possible to estimate from the recurrence of a

‘species in each or fewer of the 10 separate sub-samples, its degree

-of “faithfulness” in the community. Faithfulness is here used upon

a different scale from that normally employed: by continental

‘sociologists, but the result is of sufficient interest to make the

record worth while. Table 3 presents a summary of the results

obtained.

TABLE 3

SUMMARY

SAMPLE AREA 6,000 SQUARE FEET

Diameter Classes

Species o-1 1-3 3-5 5-7 Over7 Total %

“Surviving Trees 40 6 2 3 8 59 2.1

Seral Relics 109 2 — —_— _ 111 4.0

ee ae 1564 éi 7 oo geet 59'2

Community: Trees s

ce aamaaial : Shrubs 268 7 1 —_ — 271 9.8

Lianes 295 2 — — _ ne! gah:

Stage: Trees 161 5 —_ ce — :

a... Shrubs 34 1 — — — 35 1.3

Lianes 99 —_ — — — 99 3.6

Dnidentified 74 4 — — — 78 2.9

Totals 2648 86 11 6 9 2760 99.9

Vol. XVI. (1958).

The number of plants recorded in the first place makes it clear

that this forest is much more dense than that investigated by

Burkill—the area is approximately half. A total of ninety-five

species was recorded together with some which could not be

identified; approximately the same total.

Secondly by making use of the concept of ‘faithfulness’ it is

possible to separate out the ‘present community’ from the succeed-

ing community or next stage. Further by the use of Holttum’s

records and Corner’s notes on the Ten Men Tree and for other

reasons, three other groups can also be distinguished.

TABLE 4a

HUMAN CASUALS

DIAMETER CLASSES

Species 0-1 1-3 3-5 5-7 Over7 Total Fid. %

Acacia auriculiformis . = 3 1 1 -—— 5 2

‘Thevetia neriifolia .. oe 2 — — — — 2 1

Nephelium lappaceum Se 1 — — a“ — 1 1

Garcinia mangostana Bt 1 — — — — 1 1

Totals ssi2 4 3 1 1 — 9 — 0.3

TABLE 4b

SURVIVING TREES

Species 0-1 1-3 3-5 5-7 Over7 Total Fid, yA

Ixonanthes reticulata = 6 — — 2 7 15 8

Alstonia sp. - fe Lh 2 1 1 — — 4 1

Gordonia singaporeana sity 11 5 —_ 1 — | 5

Horsfieldia sp. i. ._ = — — — 1 1 1

Gynotroches axillaris saatie DT — 1 — — 22, 5

Totals .. 40 6 2 3 Sie) peepee s 24

TABLE 4c

SERAL RELICS: TREES & SHRUBS

Species 0-1 1-3 3-5 5-7 Over7 Fid. Total A

Macaranga javanica ~ Sel 26 — — —_ — 26 8

Adinandra dumosa .. an 29 2 — — — 25 4

Fagraea fragrans site ed 19 — — — — 19 5

Tetracera assa . a 19 — — — — 19 7

Archytea vahlii af - 8 — — — — 8! 2

Vitex pubscens - 3 — — — — 3 2

Ficus chrysocarpa .. sta 8 —_ — —_ o 8 2

Macaranga triloba .. 1 = — — — 1 1

Adinandra sp. 2 _ — —_ — 73 2

Totals .. 109 2 = — — 111 — 4.0

Together with the following ground flora of herbs:

Stenochlaena palustris, Cleidemia hirta, Lygodium microphyllum, Melas-

toma malabathricum, Gleichenia linearis, Nepenthes ampullaria, Gahnia

tristis, Nepenthes rafflesiana, Scleria sp., Phyllanthus niruri, Bromheadia

palustris.

Gardens Bulletin, S.

Table 4 presents the detailed records of the Surviving Trees, the

Seral Relics and the Human Casuals.

The surviving trees are so called from their presence in mature

form coupled with a poor representation amongst the seedling

stages. The Ten Men Tree is typical and it will be recalled that

Corner records this as being present in 1934. It is suggested that

during the period of the disturbance and subsequently, it has been

unable to reproduce and that only now with the return of forest

conditions is it beginning to be able to reproduce effectively. It

produces an abundant crop of fruit and seed every year but as yet

the seed have not been able to germinate and gain a foothold.

The seral relics are those recorded as making up part of the

‘Adinandra belukar’ or pioneer community and these, with the

possible exception of the ‘tiup-tiup’ (Adinandra dumosa) itself,

which seems to grow fast enough to avoid shading, are now on their

way out as the result of the re-establishment of forest conditions.

Macaranga javanica—the Blue Mahang—although present and

with a high degree of faithfulness, in my opinion will not grow to be

larger thax. a seedling in the conditions of shading which now ob-

tain. With this list of trees and shrubs should be read the list of

species in the undergrowth. These likewise are members of the

‘Adinandra belukar’ and will not long survive the shading. Holttum

specifically notes that the Orchid Bromheadia palustris does occur

in shade but never manages to flower in such conditions.

The Human Casuals are interesting. The first two are com-

monly cultivated plants; the first as a street tree and the second as

an ornamental shrub while the next two are the ‘rambutan’ and

the ‘mangosteen’, both everyday tropical fruits.

TABLE 5a

PRESENT COMMUNITY: TREES

DIAMETER CLASSES

Species 0-1 1-3 3-5 5-7 Over7 Total Fid. wa

Rhodamnia trinervia seen 265 31 3 1 1 301 10 10.9

Elaeocarpus petiolatus ay 38 8 3) — — 49 8 1.4

Elaeocarpus mastersii 5 aes — 1 ~s — 99 10 3.6

Oncosperma sp. ba us 73 _- — 1 — 74 7 BAT |

Eugenia longiflora .. in 4 3 — — — 494 10 17.9

Garcinia griffithii .. ay 76 2 _— — — 78 2.8

Cinnamomum iners mS 1 — — — 33 10 122,

Pithecellobium clypearia ot 27 2 — — — 29 1 st |

Eugenia grandis ba aus PA 4 — — — 25 8 0.9

Guoia pubescens... os 16 4 — — — 20 8 0.8

Timonius wallichianus ae 18 2 —_ ae oa 20 9 0.8

Angelesia splendens. . Ne 17, 3 — — —_ 20 8 0.8

Artocarpus kunstleri ah 17 1 — — — 18 8 0.7

Calophyllum floribundum oa) 1328 — — — — 328 10 11.9

Garcinia cowa a i Be — — — — 35 7 1.3

Xylopia ferruginea .. 12 —- - os a 12 7 0.4

Totals .. 1564 61 7 2 1 1635 — Sp die

fe)

Vol. XVII. (1958).

TABLE 5b

PRESENT COMMUNITY: SHRUBS

Species 0-1 1-3 3-5 5-7 Over7 Total Fid. we

‘Champereia manillana ~. PLOZ Z — — — 104 10 eT

Anisophyllea disticha ae 80 — — — — 80 8 2.9

Txora congesta i: Be: 58 — 1 — —- 59 10 Don

Lasianthus sp. ~ a 28 — —_- —_ — 28 7 1.1

Totals .. 268 2 1 1 — 271 —_— 9.8

PRESENT COMMUNITY: LIANES

Rourea splendens .. Aa 96 2 — —_— — 98 8 35

Fibraurea chlorcleuca ne 86 — — — — 86 0 Sol

Ericybe princei * z 64 — — — — 64 10 22

Calamus sp. a Py 30 — — — — 19 ¢) 0.7

Strychnos malaccensis ie 19 — — — — 19 8 0.7

Totals <2 295 Z, — — — 297 — 10.7

UNDERGROWTH CONSISTING OF:—-Davallia sp.; Taenitis blechnoides;

Selaginella sp.; Curculigo villosa; Dianella ensifolia; Hornstedtia martini-

censis: Centotheca lappacea; Panicum nodosum.

This list represents what I would consider to be, with the aid of

the use I have made of faithfulness, the present community. AJ

the plants noted have a fidelity to the sub-samples of the transect

of seven or over. (See column 7 of table 5).

The undergrowth consists of plants typical of forest conditions

on Singapore Island; the lianas will not excite comment and the

leechwood appears again amongst the shrubs, though the False

Olive (Champereia manillana) exceeds it here in number of indi-

viduals and development.

The trees are most interesting in that amongst them there is only

one species which shows a distribution over all the size classes;

seedling; sapling; young tree; mature tree and the large tree,

namely the Silver Back (Rhodamnia trinervia). It is clear that it

was present at the time of the disturbance and has not only sur-

vived that disturbance but has been able, in doing so, to extend

its territory and to become the characteristic plant of the present

community. This is in keeping with its record by Holttum as

amongst the pioneer shrubs. The Broad-Leaved Oil Fruit in this

instance has been much more successful than in the case of

Burkill’s sample. Considering the comparative success of the False

Olive (Champereia manillana) we might legitimately describe the

present community as a Rhodamnia/Champereia forest. We would

note that both the Common Kelat (Eugenia longiflora) and the

Bintangor (Calophyllum floribundum) are commencing to play a

significant role but that as yet they are present only as seedlings

and saplings.

Gardens Bulletin, S.

TABLE 6a

NEXT STAGE: TREES

DIAMETER CLASSES

—

Species 0-1 Over7 Total Fid. 7,

Millettia atropurpurea ds 38

Cratoxylon formosanum ca” yee

Calophyllum sp.I .. Se i!

Calophyllum sp. II ze 13

Garcinia sp. ie 11

Onchostachys amentacea

Elaeocarpus sp.

Lindera lucida

Arthophyllum diversifolium

Daemonerops sp.

Tlex cymosa

Litsea grandis

Plectocomia sp.

Randia anisophyllea

Palaquium sp. :

Eugenia garcinifolia

Baccaurea parviflora

Aquilaria malaccensis

bo ee CD Ld GY BBN) 00 ~1 00

(Bil es May pe ae eee pe

Reb Wet helciah ie ae

Gr IS tet IE del ae Sets

tle Slt Teta erat

mr WW Lo & f B ~1~1 00 00 10

— i WWNWW A RWW bLADAU WW

166 6.0

Totals: 2.) 161

TABLE 6

NEXT STAGE: SHRUBS

Species 0-1 1-3 3-5 5-7 Over7 Total Fid. %

Wikstroemia ridleyi 22 —_ — — —_ oR: 5

Psychotria sp. 3 — — — — 3 i}

Gnetum macrostachyum 2 — _ — —_ 2 Z

Canthium sp. 2 — — — — p) 2

Clerodendron diversifolium 2 _— — — — 2 2

Tetracera sylvestris . 1 — — — —_— 1 1

Chasalia chartacea .. 1 — — —_ 1 1

Dracaena sp. _— 1 — _ —_ 1 1

Strophanthus sp. 1 — — — — 1 1

Totals 34 1 — _— — 35 — 1.3

NEXT STAGE: LIANES

Willughbeia coriacea opel — — — — 91 3

Piper sp. py ae 3 — — — — 3 2

Artabotrys sp. +3 eee —_— — — —_ 2 2

Thottia grandiflora .. oe 2 — — — — 2 2

Paedera verticillata .. 1 —_ — oe — 1 1

Totals. 23 ,(99 — — — — 99 — 3.6

Unknowns e.. Me ae f 4 — — — 78 a 2.9

This list represents those species of trees, shrubs and lianes

which, by virtue of their low fidelity to the sub-samples of the

transect, constitute evidence of invasion of the present community

by the elements which will constitute the next phase in the suc-

cession back to high forest. The majority of plants recorded are

seedlings and all have in common a considerable shade-tolerance

as seedlings. There may be species whose natural distribution fol-

lows a pattern of such wide dispersion that they would not be

caught in a sub-sample of 600 square feet. For that reason they

would normally be expected to show a low fidelity figure in a

Vol. XVII. (1958).

transect such as this. We may be confident, however, that such

species will be revealed by continuing work along these lines for

they will always be distinguished by a low fidelity.

In these analyses certain species have been brought into promin-

ence by means of transect enumeration. These species appear to

characterise particular stages of the succession which will, in course

of time, restore the high forest. By analogy with the behaviour of

those in other less fortunate climates, where dominance develops,

they may well be used to name the stages involved. Three stages of

secondary forest succession on Singapore island then emerge:

1. The Adinandra Belukar (Holttum, 1954)

Clearly the length of time that this will occupy the ground de-

pends largely upon the degree of prior destruction of the habitat.

Over on Kent Ridge where it occupies the site of abandoned

plantations it will be many years before the reaction of this pioneer

community is able to ameliorate the habitat sufficiently for further

development to occur, while in the instance at the Lornie Road/

Golf Course site it is just as clearly already on its way out after

some ten to fifteen years.

2. The Rhodamnia/Champereia stage

The Silver Back/False Olive community now holds the ground

but clearly has within it the seeds (or better seedlings) of the next

stage.

8. The Arthrophyllum/Anisophyllea stage (Burkill, 1919)

This Common Ivy Palm/Leechwood community is the one

which Burkill so carefully annotated thirty-five years ago and which

then represented the progress of belukar which had held the

ground for thirty years.

Discussion

Comer (1940), Holttum (1930: 1940) and Koriba (1947)

have given a great deal of attention to the phenological behaviour

of our plants with special reference to time of flowering and the

formation of new leaves—to the beginnings more especially of

deciduousness—while Corner (1951) particularly in developing

his Durian Theory, has given prominence to the concept of the

moist tropical forest environment as the primal land habitat. It

has been suggested that the deciduous habit and the seasonal

flowering condition may have become consolidated by migration

to north and south, requiring those functions to become seasonal

in pattern. Let us suppose that Singapore were to suffer a severe

Gardens Bulletin, S.

drought of say three months duration—that in fact that the cli-

mate were suddenly to become monsoonal. The plants which

would be most severely affected would be those with the shallowest

and least developed root-systems—in fact it would be the seedlings

which would suffer first. The Silver Back is recorded among the

pioneer plants of the Adinandra Belukar which means that it is

relatively hardy. The figures show clearly that it is the best adapted

to the earlier conditions of the present community. Hence the

effect of such a calamitous drought would be to bring the Silver

Back into vastly greater relative prominence. It would begin in

fact to display the characters of a dominant from a less favoured

climate than our own. The concept of a primal vegetation in the

humid tropics fits these facts of community structure, and this

analysis of the Rhodamnia-Champereia community lends support

to that hypothesis by indicating how a development of dominance

in response to the demands of a harsher climate could come

about.

The use of the term “faithfulness” — “fidelity” in this context

unhappy; a future paper will propose a more neutral term.

References

BurKILL, I. H. Gardens’ Bulletin, Straits Settlements 2: 5: 145

1919.

Corner, E. J. H. “Wayside Trees of Malaya” Govt. Printer,.

Singapore, 1940. The M.A.H.A. Magazine XI, 2: 59

(1941). Ann. Bot. 13. 367 (1949).

GILLILAND, H. B. Jour. Ecol. 40: 1: 91 1952.

Ho.tttum, R. E. Gardens’ Bulletin, Straits Settlements 5: 7: 173

1930. Ibid 11: 2: 119 1940. Malayan Journal of

Tropical Geography 3: 27 1954.

Korisa, K. Physiology and Ecology (Seiro-Seitai) 1: 2: 93 &

160 1947.

Juncaceae, a new family record for Malaya

By J. H. KERN

Foundation Flora Malesiana, Leyden

AMONG SOME unidentified sedges from the Malay Peninsula kindly

sent to me for identification, I found two sheets of Juncus prismato-

carpus R.BR., collected by H. M. BURKILL in the Cameron High-

lands. It seems worthwhile to mention these collections in a short

note, as up to the present Juncaceae were unknown from the Malay

Peninsula.

Juncus prismatocarpus extends from Ceylon through S.E. Asia

to Japan and Kamtchatka, and southward to Australia, Tasmania,

and New Zealand. In Malaysia it is known from N. Sumatra, West

and Central Java, the Philippines, and New Guinea (see BACKER

in Flora Mal. I, 4: 213. 1951).

MALAY PENINSULA. Pahang: Cameron Highlands; summit of Batu

Brinchang, open rough waste, 6,660 ft. Sept. 1, 1956, H. M. BURKILL

783 (L); Break Pressure Tank Hill, open waste land, 4,900 ft. Sept.

5, 1956,,H: M. BURKILL 823-(E).

The habitat of J. prismatocarpus is in Malaysia in the mountains,

between 3,000 and 9,000 feet above sea-level, along pools, on

marshy plains, along swampy banks of brooks and rivulets, but

occasionally on drier heath lands, always in open places.

The anthropogenous habitat in the Cameron Highlands might cast

some doubt about its being native in the Malayan uplands, where

it now occurs as a ruderal. The fact that it has never been found

before in Malaya and this new habitat would at first glance point

to a new, rather recent introduction.

If the species was introduced it is most likely that it is due to man

who, after the rather recent opening of the Cameron Highlands, the

building of bungalows, etc., has frequented this pleasant resort.

The tiny, oblong ribbed seeds measuring only 4+—? mm. in length

may easily have been brought along with luggage or, more possibly,

have adhered to shoes or clothes of people who have formerly

either paid a visit to mountain resorts in Ceylon, India, Burma,

North Sumatra or Java. One could imagine for example visits of

botanists, tea-planters, or tourists. How long the seed will retain

its viability under these circumstances is unknown.

Another explanation is more likely, however, namely that it is

native and was hitherto overlooked because of its scarcity due to

Gardens Bulletin, S.

the paucity of its ecological niche. Open somewhat marshy locali-

ties must have been extremely scarce before the opening of Came-

ron Highlands. Once the opening of the highlands started, man

greatly extended the area of its potential occurrence.

A similar swarming out of native mountain plants of restricted.

occurrence over newly opened land occupied by man has been des-

cribed by Dr. VAN STEENIS)* from various mountains in Java

in which several native species invaded plantations as weeds in

large quantity or acted as roadside ruderals. A similar behaviour

has been observed in the European Alps and THELLUNG has.

called such plants on new, anthropogenous habitat, apophytes) +.

Juncus prismatocarpus may in Malaya well belong to this class.

and may originally have occurred very locally in the Camerons

from which focus it has spread to the newly opened land, thereby

greatly increasing in number. It is quite possible that other eco-

logically heliophytic plants may ‘escape’ from their small, precari-

ous niches in the densely forested hills and show their existence in

this unexpected way.

* Bull. Jard. Bot. Btzg III, 13 (1935) 303.

+ Bot. Jahrb. 53 (1915) Beibl. 116, p. 38 footnote.

Ararocarpus —-A Monstrosity

By JAMES SINCLAIR

SCHEFFER in Ann. Jard. Bot. Buit. 2 (1885) 10 described a mono-

typic Annonaceous genus, Ararocarpus from material collected by

himself in Java. He named it Ararocarpus velutinus and compared

it with Annona which differs from it in having only one seed to the

carpel but which is similar in respect of its fused carpels. The car--

pels of Ararocarpus are, however, free at their apices for a short

distance.

For a long time I tried unsuccessfully to locate the type specimen

as I thought that there might be a possibility of the plant occurring

in Malaya. Boerlage’s picture of it in Icones Bogoriensis 1 (1899)

Tab. 40 was then all I had seen of it. I followed Hutchinson, Kew

Bull. (1923) 255 and placed it in the Annonineae along with

Annona, Raimondia and Rollinia (non Malaysian genera) on

account of the fused carpels [Sinclair in Gard. Bull. Sing. 14 (1955)

167 and 177]. I failed, however, to see that it ought not to be

placed in that group on account of the several seeds per carpel.

I eventually found the type specimen of Ararocarpus velutinus

in Leiden in April 1956 and then the picture was clear. I saw what

Ararocarpus was. Boerlage, l.c. page 84, was not so far wrong

when he placed it next to Meiogyne in his scheme of classification

on account of the numerous seeds per carpel and the similar petals.

He did. however, associate it with Annona in the same scheme and

not unnaturally because of the fused carpels. He also described

(with a querry) a variety tenuifolia of Ararocarpus from a sterile

specimen in Bogor, l.c. page 123.

Ararocarpus is actually a freak of Meiogyne virgata (Bl.) Mia.,

a species which has a wide distribution, occurring in Malaya, Bor-

neo, Java and Sumatra. It is rather variable in the size of the leaves

and the breadth of the petals. The carpels are sessile or nearly so

and often touch each other at the base. The average number is 3

but sometimes 1, 2, 4 and 5 occur. In Scheffer’s specimen there are

8—12 carpels and here, apparently, on account of their large num-

ber, they had no room to expand laterally so a fusion took place,

resulting in the abnormal fruit. Naturally such monstrosities are

rare so one need not be surprised when the plant was never

collected again.

4cm

Fig. 1. Some Annonaceous carpels.

A, Fissistigma lanuginosum. B, Goniothalamus ridleyi. C, Meiogyne

virgata with a single carpel. D, the same with five carpels. E, F,

Ararocarpus velutinus. G, the same in longitudinal section. A from

Kiah S.F.N. 32199. B from Sinclair, Seletar Forest. C from Hender-

son S.F.N. 19609. D from Daud & Tachun S.F.N. 35692. E-G

copied from Boerlage’s figure in Icon. Bog. 1 (1899) Tab. 40.

Vol. XVII. (1958).

There are other species of Annonaceae with numerous, sessile or

almost sessile carpels clustered together into a ball. Such examples

are seen in Fissistigma lanuginosum, Goniothalamus ridleyi and

Uvaria sphenocarpa. One might very well expect to find these with

fused carpels since there is little room for lateral or basal expansion

of the carpels. However, let no more new names be invented for

such abnormalities or monstrosities.

Florae Malesianae Precursores—XX

The Genus Gymnacranthera (Myristicaceae) in

Malaysia

By JAMES SINCLAIR

SUMMARY

THERE ARE Six species and four varieties in the genus Gymnacran-

thera. Some taxonomists may prefer to regard the varieties as

subspecies since they occupy only certain geographical areas in

some reasonable distributional pattern. One of the six species,

namely G. farquhariana (Hk. f. et Th.) Warb., synonym G. canarica

(King) Warb., is confined to southern India, while the rest are

Malaysian. I have already dealt with the Malay Peninsula species

in Gardens’ Bulletin, Singapore 16 (1958) 434 in which publica-

tion will be found descriptions and details of distribution in Malaya.

The distribution of these Malayan species outside Malaya is

dealt with now in the present paper but descriptions and other

details are not repeated. Descriptions, however, are given for the

remaining species which do not occur in the Malay Peninsula.

Species on loan from a few herbaria were received after my paper

“A Revision of the Malayan Myristicaceae” went to the press so

these are briefly enumerated in the present account under Malay

Peninsula, extra specimens. These herbaria are BO, LE and NSW.

Vernacular names given here are those used outside Malaya. The

only common vernacular names for Gymnacranthera in Malaya

are pendarah, chendarah, penarah, darahan, chendarahan, pen-

darahan and penarahan and these are equally applicable to the

other genera of the Myristicaceae. It is not say therefore, to

repeat them under each species.

In this paper the following receive new status:—G. crassinervis

is reduced to a variety of G. forbesii and G. zippeliana to a variety

of G. paniculata. The following are reduced to synonyms of G.

paniculata var. paniculata:—-G. acuminata and G. macrobotrys,

while G. suluensis is made a synonym of G. paniculata var.

zippeliana.

Vol. XVII. (1958).

KEY

a. Leaves larger than in the other species, 20-40 cm. long

and 7:5-16:5 cm. broad (leaves from young or vigorous

shoots of G. forbesii var. crassinervis may equal their dimen-

sions but average leaves are smaller), covered beneath with

rusty or ashy tomentum which persists for some time; nerves

19-23 pairs. Inflorescence axis rusty-tomentose with hairs

1 mm. long. Flowers rusty-tomentose, larger than those of

the other species, 5-65 mm. long. Fruit rusty-tomentose,

the apex sometimes sharply acute or slightly uncinate

b. Leaves rounded or slightly rounded at the base. Male

flowers 5 mm. long and 3 mm. broad

(1) G. bancana var. bancana

b. Leaves acute at the base. Male flowers slightly larger and

broader, 5-65 mm. long and 4 mm. broad

(1) G, banecana var. borneensis

a. Leaves smaller, not over 28 cm. and up to 10 cm. broad; not

covered with rusty tomentum; nerves not more than 21 pairs.

Inflorescence axis not tomentose but often puberulous to

adpressed-pubescent, flowers scarcely tomentose, mostly

pubescent, less than 6 mm. long. Fruit scarcely tomentose,

mostly glabrous when mature, the apex obtuse or slightly

acute

c. Twigs generally stout, 3 mm. thick at the apex and 5 mm.

thick 12 cm. down, grey or pale straw-coloured. Leaves

10-28 cm. long and 3-3-10 cm. broad; nerves 13-21

_ pairs

d. Leaves often acuminate at the apex, the undersurface

with a rusty tinge, less often glaucous; nerves usually

fine beneath, not forming a distinct double loop.

Inflorescence axis rusty-tomentulose, female 1:5 cm.

long, the flowers very densely crowded and touching

each other. Fruit oblong, rusty-tomentulose, becom-

ing glabrous (2) G. contracta

d. Leaves generally less acuminate, the undersurface

more glaucous; nerves more prominent beneath,

forming a double loop. Inflorescence axis pubescent,

the male less branched, the female 2-5-3 cm. long,

the flowers not densely crowded, scarcely touching

each other. Fruit ellipsoid or obovoid, with some

rusty scurf when young, soon glabrous

Gardens Bulletin, S.

e. Leaves 10-22 cm. long and 3-3-9 cm. broad;

nerves 13-17 pairs, prominent beneath, 0:3—0-5

mm. thick on the undersurface; midrib 1-1-5

mm. thick at base below

(3).G. forbesii var. forbesii

e. Leaves more coriaceous, 15—28 cm. long and

4:5—9:5 cm. broad; nerves 14-21 pairs, more

prominent, 0-7-1 mm. thick on the undersur-

face; midrib 2—3 mm. thick at base below

(3) G. forbesii var. crassinervis

c. Twigs more slender, 1-2 mm. thick at the apex and 3-4

mm. thick 12 cm. down, brownish towards the apex,

grey lower down. Leaves smaller, 6-20 cm. long and

1-5-8 cm. broad (sometimes up to 25 cm. long in some

of the Celebes specimens of G. paniculata var. zippe-

liana); nerves 6—15 pairs

f. Leaves 10—16-(25) cm. long; nerves 8—15 pairs, pro-

minent beneath, often with distinct interarching

g. Leaves lanceolate, 2—5:5 cm. broad; nerves 8—11

pairs. Fruit globose

(4) G. paniculata var. paniculata

g. Leaves lanceolate to elliptic, 4-8 cm. broad;

nerves 10-15 pairs. Fruit oblong

(4) G. paniculata var. zippeliana

f. Leaves smaller, 6-15 cm. long; nerves 6—9 pairs, less

prominent or faint below, the interarching not

usually distinct

h. Leaves lanceolate or elliptic-lanceolate. Nerves

not raised above, very fine and faint on the

lower surface and not raised, cannot be felt by

rubbing with the finger. Male flowers 3—4 mm.

long. Fruit ovoid-elliptic, 2 cm. long and

1-3-1:5 cm. broad

(5) G. eugeniifolia var. eugeniifolia

h. Leaves elliptic to obovate. Nerves raised above

and below, fine but those below can be felt by

rubbing with the finger. Male flowers 4-5 mm.

long. Fruit oblong or sub-globose, larger, up to

2:8 cm. long and 2—2:5 cm. broad

(5) G. eugeniifolia var. griffithii

Vol. XVII. (1958).

(1) G. bancana (Miq.) J. Sinclair in Gard. Bull. Sing. 16 (1958)

436 Fig. 53. Pl. XIIIA.

Basionym: Myristica bancana Miq., Fl. Ind. Bat. Suppl. 1

(1861) 383; Warb., Monog. Myrist. (1897) 518.

Synonyms: Gymnacranthera murtonii (Hk. f.) Warb., Monog.

Myrist. (1897) 357 T. 20 Figs 1-3; Gamble, Mat. F.M.P. 5,

23 (1912) 223; Ridley, F.M.P. 3 (1924) 61. Myristica mur-

tonii Hk. f., Fl. Br. Ind. 5 (1886) 105; King in Ann. Roy. Bot.

Gard. Calc. 3 (1891) 297 Pl. 124 ter. M. ferruginea King in

Ann. Roy. Bot. Gard. Calc. 3 (1891) 298 Pl. 125. M. ampli-

folia Warb., Monog. Myrist. (1897) 517.

var. bancana

SUMATRA INpbRAGRI:

JAMBI:

PALEMBANG:

BANKA:

MALAY PENINSULA:

DISTRIBUTION:

TYPE MATERIAL:

VERNACULAR NAMES:

(Indrag. Bovenlanden) Kuala _Belilas,

bb27590 (L); Sungei Akar, bb28624

(BO, 1).

bb13659 (BO, L).

Sine coll. as No. 16 Medang Simpai (L) 3

sheets Nos. 2, 22 and 25 type material of

M. amplifolia; Banjuasin & Kubustreken,

bbE580 (BO, L); bbE581 (BO, L); Gra-

shoff 645 (BO, L); Bajunglintjir, Banjua-

sin, DbS8E, 2P, 697 (BO) and bb58E,

1P, 630 (BO); Rawas, Dumas 1575 (BO,

Gunong Mangkol, Kostermans & Anta 694

(BO, L, PNH); Rindik, bb11822 (BO,

L); near Jebus, Teijsmann 3279 (U);

Ayer Limau, Muntok, bb7609 (BO).

Johore and Singapore. For list see Gard.

Bull. Sing. 16 (1958) 438. Extra speci-

mens:—Corner S.F.N. 29945 (BO); Kiah

Sun. ee193 (BO); Simelair “S.FN.

89502 (BO); Wall. Cat. 6803 (LE).

Sumatra, Banka, Malay Peninsula.

M. bancana Miq., Banka, Teijsmann 3279.

M. amplifolia Warb., Palembang, S. coll.

No. 16 Medang Simpai, M. ferruginea

King, Wall. Cat. 6803 (BM, G, K, LE);

Ridley Nos. 1835 (BM, CAL, MEL,

SING); 4815 (CAL, MEL, SING) and

3364 (CAL, K, SING) all Singapore; M.

murtonii Hk. f., Singapore, Murton 13

(K).

Sumatra:—medang simpai, kayu_ asap,

punggung kidjang.

Gardens Bulletin, S.

var. borneensis (Warb.) J. Sinclair in Gard. Bull. Sing. 16 (1958)

439.

Basionym: G. murtonii (Hk. f.) Warb. var. borneensis Warb.,

Monog. Myrist. (1897) 359.

Synonym: M. murtonii Hk. f. var. borneensis (Warb.) Boerl.,

Handl. Fl. Ned. Ind. 3, 1 (1900) 88 nom. alt.

BORNEO Sarawak: Kuching, Beccari Nos. 1211 (A, C, FI, G &

Boiss, K, M, P, S) and 3977 (FI, G, K) both

as type material of this variety.

This variety is not very much different from the typical form but

the leaves are cuneate at the base and the male flowers slightly

larger, 5-6-5 mm. long and 4 mm. broad in dried material. Those

of var. bancana are 5 mm. long in dried material and up to 6 mm.

in fresh.

(2) G. contracta Warb., Monog. Myrist. (1897) 360 T. 20 Figs

1-4; Sinclair in Gard. Bull. Sing. 16 (1958) 439. Fig. 54.

Synonym: Myristica contracta (Warb.) Boerl., Handl. Fl. Ned.

Ind. 3, 1 (1900) 88 nom. alt.

SUMATRA BILLITON: van Rossum 63 (BO, L); sine coll. Herb.

Billiton 30 (L).

MALAY PENINSULA: Malacca, Singapore. For list see Gard. Bull.

Sing. 16 (1958) 441.

BORNEO SARAWAK: Near Kuching, Haviland Nos. 1650 (BO,

CAL, L, SING); and 2253 (SING);

Haviland & Hose 3308 (CAL, K, L,

SAR); Beccari Nos. 321 (BO, C, FI, G

& Boiss., K, M, NY, P, S); 419 (FI, G,

K) and 2999 (FI, G, K, P); Sungei Se-

mengoh F.R., Muas SAR 179 (SING);

Gunong Kayangeran, Lawas, Omar 81

(SING); S. Kenaban, Upper Plieran,

Pickles SAR Nos. 3446 (SING); 2496

(SING); 3526 (L); 3531 (SING).

BRUNEI: Andalau F.R., Ashton BRUN 568 (SING);

Cpt. 5, Andalau F.R., Kuala Belait,

Ashton, Smythies & Wood SAN 17514

(L, SING).

BRITISH

N. BORNEO: Mengalong F.R., Sibubu River, 34 miles

S.S.W. of Sipitang, G.H.S. Wood SAN

15155 (L, SAN, SING); Ulu Moyah, 8

miles S.S.E. of Malaman, Sipitang,

G.H.S. Wood SAN 16266 (BO, SAN,

SING); Tambato, Tambunan; Puasa &

Angian 4023 (L); 4 miles from Ranau on

path to Poring Hot Springs, G.H.S.

Wood & Charrington SAN 16368 (SAN,

SING).

PULAU SEBATIK: Kostermans 9154 (L, SING).

100

CC — a

Rees Lt

Vol. XVI. (1958).

PULAU

NUNUKAN:

PULAU

TARAKAN:

EAST AND

NorRTH-EAST

BORNEO:

SOUTH AND

SOUTH-EAST

BORNEO:

DISTRIBUTION:

TYPE MATERIAL:

VERNACULAR NAMES:

Bulungan, bb Nos. 26195 (BO, L); 29349

(BO, L) and 29364 (BO, L); Simeng-

kadu, Meijer 2395 (PNH).

Gil-Diola Pannusion, Meijer 2475 (PNH).

The following all West Kutei:—Sabintu-

lung, bb15807 (BO); Longbleh, bb16048

BO. -L); and bbiéres (BO, .L); Sl.

bb16792 (L); 6bb16838 (L); Kahala,

bb28356 (BO, L, SING) and Sei Ritan,

bb32494 (BO, L); the following 5 are

East Kutei:—Kostermans 7241 (BO, L,

PNH, SING); Sungei Bambangan, Kos-

termans 6086 (BO, L, PNH); Sungei

Sangasanga near Samarinda, Kostermans

77380 (BO, PNH, SING); Gunong Sek-

rat, S. of Sangkulirang, Kostermans 5955

(BO, L); Sungei Boi, bb14886 (BO);

Sungei Wain, Balikpapan, Achmat bb

Nos. 34310 (BO, L) and 34363 (BO,

L); Gunong Haping, Riko, Balikpapan,

bbi14971 (BO, L); Muan region near Sg.

Riko, Kostermans 4288 (BO, PNH);

Berouw, bb19027 (L) and bb19088

(BO, L); Beneden Dajak, bb19456 (BO,

L); Puruk, Tjahu, bb10029 (BO).

Karuing, Sampit, bb9928 (BO); Tehang,

Sampit, bb10233 (BO).

Billiton, Malay Peninsula and Borneo.

Gymnacranthera contracta Warb. Beccari

Nos. 321; 419 and 2999. Motley 1284

(K) ‘quoted by Warburg as a syntype is

G. forbesii var. crassinervis.

Billiton:— salak, Sarawak:— Kumpang

(Malay); badok, lahu, hungi (Kayan).

N. Borneo:— lunau (Dusun Tambato);

lupi (Kedayan). Indonesian Borneo:—

mandarahan, dara-dara, tabulah (at

Berouw).

(3) G. forbesit (King) Warb., Monog. Myrist. (1897) 363 T. 20

Figs 1—2; Gamble, Mat. F.M.P. 5, 23 (1912) 224; Ridley,

F.M.P. 3 (1924) 61; Sinclair in Gard. Bull. Sing. 16 (1958)

441 Fig. 55. Plate XIIIB.

Basionym: Myristica forbesii King in Ann. Roy. Bot. Gard.

Calc. 3: (1891). 306° Pl. 137.

var. forbesii

SUMATRA West Coast: Kampong Sipisang, Agam, bb2876 (BO).

101

Gardens Bulletin, S.

East Coast: Lumban Ria, Asahan, Rahmat Si Boeea

7984 (SING); Aek Kanopan, Lundut

Concession, Kualu, Bartlett Nos. 7052

(NY) and 7299 (NY).

INDRAGIRI: Belimbing (Indrag. Bovenlanden), bb28501

(BO, L); Danau Menkuang, bb27559

(BO, L); Kuala Belilas, bb27644 (BO,

L); Pagarumbei Tjenako, bb25783 (L).

BENKULEN: Suka Radja, Forbes Nos. 2976 (A, BM,

CAL, K, L, LE) and 3061 (A, BM, CAL,

BLL. LEY:

PALEMBANG: Lematang Ilir, Semangus, bb32217 (BO,

L) and bbT512 (BO, L); Banjuasin &

Kubustreken, Endert Nos. 62E.1P.595

(BO, L); 62E.1P.604(L); 62E.1P.606 (L);

62E.1P.610 (BO); 62E.1P.619 (BO, L);

62E.1P.620 (L); 63E.1P.620 (BO, L)

and 86E.1P.606 (BO).

PULAU

SIMALUR: Achmad Nos. 1258 (BO, L); 1316 (L);

1876 (L); 1442 (L); 1452 (L) and

1552. (L).

PULAU

ENGGANO: Near Boea-boea, Liitjeharms 4365 (A,

BQ; Lb, .NY, SENG).

BANKA: Gunong Mangkol, Kostermans & Anta 813

(BM, L, PNH); Toboali, 5b1945 (BO);

Ayer Limau, Muntok bb7843 (BO).

Riouw Ar-

CHIPELAGO: P. Bintang, Teijsmann s.n. (BO).

MALAY PENINSULA: Penang, Province Wellesley, Perak, Treng-

ganu, Pahang, Selangor, Negri Sembilan,

Malacca, Johore, Singapore. For list see

Gard. Bull. Sing. 16 (1958) 443. Extra

specimens:—Corner S.F. Nos. 28712

(BO); 28970 (BO); 36961 (BO); King

Nos. 6591 (BO, LE); 6784 (BO); 6973

(BO, LE): 7219 (BO); %756° (Eee

Negadiman S.F.N. 36866 (BO); Sinclair

S.F.N. 37936 (BO); Wray 1429 (LE).

DISTRIBUTION: Siam, Sumatra, Banka, Riouw, Malay

Peninsula.

TYPE MATERIAL: Myristica forbesii King, Penang, Maingay

1293 (CAL, K); Malacca, Maingay

1295 (K); Singapore, Ridley Nos. 6157

(SING) and 6270 (K, SING).

VERNACULAR NAMES: Sumatra:—edaran uding, salak. Banka:—-

salak.

var. crassinervis (Warb.) J. Sinclair, stat. nov.

Basionym: G. crassinervis Warb., Monog, Myrist. (1897)

362 T. 20 Figs 1-3.

Synonym: Myristica crassinervis (Warb.) Boerl., Handl. FI.

Ned. Ind. 3, 1 (1900) 88 nom. alt.—Fig. 1.

102

Vol. XVII. (1958).

Fig. 1. A, Gymnacranthera forbesii (King) Warb. B, G. forbesii var. crassi-

nervis (Warb.) J. Sinclair. C, G. forbesii. A, from Ridley 9464.

B, from Sinclair, Kadim & Kapis 9248. C, from Symington

24081 all in Herb. Sing.

Tree 10—20 m. high. Bark greyish brown, slightly rough but not

fissured, brittle, inner bark brown; sap red, copious. Leaves coria-

ceous, dark to medium green above and slightly glossy, glabrous

beneath, oblong-elliptic to oblong-lanceolate; nerves 14—21 pairs,

very prominent below and there 0-7—1 mm. thick; midrib also pro-

minent, 2—3 mm. thick below at base; length 15—28 cm.; breadth

4-5-9:5 cm.; petiole stout, 1:5—2:5 mm. thick. Inflorescence axis

rusty-puberulous. Flowers yellow, male and female more or less as

in var. forbesii. Fruit as in var. forbesii, ellipsoid or slightly obo-

void, obtuse at the apex, 2 cm. long and 1:3 cm. broad.

BORNEO Sarawak: Kuching, Beccari 1119 (Pl G; K, LE, P);

near Long Kapa, Mount Dulit (Ulu Tin-

jar), 4th Div., Richards 2289 (A, K,

SING).

BRITISH

N. BoRNEO: Sugut, Fraser 241 (CAL, K); Ulu Moyah,

8 miles S.S.E. of Malaman, Sipitang,

G.H.S. Wood, SAN 16689 (SAN, SING);

Gunong Temporangan, Ranau, Sinclair

9248 (A, B; BM, E, K, L, M,; SAN

SING).

103

Gardens Bulletin, S.

EAST AND

NORTH-EAST

BORNEO: Paru, Thony,. Tidungsche Landen, bb Nos.

17806 (L); 17837 (BO, L) and, 18256

(A, BO, L); Berouw, bb18463 (BO, L,

SING); Mentawir, Balikpapan, bb13918

(B, BO, L, SING).

SOUTH AND .

SOUTH-EAST

BORNEO: Banjermasin, Motley 1284 (K); Maram-

pan, bb11437 (BO).

WEST BoRNEO: Perigi Limus, Gunong Sendjudjuh, Sambas,

B. de Jong bb7045 (BO); Sungei Labai,

Sanggau, B. de Jong bb7859 (BO).

DISTRIBUTION: Borneo.

TYPE MATERIAL: G. crassinervis Warb., Syntypes, Beccari

1119 and Fraser 241.

VERNACULAR NAMES: Berouw:—uwkut.

I have to reduce this species to a variety of Gymnacranthera

forbesii. It is a more magnificent version of forbesii with larger,

more coriaceous leaves, and thicker nerves, midrib and _ peticle.

The records show that it is confined to Borneo whereas the var.

forbesii has not been found there.

(4) G. paniculata (A. DC.) Warb., Monog. Myrist. (1897) 370.

T. 20; Merrill in Philip. J. Sc. 1 (1906) Suppl. 55 et Enum.

Philip. Fl. Plants 2 (1923) 181; Elmer, Leafl. Philip. Bot. 3

(1911) 1059.

Basionym: Myristica paniculata A. DC. in Ann. Sci. Nat. Bot.

4, 4 (1855) 31 et Prodr. 14 (1856) 200; F.-Vill. Nov. App.

(1880) 177; Vidal, Phan. Cuming. Philip. (1885) 132 et Rev.

Pl. Vasc. Filip. (1886) 221.

Synonyms: Gymnacranthera acuminata Merr. in Philip. Journ.

Sci. Bot. 12 (1917) 265. G. laxa Elm., Leafl. Philip. Bot. 8

(1915) 2772. G. macrobotrys Merr. in Philip. Journ. Sc. Bot.

13 (1819) 284.

var. paniculata—Fig. 2.

Tree 12-30 m. high with lax, horizontally spreading branches.

Bark rather smooth, yellowish grey; sap red. Twigs slender at the

apex, reddish brown and 2 mm. thick there, lower down straw-

coloured and 3—4 mm. thick. Leaves thinly coriaceous, dark green

and shining above, glaucous beneath with yellowish green midrib,

drying olive green or brownish green above and pale brown or

glaucous beneath, lanceolate, base acute, apex acuminate; nerves

8-11 pairs, average 8, faint or indistinct above, fine but prominent

beneath, drying chocolate brown, curving and arching near the

104

Vol. XVII. (1958).

Fig. 2. Gymnacranthera paniculata (A. DC.) Warb. var. paniculata.

A, Leafy twig with male flowers. B, male flowers enlarged. C, fruit. A—B,

from Cuming 901. C, from Borden 669.

105

Gardens Bulletin, S.

margins; reticulations invisible; length 10-16 cm.; breadth 2-5-5

cm.; petiole slender, 1 cm. long. Male inflorescence much branched,

4—10 cm. long. Male flowers minutely pubescent outside and inside,

3—4 mm. long and 2—3 mm. broad; pedicels 2—4—(5) mm. long,

average 3 mm.; column 2 mm. long with 8 anthers. Female inflo-

rescence short, 1 cm. long. Female flowers not seen. Fruit globose

or sub-globose, orange outside, pinkish or pinkish orange inside,

2 cm. long and 1-8-2 cm. broad; stalk about 1 cm. long. Aril dull

pinkish red. Seed banded light brown and black.

PHILIPPINES Stine Loc.: Vidal Nos. 3552 (K) and 3558 (K).

MINDORO:

BABUYAN

Baco River, McGregor 231 (K, NY);

Paluan, Ramos Nos. 39587 (A, CAL,

NSW, NY) and 39655 (BO, CAL,

NSW); Mt. Halcon, Edafio 3584 (A);

Pinamalayan, Ramos 40816 (SING);

40819 (A, SING) and 41102 (A, DD,

G, K, UC); Bongabong River, Merritt

8663 (K).

IsLANDS: Calayan Island, Velasco 26647 (A, BO, K,

LUZON:

NY); Pefas 26705 (UC).

Bangui, Prov. Ilocos Norte, Ramos 27463

(A, BO); Bangui to Claveria, Prov. Ilo-

cos Norte, Ramos 33100 (BM, BO, CAL,

L, SING); Burgos, Prov. Ilocos Norte,

Ramos 32727 (BRI, NSW, SING);

Paraiso 30242 (NY, UC); Prov. Caga-

yan, Bernardo 13105 (BM, NSW):

Pinagsongayan River, Prov. Cagayan,

Edano 7856z. (BO; .K, NY; SENG);

Prov. Cagayan, Ramos 14552 (BM, BO,

E, K, L); Prov. Isabella, Darling 14843

(BM, NSW); Prov. Nueva Vizcaya,

Cenabre & Porte 28547 (BM); Lamao

River, Prov. Bataan, Barnes 174 (K.

NSW, NY, SING); Borden Nos. 669

(BM, K, NSW, NY, SING); 1138 (UC)

and 2940 (NY); Mt. Mariveles, Lamao

River, Prov. Bataan, Williams Nos. 566

(NY) and 640 (NY); Montalban, Prov.

Rizal, Loher Nos. 12310 (M, UC) and

13962 (BO); Balabac, Prov. Rizal, Lo-

her 14942 (A, BM); Prov. Laguna,

Amarillas Nos. 24664 (BM, TOFO) and

24940 (BO, UC); Prov. Laguna, Curran

19280 (BRI); Los Bafios, Mt. Maquiling,

Prov. Laguna, Elmer Nos. 17606 (A,

BM, BO, CAL,’G, K, L; NY, PN#H, 8S,

UC); 17962) (A. °SM, BOC.

NY, PNH, UC) and 18381 (A, BO, G,

K, L, NY, UC); Navarro 9475 (PNH)

and Perdido 9476 (PNH); Sinclair 9458

(material not yet distributed); Prov.

Laguna, Vidal 508; Baler, Principe, Prov.

Quezon, Merrill 1029 (K) and Quisum-

bing 2457 (A, PNH, SING); Casiguran,

106

Vol. XVII. (1958).

SIBUYAN:

TICAO:

SAMAR:

LEYTE:

PANAY:

SULU

ISLANDS:

BASILAN:

MINDANAO:

BORNEO BnritTIsH

N. BORNEO:

DISTRIBUTION:

TYPE MATERIAL:

VERNACULAR NAMES:

Prov. Tayabas, Ramos & Edano 45422

(B, BM, BRI, G, MEL, NY, UC); Prov.

Camarines, Ramos 1544 (A, BM, BO,

BRI, CAL, G, L, NSW, NY, PNH);

Albay, Prov. Sorsogon, Curran 10606

(BO) Albay Prov., Cuming 901 (BM, C,

Geko LE; MzsMELA NY, UPS);

Irosin, Mt. Bulusan, Prov. Sorsogon,

Elmer 15769 (A, BM,,BO, G, K, L, NY,

UC); 'Lake Bulusan, Sinclair 9586

(material not yet distributed).

Magallanes (Mt. Giting-giting), Elmer Nos.

12068 (A, BM, BO, BRSL, CAL, E, FI,

GK EE, LE, NSW, NY) and: 1273382

(A, BM, BO, BRSL, CAL, E, FI, G, K,

i LE. NSW, NY );

Clark 1016 (NY).

Sherfesee, Cenabre & Cortes 21074 (K).

Near Dagami, Ramos 1171 (BM, BRSL,

G, ‘KE; My.

Mt. Macosolon, Prov. Capiz, Ramos &

Edafio 30766 (BM, NSW, SING); Lila-

cao, Prov. Capiz, Ramos & Edajio 31450

(BM, BO, CAL, L); Prov. Capiz, Hirro

23954 (NY).

Tawitawi, Sulu Prov. Ramos & Edafio Nos.

44047 (LE, NY, UC) and 44284 (A,

5, By BO; G, Ky NY, PN. SING,

UC).

Miranda 18969 (BO, BRI, C, CAL, E, L);

Reillo Nos. 15460 (BM, BO, L, NSW,

NY) and 15489 (BM).

Santa Maria, Distr. of Zamboanga, Reillo

16444 (K, L, NY); Cabadbaran, Mt.

Urdaneta, Prov. Agusan, Elmer 13715

(A, BM, BO, E, G, K, L, LE, NSW, NY);

Butuan Sub. Prov., Miranda 20529

(BM); Surigao, Wenzel Nos. 2672 (A,

BO, G, K, M, NY, UC) and 2858 (A,

B, BO, G, K, M, NY, SING, UC).

Bukit Kasiladan, Ranau, Sinclair 9266 (B,

E, K, L, SAN, SING) sterile.

Philippines and probably North Borneo.

Lowland forests.

M. paniculata A. DC., Cuming 901. G.

acuminata Merr., Sherfesee, Cenabre &

Cortes 21074. G. laxa Elmer, Elmer

18715. G. macrobotrys Merr., Ramos

BLT Ls

anuping (Sul.); batu-batu (Bik.); dagang

(Fag:);. dumadaka (ilk.): laguan

(Mang.); magandau (Mbo.); malakapa

aa pamitogan (Ibn.); yango, (P.

is).

107

Gardens Bulletin, S.

Gymnacranthera acuminata, macrobotrys and laxa are not dif-

ferent species. It is true that there is some variability in size of

flowers and length of pedicels in G. paniculata and at first sight

laxa with its loose spreading panicle, may at least appear to be a

variety. Specimens of paniculata with young unopened flower buds

have short pedicels and look very different from those with mature

‘lowers. Such immature specimens, from what I have seen, are more

frequent in herbaria than the last-mentioned. The same observa-

tions on the compactness and laxity of young and old inflorescences

also apply to the var. zippeliana and such progressive stages of

growth ought not to mislead one. The bark of var. paniculata is

lighter and more yellowish than that of eugeniifolia or any of the

Malayan species.

Sinclair 9266 from North Borneo is sterile and is most probably

G. paniculata. It is quite reasonable that it should occur in North

Borneo.

var. zippeliana (Miq.) J. Sinclair, stat. nov.

Basionym: M. zippeliana Mig., Ann. Mus. Bot. Lugd.-Bat. 2

(1865) 50; Scheff., in Ann. Jard. Buit. 1 (1876) 45.

Synonyms: G. zippeliana (Miq.,) Warb., Monog. Myrist.

(1897) 372. G. suluensis Warb., Monog. Myrist. (1897) 373;

Elmer, Leafl. Philip. Bot. 3 (1911) 1058; Merr., En. Philip.

Flowering Plants 2 (1923) 181.—Fig. 3.

Tree 10-25 m. high, very similar to var. paniculata. Bark grey-

ish or medium brown, slightly striate; sap red. Leaves more variable

than in var. paniculata, generally longer and broader but not

always, lanceolate, oblong-lanceolate, elliptic or oblanceolate;

nerves 10—15-(17) average 12 pairs; length 10-21 cm. long;

breadth 4-8 cm. Male inflorescence branched, usually slightly

shorter than in var. paniculata. Female inflorescence 2 cm. long.

Male and female flowers 3 mm. long, adpressed pubescent inside

and outside; pedicels 3 mm. long in the male and 2 mm. long in the

female; ovary 2 mm. long, rusty tomentose; flowers of both sexes

yellow. Fruit oblong, yellowish green, ridged, 2—3 cm. long and

1-1-5 cm. broad. Seed dark brown.

PHILIPPINES Minporo: Lubang Island, Rosenbluth 12237 (CAL).

BAsILAN: Vidal Nos. 3546 (K) and 3561 (K).

MINDANAO:: Surigao Prov., Ramos & Pascasio 34462 (A,

CAL, BRI, K, NSW, NY, SING, UC);

Todaya, (Mt. Apo), District of Davao,

Elmer 10941 (A, BM, BO, BRSL, CAL,

E, FI, G, K, L, LE, NSW, NY).

108

Vol. XVI. (1958).

JURAIMI DEL

Fig. 3. Gymnacranthera paniculata var. zippeliana (Miq.) J. Sinclair.

A, Leafy twig with male flowers. B, leafy twig with female flowers. C, male

flowers in bud. D, fruit. E, male flowers enlarged. F, female flowers

enlarged. A and E, from Forbes 329 (L); B and F, from Carr 16172

(SING). C, from Hoogland 4618 (L); D, from Carr 12064 (SING).

109

CELEBES NorTH PENIN-

SULA:

CENTRAL

CELEBES:

SOUTH-EAST

PENINSULA:

PuLAU MUNA:

PULAU BUTON:

MOLUCCAS Ta.Laup

ISLANDS:

OBI

ISLANDS:

BuURU:

AMBON:

Kal

ISLANDS:

NEW

GUINEA VOoGELKOP

(DUTCH WEST

NEw

GUINEA):

DutTcH NorTH

NEw

GUINEA:

DutcH SOUTH

NEW

GUINEA:

Gardens Bulletin, S.

Manado, Minahassa, Koorders Nos. 18147

(BO, L) and 18150 (BO); Bongomeme,

Gorontalo, Manado, bb19416 (BO, L).

Poso, Lapé, bb29466 (BO, L); Usu, Mallili,

bb Cel/II 303 (BO, L); bb Cel/II 352

(BO, L) .and bbeCel/ iT 216 ABO; 2).

Thawtta, Tole-tole, Kawata, Malili, bb

Cel V/255 (BO, L); La Roua, Malili,

bb1880 (L); Malili, bb Cel/II 402 (BO,

L); Lingkomomo, Malili, bb8567 (BO);

Lobose, Malili, bb11421 (BO).

Sangona, Kjellberg 950 (BO, S).

Wakadea, bb5870 (L).

Kambowa, bb6643 (BO) and bb6645 (BO,

ic).

S.W. slope of G. Duatu, Karakelong, Lam

2919 (L, PNH) and 2942 (L); E. slope

of G. Piapi, Karakelong, Lam 32329 (L.).

Laiwui, bb23778 (A, BO, L, SING) and

5b28779 (A, BO, L, SING).

Balo-balo, bb25191 (L); Walpangat,

bb223875. (1).

Robinson 289 (A, BM, BO, K, L, NY).

Jaheri 80 (BO) and 714 (L).

Sorong, Beccari 221 (FI); Andai, Beccari

705 (FI); Pikpik, bb22252 (BO) and

bb22273 (A, BO, L); Modan, bb22311

(BO); Manokwari, Dessa, Momi,

bb33490 (A, L, SING) and Kostermans

800 = bb33490 (A, BO, L, SING);

Dossi, Manokwari, bb15907 (BO); Ta-

nindi, 3 km. from Manokwari, Koster-

mans 2975 (A, BO, K, L); McCluer

Gulf, Warburg 20723 (A); P. Aiduma

(Aituma) Zippelius s.n. (CAL, L);

Rauna, bb22537 (BO); Sennen, Nabire,

Kanehira & Hatusima s.n., 7th March,

1940 (BO); Fakfak, Onakasi, bb32694

(BO) 1):

Bernard Bivak, Hollandia, bb25748 (L,

BO, SING); Pionier Bivak, Hollandia,

bb31329 (BO).

s.l., Versteeg 1472 (BO, K, L).

110

Vol. XVII. (1958).

PAPUA:

MANDATED

TERRITORY

OF NEw

GUINEA

(KAISER

WILHELMS-

LAND):

Thu, Vailala River, Brass 971 (A, BRI, K,

SING); Wuroi, Oriomo River, Western

Division, Brass 5903 (A, BRI, BO, NY);

Lake Daviumbu, Middle Fly River, Brass

7897 6((A, BM; BO,. BRI, L,. LE);

Kokoda, Carr Nos. 16141 (SING);

16172 (CANB, L, SING) and 16467

(BO, SING); Koitaki Carr 12064 (L,

SING); Sogeri Region, Forbes Nos. 236

(A, BM. CA BE KL, LE, PNH,

SING); 329 (A, BM, CAL, E, FI, K, L,

LE. OPNH, SING); 350 (BO, CAL, E,

Pe LL, LE): 37 4BM, CAL, E, FI,

K); 396 (BM, E, FI, L); 646 (BM,

CAL, FI, K, L) and 709 (BM, FI, LE);

about 1 km. S.E. of Bundi Barracks,

Tufi sub-district, Northern Division,

Hoogland 4618 (A, CANB, L, LAE);

Vaimuru, Vanapa River, Central Div.,

McDonald N.G.F. 8166 (CANB, SING);

Yalu, Atzera Range near Lae, Womer-

sley N.G.F. 3205 (BRI, CANB, LAE).

Sepik, Ledermann Nos. 6551 (SING); 6917

(SING); 7790 (SING); 9659 (L) and

10456 (L); Quembung, Morobe District,

Clemens Nos. 2149 (A, B, SING); and

2189 (A, B, SING); above Gabensis,

Morobe District, Floyd N.G.F. 7251

(BRI, CANB, LAE, NSW, SING); Sat-

telberg, Nyman 750 (BRSL); Wobbe,

Schlechter 16338 (A, E, G, K, L, NY,

S, SING); Gati Mts., Schlechter 17186

(A, E. G, K, L; NY, S,~SENG); Ramu

Valley about 5 miles S.E. of Faita Air

Strip, Madang sub-division, Saunders

Nos. 208 (CANB, LAE); 229 (CANB,

LAE); 236 (CANB, LAE); 247 (CANB,

L, LAE); 255 (CANB, L, LAE); 260

(CANB, LAE); 265 (CANB, L, LAE);

293 (CANB, L, LAE); 297 (CANB, L,

LAB): 308 (CANE. EL, ~LAE): .338

(CAND. LL, LABRY 7-252 (CANB,. L,

LAB): S6i (CANBY E, “LAE);* 373

(CANB, L, LAE); 394 (CANB, LAE);

405 (CANB, L, LAE); 451 (CANB, L,

LAE); 477 (L, LAE); 480 (A, CANB,

LL, LAE); 492 (CANB, LAE); 497

(CANE, 1; LAE); 502. (CANB, L);

504A (LAE); 507 (CANB, LAE); 512

(CANB, L, LAE); 515 (CANB, LAE);

517 (CANB, LAE); 521 (CANB); 522

(CANB, LAE) and 526 (LAE); Bengi,

Lane-Poole 805 (BRI); Dobodura Area,

N.G.F. 2036 (BRI, LAE).

bit

Gardens Bulletin, S.

NEw BRITAIN: Keravat, Coppack N.G.F. 7036 (BRI,

CANB, LAE, SING); Floyd Nos. 3449

(CANB, BRI, BO, L, NSW, PNH,

SING) and 7009 (BRI, CANB, L, LAE,

NSW, PNH, SING); Womersley N.G.F.

8409 (BM, BRI, CANB, LAE, NSW,

PNH, SING); Vudal Divide, Womersley

N.G.F. 7929 (BRI, CANB, NSW,

SING).

SCHOUTEN

ISLANDS: P. Biak, Feuilletau de Bruyn 446 (BO);

bb. Nos. -80770:{A, BO, LL, SING?)

30805 (BO; 13730819 (A, BO? A.

SING).

PULAU JAPEN: Aet & Idjan 798 (A, BO, K, L); bb Nos.

80254 (A, BO, L, SING); 30490 (BO,

L, SING); 30493 (A, BO, L, SING);

bb30528 (A, BO, L); 30532 (A, BO, L,

SING); 30536 (A, BO, L, SING) and

30591 (A, BO; LL; ‘SING): ‘Sera

bb30657 (BO); Mariarattu, bb. Nos.

80378 (BO, L, SING); 30396 (A, BO,

L); 30420 (A, BO, L); 30427 (BO, L,

SING); 30430 (A, BO, L, SING) and

30459 (A, BO, L, SING).

DISTRIBUTION: Philippines, Celebes, Moluccas, New

Guinea.

TYPE MATERIAL: M. zippeliana Miq., P. Aiduma, Zippelius

s.n., G. suluensis Warb., Syntypes Vidal

Nos. 3546 and 3561.

VERNACULAR NAMES: Talaud Islands:— lahu. New Guinea:—

saksak, hokol (Amele); mobo, gadagod,

kini (Bilia); gisek, sarenki, gaigihab,

dzidzir, minip, bisip (Dumpu); gamu-

kua (Faita); koreaf (Onjob); New Bri-

tain:-—- goma (Maprik); kokomo

(Pidgin).

I cannot accept G. paniculata and zippeliana as two distinct

species after having examined the large list of specimens enume-

rated above. The former has a globose fruit and the latter an

oblong one and this seems to be the best character for distinguish-

ing them. The leaves of the variety zippeliana are generally larger

and broader with more veins, but this is not always the case and

one may have to rely on geographical distribution to identify

sterile material.

(5) G. eugeniifolia (A. DC.) J. Sinclair in Gard. Bull. Sing. 16

(1958) 444. Fig. 56. Pl. XIV.

Basionym: Myristica eugeniifolia A. DC., Ann. Sc. Nat. 4, 4

(1855) 29 et Prodr. 14, 1 (1856) 190; Migq., Fl. Ind. Bat. 1, 2

(1858) 58; Hk. f., Fl. Br. Ind. 5 (1886) 113.

Synonyms: Gymnacranthera farquhariana (Hk. f. et Th, sensu

King) Warb., Monog. Myrist. (1897) 365 T. 22 Figs 1-2;

112

Vol. XVII. (1958).

Gamble, Mat. F.M.P. 5, 23. (1912) 225; Ridley, F.M.P. 3

(1924) 62 [omnino pro parte quoad specimina malayana tan-

tum]. Myristica farquhariana Hk. f. et Th. sensu Fl. Br. Ind. 5

(1886) 108 pro parte; King in Ann. Roy. Bot. Gard. Calc. 3

(1891) 305 pro parte [non Hk. f. et Th., Fl. Ind. (1855) 162

sensu stricto — A. DC., Prodr. 14, 1 (1856) 200 et Migq., FI.

Ind. Bat. 1, 2 (1858) 63 = species ex India tantum — Synonym

G. canarica (King) Warb.]

var. eugeniifolia

SUMATRA TAPANULI: Sibolga, 5b19305 (A, L).

WEsT CoasT: Painan, Br. Belantai, bb3995 (L).

East Coast: Masihi F.R., Asahan, Krukoff 4052 (A,

BRI, BO, G, L, LE, NY, SING); vicinity

of Lumban Ria, Asahan, Rahmat Si

Boeea 8035 (A, S, SING); Ayer Kandis

near Rantau Parapat, Bilia, Rahmat Si

Toroes 2407 (A, NY, UC).

INDRAGIRI: (Indrag. Bovenlanden) Sungei Akar,

bb28598 (BO, L, SING); Hutan Pulau

Lawas near Taratak Ayer Hitam, Taluk

region, Meijer 4394 (SING).

LINGGA: P. Singkep, bb5891 (BO).

MALAY PENINSULA: Kedah, Penang, Perak, Selangor, Negri

Sembilan, Malacca, Johore, Singapore.

For list see Gard. Bull. Sing. 16 (1958)

446. Extra specimens:—Corner S.F.N.

29955 (BO); Hervey Aug. 1886, Malacca

(BO); King Nos. 4640 (BO, LE); 5801

(BOLE); 6447 (LE); 6620 (BO, LE);

6631 (BO) and 6932 (LE); Ngadiman

wieIN: 84628 (BO).

DISTRIBUTION: Sumatra, Lingga and Malay Peninsula.

TYPE MATERIAL: M. eugeniifolia A. DC., Penang, Gaudi-

chaud 116 (FI, G & Prodr. holotype, P).

VERNACULAR NAMES: Sumatra:—mandaharan, kayu gadjah.

var. griffithii (Warb.) J. Sinclair in Gard. Bull. Sing 16 (1958)

447 Fig. 57.

Basionym: G. farquhariana var. griffithii (Hk. f.) Warb.

Monog. Myrist. (1897) 368; Gamble, Mat. F.M.P. 5, 23 (1912)

226 pro parte; Ridley, F.M.P. 3 (1924) 62.

Synonyms: Myristica griffithii Hk. f. Fl. Br. Ind. 5 (1886)

109; King in Ann. Roy. Bot. Gard. Calc. 3 (1891) 304 Pl. 135

excl. Curtis 2406 and 2458. M. farquhariana Wall. Cat. 6795,

sphalmate 6798, nom. nudum; Hk. f. et Th., Fl. Ind. (1855)

162; Hk. f., Fl. Br. Ind. 5 (1886) 108; King in Ann. Roy. Bot.

Gard. Calc. 3 (1891) 305 Pl. 135 [omnino pro parte]. G. far-

quhariana (Hk. f. et Th.) Warb. sensu Warb., Monog. Myrist.

113

Gardens Bulletin, S.

(1897) 365; Gamble, Mat. F.M.P. 5, 23 (1912) 226; Ridley,

F.M.P. 3 (1924) 62. M. farquhariana var. major King in Ann.

Roy. Bot. Gard. Calc. 3 (1891) 306 Pl. 136 Fig. 4. G. farqu-

hariana var. major (King) Gamble, Mat. F.M.P. 5, 23, (1912)

226; Ridley, F.M.P. (1924) 62. G. apiculata Warb., Monog.

Myrist. (1897) 359 T. 20 Figs 1-2. M. apiculata (Warb.)

Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 88 nom alt.

SUMATRA East Coast: Langga Pajung, Sungei Kanan, Kota Pi-

BANKA:

MALAY PENINSULA:

BORNEO Sarawak:

BRUNEI:

BRITISH

N. BORNEO:

EAST AND

NORTH-EAST

BORNEO:

DISTRIBUTION:

TYPE MATERIAL:

nang, Rahmat Si Toroes 3425 (A, NY);

Gunong Si Papan, Kalubi, Kota Pinang,

Rahmat Si Toroes Nos. 3886 (A, NY)

and 3944 (A, NY).

Bihang, S. Banka, bb Nos. 15100 (BO)

and 15104 (BO).

Perak, Selangor, Malacca, Johore, Singa-

pore. For list see Gard. Bull. Sing. 16

(1958) 447. Extra specimens:—Corner

S.F.N. 29499 (BO); Griffith 4355 (LE);

Kiah S.F. Nos. 32157 (BO) and 37718

(BO); King 7928 (BO, LE); Wall. Cat.

6795 (LE); Ridley 3365 (NSW).

Beccari 2246 (FI, G, K, LE, S); Hose

(probably), June 1906, PNH acc. No.

23664 (PNH); Loba Kubang Protected

Forest, Jara SAR 540 (SAR, SING);

Anderson SAR 2723 (SAR, SING);

Baram, Anderson SAR Nos. 2017 (SAR,

SING) and 3251 (SAR, SING); 6th

mile F.R., Kuching, Muas SAR 98

(SING).

Berakas F.R., Ashton BRUN 837 (SING);

Labu F.R., Temburong, Ashton, Smy-

thies and Wood SAN 17434 (SAN,

(SING).

Damit River, Md. Tahir 798 (A, NY,

SING).

Sebakis R. region, Kostermans 9301 (L,

SING).

Sumatra, Banka, Malay Peninsula and

Borneo.

G. farquhariana var. griffithii (Hk. f.)

Warb., Malacca, Griffith 4356 (A, CAL,

K, holotype, P). M. farquhariana var.

major King, Perak, King Nos. 6548

(CAL; DD,, E,.. FLAK, L);- 6622. .8M,

CAL, DD, G, K, L, SING); 6736 (CAL,

MEL, UPS):.7928 (GAL, FI) G, Ko)

and Perak Wray Nos. 23899 (CAL, K,

SING) and 2695 (CAL, FI, K, SING);

Malacca, Griffith 4355 (A, CAL, FI, K,

LE, M); G. apiculata Warb., Sarawak,

Beccari 2246 (FIfG)K, LES):

VERNACULAR NAMES: Sarawak:—Kumpang (Malay).

114

Vol. XVII. (1958).

Excluded Species

Gymnacranthera_ cryptocaryoides Elmer — Knema_ kunstleri

(King) Warb.

G. ibutii Holth. — Horsfieldia macrocoma (Miq.) Warb.

G. lanceolata Merr. = Myristica agusanensis Elmer

G. negrosensis Elmer = M. cumingii Warb.

G. stenophylla Warb. — K. stenophylla (Warb.) Sinclair

G. sulphurascens Elmer — M. elliptica var. simiarum (A. DC.)

Sinclair.

G. urdanetensis Elmer = M. cumingii Warb.

LIST OF COLLECTORS’ NUMBERS

ACHMAT—1258; 1316; 1376; 1442; 1452 and 1554 forbesii var.

forbesii; bb34310 & bb34363 contracta.

AET & IDJAN—798 paniculata var. zippeliana.

AMARILLAS—24664 & 24940 paniculata var. paniculata.

ANDERSON—SAR Nos. 2017; 2723 & 3251 eugeniifolia var.

griffithii.

ASHTON—BRUN 568 contracta; BRUN 837 eugeniifolia var.

eriffithii.

ASHTON, SMYTHIES & Woop—-SAN Nos. 17434 eugeniifolia var.

grifithi; 17514 contracta.

BARNES—174 paniculata var. paniculata.

BARTLETT—7052 & 7299 forbesii var. forbesii.

bb Nos.—1880 paniculata var. zippeliana; 1945 forbesii var.

forbesii; 2252 paniculata var. zippeliana; 2876 forbesii var.

forbesii; 3995 eugeniifolia var. eugeniifolia; 5870 paniculata

var. zippeliana; 5891 eugeniifolia var. eugeniifolia; 6643 &

6645 paniculata var. zippeliana; 7045 forbesii var. crassi-

nervis; 7609 bancana var. bancana; 7843 forbesii var. forbesii;

7859 forbesii var. crassinervis; 8567 paniculata var. zippe-

liana; 9928 contracta; 10029 & 10233 contracta; 11421 pani-

culata var. zippeliana; 11437 forbesii var. crassinervis; 11822

& 13659 bancana var. bancana; 13918 forbesii var. crassi-

nervis; 14886 & 14971 contracta; 15100 & 15104 eugeniifolia

var. griffithii; 15807 contracta; 15907 paniculata var.

zippeliana; 16048; 16146; 16792 & 16838 contracta; 17806;

17837; 18356 & 18463 forbesii var. crassinervis; 19027 &

19088 contracta; 19305 eugeniifolia var. eugeniifolia; 19416

{15

Gardens Bulletin, S.

paniculata var. zippeliana; 19456 contracta; 22273; 22311;

22537; 22815; 23778; 23779; 25191 & 25748 paniculata

var. zippeliana; 25783 forbesii var. forbesii; 26195 contracta;

27559 forbesii var. forbesii; 27590 bancana var. bancana;

27644 forbesii var. forbesii; 28356 contracta; 28501 forbesii

var. forbesii; 28598 eugeniifolia var. eugeniifolia; 28624

bancana var. bancana; 29349 & 29364 contracta; 29466

paniculata var. zippeliana; 30254; 30379; 30396; 30420;

30427; 30430; 30459; 30490; 30493; 30528; 30532; 30536;

30591; 30657; 30770; 30805; 30819 & 31329 paniculata var.

zippeliana; 32217 forbesii var. forbesii; 32494 contracta;

32694 & 33490 paniculata var. zippeliana; 34310 & 34363

contracta; E580 & E581 bancana var. bancana; T512 forbesii

var. forbesii; Cel/II 303; Cel/H 316; Cel/Tl 352; Cel/il

402; Cel/V 255 paniculata var. zippeliana; 58E 1P630 & 58E

2P697 bancana var. bancana; 62E 1P595; 62E 1P604; 62E

1P606; 62E 1P610; 62E 1P619; 62E 1P620; 63E 1P620;

86E 1P606 forbesii var. forbesii.

BrEccARI—119 forbesii var. crassinervis; 221 paniculata var.

zippeliana; 299; 321 & 419 contracta; 705 paniculata var.

zippeliana; 1211 bancana var. borneensis; 2149 & 2189 pani-

culata var. zippeliana; 2246 eugeniifolia var. griffithii; 3977

bancana var. borneensis.

BERNARDO—13105 paniculata var. paniculata.

BORDEN—669; 1138 & 2940 paniculata var. paniculata.

Brass—971; 5903 & 7897 paniculata var. zippeliana.

CARR—12064; 16141; 16172 & 16467 paniculata var. zippeliana.

CENABRE & PORTE—28547 paniculata var. paniculata.

CLARK—1016 paniculata var. paniculata.

CLEMENS—2149 & 2189 paniculata var. zippeliana.

CopPpAckK—N.G.F. 7036 paniculata var. zippeliana.

CURRAN—10606 & 19280 paniculata var. zippeliana.

DARLING—14843 paniculata var. paniculata.

DumMAs—1575 bancana var. bancana.

EpaNo—3584 & 78564 paniculata var. paniculata.

ELMER—10941 paniculata var. zippeliana; 12068; 12138a;

13715; 15769; 17606; 17962. & 18381 paniculata var.

paniculata. .

FEUILLETAU de BRUYN—446 paniculata var. zippeliana.

116

Vol. XVII. (1958).

FLoyp—N.G.F. Nos. 3449; 7009 & 7251 paniculata var.

zippeliana.

ForBES—236; 329; 350; 375; 396; 646 & 709 Meer var. Zip-

peliana; 2976 & 3061 forbesii var. forbesil.

FRASER—241 forbesii var. crassinervis.

GRASHOFF—645 bancana var. bancana.

HAVILAND—1650 & 2253 contracta.

HAVILAND & HosE—3308 contracta.

Hirro—23954 paniculata var. paniculata.

HooGLAND—4618 & 4896 paniculata var. zippeliana.

Hose—June 1906, PNH acc. No. 23664 eugeniifolia var.

eriffithil.

JAHERI—80 & 714 paniculata var. zippeliana.

JARA—SAR 540 eugeniifolia var. griffithi.

B. de Jonc—bb 7045 & bb 7859 forbesii var. crassinervis.

KJELLBERG—950 paniculata var. zippeliana.

KoorpERS—18147 & 18150 paniculata var. zippeliana.

KOSTERMANS—300 & 2975 paniculata var. zippeliana; 4288;

5955; 6086; 7241; 7730 & 9154 contracta; 9301 eugeniifolia

var. griffithii.

KOSTERMANS & ANTA—694 bancana var. bancana; 813 forbesii

var. forbesii.

KRUKOFF—4052 eugeniifolia var. eugeniifolia.

LamM—2919; 2942 & 3329 paniculata var. zippeliana.

LANE-POOLE—805 paniculata var. zippeliana.

LEDERMANN—6551; 6917; 7790; 9659 & 10456 paniculata var.

zippeliana.

LOHER—12310; 13962 & 14942 paniculata var. paniculata.

LUTJEHARMS—4365 forbesii var. forbesii.

MCcCDONALD—N.G.F. 8166 paniculata var. zippeliana.

Mc Grecor—231 paniculata var. paniculata.

Mp TAHIR—798 eugeniifolia var. griffithii.

MEIJER—2395 & 2475 contracta; 4394 eugeniifolia.

MERRILL—1029 paniculata var. paniculata.

MERRITT—3663 paniculata var. paniculata.

MiraNnpA—18969 & 20529 paniculata var. paniculata.

117

Gardens Bulletin, S.

MoTLeyY—1284 forbesii var. crassinervis.

Muas—SAR Nos. 98 eugeniifolia var. griffithii; 179 contracta.

NAVARRO—9475 paniculata var. paniculata.

N.G.F.—2036 paniculata var. zippeliana.

NyMAN—750 paniculata var. zippeliana.

OMAR—S81 contracta.

PARAISO—30242 paniculata var. paniculata.

PERDIDO—9476 paniculata var. paniculata.

PENAS—26705 paniculata var. paniculata.

PICKLES—SAR Nos. 3446; 3496; 3526; 3531 contracta.

Puasa & ANGIAN—4023 contracta.

QUISUMBING—2457 paniculata var. paniculata.

RAHMAT SI BOEEA—7984 forbesii var. forbesii; 8035 eugeniifolia

var. eugeniifolia.

RAHMAT SI ToroES—2407 eugeniifolia var. eugeniifolia; 3425

& 3944 eugeniifolia var. griffithii.

RaMos—1171; 1544; 14552; 27463; 32727; 33100; 39587;

39655; 40816; 40819; 41102 paniculata var. paniculata.

Ramos & EpDANO—30766; 31450; 44047; 44284 & 45422 pani-

culata var. paniculata.

Ramos & Pascaslo—34462 paniculata var. zippeliana.

REILLO—15460; 15489 & 16444 paniculata var. paniculata.

RICHARDS—2289 forbesii var. crassinervis.

ROBINSON—239 paniculata var. zippeliana.

ROSENBLUTH—12237 paniculata var. zippeliana.

van RossuUM—63 contracta.

SAUNDERS—208; 229; 236; 247; 255; 260; 265; 293; 297; 308;

338; 352; 361; 378; 394; 405; 451; 477; 480; 492; 497;

504; 504A; 507; 512; 515; 517; 521; 522 & 526 paniculata

var. zippeliana.

SCHLECHTER—16338 & 17186 paniculata var. zippeliana.

SHERFESEE, CENABRE & CoRTES—21074 paniculata var.

paniculata.

SINCLAIR—9248 forbesii var. crassinervis; 9266, 9458 & 9586

paniculata var. paniculata.

TEIJSMANN—3279 bancana var. bancana; s.n. forbesii var. forbesii.

VELASCO—26647 paniculata var. paniculata.

VERSTEEG—1472 paniculata var. zippeliana.

118

Vol. XVII. (1958).

VipaL—508 paniculata var. paniculata; 3546 paniculata var. zip-

peliana; 3552 & 3558 paniculata var. paniculata; 3561

paniculata var. zippeliana.

WARBURG—20723 paniculata var. zippeliana.

WENZEL—2672 & 2858 paniculata var. paniculata.

WILLIAMS—566 & 640 paniculata var. paniculata.

WoOMERSLEY—3205 & 7929 paniculata var. zippeliana.

G.H.S. Woop—SAN Nos. 15155 & 16266 contracta; 16689 for-

besii var. crassinervis.

G.H.S. Woop & CHARRINGTON—SAN 16368 contracta.

ZIPPELIUS—s.n. paniculata var. zippeliana.

119

Index To Gymnacranthera

Synonyms are printed in italics and new combinations in bold-faced type.

Gymnacranthera, 96.

acuminata Merr., 104.

apiculata Warb., 114.

bancana (Migq.) J. Sinclair, 99.

var. bancana, 99.

var. borneensis (Warb.) J. Sin-

clair, 100.

canarica (King) Warb., 96, 113.

contracta Warb., 100.

crassinervis Warb., 102.

cryptocaryoides Elmer, 115.

eugeniifolia (A. DC.) J. Sinclair,

at2:

var. eugeniifolia, 113.

var. griffithii (Warb.) J. Sinclair,

3S,

farquhariana (Hk. f. et Th.) Warb.,

farquhariana (Hk. f. et Th. sensu

King) Warb., 112.

farquhariana (Hk. f. et Th.) Warb.

sensu Warb., 113.

var. griffithii (Hk. f.) Warb., 113.

var. major (King) Gamble, 114.

forbesii (King) Warb., 101.

var. crassinervis (Warb.) J. Sin-

clair, 102.

var. forbesii, 101.

ibutii Holth., 115.

lanceolata Merr., 115.

laxa Elmer, 104.

macrobotrys Merr., 104.

murtonii (Hk. f.) Warb., 99.

var. borneensis Warb., 100.

negrosensis Elmer, 115.

paniculata (A. DC.) Warb., 104.

var. paniculata, 104.

var. zippeliana (Migq.) J. Sinclair,

108.

stenophylla Warb., 115.

sulphurascens Elmer, 115.

suluensis Warb., 108.

urdanetensis Elmer, 115.

zippeliana (Miq.) Warb., 108.

Horsfieldia, 115.

macrocoma (Mig.) Warb., 115.

Knema, 115.

kunstleri (King) Warb., 115.

stenophylla (Warb.) J. Sinclair, 115.

Myristica, 115.

agusanensis Elmer, 115.

amplifolia Warb., 99.

apiculata (Warb.) Boerl., 114.

bancana Migq., 99.

contracta (Warb.) Boerl., 100.

crassinervis (Warb.) Boerl., 102.

cumingii Warb., 115.

elliptica var. simiarum (A. DC.) J.

Siclam,. 115.

eugeniifolia A. DC., 112.

farquhariana Wall. Cat. 6795, 113.

farquhariana Hk. f. et Th. sensu FI.

Br. Ind., 113.

var. major King, 114.

ferruginea King, 101.

forbesii King, 101.

griffithii Hk. f., 113.

murtonii Hk. f., 99.

var. borneensis (Warb.) Boerl.,

100.

paniculata A. DC., 104.

zippeliana Migq., 108.

BOOK REVIEW

Common Malayan Plants—(Selected Drawings with notes) by H. B.

Gilliland, Professor of Botany, University of Malaya. Published

by the University of Malaya Press, Singapore 1958—228 pages and

222 line drawings. (Price M$5).

This handy pocket-size volume consists of line drawings of

common Malayan plants most of which are selected from a war-

time publication in Japanese by Professor Watanabe of the Chiba

University entitled “Illustrations of Useful Plants of the Southern

Regions”. It is more or less an abridged edition of the Japanese

book with short notes in English, while a few new drawings, pre-

pared by ’Che Juraimi bin Samsuri, artist at the Botanic Gardens,

Singapore, have been added. The paper is good, of much better

quality than that of the original book, and the general set-up of the

type is clear. There is an attractive cover.

The purpose of the book is to interest younger students in the

plants which grow around them. The pictures should inspire them

with a genuine and spontaneous interest in a far better way than

text books with difficult words and scientific terms can ever do.

Thus drudgery, boredom and tears are avoided and their early

interests are not frustrated but fostered. The book ought also to be

of some interest to the tourist passing through Malaya, who inevit-

ably sees some strange fruit in the market and wishes to know more

about it. Teachers repeatedly visit the Botanic Gardens and ask

for simple publications on plants for school lessons. There is a

shortage of such and it is hoped that the present one may meet

their needs. The choice of plants for illustration is a good basic

representation of what is commonly met with round villages and is

more likely to interest the layman than a selection of the rare ones

or those from the remote jungles.

If the book falls short of anything at all, then perhaps it might

be criticized a little by Malay speaking students on the author’s

choice of Malay names. He has not always selected the commonest

Malay name. His choice is often with emphasis on names used in

the north. For example, he calls the well-known bunga tanjong,

Mimusops elengi, mengkula. Sesbania grandiflora is getih, but it is

far better known as turi in Singapore and Johore, the leaves being

used as a vegetable. Paku laut or paku hadji would have been

better for Cycas rumphii than bogak hutan which is probably

seldom used. Saga is sufficient for Adenanthera pavonina without

121

Gardens Bulletin, S.

the epithets daun tumpul and likewise Durio zibethinus is simply

called durian and not durian kampong. Finally the three species of

Ocimum namely basilicum, sanctum and canum are often confused

on account of their similarity. The name selaseh is more correcily

applied to O. sanctum, the sacred tulasi of the Hindus and not

to O. basilicum which is selaseh hitam or jantan.

There has been more uniformity and standardization recently in

the use of botanical names of plants and most of Professor

Gilliland’s names are correct and up-to-date according to the rules

of botanical nomenclature. The following are the exceptions

noted:—Rhizophora conjugata should now be R. apiculata while

Bruguiera conjugata becomes B. gymnorrhiza. The better known

name Scaevola frutescens ought now to be changed to S. sericea

and Bouea macrophylla to B. gandaria. Neptunia oleracea is older

than N. prostrata while Lagerstroemia speciosa takes precedence

over L. flos-reginae.

The book is almost free from misprints. On page 25 there is

Averrhoa calambola for carambola but here the sound is not really

very different and anyway neither the Chinese nor certain English

people ever lay much stress on the letter r. Adenostemma lavenia

(lavenia a classical Ceylonese name) is misspelt laevinia.

The notes accompanying each plate are short but very interesting

and there is often information which trained botanists would do

well to read. The author states that the flowers of Dillenia indica

and Momordica cochinchinensis are yellow. This is not quite cor-

rect as the petals of the former are white but the stamens are

yellow. The petals of the latter are white or cream or white with a

yellow tinge. Not-with-standing these slight defects the book is quite

useful and we hope to see more of its kind at some future date.

J. SINCLAIR.

122

De,

em _

fics

= at

OTHER PUBLICATIONS OF THE BOTANIC GARDENS

SINGAPORE

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Reports for many years 1886 onward remain available.

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Vols. 1-10, 1901-12, monthly issues.

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4. The Gardens’ Bulletin, Straits Settlements (Series III).

Vol. 1 (1-5) Jan.-May 1912, issued under title of

Agricultural Bulletin of the Straits and F.M.S. and

Vol. 1 (6-12) Dec. 1913—March 1917 as The Gardens’

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Present subscription per volume payable in advance,

$20 in Malaya, or $21 outside.

6. Malayan Garden Plants: a series of six booklets each

illustrating 10 useful garden plants.

10 useful garden plants.

All available. Price 50 cents each.

7. An illustrated guide to the Botanic Gardens Singapore.

Price $1.

8. Materials for a Flora of the Malay Peninsula.

Pt. I, II and III remain available.

9. A Revised Flora of Malaya, Vol. 1 Orchids by R. E. Holttum.

Government Printing Office, Singapore, 1953 ed. I,

1957 ed. II. Price $20, second edition.

10. A Revised Flora of Malaya, Vol. 2, Ferns by R. E. Holttum.

Government Printing Office, Singapore 1955. Price $20.

11. Malayan Orchid Hybrids, by M. R. Henderson and G. H.

Addison.

Government Printing Office, Singapore 1956. Price $21.

12. Wayside Trees of Malaya by E. J. H. Corner, 2 Vols.

Government Printing Office, Singapore. Price $25.

Items 1-8 can be obtained from the Botanic Gardens,

Singapore.

Items 9-12 can be obtained from the Government

Printer, Singapore.

Prices quoted in Malayan Dollars.

In certain cases postage is extra.

2758 —-400—11/58

Andssue to mark the

~ occasion of the

100th Anniversary

of the founding of the

Botanic Gardens, Singapore

THE

GARDENS’ BULLETIN

SINGAPORE

Volume XVII

-Part Il

Sth December, 1959

— NOLD ARSO ——,

Oe” RECEIVED =7U

' APR15 1960

LIBRARY

Published by Authority

To be purchased at the Botanic Gardens, Singapore

Price: $12.50

CONTENTS

DEDICATION

PURSEGLOVE, J. W.: History sed Punedons of Botdvic Galens with

special reference to Singapore 3 : , P

RussEL, T. A.: Kew and Singapore .

VAN STEENIS, C. G. G. J.: Singapore and Flora Salesian

Lam, H. J.: A tale of two cities: Singapore and Leiden

BuRKILL, I. H.: A note on the Gardens’ Jungle

WYATT-SMITH, J.: The Seance Botanic Gardens and Forestry in

Malaya : ;

WYCHERLEY, P. R.: The Singapore Botanic Galle and Rubber in

Malaya :

ANONYMOUS (A Peele A ceautye The Contiibuieeh| to Agri-

culture in Malaya by the Singapore Botanic Gardens ,

Hotttum, R. E.: Orchids, Gingers and Bamboos; Pioneer work at

the Singapore Botanic Gardens and its ee for * {ieee and

Horticulture :

FurTapbo, C. X.: Singapore’s Gantabniies to the Study of ble

QUISUMBING, E., Director, National Museum, Manila: Manila and the

Singapore Gardens)

BuRKILL, H. M.: The Botanic Gatton and Generar in | wlan

Kusnoto SETYODIWIRYO: The Singapore Botanic Gardens and the

Central Institute for Nature Research in Indonesia at Bogor.

Comments on Past and Present a and the Need to

continue it . , |

Corner, E. J.:H.: The Importance of Tropical Taxonomy to modern

Botany :

Gi_mour, J. S. L.: The Misrnational Code of Nomencineae tae Cul-

tivated Plants, with Special Reference to Tropical Botanic Gardens

FosBerG, F. R.: The cibiticanek of piace Research in the Pacific

Region

MaTTSSON, L.: Role of hoeanied! pandcas in cae Ena Tropics and

UNESCO's Programme related to them :

GILLILAND, H. B. and WANTMAN, M. J.: Regenerating High Bones on

Singapore Island :

ROBINSON, R. A.: The pH of Rain ae on the Botan Garten

MOLESWORTH ALLEN, B.: Malayan Fern Notes

MOLESWORTH ALLEN, B.: Malayan Fern Notes, II .

KERN, J. H. and vAN STEENIS, C. G. G. J.: An interesting new Record

from the Malayan Beach: ssa a urens ae its synonymy

and distribution : 7

FurtTapbo, C. X.: A New rer Goi Spal

Furtapo, C. X.: Some New or Noteworthy Species of Wea sta

JOHNSON, A.: The Genus Sphagnum in Malaysia

JOHNSON, A. and TAN KiAP SENG: Cleome ciliata Schum. et Thonn.

in Singapore ‘ : ’ :

OBITUARIES:

Haji Mohamed Nur bin Mohamed Ghous, B.E.M. .

Ngadiman bin Haji Ismail . ;

Kwan Koriba

PAGE

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VOLUME XVII STH DECEMBER, 1959 PART II

THE

GARDENS’ BULLETIN

SINGAPORE

AN ISSUE TO MARK THE OCCASION OF THE

HUNDREDTH ANNIVERSARY

OF THE FOUNDING OF THE

BOTANIC GARDENS, SINGAPORE

1859 - 1959

History does not record the exact date.

The nearest one can get is that the

period of formation was between the

first half of November 1859, when a

body of citizens was formed with the

express purpose of establishing a gar-

den, and Ist January, 1860 when the

first subscription became valid.

lat

y E

ae »® r

History and Functions of Botanic Gardens with

Special Reference to Singapore

By J. W. PURSEGLOVE

Director of the Botanic Gardens,

Singapore (1954-57)

MANY PEOPLE THINK of botanic gardens as public parks in which to

stroll amid pleasant surroundings and admire beautiful and rare

flowers. Others consider them a convenient exercising ground for

the baby or the dog, a good place for picnic. and, in Singapore,

somewhere to feed the monkeys.

Definition of Botanic Gardens

Any public park can provide these amenities. Why then go to

the expense of maintaining botanic gardens with their staff of

botanists, horticulturists and other specialists? The answer is that

botanic gardens, to merit the name, are gardens maintained for the

scientific study of the plants. As soon as this vital function is neg-

lected botanic gardens change to public parks. This has been the

fate of many botanic gardens throughout the tropics, while others

have been totally neglected or have disappeared. It is true that they

sometimes sell or give away planting material of fruit trees and

ornamental plants, but this function can be undertaken by agricul-

tural stations or commercial nurseries.

Botanic or Botanical

Some gardens such as Kew and Singapore are proud of the name

Botanic Gardens, all too frequently called Botanical Gardens by

the public, while others, especially in America, style themselves

Botanical Gardens. Why is this? Botanic is the older form of ad-

jective, but is now mostly superseded by botanical, except in the

names of institutions founded long ago. It is advisable to ascertain

which adjective is used by any particular gardens in order not to

give offence. Some such as the Royal Botanic Garden, Edinburgh,

do not use the plural of the noun.

The first Botanic Gardens

_ The oldest botanic gardens in existence today are those of Pisa

in Italy founded in 1543, closely followed by Padua. They were

founded by the Universities and were herb gardens in which the

Reprinted with minor amendations from Tropical Agriculture 34 (3) 1957

125

Gardens Bulletin, S.

future physician was taught the medicinal use of herbs and how to

recognize them. The Jardin des Plantes was founded by Louis XIII

in Paris for this same purpose in 1626, as was the Oxford Botanic

Gardens by the University in 1632, and the Chelsea Physic Garden

in 1673 by the Apothecaries Society.

Systematic study of plants

In the eighteenth century Linnaeus laid out a garden showing his

new system of classification and the plants were labelled in accord-

ance with his binomial nomenclature. Other gardens followed suit

and were designed to illustrate a purely scientific concept. Living

plants, as well as herbarium specimens, were brought from all over

the world for study in temperate countries. Botanists were often

attached to voyages of exploration to collect and bring back this

material. Many of the famous gardens of Europe, including the

Kew Royal Botanic Gardens, founded in 1759, carried out these

early taxonomic studies. (The first Botanic Gardens in the United

States were not founded until the nineteenth century.) The herbaria

and libraries which are so essential for this work were accumulated

and the systematic study of plants, sometimes on a local basis,

sometimes on a world basis, is still one of the most essential func-

tions of modern botanic gardens.

Founding and work in the early tropical Gardens

As far as I can ascertain the oldest botanic gardens in the tropics

are those of Pamplemousses in Mauritius which were established

by the French in 1735. Hart’s (1919) history of these gardens

makes fascinating reading and is typical of many tropical gardens

elsewhere. They started as a pleasure garden, although cassava

was first introduced as early as 1736. After periods of neglect their

principal function was to obtain, grow and distribute seeds of

economic plants, particularly of spices, hitherto only cultivated by

the Dutch in the Moluccas. Nutmegs were first introduced in 1765,

but died. Among the introductions in 1767 were pepper, cinnamom,

fruit trees and other plants yielding dyes and varnishes. Expeditions

in- 1770 and 1772 brought back nutmegs and cloves. The gardens

were laid out in a picturesque manner and planted with native,

African, Indian, Malayan and Australian plants. They suffered

from subsequent periods of neglect, but later played a prominent

part in the founding of the Mauritius sugar industry. They were

taken over by the newly-formed Agricultural Department in 1913.

The first British tropical botanic gardens are those of St. Vincent

founded in 1764 (Chipp, 1920). Bligh was commissioned to ob-

tain bread-fruit for these gardens from Tahiti and his expedition

126

Vol. XVII. (1958).

ended in the famous mutiny of the Bounty in 1787; he was more

successful in 1791 and brought back bread-fruit plants to the West

Indies. These gardens were abandoned for some 60 years later in

their history. Among the other early botanic gardens founded in

colonial territories are Jamaica in 1774, Calcutta in 1786, the first

of the three Penang Botanic Gardens in c.1796, Buitenzorg (now

re-named Bogor) in 1817; Trinidad in 1819, Peradeniya (Ceylon)

in 1821 and the first Singapore Botanic Gardens in 1822. Typical

of all these gardens is the founding of the first botanic gardens in

the north American continent when Charles III of Spain sent a

scientific expedition to Mexico in 1781, when “one of its objects —

was to establish a botanic garden in which instruction should be

given in botany and in which the native products of the country

should be cultivated and preserved” (Rickett, 1956).

In addition to the very important function of collecting, growing

and distributing indigenous and exotic economic crops of promise,

as well as seedlings of timber trees, and giving advice to the early

planting communities on methods of cultivation and the control of

pests and diseases, many of these botanic gardens laid a sound

foundation of botanical research in the tropics in the nineteenth

century. Many of the early directors were men of outstanding ability

and industry. First and foremost they were keen botanists and could

not help but be interested in the wild plants growing around them,

many of which were as yet undescribed. They made arduous collect-

ing trips into little-known country bringing back living plants for

trial in their gardens, as well as large numbers of herbarium speci-

mens. These were exchanged with other institutions throughout the

world, and as time went on local floras and monographs of parti-

cular groups were prepared. The world owes these great pioneers

a considerable debt.

By the second decade of the twentieth century most tropical terri-

tories had inaugurated independent agricultural and forestry de-

partments, and much of the work on crops and forest products was

handed over to them. Many of the botanic gardens were absorbed

into the new agricultural departments, often with disastrous results

to the research in botany which they had hitherto undertaken. The

role of botanic gardens in fostering agricultural development in

many tropical countries, usually achieved with very small staff and

slender financial resources, is now often forgotten.

The movement of crops in the tropics

I have already shown how botanic gardens played a prominent

role in the distribution of the principal economic crops grown today

in the tropics. It is worth examining the subject a little further. It

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Gardens Bulletin, S.

has always seemed strange to me that many of the world’s major

tropical plant products are produced largely in countries far re-

moved from their region of origin, e.g. South American rubber in

Malaya and Indonesia, South America cocoa in West Africa, South

American quinine in the East Indies, African coffee in Brazil,

cloves and nutmegs from the Moluccas in Zanzibar and Grenada

respectively, sugar, bananas and limes from South-East Asia in

the West Indies, and vanilla from Central America in Madagascar.

Obviously this cannot be attributed to any one particular reason

and many factors are involved, including economics, available land

and labour supply, technical skill in processing, suitability of the

crop for plantation or peasant agriculture, etc. Nevertheless, one

would have expected that a crop was more suited ecologically to its

country of origin than to its new home. I suspect that one of the

major reasons is that when a new crop has been introduced with-

out many of its normal pests and diseases, it has more chance of

flourishing and giving high yields. Many of the crops are introduced

as seeds, which limits the number pests and diseases which they

can carry with them, and this is further enhanced by the plant

quarantine regulations now enforced in many countries. One can

only contemplate what would happen to the Malayan rubber in-

dustry if the South American Leaf Blight (Dothidella ulei) were

accidently introduced.

In this digression I have limited myself to economic crops, but

further examples can well be given from food crops, too. Many of

the staple crops now grown in parts of tropical Africa are of New

World origin, e.g. cassava, sweet potatoes and maize.

The early Penang Botanic Gardens

Ridley (1910a) gives the date of the founding of the first Botanic

Gardens, Penang as 1800, although in the same article he states

that, soon after the first settlement in Penang in 1786, the East

India Company decided to start spice gardens with a view to

breaking down the Dutch monopoly of the spice cultivation and

trade. A botanist, Christopher Smith, was appointed in 1794 and

in 1796 was sent to the Moluccas to collect living plants of nut-

megs and cloves. Ridley then adds that by 1800 the Gardens con-

tained 1,300 plants and that they were greatly enlarged in that

year, when 15,000 clove and 500 nutmeg trees arrived from

Amboina. It thus appears that the first Gardens were begun about

1796 and not in 1800 as is usually stated. Ridley goes on to say

that Smith sent in all 71,266 nutmeg plants, 55,264 cloves and

large quantities of canary nut, (Canarium commune) and Kabong

palm, (Arenga saccharifera).

128

Sh 4

Vol. XVII. (1958).

This transporting of very large quantities of plants in the tropics

was to continue throughout the nineteenth century. Nor was the

movement confined to economic crops, as plants of horticultural

merit were freely distributed between botanic gardens, while scien-

tific institutions, wealthy patrons, seedsmen and nurseries sent col-

lectors to the far corners of the earth to bring back novelties for

their hothouses and for sale. The scale on which the exchanges

took place is hardly conceivable today and the expense, difficulty

and length of time spent transporting them must have been very

considerable.

The first Penang Botanic Gardens were sold in 1806. Raffles

founded the second Gardens in Penang in 1820. Ridley states that

these existed until 1834, “when Governor Murchison, who took no

interest in gardens or agriculture, sold them for 1,250 rupees, be-

cause his wife could not get enough vegetables from them to

diminish her cook’s bill.”

HISTORY OF THE SINGAPORE BOTANIC GARDENS

The Singapore Botanic Gardens are the last gardens in the

British tropics which function as a separate and self-contained de-

partment and have never been under the control of any Agricul-

tural Department, Municipality or University. I now propose to

confine my attention to these Gardens and describe some of the

work carried out there during their long history.

The first Singapore Botanic Gardens

Shortly after the founding of Singapore in 1819, the founder,

Sir Stamford Raffles started the first Botanic Gardens on the Gov-

ernment Hill at Fort Canning in 1822. Nathaniel Wallich, who was

visiting Singapore from the Calcutta Botanic Gardens, advised on

this project. Raffles introduced nutmegs, cloves and cocoa. These

gardens were closed in 1829 on instructions from India to retrench

expenditure. They were re-opened later by an Agri-Horticultural

Society in 1836, who covered the cost from the sale of nutmegs

and cloves growing there. They were finally abandoned about 1846.

The principal crops of Singapore at that time were nutmegs and

gambier; the former failed because of pests and diseases; the latter

was too exhausting to the soil and required large quantities of

charcoal for its preparation.

In a recent popular article in a Singapore paper a writer (H.T.S.,

1956) says, “When critics carp at British colonialism, they should

stop for a moment and ponder on the old circumstance that one of

£29

Gardens Bulletin, S.

the first matters which occupied the earnest attention of the foun-

ders of Singapore was to find a site for a botanical (sic) gardens.

No great plan for exploiting the local people was afoot here.”

The Second Singapore Botanic Gardens

The First Phase: 1859-1875

In 1859 a new Agri-Horticultural Society was formed in Singa-

pore, the third in the history of the island. The Society was given

a grant of some 60 acres of land in Tanglin by the Government on

which to found a garden. Convict labour was also provided. Mem-

bers of the society paid an entrance fee and a monthly subscription,

while there were second-class subscribers who might enjoy the use

of the gardens by paying a monthly fee. They also raised money

by public entertainments. The southern end of the Gardens was

probably abandoned gambier land, a haunt of tigers, while the

north was still virgin tropical evergreen rain forest, 11 acres of

which are still preserved in their original form, a most valuable asset

in the centre of a great city. The first object of the committee was

to create a pleasure garden with roads, terraces and a band stand,

where military bands performed fortnightly, the gardens providing

an alternative evening drive to the Esplanade for ‘the quality’.

Laurence Niven, who supervised a privately-owned adjoining

nutmeg plantation, was employed part-time as Superintendent. He

had a fine eye for design and laid out the Gardens much as they

are today. As early as 1861, flower shows were organised in the

hope of encouraging the local cultivation of vegetables, fruits and

flowers. In 1866 the Society purchased a further 25 acres of land

and built the present Director’s house, as well as constructing the

Gardens’ lake. This depleted the Society’s funds and in 1869 they

approached the Government for a further grant, the first being $50

towards Niven’s salary in 1866. The second grant was agreed to on

condition that the Society should exhibit in the gardens living eco-

nomic plants. At the same time a small zoo started. (There has

been a menagerie at the Jardins des Plantes in Paris since 1794.)

By 1874 the Society was in debt and could carry on no longer.

They asked the Government to take over their Gardens and their

debts and to maintain the Gardens for the benefit of the people of

Singapore. Government agreed to do this and in 1875 the manage-

ment committee asked Sir Joseph Hooker, the Director of the

Royal Botanic Gardens, Kew, to recommend a suitably trained

candidate for the post of Superintendent, who “‘was to be a prac-

tical, as well as systematic botanist, and to travel in the Malay

Peninsula not a little for the purpose of investigating its vegeta-

tion”. (Burkill, 1918).

130

Vol. XVII. (1958).

Thus the second Singapore Botanic Gardens, which celebrates its

centenary in 1959, originating as a little more than a public park,

began the second phase of its history and was changed into a

working Botanic Garden.

The Second Phase: 1875-1888

Sir Joseph Hooker sent out James Murton, a Kew trained horti-

culturist. He was energetic and enthusiastic, but unfortunately was

very young and little suited for control. He added greatly to the

collection of plants growing in the Gardens, both economic and

horticultural, from the principal Botanic Gardens in the East, as

well as Kew. Kew sent the first live Para rubber seedlings in 1877

(see below).

Under the Raffles Society Ordinance of 1878 the Gardens were

legally taken over by the Government of the Colony and vested

in Her Majesty the Queen, her Heirs and Successors. In 1879 the

Government allocated a further 102 acres of land, now occupied

by the University of Malaya, for the cultivation of economic crops,

particularly gutta-percha and rubber-producing trees. Also growing

in these Economic Gardens at this time were Cinchona, arabica

and liberica coffee, Eucalypti, Ipecacuanha, tea, maize, sugar cane,

cola nuts, mahogany, etc. Murton added greatly to the Gardens’

menagerie. He collected plants in Singapore and in the Peninsula,

but his duplicates in Singapore Jater disappeared, when he was

dismissed in 1880. He died in Bangkok the following year and

Burkill (1918) says “his death perhaps hastened by his having

burnt the candle at both ends.”

Murton was succeeded by Nathaniel Cantley, who was also a

Kew trained horticulturist. He had been Assistant Superintendent

of the Mauritius Botanic Gardens and was an older man, a great

lover of order. He did much planting and named and labelled all the

plants in the Gardens. He surveyed the forests of the Straits Settle-

ments and was responsible for gazetting the first forest reserves. He

planted large numbers of Malayan and introduced timber trees in

the Economic Gardens, and his arboretum was arranged in a bota-

nical sequence on the system of Bentham and Hooker. Cantley was

a keen botanist; he collected local plants and started the herbarium.

He also experimented with European vegetables. The grounds of

Government House Domain were placed under the Botanic Gar-

dens and still remain so today. The Gardens also helped in horti-

cultural work in the city and much of the tree planting in Singapore,

which greatly adds to its attraction, was supervised by the Gardens’

staff. Cantley, who had never been a very fit man, went on sick

leave in December 1887 and died the following year.

131

Gardens Bulletin, S.

The Third Phase: 1888-1912

With the appointment of Henry Nicholas Ridley as Director in

1888 the Gardens entered upon the third stage in their history

and a most productive period it was. “Ridley’s capability as a re-

search worker and his power to demonstrate by his publications the

practical and beneficial effects of photography on plant science in

all its aspects . . . succeeded in a surprisingly short time in pushing

the Singapore Gardens to the fore as a centre of Malaysian phyto-

graphy” (de Wit, 1949). Ridley was a tireless worker and his

phenomenal output of work in the natural sciences and his appli-

cation of this to economic problems is quite remarkable.

He has been rightly named “the father of Malaya’s rubber in-

dustry”. His early experiments on the tapping and preparation of

rubber and his constant advocacy of it as a plantation crop, often

in the face of opposition, led to the foundation of the present in-

dustry upon which so large a share of the wealth of Malaya and the

eastern tropics has rested (Purseglove, 1955 a). Nor was his work

on economic crops confined to rubber; he carried out experiments

with most of the actual and potential crops known. As early as

1889 he reported that cocao would probably be profitable on the

better Malayan soils, while in 1907 he directed attention to the

possibility of oil palm as a plantation crop. The local planting com-

munity was largely dependent upon the Economic Gardens for

planting material and seeds of economic crops were distributed to all

parts of the tropics. In the case of rubber alone, over seven million

seeds were distributed from the Singapore Botanic Gardens, as well

as many seedlings. The Gardens obtained a considerable income

from the sale of these, as well as the rubber produced, thus allowing

them to be maintained without any increase in Government vote.

Visitors from all over the world visited the Economic Gardens,

while Ridley and his three assistants in Singapore, Malacca and

Penang, were the only trained men whom the planting community

could approach for technical advice and help. It was Ridley who

was responsible for and then administered, the Coconut Trees Pre-

servation Ordinance of 1890 for the control of the rhinoceros

beetle. In addition to being Director of the Botanic Gardens, Ridley

was also Director of Forests, Straits Settlements and was respon-

sible for gazetting additional forest reserves. He produced the first

published accounts of Malayan timbers, vegetables, fruits, fibres,

dyes and drug plants, while his book on Spices (1912) is the

standard work.

Ridley’s work as a field botanist is equally remarkable. He tra-

velled far and wide in the Malay Peninsula, often in remote regions,

132

Vol. XVII. (1958).

as well as visiting Cocos and Christmas Islands, Borneo, Java and

Sumatra. On all these expeditions he studied the vegetation and

brought back vast numbers of herbarium specimens, as well as

many living plants for trial in the Gardens. In all he collected many

thousands of specimens, of which a large number were unknown to

science. These specimens are now in our herbarium, but duplicates

were sent to Kew and other botanical institutions. He named and

described for the first time a few thousand plant species and a sur-

prising number are still only known from his original gatherings.

Specimens were sent to experts on certain groups elsewhere and

many new species were named in his honour. Ridley realised very

early in his career that the flora of a country cannot be studied

in vacuo and that it is essential to take into account plants from

neighbouring territories as well. After his retirement he wrote his

five-volume Flora of the Malay Peninsula (1922-25).

Nor was Ridley’s attention confined to plants, of which he col-

lected in all groups. He belongs to that great pre-specialised age of

scientific natural history and he collected and studied many ani-

mals, as well as distributing specimens and writing about them.

Ridley’s work as a naturalist has been more fully described else-

where (Purseglove, 1955 b). He took a keen interest in the Gardens’

menagerie, which was disbanded in 1904—5, much to Ridley’s

obvious regret.

In addition to publishing in many scientific journals in Singapore

and overseas, Ridley founded the Agricultural Bulletin of the Malay

Peninsula in 1891, so that he could supply the planting community

and others with information resulting from his many investigations.

In all he published over 500 papers, articles and books, totalling

over 10,000 printed pages, so that the work of the Singapore

‘Botanic Gardens became widely known throughout the world.

Details of his publications and other work will be found in Gardens’

Bulletin, Vol. IX, 1935, which was dedicated to him on his 80th

birthday. Attention should be drawn, however, to his encyclopaedic

volume, The Dispersal of Plants throughout the World, published

in 1930, which still contains much of value for the botanist and the

agriculturist.

Few people have accomplished so much in twenty-three years of

tropical service as Ridley did—he retired in 1912—nor have com-

pleted their work after retirement as he had done. He was elected

a Fellow of the Royal Society in 1907 and, apart from the honour

of C.M.G. in 1911, he received comparatively little public recog-

nition, until his hundredth birthday, which was celebrated on the

10th December, 1955. On this occasion he wrote “It is a great

delight to me to have lived to see the Gardens, the best tropical

133

Gardens Bulletin, S.

Gardens in the world,” and that “my hundredth birthday was one

of the most enjoyable days of my life”. It is with regret that I record

the death of Mr. Ridley on the 24th October, 1956.

The founding of Malaya’s rubber industry

Accounts of the transfer of Para rubber from Brazil to the east-

ern tropics have often been given, but frequently they are inaccu-

rate in detail. As the Singapore Botanic Gardens played such an

important part in this introduction, it is worthwhile recording briefly

the facts at this juncture.

Sir Clement Markham of the India Office, who had been res-

ponsible for bringing Cinchona from South America to the east in

1865, pointed out in 1870 that there was a danger of exhausting the

Indo-Malaysian supply of wild rubber and recommended that

plantations should be put down in the east. James Collins, who was

Government Economic Botanist and Librarian in Singapore, was

asked to collect all the available information about rubber-produc-

ing plants and his report was published in 1872. Through Mark-

ham’s efforts and those of Sir Joseph Hooker, the Director of

Royal Botanic Gardens, Kew, a few seeds were collected and sent

to Kew in 1873, when a dozen plants were raised. Six of these were

sent to the Calcutta Botanic Gardens and were later propagated

and sent to Sikkim, where they were a failure, the climate being

unsuitable. A consignment of seeds was sent to India in 1875 and

failed to germinate.

Markham then arranged for Robert Cross to collect rubber seeds

in South America, while Hooker obtained the services of H. A.

Wickham, later Sir Henry, who was then on the Amazon. Wickham

collected 70,000 seeds. He did not smuggle the seeds out as is often

suggested; it is now known that they were brought out with the

goodwill and co-operation of the Brazilian Government. Wickham

chartered a ship and the seeds arrived at Kew on 15th June, 1876,

where they were planted immediately. Less than 4 per cent germi-

nated and some 2,800 seedlings were raised at Kew. Many large

tropical oily seeds retain their germination powers for short periods

only and it was Ridley who later discovered that they could easily

be transported in damp powdered charcoal. On 9th August, 1876

Kew sent 1,919 plants to Ceylon in 38 Wardian cases. Two days

later 5O plants were sent direct from Kew to Singapore. Thiselton-

Dyer (1878), and later quoted by Ridley (1910b), states that

“owing to the delay in payment of freight (in Singapore) these

plants all perished”. Burkill (1935) says that Murton reported

that “five plants were alive when the cases were received at the

Gardens”, although they appeared to have perished subsequently.

134

Vol. XVII. (1958).

On 10th June, 1877 Kew sent a further 22 plants direct to Singa-

pore and these thrived. They were not sent from Ceylon as is often

published.

In the meantime Cross returned to England with a consignment

of plants in November, 1876. Burkill (1935) states that “not one

of Cross’s plants collected in Para is known to have survived the

seedling stage or to have reached the east’. This was good fortune

in disguise as the east was stocked from Wickham’s seeds, which

were collected in the central valley of the Amazon, where the trees

are now known to have been of better quality than those nearer the

mouth. Murton planted eleven of the rubber plants near our pre-

sent nursery and then Fox in 1879 transferred them to swampy

ground in the new Economic Gardens, as it was thought that this

was the natural habitat of the tree. He took nine of his seedlings

to Perak in October 1877 and these were planted at the Residency

at Kuala Kangsar. Most of the rubber in Malaya has come from this

original introduction, although some seeds were received from

‘Ceylon at a later date.

The Singapore trees first fruited in 1881. When Ridley arrived

in 1888, there were 9 trees of the original introduction, some 50

trees 2—4 years old and over 1,000 young seedlings. He quickly

planted more. The subsequent exploitation and establishment of

rubber as a plantation crop in Malaya was due almost entirely to

Ridley. Within four months of his arrival in Singapore he drew the

Government’s attention to this possibility. With great perseverance,

little encouragement or help and slender financial resources, he set

to work. He has since recalled how he was “carpeted by the

‘Governor, Sir Frank Swettenham, who admonished him to waste

less time on an uneconomic product such as rubber.” He experi-

mented with various rubber-producing trees and found that Hevea

was superior to all others. Starting with a mallet and chisel, and

later using a modified farrier’s knife, he developed the tapping tech-

nique which has been but little changed since. He discovered

wound response, which he terms “calling the rubber’, and he also

demonstrated that the bark regenerated and could be tapped again.

He exhibited coagulated rubber in Singapore in 1890 and the fol-

lowing year sent specimens to England which were well reported on.

Having shown that rubber was a commercial proposition, Ridley

then tried to persuade the reluctant planters to give the new crop a

trial, but at first met with little success, as coffee was considered

to be the only important crop in Malaya. For his efforts he was

known as “Mad Ridley”. In 1893 he distributed the plants and

seeds to Residents and District Officers in the Federated Malay

‘States to plant near their houses. It was fortunate for Malaya that

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when rubber planting began in earnest—coffee prices had slumped

and the trees had ben attacked by diseases and pests—that there

was at the Singapore Botanic Gardens a good supply of planting

material and Ridley, the expert, with the latest ‘know-how’ to advise

and to help planters. Ridley stayed long enough in Singapore to

see the great rubber boom in 1910 and for Malaya to reach a pre-

dominant position in world rubber production, which it has never

lost.

The Fourth Phase: 1912-41.

During Ridley’s term of office the Gardens established for them-

selves an international reputation, which was adequately main-

tained during the next stage in its history. With the forming of the

new Agricultural Department in 1904, much of the work on the

economic crops passed from the Gardens and the Economic Gar-

dens were finally closed in 1925. Gradually during the period more

staff were recruited, especially botanists, so that the work on taxo-

nomy could be extended. Some work was done on the lower plants,

particularly the Fungi. In the horticultural field the hybridising of

orchid and other cultigens was undertaken.

I. H. Burkill, who had been on the staff of the Calcutta Botanic

Gardens and had collaborated with Watt on the study of the eco-

nomic products of India, was appointed Director in Singapore in

1912. During World War I the Gardens were understaffed and

there was little time for expeditions and collecting. Anderson’s

Catalogue of Plants growing in the Gardens was published in 1912

and lists 1739 species, which included 276 orchids, 261 ferns and

245 palms. T. F. Chipp was appointed Assistant Director in 1915,

a new post having been created, but he did not take up his duties

until 1919. He was transferred to West Africa at the end of 1920,

in which year he published his List of the Fungi of the Malay

Peninsula. Until the Economic Gardens were taken over by Raffles

College, now the University of Malaya, in 1924—25, they continued

to play an important role in the distribution of economic plants.

In 1916 two hundred species were listed as available for sale at the

Gardens. The loss of the Economic Gardens was a grievous blow,

as the land available for planting was more than halved.

Burkill continued his interest in medicinal and economic plants

generally and collected much information on these in Malaya. He

continued this work for 10 years after retirement and published in

1935 his large two-volume Dictionary of the Economic Products of

the Malay Peninsula, a work of great scholarship, “a masterpiece

of synthesis”, and a model for all works of this type. It is of great

value to the botanist and the agriculturist, not only in Malaya, but

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Vol. XVII. (1958).

throughout the tropics. Unfortunately it is now out of print, but I

hope that it may be reprinted soon. Burkill also made a detailed

study of the genus Dioscorea (yams), a genus on which he is still

working.

The Gardens Committee ceased to have their own banking ac-

count in 1919 and their balance was paid into the Treasury. There-

after the Government increased the Gardens’ vote, which was

provided under several separate heads, where formerly a lump sum

had been given.

R. E. Holttum was appointed Assistant Director in 1922. He

was promoted Director in 1925 and continued in this post until

1949, when he became the first Professor of Botany in the Univer-

sity of Malaya. Holttum’s principal interest were ferns and mono-

cotyledons and he added greatly to our knowledge of these plants,

describing many new species. He showed that wild orchids may

hybridise and produce ‘species’ which had hitherto never been sus-

pected to be of hybrid origin (de Wit, 1949). He began work in

1928 on the production of spectacular new free flowering hybrids

of cultivated orchids and this work is still being continued at the

Gardens.

C. X. Furtado joined the staff in 1923 and is still Botanist here.

During this period he has become an acknowledged authority of the

International Rules of Botanical Nomenclature and on Malaysian

palms and aroids.

M. R. Henderson was appointed Curator of Herbarium in 1924

and held this post until he was appointed Assistant Director in 1946

and Director in 1949. He worked on Malayan flowering plants

generally, his special interest being limestone plants and the family

Myrtaceae. .

E. J. H. Corner arrived in Singapore in 1929 as Assistant Direc-

tor, the post which he held until 1946. He specialised in Fungi and

trees and carried out research on the morphological and ecological

aspects of the vegetation. Corner trained berok or pig-tailed mon-

keys (Macacus nemestrina) to collect botanical specimens, thus

partly solving the problem of obtaining specimens from tall trees

and climbers otherwise inaccessible. Corner states that one of these

monkeys knew the meaning of 18 words of Malay and was able to

find at the tree tops fruits and flowers which had been shown to

him on the ground. It has been said that these botanical monkeys

were “the first apes to enter Government Service”. Corner’s two-

volume Wayside Trees of Malaya was published in 1940, a work

full of out-of-the-way information and original observations and

is one of the most fascinating books ever produced on the trees of

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any country. Corner, now Lecturer at Cambridge University, is

continuing his study of the genus Ficus in Malaysia.

Mention must also be made of C. E. Carr, a Malayan planter,

who worked in conjunction with the Gardens, and added greatly to

our knowledge of Malaysian orchids. He died in New Guinea in

1936 when returning from a collecting expedition.

From 1923 to World War II the senior staff of the Gardens con-

sisted of Director, Assistant Director, Curator of the Herbarium

and Assistant Botanist, all actively engaged on botanical research

and three horticulturists as Curators, two in Singapore and one in

Penang. In Singapore one Curator supervised the horticultural work

at the Gardens and the other looked after Government House

Domain and the outside commitments in the City.

The Gardens’ collection of living plants and the herbarium were

added to greatly during this period.

The Fifth Phase: The Japanese Interregnum: 1942—46

During the early years of the World War II before the Japanese

attacked Malaya at the end of 1941, most of the senior Garden

staff were seconded for at least part of their time to the Depart-

ment of Food Control and Information. The Gardens maintained

demonstration plots of vegetables to encourage local production.

J.C. Nauen and G. H. Addison, Curators at Penang and Singapore,

were mobilised with the local defence force and were later taken

prisoners and sent to work on the Siam-Burma railway, where

Nauen lost his life. Holttum, Corner and Furtado remained in

Singapore during the occupation and were permitted to carry on

with their botanical research. Immediately following the capture of

Singapore Island, Professor Tanakadate, of Tohoku University,

took over the control of the Gardens, and took care that they were

not despoiled by the Japanese military forces. Holttum remained in

executive charge until the appointment of Professor Kwan Koriba,

of Kyoto University, as Director of Gardens in December 1942.*

During his stay in Singapore Koriba undertook research on the

growth-behaviour of some Malayan trees. It was particularly fortu-

nate for the Singapore Botanic Gardens that the Japanese scientific

officers and the British staff who remained were able to preserve

the herbarium and the library as part of the cultural heritage of

Malaya and that no loss whatsoever was suffered in them, except

for a few specimens which were on loan in Germany. Losses did

occur, however, among the living collection and garden work dete-

riorated generally, as more than half of the out-door staff (49 men)

. * An orbituary of Professor Kwan Koriba is published in this Bulletin.

Editor.

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Vol. XVII. (1958).

were sent to work on the Siam-Burma Railway, where 22 of them

lost their lives. There was comparatively little damage in the Gar-

dens, although the Director’s house received a direct hit from a

shell, but, even so, the damage was not serious.

From September 1945 to March 1946 the Gardens came under

the British Military Administration.

The Sixth Phase: 1946-56

After the war the staff was gradually re-assembled, but did not

reach its full compliment until the end of 1954, M. R. Henderson

was appointed Assistant Director and later Director (1949-54).

J. Sinclair arrived in 1948 as Curator of the Herbarium, the title

later being changed to Keeper. J. W. Ewart, Horticultural Curator,

served as Agricultural Officer, Singapore, as well as carrying out

his duties at the Gardens. Henderson retired in 1954 and J. W.

Purseglove was appointed Director, while H. M. Burkill (son of

I. H.), was appointed Assistant Director in December of that year.

Gradually during this period the Gardens were brought back to

their former good condition, the living collection of plants was in-

creased and collecting in Malaya and Borneo was resumed. The

first two volumes by R. E. Holttum of the Revised Flora of Malaya

were published. Burkill has begun a study of Malayan seaweeds.

On the 10th December, 1955, the Gardens celebrated the hun-

dredth birthday anniversary of H. N. Ridley, with an exhibition of

Ridleyana and current work, while the Gardens were floodlit for

one week. A special brochure for private circulation was produced

for the occasion (Purseglove, 1955c).

Early in 1955 a scheme was drawn up in conjunction with the

University of Malaya with the aim of attracting research workers

from Britain and elsewhere to Singapore for one or two years to

carry out the botanical research in their specialised fields. By this

means the output of research would be increased and knowledge

would be obtained which would be of great value to the tropics as

a whole. Furthermore, workers from temperate countries would

become familiar with tropical botany and their interest in our

problems would be stimulated. In 1955 P. B. Tomlinson of Leeds

University was awarded an Agricultural Research Council Schoiar-

ship and spent one year here studying the anatomy of palms. T. C.

Whitmore, a Ph.D. student of Cambridge University, arrived in

December 1956 to spend a year working on the anatomy of Dipte-

rocarp barks in relation to taxonomy. He is receiving a Colonial

Development and Welfare grant. It is hoped that more research

workers may come in the future.

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[Jn order to bring this paper up-to-date as a history of the first

hundred years of the Botanic Gardens, the footnote below

headed: The Seventh Phase: 1957 and on, has been added to the

original paper in concurrence with the author.—EDITorR.]

Waterfall Garden, Penang

The third Botanic Gardens in Penang were founded by Charles

Curtis in 1884 as “a nursery for the planting of colony products”,

and were under the charge of the Gardens and Forest Departments

Straits Settlements, with its headquarters at the Singapore Botanic

Gardens. Curtis, who had been employed as Messrs. Veitch’s plant

collector in Mauritius, Madagascar and Malaysia, was appointed

Superintendent and remained there until his retirement in 1903.

Holttum (1934) records that Curtis “was an enthusiastic plant-

lover and a born gardener, and the development of the Waterfall

Garden is a monument of his energy and skill”. Curtis also made

large collections of herbarium specimens in Northern Malaya. The

Penang herbarium was later incorporated with that at Singapore.

The Seventh Phase: 1957 and on.

The penultimate constitutional step in the introduction of full internal

autonomy of Singapore (effccted in May 1959) was operative from April

1955. As befits a country in charge of its own affairs, the Singapore Govern-

ment adopted a policy of “malayanisation” of the public service from Jan-

uary list, 1957. J. W. Ewart retired in March 1957 in accordance with this

policy, and A. G. Alphonso, who had returned in 1956 from a two year

course of training in horticulture at the Royal Botanic Gardens, Kew,

England, was promoted Curator. J. W. Purseglove retired voluntarily in

March 1957, and H. M. Burkill became Director. G. H. Addison retired in

February 1959. Chew Wee Lek was appointed Botanist in 1956 and went to

‘Cambridge, England, in 1957 for three years to study for a higher degree.

Lam Hin Cheng was appointed Horticultural Assistant in July 1957 and

went to the Royal Botanic Gardens, Kew, for a two year course in horti-

culture. The new post of Librarian was eventually filled in April 1958, and

Tan Kim Ho went to Melbourne, Australia, in 1959 for training in library

management on an Australian Government award under the Colombo Aid

Programme. Attempts to fill the post of Assistant Director were unavailing,

and finally Miss Chang Kiaw Lan was appointed Botanist (vice Assistant

Director) to take up a study of mycology. In 1958 Bajuri bin Sappan,

Laboratory Assistant, was sent to various institutions in England for one

year to study plant breeding techniques with a view to furthering the De-

partment’s plant breeding work.

The last three years, 1957 to this centenary year of 1959 have been a

period of transition characterised by an acute shortage of senior personnel

through loss of qualified staff and temporary loss of the services of officers

‘sent overseas for training. The most severe effects of this will be over by the

end oi 1959, when only one officer remains overseas, and he will be back

in 1960. Thus will close a century of expatriate know-how in the senior

botanical and horticultural posts, and there will open the second century of

the Gardens’ history, every bit as promising in the comity of international

botany as the first has been successful, with the majority of the senior posts

held by Malayan personnel. H.M.B.

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In 1910 it was decided to convert the Gardens into a reservoir,

but fortunately this did not materialise. With the formation of the

Malayan Union in 1945, later renamed the Federation of Malaya,

and the separation of Singapore as an individual colony, the con-

trol of the Waterfall Gardens passed from the Singapore Botanic

Gardens to the Federation Agricultural Department. This was a

great loss to the Singapore Gardens as the climate of Penang is

more seasonal than that of Singapore and was therefore better

suited for the growth of certain plants, which flower and fruit there,

but will not do so in Singapore. Furthermore, the Penang Gardens

lost the stimulus of direct contact with the parent institution where

botanical research is carried out.

THE WORK OF BOTANIC GARDENS

The principal work carried out by botanic gardens can be divided

under three broad heads, namely, botanical, horticultural and edu-

cational, although inevitably there is some overlapping between

them. In addition to these functions most botanic gardens are

opened to the public and thus provide the facilities of a public

park. For this purpose they should be laid out as attractively as it

is possible to make them, without jeopardising the work of the

scientific study of plants. The Singapore Botanic Gardens are

opened to the public from 6 a.m. until 7 p.m. daily; no charge is

made and the popularity of the Gardens can be gauged from the

very large number of visitors to them. Most tourists visiting Singa-

pore also come to see the Gardens.

I now propose to give a more detailed account of the work under

the various heads given above, illustrating this from the Singapore

Botanic Gardens.

BOTANICAL WORK

The living collection

A botanic gardens should have as large a collection of living

plants as it is possible to grow, without sacrificing an attractive lay-

out. In fact a Gardens should be a great museum of living plants.

All the plants should be correctly labelled with their scientific name

and family, together with the country of origin. If there is a common

English or vernacular name, this may be added, but this is often

difficult when various languages are spoken as in Singapore. Care

should be taken to keep the names up-to-date according to the

latest nomenclature changes. As many different groups of plants as

possible should be represented. Inevitably, certain gardens special-

ise in certain groups, particularly those on which specialists have

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been working. Thus in Singapore our collection of palms, orchids

and ferns are particularly good. Inevitably many plants of bota-

nical interest, but of little value horticulturally, must be included in

a botanic gardens, and they are not always as colourful as the

layman expects. It is desirable that closely related species or even

groups, should be planted near each other, which is very useful for

comparative purpose, but this should not be carried to the extreme,

as it results in a deterioration in the lay-out. Herbs and woody

plants of varying stature, although belonging to the same family,

are often difficult to grow together and to maintain and are un-

sightly when so planted. A lot of plants can be grown in pots. For

this a burnt earth-compost mixture has proved to be the most suit-

able for our conditions for many plants. This method is described

by Holttum (1953 a).

The Singapore Botanic Gardens have some 3,000 perennial

species growing in them at present and this does not include hy-

brids. Even so, there is a limit to the number of plants that can be

grown in our eighty-five acres and there is little room for further

planting, particularly of large trees. The work of labelling such a

large collection presents difficulties and label printers are constantly

at work repainting old labels and preparing new ones. The majority

of our labels are made of wood or metal, 6” « 4”, with an iron

prong 8” long for sticking in the ground. They are painted black

with the plant name clearly printed in white. For pots and small

species printed aluminium labels are used. These are also attached

to trees in the Gardens’ jungle, but unfortunately they are often

removed by monkeys.

The Herbarium

In addition to the living collection, a botanic garden should have

a hortus siccus—the herbarium, or at least one reasonably acces-

sible to it. In this, dried mounted specimens are preserved. The

Singapore Herbarium is situated in the Gardens and has over

400,000 sheets. The specimens are mainly confined to plants of the

Malaysian region, which extends from Sumatra and Malaya to New

Guinea, but there are a few from surrounding countries. Our oldest

specimens are over 150 years old and are still in a good state of

preservation, despite the fact that they are housed in wooden cup-

boards and the building is not air-conditioned, although the latter

is preferred if possible. The collection is arranged according to the

Bentham and Hooker system, and, within the species folders, ac-

cording to the various Malay states, Malay islands and exotics.

There is also a smaller and separate Gardens’ Herbarium of culti-

vated plants.

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Vol. XVII. (1958).

In addition to being in constant use by the botanists, both at the

Gardens and the University, the specimens are sent on loan to

botanical institutions elsewhere for study. In 1956 a total of 4,525

sheets were sent on loan, mainly to Leiden, Bogor and Kew, to

authors of the Flora Malesiana, while 4,903 sheets were sent on

loan in 1955. The value of the specimens is enhanced considerable

when the sheets are returned correctly annotated. Inevitably some

mixing occurs in folders until the genus has been currently revised,

but this is not peculiar to Singapore.

There is also a good collection of spirit material, particularly of

orchids, and this is invaluable for detailed study.

Collecting, naming and distribution of specimens

Both the living and dead collections of plants at the Botanic

Gardens are added to constantly by collection, donations and ex-

change. Plants do not recognize political boundaries and in studying

the plants of a country such as Malaya, it is essential to study the

plants of surrounding countries, if one is to understand the varia-

tion and distribution of the local species. Thus botanists from the

Singapore Botanic Gardens, although the bulk of their collection

has been made in Malaya, have also collected extensively in

Borneo and to a lesser extent elsewhere, where the flora shows

regional affinities. We also try to collect as many duplicates as pos-

sible of each gathering, so that these may be distributed to other

institutions elsewhere. In these uncertain days spacial separation

of duplicates is very desirable, so that if one or more sets are des-

troyed, as happened in Berlin and the Philippines during the war,

others are available elsewhere. 5,397 duplicates were distributed

from Singapore in 1956, many of which were plants collected by

the writer in Sarawak. 6,768 duplicates were received, of which a

large portion came from the Forestry Departments of Sarawak,

Brunei and British North Borneo. Many of these were unnamed

and have since been determined by the staff. Private collectors are

also encouraged to send in specimens.

The importance of adequate field data on all specimens cannot

be over-stressed. Each specimen should be given a serial number

and notes are made of the place of collection, habitat, altitude,

relative abundance, as well as information on habit, colour of

flowers, etc., which cannot be ascertained from the dried specimens.

Special herbarium labels are provided for listing this data. In the

past this type of information, which can be of very great value to

the taxonomist, was frequently neglected. Sterile material is often

of doubtful value.

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With the aid of a good modern flora, a botanist should be able to

determine most of his specimens in the field—but unfortunately

how many tropical countries have complete up-to-date floras! Even

when they are available, certainty can only be reached by com-

parison with authentic named material. Agricultural and forestry

departments, teachers, students, planters and others from Malaya,

Borneo and elsewhere send us specimens for determination. Small

plant fragments are sometimes sent for identification in connection

with police investigations, medical enquiries, etc. We receive fre-

quent requests for information on certain plants. Thus we provide

an important service, which cannot be obtained elsewhere in

Malaya.

Taxonomic studies

A very important function of any botanic gardens is research in

systematic botany. This critical and highly specialised work in-

volves detailed revisions of genera and families and is an essential

preliminary to the publication of any flora. Previously named spe-

cies are critically examined, questions of synonomy are finalised,

and new species are described. Furthermore such work can never

be completely perfected, as a critical study of the great amount of

material collected during the past 40 years reveals large numbers

of undescribed species, as well as new information on distribution

and variability, while future collecting will yield yet more. It is

necessary that the names and descriptions of Malayan plants should

be brought into line with those of neighbouring countries. | am

convinced that a study of Malaysian plants as a whole is an essen-

tial preliminary in the preparation of the local flora. Consequently,

the staff of the Singapore Botanic Gardens is now collaborating in

the production of the Flora Malesiana, edited by C. G. G. J. van

Steenis of Leiden University and financed by the Indonesian Gov-

ernment.* This major work will occupy many botanists from many

countries for many years.

So much new material and new knowledge has accumulated

since the publication of Ridley’s Flora of the Malay Peninsula in

1922-25, that it is now out-of-date and a Revised Flora of Malaya

is being prepared. Two volumes have been published so far, both

by R. E. Holttum, namely, Orchids of Malaya (1953) and Ferns

of Malaya (1955). This work is designed to be of use to the field

* From 1st January, 1958 funds from this source have ceased. The Flora

Malesiana Foundation is currently sponsored by the Netherlands Organisa-

tion for Pure Scientific Research “Z.W.O.” with contributions from British

Government and Commonwealth sources, including Singapore.

Editor.

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naturalist and the gardener, as well as the specialist, and includes

introduced, as well as indigenous plants.

The great complexity of the Malaysian flora, which is one of the

richest in the world, makes the task of the taxonomist a very formid-

able one. Some idea of this can be obtained if one realises that the

island of Singapore, with an area of 225 square miles and a popu-

lation of one and a quarter million, has some 2,000 indigenous

species of plants, a total which is greater than that of the British

Isles; the Malay Peninsula has about 10,000 species, while the

estimate for the Malaysian region as a whole is 30,000 species.

The taxonomist working in the region whose flora he is describ-

ing has the great advantage of being able to study many of the

plants in his group living in their natural habitat, which is an in-

valuable aid to his study of herbarium material. Some of the plants

may also be growing in the local botanic gardens. For modern

taxonomic study it is essential for the worker to see as many as

possible of all the collections that have ever been made of the

species he is studying. Thus most herbaria are willing to lend their

specimens to experts wherever the latter may be working.

The importance of taxonomic research

The great Linnaeus has said “Science, and in the first place

botany, is the only reliable basis of private as well as national

economy.” Many people would regard this statement as too sweep-

ing; nevertheless taxonomic research is of vital importance to the

agriculturist, the forester and other applied scientists, as well as

all those who wish to assess and develop the natural resources of a

country. Many people fail to realise the importance of giving plants

their correct scientific names, so that their results can be under-

stood in all countries regardless of the language used. Their re-

search would often be of greater value if they took the trouble to

ascertain the correct names of the plants they are working with,

and, in the case of plants not widely known, would lodge authentic

specimens at some recognized institution for reference purposes.

Anthropologists and others frequently write of plants used by indi-

genous people as food and medicine and in magic, etc., and are

content to give the vernacular names only, thus rendering their

work nearly worthless for the botanist, chemist and others, who

might well follow it up had they taken the trouble to collect speci-

mens and get them properly identified. I have long felt that there is

far too little active collaboration between the agriculturist and the

botanist to their mutual disadvantage.

The knowledge of the correct names of plants is essential for the

ecologist. I would like to see the collecting numbers of plants

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quoted in ecological papers, so that future workers have access to

the specimens of which authors have based their results. Some eco-

logical studies have been carried out by the staff of the Singapore

Botanic Gardens in the past and I would like to see this extended.

I believe that ecological studies in the widest sense provide the only

sound basis for the preparation of rational plans of land utilisation.

Botanists from their collections and research discover plants

which are closely related to those in cultivation and thus can pro-

vide material of great value to the plant breeder. An Indian sugar-

cane breeder visited Singapore in 1956 and found in our herbarium

one sheet of a variety of Saccharum spontaneum, which had been

collected by Ridley in 1891 in a remote area of Pahang. Travelling

by plane, car and boat he was able to obtain planting material of

the variety and returned to Singapore within four days.

This section could be extended considerably. I hope I have said

enough to convince the Philistines that the plant collecting and

taxonomic research carried out by the staff of the botanic gardens

has a wider application than is often realised.

HORTICULTURAL WORK

Exchange of Planting material

Most botanic gardens produce an annual Index Seminum, or

seeds list, which they distribute to other botanic gardens for favour

of exchange. As the number of species of which seed is available in

Singapore varies but little from year to year, our list is produced

at irregular intervals for distribution and is also available on re-

quest. We receive. lists from all over the world, including most

communist countries. In this way a free exchange of planting

material is effected, thus enriching the living collection of plants in

all countries. In 1955 the Singapore Botanic Gardens sent out 515

packets of seeds and 113 plants on an exchange basis and received

362 packets of seeds and 371 plants, including gifts. 368 packets

of seeds and 87 plants were despatched in 1956, while 575 packets

of seeds and 185 plants were received. Only a small proportion of

the introductions may be really successful in Singapore; neverthe-

less, our collection of plants is being added to constantly and new

plants of horticultural value are available for the country as a

whole.

The immediate supply of seeds from the Singapore list cannot be

guaranteed as many local seeds are viable for short periods only

and stocks cannot be kept in good condition, while many species

fruit at irregular or long intervals. Every effort is made to meet

requests, however, and lists are kept of requirements and the seeds

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Vol. XVII. (1958).

are collected and despatched at the earliest opportunity as they

become available. Our current list gives 864 species, some of which

are grown in the Gardens, while others are collected in the natural

vegetation on the island. Many of the larger seeds of short viability

are sent out in damp powdered charcoal or damp coir dust. Ger-

mination begins during transit and we receive very favourable

reports on palms and other seeds sent out in this way.

In addition to this normal exchange of seeds between botanical

institutions, many requests are received for living plants, as well as

spirit and fixed material, from specialists elsewhere. Wherever pos-

sible it is provided, except for unreasonable requests which would

require a major expedition to fulfil or take up too much time. In

all this exchange care is taken to see that only authentic material

1s sent out.

During the past few years a collection of over 400 species of

cacti and other succulents has been built up. These grow surpris-

ingly well in Singapore, provided they are sheltered from the rain

and receive plenty of sun. They are grown in open-sided houses

with glass roofs. Succulents are becoming very popular among

local growers.

Members of the staff on collecting expeditions bring back local

plants of horticultural value for trial in the Gardens, while friends

and correspondents send in similar material. Living material is

usually packed in polythene bags, in which it remains fresh for a

considerable period. One of the most successful introductions in

recent years has been the New Guinea Creeper (Mucuna bennet-

tii), which produces hundreds of large spectacular trusses of

brilliant flame-coloured flowers several times a year.

Although most of the work on economic plants has been taken

over by the Agricultural and Forestry Departments, there is still a

good collection in the Singapore Botanic Gardens and new ones of

special interest, such as Rauwolfia spp., are introduced for trial.

Breeding of Orchids and Other Plants

The seasonless climate of Singapore, with an average annual

rainfall of 105 inches per year, a mean temperate of about 80°F.

with little diurnal change and a mean relative humidity of 82 per

cent, makes it difficult to find plants which will flower regularly and

provide colour in the Gardens. The aspect of the natural vegetation

is varying shades of green and offers little in the way of good hort-

icultural plants. Consequently, most of the garden plants grown in

Singapore have originated in countries other than Malaya and

where there are distinct cold or dry seasons. Under our conditions

there is no special season of flowering or leaf-fall and individual

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Gardens Bulletin, S.

plants of the same species adjust themselves quite differently. Con-

sequently, we never get the spectacular bursts of flowering of such

trees as Delonix regia or Jacaranda as can be seen in India or East

Africa.

One of the solutions to this problem is the breeding of new varie-

ties of plants suited to local conditions and this work has been

actively pursued at the Singapore Botanic Gardens for many years.

Although hybridisation has been done in various groups such as

Bougainvillea, Cannas, etc., by far the most attention has been

devoted to orchids. This work was begun in 1929 and has produced

very successful and spectacular results. The aid is two-fold, namely,

to produce hybrids of horticultural merit and commercial worth

and to investigate the relationship between the various groups of

orchids. Malaya has some 800 species of indigenous orchids, but

only a few of these are suitable for garden plants. By hybridising

these, together with imported species from Burma, Java, the

Moluccus, New Guinea and the Philippines, a very wide range of

free-flowering and beautiful hybrids have and are being produced.

The principal genera used in crossing are: Arachnis, Dendro-

bium, Renanthera, Spathoglottis and Vanda. The first Singapore

hybrid was Vanda Miss Joaquim raised in 1893, but the bulk of

the hybrids now grown have been produced in the last 30 years

and many of these since the war. Some 2,800 crosses have been

made to date and every year new hybrids are coming into bloom.

Local growers also make crosses and bring their seeds for raising

at the Gardens, while a few have now mastered the technique and

raise them themselves.

The seedlings are first grown in conical flasks under sterile con-

ditions. The medium used in Singapore is agar-agar with Vacin’s

solution and 2 per cent sugar, at a pH of about 5-2. Full details of

the method are given by Holttum (1953b). In 1956 experiments

were begun using coconut milk (the water inside the mature nut)

in the medium and this is giving a considerable stimulus to the

early growth of the seedlings. The mature orchids are grown in the

open or with slight shade, either in pots or in beds, depending on

the type, and details of the cultivation can be found in Holttum

(1953b) and Henderson and Addison (1956).

Mention can only be made of one or two of the hybrids raised in

Singapore. A fuller account of the principal hybrids, except for the

very recent ones, with photographs of a spray and individual flower

of each hybrid, has recently been published in Henderson’s and

Addison’s Malayan Orchid Hybrids (1956). Vanda Tan Chay

148

Vol. XVII. (1958).

Yan, which first flowered in 1952, was awarded a First Class Certi-

ficate of the Royal Horticultural Society at the Chelsea Flower

Show in 1954 and is considered to be one of the best Vanda

hybrids bred anywhere in the world. Arachnis Maggie Oei, now

one of the principal cut flowers in Singapore, flowers continuously,

whereas its larger parent A. flosaeris flowers but twice a year.

Among the inter-generic hybrids bred in Singapore are Aranda

(Arachnis * Vanda), Aranthera (Arachnis * Renanthera)

Renantanda (Renanthera x Vanda), while our first tri-generic

hybrid flowered in 1956 and has been named Ridleyara Fascad, in

honour of H. N. Ridley. Mr. Ridley was informed of this shortly

before his death.

There are a large number of enthusiastic orchid growers in

Singapore and the Federation and there is a considerable demand

for plants and seedlings grown in the Gardens. The sale of cut

orchids, both local and for export, is rapidly increasing. An ex-

perienced cytogeneticist is expected to visit Singapore shortly to

work on problems of orchid hybridization.

Sale of Plants

Although the sale of plants is not an essential function of a

botanic gardens, it is desirable in those countries where there are

few nurseries. The Singapore Botanic Gardens sells plants daily,

but limits the sales to those plants which are not readily available

in the local nurseries. The revenue from the sale of plants in 1956

was $27,245 (Straits), of which the major share came from the

sale of 3,446 orchid plants and seedlings, while 26,092 other plants

were also sold.

Advisory Work

The horticultural staff of the Singapore Botanic Gardens advise

and assist various government departments in planning the lay-out

and planting of roadsides, open spaces, school compounds, etc.,

- and usually supply free material for this purpose. They also main-

tain close contact with the City Council Parks Department and

many requests for advice are also received from private firms and

individuals. Thus the Gardens play a large part in beautifying the

city and island.

Men from the Rehabilitation Centre of the Labour Department

are accepted for training as gardeners. The Director of the Singa-

pore Botanic Gardens is the Imperial War Graves Commissioner’s

representative in Singapore and the staff of the Gardens take charge

of the care and maintenance of the Kranji War Cemetery.

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Gardens Bulletin, S.

EDUCATIONAL WORK

General

The maintenance of a large, named and labelled collection of

local and introduced plants attractively laid-out in the Botanic

Gardens enables all visitors, who are interested, to increase their

knowledge of plants. Parties of school children and students, both

from Singapore and the Federation pay regular visits to the Gar-

dens and occasionally parties are taken round by members of the

staff. Special exhibits are arranged from time to time. One which

proved popular with teachers and students was an exhibit of seven

common plants which were changed weekly. The plants were

labelled with their scientific, English and Malay names and family

and brief notes were appended. Members of the staff are often

requested to give lectures to local societies, a valuable method of

educating the public in our work.

The staff of the Gardens co-operates fully with the staff of the

University of Malaya’s Department of Botany. The University is

adjacent to the Gardens and so the latter are very handy for lec-

turers and students who wish to use them, while much material is

provided for class work and research.

Singapore Gardening Society

The staff of the Singapore Botanic Gardens was largely respon-

sible for the founding of the local Gardening Society, which at pre-

sent has about 250 members. An active interest is taken in its work,

as well as that of the Malayan Orchid Society. Some monthly

meetings are held at the Gardens and emphasis is usually placed on

practical demonstrations. A very successful annual flower show is

held and the standard of blooms and plants exhibited is high. In

these ways, the interest of local horticulturists is stimulated and the

standard of gardening is raised.

PUBLICATIONS AND LIBRARY

It is very desirable that botanic gardens carrying out research |

should have their own journal for the publication of their results.

Ridley founded the Agricultural Bulletin of the Malay Peninsula

in 1891. The title was changed to the Agricultural Bulletin of the

Straits and Federated Malay States in 1900 and it was then issued

monthly. This journal was largely devoted to work on economic

plants. Ridley published many taxonomic papers in the journal of

the Royal Asiatic Society, Straits Branch, of which he was editor

from 1889 to 1911, as well as being secretary of the Society. The

title of the Agricultural Bulletin was changed in 1913 to the

Gardens’ Bulletin, Straits Settlement and in 1947 to the Gardens’

150

Vol. XVII. (1958).

Bulletin, Singapore. This is published at irregular intervals and an

attempt is now made to produce a volume per year. It is largely

devoted to papers on local taxonomy. It is distributed to the major

botanical institutions in the world and exchange arrangements now

exist with 160 scientific bodies. It can also be purchased direct

from the Botanic Gardens, Singapore. In exchange we receive valu-

able periodicals and reprints, which are so essential for our work,

and which we would not be able to purchase from our slender

library vote. The Annual Report is also printed and distributed.

In addition to these and other publications already referred to in

this article, the Gardens staff prepares from time to time more.

popular works which are of general interest to the amateur. Such a

work is Henderson’s Malayan Wild Flowers (1949-54) published

by the Malayan Nature Society. A series of small illustrated pam-

phlets, each containing drawings and brief descriptions of ten

Malayan Garden Plants are also produced. Popular horticultural

articles are submitted regularly to the Malayan A gri-Horticultural

Journal and the Malayan Orchid Journal. Names and descriptions

of our new orchid hybrids are published in the latter.

Adequate library facilities are essential for the research workers

at botanic gardens. The taxonomist must consult original publica-

tions for the names and descriptions of the plants on which he is

working, and for this the older, as well as modern literature is

required. The Singapore Botanic Gardens are particularly fortunate

in that they have one of the finest collections of old botanical books

in South-East Asia, many of which are of considerable value and

rarity. In all, the library contains some 8,000 bound volumes, as

well as many periodicals.

Since 1890 there has always been an artist on the Gardens’ staff,

who prepares drawings and paintings for the specialist staff. The

paintings done by James and Charles de Alwis between 1890 and

1908 are of outstanding beauty and accuracy and are among the

treasured possessions of the Singapore Botanic Gardens. Over the

years a large and valuable collection of plant drawings and paint-

ings has been built up.

NATURE RESERVES

I have already shown how Cantley and Ridley were responsible

for gazetting and maintaining the first forest reserves in the Straits

Settlement. These were later taken over by the Forest Department.

The Singapore forest reserves reverted to the control of the Botanic

Gardens in 1937 and have since been maintained as nature re-

serves. Under the Nature Reserves Ordinance of 1951, some 8,000

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Gardens Bulletin, S.

acres in Singapore were “set aside for the purpose of the propaga-

tion, protection and preservation of the indigenous fauna and flora

of the Colony”, as well as “providing, under suitable conditions and

control, facilities for the study of and research into matters relating

into the fauna and flora and the physical conditions in which they

live’. The reserves are administered by a statutory board of

management of six nominated trustees, with the Director of the

Botanic Gardens as ex-officio chairman. The day-to-day adminis-

tration is carried out by the staff of the Singapore Botanic Gardens.

Inevitably, with the spread of the rapidly increasing population,

plants and animals, which were known to have occurred in Singa-

pore, have disappeared completely. Every effort is being made to

preserve the remnants which are left for present and future genera-

tions to study and enjoy. Only a comparatively small area of the

nature reserves consists of primary vegetation. The most important

is undoubtedly the Bukit Timah Reserve covering 163 acres on the

highest hill on the island. It has been a botanical collecting ground

for more than a century, and it is the type locality of many Malayan

plants. It consists of primary tropical evergreen rain forest, with

many trees 50—70 m. high, and for the greater part it has not been

cut over nor damaged. There are also important areas of mangrove

on the south coast preserved as a nature reserve.

Thus the Botanic Gardens have guarded jealously this heritage

of the people, which might so easily have been lost for ever.

THE FUTURE

And what of the future? Singapore is shortly to receive inde-

pendence. Already the Government has embarked upon a pro-

gramme of rapid Malayanisation of its staff, although it is not

likely that this will be fully achieved at the Gardens as rapidly as

in some other departments. A local man has returned from training

as a horticulturist at the Royal Botanic Gardens, Kew, while bota-

nists and others are to be sent overseas for further training and on

their return will take over from the expatriate officers. Much will

depend on these new recruits to maintain the Gardens and collec-

tions and to carry on the tradition of research which has been built

up during the past hundred years. It is hoped that the scheme re-

cently devised to attract research workers from Britain and else-

where to Singapore to carry out botanical research in their

specialised fields will be continued and extended.

The Gardens are financed entirely by the Singapore Government

and now come under the Ministry of the Chief Secretary.* In the

* In 1959 the Gardens came under the newly formed Ministry of National

Development. Editor

152

Vol. XVII. (1958).

1957 estimates a total of $546,780 (=£63,791) is voted to the

Gardens and this includes the votes for the upkeep of Government

House Domain and the Nature Reserves. If they are to survive, it

will be necessary to convince the politicians and the general public

that the Gardens are fulfilling a worthwhile rdle, not only as a

public park, but also that the research carried out is of value to

the economy and prestige of Singapore. As Commelijn said in 1701

(de Wit, 1949) “It is certain, however, that this Science, like all

Sciences, flourishes sometimes more and sometimes less, all in

accord with the inclination of Rulers and the Favour of Govern-

ment.”

A recent distinguished visitor wrote in my visitor’s book “The

Singapore Botanic Gardens are unique. Civilisation would be the

poorer if they were not maintained.” In discussing the history of

botanic gardens in the tropics, I have shown how many of them

were neglected or abandoned and failed in their primary function,

which is the scientific study of plants. New botanic gardens are

being founded at some of the new tropical universities and univer-

sity colleges, but it will take them many, many years to build up

the type of collection of living and herbarium specimens which is

now available in Singapore. It is my fervent hope that the Singa-

pore Botanic Gardens may be fully maintained for many long years

to come and to continue as a leading centre of tropical botany and

a worthy memorial to Ridley and those other members of the staff

who have helped to raise them to their present position.

I have shown how in these uncertain and unsettled days there

still exists much goodwill and active co-operation between botanic

gardens, wherever they may be situated. As Baas Becking says in

the Preface to Flora Malesiana “as long as sentiment, politics,

greed and bigotry rule this world, a purely scientific endeavour

may become a binding force between individual groups, and may-

be, even between nations’. If the aims and ideals which motivate

the work of botanic gardens could be extended to certain other

fields of human endeavour we could make this planet a better place

on which to live.

ACKNOWLEDGEMENTS

In addition to the works quoted in the bibliography, reference

has been made to all the published Annual Reports of the Botanic

Gardens Department, Singapore from 1875 to the present day, as

well as unpublished data in the files of the Department.

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Gardens Bulletin, S.

BIBLOGRAPHY

BURKILL, I. H. (1918)—The Establishment of the Botanic Gar-

dens, Singapore. Gardens’ Buil., II, 2, 55-72, & 3,

93-108.

BURKILL, I. H. (1935)—A Dictionary of the Economic Products

of the Malay Peninsula. Crown Agents, London.

CuHiPpPp, T. F. (1920)—Botanic Gardens. Gardens’ Bull., I, 8,

306-7.

DE Wirt, H. C. D. (1949)—Short history of the phytography of

Malaysiana Plants. Flora Malesiana, I, 4, LXXI-

CLXI.

Hart, W. E. (1919)—The Botanic Garden of Pamplemousses.

Kew Bull. of Misc. Information, 279-286.

HENDERSON, M. R. & Appison, G. H. (1956)—Malayan Orchid

Hybrids, Govt. Printer, Singapore.

Ho.LtTtTum, R. E. (1934)—The Waterfall Gardens, Penang. IIlus-

trated Guide. Govt. Printer, Singapore.

Ho.tttum, R. E. (1953 a)—Gardening in the Lowlands of

Malaya. Straits Times Press, Singapore.

Ho.tttum, R. E. (1953 b)—A Revised Flora of Malaya, vol. 1

Orchids of Malaya, Govt. Printer, Singapore.

H. T. S. (1956)—The first Botanical Gardens in Singapore.

Straits Times, 22-12-56.

PURSEGLOVE, J. W. (1955 a)—Mr. H. N. Ridley’s Hundredth

Birthday. Nature, 176, 1092—93. —

PURSEGLOVE, J. W. (1955 b)—Ridley, Malaya’s greatest natu-

ralist. Malayan Nature Jour. 10, 2, 43-55.

PURSEGLOVE, J. W. (1955 c)—The Ridley Centenary. Govt.

Printer, Singapore.

RicKETT, H. W. (1956)—The Origin and Growth of Botanic

Gardens. Garden Jour. of New York Bot. Gard. 6, 5.

RIDLEY, H. N. (1910 a)—The abolition of the Botanic Gardens

of Penang. Agric. Bull. of Straits and Fed. Malay

States. IX, 3, 97-105.

RIDLEY, H. N. (1910 b)—Historical Notes on the Rubber In-

dustry. Agric. Bull. of Straits and Fed. Malay States,

XI, 6, 200-214.

154

Kew and Singapore

T. A. RUSSELL

Senior Scientific Officer,

Department of Economic Botany,

Royal Botanic Gardens, Kew

“The gift of a useful plant seems to me more precious than

the discovery of a gold-mine’—BERNARDIN DE ST. PIERRE

(translated)*

IT Is A pleasing thing that the Botanic Gardens at Singapore and

the Royal Botanic Gardens at Kew should be celebrating in the

same year, the one honouring 100 years of existence on the present

site, the other commemorating the 200th anniversary of its found-

ing. As we congratulate each other on the happy occasion and

wish each other well for the future, it is also fitting that we recall

the close bonds which have been woven between Kew and Singa-

pore during their common life. In order to understand how these

bonds came to be fashioned, it may be helpful to retrace some

part of the history of the gardens at Kew.

The Royal Garden of Princess Augusta

Two hundred years ago Kew House, situated near the River

Thames 7 or 8 miles above Westminster, was the residence of

Augusta, Princess of Wales and daughter-in-law of King George II.

Here this German princess had been brought by her husband,

the Prince of Wales, soon after their marriage, and here she con-

tinued to live after his early death. In 1759 she decided on making

improvements to the place. She engaged a prominent architect,

Sir William Chambers, to re-design the grounds, and employed a

skilled gardener, William Aiton, to make a garden of exotic plants

or botanic garden. Chambers laid out the grounds in the fashion-

able mode of the day, building a handsome Orangery, a Great

Stove (the largest heated greenhouse then in existence), a Pagoda,

and various Temples; and some of his buildings are features of the

gardens still. Aiton was no less industrious, and, in seeking plants

for the botanic garden, Princess Augusta was greatly helped by

the third Earl of Bute, whose influence was exerted to secure plants

from many distant lands. Soon the garden became famed through-

out Europe for the unrivalled variety of its plants.

* From the inscription on a memorial erected in 1860 in the Botanic

Gardens of Pamplemousses, Mauritius.

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Gardens Bulletin, S-

New Functions for the Royal Gardens

It may be noted that, at the start, no more was desired of the

garden than that it should provide pleasure and interest to its royal

owner and her friends. In 1772 when Princess Augusta died, the

garden was taken over by her eldest son who, by this time, had

succeeded to the throne as King George III, and he increased the

area of the property by addition of adjoining lands. For guidance

in the care of the garden the King sought the advice of Sir Joseph

Banks, a man of outstanding ability and scientific knowledge, who

was to exercise a directing influence on its activities over a period

of many years. His view was that the royal garden should become

a place where the plant-resources of the world would be studied

and made known, and where men might be trained who would go

abroad to seek and collect new plants. These proposals, being

approved by the King, set the Gardens on a course which has been

followed to the present day. There have been times when the policy

wavered, and indeed a bad period followed the death of Banks and

George III when the closure of the Botanic Gardens was under

consideration. But this, causing an outcry from a number of people

who appreciated their value, led in 1838 to the appointment of a

Commission to advise on the future of Kew. This Commission not

only recommended the continuance of Kew as a State institution,

but urged a more vigorous direction of its activities, in particular

the study of plants and their uses, and the supplying of useful

plants to territories overseas. To make effective these recommen-

dations the first Director, William Hooker, was appointed in 1841.

In the years since then under successive directors, the study ot

the world’s plants has been the main task of Kew. Here in the

course of years, has been built up an unrivalled collection of plants,

some in living state in the gardens and glasshouses, but many more

in dried state in the Herbarium, providing a rich store of material

for study of the identification and affinities of plants, their distribu-

tion over the world’s surface, and the uses to which they can be

put. Amongst this collection the flora of Singapore and of the

neighbouring territories is well represented, thanks to the efforts

of past collectors, and as a result Kew is able to be of assistance in

the compiling of the great Flora Malesiana now being undertaken.

Kew Collectors

But it is in Kew’s other functions, the sending out of men trained

in the knowledge and care of plants, and the supplying of plants

of economic use, that the strong links between Kew and Singapore

have been forged. As early as 1772 men trained at Kew were being

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Vol. XVII. (1958).

sent abroad, the first, Francis Masson, being appointed by Banks

to go to South Africa there to travel and collect seeds and plants

for Kew. Masson proved a diligent collector sending back to

England many plants hitherto unknown, of which some have since

become garden-favourites. He subsequently made expeditions in

Europe, the West Indies, and Canada, with such good results that

Banks was encouraged to send out others.

Such a one was David Nelson who went as botanist in 1776 on

the third voyage which Captain Cook made to the South Seas, and

collected plants at many places where the ship called. Returning

safely from this voyage, he was then appointed to Captain Bligh’s

ship H.M.S. Bounty sailing for Tahiti. At this place breadfruit

plants were to be taken on board and Nelson’s job was to see that

they were properly stowed and tended during their voyage to the

West Indies. The story of that ill-fated voyage and its mutiny is

well known. Loyal to his captain, Nelson shared the privations of

the open boat during the long journey to Timor, only to die a few

days after reaching this haven, commended by Captain Bligh for his

“manly fortitude in our late disastrous circumstances.”

To retrieve this disaster a second voyage was made by Captain

Bligh in 1791. On this occasion two Kew men accompanied the

expedition, which was completely successful in conveying a large

supply of breadfruit trees to the West Indies where they were es-

tablished without difficulty. One of these men, Christopher Smith,

deserves mention here as being probably the first man trained at

Kew to give his service, if not to Singapore, at least to Malaya.

In 1794 Smith was appointed botanist of the Honourable East

India Company at Calcutta and, three years later, he was sent to

the Molucca Islands to collect plants. These islands were then the

centre of the spice trade which had been held as a strict monopoly

first by the Portuguese and latterly by the Dutch. In 1797, through

fortunes of war, Britain had access to the spice islands and

Christopher Smith’s commission was to obtain planting-material

of valuable spice-trees for growing in the East India Company’s

territory. In this he was extremely successful, obtaining no less

than 70,000 nutmeg trees and 55,000 clove trees of which the bulk

went to Penang whence someone was able to write “we are now

become masters of every kind of spice plant valuable or un-

common.” Smith himself went to Penang to supervise their cultiva-

tion and died there in 1806. Doubtless from some of his trees

were descended the nutmegs and cloves introduced in 1819 into

the Gardens of the Agri-Horticultural Society in Singapore, which

provided a useful source of revenue to the Society for many years.

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Gardens Bulletin, S.

About this time, botanic gardens were being opened up in many

British colonial possessions. The first were in West Indies, at St.

Vincent in 1764 and Jamaica a few years later. Then came gardens

of the East India Company, at Calcutta in 1786 and Madras soon

after, the spice garden at Penang about 1800, and the garden of

Peradeniya, Ceylon in 1821. These places looked to Kew to pro-

vide men trained in horticulture, and Kew men soon filled a majo-

rity of posts in the colonial gardens and were responsible for

starting horticultural work in many new territories. These men

commonly remained in touch with Kew and there was often a

frequent exchange of plants, Kew sending out plants of economic

or ornamental value for the new gardens and receiving in return

specimens of the indigenous vegetation.

Kew Men in Singapore

Singapore was added to the botanic gardens served by men of

Kew in 1875, when the Director of Kew was asked to supply a

practised botanist for the post of Superintendent. He chose Henry

James Murton, a young man for the job, but a skilled and keen

horticulturist. The Gardens at Singapore up to this time had served

chiefly as a pleasure-park, and Murton deserves recognition for

altering its character, for improving the collection of plants, seeing

to their naming, and making trials of new economic plants. This

work was continued from 1880 by another Kew man, Nathaniel

Cantley, a great believer in system and orderliness, whose notable

contribution to the Gardens was cut short by his early death. Both

these men had the assistance of a third trained at Kew, Walter Fox,

whose service in Singapore and Penang extended over 31 years.

These three did much to set the Botanic Gardens on the right lines

and prepare for the progressive development which followed under

H. N. Ridley and his successors. In more recent times men trained

at Kew have continued to be appointed to Singapore and, amongst

them, we would pay tribute to a splendid horticulturist J. C. Nauen,

whose service to his fellows and gallant death as a prisoner-of-war

are remembered at Kew, as doubtless at Singapore.

It must not be suggested, however, that movement between Kew

and Singapore has been all one way, for it was to the vicinity of

Kew that Ridley came on retiring from Service in Singapore and,

so long as he remained in active health, he visited the Gardens at

Kew almost daily. Another former Director from Singapore, R. E.

Holttum, is also a neighbour at Kew, and both I. H. Burkill and

E. J. H. Corner are sufficiently near to be occasional and welcome

visitors. It is a happy occurrence that in this year of celebration

the Kew Guild, the association of present and past members of the

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Vol. XVII. (1958).

_ Kew Staff should have as its President F. Flippance, a Kew man

whose long experience in the East has included service in the

Singapore Botanic Gardens. Moreover at Kew we have been glad

to welcome as student-gardeners young men from the Gardens at

Singapore who, after a period of training at Kew, return east with

a widened knowledge of plants and of horticultural skill. These

too, it may be hoped, will help to strengthen the bonds between the

two Gardens.

New Plants for the East

For these are the strong links between Kew and Singapore, the

men such as have been mentioned whose skill and loyal service

have been shared by both places. Nor must we forget the plants

distributed from Kew which have formed so useful a link with

other gardens.

Reference has been made to co-operation between the Honour-

able East India Company and the Royal Botanic Gardens, and this

had as one of its most productive results the introduction of qui-

nine into the Far East. A century ago, while the effectiveness of

quinine against malaria was well known, the drug itself was scarce

and not generally available, since the only source was the wild

growth of Cinchona trees in the Andes. An expedition to South

America in 1860 was successful in obtaining seed of several spe-

cies, and the seedlings, raised at Kew, were sent to India, Ceylon,

and other places. In India particularly the plants were readily

established and plantations were made on a large scale with the

result that quinine was soon obtainable throughout the East at low

cost. The influence this introduction had on general health and

well-being can hardly be estimated.

So encouraging was the outcome of this venture that it was fol-

lowed by a second, an expedition to Brazil to collect seed of the

Para rubber-tree. A supply of seed was sent to Kew in 1876 and

gave rise to seedlings which, as soon as they were large enough,

were distributed to countries where it was thought they might suit-

ably be grown. A case of these plants sent to Singapore in 1877

came into the hands of H. J. Murton, and 12 plants survived to be

planted in the Gardens at Singapore and 9 at Kuala Kangsar in

Perak. From these plants, and others which came subsequently

from Ceylon, was built up the great rubber industry of Malaya,

which brought a new commodity into every civilised home and

made possible a new era in road-transport.

It may seem idle to conjecture what the history of Singapore

would have been if Hevea had not arrived when it did, if the

threatened closure of Kew in 1838 had in fact taken place, if the

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Gardens at Singapore had not been in running order with a Murton

and a Ridley to take charge. Such thoughts are not altogether vain

if they bring home to us what great service these botanic gardens

have rendered through the exchange and development of useful

plants. Nor must this be regarded as their only, or even their chief,

function, which, as has been shown, is the study of plants and

plant-resources in its widest sense.

Botanic gardens have not been without their critics, who, at

different times and places, have pointed out how much money is

spent and how little comes back as revenue, urging that they be

closed or their activities trimmed in the interest of economy. Look-

ing back 100 years, and 200 years, we can clearly see how time

has fully justified the men of vision, men like Banks, the Hookers,

and Ridley, whose faith in botanic gardens led them, not to close

them down, but to develop and shape them to serve the community

to best effect. Having viewed the splendid record of the past, we

can look to the future of these Gardens with confidence, believing

that the skill and devoted service, which have already achieved so

much, may have no less a contribution to make in years to come.

160

Singapore and Flora Malesiana

C. G. G. J. VAN STEENIS

Professor of Tropical Botany,

Leyden and Amsterdam Universities, Netherlands

IN MY TRIBUTE to the healthy centenarian, the Singapore Botanic

Gardens, I feel that the intimate relation which has grown in the

past years between the Singapore Botanic Gardens and the Flora

Malesiana requires an introduction and an explanation. Many of

the Singapore citizens will know this work only by name but not

by personal experience.

Let me first explain this Flora Malesiana. It intends to be a

modern critical inventory of the native, introduced, and major cul-

tivated plants of the entire Malaysian region. This is the whole

archipelago between the Asiatic and Australian continents includ-

ing also Singapore and the Malay Peninsula up to about the isth-

mus of Kra where the flora rather abruptly changes into that of

continental South-East Asia. It wants to give critical data on the

ecology and distribution of all the plants, of the altitude at which

they occur, the soils which they can stand or prefer, the climates

which they tolerate, their mode and period of flowering and fruiting,

the manner in which they are pollinated, the mode of dispersal

and germination of their seeds, etc.

The knowledge to be accumulated must be carefully collected

by qualified, scientific botanical specialists and we are happy to

put on record that many prominent persons all over the world

have been persuaded to add to the effort. Malaysia is botanically

one of the richest countries of the tropics, it can boast of probably

not far from 25,000 different species of flowering plants in all,

from the tiny herbaceous up to the colossal tualang and the majestic

jelutong, unique creations of tropical nature beside which man

feels his smallness.

The exact knowledge of this vegetable wealth has grown slowly

and gradually, in former times largely from curiosity of disinter-

ested scientists, local and overseas, but it appears more and more

in modern time that this knowledge is intimately connected with

far-going aspects of human welfare, land-use, economy, and in-

dustry. The fast-growing world population, with its increasing

needs and its stupendously developing techniques, is steadily ask-

ing more information about the three kingdoms of nature, the

earth, the animals, and the plants. Among these kingdoms the

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plants play an enormous role by their bulk and by the easy way in

which man can manage them. But before man can use them effici-

ently he must know their identity, their name. The critical name of

a plant species is the alpha of botanical knowledge, it is the key

to our knowledge of it. And that is what we intend to accomplish

with the Flora Malesiana. Following this trend of thought it will

also be clear that, though elaborated by disinterested research, the

Flora Malesiana is automatically becoming a most useful tool for

humanity in the East.

How is it then, the readers will ask: have we not already a

Flora of Malaya? This is admittedly true, there are even three with

a start of a fourth. The flora of Malaya was in the past century in-

corporated in the Flora of British India. Shortly afterwards it

was the subject of the excellent work by King & Gamble, Mate-

rials towards a Flora of the Malay Peninsula, and finally there is

the more recent Flora of the Malay Peninsula by Ridley.

The first two treatments are good but more than half a century

old and naturally very incomplete. The third one unfortunately

does not come up to the required standard of accuracy. All of them

suffer from two inadequacies, viz the lack of correlation with

plants described from other parts of Malaysia and the lack of

general data, on uses, distribution, and ecology of the species. The

second desideratum was to be filled in the Revised (fourth) Flora

of Malaya initiated a few years ago of which two volumes were

issued, viz on the orchids and ferns, by Professor R. E. Holttum.

Admirable as these volumes are, there is still the lack of correla-

tion notwithstanding their author’s speciality on these two groups.

A large plant geographical study has revealed that Malaya,

Sumatra, Borneo, the Philippines, Java, the Moluccas, and New

Guinea form for plants one unbreakable whole. Plants do not

keep to political boundaries, they have a distribution of their own

and attained their area through the geological ages of the past.

This means that for an adequate knowledge of their identity they

should be pursued throughout that area.

We should strive then first towards a general Flora of the entire

Malaysian province, as Nature made it, before giving an account

of a politically defined part of it, a so-called local Flora.

A few days ago I was struck again by the truth of this basic

principle, in trying to arrange the species of a rather important

timber genus Campnosperma, téréntang or séréntang as it is known

by its vernacular name; it is common in the lowland and swamp

forests of Malaya. Mr. Corner, who in 1939 made a study of it

on the basis of material from Malaya only, concluded that there

was an undescribed new species, téréntang jantan of Johore, which

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Vol. XVII. (1958).

he called Campnosperma minor and which would be unique to

Malaya. But revising the genus over the whole of Malaysia a few

weeks ago, it appeared to two of us that this same new species had

already been described from the swamp forest of Borneo, under

the name Campnosperma squamata by Ridley in 1933, but

occurred also in the Moluccas and New Guinea and from there

had been described still earlier as Campnosperma montana by

Lauterbach in 1920. The latter is obviously the oldest and correct

name for it. Once that is known we can properly proceed to judge

its ecology and uses over its entire range and we know where

we are.

This example is no exception, unfortunately it is the rule as soon

as groups are revised regionally. The background of this rule is the

fact that in the past the study of the flora of the Malaysian region

has been approached by local work, in Malaya, in Java and other

parts of Indonesia, in the Philippines, in West New Guinea, and in

Papua, instead of by regional attempts covering the entire archi-

pelago. This serious lack of correlation we try to remedy by the

Flora Malesiana attempt, from which all separate states and terri-

tories will have the same benefit. In this whole, Malaya, with its

very rich flora, forms an essential integral part.

The senior leaders of Malayan botany, who were or are directors

or staff members of the Singapore Botanic Gardens, have in the

past decade realized the scientific necessity to give priority to a

regional flora over a local flora, large as it may be, and Flora

Malesiana is happy to have their authoritive collaboration.

Mr. Burkill Sr., the author of the standard work on the Economic

Products of the Malay Peninsula, revised the yam _ family

(Dioscoreaceae), Mr. Corner is doing the colossal job of tackling

the fig family (Moraceae), Mr. Sinclair revised the nutmeg family

(Myristicaceae) and intends later to work on the kenanga family

(Annonaceae). Mr. Furtado has also, in his studies on aroids and

palms, felt the need to step far beyond the borders of Malaya.

Last not least, Prof. Holttum is organising the Pteridophytes, a

lofty attempt; it will take him at least ten years to come to achieve

this, his final magnum opus.

One might query: but what is the use for Malaya to know all

about the floras of the other islands, even as far as Papua? This

question has already partly been answered by the example of

Campnosperma given above. It is curious and possibly of interest

to Malayan readers to know that the closest relative of the genus

Hevea, to which the pararubber tree belongs, is a tree native in

Papua, Annesijoa novoguineensis. In wider scope this question is

part of the general issue: what is the potential use of plants?

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Frankly, no botanist is capable to say which plant can be of what

special service to man and which not. It is certainly not determined

by size: most humble annual plants like flax have shown their

importance whilst woody giants are sometimes of hardly more

value than as firewood. Use depends on the needs of man and that

varies with place and time. The tuba plant (Derris) was until the

twenties only locally used as a fish-poison until it appeared that

this fish-poison contained a powerful insecticide harmless to man.

Then it became at once an industrial plant from which money

could be made. Nowadays interest in the plant has waned

as during the war other, artificial insecticides have superseded it.

When in the thirties it appeared necessary to have pararubber latex

shipped in bulk, without coagulating the milk, the tuber of a fetid

aroid containing a special kind of starch which could do that trick,

but to which no use had ever been ascribed, came into great

demand.

Some other Malaysian plants have never reached the industrial

phase, possibly because they have never been given a trial, such as

the bark-cloth yielding figs and champedaks. Nothing is yet known

about possible industrial qualities of these fibres.

Timber and wood-pulp are naturaily of basic value, but it is not

certain that Malayan trees used for that purpose are the most eco-

nomical that grow on Malayan soils; possibly the Papuan eucalypt

(Eucalyptus deglupta), a rain-forest species, is superior.

Useful plants for specialized purposes, specially for medicinal

use, are still hidden among the native flora, sometimes even in

Malaya itself and the vivid interest in Strophanthus, Rauwolfia,

Kopsia, and other alkaloid-containing plants are contemporary

examples. But the phytochemist cannot work with anonymous

material: he must know the exact name of the plant which he is

analyzing, under what conditions it can grow, and where it occurs.

That information he will ultimately find compiled in the Flora

Malesiana.

This seems enough to demonstrate to non-botanists the value of

that scientific basis for applied work in botany: Flora Malesiana.

I have alluded to above to the heart-warming fact that all former

and recent systematists of the Singapore Botanic Gardens partici-

pate in this great work and that through the farsighted policy of its

director, Mr. H. M. Burkill, it figures on the official programme of

the Gardens. It is evident that for its completion we will need at

least another twenty-five or even more years, depending on money

and manpower of specialists. Though in the past ten years three

thick volumes have appeared the major part of the work still

remains to be done and our sincere hope is that not only retired

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Vol. XVII. (1958).

officers of the Singapore Botanic Gardens will collaborate in an

honorary function, but that the number of active staff members will

be increased for this purpose and brought at least to pre-war

strength.

The eastern tropics have always been much better provided with

botanic gardens than Africa and South America. Unfortunately the

Calcutta Gardens, the oldest, have since long lost their standing as

a scientific botanical research centre and the same can be said of

Peradeniya. In the past few years the Bogor Botanic Gardens have

almost been deprived of scientific personnel and no qualified sys-

tematist is any longer on the staff of the Herbarium, to the great

regret of its director. The Singapore Gardens suffer in less degree

from the same evil of having several vacant posts. I trust that this

situation will be improved in the near future, as whatever high

qualification can be given to a Garden in the way of collections of

living plants, herbarium, buildings, space, and library, its scientific

activity which is decisive for its importance depends on the quality

and size of the scientific staff; first brains, then books, then bricks.

It is this scientific activity which is of the highest importance in

the world of today, and upon which progress depends more than

ever.

With the occasion of a centenary one is apt to look backward;

there we see a dignified growth of an outstanding botanical centre

in the East; gradually as a matter of fact, but all good things grow

slowly.

But we have also to look forward, in the future. There we see a

vista of intensified and extended scientific research to be achieved

for future generations, for their education and welfare, for adequate

use of the land, appropriate utilization of plants and their products,

made possible by critical knowledge and study of them.

The Flora Malesiana is a useful tool towards that aim intended

for the future generations of the eastern peoples. The sooner it will

be completed and be available the more profit it will yield. I trust

that the Administration, convinced of this profit, will find ways

and means to increase Singapore’s contributions towards this goal

and be proud of it.

165

A tale of two cities: Singapore and Leiden

By H. J. Lam

Director, Rijksherbarium, Leiden and

Professor of Systematic Botany of the Government University

I FEEL BOTH pleased and honoured to find myself amongst those

who were invited to contribute towards the Centenary number of

this Bulletin, because “Singapore” (meaning the Botanic Garden)

is one of the important centres of botanical research in insular Asia.

There is, and was, a marked parallel relationship in that Singapore

was, in a way, an affiliation of Kew and Bogor of Leiden. In addi-

tion, there were close bonds between Singapore and Bogor in the

Far East and between Kew and Leiden in Europe.

When speaking of these four places it must be stated that the

relationships between them were, in more recent times, principally

maintained by the respective herbaria, all of which are younger

than the botanical gardens they belong to or are connected with.

And since the present commemoration actually deals with the

garden it may be stated first that in this respect Leiden is the senior

institution, since its botanical garden was created shortly after the

establishment of the University (1575), viz. in 1587. Followed

Kew in 1759 (as a Royal Garden; 1841 as National Gardens),

Bogor (Buitenzorg) in 1817, and Singapore in 1859.

From times immemorial botanical gardens were institutions with

a prevailingly practical aim, viz. the study of medicinal plants. As

far as means of conveyance permitted there was an ever increasing

exchange of material and many gardens organised collecting trips

in order to augment their assortment. In this way a great number

of useful plants, medicinal and otherwise, became known or better

known and in the nineteenth century quite a number of large tro-

pical crops owed their origin and extension to being first studied in

botanic gardens, e.g. pararubber (Hevea), quinquina (Cinchona),

coffee (Coffea), tea (Thea) and numerous others.

In later years when experimental gardens and stations were

created, the original aims of botanical gardens gradually shifted to

these specialised institutions, and while the study of the flora was

transmitted to the herbaria which naturally grew out of collections

of dried specimens of the local flora in and around the gardens

proper, the latter more and more served as open air (and hothouse)

museums of the wealth of plant forms with both decorative and

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a a oo es

RIJKSHERBARIUM

LEIDEN

OTRO ROI LEADS RA IO

The Dutch Herbarium and Cryptogamic Herbarium

An annexe to the General Herbarium

Vol. XVII. (1958).

scientific purposes; decorative so as to indulge in the growing

demand of recreation by town-dwelling people and their natural

craving to admire the wonders of plant life; scientific so as to meet

the increasing necessity of developing sciences such as plant phy-

siology, biochemistry, experimental taxonomy, and the like.

As a matter of course the aims and purposes of the temperate

gardens and tropical ones soon diverged. The first-named ones, in

casu Kew and Leiden, grew next to indigens such plants from as

many different vegetational and floristic areas as they could lay

hands on, and various artificial habitats were created for these

according to their ecological preferences and exigencies. The tro-

pical gardens, i.e. Singapore and Bogor, naturally showed first of

all a display of the surrounding vegetational riches and furthermore

strove for the cultivation of tropical plants other than their own.

In the meantime, as we said, herbaria came into operation. Old

collections of dry plant specimens, some of these as old or even

older than the garden whose property they were or became, were

assembled and arranged. Much new material was added, more and

more intentionally and methodically, as collectors or expeditions

were sent out, and gradually a worldwide system of exchange of

duplicates as well as of borrowing and lending on behalf of mono-

graphic work was brought about.

Whereas thus a wealth of information was obtained, catalogued,

and systematically published, it soon appeared that the floras of

all countries and particularly those of tropical areas whose study

had been tackled at a much later date than that of temperate ones,

was unbelievably richer in every respect than had been long anti-

cipated. It is now recognized that our knowledge of the more inti-

mate details of the vegetable kingdom is still in its infancy. Again

the temperate situation differs from the tropical one. In temperate

countries the higher plants are generally fairly well known specifi-

cally although much work has still to be done on variability, specific

and infra-specific limits, and nomenclature. The study of lower

plants (Thallophyta), however, which started little over one hun-

dred years ago when the microscope got its first modern construc-

tion, is now well under way and shows an unexpected and bewilder-

ing wealth of forms and products, particularly in the fungi.

The tropical countries are, also in this respect, still very much

“underdeveloped”. The causes of this are not only that western

science started its investigation there at a relatively late date, but

that their flora appeared to be infinitely more diversified than that

of e.g. N.W. Europe (which, among its temperate sisters, is parti-

cularly poor as a consequence of the comparatively recent glacial

periods). It has taken some considerable time to arrive at the

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recognition that on studying life phenomena, biologists should have

started in the tropics with their optimal environmental conditions

rather than in any temperate climate. Unfortunately at the time

when western science consciously penetrated in those areas, the

basis of many sciences and their terminology had already been

laid and this has been the cause of the fact that all too long tropical

life has been considered a special case of temperate phenomena,

instead of the reverse. |

Today, our knowledge of tropical plants is still scanty. That of

higher plants, though tolerably good in and near cultural centres

such as in S. Malaya, W. Java, and S. Luzon, is very much inade-

quate regarding such large areas as central Borneo, New Guinea

and a number of remote smaller islands. Practically every explorer

in those parts collects numerous unknown species and is even likely

to discover new genera.

As to the lower plants such as fungi and algae (including kelps),

our knowledge is entirely haphazard and we still have to find our

way in this enormous terra incognita. Everybody knows how much

the life of tropical peoples depends on their surrounding plant

world. But even though their very intimate daily contact with that

plant world is the outcome of an association of tens of thousands of

years, it should not be believed that nothing in it can be improved

and that nothing is to be added. On the contrary. Already science

has revealed many useful particulars of plants so far unrecognized.

A striking example is that of the unpretentious shrub Rauwolfia,

practically unknown only a few years ago, now of extensive fame

as a producer of a very effective remedy against high blood pres-

sure. Numerous other plants containing substances which are sur-

mised to be of potential value to men are still. unknown phyto-

chemically, numerous others doubtless are still to be discovered

and native plantlore may serve as a guide where to look for them.

But also among the lower plants untold riches and possibilities

are undoubtedly stored away. Already certain fungi play a very

important part in native food stuffs but our knowledge of their

processes is extremely scanty and many of them are waiting to be

accurately investigated, cultivated, hybridized and segregated in

order to give them their optimal efficiency.

For all this the botanic garden, its affiliations and its incorporated

institutes, including the herbaria, are there to serve as an indispens-

able tool and medium to be used for the benefit of the people.

At present, the relationships between the four botanic gardens

is rather more platonic than practical, though I do not underrate

the value of occasional exchanges. But the emphasis of the co-

operation clearly lies on the work of the herbaria. Of these again

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Vol. XVII. (1958).

Leiden is leading as far as the date of origin. is concerned. The

Rijksherbarium as it is called—or national herbarium—was founded

in 1829 in Brussels and it was moved to Leiden in 1830. Kew,

now by far the largest herbarium in the world, followed in 1841,

Bogor in 1844 (the date of 1817 mentioned in Index Herbariorum

is that of the garden, not of the herbarium), and Singapore in

1875. Naturally Kew got the best opportunity for a rapid exten-

sion of its collections from all over the British Empire. The Dutch

national herbarium in its modern form was based on the extensive

collection made by BLUME and some others in western Java after

the Dutch had retaken supremacy from the British interregnum in

Napoleonic times. To these some older garden collections were

added, some even older than the university.

During the whole of the nineteenth century numerous collections

were added from the Malay Archipelago as weil as from many other

parts of the world, notably the West Indies. Around the secular turn

the study of the West Indian flora was taken over by the Utrecht

School, founded by PULLE, but the Rijksherbarium received

more and more acquisitions from the Malaysian area, as a conse-

quence of a very close co-operation between Buitenzorg (now

Bogor) and Leiden. Bogor has grown to be a very important tro-

pical herbarium and it must be stated with great satisfaction that

in spite of unfortunate political misunderstandings the mutual insti-

tutional and personal relations with Leiden have remained excellent,

thanks to our colleagues KUSNOTO and DILMY.

After the second world war the Rijksherbarium has again in-

creased rapidly. Next to Bogor it is the recognized world centre

for the study of the Malaysian flora, particularly important because

of the great many types, especially older ones. In addition, after

Berlin had lost its leading position regarding the flora of New

Guinea, Leiden has taken over here, and a new scientific co-opera-

tion has arisen between Lae, Canberra and Leiden. The total

average yearly acquisitions now amount to some 40,000 specimens.

The activity of the Leiden herbarium has greatly increased as

the staff grew to the present number of 17 full-time botanists and

6 honorary fellows. Of the former 8 devote their time to the study

of the Malaysian flora and 4 of them, now still in the service of the

Flora Malesiana Foundation of which VAN STEENIS gives an

account elsewhere in this Bulletin, will possibly be taken over

gradually by the Rijksherbarium now that the Bogor staff, tempo-

rarily I trust, has been curtailed in a most lamentable way.

Naturally the co-operation between the two tropical and the two

temperate institutions has been mutual. Either party would be

greatly hampered in its activities without the assistance of the other.

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Gardens Bulletin, S.

The older collections and types are preserved and best represented

in Europe, more recent ones in the tropics, though duplicates have

been liberally ceded. The indispensable literature is generally more

copiously available in Europe (though Bogor can boast of a re-

markably rich library). Living specimens, on the other hand, can

only be studied in the tropics and the same is true for all characters

which lose their value on the plants being dried.

As has been stated above Singapore has ever maintained a simi-

lar bond with Kew as that which exists between Bogor and Leiden.

Yet the direct relationship between Singapore and Leiden has by

no means been negligible. The exchange of duplicates from 1948

(inclusive) to May 1959 amounts to 2,600 from Leiden and 6,700

from Singapore, material on loan in the same period to 1,300 from

Leiden and 21,000 from Singapore. These figures are significant:

on the one hand they show the main source of the material, and

on the other the place where it is mainly investigated.

Thus, the Singapore Gardens and their valuable herbarium re-

present a rich source of important information and the Rijksher-

barium is proud to be allowed to co-operate with its staff. We on

our side participate in this work out of pure scientific interest in

the overwhelmingly rich tropical flora and vegetation with its count-

less and inexhaustible scientific problems. Our tropical friends on

the other end of the line, who daily enjoy the inspiring neighbour-

ship of their flora, know that ultimately they work to the profit of

their country. To both parties co-operation is a living necessity, for

better and for worse.

On offering the Singapore Gardens our hearty congratulations on

their first centenary, I venture to express the hope that our co-ope-

ration will hold, nay, be intensified for many years to come, to the

benefit of science and the welfare and happiness of the human race.

170

A Note on the Gardens’ Jungle

By I. H. BURKILL

Director, Botanic Gardens, 1912—25

THIS IS AN INCIDENT of 35 years ago at which time search was being

made for a site that could be used for the proposed University

buildings in Singapore, and the Economic Garden has been men-

tioned. On one afternoon I met in the Economic Garden a friend

from Johore who was taking the opportunity of a visit to Singapore

to form an independent opinion. So we walked round the Economic

Garden together and discussed the subject. It was not difficult to

agree that the site, having become surrounded by houses was now

a town site with a value greater than that to which it was put as a

nursery. But when we returned to the Botanic Garden he dropped

a remark which was astounding,—pointing to the Gardens’ jungle

he recommended that it should be used for houses. It was useless,

I found, to point to the beauty of the trees: he could not see it.

He grudgingly consented that these as trees added certain species

to the display in the Gardens; but could they not be seen in plenty

in the forests of the Peninsula? It was quite beyond him to think

of one tree helping another to live; are they not competitors; he

claimed. At this I gave up my pleading.

Those beautiful trees, mainly of three species of Shorea, when

they were but seedlings grew in the shade of others and as they

surmounted the ground-vegetation of their habitat gradually fitted

themselves for drier air and stronger light. Now they are the means

of preventing the death of various tender companions and more

than that they are the front line in the preservation of the humidity

not only of the soil under them, but of the air necessary for the

healthy growth of the palms in the Palm Valley.

The Dell at the head of the Gardens’ Lake furnishes another

illustration of interaction between different items of maintenance

within the Gardens’ boundaries; or the humidity maintained by the

Lake makes the display of the Dell possible.

Any interference with either source of humidity could do more

damage than the man in the street can realize.

The Singapore Botanic Gardens and Forestry

in Malaya

By J. WYATT-SMITH

Forest Department, Federation of Malaya

ALTHOUGH Singapore and the Federation of Malaya are today two

separate countries, and the Singapore Botanic Gardens and the

Malayan Forest Department two entirely separate organisations, it

is, perhaps, not realised except by a few that the first forest depart-

ment in Malaya was that of the Straits Settlements and that it was

formed in 1883 under Cantley, the then Superintendent of the

Botanic Gardens, Singapore. It remained under the Botanic Gar-

dens until 1901 when the administration of forests was handed over

to the Land office. It may also not be realised by many that it is

mainly due to Ridley who became Director of the Botanic Gardens

and Forests in 1888 that Malaya owes the survival of many of the

existing forests in Malacca, Penang, and the Dindings.

During this period the forests of Negri Sembilan, Pahang, Perak,

and Selangor, the former Federated Malay States, were under the

charge of the Land Officers, but in 1895 a full-time Forest Officer

was appointed in Perak. Forestry in Malaya owes another debt to

the Singapore Botanic Gardens when a year later Ridley, in a report

on the forests of Selangor, recommended the establishment of a

properly organised forest department under a forestry trained offi-

cer for the four Federated Malay States. This was unfortunately

not acted on, though special officers were appointed for Selangor

and Negri Sembilan. Four years later in 1900 Hill of the Indian

Forest Service was commissioned to advise on the forest adminis-

tration of the Straits Settlements and Federated Malay States, and

a year later Burn-Murdoch was transferred from Burma to form a

department.

One of the early duties of any forest department is to obtain a

census of the forest resources available both in existing forest re-

serves and on State Land, a census not only of the species that

occur, but of their distribution, frequency, size class distribution,

and subsequently of their ecological characteristics. Such a census

or stock taking is, however, of very reduced value if the identity

of the material is not known and if it is uncertain whether any

material collected is composed of many or a single species. Plant

collecting and the setting up of a small forest herbarium are there-

fore logica] developments, and in 1918 F. W. Foxworthy, a botanist

Lf2

Vol. XVII. (1958).

and wood technologist of international repute who had spent many

years’ service in the Philippines, was engaged as Forest Research

Officer. Determinations were naturally, however, still largely sought

from botanists working in the established herbaria of the world that

were concerned with plants of this region, and particularly those at

Kew and Singapore. The establishment of the Forest Research Ins-

titute at Kepong in 1929 with increased research staff saw the

growth of the departmental! herbarium with particular attention

being paid botanically to this region’s important timber family, the

Dipterocarpaceae. The Institute, however, was still dependent on

other herbaria and particularly Singapore for assistance with deter-

minations; and, in the case of Singapore, which is near at hand, for

reference to their excellent library. It is frequently not realised by

the layman how important these old works are to taxonomists and

also how important it is to have easy access to type sheets or

authentic material. Adequate collections and distribution of dupli-

cate material to many of the well-known general herbaria in the

world and to those specialising on the flora of the area are un-

doubtedly the means of overcoming this, as is readily done today.

But there is no such solution for old plant collections nor for many

of the old books or journals published which, even if they do appear

periodically on the market, are usually so expensive that they are

completely beyond the means of any new herbarium. It might

almost be stated that existing and long established herbaria such as

the Botanic Gardens Singapore, in view of their old collections and

their libraries, have an obligation to the scientific world and parti-

cularly to their region to maintain their activities, since no newly

formed institution can ever hope to obtain the same essential

foundations. And this obligation, we are pleased to say, is fully ac-

cepted and met by the Singapore Gardens.

The herbarium at the Forest Research Institute has naturally

specialised in woody species and is really no more than a reference

herbarium for the Forest Department. Up to World War II interest

was focussed particularly on the dipterocarps, and a collection has

been built up, mainly through the efforts of Symington, which apart

from early material and types is possibly the best in the region.

Since World War II the emphasis has been on the other important

timber producing families and on many of the smaller woody plants,

the latter being required in connection with ecological studies of

rain forest. Many of the final determinations and all the preliminary

determinations are being done at the Institute, but continued refer-

ence is still made to the herbarium and assistance required from the

qualified staff of the Singapore Botanic Gardens. It is certainly not

173

Gardens Bulletin, S.

envisaged that the Forest Research Institute with its present bota-

nical facilities can ever dispense with the resources of the Botanic

Gardens, Singapore, and it is earnestly hoped that it will be found

possible to maintain the herbarium, library, and staff of the Gar-

dens at the highest level for the next hundred years as has been so

well done during the past century.

Apart from the early guidance in direct administration of the

forests of the former Straits Settlements, the subsequent assistance

with determinations and permission to use the library and herba-

rium, the Forest Department has also benefitted in the extreme

interest shown by these early and subsequent botanists in the locai

flora and the large number of publications concerning trees and

forest produce that they have published. Works that immediately

spring to mind and with which all Malayan forest officers are

acquainted are Ridley’s Flora of the Malay Peninsula which was

based on King and Gamble’s Materials for a Flora of the Malay

Peninsula; Burkill’s Dictionary of the Economic Products of the

Malay Peninsula; Corner’s Wayside Trees of Malaya; Holttum’s

works on orchids and on ferns which form Volumes I and II of

the new Flora of Malaya; many articles in the Gardens’ Bulletin

including Holttum’s works on bamboos and gingers, Furtado’s

work on the rattans and canes, Sinclair’s on the two tree families

Annonaceae and Myristicaceae, Henderson’s on the important

timber genera Calophyllum and Eugenia; and MHenderson’s

Malayan Wild Flowers published by the Malayan Nature Society.

In addition numerous vegetational papers by members of the Bota-

nic Gardens’ staff have appeared in such Journals as that of the

Royal Asiatic Society and the Federated Malay States Museum,

papers which are of great interest and value to forest officers. We

may mention here that the traffic has not been all one way, for

Departmental botanists, particularly Symington in connection with

dipterocarps, have contributed to the Bulletin of the Singapore

Gardens. And some of the series of Malayan Forest Records are

valuable items in the botanical literature of Malaya.

174

The Singapore Botanic Gardens and

Rubber in Malaya

By P. R. WYCHERLEY

Botanical Division

Rubber Research Institute of Malaya

NATURAL RUBBER PRODUCTION in Malaya and other tropical Asian

countries depends upon the cultivation of Hevea brasiliensis. This

tree was introduced into the East and its exploitation developed

through the agency of various botanical institutions and, as will be

shown below, the part played by staff of the Singapore Botanic

Gardens has proved exceptionally important.

The changing fortunes of the rubber producing industry have

reflected the trends of world economic history during the first half

of the twentieth century. Although during times of trade depres-

sion the plantation industry has done little more than provide a

survival existence for its many employees, on balance natural rub-

ber production has made one of the greatest contributions to pros-

perity in Malaya. This has been achieved in the first place by the

attraction of capital and latterly because the industry has become

the largest single source of employment and revenue in the Fede-

ration of Malaya. At present rubber plantation employees number

over a quarter of million and it is estimated that nearly 400,000

smallholders are supported wholly or partly by rubber cultivation.

Direct taxation of rubber exports has provided about 15 per cent

of the total Federal revenue during the last five years. This figure

was swelled by taxes on company profits and by other less direct

means.

_ This enterprise, which has given a livelihood to a large section of

the population representative of all races in Malaya and provided

wealth for the country’s development, owes its origin mainly to

the Singapore Botanic Gardens.

The Introduction of Hevea to the East and Establishment in

Malaya

The distribution of potential crop plants to enhance the utilisa-

tion of natural resources was a conspicuous feature of the work of

the Royal Botanic Gardens, Kew, and its daughter institutes while

Joseph Hooker was Director. He was knighted in 1877 and in

1897 honoured for his editorship of the Flora of British India.

[75

Gardens Bulletin, S.

Mr. James Collins had reviewed the rubber producing species of

plants at the request of Sir Clements Markham of the India Office,

the latter recommended the collection of Hevea from Brazil and its

despatch to oriental territories, and Hooker implemented this

recommendation.

Although several attempts were made under Hooker’s guidance

or by the India Office, only one—the famous Wickham collection—

succeeded. The others failed either because the seed did not ger-

minate or owing to the selection of unfavourable reception sites

such as Calcutta and Sikkim where the plants died.

The success of the Wickham collection owed much to the ex-

peditious handling of the material. Joseph Hooker commissioned

Mr. H. A. Wickham (later Sir Henry) to collect seed, which he

did in the area between the Tapajos and Madeira rivers of the

Central Amazon basin. Wickham chartered a cargoless ship to

rush his 70,000 seeds to England. At the expense of digression

from the main theme the charge against Wickham of smuggling

should be refuted here. Although he may have disbursed hospitality

to speed official clearance of his perishable cargo, there is no

evidence that he resorted to corrupt evasion of the law; in fact the

shipment of seeds of Hevea from Brazil was not prohibited at that

time and an official statement—Borracha no Brasil, Rio de Janeiro,

1913—reported that Wickham’s and other exports were made

with the goodwill and co-operation of the Brazilian government.

The seed reached Kew on 14th June, 1876 and Hooker ensured

careful treatment; he obtained about 4 per cent germination,

which was good considering the long journey and that it was un-

known how the seed should be packed or treated. The bulk of

the seedlings were despatched in Wardian cases (miniature green-

houses) to Ceylon, where extensive nurseries had been prepared

at Henaratgoda on the orders of the India Office. Further shipments

from Kew in 1876 were 18 plants to Java and 50 to Singapore,

but the latter arrived at the Gardens in poor condition and the

five survivors appear to have died subsequently. The following

year 22 plants were sent from Kew to Singapore and successfully

established by Mr. J. H. Murton, Superintendent of the Gardens.

The distribution of these plants is not fully known. Murton went

plant collecting in Perak during October to December 1877, when

he planted nine Hevea seedlings at Kuala Kangsar behind the old

Residency on the bank of the Perak River. The Resident was

Mr. (later Sir) Hugh Low, who, like Sir Stamford Raffles, was an

active naturalist and very interested in economic botany. Murton ‘is

reported to have planted one of the ‘Wickham’ seedlings with other

rubber bearing plants at Durian Sabatang. This may have been the

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Vol. XVII. (1958).

Durian Sabatang near Telok Anson, but his intinerary did not

include Lower Perak unless he made an unrecorded call during

the sea voyages between Singapore and Penang. He passed near

the Durian Sabatang about twenty miles south of Matang and

Taiping. Both these places together with Telok Anson have been

quoted as sites for one or more of the first introductions, probably

all referring to the single plant, which appears to have been soon

lost for no traces could be found by Low in 1878. At least nine

seedlings were retained at Singapore. The important point is that

Murton established Hevea brasiliensis in two centres where the

trees could be tended and observed, namely, at the Singapore

Botanic Gardens and under Low’s good care at Kuala Kangsar.

The plants at Singapore were multiplied to a limited extent by

cuttings. However, Murton found, as has been confirmed repeat-

edly since, that only cuttings from young seedlings rooted easily.

Thus there was a limit to the amount of material which could be

propagated by this means. Nevertheless it is interesting and to

Murton’s credit that he was alive to the potentialities of the method

and investigated its possibilities. The most suitable site was sought

by Murton, his successor, Mr. N. Cantley and Mr. W. Fox, and,

after two transplantings, the Hevea collection found a home in the

newly acquired Economic Garden in 1879. Unfortunately the

Government provided no funds for the upkeep of the Economic

Garden and scrub invaded; thus by 1885 the trees were in a poor

state of upkeep. The numbers were reinforced from 1884 onwards

with the progeny of the originals and new imports from Ceylon.

This was most important as these provided the material for the

classic experiments by Ridley and Derry.

Meanwhile the trees at Kuala Kangsar had flowered in March

1880 and set seed the following year. Mr. (later Sir) Frank Swet-

tenham, acting for Low in 1884, planted out about 200 seedlings

on the banks of the Perak River and on the hill behind, where the

new Residency was built. A group of seedlings was planted around

the grave of British soldiers killed in the Perak expedition of 1876.

These proved a valuable source of seed in later years.

The Search for a Means of Exploitation

The first tapping in Malaya was on the Kuala Kangsar trees by

aborigines; they used an unsuccessful incision method probably

similar to that employed to extract poisonous sap from the Ipoh

tree. It is not clear when this was done, as some of the early lite-

rature is contradictory and includes evident typographic errors in

the dates. Low is reported to have sent rubber samples to London

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Gardens Bulletin, S.

in 1884, which were not received favourably, but these may have

been Ceara Scrap (from Manihot glaziovii, which was also planted

at Kuala Kangsar); at any rate the confusion did not enhance the

reputation of cultivated rubber. Swettenham may have directed an

attempt in 1886 and—or—Low an attempt in 1888.

Another attempt was made in 1889 when Swettenham had suc-

ceeded Low and the results were again disappointing. Swettenham,

having decided that there was no future for Hevea in Malaya,

ordered the destruction of several of the original Kuala Kangsar

trees and discouraged further interest. This remained the view in

government circles for some years.

The year 1888 marked the arrival in Singapore of Mr. H. N.

Ridley as Director of the Botanic Gardens. He shared Hooker’s

faith in natural rubber production by Hevea cultivation and, des-

pite government opposition and lack of funds, set about rehabilita-

tion of the Economic Gardens and experimented to find a practical

means of extracting latex. As early as 1889, while Swettenham

made his gloomy predictions, Ridley made his first basic discovery.

Hitherto virtually all forms of and attempts at tapping Hevea

for latex were incision methods, by which a cut was made through

the bark but no bark was removed. (We use bark here in the

popular sense to denote all tissues outside the wood). Incisions

tend to close, there is no proper channel for the trickle of latex to

flow along into a receptacle and it is hard to judge the depth of

the cut—the result is much wounding of the tree.

The excision method invented by Ridley involved removal of a

paring of bark from the cut, which in the first place overcame the

main defects of incision noted above, but lead to the further dis-

covery that by repeated paring away of bark from the one cut at

intervals a greater flow of latex could be stimulated. Rubber ob-

tained by this method was exhibited at the Singapore Agricultural

Exhibition in 1890 and samples were sent to London the next year,

where they were pronounced excellent. Other members of the

Gardens staff joined in the tapping experiments, C. Curtis in 1896

at Penang, L. Wray in 1897 at Taiping and R. Derry in 1898 at

Kuala Kangsar, whence rubber was sold in 1899; the various

botanic gardens in Malaya were then all under the direction of

Singapore.

In 1904 Derry joined Ridley in Singapore and together they

conducted a series of experiments, which established a number of

principles and carried the technique far towards that currently

employed. Mr. A. D. Machado and Mr. C. Boden Kloss assisted

in the experiments and yield recording during some years. Only

the main trends are described here to illustrate the progress made.

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Vol. XVII. (1958).

The first tapping was done with a pruning knife, chisel and

mallet but these were superseded by a modified farrier’s knife or

gouge, a form of which was adopted on one of the first plantations

—Jebong Estate—and the “Jebong” is still among the most useful

tapping implements. The advantage of morning over afternoon

tapping was first established at Singapore. The relative perform-

ance of wide- and close-planted trees was investigated; the conclu-

sion was reached that a relatively wide spacing giving a stand of

150 trees per acre would give more economic tapping and a longer

useful life than much denser plantings.

The arrangement of the cuts fell into two patterns: (i) herring

bones, in which sloping lateral cuts branched upwards from a

central vertical channel and (ii) spirals encircling the tree several

times. The Singapore workers soon abandoned the more fanciful

systems, and although they do not seem to have reached the ulti-

mate simplifications, i.e. neither reduction of a double fishbone to

a single V-cut and of a half fishbone to a single half spiral, nor

restriction of the number of turns in a helical cut to one full or half

circumference cut as now, their conclusions were moving in that

direction.

At first there was much apprehension that if tapping were con-

tinued too long, the bark would not renew over the ever extending

area of excision. Ridley and Derry grew bolder as their experience

confirmed that continuous tapping—indefinitely repeated excision

at regular intervals-—was possible, the bark renewed and moreover

greater yields were induced. The simplification of the original her-

ring bone was largely to allow adequate time (four years was at

first thought sufficient) for bark renewal before one cut encroached

on the next under continuous tapping. They compared daily and

alternate daily tapping and expressed a preference for the latter.

Changing Interests and the First Plantations

The tapping experiments described above continued until 1909

and we must revert to the period immediately after Ridley’s dis-

covery of a practical method of exploitation to carry on the general

history of the subject. The depredations of disease were rapidly

making coffee cultivation uneconomic and the owner-managers of

the first estates were looking for alternative crops. At first Ridley

was still a voice crying in the wilderness as far as rubber was con-

cerned, some of his critics referring to him openly as ‘mad’ Ridley.

Although some planters such as Mr. Hislop Hill and the Kindersley

brothers continued to plant scattered trees of Hevea for observa-

tion and to provide seed should the crop realise its potentialities,

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Gardens Bulletin, S.

the first plantation development was the planting of 40 acres of

Hevea by Mr. Tan Chay Yan at Bukit Lintang. There is some con-

fusion about the date of this. Ridley has stated 1896 but some

later authors quote 1898. One of the Kindersley ventures in 1895

is reputed to have been virtually a plantation of Hevea. Mr. Tan

exhibited rubber in 1898, presumably prepared from his older

parent trees and not from the 40-acre block. At any rate this was

the turning point; there had been little interest before but now

Mr. Tan went ahead with a larger planting at Bukit Asahan and

many other planters followed suit.

The decline of coffee cultivation, the increasing demand for and

price of rubber on the world market were important factors:

nevertheless Ridley’s discovery of excision tapping and persistent

propaganda were essential contributions.

We must note here that Murton’s choice of two sites, Singapore

and Kuala Kangsar, for the first introductions had several impor-

tant results relating to the rapid expansion of plantations. Mr.

Cross, who made an unsuccessful shipment of seed from Brazil to

the East, had stated that damp, swamp soils were the best for

Hevea, especially land subject to flooding. Wickham had in fact

collected his seed from an undulating plateau probably with a well

drained soil, which he confirmed in 1902. However, unsuitable

water-logged sites would probably have been chosen for many years

if Murton had not established the Kuala Kangsar trees on what

would now be recognised as a typical soil for Hevea, i.e. moist but

well-drained. Thus the range and suitability of habitats for Hevea

in Malaya was demonstrated by this and subsequent early plantings

at Kuala Kangsar, Taiping and elsewhere by Low, Swettenham and

Hill.

Furthermore these trees provided a local source of seed for

North Malaya, some of which were planted by Hill in Negri Sem-

bilan, and in 1893 he had seed available in quantity. Seed was des-

patched to the many new plantations in Malaya and other terri-

tories; Ridley had found that the best packing for long distance

travel was damp charcoal in sealed biscuit tins.

Research and Advice

The new development raised many problems, first in cultivation

and later in processing of the crop—in other words there was a

need for research and advice. This had been anticipated by Ridley

and his assistants, who had devised means of coagulating latex for

shipment and studied conditions of tree growth and many diseased

specimens.

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Vol. XVII. (1958).

As early as 1891 the first Agricultural Bulletin had been pub-

lished; but owing to lack of funds and delays at the Government

Printing Office this did not appear regularly until 1901. When

Ridley was preparing to depart in 1911 and it appeared that publi-

cation might cease, the Planters’ Association of Malaya pleaded

that his successor should continue it, remarking that ‘Ridley’s con-

tributions have been highly valued by the Planting Community and

it would be a distinct loss if the Bulletin were discontinued.’ Feeling

towards ‘mad’ Ridley and the other rubber enthusiasts among his

colleagues had changed considerably over twenty years. When leav-

ing Malaya Ridley left pleas in the Agricultural Bulletin and

Greniers Rubber News for an expansion of research and advisory

services.

By this time the Department of Agriculture F.M.S. (founded

1905) had assumed responsibility for investigations on rubber,

although there had been a transition period during which the ex-

perience and current experiments of the Gardens’ staff were avail-

able until the new Agricultural staff were conversant with the

problems. For instance the Singapore Botanic Gardens was for

many years among the few institutions with an adequate number of

trees in tapping for experiments on the preparation of rubber from

latex; Ridley had made the first ‘biscuits’ of rubber coagulated in

shallow plates and Derry continued for many years to make im-

provements in processing, including coagulation, sheet and crepe

manufacture.

We will review here only the contributions made by the Gardens’

staff—in particular Ridley’s. There was activity along parallel lines

elsewhere, particularly in Ceylon and Java, the importance of

which we do not wish to ignore or gainsay; moreover the Gardens

enjoyed the collaboration of Mr. P. J. Burgess, the Government

Analyst, who made several contributions on tapping, coagulation

and processing. Nevertheless the extent of activities and number of

first records by the Gardens’ Staff adequately illustrate their pre-

eminence in this field.

Pests and diseases are always among of the first difficulties en-

countered in new crop culture. Termites were the first insects to

trouble Hevea in Malaya; apart from ‘plantation hygiene’-—which

was only partly helpful—Ridley was unable to solve this problem.

However, the full control of termites has had to await the dis-

covery of modern insecticides. Crickets and mites appeared in due

course; for the latter Ridley and Derry recommended sulphur

dusting.

The white and brown root diseases of rubber were first described

in 1904 and 1909, respectively, by Ridley who advocated detection

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Gardens Bulletin, S.

and destruction of sources of infection. He reported on ‘pink’ dis-

ease of the stem and branches in 1905 and advised destruction of

diseased tissue and treatment with copper sulphate and lime. Birds

eye spot was recorded as an emphemeral disease of young seed-

lings in 1906. The present-day planter will be struck by the small

changes in the principles of disease control recommended today

and those suggested by Ridley.

The first report on the deleterious effects of applying lime to

Hevea was made by Ridley in 1903, when he conducted a

manurial trial.

Prophecies and Warnings

The early planters were, almost without exception, keen advo-

cates of clean weeding; the majority would not tolerate any catch

crop which might deter from the main crop nor any cover crop for

soil protection. During the first decade of the century many plan-

ters had adopted the practice of eradicating all plant growth by

spraying sodium arsenite repeatedly. Ridley saw further and espe-

cially in his last editorials addressed serious warnings on the risks

of soil erosion to the plantation managers. He suggested chillis,

pineapples, bananas and bowstring hemp (an agave) as possible

catch crops, which—except the last—can all be seen put to good

use on small properties today especially by Chinese. Tapioca, the

catch crop favoured by the minority group of ‘maximum exploita-

tion—minimum time and effort’ planters, was considered by Ridley

to be liable to exhaust the soil and to spread root disease; therefore

he advised caution and experiment before using tapioca as a catch

crop, which attitude has been amply justified. In 1910 Ridley pro-

phesied that contour terraces would be necessary to control erosion

on slopes and that cover crops such as the local wild Crotalaria

would be needed for soil protection and enrichment. He realised

that not all lands were suitable for rubber cultivation and con-

demned planting Hevea on deep peat soils.

We note too that Ridley advised against cutting out coconuts in

order to plant rubber, stating explicity that neither individual

estates nor the country should rely exclusively upon one crop. He

was one of the first to suggest the oil palm as a plantation crop and

distributed over 3,000 seeds.

There have been and there are—as there should be—critics of

the unbalanced state of Malayan agriculture, meaning in particular

the predominance of and dependence on rubber. Critics of the

plantation industry have pointed to clean weeding and other bad

practices of the early planters, whose rapacious exploitation of the

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Vol. XVIT. (1958).

soil has bequeathed us many problems in its rehabilitation. Never-

theless, these critics should realise that the practical founder and

advocate of the rubber plantation industry—Ridley—warned

against these errors; indeed the slump of 1913 came as an early

reminder of the risks involved in dependence on one crop, but

neither the sound business men nor officialdom took much heed.

Ridley was ably assisted and—perhaps to greater extent than we

realise—guided by a team of Gardens’ staff in Singapore and

throughout Malaya, who were all devoted to the study of plants and

fully appreciative of their value in preserving and developing the

resources of the land they had come to serve.

We come now to the close of the period during which the Singa-

pore Botanic Gardens were directly responsible for the develop-

ment of the rubber industry in Malaya. Mr. I. H. Burkill became

Director in succession to Ridley in 1912. Rubber development had

then become entirely the responsibility of the Department of

Agriculture. Yet Burkill maintained tapping of the Hevea at

Singapore and yield recording of the individual trees, so that the

most productive could be selected as seed parents for future im-

provement of planting material.

The Present Situation

In 1926 the Rubber Research Institute of Malaya was created

by enactment and in turn took over responsibility for rubber. The

Botanic Gardens have continued with the basic studies of syste-

-matic research into the Malayan flora and the nature of the vegeta-

tional formations in the region. It might seem that we have gone

our separate paths and the Botanic Gardens are of no further

service to the rubber industry, but such a view ignores one of the

greatest problems still lacking a solution. We need the help of the

fundamental botanist, who is found in institutions such as the

Botanic Gardens, which we wish well—not for sentimental reasons

because of the great contributions of the past—but in expectation

of further assistance in the future.

Malaya and most of the other countries where Hevea is cultivated

lie in the region of the tropical rain forest. This type of vegetation

is very complex; frequently over one hundred different tree species

are found in a single acre of Malayan jungle with even more

species amongst the epiphytes, lianes and ground flora. The rain

forest has developed and perpetuated itself for millenia, but the

whole formation is now rapidly disappearing under the pressure of

man’s advance before he has discovered how it functions.

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Gardens Bulletin, S.

This state of affairs has been frequently acknowledged recently,

for instance at the Ninth Pacific Science Congress, Bangkok, 1957,

at the Centenary and Bicentenary Congress, University of Malaya,

Singapore, 1958, and at the UNESCO Symposium on Humid

Tropics Vegetation, Bogor, 1958. Every appreciation of the situa-

tion emphasises that the effects of rain and insolation on exposed

tropical soils can lead to rapid impoverishment and erosion.

Phosphate availability in most Malayan soils is notoriously low and

other nutrients are easily leached out of disturbed soil; there are

indications that these can be alleviated or prevented by a suitable

vegetational cover which maintains the organic content of the top

soil. Such vegetation controls physical erosion and through its

many members circulates nutrients also.

Copious rainfall, constant warmth and generous irradiation with

sunlight are all factors which contribute both to rapid plant growth

and to rapid soil degradation. The success of the tropical rain

forest derives from its ability to use the benefits of these factors to

provide a luxuriant vegetation on generally poor soils, despite the

deleterious potentialities due to these same factors. The object of

agriculture in the tropics is to produce a luxuriant crop without

soil degradation.

Although the agriculturalist can apply fertilisers to make good

losses and deficiencies and can use physical means such as terraces

and water channels to control erosion and water distribution, basic-

ally he has to solve the same problems which the forest has sur-

mounted. This will probably be achieved most effectively and

economically by the same means as those which are successful in

nature; it is unlikely that totally different principles will succeed

where we must contend with tropical climate, poor soil and undu-

lating terrain.

A single crop culture, even in association with a cover crop

which is often limited to a narrow spectrum in floristic composition

and growth form, is unlikely to perform all the functions carried

out by the natural forest vegetation. Plantation crops such as

rubber will continue to be grown in stands of one crop because

this enables more economic cultivation and harvesting. Neverthe-

less there is a need to enrich the ecological association of these

crops by introducing into the cover further elements which can

fulfil the processes of soil conservation. Currently we are approach-

ing this problem by growing various plants in association with the.

main crop and endeavouring to assess their effects upon the main

crop and upon the soil. However, we would be better guided in

our choice of plants and factors to study if we knew how the natural

tropical rain forest functions.

184

Vol. XVI. (1958).

The Future

It is often stated that the future of the natural rubber industry

depends upon reducing the cost of production; nevertheless if the

existing advances, especially in the selection of planting material of

improved yield, are exploited fully, as is attempted in the current

replanting programme, a production cost competitive with syn-

thetic rubber can be achieved. Further research will be necessary to:

keep the cost of production down to this level despite rising wage

trends and possible cheaper synthetic production. In the preceding

section we envisaged an ideal tropical agriculture harnessing the

natural processes of the forest; such a system for rubber cultiva-

tion would doubtless form a valuable contribution towards the long.

term reduction of production costs, but would moreover preserve

the fundamental asset—the soil itself. Investigations on the func-

tioning of the tropical rain forest have scarcely been essayed, yet

this is a most important field both for its scientific interest and in

order to evolve a rational form of agriculture for land utilisation

in the tropics. The scientific manpower available is limited but the

destruction of the forest is continuing apace. No one institute can

tackle this problem alone, because the facilities built up over long

periods such as herbaria, experimental gardens and forest reserves

of known history and laboratories proper to different institutes are

all needed. Moreover scientists trained in various disciplines who

have experience in institutes devoted to every branch must co-

operate in this undertaking.

The Departments of Forestry and Agriculture in the Federation

of Malaya are daughters of the Singapore Botanic Gardens and the

Rubber Research Institute is in effect a grandchild. Within this

family there is scope to tackle the fundamental problems of tropical

ecology; may the centenary of the Singapore Botanic Gardens be

marked by revived interest in this subject. The difficulties which

we have inherited and the problems we may bequeath owing to

poor land management will be best solved and prevented by this

means.

References

Accounts of the introduction and early history of Hevea in

Malaya will be found in:—

(1) A Dictionary of the Economic Products of the Malay

Peninsula, Vol. I, p. 1143 by I. H. Burkill. London,

1935.

(2) Journal of Heredity, Vol. XIX, p. 193, 204 and 485.

1928.

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Gardens Bulletin. S.

(3) Various articles in:—

(a) Agricultural Bulletin of the Malay Peninsula,

Singapore, 1891-1900.

(b) Agricultural Bulletin of the Straits and F.M.S.,

Vols. I to X. Singapore, 1901-1911.

An index to these publications, including articles on

research and advice on Hevea cultivation, will be

found in:—Gardens Bulletin S.S., Vol. I, p. 48.

Singapore, 1912 (which was originally published as

Agric. Bull. S. & F.M.S. Third Series, Vol. I.. 1912).

(4) Gardens Bulletin S.S., Vol. I, p. 247. Singapore, 1912.

Biographical notes on botanists resident in Malaya, including the

itinerary of Mr. H. J. Murton’s journey, may be found in:—

Flora Malesiana, Series I, Vol. I. Djakarta, 1950.

The climate of opinion among early planters may be judged from

minutes of their meetings given in the Agricultural Bulletin, also

by the correspondence and reports in Grenier’s Rubber News,

Kuala Lumpur, 1909 et seq.

The proceedings, when published, of the undermentioned may

be consulted for accounts of the significance of the tropical rain

forest in land utilisation today.

Ninth Pacific Science Congress, Bangkok, 1957. (Especially the

Symposium Climate, Vegetation and Rational Land Utilization in

the Humid Tropics Brief review in:—Planters’ Bulletin R.R.I.M.

No. 35, p. 25, March 1958).

Centenary and Bicentenary Congress, University of Malaya,

Singapore, 1958. Bulletin Raffles Museum, in press, (especially

Dr. Dwight Davis on ‘ The naturalist in the tropical rain forest’).

Symposium on Humid Tropics Vegetation, Tjiawi, Bogor, Indo-

nesia, 1958. Sponsored by UNESCO and the Council for Sciences

of Indonesia. |

Aspects of the Malayan natural rubber industry’s position are

reviewed in:—

Federation of Malaya Annual Report, 1957, Kuala Lumpur,

1958.

Planters’ Bulletin R.R.I.M. No. 36, p. 49, May 1958.

186

The Contribution to Agriculture in Malaya

by the Singapore Botanic Gardens

By a Malayan Agriculturist

A GARDEN: A piece of ground where flowers, etc. are cultivated.

A pleasant spot. (Chambers’s Dictionary).

THE SINGAPORE BOTANIC GARDENS are a very pleasant spot indeed

and a good place to relax. So much so, that the casual visitor would

have no idea of their importance to Malayan agriculture and the

valuable scientific research that is carried on there.

It has been said* that the experimental and scientific work in a

botanic garden “cannot by its very nature be directly remunerative

but may indirectly repay its cost many times by establishing new

industries, by saving old ones from losses by disease or otherwise,

introducing new and improved methods and so on”. The validity

of this statement is well demonstrated by development of the

rubber industry in Malaya, but even if this were not so, even if

rubber had failed to grow here at all, the Botanic Gardens have

made many other less spectacular but nevertheless notable contri-

butions to Malayan Agriculture.

In the first place it should be recalled that when the Gardens

were taken over by Government in 1875, it was intended that they

should be concerned with agriculture and the economic develop-

ment of the Straits Settlements and not merely a pleasure garden or

a base for collecting material for the Botanic Gardens at Kew in

England. The Director of the Gardens was also the Director of the

Forest Department and regarded, by the Government at least, as

the pre-eminent authority in Malaya on anything whatever con-

cerned with plants.

Starting in 1891 a Bulletin dealing with Agri-Horticulture, and

other cognate subjects, was published by the Gardens and this was

issued as a monthly journal from 1901 to 1911. (Agricultural

Bulletin of the Straits and Federated Malay States), Opening

Volume I of this Journal at random one may find articles on rubber

cultivation, a series on the Timbers of the Malay Peninsula, Trade

and Market Reports and, among the correspondence, a letter from

Mr. Carey claiming that coconuts will yield well on the coastal clay

soils of the West Coast. Even today, the scientific reports are of

* J. C. Willis, 1907.

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Gardens Bulletin, S.

value and although the price of opium (11s. 6 d. per lb., spot)

and Mr. Carey’s well justified optimism are only of historical

interest, one can appreciate how this Journal must have made up

for the lack of communications and expert advice to which we are

now accustomed.

The Bulletin continued to be published even after a Department

of Agriculture for the Federated Malay States was set up in 1905,

but its name was changed to the Gardens Bulletin in 1912 when

the F.M.S. started to produce its own agricultural magazine. In

1919, the Department of Agriculture, F.M.S., also took over the

responsibility for agriculture in the Straits Settlements, and the

Gardens Bulletin gradually began to put less emphasis on practical

agriculture until 1924 when this aspect of the Garden’s work finally

came to an end.

This is not to say that the Gardens’ staff were no longer interested

in agriculture. The Economic Gardens of 102 acres had been

opened in 1879 and used as a preliminary trial area for numerous

local and introduced plants which were considered to have some

potential value. In addition to rubber, it is worth noting that the

first oil palms in Malaya were planted there about 1875 and an

excellent review of the possible value of these palms in Malaya

was published in the Bulletin in 1907. The Economic Gardens

were unfortunately closed in 1925, but despite this I. H. Burkill

was able to publish his Dictionary of the Economic Products of the

Malay Peninsula, which is still in every-day use as a reference book

at the Department of Agriculture in Kuala Lumpur and all over

the world. Other books which have been invaluable to agricultural

research and have an international reputation include the floras by

Ridley, Holttum and Henderson and the well illustrated two

volumes on Wayside Trees of Malaya by Corner.

It is our good fortune that the work of the Gardens has been so

well documented but no library, however comprehensive, could

replace the collection of living and preserved material which is

available there. The value of these specimens to the teacher, the

student and amateur gardener need not be mentioned here, but to

the scientist from tropical and sub-tropical countries, the herba-

rium type specimens and the easily accessible living material, form

an invaluable point of reference. For instance, when the Depart-

ment of Agriculture was recently anxious to learn the types and

distribution of citrus trees growing in Malaya in order to prepare a

scheme to improve the planting material of this crop, the Plant

Breeder first visited the herbarium at the Gardens and was not only

able to gather much of the information he required but was also

able to clear up some difficulties over citrus taxonomy in general.

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Vol. XVII. (1958).

Difficulties are, in fact, quite frequently encountered by anyone

concerned with taxonomy and since this is a science in itself there is

a small traffic of specimens from Kuala Lumpur to the Gardens

for identification.

It would perhaps be appropriate to emphasise here how useful it

is for the agriculturist to know the correct botanical name for a

plant he is growing. It would be useless for anyone wanting oil

palm seed to write to West Africa asking for seed of “kelapa bali”

or “kelapa sawit” but if the palm were given its accepted botanical

name, Elaeis guineensis, the type of seed required would be clearly

understood. Similarly root-disease of fruit may be due to a number

of causes, but if the plant pathologist can isolate and identify, say,

the fungus Phytophthora citrophthora from the diseased roots, he

can at once be almost certain that he has found the cause of the

trouble, and can consult the literature describing methods for con-

trolling this same fungus in other parts of the world.

Until, perhaps, electronic translating machines are invented, it is

too much to hope that scientists throughout the world will publish

all their works in a universal language. For the moment we must be

thankful that Linnaeus invented his binomial system for naming

plants and animals. Without this system, and the support of institu-

tions such as the Singapore Botanic Gardens, which are mainly

responsible for its propagation, there is no doubt that agriculture

would be immeasurably the poorer.

189

Orchids, Gingers and Bamboos: Pioneer work at the

Singapore Botanic Gardens and its significance for

Botany and Horticulture

By R. E. HOLTTUM

Director, Botanic Gardens, Singapore 1925-1949

A BOTANIC GARDEN is essentially a museum of living plants main-

tained for botanical study. Out of that study comes new knowledge

of various kinds, much of which can find an application in horti-

culture.

The maintenance of any garden involves thought about horti-

cultural technique; and the maintenance of the great variety of

plants in a botanic garden should result in experiment in techniques

and in new understanding of the diverse cultural needs of the

many plants concerned. Thus in a botanic garden there is a two-

way traffic of knowledge and its application; new botanical know-

ledge finding applications in horticulture, new horticultural tech-

niques aiding the cultivation of plants for botanical study.

A botanic garden may serve also as a park, but that is not its

primary function. It should be the source and storehouse of new

knowledge, both horticultural and botanical, which can find its

application in parks and in private gardens; and without that con-

stant pursuit of new knowledge the parks and gardens lose the

impetus to progress.

Botanical study possible in botanic a due may be divided

broadly into two aspects: study of the structure and classification

of plants, and experimental work of various kinds. The study of

classification is basic to all other, and I would regard it as the

primary function of a botanic garden. Such study may seem very

theoretical; but it has immediate application in plant breeding and

also in other ways. Plant breeding is one kind of experimental

work which may be undertaken at a botanic garden; it can pro-

vide new kinds of plants for horticultural use. Other kinds of ex-

perimental work are concerned with the physiology of plants, and

such may result in new knowledge useful in horticultural techniques.

The orchid family is the largest plant family (in number of

species) in the Malayan region, and includes many beautiful plants

which are valued horticulturally. Botanically, the great variety and

complexity of floral structure presents very intricate problems to

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Vol. XVII. (1958).

the classifier. The botanist who explores the forests of Malaya for

orchids (or other plants) cannot make a final study of them as he

finds them; he needs to take them away for more detailed examin-

ation at leisure. He may either preserve them as herbarium speci-

mens, or he may take living plants for cultivation. The herbarium

at the Botanic Gardens, Singapore, is thus a vast storehouse of

information, much of which has still not been thoroughly investi-

gated. For orchids it is much more satisfactory to have living

plants in cultivation, and to examine the fresh flowers, as dried

ones lose some of their characteristic shape as well as their colours.

Mr. H.N. Ridley, who was Director of the Gardens from 1888

to 1912, had made a special study of the orchid family before his

arrival in Singapore, and during his many travels of botanical ex-

ploration he brought to the Gardens a large number of orchid

plants, many of which he described as new species, preserving also

specimens of them. He also had the services of an excellent artist

(Charles de Alwis) who made accurate coloured drawings of the

plants and their flowers. Thus the records of Mr. Ridley’s study

of native Malayan orchids are preserved in Singapore in the form

of specimens, illustrations and descriptions. Upon these records

depends stability in the use of names; if a name is to be useful it

must always mean the same thing.

Similar work on orchids was carried out in neighbouring coun-

tries (India, Siam and Indo-China to the north, and in Indonesia).

In order that there shall not be confusion and duplication in the use

of names, it is necessary to correlate study of plants in Malaya

with those in the surrounding countries (where many of the same

species occur, along with others). This has been achieved by the

accumulation in Singapore of a valuable botanical library and by

exchange of specimens.

The most important later collector and describer of new orchids

in Malaya was Mr. C. E. Carr, a rubber planter who made this

work his chief recreation. He used the herbarium and library at

the Botanic Gardens, Singapore, and contributed specimens which

he had gathered. After his death, his many manuscript notes and

drawings were also acquired for the Gardens, and are a most

valuable record of his work.

In this kind of way a great body of records concerning the native

orchids in Malaya and neighbouring countries was built up. Much

of this recorded knowledge became of great importance when the

breedings of new orchids, for use as garden plants and for their

cut flowers, was begun about the year 1928. It is true that only a

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Gardens Bulletin, S.

small proportion of orchid species are useful for exploitation in

this way; but for a proper appreciation of the possibilities, the full

survey is necessary, showing the more useful plants in perspective

as parts of a whole, and thus showing their own inter-relations.

Such a survey also shows what other plants exist, not yet in cultiva-

tion, which might be valuable for breeding purposes.

The impetus to orchid breeding was provided by Mr. John

Laycock, who imported to Singapore many species from neigh-

bouring countries, thus providing valuable breeding stock, and by

a visit from Prof. H. Burgeff, a pioneer of the study of orchid

seedlings. Up to that time, no orchid seedlings had been raised in

Singapore by modern methods of pure culture which ensure a

high percentage of germination; and nobody had any experience

of the behaviour and cultural needs of young orchid seedlings

newly removed from culture tubes. As a result of trials and errors,

methods of dealing with these plants were gradually discovered,

and the first seedlings flowered in 1931 (Spathoglottis Prim-

rose). A succession of new hybrids followed, and at the time of the

Japanese occupation of Singapore some 4,000 seedlings were in

pots at various stages of development, in addition to those in

flasks and to larger plants. A considerable proportion of these

plants was maintained during the occupation period, though there

were many losses, and some new crosses were made. Methods of

raising seedlings on agar-agar prepared directly from seaweeds

were developed. Progress after the war was rapid, and today the

production and culture of fine new orchids has become of con-

siderable commercial importance, as well as a recreation for a

large number of people.

The significance of this development is that nature provides com-

paratively few kinds of orchids well suited to the local climate in

vigour of growth, and in freedom of flowering, which have also

the qualities of floral form and durability required for horticultural

purposes. The production of large numbers of new kinds of plants

by crossing, by new seedling techniques, and by selection of the

best, has resulted in a complete transformation of Singapore gar-

dens, and in the production of a wealth of cut flowers throughout

the year in considerable variety, and of kinds previously unknown.

It is an example of the kind of change which is possible also with

other families of plants.

The Ginger family provides a rather different story. There are

about 150 known species of the family in Malaya, most of them

wild in the forests, a few others being cultivated in villages for

flavouring food or for medicine. Nearly all plants in the family

have aromatic properties, and more of them are potentially useful

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Vol. XVII. (1958).

than are yet in use. Little or no breeding or selection, even of the

kinds now in use, has yet been undertaken. The family has its

main centre of distribution in Malaysia, and the only full botanical

monograph upon it was prepared fifty years ago, in Europe, from

imperfect material. This monograph is very unsatisfactory, and a

careful study of native Malayan species, made in recent years at

Singapore, led the way to revised concepts of basic classification.

This study was possible through field and garden work similar to

that undertaken for orchids, as above described. The flowers of

the ginger family are delicate and short-lived, and also complex in

structure, so that dried specimens are difficult to interpret; one

needs to see either fresh flowers or specimens preserved in alcohol.

The revised survey of the family in Malaya gives a full statement

on classification as well as detailed descriptions of all species

known to occur in the country. It was in part based on pioneer

studies made of some allied species in Java; and in its turn the

Malayan survey has helped in the understanding of other Indo-

nesian members of the family. This study, made in Singapore

through a combination of botanical and horticultural activities

(especially the collections of plants and specimens made by Mr. E.

J. H. Corner) is basic to any future exploitation of this family,

which contains so many useful plants.

Bamboos are used daily, for many purposes, by almost all people

in Malaya; there are no other plants which have such varied uses.

But until recently no descriptive account of the various kinds of

Malayan bamboos had been published. The reason is that there are

many peculiar difficulties in the botanical study of bamboos. In

the first place, botanical classification must be based on floral

structure, and most bamboos flower rarely (one plant from north-

ern Malaya, in cultivation in Singapore for fifty years, has never

been known to flower). Thus it is obviously necessary also to study

other parts of the plants, so that recognizable descriptions, based

on the non-flowering parts, may be drawn up. In practice, such

descriptions are not easy to make, and no real attempt to make

them was tried in Malaya until quite recent years. This attempt

was only possible because from time to time native Malayan bam-

boos of various kinds, and also species from India and elsewhere

outside Malaya, had been brought into cultivation in the Botanic

Gardens, Singapore. These plants were available for detailed study

at leisure, and their flowering could be observed when it occurred.

As a result of such study, further field work could be effectively

carried out, and resulted in more plants being brought into culti-

vation, as well as in specimens and in recorded observations on

plants found in various parts of Malaya.

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The results of this first survey of Malayan bamboos, made pos-

sible by the cultivation of plants in the Botanic Gardens, Singapore,

have now been published. It is hoped that this publication may be

made the basis for a comparative study of the useful characters of

Malayan bamboos. In the north of Malaya there are many kinds of

native bamboos which have never been brought into cultivation,

and there have been no experiments to test the usefulness of parti-

cular kinds of bamboos for different purposes, or to select the best

kinds for propagation and wider use. Here is a subject which

touches the daily lives of all people living in the rural parts of

Malaya, basic work on which has been carried out at the Botanic

Gardens, Singapore, and which awaits that further investigation

which is outside the scope of the Botanic Gardens.

194

Singapore’s Contribution to the Study of Palms

By C. X. FURTADO

Botanic Gardens, Singapore

BOTANICALLY PALMS are considered as the “Princes” of the vege-

table kingdom, but they are also important economically. In some

cases like the coconut palm it is possible not only to make a boat

out of the timber but also to provide it with ropes, sails, cloth, .

flour, oil, sugar, wine, vinegar, spoons, brushes, brooms, drinking

vessels, etc. all derived from the palm itself, showing thereby a

variety of uses some palms have. Others like the Nibung made the

Malaysian civilisation possible since its timber was needed not only

for the boats but also for the houses, for no other cheap timber

was available that could stand salt water and also resist the attack

of termites and other destructive organisms. Similarly were it not

for the leaflets of the two palms Nipah and Rumbia (Sagu),

Malaysians would have experienced great difficulties to find for

their houses and boats suitable attaps that would resist weather

and the destructive action of white ants and other insects. Much

of the sugar that .was available to the original Malaysians in low-

lands was derived from coconut and Kabong palms, since sugar

cane could not be grown without being protected against the

ravages of wild animals. The “dragon-blood” which was formerly

exported to Europe and elsewhere for medicinal and other uses,

was derived from some species of climbing palms known as rat-

tans or rotang, a group of palms that provide what is commer-

cially known as the Malacca cane used for making walking sticks,

wicker chairs and other articles.

Now every species of a particular genus of plants has different

properties and uses. This is one of the reasons why systematic bota-

nists keep themselves busy in studying the characters by which

different species. and varieties of plants might be recognized. Sys-

tematic studies help to correlate the results of research workers

in different branches of science or knowledge. Thus a dietician or

nutritional chemist will want to classify the results on the different

vegetable products used in the diets of the people, the pharmacist

to know the plants used as medicines by the aborigines or the rural

people; the police to know the plant poisons used in murders,

suicides and other crimes; the veterinary officers to know the dif-

ferent kinds of plants used to feed, to poison, and to cure the

animals; the fisherman to know the small plants that help the fishes

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Gardens Bulletin, S.

in pisciculture; the malaria controllers to know the plants that

encourage or hinder the propagation of the mosquitoes; the silvi-

culturists, the trees that yield the timbers suitable for different

purposes, and so on. The systematic botanist is therefore an indis-

pensable person for research workers and commercial men who

have anything to do with plants or their products.

To classify plants that are not microscopic a large collection of

dried specimens to illustrate the leaves, flowers and fruits as well as

the natural variation between these is necessary, for only with a

carefully preserved material in herbarium can a systematic botanist

concentrate to detect the characters that distinguish one species or

variety of plant from another. The need for such a collection is

easily satisfied in case of plants which have fairly small twigs,

leaves and fruits. But in the case of palms the difficulty is often a

formidable one, both for the field botanist who collects the speci-

mens in the field and for the systematist who arranges and studies

the material in the herbarium. Many palms have large leaves and

their stems are very bulky too. If one were to collect only one leaf

and flowering branch of a large palm, they would form a great load

for a person, whereas within the time and labour several dozens of

specimens could have been made of large trees having small twigs,

leaves, flowers and fruits.

Palms also show a good deal of difference in age. The leaves

produced when the plants are young and not flowering are often

completely different from those produced by a flowering stem. Fur-

ther an inflorescence might appear completely male at one stage

and completely female at another stage. Moreover in the early

stages of flowering of a palm that produces both male and female

flowers on the same spadices, often no female flowers are produced

so as to make one imagine that the sexes are distributed on

different plants.

In case of rattan palms, additional difficulties occur often in the

better varieties that produce long climbing stems. The long, hooked

whips which enable the plant to climb can entangle a collector

tearing his clothes and wounding his skin in a way that it may be

difficult to extricate him unless armed with a sharp knife. Since

such climbing rattans produce flowers and fruits only when they

reach to a great height on the trees which they climb, it becomes

impossible to get fertile specimens essential for the herbarium ex-

cept by cutting the rattan; but since the rattan does not climb

straight and since the hooked whips are sometimes 15 ft. long or

more and the hooks are reversed, it is not possible to pull such a

rattan down even with a force of 10 to 15 men. Several times I

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Vol. XVI. (1958).

have had to fell 7 to 8 trees and treelets before getting the speci-

mens of one rattan. Further all rattans have thorny sheaths on

the leafy stem and the leaves themselves are also armed with hooks.

This means that one has to use leather gloves and handle the

whole material with care lest it injures the hands and fingers.

Now having obtained a rattan plant, how is one to make the

specimens since each leaf or spadix is very large? In the early days

when forest officers knew little about collecting palms, the Berlin

palm specialist wrote to a forest officer in New Guinea to send

him a good specimen of a palm that looked to be a new species

though well known to the aborigines. The silviculturist immediately

got his men active. He cut a palm with 30-50 ft. tall trunk and 18

inches thick, and had his men transport it to the nearest port

whence the plant was shipped to Germany. On its arrival in Ham-

burg special arrangements had to be made in order to transport it

to Berlin, and it was kept in the Museum for the benefit of the

school children. But since during the transport the flowers and

fruits were lost, the specimen was useless for a botanical study,

though several hundreds of dollars were spent in getting the speci-

men from the jungles of New Guinea to the Botanical Museum of

Berlin!

This digression will help to emphasize the need for the botanists

to study the palms in living state in order to decide what parts are

essential to enable him to distinguish the one species or variety of

rattan from another, that is, the specimens that will not burden

unnecessarily the collector and that, without occupying unneces-

sary space in the herbarium, will provide the necessary distinctive

clues to the systematist. In this regard the observations made by

the botanists in the field as well as on the plants grown in the

Botanic Gardens of Bogor and Singapore have provided useful

clues for making satisfactory specimens. And as far as the Malayan

rattans are concerned, the Botanic Gardens of Singapore may be

given the credit of having played the major role in the classifica-

tion of the rattans, having secured in the initial stages the help of

the great palm expert, the late Dr. O. Beccari of Florence, Italy,

who had also studied the palms in the forests of Borneo and New

Guinea and those collected by the staff of the Botanic Gardens of

Bogor and Singapore.

In the course of these studies the status of several doubtful ones

has been cleared, some passing under two or more names because

of some variation due to age or ecological conditions have been

merged into one, several new species and varieties have been des-

cribed and the generic distinctions have become better understood.

In 1907, for instance, Ridley who, as said above, had enlisted

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the help of Professor Beccari of Florence, described in his

Materials 86 species of rattans under 6 genera, of which 16 spe-

cies were new, that is, in addition to those new species described by

Beccari from the specimens collected by the Singapore Gardens’

Department. Later in his Flora of the Malay Peninsula V (1925)

Ridley described 94 species of rattans (with 5 new species) under

7 genera. In 1951-56, Furtado described and illustrated in the

Gardens’ Bulletin, Singapore, 127 species of Malayan rattans under

10 genera, of which one was a new genus and 28 new species and

several varieties and new records of species previously known only

from non-Malayan regions.

However the study of the Malayan palms is not yet complete. A

proper field survey of all the palms is needed in order to build up

an adequate herbarium in Singapore, and in this the co-operation

of the Forest Officers is valuable, since they make frequent trips

to the jungles and can make valuable collections whenever a coupe

is felled for timber. If a comparative systematic study were made

of the palms from the Malaysian region including Assam, Burma,

Lower Siam, Malaya, Sumatra, Java and the neighbouring islands,

it should yield much valuable knowledge of the palm genera and

species occurring in this region, their distribution, and the range

of intra-specific variation and other characters.

198

Manila and the Singapore Gardens

By EDUARDO QUISUMBING

Director,

National Museum, Manila

THE NATIONAL Museum of the Philippines, in Manila, greets the

Singapore Botanic Gardens on the occasion of its centenary. We in

the Philippines, who have immensely and continuously profited by

its goodwill and co-operation, have every reason to feel proud of its

achievement.

For the last 100 years, the Singapore Botanic Gardens and Her-

barium have been known as institutions which are not only useful

to Malaya but to all scientists the world over. While both have

served as places of recreation and education for the people of

Malaya, they have contributed their own share in the dissemination

of plant knowledge and distribution of plant species to all parts of

the world. The Herbarium itself is well known for its valuable col-

lections and adequate library and serves as a seat for research and

study of the flora of Malaya.

Many famous names have served as directors of the Gardens and

many of its botanists have made invaluable contributions not only

to the flora of Malaya, but also to that of Malaysia. Their works

appear in the Gardens’ Bulletin, which, likewise, has been con-

sidered as an invaluable scientific publication.

Our first scientific contact with the Singapore Botanic Gardens

was established after the year 1910 by the late chief botanist (later

director) Dr. Elmer D. Merrill, of our Bureau of Science. He had

started the exchanges in botanical knowledge as well as in botanical

specimens, which are equally valuable to both institutions. This co-

operative work was continued by me after 1923 when Dr. Merrill

left the Philippines and I became an official of the Philippines

National Museum, which handles botany as well as all divisions of

natural history. Besides our exchanges of specimens we also ex-

changed our publications—the Gardens’ Bulletin for the Philippine

Journal of Science, and vice-versa—and loaned Philippine and

other Indo-Malayan mounted herbarium material to Malayan spe-

cialists for their study of Philippine representatives of various plant

families or for their preparation of monographs on _ botanical

groups.

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It is a pleasure to recall at this time my personal visits to the

Gardens—one in August 1928 and another in November 1938.

From this latter visit I brought home with me some cuttings of the

various varieties of bougainvilleas and a seedling of Lagerstroemia

floribunda. The former now adorns the gardens of Malacanang,

official home of the President of the Philippines, while the latter is

a full-grown tree now serving as a graceful ornament in my private

garden. The late Mr. Wester also brought many ornamentals from

the Singapore Botanic Gardens.

The Singapore Botanic Gardens was and still is the centre of

botanical work on the Malayan flora, just as Manila was and is for

the Philippine flora. Despite the complete destruction of our her-

barium during the war, the Philippines is at present in a position

to begin—as it has already begun—to rebuild its collection and

regain its place in its own field.

With such thoughts as the above on our past contacts and ex-

changes and close associations and co-operation in our particular

branch of science, the centennial celebration of the Singapore

Botanic Gardens has become truly significant to us. It is my hope

that we can continue to carry on our present harmonious relation-

ship for the mutual benefit and satisfaction that such relationship

can bring us.

200

The Botanic Gardens and Conservation in Malaya

By H. M. BuRKILL

Director,

Botanic Gardens, Singapore

RESOURCES are not only fodder for industry but include the maté-

riel for man’s existence and his mental and physical well-being.

Conservancy is a rational policy to avoid wasteful destruction of

resources, to ensure that what is renewable, such as plant and

animal life, is not utilised (destroyed) beyond its capacity to

maintain itself, and to put land to the best possible use for the

country, thus avoiding conflicting demands on it.

The Botanic Gardens have been involved in various conservation

projects which have contributed very significantly towards the deve-

lopment of modern Malaya*. The most important of these in the

early days has without doubt been in the field of forestry, but there

have been other matters the relevance of which to botany does not

appear so obvious, as for example town water supplies and the re-

settlement of aboriginees. But times and accents change and the

present trend is towards nature conservation to protect the little of

the natural landscape that is left to us.

Early writers on eighteenth and nineteenth century Malaya em-

phasised the extensiveness of the forests. In their extensiveness

authority saw no need for protection. When the Botanic Gardens

were first established in 1859, the Singapore forests had already

been so destroyed that Government had on its hands no less than

45,000 acres of abandoned land. (I. H. Burkill, 1918.) That was

over a quarter of the Colony. Such reckless destruction had spread

to Malacca and Penang and into the Malay States.

Murton, Superintendent of the Botanic Gardens, 1875-80,

pointed out in his first annual report (1876) that good timber trees

had almost entirely disappeared from Singapore. Government be-

came alarmed at the increasing damage done to the forest of the

Straits Settlements and Cantley, who succeeded Murton, 1880-87,

was commissioned in 1882 to report on the forests of the three

colonies.

* Malaya is used here to mean both Singapore and the Federation of

Malaya.

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Cantley (1883) recommended the formation of forest reserves

as much for the protection of the timber resources they contained

as for improvement by re-afforestation with valuable species. In

addition he specified the need of proper control over water catch-

ment areas supplying public water services, and for the drafting of

reliable maps. It is significant that the world over forestry officers

are styled ‘conservators’.

In 1884 Government set up a Forest Department under the

Botanic Gardens establishment. By 1895 when the Gardens relin-

quished control of the Forest Department, Cantley and Ridley, who

followed him, had secured 88,336 acres of forest reserves in the

Straits Settlements. There is no doubt that the official implementa-

tion of Cantley’s recommendations marked the beginning of a

constructive policy of land utilisation and conservation in Malaya.

One of the most besetting obstacles to forest regeneration in

Singapore was the frequency of lalang fires over the areas adjacent

to established reserves or in the pockets within the reserves. Cantley

selected Eugenia grandis as a fire belt tree. Several of his avenues

still exist.

During Cantley’s time, the Gardens were concerned with the

conservation of water resources. He became adviser to the Malacca

and Penang Municipal Councils. Curiously the Gardens had no-

thing to do with Singapore’s supply till very much later.

In the course of his duties with the water supply at Malacca,

Cantley had to advise on a land settlement scheme for a part of an

aboriginee tribe, the Jakun. This must rightly be considered a con-

servation of resources, not only because of the water scheme, but

because people are a part of a country’s resources and because

conservation in a wider sense applies equally to a cultural heritage

to which undoubtedly the aboriginee tribes of Malaya do con-

tribute.

At a risk of altering the sequence of events, it is convenient here

to mention the Gardens’ contribution in the post war years to the

land utilisation survey which culminated in the drawing up of the

Master Plan for the development of Singapore Island. J. W. Ewart,

Curator of the Gardens, was as well Agricultural Officer, Singapore,

for the period 1948-54 during which time the survey was under-

taken. Ewart working in conjunction with R. A. Wright, Singa-

pore’s Veterinary Officer, and the officers of the Land Office

prepared plans for rural development and the demarcation of

agricultural areas. With the creation of many new housing projects

a large number of market gardeners were displaced and had to be

resettled in the rural areas. These land utilisation plans brought up

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Vol. XVII. (1958).

many problems, not least of which was the loss of fertility of the

upper slopes and tops of hills which had been bare of all but the

most impoverished vegetation for decades.

In Singapore the water catchment area of the MacRitchie Reser-

voir had been extended by 1910 to the land surrounding the Peirce

Reservoir. Much of it had previously been alienated for cultivation.

The vegetation which returned was an Adinandra scrub which is

typical of secondary growth on Singapore Island. In 1930 the

Governor, Sir Cecil Clementi, saw in this land the makings of a

public park. Therein lay a significant widening of the conceptions of

conservation by the creation of a public recreational amenity. The

Governor instructed that attempts be made to improve the forest

and to create bridle paths and to beautify various vistas, which

were afforded, with stately trees. The arboricultural work fell to the

Botanic Gardens, but it was beset with obstacles and only a little

success was achieved before the outbreak of the 1939-45 war

stopped this work.

In the 1930’s there was a growing public awareness that all was

not well with the conservation measures for Malaya’s wildlife. A

commission of enquiry was established under the chairmanship of

T. H. Hubback whose report (1930) advocated measures to pro-

tect game within delineated reserves. The Botanic Gardens took

no part in this enquiry which confined itself exclusively to animals

and Malaya’s vegetation was mentioned only insofar as it provided

a habitat for wild animals. Singapore itself got but passing refer-

ence—that the whole Island should be declared a refuse for most

species of birds. The Botanic Gardens however saw the matter in a

wider perspective and Holttum advocated to authority that since

the F.M.S. enactments protected only animal life but not plant life,

the principle that both should be considered together ought to be

recognised. Holttum also advocated the reservation of areas con-

taining uncommon plant communities before they were lost for all

time under some other form of development. In particular he re-

commended the protection of Taman Sedia in the Cameron High-

lands and Gunong Panti in Johore. After the war Henderson

advised in the setting up of Templer Park just north of Kuala

Lumpur which has within its limits Bukit Batu Takun (1,387 ft.)

a limestone outcrop with its peculiar flora.

In Singapore the Forest Reserves were deleted in 1935 and

Holttum and Corner recommended Government to put those at

Bukit Timah, Kranji and Pandan, under the Botanic Gardens. In

1939 these areas were reserved again under the Forest Reserves

Ordinance and the Director of Gardens, after a lapse of 46 years,

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became again the Conservator of Forests. But there was no inten-

tion this time of commercial exploitation: it was for absolute pro-

tection to provide areas for research, education, recreation and as

samples of the country’s biographic history and heritage.

These reserves survived the war thanks to the interest of the

Japanese Director of Gardens, successively H. Tanakadate and

K. Koriba. But in the years after the war the reservation of Bukit

Timah came into conflict with granite quarrying interests. Govern-

ment set up a commission and its reports (Harvey 1950, 1951)

recommended the enactment of legislation to put the reserves on a

modern footing under a Nature Reserves Ordinance. This recom-

mendation was implemented in 1951. To the three reserves under

the control of the Botanic Gardens were added Labrador Cliff,

and the Municipal Water Catchment Area.

The intention of the new ordinance was: “To provide for the

dedication and administration of certain lands in the Colony of

Singapore as Nature Reserves’. A Board of Trustees was created

with the Director of Gardens ex-officio Chairman, and the Board

was directed to undertake the propagation, protection and preser-

vation of the indigenous fauna and flora of Singapore and to pre-

serve objects and places of aesthetic, historical and scientific

interest. The Board is empowered to provide facilities for study

and research.

The reserves when they were created, and still are unique in

Malaya. They put Malaya in fact in the forefront of the countries

of South-east Asia from the point of providing field research faci-

lities as well as providing educational and recreational amenities

so close to the centre of population.

Nature conservation is in itself a development project just as are

town planning or the building of aerodromes, roads or railways.

But since its stock-in-trade is things which live, as a project it is

not amenable to constant change and interference and so it must be

considered as a long term one. Allowed to grow on its own it is a

complementary anodyne for a modern society in this age of

concrete.

This brings the matter up to the present time. The story has had

an expanding theme from natural bounty to destruction, through

exploitation for commercial benefit to cultural and educational

ends. All these stages remain with us in varying degree, but it is to

human advantage to eliminate for all time wastefulness. The part

played by Botanic Gardens in this endeavour has been fundamental

and of great significance in Malaya’s development. These achieve-

ments have been and will continue to be of the greatest importance

for the future.

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Vol. XVII. (1958).

REFERENCES

BuRKILL, I. H. 1918. Gardens’ Bulletin, S.S. 2 (2) 1918, 55-72.

CANTLEY, N., 1883. Report on the Forests of the Straits Settle-

ments.

Hupssack, T. H., 1932. The Report of the Wildlife Commission.

HARVEY, J. A., 1950. Interim Report of the Select Committee on

Granite Quarries and Nature Reserves, No. 45 of

1950, Colony of Singapore.

Harvey, J. A., 1951. Final Report of the Select Committee on

Granite Quarries and Nature Reserves, No. 6 of

1951, Colony of Singapore.

MurtTon, H. J., 1876. Report on Government Botanic Gardens

for 1875.

Use has also been made of unpublished papers in the Botanic

Gardens Archives.

205

The Singapore Botanic Gardens and the Central

Institute for Nature Research in Indonesia at Bogor

Comments on Past and Present Copan

and the Need to Continue it

By KUSNOTO SETYODIWIRYO

Central Institute for Nature Research

Bogor, Indonesia

THE WELFARE and progress of a country, especially of one with an

economy and industry still in its infancy, depend a great deal on its

natural resources and on the ability and good sense of its leaders

to exploit and to make use of this wealth in such a way as to safe-

guard the well-being of its people from generation to generation.

The ability to make responsible use of the natural resources

rests on a vast and profound knowledge of the soils, the waters,

the flora and fauna of the country as well as of its wealth of mine-

rals that are contained in the soils.

A wise and responsible management should be based upon a

good understanding of the kind and the potency of the natural

resources and of the best way of utilization for the prosperity of

the country and the people now and in future.

In view of the welfare and development of a nation the leaders

should attempt to know their colleagues in other countries and in

the neighbouring states in particular, to create and maintain a

mutual understanding, trust, respect and tolerance.

If this basic principle were followed in the field of research, it

might become an endeavour to create an atmosphere of inter-

national co-operation and goodwill, a sincere spirit of “give and

take” with respect to research projects, achievements, views and

experiences, exchange of material, and reciprocated visits for re-

search purposes, all for a mutual benefit and improvement. Only on

this basis of goodwill and readiness to trust one another can the

ideals of international co-operation be truly attained.

The prosperity and achievements of a neighbour state should be

greeted with sympathy and rejoicings rather than with a complex of

superiority or inferiority which will undoubtedly mar the friendly

relations. On the other hand, the misfortunes that befall a neigh-

bour state should be shared, looked into, and relieved as well as

possible together as members of a family.

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Vol. XVII. (1958).

It is in this spirit of friendliness that I wish, on behalf of the

Central Institute for Nature Research in Indonesia, to convey our

sincere congratulations on the occasion of the centenary anniver-

sary of the Singapore Botanic Gardens. The relations between the

Central Institute for Nature Research in Indonesia and the Singa-

pore Botanic Gardens, which has always been characterized by a

mutual readiness to help one another, should, or rather needs be

continued according to the good, extant tradition, and should even

be expanded and intensified in the years to come.

Since the Central Institute for Nature Research in Indonesia,

better known by the name of “Botanic Gardens of Indonesia” since

its foundation in 1817, has a wider field of activities covering all

that lives in nature, both on land and in the waters, this Institute

would like to invite the Singapore Botanic Gardens to keep up a

more extensive co-operation than the one dealing with the plant

world only. I feel that the Singapore Botanic Gardens has a similar

function in the State of Singapore as the Central Institute for

Nature Research has in the Republic of Indonesia.

The integral task of the Central Institute for Nature Research in

Indonesia can be outlined as follows:—

(a) To conduct basic research and to give expert training in

the field of natural sciences (Botany, Zoology, Ichthyo-

logy, Oceanography) under conditions prevailing in tro-

pical areas, for the benefit of practical agricuiture,

horticulture, landscape architecture, forestry, conserva-

tion of nature and natural resources, veterinary science,

fisheries and medicine.

(b) To collect and to catalogue the plant and animal species

occurring in Indonesia and the neighbouring areas, to

select scientific names for them and to prepare mono-

graphs.

publications on the basis of international co-operation.

(d) To provide Indonesian and foreign scientists with the

accommodation and the facilities to perform research

work in the various institutes.

(ec) To expedite the long-term project of the “Flora Male-

siana” Foundation, i.e. to compile a standard work on

the vegetation of the Republic of Indonesia and the

neighbouring areas on the basis of international co-

operation.

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Gardens Bulletin, S.

(f) To protect and care for the natural resources of Indonesia

as well as possible and to disseminate knowledge of

nature conservation among the general public in ac-

cordance with the principles of the International Union

for the Protection of Nature and Natural Resources, of

which the Botanic Gardens of Indonesia are a member.

(g) To maintain a Zoological Museum, a Marine Aquarium,

a Freshwater Aquarium, and a Biological Station for

Animals Protected by National Laws, for the benefit of

popular education.

(h) To render advice on the lay-out of parks and gardens,

their upkeep and embellishment.

(i) To build up a Central Library for the Natural Sciences

and to supply books and periodicals needed by the

different institutes of the Ministry of Agriculture.

(j) To carry out orders from the various institutes for making

scientific pictures, films and drawings, and to train

candidates in scientific photography and drawing.

The above mentioned task is carried out concordantly by the

following institutes of the Central Institute for Nature Research:-—

1. Hortus Botanicus, founded on 18th May, 1817.

2. Herbarium Bogoriense, founded in 1817.

3. Flora Malesiana Foundation, founded on 21st October,

1950.

. Treub Laboratory, founded in 1884.

. Institute for Microbiology, founded in 1956.

. Museum Zoologicum Bogoriense, founded in 1894.

. Institute for the Conservation of Nature, founded in 1937.

. Biological Station for Animals Protected by National Laws,

founded on 17th August, 1957.

9. Institute and Aquarium for Marine Research, founded in

1904.

10. Bibliotheca Bogoriensis, founded in 1842.

11. Photographic and Drawing Studios, founded in 1878.

oN N NN

All of which form one harmonious and inseparable unit.

208

The Importance of Tropical Taxonomy to

Moder Botany

By E. J. H. CORNER

Botany School, Cambridge, England

(lately of the Botanic Gardens, Singapore)

EFFORTS TO LEARN about plants are called botany, when they are

sufficiently rational. Efforts to grow plants and make new ones and

to destroy the harmful and unwanted are parts of agriculture and

sylviculture, when they are sufficiently profitable. Our enjoyment

of plants, wild or in the garden, has no name, but it is an aspect of

civilisation as essential as art and literature; and horticulture is a

sign of progress. Thus, in concourses of men, the academic, the

applied, and the recreational sides of plant-lore have developed

into the botany schools of universities, the plant-breeding stations

of agriculture, the research institutes of forestry, the nature-reserves

of wild-life services, the parks and botanical gardens of cities, and

the flower-beds, window-boxes, vases, and books of the home.

Wretchedly barren is that community unmindful. In this concrete

age, which hardens our lives, we should reflect upon the appeal of

kampong, sawa, pasir panjang, gunong hijau, sungei berassau, and

kayu chondong. The beauty of Rio de Janeiro, so inspiring to the

visitor, is the city in the bay of forested mountains. Seeing then

that botany, if we use that word for all ways in which plants enter

our lives, is a subject vital to learning, practice, and recreation, let

us consider the harder word taxonomy.

Taxonomy. All sciences split up as particular methods are used

in their study. Botany, too, is sundered into an increasing number

of “-ologies”, many of which require not a knowledge of plants so

. much as of other natural sciences, and an electron-microscopist or

radiographer may be a botanist. One of these divisions is now

called taxonomy. It is the old core of professional botany, derived

from the herbalists through the systematists who variously im-

proved the classification of plants. It is regarded as old fashioned,

being the parent, but rather is it the trunk to the branches, ever

swelling as they multiply, gathering the information, and, I believe,

still the core and heart of botany. Several “-ologies” and “ogra-

phies” are indeed, attributed to taxonomy, which means the method

of orderly classification but we can understand better what it is

from its outcome. Nowadays taxonomy is the classification of plants

according to their evolution. The early botanists and zoologists

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discovered that plants and animals were constructed in definite

ways, from which an orderly classification could be thought out

and named, so as to give a better and international understanding

of biology. A good classification is needed in science to pigeon-hole

the enormous and ever-increasing number of facts: that it is objec-

tive and satisfactory, international recognition proves. Until a hun-

dred years ago, however, this classification was an enigma because

there was no adequate explanation of the expanding, varying, and

yet coherent, marvellous pattern of life which it revealed, or why it

should have been satisfactory. Darwin’s theory of evolution was not

a flash in scholastic darkness, but the sun which rose on biology.

The orderliness of classification was seen to express the evolution

(and extinction) of plants and animals in the past. They were being

classified on the peculiarities of their various lines of descent, or

heritages; being founded on “raison d’étre”’, this was the order so

satisfactory. The sun, I think, was the spontaneous combustion of

taxonomy, but that is another story. To appreciate the importance

of taxonomy, as evolutionary classification, we must consider a

method of scientific enquiry which it has introduced. This is the

distinction between homology and analogy. The method is most

useful in other branches of biology which have not been able,

because of the baffling complexity of protoplasm, to advance as far

as morphology, or the study of structure, on which taxonomy is

based. I refer particularly to physiology, genetics, biochemistry,

and other experimental subjects which are incapable of classifying

living things, of correlating the facts of biology, and of organising

biological thought.

Homology and Analogy. There are two ways of studying plants

and animals, namely the particular and the comparative. In the

particular we investigate one kind of plant or animal, or one part

of it, intensively. Thus, we may study the respiration of a root or

muscle, or the absorption of mineral salts by root hairs, or the in-

heritance of certain characters such as flowering-time, grain-size, or

seed-production in a plant such as the rice-plant. As any of these

subjects, or the means by which they are studied, can be developed

into a particular branch of science, it can be seen how these bran-

ches multiply in modern research. Sooner or later comparison must

be made with other plants or animals, but what ones? Obviously

comparison of objects superficially similar, as fish and whales, or

ferns and palms, may be exceedingly misleading, for their similari-

ties are analogous, not fundamental; and, as we are dealing with

the minute and intricate behaviour of this exceedingly complex,

largely unknown, protoplasm of living things, we must avoid as

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Vol. XVII. (1958).

many unknown quantities as possible. The best comparison, there-

fore, will be between species which have been shown by modern

taxonomy to be closely allied: their protoplasm will differ the least

from each other and these slight differences may be exactly what

the researcher needs. Basically, plants have a certain and not im-

possibly large number of properties, or things which they can do.

Respiration, photosynthesis, cell-differentiation, and reproduction,

for instance, are essentially the same in all green plants, but the

species differ in the way in which these processes are variously

amplified, side-tracked, baffled, distorted, suppressed, and so on.

This baffling of living processes, as we may call it, occurs largely

through the solid parts of the plants making up their visible struc-

ture, on which morphology, and so taxonomy, are built. Now any

one living process consists of many steps. One species of plant may

have steps A—-D, another A—H: one may exaggerate B—D at

the expense of A and E, or vice-versa: another may add N—Z, and

another eliminate A—-P: some steps may be passed over so rapidly

as not to be detectable, and so on in endless variation. Thus, when

we study one plant in particular, we cannot know how the process

under investigation is baffled: probably we have only part of the

picture, and we do not know which part. To understand more

fully we must compare taxonomically related species (of the same

general heritage) which will, through the very fact that they are

not the same species (or genus) and have not precisely the same

protoplasm, reveal other parts of the picture. This is homologous

comparison as distinct from the artificial and analogous comparison

which will introduce a multitude of unknown quantities; and homo-

logous comparison depends on good taxonomy: it is, in fact, taxo-

nomy. Obvious and simple as this important method of enquiry

seems, it is far from being generally appreciated, and a great deal

of botany is confused by analogous comparison. Nevertheless, it is

being tacitly adopted because it works, and it is my purpose in this

article to make sure that we understand why it works. Thus, com-

parative cytology, dealing with chromosome-numbers, structure,

and behaviour, could never have been developed if there had not

been the taxonomic work for it to follow: the tendency is to extol

the new cytology and forget, if not to ridicule, the service of taxo-

nomy. Laboratories have begun to look for new “guinea-pigs”’:

hitherto they have adopted what was at hand, but they should be

advised to enlarge their stock-in-trade taxonomically. Plant-breed-

ing is scouring the world for wild species allied to crop-plants,

whereby to breed new strains. Sylviculture is considering other

timbers as world-demand exhausts supply. Horticulture is experi-

menting with the vigour and novelty of hybrids. In every case the

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guide to requirements is in the pages of taxonomy, wherein the

world’s supply of species are catalogued for ready reference on the

principle of hereditary construction. When we consider that this

research demands taxonomic information from microscopic plants,

as algae and yeasts, to timber-trees, orchids, and grasses, the im-

portance of taxonomy can be understood. It has well been called

the hand-maid of biology: in my experience, the higher the office,

the greater the servant, and that applies both ways to taxonomy.

Tropical Taxonomy. I have assumed that our evolutionary

classification is perfect. Of course it is not. Much is indisputably

true, such as the recognition that aroids are not orchids, or casua-

rinas are not pines, but there are many plants, including casuarinas,

the proper position of which in classification we do not know. In

many groups, such as orchids, grasses, sedges, palms, and many

trees, we do not know how to draw generic limits and, often, we

have not the species properly arranged in the genus. We have no

satisfactory classification of the orders of dicotyledons, and that of

the larger fungi is extremely empirical. There are four great diffi-

culties: firstly, the multitude of organisms is almost inconceivable:

secondly, there have never been enough taxonomists to do the

work thoroughly: thirdly, the material on which the taxonomist

has to work is too often insufficient: and, fourthly, it is extremely

difficult to distinguish analogy from homology, or, as we say,

parallel from phyletic evolution. The result is that, when a new

taxonomist has enough material to re-examine a deficient group, he

almost invariably discovers that previous classification has mis-

taken analogy for consanguinity. One of the greatest revisions in

this respect was the re-classification of the algae towards the end

of last century: though scarcely mentioned in the history of botany,

it has revolutionised the teaching. Now, nowhere are these defects

sO numerous as in the tropical countries. Fungi, ferns, and flowering

plants have their headquarters in the tropics because here the

broad-leafed trees have built up the richest biological environment

of all—the tropical forest—wherein tens of thousands of flowering

plants, ferns, and fungi dwell. You may think that north temperate

students of fungi have a good knowledge of Boletus, but if we

knew the boleti from Assam to New Guinea we would re-do the

classification of Boletus! The “mushroom-soup” of Europe, which

is a prized and limited commodity, is made from Boletus edulis,

many allies of which grow in Malaya. If we understood the tro-

pical polypores, which abound on wood from rubber-roots and

tea-bushes to meranti and oil-palm, and in thousands in the forests,

we would entirely revise the present artificial classification of these

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fungi, with results of great significance for research: in fact, re-

search on these fungi is baulked through lack of satisfactory classi-

fication. If we could revise the classification of the trees and lianes

of the tropics from living plants and bring this improved knowledge

into general botany, we could resolve many of the major problems

of flowering plants. Living plants? So far, botany has had to deal

mainly with dried specimens and pickled flowers of tropical plants;

and more than half the plants of the world are known, accordingly,

fragmentary and dead. There is the practical difficulty how to study

these plants nearer to the places where they grow than the

museums, herbaria, and botanic gardens of Europe and North

America. The interest of most of these plants may be only aca-

demic, but we do not know which may not become a subject of

intense laboratory or commercial interest. Thus Rauwolfia sprang

to fame, and luckily it had been studied taxonomically. It has been

said that, as timber supplies dwindle, bamboos of rapid growth will

be required, and how many of us feel content with the meagre

knowledge of bamboos? Indeed, with new methods of fibre-boards,

sylviculture may turn to entirely new practices with the small

quick-growing trees of the tropics and the manager, or taxonomist,

of these plants will be needed to advise. Most tropical fruits could

be improved and diversified, but nearly all the general of tropical

fruits need taxonomic revision for adequate research. The culti-

vated banana has demanded extensive botanical exploration in the

last twenty years to collect as many wild species of its genus,

Musa, as could be obtained. Solanaceous fruits, particularly of tro-

pical America, may be called upon next, but we have no modern

guide to the genus Solanum. In contrast, the durians have recently

been monographed and here is the prospect for the improvement

of this popular fruit.

Tropical Botanical Institutes. There is no fundamental difference

between tropical and temperate botany, but the richness of the

tropical flora means so much more to be discovered and the ex-

pansion of our thoughts. As rubber, coffee, cocoa, banana, and

other tropical commodities in daily life, botany also should thrive

on tropical ideas. Our textbooks, however, written from the north

temperate angle, seem to me like the splendid treatises the Romans

could have written about the world, ignorant of three-quarters of

it; and it was the “barbarians” who took over! In tropical botany

we are passing from the era of exploration, when we had to be

content with the dead records of expeditions, to that of the scientific

appreciation of the living plants now made possible by expanding

civilisation. The former has still much to contribute, which tropical

countries cannot by themselves achieve, as may be learnt from the

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Flora Malesiana and similar works on Africa in process of publi-

cation. But, for the taxonomic progress with living plants and all

the impetus to botany, theoretical and applied, which will follow,

we must look to the botanical institutes in tropical countries. I am

impressed by the growing number of young students in the tropics.

The thirst for knowledge cannot be quenched. They will build, and

civilisation will expect it. Western science has led the way to a

better appreciation of nature, but the tropical countries must now

help their eager students to extend this knowledge in their own

rich heritage for the benefit of mankind. I have often thought that

what has been started in botany in Europe and North America

will eventually have its headquarters in the tropics where the plants

themselves orientate thought. On the tropical students now falls

the responsibility for writing their biological floras, as is the newest

taxonomic development in Europe. Theirs will be the responsibility

of preserving the native vegetation and the beauty of the country

by wayside and in national park, and of collecting the living assem-

blage of economic, ornamental, and rare plants for research and

recreation in botanic gardens. Theirs will be the opportunity to

step up all botany by study of new sorts of protoplasm. It is a hope-

ful glimpse. At present they may lean on outside support, but |

Jook to the time when students from outside will learn in the tro-

pical institutes.

In conclusion, tropical countries have a duty to promote botany.

They inherit a rich share of plant-evolution, which has yet to be

incorporated into the wisdom, practice, and enjoyment of civilisa-

tion. Their students will expect intellectual progress. In practice,

success will depend on taxonomic efficiency, which is the chief

scientific work of botanic gardens; other subjects will be developed

by universities, special institutes, and amateur societies. It is not a

pole or bush to be grown, but a tree of knowledge, the trunk

whereof is taxonomy.

214

The International Code of Nomenclature for

Cultivated Plants with Special Reference to

Tropical Botanic Gardens

By J. S. L. GILMOUR, M.A., F.L.S., V.M.H.

Director, University Botanic Garden, Cambridge

Rapporteur of this International Commission for the

Nomenclature of Cultivated Plants

THE NAMES OF PLANTS are of three kinds, botanical or scientific

names (e.g. Mangifera indica), common or vernacular names

(e.g. Mango), and the names of the cultivated varieties or cultivars

(e.g. the Mango ‘Jafina’). The use of botanical names is governed

by The International Code of Botanical Nomenclature, under the

control of successive International Botanical Congresses; common

names are gloriously free from all control, except that of common

usage; cultivar names of horticultural plants have for many years

been, in theory, subject to an International Code, but, until recently,

the Code was virtually unknown to most horticulturists, and it is

only since 1953 that a widely circulated Code has been available

to those concerned with cultivated plants. It is this Code that I

want to discuss in this article, and especially its importance to the

naming of plants cultivated in the tropics.

The International Code of Nomenclature for Cultivated Plants,

published in 1953, was really a combined effort of the International

Botanical Congress held at Stockholm in 1950 and the Inter-

national Horticultural Congress held in London in 1952. Though

entitled “for Cultivated Plants”, it was drawn up primarily with

horticultural plants in mind, and it soon became clear that several

modifications would have to be made if it was to be equally service-

able to agriculturists and foresters. Accordingly, a special /nter-

national Commission for the Nomenclature of Cultivated Plants

was established under the International Union of Biological Sci-

ences, and this Commission set to work to re-formulate the Code

so that it would apply satisfactorily to all cultivated plants. The

full history of the work of the Commission is set out in the Preface

to the new (1958) edition of the Code that it produced, together

with the names of the agriculturists, horticulturists, foresters and

botanists responsible.

The Code is obtainable from the Royal Horticultural Society in

London and, since its publication, is gradually becoming known

215

Gardens Bulletin, S.

and accepted by those concerned with the cultivation of all types of

plants throughout the world.

I would strongly recommend that all those interested should

obtain a copy of the Code, but it may be helpful to summarize

here some of its main provisions.

(1) The Code defines a cultivar or cultivated variety as “an

assemblage of cultivated individuals which are distinguished by

any characters (morphological, physiological, cytological, chemical,

or others) significant for the purposes of agriculture, forestry, or

horticulture, and which, when reproduced (sexually or asexually),

retain their distinguishing features.” It recommends the use of the

international word “cultivar’’, but allows, of course, for the use of

other terms normally employed as equivalents in various languages

(e.g. Variety in English, Sorte in German, etc.).

(2) The various different kinds of cultivars are set out and

defined (i.e. clones; lines; assemblages of individuals showing gene-

tical differences but united by one or more characteristics differ-

entiating it from other cultivars; and fi hybrids of the type now

common in maize and other crops).

(3) The category strain is not adopted in the Code; any selec-

tion showing sufficient differences from a parent cultivar to render

it worthy of a name is regarded as a distinct cultivar.

(4) It is recommended that, in future, all cultivar names should

be “fancy names”, that is, names markedly different from botanical

names in Latin form, so as to distinguish them from the names of

botanical varieties. However, existing cultivar names in Latin form

are not to be altered.

(5) Various rules are set out governing the choice of new cul-

tivar names. For example, they should preferably consist of only

one or two words, and must not consist of more than three; words

that are excessively long or difficult to pronounce in other languages

should be avoided and there are many other similar common-sense

provisions.

(6) Names may, of course, be in any language, and there are

regulations for transliteration and translation into other alphabets

and languages. Where a cultivar name, even though transliterated

or translated, is “not commercially acceptable” in other countries,

it may be changed and a “commercial synonym”’ substituted, pro-

vided the synonym is approved by the originator of the cultivar

and by the official Registration Authority concerned (see below).

(7) For cases where two or more names have been given to a

single cultivar, rules are laid down for choosing the correct name.

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Vol. XVII. (1958).

These are, broadly speaking, based on priority of publication, but

exceptions are allowed for when strict application would lead to

the changing of widely used names.

(8) Rules are provided for the description of cultivars and the

publication of cultivar names. Any language may be employed

for the description, but the use of English, French, German, Rus-

sian or Spanish is strongly recommended.

(9) The method of writing cultivar names is fully dealt with.

All names should be written with capitals, unless the custom of the

language concerned dictates otherwise. When printing a cultivar

name “‘in full”, it should follow the botanical (or common) name

of the species or hybrid concerned and should be distinguished

typographically from it, preferably by presenting the botanical name

in italics and the cultivar name in roman type, and by enclosing

the cultivar name in single quotation marks, e.g. Mangifera indica

‘Jaffna’.

(10) Full regulations are included for the naming of hybrids

produced in cultivation, and their cultivars.

(11) One of the most important sections of the Code deals with

the establishment of official Registration Authorities for all the

main groups of cultivated plants. The primary function of these

authorities is to register the names of new cultivars in their groups,

in much the same way as the names of race horses or pedigree

dogs are registered, thus preventing duplication and ensuring that

names are in accordance with the Code; but the authorities would

also be responsible for publishing basic lists of all the cultivar

names in their group (to be brought up to date from time to time),

approving commercial synonyms and names retained against the

rule of priority, and, in general, for acting as a central clearing

house and “advisory bureau” for the cultivars with which they are

concerned. The aim is to establish one International Registration

Authority for each group, but either as a stage towards this, or, in

the bigger groups, perhaps in addition to the International Autho-

rity, National Authorities may also be desirable. For horticultural

plants, International Registration Authorities are appointed by an

International Horticultural Congress and national authorities by

agreement between those concerned.

A number of International Authorities has already been ap-

pointed by the Horticultural Congress held at The Hague in 1955

and at Nice in 1958. These are mostly for plants grown in tem-

perate climates (e.g. The American Iris Society for the genus Iris

and the Royal Horticultural Society for Rhododendron), and there

will be a great need in the future for suitable bodies to undertake

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Gardens Bulletin, S.

the registration of names of cultivars of ornamental and useful

genera grown in the tropics. This is where tropical botanic gardens

might play a very useful role, especially in acting as registration

authorities for ornamental genera. An increasing number of culti-

vars are being produced in these genera and it is important to

begin registering their names, and printing basic lists, at as early a

stage as possible, so as to prevent the development of overlapping

and duplication. Bougainvillea, Hibiscus, Nerium, and Zinnia are

genera that spring to mind, and there are, of course, many others,

both ornamental and useful, that should be included in a compre-

hensive scheme. It should be emphasised that, although it is highly

desirable to combine cultivar trials with registration, this is not

essential, and the registration of names can be carried out inde-

pendently of such trials. If any botanic garden or other suitable

institution in the tropics feel that they are able and willing to act

as a registration authority, they should communicate with the

Secretary of the International Commission, Dr. H. R. Fletcher,

The Royal Botanic Garden, Edinburgh.

I hope that this very brief account of the International Code and

of the registration scheme that it embodies will be of interest to

those concerned with the cultivars of tropical plants, and that they

will promote international co-operation in this important field by

following the provisions of the Code and by helping to set up

registration authorities where they are needed.

218

The Importance of Biological Research in the

Pacific Region

By F. R. FOSBERG

Chairman, Standing Committee

for Pacific Botany

The Pacific Science Association

AT FIRST GLANCE The Importance of Biological Research in the

Pacific Region seems such an obvious topic that anything one

might write about it would be a truism. The world, including the

Pacific, is dependent for its food supply on the products of several

branches of applied biology. Large parts of industry depend for

raw materials on applied biology. We all depend for cures for our

ills and injuries on applied biology.

Before going on it may be well to define our terms. Biology is

the field of science that deals with living things. Its scope runs

from minute bacteria and even viruses to flowering plants and from

the simplest one-celled animals to dinosaurs, elephants and man

himself. It deals not only with the kinds, nature and physiology of

organisms, but with their relationships to each other and to their

non-living environments. Research is the attempt to learn new facts

and to determine their significance in relation to what is already

known. It is biological research, of course, when the facts learned

are about living things.

Hence, agriculture, husbandry, fisheries, forestry, and medicine

are branches of applied biology, and are only some of the branches

that affect our daily lives. And biological research is the search for

new knowledge bearing on these vital activities. Who could ques-

tion the importance of this? In view of the facts that we do not have

enough food for all the people, that raw materials are often scarce,

that we still fall ill and die before our normal life span is over, it

would seem obvious that biological research would be regarded as

perhaps the most important of all activities, that the demand for it

would be unlimited and the amount of research would only be

limited by the supply of competent and willing research workers.

Curiously enough, this is not generally the case. What we know

about agriculture, forestry, fisheries and other applied fields in

biology is taken for granted. The research worker is commonly

regarded as a strange man, perhaps a little mad. That there should

be a reason to pay him a salary, or that he should need funds for

his work, are hard for the politician who controls the spending of

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Gardens Bulletin, S.

public money to understand and harder for him to accept. And

when the budget must be reduced, the first and easiest item to cut

out is research. This may seem incredible and incomprehensible

but is a phenomenon observed repeatedly. It may be worthwhile to

seek an explanation for it.

Clearly basic to the problem must be a lack of awareness that all

is not going as well as it could. The politicians and the influential

people that control them are not the hungry ones. Their economic

situation is such that they can buy food when they need it regard-

less of its scarcity. If they are users of raw materials, scarcities only

show up, during peace-time, at least, as increases in costs which are

simply passed on to the consumer in the form of increased prices.

Deficiencies in medical knowledge, indeed, do become evident to

even the most comfortably situated and influential politician or

magnate, at least when he gets old. And it is significant that medical

research is by far the best supported of the research activities of

_ biology.

Perhaps the best way to approach the subject of the importance

of biological research is to examine the general system in which we

live, to see just what troubles that result from an insufficiency

of biological knowledge might be corrected by research. If

attention can be directed to these and some indication given of how

biological research is pertinent toward remedying them the general

question may be clarified.

The fact that people are inadequately nourished was mentioned

above. It has been said that this is due to inadequate distribution

rather than under-production. However, if distribution of food were

really equalized, probably the well-fed among us would gain a

slight knowledge of the pangs of hunger and the presently hungry

would only be a little less so. And with this somewhat more ade-

quate nourishment more of the presently starving would survive,

susceptibility to disease would be less, and population would in-

crease to where the food supply would be on the whole more inade-

quate than now. Research in agriculture, and in the various basic

aspects of biology that are applied by agriculture, can contribute

greatly toward the alleviation of current food shortages, also of

shortages of raw materials produced by agriculture. Few in Malaya

will have to be told of the part played by botanical research in the

present prosperity of the area. The entire rubber industry in Malaya

stems from the experimental introduction of the Hevea rubber

tree into the Singapore Botanic Gardens where it was nurtured by

the botanist Ridley. Research has been the constant accompani-

ment of every step in the development of this industry from the

stage of gathering gum from the wild trees in Amazonia to the

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Vol. XVII. (1958).

highly efficient plantation culture in modern Malaya. One hundred

fifty years ago Hevea was only a botanical curiosity. Palm oil,

cane sugar, quinine, and pineapples are only a few of the plantation

crops that are produced in a very efficient and economical manner

today because of large investments in biological research.

Although medical research has changed the lives of people in the

tropics from continual struggles with a host of parasites no one

would pretend that we have reached the point where long healthy

lives are to be assumed as normal. Tropical medicine and public

health are more than ordinarily dependent on basic biological re-

search because they lack the long background of research possessed

by temperate medicine. We do not even know the complete ecology

and life history of some of our serious parasites and disease vectors.

A few years ago there was a fairly widespread idea that synthetic

chemistry had pretty well taken over the drug field—that plant

products were interesting only as examples of primitive medicine,

not to be compared with sulpha and other synthetic miracle drugs.

Then came penicillin, cortisone, and reserpine and a sudden inten-

sive preoccupation arose with a large number of obscure plants,

from microscopic bacteria and fungi to forest trees. These plants

occur in all parts of the world, but the tropical Pacific has more

than a small share of them, because of the enormous flora of this

tropical region. Research on medicinal plants is slow tedious work,

and much of it never pays off, but the occasional important dis-

covery much more than justifies the entire outlay in work and

money. This is, of course, much more apparent to the sick man

than to the healthy one. Few people who have been cured of pneu-

monia by antibiotics would doubt the wisdom of money spent on

pharmaceutical microbiology.

One of the serious difficulties in tropical countries is the degra-

dation of the soils. Originally it is probable that most tropical soils

were in a state of equilibrium with the biological communities that

occupied them. When man came into the picture this equilibrium

tended to become more and more disturbed. As man’s ability to

affect this environment increased, through his development of im-

plements and technology, his numbers grew. More strain was put

on his environment as more demands arose for its productions.

One only needs to look at the waste places on Singapore Island to

see in aggravated form the results of this pressure on the resources

of the soil. Degraded vegetation and exposed subsoil, capable of

producing almost nothing of value are almost the rule here as in

many thickly populated tropical areas. Although great steps have

been made in preventing and alleviating such conditions in the

temperate zone little has been done in this direction in the tropics.

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Gardens Bulletin, S.

This is primarily a biological problem, one of managing vegetation

and soil biota, of working out wiser methods of using the soil, and

more importantly, of managing or slowing down the growth of

human populations. Intensive research in the biological fields basic

to all of these activities is urgently needed, especially as environ-

mental failure is cumulative.

A specter haunting agriculture, roe and all activities depend-

ing on plants is plant disease. Crop losses and landscape degrada-

tion from plant disease reach enormous proportions. In temperate

areas vast amounts of research in plant pathology are required just

to keep pace with the increasing ravages of plant diseases. In the

tropics there are, at present, more diseases and less research. And

as agriculture and forestry become more intensive, disease pro-

blems increase, rather than decrease. Monocultures provide ideal

conditions for the spread of diseases. New strains of disease organ-

isms come into existence and become established. The same can be

said of insect pests. At the present level of research in plant patho-

logy and entomology we can not hope to hold our own, especially in

the tropics.

So far the discussion has concerned only areas where serious

difficulties are to be corrected or averted. Examples of these could

be multiplied indefinitely, especially by going more into particular

cases. Of importance, also, are areas where improvements in al-

ready tolerable existing conditions may be brought about by re-

search. We have palatable fruits and other food crops in our.

gardens and fields. This does not say, however, that they are the

best possible. Plant breeding has in the past made the difference

between the inferior, often almost inedible wild products, and the

fine domestic varieties that we have. This was accomplished without

much knowledge of the principles underlying the evolution of new

kinds of plants. Now the science of genetics is an advanced discip-

line and future possibilities in plant breeding dwarf completely past

accomplishments. In the past necessity guided the groping plant

breeder. Now, knowledge of the possibilities should provide the

stimulus and imagination can furnish the direction. The same can

be said of breeding of other classes of economic plants and animals.

The ways in which man’s environment can be influenced to make

it more agreeable for him to live in are countless, and many of them

depend on biological research. Most of them, unfortunately, will

take the form of counteracting detrimental changes brought about

by man’s own activities. Man was well adapted to his primitive

habitat. As he has changed this environment it has in many res-

pects become more unsuitable for his occupancy. This does not

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Vol. XVII. (1958).

seem to be necessary but it has happened and is continuing in ac-

celerated fashion. The advice of ecologists is vital to retarding or

reversing this trend. This advice must be based on extensive re-

search, most of which has not yet been done. If man is not to foul

his nest so completely that it will eventually be uninhabitable he

must understand his own relation to his environment. This will

come about by properly oriented ecological research.

So far this discussion has been devoted strictly to matters that

affect man’s material needs, comfort, and well-being. These are

important, but there is all too great a tendency to consider them to

the exclusion of everything else. Traditionally, the peoples of tro-

pical Asia have been able to strike a better balance between the

material and the higher aspects of life than most others. With the

impact of Western culture there is a tendency for the material side

to receive more emphasis. This is not necessarily good. It entails a

loss in some of the virtues inherent in some Asiatic cultures.

Though science is generally thought of as strictly a materially

oriented activity it is fortunately not completely so, either in its

motivations or in the benefits it yields. One or two of the other

facets may be examined to give us a truer picture of the place and

importance of biological research in the present phase of the cul-

tural history of the tropical Pacific region.

The greater part of the people of this part of the world belong to

newly independent nations, after varying periods of colonialism.

During the colonial period the cultural patterns changed, here as

indeed in western Europe, from a mediaeval, essentially feudal

state, more or less self-contained and in equilibrium with local en-

vironments, to modern membership in a highly integrated world

community, completely dependent on the application of scientific

knowledge and accomplishment for its normal functioning. A

heroic, if somewhat frantic, effort is being made by these peoples

to participate in the material and social benefits of modern scien-

tific achievement and technology. This is regarded as an inherent

right. Most of the science at present emanates from the western

world, much of it from former (and present) colonial powers. The

question has been asked if the newly independent peoples are

really ready for full participation in the modern scientific world

community, whether an essentially parasitic status, scientifically, is

proper or justified. The question has even been asked whether the

members of these non-western cultures are capable of scientific

achievement equal to that of their western colleagues.

On the basis of actual quantity and quality of work done in these

countries at the present time there is a certain amount of valid

reason for these questions. Rather little work is done and much of

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it does not measure up to the best produced elsewhere. However,

those of us with experience in these countries and with personal

acquaintance with indigenous scientists there mostly realize that

present inferiority in output results from the cultural situation rather

than from any inherent lack of ability among the scientists. In cul-

tures where in the past there have been few opportunities for scien-

tific work among the local people, poor educational facilities for

them, and no scientific tradition, there is not likely to be immediate

superior scientific accomplishment. If the intellectual climate is not

right for it, and if no prestige is attached to it, science will not flour-

ish. At the present time in southern Asian countries there are few

scientists and these are smothered by administrative duties. This is

especially true of biologists. It is obviously of the utmost import-

ance to correct this situation, and clearly the way to do it is to

emphasize, in every way possible, officially and privately, the im-

portance of biological research. It is clearly essential to show the

_world that countries newly emerged from a dependent status are

capable, both of attracting first-class foreign scientists and of even-

tually training local scientists who are the equals of those in other

countries. This is to be done, obviously, not by creating scientists

by official decree, but by placing such emphasis on research that

young people with high intelligence and with scientific inclinations

will be attfacted into scientific careers and will realize the necessity

of achieving training and standards equivalent to those prevalent

abroad. This is a long-term project and crash programmes are not

the answer. Rather, steady emphasis on research and well-directed

support for it will bring about sound progress.

Finally, it must be pointed out that, important as are the prac-

tical, economic, and political benefits of biological and other scien-

tific research, the fundamental importance of this, as well as other

intellectual activities, lies in the development of the human mind

and in the satisfaction of man’s curiosity and his desire to under-

stand his universe and his place in it. This is the motivation for

all research beyond the solution of obvious immediate practical

problems. The place of pure research as an intellectual activity

and the strength of the urge toward understanding in a culture is

perhaps the best measure of its degree of advancement in the scale

of civilization.

224

Role of Botanical Gardens in the Humid Tropics

and UNESCO’s Programme related to them

By LENNART MATTSSON

Director, U.N.E.S.C.O.,

South-East Asia Science Co-operation Office,

’ Djakarta

LUXURIANT VEGETATION is a prominent feature of the humid

tropical regions and yet very little of it is known. The nature of the

plants making up this vegetation, methods of propagation, growth

requirements, potential use for the benefit of man—timber, pro-

ducts, barks—have not been studied systematically so far. This

handicaps planning and development of agriculture and forestry,

important contributors to the economy of a tropical country.

The little information available at present has chiefly come from

botanic gardens maintained by departments of agriculture or scien-

tific societies. There are at present only a small number of botanical

gardens in the tropical regions. Some of them are:

— Botanic Gardens, Peradeniya, Ceylon

— Central National Botanic Garden, Calcutta

— National Botanic Gardens, Lucknow

— Botanic Gardens, Singapore

— Jardim botanico do Rio de Janeiro

— Botanic Gardens, George Town, British Guiana

— Royal Botanic Gardens, Hope, Kingston, Jamaica

— Botanic Gardens, Bogor, Indonesia.

The gardens are of different size and their activities include

maintenance of a representative collection of the vegetation of a

region, demonstration of principles of conservation of vegetation,

maintenance of nurseries of plants, collection, preservation and

exchange of dried plant specimens for purposes of identification.

It is commonly acknowledged that much of the natural vegetation

of the humid tropical regions is cut down or burnt unmethodically

resulting in erosion of the soil and this results in subsequent

damage by floods. The botanic gardens—the large ones with ex-

tensive enclosures—have a definite role here to play in demons-

trating to the general public the useful role of vegetation in

binding soil, thus preventing erosion and making them useful for

agriculture. It may be added here that in the desert tropical regions,

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Gardens Bulletin, S.

where the advance of sand dunes threatens valuable agricultural

land, demonstration of fixing the sand dunes by appropriate vege-

tation is a definite function of a botanical garden located in a

desert tropical region.

Secondly the gardens can play a valuable role in breeding plants

resistant to various types of plant diseases in nurseries and trial

plots and then distributing such varieties to agriculturists. It can

and does help in introducing plants which thrive in similar environ-

ments in another region to its nursery for the first time and breed-

ing one or more generations of them, thus making them adapted

to their new surroundings. Further some plants are in danger of

complete extinction from the surface of the earth owing to their

inability to adapt to possible changes in the environments in which

they grow. It would be valuable to protect them and to increase

their numbers if possible, and the botanical gardens can play a

vital role in this matter. Several botanic gardens undertake such

_ projects and examples that come to the mind are the Medicinal

Garden in Ganewatte in Ceylon, the Botanic Gardens of Hakgala

and Gampahe, the Botanic Gardens in George Town, British

Guiana, the Royal Botanic Gardens, Hope, Kingston, Jamaica.

Most of the botanic gardens maintain a herbarium wherein type

specimens of plants are preserved for taxonomic studies. This mate-

rial is annotated by competent botanists describing their chief

characteristics, places from where they have been collected and

their scientific names. In order to do this regular expeditions are

undertaken to different areas to collect the plants. Several sets per

species are collected and the specimens are suitably mounted on

sheets of paper to preserve them. Identified duplicates of the spe-

cimens are exchanged with herbaria in other parts of the world.

Preparation of floras of the region are undertaken utilizing the

material preserved in such herbaria. The preparation of the flora

of the Malaysian region is currently in progress and a large number

of herbaria and taxonomists all over the world are co-operating

in the project.

The humid tropics programme of UNESCO intends supporting

the collection, identification, preservation and exchange of mate-

rial in herbaria. Until now, material collected had to be sent

abroad for identification purposes. This has naturally retarded the

regular publication of the floras of tropical regions, important in

knowing the features of the vegetation in the regions.

The first step is the organization of regular expeditions for col-

lection purposes from a large number of selected institutions

located in the humid tropical regions. In order to do so, a number

of specialists have to be trained to undertake regular expeditions.

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Vol. XVII. (1958).

Relatively well-established institutions in the tropics would serve

as key centres for organizing training programmes. UNESCO as-

sistance would be utilized to finance this training programme, by

provision of foreign specialists, equipment, etc. It would also bear

the costs of publication of results.

UNESCO’s Advisory Committee for Humid Tropics Research

at its first session in Manaus (Brasil) in July 1957, recommended

that the Secretariat study the practicability of establishing such

key herbaria. Information from a large number of herbaria in the

humid tropical regions was collected regarding the nature and

total number of specimens, annual additions, sources of specimens,

identifications carried out and exchange of specimens effected

during the last three years. More than 100 institutions furnished

this information, and it is hoped that a short list of suitable institu-

tions for development as key herbaria will be established by the

Advisory Committee.

Depending on the finances available, it is hoped to conclude

agreements with these institutions to make arrangements for

organizing training programmes.

Another form of UNESCO’s activities in the field of Humid

Tropics research in which we collaborate closely with the botanic

gardens, is the convening of scientific symposia under the auspices

of this UNESCO Office, e.g. the Symposium on the Study of

Tropical Vegetation in Kandy, Ceylon, in March 1956, which was

organized jointly by this UNESCO Office and its sister office in

New Delhi; the Symposium on the Phytochemistry of Tropical

Plants in Kuala Lumpur, Malaya, in December 1957, and the

Symposium on the Vegetation of the Humid Tropics, held in

Bogor, Indonesia, in December 1958.

It may also be mentioned that this UNESCO Office has on

several occasions awarded grants to facilitate the exchange of re-

search workers between botanical gardens and herbaria in the

tropical regions.

It will be evident from the foregoing that UNESCO is deeply

concerned with the development of botanic gardens. Their colla-

boration is indispensable for the implementation of the UNESCO

programme in Humid Tropics research. The UNESCO Science

Co-operation Office for South-east Asia is happy to express its

appreciation of the scientific work done by the Singapore Botanic

Gardens. We congratulate the Gardens on the Centenary and

express our hope for fruitful collaboration in the future.

227

Regenerating High Forest on Singapore Island

By H. B. GILLILAND AND M. J. WANTMAN

University of Malaya, Singapore

Introduction

NOTES ON SECONDARY or subseral vegetation on Singapore Island

were given in a previous paper (Gilliland, 1958). Most of the

island is covered with such vegetation today and the causes of this

are made patent from the history of gambier, pepper and coffee

cultivation and timber and fire-wood exploitation as recorded by

Burkill (1935, p. 2201). The principal surviving relic of primary

high forest has long been preserved first as a forest reserve by the

Forest Department of the Straits Settlements, and later as a nature

reserve by the Nature Reserves Board, Singapore. This relic of

some 152 acres covers the upper slopes of the island’s highest hill,

Bukit Timah, and is the type-locality for a remarkably large num-

ber of species. The possibility of the existence elsewhere on the

island of small relics of primary high forest was made manifest

from a study of aerial photographs. A large area jointly maintained

as a Nature Reserve by the City Council and the Nature Reserves

Board comprises the catchment area of the three principal reser-

voirs of the City’s water supply. The southernmost of these is the

MacRitchie Reservoir and the photographs suggested that the vege-

tation along its southern shore would warrant study. Such study

reveals the presence of a forest of Dipterocarp dominated type

which has not been so exploited as to alter radically its character.

This paper discusses the results of the application of a line transect

sampling technique to this forest.

History

Enquiry from the City Water Engineer reveals that this Reser-

voir first came under the control of the City Council Water Depart-

ment in 1857.

The Office of the Commissioner of Lands, Singapore, records

that in the same year the piece of land on which this forest now

stands was granted to Mr. Sim Wye Tye (Indenture No. 19) and

Mr. Tan Ah Pok (Indenture No. 84).

Further sales of this land, of the first portion in 1876 and of the

second portion in 1858, to further owners are recorded but in

1890 the Municipal Commissioners acquired all the land in this

area. The acquired land was surrendered to Government and on

228

Vol. XVI. (1958).

2nd June, 1906 was re-issued with new title to the Municipal Com-

missioners. .

In 1910 the bridle path, which currently traverses the forest

from west to east, was cut and in 1933 the road “Lornie Road”

which, today, forms the southern boundary of “MacRitchie Reser-

voir Jungle” was constructed.

It is reasonable to conclude, especially in light of the more recent

building of the paths and roads, that such occupation of the land as

did occur was not wholly destructive of the then-existing vegeta-

tion. However, the existence of an old road crossing the reservoir

from north to south,-recently revealed during a drought period,

indicates that communications, even if of different pattern from the

present, did exist previously.

Principal Trees

There are many large trees in the area and specimens of the

following with a diameter at breast height, or above buttress, of 2

feet or more (76 in. girth) were noted.

1. Adina rubescens 20. Ficus consociata v. murtonii

2. Alstonia angustifolia 21. Gordonia singaporeana

3. Anisoptera megistocarpa 22. Heritiera simplicifolia

4. Aquilaria malaccensis 23. Irvingia malayana

5. Artocarpus scortechinii 24. Koompassia malaccensis

S Rep a pia 25. Melanorrhoea woodsiana

8. Artocarpus rigidus 26. ae aise

9. Calophyllum floribunda 27. Palaquium rostratum

10. Campnosperma auriculata 28. Palaquium obovatum

11. Castanopsis hullettii 29. Pygeum polystachyum

12. Cinnamomum iners 30. Quercus brevistyla

13. Cryptocarya griffithiana 31. Santiria laevigata

14. Dipterocarpus hasseltii - 32. Santiria tomentosa

15. Dipterocarpus apterus 33. Santiria griffithii

16. Dipterocarpus kunstleri 34. Shorea ovalis

17. Dyera costulata 35. Shorea glauca ©

18. Dysoxylum mollissimum 36. Shorea paucifolia

19. Fagraea fragrans 37. Vitex pubescens

It seems reasonable to conclude on the basis of such mean annual

increment data as are available (Edwards, 1930) that none of

these trees is likely to be older than 100 years. This finding agrees

with the supposition that following the original alienation of the

land little more than the extraction of standing timber was done—

the present trees would then represent the successful subsequent

growth of existing regeneration (Richards, 1952).

Sampling Method

To contrast this with the secondary forest already examined

(Gilliland, 1958) it was decided to make a similar line transect

229

Gardens Bulletin, S.

sample. The sample consists of a belt or transect 1,000 feet long

and 6 feet wide and is obtained as follows:—

(a) It is divided into 10 sub-samples each 100 feet long, each

sub-sample taken at random.

(b) Each sub-sample is obtained by laying down a surveyor’s

100 foot chain. Thereupon the No. 1 of the pair of ob-

servers prepares four 3 feet wands which are spaced

and respaced along the chain to demarcate convenient

portions 6 feet wide. No. 2 of the pair of observers acts

as amanuensis recording data, and the team works its

way along the chain.

that they may be recorded in the size categories 0-1;

1—3; 3-5; 5—7; and > 7 inches.

(d) A specimen of any seedling, sapling, or tree which cannot

immediately be identified is numbered and collected for

identification.

It is wise to make two practice 100 foot samples before commenc-

ing on the serious record in order to become familiar with the

principal plants present in all their stages.

A sub-sample takes between 1 and 2 hours to record in the field.

As soon as the data for a particular sub-sample have been obtained

work on the “unknowns” begins. This may occupy a further week.

Not till this is complete does work on the second sub-sample begin.

Identification was greatly assisted by the knowledge and help of

‘Che Ahmad bin Hassan and Tuan Haji Mohammed Nor bin

Mohd. Ghous both of whom had served on the Botanic Gardens

staff for over 40 years. Much time and patience is clearly required

to complete a sample.

As noted above 10 sub-samples were used for the collection of

the data. The question then arose as to whether or not one can

generalise from the 10 sub-samples and draw conclusions about the

total area from which the sub-samples were taken.

The first step was to determine whether or not these 10 sub-

samples could be treated as a random group of samples from the

general area. The intention was to test this by the analysis of vari-

ance technique. One of the requirements of the analysis of variance

technique is that the variability should be similar from one sub-

sample to the next. Putting it another way the 10 sub-samples

should be homogenous with respect to the variability of the fre-

quencies. This test for homogeneity was carried out by means of

Bartlett’s test and the samples proved not to be homogeneous by

this test. The results are presented in table I.

230

Vol. XVII. (1958).

TABLE I

MEANS AND VARIANCES OF THE FREQUENCIES OF THE 205

SPECIES IN THE 10 SUB-SAMPLES.

Sub-sample Mean Vi 6s

ig ax hurtes Frequencies

1.33 21.90

1.53 19.45

3 2.43 52.98

4 2.42 41.70

5 2.63 79.87

6 2.74 49.16

7 2.80 67.40

8 2.05 30.81

9 1.82 17.86

10 2.43 44.10

Bartlett’s Test: X*= 236.64

P<.001

The result of Bartlett’s test therefore made the application of the

analysis of variance “F—test’” questionable. The decision was

made then to test all the possible differences between the means

of the sub-samples by the “t’—test of significance. Forty-five

t-tests then had to be carried out. With the 0-05 level of significance

as the critical value, if the hypothesis of random sampling is ten-

able, there should be no more than 3 differences large enough to

be judged significant at the 0-05 level and beyond. When the 45

tests of significance were carried out 3 differences barely reached

the -05 level of significance. These differences were between sub-

sample 1 and sub-sample 6, sub-sample 1 and sub-sample 7, and

between sub-sample 2 and sub-sample 6. No difference reached

the 0:01 level. Thus with respect to the means of the 10 sub-sam-

ples one can retain the hypothesis that they are random samples

from the same population.

It is interesting to note that in spite of the violation of the homo-

geneity of variance test and in spite of the distributions being non-

normal the overall F-test shown in table II clearly supports the

conclusion above that the means can be assumed to be drawn

from the same population.

TABLE II

ANALYSIS OF VARIANCE TEST OF RANDOM SAMPLING.

Sums of Squares d.f Variance

Estimate

Between sub-samples gt 485.95 9 53.99

Within sub-samples oR 86,742.58 2,040 42.52

Pa" 27

P> +05

231

Gardens Bulletin, S.

The data were then rearranged from the index cards and each of

the 205 species was classified according to its “B” (belongingness )

value where the value of “B” is the number of sub-samples in

which that species occurred; thus “B” = 7 means the species has

been recorded in 7 out of 10 sub-samples and “B” = 2 means the

species has been found in 2 sub-samples only. A “B” value was

determined for each species. |

It is clear from the above definition of “B” that one would expect

a positive correlation between the frequency of a plant and its “B”

value, but there is no reason to expect this relationship either to be

linear or perfect. Table III shows the scatter diagram of the two-

way plot of “B” values and frequencies. The positive relationship

and its non-linearity can readily be seen from the table.

TABLE III

SCATTER DIAGRAM OF ‘“‘B’? VALUES AND FREQUENCIES.

*“B’’? VALUES

“*frequencies”’ 1 2 3 4 5 6 i 8 9 10

200-299... ae ae et bs iy aie Yes rT , 2

ig0- r as ts $e > a - + 3h dh

160-179... on are oe a3 $3 i. Zi dita: 2} 1

140-159... - fs ay, ae bb nis 7 eee 1

We Ie > RE ois a Ay ae ie of os LT 1

100-119... ms - Ee Be a 33 st Ft 6 3

90-99 + as ite Lid ys ie sf a ™ a I

80-89 UE nes a bre 2 a Me re sa 2

70-79

60-69 st iF ce ‘3 ban ok

59-59 ra ” sa of a |

40-49 i Es ee

35-39

30-34 is vs eT

25-29 de on I

20-24 -* .s es

15-19 #5 edi I

10-14 ot se ,

mem bh

mh ee

—

= a ee eee oS

creeee: «> lane

Bee ap tee

—=hv

KH NWAUAD~ 100

~ WNN Ww .

—__ -- | —_ — _ -| _.. ——— | —__. | —— — | —— | | | |

Totas..: Ta) 2h 2S 16 14 {9 9 Ole as

(Note:—The grouping interval of the frequency variable has not

been kept constant in order to show the non-linearity more clearly).

Loe

Vol. XVII. (1958).

A priori considerations had led to the tentative conclusion that

plants having a “B” value of 7 and higher were more likely to

“belong” to that particular area than those with “B” values of 6

and lower. This hypothesis suggested dichotomizing the “B” vari-

able and summarizing the relationship between frequencies and B

values by means of a statistic such as the biserial product moment r.

The formula for r is as follows:—

ee Ge = —

/ q 6r

where p = proportion of plants in the group with the higher B

values

hamid oP

My = Mean frequency of the higher group

M+ = Mean frequency of the total sample of plants

6, = Standard deviation of the frequencies of the total

sample.

An alternative formula for the biserial product moment correla-

tion coefficient may be written as follows: —

a OT’

where the symbols are defined as they were for the formula above

and My; equals the mean frequency of the group with the lower

B-values. From the alternative formula it can be seen that the

magnitude of the r is a function of the difference between the

mean frequencies of the higher and lower groups. The use of

the biserial product moment r has the advantage over the simple

difference in that it takes into account the variability in the total

group and the proportions in the two groups thus making possible

comparisons from one investigation to another.

Even though the original hypothesis had suggested dichotomizing

at B=7 and higher to obtain the higher group, it was decided to

investigate the correlation at other possible points of dichotomy.

The results are presented in table IV.

233

Gardens Bulletin, S.

TABLE IV.

BISERIAL PRODUCT MOMENT CORRELATION AT DIFFERENT

POINTS OF DICHOTOMY.

Pt. of dichotomy p. M rum ir

B=10 a fi, 0537 110.66 .60

B= 9 and 10.. Si .0829 100.47 —-.68

B= 8 and higher eh .1268 74.20....5al

B= 7 and higher i .2195 56.71 .68

B= 6 and higher Ped .2878 43.24 bz

B= 5 and higher bi 3659 3759.

B= 4 and higher .4000 2607 ae

(Note:—Computations based on ungrouped data of table IIT).

The highest correlation viz. -68 is obtained with the dichotomies at

B=9 and B=7. It should be obvious that the B=7 is a more

desirable point of dichotomy.

The biserial product moment r as a summary statistic of the

relationship between frequency and B value was examined for

data from two earlier investigations in order to check whether or

not the dichotomy at B=7 and higher would have been best for

those data. In one of the earlier investigations (Gard. Bull.

XVII 1958) the r’s for dichotomies at B=8, B=7, B=6, and

B=5 were -49, -46, -45 and -39 respectively and in the second earlier

investigation, the r’s for dichotomies at B=8, B=7, B=6 and

B=5 were -61, -56, -49 and -41 respectively. (Sarawak cf. Trans.

Bot. Soc. Edn. 1959). These results would lead one to retain the

B=7 dichotomy since they do not conflict with the present study

and since the B=7 dichotomy was chosen on the basis of other

considerations prior to the statistical analysis.

Having established the dichotomy at B=7 the 205 species can

now be separated into two groups:—

Group I those species with B=7 and higher.

Group II_,, s » B=6 and lower.

This dichotomy resulted in 45 species being classified in group I

and 160 in group II. The species in group I clearly represent the

important plants of the present community; the group II species are

those occurring less regularly and in smaller frequencies.

The 45 species in group I were further classified into A. Trees,

B. Shrubs, C. Lianes, and D. Herbs.

The 160 species in group II were classified into Group G—22

species and Group H— 138 species. This further classification is

presented below.

The main defect in the description of the community is the

absence of quantitative data on epiphytes.

234

Vol. XVII. (1958).

Since the data for the 10 sub-samples were combined for each of

the six groups A, B, C, D, G and H, the question arises as to

whether or not each sub-sample can be considered with respect to

the group under discussion as random sub-samples from the total

sample.

The results are summarised in Table V.

Results

TABLE V.

No. of | No. of % of total

specs. pits. pits.

( Group A. Trees ii 16 1,459 32.0)

“B” = 7 ~ 3B. Woody Planis .. 17 1,096 24.1 179

or more «» <.. Lianes “ 11 987 Aled :

4 » D. Herbs s I 8 0.1 J

mpeg | GEG. —~. BAS CMO» EA 60)

ofl 1 ont et im bya; ios)

205 4,546 99.8

Bartlett’s test for homogeneity of variance again showed a lack

of homogeneity in the five groups tested (the D group had to be

omitted because of only one species in it). Thus the application

of the F-test for an overall test of differences was again question-

able. The forty-five mean differences were therefore tested for sig-

nificance in each of the five groups. A summary of results is given

in table VI. With the chance expectancy of no more than

3 “significant” differences among 45 at the -05 level and beyond,

the results showed no differences at the -05 level or beyond for the

A, B and C groups. For the G and H groups there were 4 and 6

differences respectively which were beyond the -05 level although

none of them reached the -01-level. When the A, B, C and D

Groups are combined, only 2 differences reach the -05 level; the

combination of G and H groups yields 4 differences at the -05 level

and 2 beyond the -01 level.

These results suggest that the species in the A, B, C and D

groups differ from those in the G and H groups in this forest area.

Thus, all of the statistical analyses confirm the conclusion that the

species in the A, B, C and D groups “belong” to this forest area

while those in the G and H groups do not.

235

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236

Vol. XVII. (1958).

Group A, comprising those species whose mature individuals reach

to canopy height, is listed in Table VII. Clearly dominance in this

group could refer to the Shorea spp. but when Table XIII with a

total of 513 Dipterocarps is taken into account the “family domi-

nance” of the Dipterocarpaceae seems clear. It should be noted

however that members of the Guttiferae (which includes Garcinia

and Calophyllum) total 682 plants. Most of these (677) however

fall into the first size class, the seedlings with a stem diameter less

than one inch, and furthermore trees of these genera rarely reach

to the height of the canopy.

TABLE VII.

GROUP A.—TREES

| DIAMETER CLASSES

Species | toiat | “3”

0-1 1-3 | 3-5 | 5-7 | Over7

Shorea macroptera... 178 ue ee he 3 181 9

Shorea parvifolia x Bie Ba ‘Wes Fea 8 eda 173 lack

Shorea parviflora oe 83 want pany pay. hs * 83 10

Gironniera nervosa .. we Z eee bo. J Dstiladins A 2 8

Pygeum polystachyum 124 3 em ae ae Se 128 10

Onchosperma horridum 99 pray 2 6 errs aid 10

GMcimeeentitie ene 3d | PIPET Mo ee Oa eas os

Santiria tomentosa ie 34 2 terre t Sate git ears 40 7

seis tan | ad oe ed bo. 6 | 8

Quercus or marr marae pelt ee, theses I 8

Picea, | (eta. |) ..d\he.. | is| 8

meaede ee ag al td) ld Re | |. nibebi@olnng

Coelodepas wallichianum| 158 Ria), AGh pe | of ws AG, 10

Melanorrhoea woodsiana iia meee Peri aay igi Senge in 10

Gomphandra affinis .. waa erat + ie Yi Bike) yd the fart 48 7

Elateriospermum tapos maya £: ee. Ti ey Bey 14 7

ren ey tad |) a eat iho) |S 1.450

Gardens Bulletin, S.

Group B, Table VIII lists the shrubs and small trees which do not

reach to canopy height or potential trees which have not yet ex-

ceeded sapling size. The shrubs characteristic of this community are

clearly Agrostistachys sessilifolia and Aprosa benthamiana (both

Euphorbiaceae ).

TABLE VIII.

GROUP B.—SMALL TREES AND SHRUBS

Species

Agrostistachys sessilifolia

Aporosa benthamiana

Cryptocarya griffithiana

Calophyllum pulcher-

rimum .. Pe

0-1

234

Calophyllum inophy-

lloide

var. singapurense

—

Calophyllum ferrugineum

Xanthophyllum discolor

Garcinia parvifolia

Garcinia eugenifolia ..

Scorodocarpus borneen-

sis <3

Anisophyllea disticha

Clerodendron dispari-

folium .. re

Myristica crassa

1-3

— ee | | | | — § | |

Acronychia porteri ..

Eugenia longiflora

Eugenia sp. ..

Aquilaria malaccensis

DIAMETER CLASSES

“iRee

Total | Value

3-5 5-7 | Over7

a; 238 | 10

54] 10

Tepe ©

182} 9

149 | 9

"ep Tah eke Beak ie

TT aim

hee 52| 9

erie roe ie

f 45.00

i %1° 9

oe | |g. RE A ak, ok cee

" 200| 7

a i9| 8

“y i2.|. 8

PS ee ewe fc cine...

Pra a eer: Gi

Vol. XVII. (1958).

Group C, Table IX gives the lianes present and the prominence of

the family Connaraceae is noteworthy and is indicated by * in the

table.

TABLE IX.

GROUP C.—LIANES AND CLIMBERS

DIAMETER CLASSES

Species ______ Total | im

0-1 1-3 3-5 5-7 | Over 7

Willughbeia coriacea .. 285 wat ne a wr 285 10

Rourea sp.* ~ 150 ite RANT) saa —... ASO. w

Rourea fulgens* ‘aaa 85 am [ Piszi9: TAS ib 85 10

Agelaia sp." pA tee Ue PEN 2 tal eeaiiae ‘eek iia

Psychotria ovata Be 109 Post "ENed ey we 109 10

Connarus sp.* ee 64 1 I Carrs Pe foe ee weir od zs

Fibraurea chloroleuca a 51 pei i Latte OR begins 51 7

Piper caninum - 35 eet Synth: PD ays sid 35 7,

Tetracera indica ss 26 ee ae. ‘aes f ov 26 2

Ventilago sp. as 19 Sa afin pie es 19 a

Vitis macrostachya wi 15 oe Ceti lee | CaN whine a

Group D, Table X the true herbaceous plants—i.e. those which are

not seedlings or saplings of trees, shrubs or lianes, are represented

here solely by the fern Tectaria singaporeana.

TABLE X.

GROUP D.—HERBS.

Species Total V fe id

Tectaria singaporeana @. 8 7

Group G, is listed in Table XI. This group is distinguished from the

remainder of those with “B” = 6 or less by the fact that some of

them have a stem diameter greater than 1” at breast height. At

least the first 8 are large enough to be reproducing themselves and

the comparative paucity of their seedlings is noteworthy.

259

Gardens Bulletin, S.

TABLE XI.

GROUP G.

Saas DIAMETER CLASSES

Of eAsSe :345.1|- S279 Oven7 | Total ae

Anispotera megistocarpa 8 ae 3. 1 1 10 5

Dialium maingayi ee. 2 e+ Si ae ity 1 ioe se 1

Dipterocarpus Fonrucosus 1 “se ey Se ee _*, niet

Dipterocarpus hasseltii 7% aan Tae egal 1 ss gee apy

Litsea grandis + PZ 3 l aera: eee toed 16 5

Macaranga lowii ed 8 eh. | = “anes 9 3

Timonius wallichianus el ar ae oa ~ pee 3

Diospyros lancifolia .. ee: PE: oe aad - 3 3

Eugenia sp. 4. a ae a Roun) pee: He Be -—§—

Litsea sp... a ey ee a a re ie er

Polyalthia anna Gsstaial eas 3 cal a eas. Zi 6

Aghia claiifiorse) li; ane be | 5

Horsfieldia subglobosa ee an aes | eee ee 17 3

Zygia jiringa eee Bae: ee posal = ce “ tg —

Ochauestachys amen iS: 1 a IRI SEE ct SEE FS

tacea) =~. rie 11 I a oe e 12 5

Bugenia;polyantha »4iell id@ates3. |» 0.1 1 aaatiana so popieeaen men

Hopea megarawan .. Igy peg i ae em st ot'y FTE 4

Elacocarpus sp... | (A to] 4.) ee oe

Baccaured parvition! o» Te i, _— ee Parner aes & 3

Galeatia'spyo- 2-00) aleetea--| ai seal a

Knoma hookerianay ../{ a2” |. | a ares

Salacia flavescens.. uae ie eae: tet dus bape 3

alu) SARE We er Re

In contrast Group H, Table XII, with a total of 138 species repre-

sents plants which are seedlings or saplings of larger plants or

rarely, and these are marked with an asterisk, Herbs of the forest

floor.

240

Vol. XVII. (1958).

Species

Actinodaphne pruinosa. .

Adinandra dumosa

Aeschynanthus sp.

Aglaia trichostemon

Aglaia sp.

Aglaonema schottianum

var. angustifolium* ..

Alsodeia floribunda

Antidesma alatum

Aporosa sp.

Apostasia nuda*

Aquilaria hirta

Ardisia teijsmanniana ..

Ardisia spp.

Artabotrys costatus

Artabotrys susveloens ..

Arthrophyllum — diversi-

folium

Artocarpus anisophyllus

Artocarpus dadah

Artocarpus integer

Artocarpus lancifolius ..

Artocarpus scortechinii

Athyrium cordifolium ..

Blumeodendron tokbrai

Byttneria maingayi

Caladium denudatum ..

Calamus ornatus_ var.

horridus

Calamus javensis

Calophyllum sp.

Canthium horridum

Centotheca latifolia*

Centrosema sp.

Champereia manillana .

Chasalia chartacca

Chisocheton sp.

Cnestis palala

Cordyline sp.

Cratoxylon formosum .

Croton laevifolium

Curculigo latifolia*

Cyathocalyx ridleyi

Cyathostemma viridi-

florum

Daemonerops sp. :

Desmos dasymaschalus. .

Demsos dumosus

Dioscorea pyrifolia

Dipterocarpus sublamel-

latus

Dracaena granulata

Dracaena umbratica

TABLE XII.

GROUP H.

Total | B Species

Elaeocarpus ferrugineus

Elaeocarpus mastersii ..

Elaeocarpus petiolatus . .

Elaeocarpus stipularis . .

Elaeocarpus sp.

Ellipanthus tomentosus. .

Eugenia garcinifolia

Eugenia grandis

Erycibe maingayi

Ficus callicarpa

Ficus sp. ths

Fissistigma latifolium

var. ovoideum

Fissistigma fulgens

Fordia filipes at

Galearia phlebocarpa ..

Garcinia gaudichaudii . .

Garcinia mangostana ..

Garcinia sp.

Glycosmis chlorosperma

Gnetum macrocarpum .

Goniothalamus macro-

phyllum ..

Goniothalamus malaya-

nus ‘

Guioa pubescens :

Gymnacanthera bancana

Gynotroches axillaris ..

Helicia excelsa

Homalomena rubra*

Hoya ridleyi

Ilex cymosa

Ipomoea maxima

Ixonanthes icosandra

Ixora congesta

Ixora sp.

Knema communis

Knema malayana

Korthalsia sp.

Labisia pothoina* ie

Lasianthus densifolius ..

Lasianthus sp. I

Lasianthus sp. II

Lasianthus sp. III

=e hon

—

et eet CU

Qe td)

We ne S&S NS na — WD

—

NNENA—fhK Lh

—

— ie NNR RWW a

peck, pe Fa Jom em fed oP UD Fed ee KN GN UD NN—NN WwW Nn

5 2 || Licuala sp.

21 6 |} Lindera sp.

1 1 Luvunga sarmentosa

3 3 || Macaranga javanica

2 2 || Mangifera sp.

Nephelium lappaceum A

1 1 || Oxymitra calycina é

1 1 || Palaquium obovatum ..

12 6 || Pandanus ornatus

241

N —

NOOR — NN OO NWR KAN SK ENE

—

—y

—

——s

WwW OORN EEE OAS ENREANDOHENWOEEENENE ES

—

mee OD ON ND

NAE RE RAWA

—

NWNRK SB KHUN NRE RK NWPhRehK WNW REE NEN

Species

Parkia speciosa

Phaeanthus opthalmicus

Philodendron sp.

Phymatodes scolopen-

drium* . Le

Abarema elliptica

Plectocomia griffithii

Polyalthia cauliflora

Psychotria helforiana

Psychotria maingayi

Psychotria sp.

Pternandra sp.

es ei tea singaporea-

na i:

Quercus sp.

Randia arisophylla

Randia incurva

Randia longiflora

Randia macrophylla

Raphidiophora sp.*

Rhodamnia trinervia

Species

Shorea macroptera ..

Shorea parvifolia

Shorea parviflora

Shorae sp. No. 4

Anisoptera megisto-

carpa

—

Hopea mengarawan

Dipterocarpus verru-

cosus yp

Dipterocarpus hassel-

tii 4

Dipterocarpus sub-

lamellatus é

Total

Gardens Bulletin, S.

GROUP H—continued

DIAMETER CLASSES

0-1

1-3

35°) Sey yee Total

3, 10° 18

} 52

1 1 10

fans WG

he 1 2

is 1 1

0 1 ne Derk

mRANN new

mmm ee et ee AQ ee Re

Total | B Species Total

511 588

3 3 || Sandoricum koetjape .. 4

1 1 || Scleropyrum ridleyi .. 1

2 1 Shorea sp. No. 4 52

Sloetia elongata a 2

2 2 || Smilax calophylla a 3

7 5 || Strombosia rotundifolia 5

4 1 || Strychnos ovalifolia 15

8 3 || Strychnos sp. 3

1 1 || Tabernaemontana malac-

1 1 censis .; &, 26

1 1 || Urophyllum glabrum .. 6

| 1 || Urophyllum _ griffithia-

num ae 1

1 1 || Vatica wallichii 1

1 1 || Vitex pubescens 1

9 4 || Vitis gracilis 2

1 1 || Wikstroemia ridleyi 1

10 5 || Xanthophyllum sp. 1

20 5 || Xylopia ferruginea 1

D 2 || Zizyphus horsfieldii 7

2 2 |} Unknowns .. z

588 722

TABLE XIII.

DIPT EROCARPACEAE.

Vol. XVII. (1958).

Table XIII summarises all the data of the family Dipterocar-

paceae, totalling 513 records. This total, 513, represents 12 per

cent of the total of plants recorded from the transect.

In conclusion the naming of the community gives little difficulty

in terms of previous proposals (Gilliland, H. B., 1958). This is a

Shorea/ A grostistachys community.

References

BurKILL, I. H. 1935. “A Dictionary of the Economic Products of

the Malay Peninsula.” London.

GILLILAND, H. B. 1958. Gard. Bull. Singapore. XVII. 1. 82-90.

1959. Trans. Bot. Soc. Edinb. 38, 56—63.

KELLEY, T. L. 1947. “Fundamentals of Statistics.” Harvard Univ.

Press. Cambridge, Mass.

GREIG-SMITH, P. 1957. “Quantitative Plant Ecology.” Butter-

worths, London.

RICHARDS, P. W. 1952. “The Tropical Rain Forest.” Cambridge

Univ. Press.

SNEDECOR, G. W. 1946. “Statistical Methods applied to Experi-

ments in Agriculture & Biology.” Ed. 4. Iowa State

Univ. Press. Ames, Iowa.

Epwarps, J. P. 1930. “Growth of Malayan Forest Trees.”

Malayan Forest Records. No. 9. Singapore.

243

The pH of Rain Water from the Botanic Gardens

By R. A. ROBINSON

University of Malaya,

Singapore

THE HYDROGEN ion concentration of rain water is always very

small, typical samples containing from 10° to 10-8 moles of hydro-

gen ion per litre. Because the values are so small, it is convenient

to represent the data in terms of pH, viz. the negative of the loga-

rithm of the hydrogen ion concentration. On this scale, we would

say that typical samples of rain water range from pH 5 to pH 8.

If the pH is 7, we say that the rain water is neutral, if it is less than

7 it is acid and if it is greater than 7 it is alkaline.

Chemically pure water has a pH of 7 but rain water in the course

of its condensation in clouds and passage through the atmosphere,

can dissolve gases such as carbon dioxide, oxides of nitrogen and

ammonia, which contribute to the pH, and also dust particles

normally present in the atmosphere or discharged into the atmos-

phere during building operations, etc. The pH is also affected by

any trace of alkali dissolved from glass containers.

The pH of the rain water falling in the Botanic Gardens has

been examined over a three year period, March 1956 to February

1959 and, during the last ten months measurements have also been

made on rain water samples collected at the University of Malaya.

pH measurements were made in the usual way with a glass elect-

rode standardised by a phosphate buffer. These pH values are not

equilibrium properties for they change with time as dissolved gases

escape and a further supply of gas is absorbed from the atmos-

phere; nevertheless, if the collection of samples and the method of

measurement are uniform, the results should have some signifi-

cance.

The results are too numerous to tabulate in detail but a sum-

mary is given in Tables I and IJ. Some features that stand out are

as follows: —

(1) There is a very wide variation in pH even in a period of

one month. The average spread in a month was 1-45

pH units but in February 1957 the pH varied from 5-03

to 7:81, a difference of 2:78 pH units whilst in July

1958 it varied only from 6-71 to 7-32, a difference of

244

Vol. XVII. (1958).

only 0:61 pH units, in the Botanic Gardens although

values between 4:85 and 6:52 were observed with the

University samples. The highest value recorded was

8-16 (19 November, 1957) and the lowest was 4:58

(14 November, 1958), the latter being at the Univer-

sity site.

(2) There is an astonishing difference between the pH values

found for samples taken in the Botanic Gardens and

those from the University site only seven chains away.

In all months from May 1958 to February 1959, the

pH was between 0:44 and 1:38 pH units lower at the

University site, the average value in the Botanic Gar-

dens being 6:80 compared with 5:86 in the University.

Whilst the total rainfall was about the same at the two

sites—89-61” in the Botanic Gardens and 87-48” at the

University, there is considerable difference in the distri-

bution from month to month. Thus in June, 1958,

11-89” were recorded at the University against 11-90”

at the Gardens but in November 1958, 18-75” fell at

the University site compared with 21-65” at the

Gardens.

An examination of the records of thunderstorms made at Singapore

airport (some distance from the Botanic Gardens) shows that on

the 139 days of thunderstorm on which samples were obtained

from the Gardens, the average pH was 6:56 compared with 6-65

for the total of 548 samples. Some diminution in pH such as this

would be expected as the result of the formation of oxides of nit-

rogen during thunderstorms.

On the days when no thunderstorms were recorded, there is

some indication that a rainfall of less than 0:2” gives rainwater of

average pH about 6-85: there seems to be no appreciable variation

if the rainfall is between 0-2” and 2:25”, the pH being about 6-43

but on the six occasions only on which a higher rainfall occurred,

the average pH fell to 5-86, approaching the value, 5-68, expected

for water saturated with carbon dioxide in equilibrium with atmos-

pheric carbon dioxide. It would be interesting to confirm this with

more samples.

(3) The month of January 1957 gave inexplicably low pH

values, averaging 5-92, at the Botanic Gardens.

245

Gardens Bulletin, S.

(4) Twenty-two times during the period May 1958 to January

1959, we were able to obtain not only composite sam-

ples from the Botanic Gardens but also the final sample

left in the Recording Rain Gauge. The results are given

in Table III. The first ten results tabulated refer to days

in which more than one inch of rain fell in the Botanic

Gardens and thunderstorms were noted at the same

time at Singapore Airport. The rest of the results refer

to days on which more than an inch of rain fell but no

thunderstorms were noted. With the reservation that

only a small number of results are available, some con-

clusions are suggested.

On days without thunderstorms, the average pH, 6-44, of the

composite samples was almost that of the final sample, 6-43. We

have noted in section (3) that over the entire period of the tests

rainfalls between 0:2” and 2-25” tend to give a pH about 6-43.

We might have expected a lower pH for the final samples from the

rain gauge, in view of the statement in section (3) that rainfalls

greater than 2-25” tend to a low pH of 5-86 but it will be noted

from Table III that the value of 6-43 for the final samples from the

rain gauge is based on twelve samples and in only one instance

did the total rainfall exceed 2”. The findings in this section and in

section (3) are, therefore, not contradictory. The average pH of

the composite samples on days of thunderstorms was 6-31: over

this period the average of all samples was 6-80. This would suggest

a considerable diminution of pH as a result of thunderstorms,

greater than the difference noted in section (3). However, the

average of 6-80 was deduced from all samples, even if the rainfall

was only a few hundredths of an inch: we have already noted that

such small rainfall has comparatively high pH, thus raising the

average value. A fairer comparison would be between days with

and days without thunderstorms, each with over an inch of rain;

i.e. we compare an average of 6:31 in the first instance with 6-44

in the second. The influence of thunderstorms is now more consis-

tent with that noted in section (3).

The average value of the final samples from the rain gauge on

days with thunderstorms was 6:46, close to that on days without

thunderstorms: this suggests that the first inch of rain succeeds in

washing the atmosphere free of oxides of nitrogen.

246

Vol. XVII. (1958).

Table I

pH OF RAIN WATER FROM THE BOTANIC GARDENS

Number Monthly

H H iH of days 7

2, fverase) (minimum) (tite) ie ieee (i ee

rain fell

1956

March a 6.41 5.70 6.95 11 4.28

April .. a 6.72 6.30 7.09 10 2.68

May .. c 6.57 6.23 i 19 10.49

Jiitte. i sd 6.59 5.81 6.87 13 5.67

duly... - 6.36 5.95 6.79 16 13.82

August aH 6.40 5.29 6.92 24 8.90

September a4 6.36 5.18 7.19 19 12.05

October ss 6.26 4.95 7.48 20 4.82

November ae 6.66 5.50 8.16 21 19.30

December of 6.73 6.19 7.09 17 8.32

1957

January rs 5.92 4.69 6.88 14 4.81

February + 6.56 5.03 7.81 8 4.09

March ‘2 6.28 5.38 7.42 13 6.98

April .. Si 6.89 5.84 7.28 9 3.91

May .. Le 7.11 6.40 wey ie 16 12.20

June .. ox 6.47 5.61 pe 10 2.92

July .. ie 6.67 5.89 125 15 8.98

August 4 6.71 6.32 7.18 15 4.65

September on 6.79 6.09 7.69 19 10.27

October 3 7.11 6.67 7.67 13 2.33

November a 6.69 5.82 7.51 21 6.45

December a 6.79 4.52 7.29 22 11.40

1958

January Y; 6.85 6.62 7.48 13 7.79

February ak 6.76 5.98 iss 13 5.68

March 4g 6.73 6.16 T.22 iH 3.68

April .. 44 6.78 6.24 7.39 12 3.45

May .. ACER 6.99 6.62 7.47 18 7.44

June .. a“ 6.85 5.82 7.69 14 11.90

July 3. 7” 6.88 6.71 7.32 8 2.19

August KE 6.91 5.88 7.18 17 12.81

September by 6.97 6.76 7.38 14 3.18

October a 6.58 5.88 7.28 18 12.35

November - 6.51 5.89 7.33 20 21.65

December Ld 6.89 6.50 7.28 12 4.43

1959

January 45 6.78 5.59 7.49 19 9.78

February As 6.63 5.98 - 7.19 10 3.88

247

Gardens Bulletin, S.

Table II

pH OF RAIN WATER FROM THE UNIVERSITY SITE

Number

pH pH pH of days M aed

(average) (minimum) (maximum) on which ( # ee

rain fell inches)

1958

May 6.56 5.14 7.29 19 5.54

June 5.89 4.95 TAZ 15 11.89

July 5.76 4.85 6.52 7 3.19

August S53 4.85 6.16 15 13.85

September 6.04 5.44 6.69 9 3.00

October 5.65 5.04 6.37 15 11.58

November 5.48 4.58 5.83 16 18.75

December 6.00 FO. 6.33 8 4.70

1959 .

January 5.92 5.56 6.39 11 10.38

February ait 4.59 6.72 9 4.60

Table III

DATA FROM RECORDING RAIN GAUGE IN BOTANIC GARDENS

Date Time of Time of Composite Sample Final Sample

Thunderstorm Rainfall Inches pH Inches pH

25-5-58 0830-0920 0810-1000 1.52 662 0.52 7.20

12-6-58 0720-0755 0630-0730 1.10 582 O10. 639

29-8-58 0615-0755 0505-1015 1.89 630 0.89 6.18

8-10-58 0510-0630 0415-0900 1.13 6.74 0.13 6.82

22-10-58 1040-1150 0930-1130 2.92 5.88 0.92 6.06

1-11-58 0240-0350: 0135-0735 1:82 6.16 082 6.10

1-11-58 we 5 1755-2055 1830-2200 1 1.48 589 048 6.49

1530-1620 1445-1630

16-11-58 “12305-2320 2145-2400 f 13% 7602. 5-087 Gs

24-11-58 0130-0345 .-0100-0335 1:25: . 6:68,..0.25...6000

27-11-58 1730-1820 1630-1800 1.31 7.03 O31 6.28

Composite Sample Final Sample

ag 2 Inches pH Inches pH

6-5-58 1.82 6.93 0.82 6.83

22-6-58 1.95 6.50 0.95 7.10

29-6-58 1.97 6.70 0.97 6.39

17-8-58 1.90 5.88 0.90 6.04

18-8-58 1.08 6.18 0.08 5.76

1-10-58 1.68 6.89 0.68 6.51

14-11-58 1.81 6.18 0.81 6.42

15-11-58 1.24 6.10 0.24 5.81

17-11-58 3.61 6.08 0.61 5.90

18-12-58 132 6.27 0.32 6.34

6-1-59 1.99 6.84 0.99 7.00

12-1-59 1.20 6.72 0.20 7.07

In the latter part of the table, the rainfall refers to a 24 hour

period commencing at 0800 on the day noted in the first column.

248

Vol. XVII. (1958).

(5) In spite of these variations from day to day and from site

to site, the graph of average monthly pH plotted against

time shows some remarkable features (Fig. I).

After the first year’s measurements had been completed, it was

thought that the variation with time was characterised by two

maxima, one in April and the other in December. It could well be

argued that the small maximum in August is due to a chance

fluctuation. This maximum appears, however, in the graphs for

each of the succeeding years. It is also remarkable that the data for

the University site show the same variation although the monthly

pH values are uniformly lower and the incidence of rainfall dif-

ferent. All four graphs show maxima one about April or May, a

second about September and a third about December. It may be

significant the first two maxima occur at the beginning and at the

end respectively of the South-West monsoon: the third maximum

occurs about half way through the North-East monsoon.

The measurements described in this paper were not designed as

an elaborate study: at first, we were interested only in answering a

question which arose from some work on the corrosion of metals:

what is the average pH of Singapore rain water? In the course of

this work a number of related problems suggested themselves,

some of which have been described. The answers we have given are

tentative because the original experiment was designed for one pur-

pose only. If this work could be continued, several modifications

could be suggested, such as the collection of separate samples

during thunderstorms, the observation of thunderstorms from the

site of sampling and not from a distant observation post, and the

examination of samples for nitrite and nitrate.

I am grateful to Professor Gilliland, Mr. Purseglove and Mr.

Burkill for the collection of samples and to Mr. Mather for the

provision of meteorological data.

249

Gardens Bulletin, S.

Figure I

OWeWOwryY

weroutu

S98 -ywew-p em

g34

930

AON

L350

d3s

oNv

NNO

AVW

Udv

YVAN

250

Botrychium daucifolium Wall.

from Cameron Highlands

Maiayan Fern Notes

By B. MOLESWORTH ALLEN

A preliminary note

BOTRYCHIUM DAUCIFOLIUM Wall. ex Rac., Flor. Btz., I. 4. (1898).

The genus Botrychium has not previously been recorded for

Malaya. Holttum however, when discussing the Ophioglossaceae

in “Ferns of Malaya” (1954), page 38, notes: “Botrychium occurs

in both Borneo and north of Malaya, so that it may occur in

mountain forests in Northern Malaya.” Thus it was not surprising

when I found it in the Cameron Highlands area of Pahang in

April 1958.

The plants agree with the description of B. daucifolium given

by van Alderwerelt van Rosenburgh in “Malayan Ferns” (1908)

page 778 (the forma typica of his supplement (1916), page 455).

Most are a little smaller than his measurements, and the segments

of the sterile frond are inclined to be coarsely toothed. I have not

seen Wallich’s description, so do not known whether the presence of

copious pale hairs on very new growth and the almost glabrous

state of the mature fern, is typical. This is not mentioned in any

description I have seen, but Beddome’s illustration in “Ferns of

British India & Ceylon” (1892), page 470, shows a very hairy

stipe and rachis of an adult plant. This is not described in the text.

None of my mature or nearly mature specimens approaches this

state.

The path to Gunong Perdah, for about one mile past the Gunong

Jasah turnoff, follows a more or less flat contour, along the sloping

sides of the hills, which are clad in tall rain forest. This is at about

5,000 feet in altitude. Here nine plants of Botrychium were found

in a small area of about two square metres, where there was a

slight opening in the tall dark forest. Apparently a tree had fallen

sometime previously widening the opening already made by the

path. In the immediate vicinity there were no plants growing be-

tween the tall saplings which were nearly two metres high, and the

scattered plants on the forest floor. The tallest of the latter were

about 45 cm., and were mostly a species of Didymocarpus and

young Cyathea species.

_ The earth was rich in humus being black and sticky, and the

situation was some distance above a stream. The Botrychium plants

were a paler green than the surrounding vegetation and showed up

Gardens Bulletin, S.

clearly. The largest plant measured was nearly 36 cm. tall and had

the remains of a very old fertile spike which projected beyond the

sterile portion of the lamina. The sterile leaves were also past

maturity, but there was one very new and uncurled frond, most

parts of which were covered with transparent spreading hairs. All

the older fronds examined were almost, or quite, glabrous. Amongst

the other plants there were only two fertile spikes, both being very

new, but many of the other fronds still showed where a fertile

spike had been. Two of these can be seen in the accompanying

photograph.

Although the adjacent ground, both above and below the path

was searched, no more plants were found here, but further on to-

wards the Jasah turnoff, eleven more were found. They occurred

sporadically on the pathside for about half a mile, in similar places

to the first lot except that there was seldom as much light on the

forest floor. Most of these had new fertile spikes which were often

nearly mature.

Further field observations are necessary to determine whether

Botrychium dies off in the same way as some of the other Ophio-

glossaceae in Malaya. The presence of bases of old fertile spikes

rather suggests that they do not do so, yet a plant may produce

more than one fertile spike in one season before a die-back. I had

hoped to return in July, but new emergency regulations have

closed this area, so no further notes have been made since the first

collection.

Botrychium may not be uncommon in these forests where the

floor is flat and where damp (not wet) conditions prevail. They

appear to require a certain amount of light so perhaps the normal

tall jungle provides only a temporary habitat after an opening has

been made, and before other plants of more vigorous growth cover

the ground. The path here no doubt, has provided this sort of

habitat, for since the emergency it has remained practically undis-

turbed. Saplings on the path edges have become tall and so have

kept that part of the floor fairly clear. The saplings are now how-

ever, at a height where the undergrowth can and probably will

start covering the ground again. So possibly Botrychium may be

on the decline here. It will be interesting to see the position in a

year’s time, and probably much will depend on whether the paths

have been cleared. Perhaps the original habitat is in the light shade

provided by the smaller trees in the padang areas of higher

altitudes.

A living plant is at Kew and herbarium specimens No. 3930

were sent to the herbaria of Kew and the Singapore Botanic

Gardens.

Zoe

, 7 ee,» tae “ieee

Malayan Fern Notes, II

By BETTY MOLESWORTH ALLEN

MALAYAN HILL STATIONS are always interesting botanically, espe-

cially so where the clearing of jungle is still taking place and there-

fore the plant formation is continually changing. In the past,

however, cleared areas such as roadsides, lawns, golfcourse and

garden edges and forest paths have been kept cut for so many

years that small, light loving plants, both native and introduced

have often become established. Now, because of the present emer-

gency, some of these areas have remained almost undisturbed for

nearly ten years, allowing a taller and different succession of plants

to take over. Many paths through the jungle received little or no

clearing, and in the Cameron Highlands district of Pahang, some

of the gardens of remoter bungalows became neglected. In the

lower part of the now deserted garden of one such place, Walker-

burn Cottage, several interesting ferns have become established

amongst short second-growth and lank grass. These include

Cyathea excavata, Cystopteris tenuisecta, Athyrium amplissium,

Merinthosorus drynarioides, Thelypteris beddomei and T. brunnea.

The fairly new car road to the summit of Batu Brinchang, a

mountain of 6,666 feet in the Cameron Highlands area of Pahang,

enables the botanist to spent more time and energy at its higher

altitudes. Here on clearings caused through roadmaking, colonisers

(seen in 1956) were small, some being rare plants which will no

doubt disappear when larger plants become established. In fact this

has already begun to happen and in two years there has been a

marked change on some of the higher mist swept ridges over

which the road runs. It is a pity that some systematic record could

not have been kept of these early stages of plant succession, for in

the past six years or so, roads have been built up to other summits

and high ridges of Malayan mountains for V.H.F. wireless stations.

In 1956 on Kedah Peak in Kedah, I found that the then un-

finished road up the last part of the very steep and narrow summit,

had already influenced plant growth below. The road was draining

water away from the flatter ground above the resthouse where pre-

viously large patches of Sphagnum were common in the wet ground.

Most of these patches had disappeared together with the little local

Schizaea malaccana Bak., which grew amongst the Sphagnum, and

the ground was very dry and almost devoid of plant growth.

253

Gardens Bulletin, S.

Again the emergency has contributed directly to a changed vege-

tation in places in the lowlands. Previously fertile ground such as

riverine areas and terrain near the bases of limestone hills (espe-

cially in the ipoh district of Perak) have for many years been

under almost continual cultivation. During the emergency these

areas were cleared of squatters and put under curfew. Thus neg-

lected for nearly ten years, the rich soil together with the moist

hot climate induced a quick succession of belukar (a local name

for secondary scrub formation). In many places this resulted in a

growth tall and thick enough for terrestial ferns, which would not

tolerate full sun, to appear. For instance, Phymatodes longissima

(Bl.) J. Sm., apparently not a common fern in Malaya, is now

abundant in belukar in these places around the Ipoh district. With

better political conditions approaching however, curfews have

been lifted in many of these places and people are going back on

to the land. Since June 1958 many areas have been cleared again

by cutting and burning, and already much is under vegetable culti-

vation. Thus possibly 1959 will see the end of this interesting

decade of undisturbed belukar near the towns.

The order of the following annotated list and the nomenclature,

follows those used by R. E. Holttum in his “Ferns of Malaya”,

1954, and the page number of the species discussed, is given in

brackets.

All collecting numbers quoted are of my personal series, the

specimens being either in my private collection or in the herbarium

of the Botanic Gardens, Singapore. Duplicates have been distri-

buted to other herbaria including Kew and the British Museum

(Natural History).

The list contains results of field observations, including some

extensions of known distributions and notes on two species of

ferns not previously recorded from Malaya.

Trichomanes motleyi v.d.Bosch, (Holttum p. 92).

Growing on the vertical side of a large boulder in a stream bed

in tall forest on the lower slopes of Penang Hill, at Batu Ferringhi.

April 1956. Previously known from Singapore, Pahang and

Selangor. Fertile fronds were common but the specimens I found

in Kepong, Selangor which were in a similar habitat, had, in

August, very few fertile fronds. I have since found it on a tree

trunk about two feet from the ground, in a stream in tall forest at

Ginting Simpah, Selangor. Penang specimen no. 2795.

254

Vol. XVII. (1958).

Cyathea excavata Holttum, (Holttum p. 121).

So far this fern is known only from Cameron Highlands in

Pahang. Holttum states (p. 122)* that only old sori have been

seen. In August 1956 I found fertile fronds with immature and

newly mature sori and the following description is taken from

several large plants growing either in open or fairly open places in

Cameron Highlands.

The largest middle pinnae measured were up to 63 cm. long

(commonly 60) by 20 cm. wide, usually short stalked. It will be

seen that these measurements are somewhat larger than those given

by Holttum. Unless the frond is past maturity the lobes are much

closer together than their own width when living and the edges are

nearly always conspicuously toothed, sometimes lobed, being en-

tire only towards the apex. Veins were forked up to four times,

two to three being usual. The smooth stipes were almost black at

the bases, but were green higher up. I found practically no scales

on the stipe, even when young; they were present only on the very

new and curled fronds. These scales were not thin, but fairly

thick and large (often over 1-5 cm. long). Sori were 1 to 4 (com-

monly 1 to 2) at the bases of the pinnule lobes near the costae.

Mature sori coalesced when dry, obscuring the lower part of the

costules. Fronds frequently produce very few sori; a pinnule being

fertile for only half its length, and the frond itself being sterile

towards its apex. The very thin transparent indusium completely

covers the young sporangium, but appears to break down at a very

early stage of maturity. I have found that in pressed specimens, if

the sporangia have burst during the process of drying, they seldom

show any traces of indusia, yet if they have reached maturity nor-

mally before drying, some vestiges can be seen as described by

Holttum. One and sometimes two, dark brown thin appendages

similar to paraphyses were noticed attached to the base of a sorus,

curving round it to about half its height. I saw these only on large

fronds when the sori were immature. Paraphyses amongst the

sporangia were not seen.

In the field this fern is most distinct from other species of

Cyathea. The fronds when living are pale green, soft and inclined

to hang laxly. A distinctive feature, not found to the same extent

in any other Malayan Cyathea, is that the old dead fronds remain

on the trunk, hanging down more or less concealing it, in much the

same way as in some Dicksonia species. Another field character,

but I am not sure how constant this is, is that all the plants seen

had more than one trunk. Two was usual, but occasionally there

* From “Ferns of Malaya’ unless otherwise stated.

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were three, and on these trunks, at their bases, and up to about

60 cm., young adventitious plants were growing (I have seen the

same in C. recommutata Copel. and in C. polypoda Bak.). One

plant had become quite large and had fertile fronds.

Three mature plants of C. excavata were found fairly close to-

gether by a stream above Sultan Abu Bakar Road near Brinchang

village in 1956, at about 5,000 feet altitude. In 1958 several more

were found in this area and others at Taman Sedia, which is about

4,500 feet. Since then more plants have been seen in open or semi-

open places, but only in the Cameron Highlands area. It appears

to prefer streamsides, well above the normal water level, on forest

edges. In Taman Sedia however, they were growing in open grassy

places where there was only an occasional small tree and several

tall specimens of Cyathea contaminans (Hk.) Copel. C. excavata

appear to be on the increase where streamsides have been cleared

or partially cleared, and some plants have grown a great deal since

first seen here. Trunks 150 cm. high were not uncommon, and

some were higher. The plants at Taman Sedia at least, are un-

doubtedly in an artificial habitat, for Holttum originally found C.

excavata in shady forest by small streams, so it will be interesting

to watch their development if they are left undisturbed.

Colysis acuminata (Baker) Holtitum, (Holttum p. 162).

Sungei Buloh, Selangor; on the base of a large tree and on the

ground beside a stream, well below the flood level; tall lowland

forest; rare. March 1952, specimen no. 1875; June 1953, no.

2587.

Previous records are from Perak and Pahang. Holttum says that

it occurs in the same situations as C. macrophylla (Bl.) Presl,

which I did find near the Selangor specimens, but I have only

found C. acuminata in association with streams and growing where

it must frequently be inundated. In Perak it grows on rocks in the

beds of small shallow streams. C. macrophylla on the other hand

is more common in flat places in moist lowland forest, but not

necessarily near streams.

Loxogramme scolopendrina (Bory) Presl, (Holttum p. 168).

Batu Caves, Selangor; January 1956; specimen no. 2686. Near

the summit of this limestone hill, probably about 900 feet altitude.

It was growing in a pocket of earth on steep ground and was rare.

Holttum lists this species from Pahang, Perak and Kedah.

Aglaomorpha heraclea (Kze.) Copeland, (Holttum p. 185).

Cameron Highlands, Pahang; altitude 5,000 feet. Although this

fern record is only 500 feet higher than the highest altitude given

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Vol. XVII. (1958).

in “Ferns of Malaya’, it has been included, as all the plants seen

in the forest were small and infertile, indicating that they may be

of recent origin. In June 1958 plants were seen on trees in tall

forest on the slopes of Gunong Jasar and G. Purdah. They were

not uncommon here. In the garden of Walkerburn Cottage at

about the same altitude there is a large clump of Aglaomorpha

heraclea perching on an Erythrina sp., and this was fertile in June.

Merinthosorus drynarioides (Hook.) Copeland, (Holttum p. 186).

The following are some field observations on this fern which

has been recorded from several localities in Malaya.

In February 1948 I first saw a plant of Merinthosorus on a rock

in a fairly open place near Robinson’s Falls at Cameron Highlands,

Pahang, at about 4,500 feet in altitude. It was small and not fer-

tile, but the rhizome scales distinguished it from Aglaomorpha

heraclea which it resembles in leaf character. When I visited this

area again in 1956 the fern was no longer there, but I subsequently

found another plant on a concrete wall, in almost full sun in the

deserted garden of Walkerburn Cottage, near Brinchang village.

Here the altitude is about 5,000 feet. The plant was young and

infertile then, and so far (1959) has not produced sori although it

has increased in size.

I have not seen Merinthosorus growing elsewhere at this altitude

in the Cameron Highlands area, but it is common as an epiphyte

on tall trees from Ringlet (circa 3,500 feet) downwards, until the

jungle gives way to rubber estates, and they can be seen from the

road. I have seen this fern in the following localities in Malaya: —

PERAK: Maxwell’s Hill, on rocks in open places near the bunga-

lows at 3,200 feet in altitude; on canopy trees in tall forest at 2,400

feet. On road to Cameron Highlands (but still in Perak), on a

vertical rock bank beside road at c.3,000 feet (mostly young

plants); high up on canopy trees in tall forest at c.1,700 to 3,500

feet. PAHANG: Cameron Highlands district on rock and wall in

open places; Fraser’s Hill, common on stone walls by bungalows*,

about 4,000 feet altitude; on large boulders in a meadow at Jeriau

Farm, 3,400 feet. SELANGOR: In tall forest common from about

1,500 to 3,000 feet, below and above the Semangkok Pass (The

Gap); Ginting Simpah, on Rain Trees (Enterolobium saman)

which have been planted on the roadside at the 17th milestone, at

about 1,000 feet altitude; on tall forest trees in the forest near this

altitude.

*In June 1958 most of the ferns from the stone walls had been removed.

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Gardens Bulletin, S.

I have not found Merinthosorus on Penang Hill (from where it

was originally collected in Malaya) although I have searched for it

during three months when I was living on the hill.

Merinthosorus in common with other acrostichoid ferns in

Malaya, only periodically produce fertile fronds. In this genus

however, it is almost certain that they are developed twice a year,

(not just once a year, as for instance is the case with Elaphoglos-

sum spp.) and possibly more frequently on rock walls. The latter

may be induced by the frequent cutting back that they receive.

The following, although not yet complete, show the months during

which fertile fronds have been observed. Fertile on rocks: —Max-

well’s Hill, February and November; Fraser’s Hill, April, August

and December. Fertile on trees:—Maxwell’s Hill, November; road

to Cameron Highlands, June and August; above and below the

Semangkok Pass, June, December; Ginting Simpah, June. Only

Fraser’s Hill has been observed during December.

In 1957 I was able to watch the development of some Merintho-

sorus plants which were growing on the previously mentioned Rain

Trees at Ginting Simpah. Although these plants must be fairly

recently established, they were mature, and had the advantage of

being easily and frequently seen. During February and March

there was no noticeable growth, only sterile fronds being present,

but during April new (sterile) fronds appeared. A few weeks later

the bases of each of these new fronds became brown and dead-

looking for less than one-quarter of their length, the rest remaining

green. This gave a remarkable resemblance to the scale or nest-

leaves of a Drynaria, and the plant from a distance would surely

be taken for D. sparsisora (Desv.) Moore. It was the beginning

of May when the bases turned brown, and a month later the fertile

portion appeared on these new fronds. Since then I have observed

similar growth on other Merinthosorus plants on trees by the

Cameron Highlands road. Although this scarious condition of the

frond has been noted previously, I am not aware that its develop-

ment has been described. This, and the fact that it is epiphytic on

uppermost branches and on crowns of very tall canopy trees in

dense forest, is probably the reason that it has escaped notice in

these places for so long. This habitat I believe is almost certainly

the natural one in Malaya.

In the north of Malaya at least, Merinthosorus is probably

quite common high up in the large trees of tall midmountain forest

where the humidity is high. Even so, field glasses are often neces-

sary to detect the fern. On rocks in open places and at hill stations

I think it is most probably of recent origin, appearing after the

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Vol. XVII. (1958).

jungle was cleared and that it will increase in these areas, for it

appears to adapt itself easily to the sides of large rocks and walls

where there is good drainage.

Polypodium prainii (Bedd.) C. Chr., (Holttum p. 204).

Holttum describes this only as an epiphytic fern. It is still very

common at Cameron Highlands in Pahang, but I have found, with

only a few exceptions, that it is terrestial there. It grows where

there is abundant moss and liverwort both on rocks and wet rocky

ground, above streams where there is often much undergrowth and

little light. It may have become a terrestial fern by adoption in

areas where the tall trees have been removed, for branches are

usually left in the forest, and it is in such places that it has been

seen most commonly. It occurs quite frequently however, on the

ground on the lower slopes of Brinchang and Jasar mountains

where the tall forest appears to be little disturbed.

Polypodium prainii appears to have a seasonal die-back. In July

only very old fronds were seen, and none were found in the same

area in August, but when again visited in late September, new

fronds were common. I have never seen it at Fraser’s Hill.

Christiopteris tricuspis (Hook.) Christ. (Holttum p. 211).

Slopes of Gunong Terbakar in the Cameron Highlands district

of Pahang, at 3,800 feet, August 1956.

Plants of this fern were growing out of the base of a large plant

of Aglaomorpha heraclea, which was on a tall tree in a very ex-

posed position. The Christiopteris had fertile fronds. It was not

seen elsewhere here, but should be looked for wherever large

clumps of Aglaomorpha grow, for it has only been found in asso-

ciation with this fern.

The accompanying photograph shows a small fourth lobe on a

fertile frond of Christiopteris from Fraser’s Hill. The frond was

taken from a plant which is more or less established on the ground

at the base of a fallen Aglaomorpha on the roadside near Pahang

bungalow. This development has not been previously observed in

Malaya, and all other fronds from this plant had the normal three

lobes.

Since writing the above, more plants of Christiopteris have been

found at Kuala Terla at Cameron Highlands, at 3,800 feet altitude.

They were young and infertile, growing from an Aglaomorpha on

a tree over the Terla River.

Grammitis crispatula Holtt., (Holttum p. 218).

Maxwell’s Hill, Perak, February 1952. Herbarium specimen

no. 1828. Although originally found in this locality, the fern does

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not appear to have been seen in recent years. My field notes state

that there were only a few plants and these were growing on a

very large moss-covered rock in tall jungle, on a ridge just below

3,000 feet, but not anywhere near a stream. The fronds were dull

and dark green with conspicuously crisped edges. My identification

was kindly confirmed by Dr. E. B. Copeland to whom I sent a

specimen with viable spores.

Ctenopteris celebica (Bl.) Copeland, (Holttum p. 233).

Robinson Falls, Cameron Highlands, Pahang, at about 4,500

feet. August 1956. This fern was not seen elsewhere and was found

on a sloping mossy tree trunk in a very damp place overhanging

the river. It was growing amongst Hymenophyllum badium Hk. &

Grev. and Grammitis hirtella (Bl.) Tuyama. Fronds when living

were pale yellowish and dull, with conspicuous red hairs on the

stipes. Some of the fronds had new sori. The tree has since been

destroyed by blasting operations for the new hydro-electric dam,

but it surely must occur elsewhere here. Specimen no. 2895.

Thelypteris brunnea (Wall.) Ching, (Holttum p. 240).

This fern has obviously increased greatly in recent years for it is

now very common at Cameron Highlands, growing in grassy places

in full sun, by streamsides in shade, and on banks by roads. It

sometimes becomes very large in the first mentioned habitat, larg-

est fronds measured being nearly 200 cm. long including stipe, 160

cm. being about average.* The stipe is always shining and rather red.

I have however, examined many plants from Camerons and from

Fraser’s Hill and have found in every case that the stock is either

erect or suberect, but never long-creeping which is characteristic

of this species. I have collected Thelypteris brunnea on Mt. Kina-

balu in British North Borneo and although the fronds appear simi-

lar, the rhizome is entirely different, being creeping, with fronds

widely spaced and not bunched as they are in Malayan specimens.

I have found what I consider to be normal T. oppositipinna

(v.A.v.R.) Ching at both Cameron Highlands and Fraser’s Hill,

but never commonly. In the field it looks very different from T.

brunnea, yet there is very little real difference, and I have found

them growing together. The late Mr. Alston of the British Museum,

to whom I sent specimens of T. brunnea said they appeared to be

that species except for the rhizome, so presumed they belonged

to JT. oppositipinna. Holttum notes that the rhizomes of the

Malayan specimens were missing.

* From the overgrown garden at Walkerburn Cottage.

260

Vol. XVII. (1958).

I believe that the true species of 7. brunnea as interpreted by

Ching (Bull. Fan Mem. Inst. Bot. 6: 1936) has not yet been

found in Malaya and perhaps this species should bear another

name, if really different from T. oppositipinna.

Thelypteris beddomei (Bak.) Ching, (Holttum 240).

Cameron Highlands, Pahang, August 1956, c.5,000 feet. no.

2876. Previously known only from Maxwell’s Hill where I found

it in 1949, and where it is still common. In the Camerons it is

fairly widespread and abundant where it does occur. It was grow-

ing above streams in open places and in semi-shade on the edges

of light forest. In some places it formed stands so thick that there

was little other growth. Old rotting fronds formed a deep loose

layer (sometimes to 45 cm.) beneath the living fronds so that the

latter had greatly elongated stipes (to 60 cm. long). The longest

lamina measured was 40 cm. The fern seemed well established in

this area just below Walkerburn Cottage, but when I returned in

June 1958, grass and the shrub Melastoma muticum Ridl. had

become tall and this fern had decreased quite considerably. It had

become abundant however, on the more recently neglected lawn

near the house, but the fronds were short.

In July 1958 I found T. beddomei growing between the tea

bushes at Sungei Palas Tea Estate at c.5,500 feet. Sometimes it

was so abundant that it covered the ground to the exclusion of

other plants. It was also common on and above the roadsides

here, and wherever the ground was sloping. As at Maxwell’s Hill,

the ferns were in full sun and the fronds were quite short, agreeing

with the measurements given by: Holttum.

I examined many fronds and found that, as Dr. Holttum notes,

indusia were always present, and were evident even when the

specimens were dry.

Ampelopteris prolifera (Retz.) Copel., (Holttum, 299).

Collected once in Malaya during last century by King’s collector,

on the Kinta River in Perak. In April 1956 I found one patch of

this fern on the Kedah-Province Wellesley border where the old

Bambong Lima bridge spans the Muda River. It was growing just

below the road in alluvial soil, on flat ground which is frequently

flooded by the river. It was scrambling over, and covering small

bushes with quite a dense growth. Buds were produced five to six

times along a lamina; these in turn became proliferous as they

matured, so tracing the overall length was difficult. Stipes and

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fronds were dark green when living. I found only two fronds bear-

ing fertile pinnae and these were on tertiary fronds. Athyrium

esculentum (Retz.) Copeland was very common nearby. In

August 1958 I found Ampelopteris in Perak. It was alongside the

Kinta River near Batu Gajah, possibly near the place of the ori-

ginal collection. For at least 150 yards along the river bank it

was common to abundant, scrambling up small trees. Although the

stipes and primary rachises were living, nearly all the pinnae,

together with the secondary and tertiary fronds, were dead. No

new nor fertile fronds were found. Here the river lies fifteen to

twenty feet below the bank which is flat and composed of heavy

soil, which was then very dry and hard. Apart from one small

patch on the river bank, Ampelopteris was confined to a narrow

band starting about ten feet from the bank edge and running

parallel. This is where the trees occured, and beyond was cropped

grass which I believe is frequently burnt back, and no ferns were

found there. Other ferns growing amongst the Ampelopteris were

Athyrium esculentum and Cyclosorus aridus (Don) Ching, both

of which it slightly resembles in the field. In the following Decem-

ber fertile plants were found here, with mature sori on secondary

fronds and very new sori on tertiary fronds.

The rootstock of Ampelopteris is very surface-rooted, being

easily pulled out; the fronds when dry could be easily burnt. So

perhaps as river edges are so often cultivated, this fern may have

been eradicated from many areas. Otherwise it is difficult to

understand why it is not so common as is Athyrium esculentum

in these places. It is apparently seasonal as it is in other countries,

dying back (in Perak) during the dry months between June and

August. I have seen this fern with mature sori in December on

riverbanks in west Siam. I have grown some plants by the Pinji

River in Ipoh to observe their development. Herbarium specimens:

2794, 4090.

Microlepia puberula v.A.v.R., (Holttum p. 312).

Kepong and Sungei Buloh, Selangor. Collections made between

1951 and 1955. Herbarium specimens: 1382, 1482, 2333, 2393,

2541; from Ginting Simpah, Selangor in 1957, no. 3473.

Lindsaya cultrata (Willd.) Sw., (Holttum 328).

Near the 14th milestone, Cameron Highlands Road, Perak, Sep-

tember 1958, 1,000 feet; no. 4017. Locally frequent on mossy

rocks in bed of large stream in jungle. Probably from plants swept

down river from the mountains.

262.

Vol. XVII. (1958).

Leucostegia pailida (Mett.) Copel., (Holttum 353).

Although this fern has been seen recently by Sinclair, its habitat

is of interest because of its doubtful native state. On a path along-

side the water pipeline at about 2,400 feet on Maxwell’s Hill in

Perak, it is locally abundant, growing on the tops of large boulders.

Here the fronds grow to at least the maximum length as given by

Holttum; new fronds, which were produced in quantity in Novem-

ber, were bright pink. This immediate area, although clothed in tall

forest has been cleared of a great deal of undergrowth, so that the

boulders are only lightly shaded. Well away from the path the fern

becomes less common, not occuring on the many similar boulders

in the untouched forest where the light is poor. It grows however,

on mossy tree trunks on, or just above the streamsides, and is

quite widespread on this side of the hill at about this altitude. The

fronds, although there were mature and old ones, were very much

smaller than those from the first mentioned habitat.

I have looked for it several times in 1956, in similar places on

Mt. Matang in Sarawak, from where it was formerly collected, but

without success.

Blechnum orientale L., (Holttum p. 446).

A bi-pinnate to tri-pinnatifid form of the above fern was first

seen on the forest edge by Girdle Road at Fraser’s Hill at about

4,200 feet. Another plant was seen later on the roadside to the

farm, but was very much smaller. In December 1956 I found this

form was not uncommon locally near Batu Lintang Road in Ku-

ching, Sarawak at sea level.

Only the Girdle Road plant showed transition fronds. There

were very old fronds which were quite normal, others had deeply

lobed pinnae and the newest (pink) fronds were bi-pinnate becom-

ing almost tri-pinnate at their bases. When this plant was revisited

sometime later, all the fronds were like the last mentioned ones,

some measuring 145 cm. long. Unfortunately this plant was des-

troyed during roadside clearing operations. All plants examined

had fertile fronds. It would make an attractive fern in cultivation.

Specimen no. 2202.

Blechnum fraseri (A. Cunn.) Luerssen var. Philippinense Christ,

Bull. Herb. Boiss. 6.149; van Alderwerelt van Rosenburgh,

Malayan Ferns, 382. 1908.

Gunong Batu Brinchang, Cameron Highlands district of Pahang;

6,500 feet altitude, August 1956. Specimen no. 2920; February

1959, no. 4145. A living plant was sent to Kew in 1956.

Not previously collected in Malaya, but this variety is known from

the Philippine Is., Borneo, Sumatra and New Guinea. Blechnum

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fraseri belongs to the sub-genus Lomaria to which, in Malaya,

only the mountain B. vestitum (Bl.) Kuhn, belongs.

Apart from the adult plants being larger, the Malayan specimens

agree quite well with the descriptions I have seen of the variety,

and with plants I subsequently saw on Mt. Kinabalu in British

North Borneo. The following description is taken from Malayan

plants from a series collected from one area on Brinchang.

Mature plants frequently (?) stoloniferous. Rootstock elongated

into an erect caudex, the tallest measured being 235 cm. high, with

a diameter of 5 cm. at approximately half way up the caudex, but

more commonly 200 cm. by 6 cm., plants becoming fertile when

caudex reached a height of 30 cm. Base of caudex thickened by a

mass of old roots to 16 cm. or more through. Caudex or trunk

rough, dark brown almost black, the greater part being scaly and

rough from the remains of old stipe bases, but from the base ex-

tending upwards to c.90 cm. covered with close thin adventitious

roots. Through these protrude scale-covered adventitious buds,

2—3 cm. long, which, near the trunk base had elongated into prop-

roots 10 cm. and more long by about 1 cm. thick. Stolons 1 to 2

(or more?) from a parent plant, horizontal for at least 20 cm.

before becoming erect and forming another plant which later be-

comes independent. Some of the thin adventitious roots from the

upper side of the stolons elc 1gate and growing downwards, anchor

the plant. Caudex of new stoloniferous plant densely clothed with

scales. Scales on caudex, stipe bases, and buds more or less similar,

dark brown, thick with narrow elongated non-clathrate cells;

length of scale to about 15 mm. and 2 mm. wide at base with subu-

late apex; edges entire with pale thin margins which are deciduous.

Fronds bipinnatifid, whorled near the apex of the caudex and

rather tufted at the crown. Fronds from shaded plants with thin

elongated stipes, and laminae large, lax and dark green above,

thin in texture; fronds in semi-shade or open places, nearly erect,

shorter, upper surface yellowish-green, firm in texture. In both

forms, upperside, shiny and glabrous; undersurface, dull, pale,

sometimes almost glaucous and glabrous, except for sparse pale

thin scales on rachises, costae and costules. Fronds dry olive brown

above and greenish below. Stipe bases thickly clothed with dark

spreading scales. Sterile fronds varying greatly in size according to

habitat, up to 100 cm. long including stipe, by 35 cm. wide in

shade specimens, but more commonly 56 cm. by 19 cm. Fronds

widest about the middle, tapered to the apex. Base narrowed.

Rachis winged throughout connecting pinnae, wing with triangular

lobes, contiguous. Middle pinnae on large fronds to 17 cm. long

264

Vol. XVII. (1958).

by 2:5 cm. wide but more commonly 10 cm. by 1:5 cm., deeply

Jobed with a long tapered apex which is entire for a length of 1 cm.

or more at its tip. Upper pinnae ascending, gradually reduced in

size to a lobed acuminate apex. Lower pinnae reflexed and reduced

in size to about 7 cm. long on large fronds to 4 cm. on small fronds.

Rachis wing continuing down the stipe gradually disappearing,

leaving the triangular lobes as small auricles, until ceasing near the

stipe base. Pinnae lobes entire, or toothed in shaded fronds, fal-

cate, blunt or occasionally with very short pointed apices; lobes

2—4 mm. wide, cut to a costule wing 1-5 mm. wide, each lobe being

separated by a hyaline membrane. Lobes becoming less deeply cut

and decreasing in size towards the apex. Fertile fronds similar in

shape to the sterile, but usually smaller and slightly contracted, to

60 cm. long by 20 cm. wide, with entire lobes gradually reduced in

size for threequarters the length of a pinna, then rather suddenly

contracted forming a long thin entire apical lobe, which is about

25 mm. long by 3 mm. wide. Sporangia covers the lower surface of

the lobes, indusia are thin and light brown, just visible at maturity.

Spores, anisopolar, pale, slightly verrucose, perispore folded.

Veins translucent in thin fronds, but mostly obscured in those of

firm texture. This description is taken from the former:—Two

veins at each side of a costule base, springing from the costa,

sometimes forked, ending within the lobe margin with a small

hydathode. Veins springing from costule, to about 7 pairs, forked

up to four times, more commonly three times, never reaching the

margin. Forks of the lowest pairs occasionally anastomosing.

Young, non-stoloniferous plants have short erect stocks, simply

pinnate fronds with the bases of the bluntly toothed pinnae decur-

rent on the rachis.

On the left hand side below the road to the wireless station on

Batu Brinchang, there is an almost vertical rock face about 150

feet from the mountain’s summit. Here a long time ago there has

been a landslide exposing the rock, and the earth from this slip has

formed a shelf some 50 feet below the road. It was here in 1956,

amongst mixed second growth of about 3 metres high, that the

mature plants of Blechnum fraseri were found. It was then impos-

sible to move through this vegetation, so an accurate count of this

fern was not made, but twenty plants were noted. The largest, with

trunks up to 200 cm. high, were concealed by a Melastomaceous

shrub, by plants of Cyathea latebrosa (Wall.) Copel. var. indu-

siata Holtt., and by Polygonum chinense Linn. A few small in-

fertile plants were growing in peaty soil on the road edge, and were

only about 15 cm. high. Nearby on the cliff edge, in loose soil, were

some slightly larger but still infertile plants; these were trunkless.

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The two latter places were newly established, having been made

during the road construction. On the cliff edge there were few

other plants apart from a sedge and a dwarf Thelypteris viscosa

(J. Sm.) Ching. In April 1958 all except one of the very small

roadside plants of Blechnum fraseri had disappeared; the others on

the cliff edge had grown considerably, but had not yet formed a

trunk, nor were they fertile. The latter were almost concealed by

the sedge, which had grown enormously but not yet flowered. In

another area, in an open place a quarter of a mile away, and about

50 feet from the summit, there is another plant of this fern. It was

of medium size, with a small trunk but was not fertile. This year

(1959) I again descended to the earth shelf and found that the

cover plants had grown so much it was now possible to crawl

under them. Blechnum fraseri had certainly increased a great deal

and within an area of approximately 35 by 25 metres, I counted

137 plants of this fern. More than 30 had definite trunks, some of

these being more than 150 cm. high. The tallest specimens were in

amongst the vegetation on the shelf, and were nearly all bearing

fertile fronds, whereas the smaller and more sturdy specimens,

which were fertile only when a trunk was developed, grew where

there was only other sparse growth. In small pockets of earth on

the rock face and in other quite open places, very young plants

were exceedingly common.

The stoloniferous habit was found only amongst those speci-

mens growing in full shade, where none of the young normally pro-

duced sporophytic plants were seen. Unfortunately this place is

being used as a repository for old tins and boxes discarded from

the wireless station above, so examination for stolons was difficult,

and it has not been ascertained how common the habit is, in this

variety of Blechnum fraseri. A new plant arising from a stolon

which was measured, had a trunk 17 cm. high with a diameter of

1-5 cm., but had not produced any fronds, but already adventiti-

ous roots were strengthening its base.

In New Zealand Blechnum fraseri is a common fern of the

Kauri (Agathis australis A. Cunn.) and other dry forests in the

north, forming colonies but the caudex seldom exceeds 75 cm. in

height.

Apart from the dendroid habit which makes this fern look like

a diminutive Cyathea, it is an interesting addition to the Malayan

fern list. The fronds show an intermediate state between simple

and double pinnation, and the production of stolons, although a

common feature in the type in New Zealand, may not have been

recorded previously for the variety. The occasional anastomosis of

266

Cyathea excavata Holtt.; part of a pinna

Cyathea excavata Holtt.; stipe base showing excavations

QQ \

is (Hook.) Christ.

s tricuspi

Christiopter

Us “Gg if

7 sige

if fe 4

< x4 ‘ ©: RE: “> wn }

» wis

Blechnum orientale L. Bipinnate form; one frond

var. philippinense Christ.

Blechnum fraseri (A. Cunn.) Luers.

var. philippinense Christ.

Sterile and fertile fronds

Vol. XVII. (1958).

the veins is interesting although this may be more accidental, than

indicative of development towards those Blechnaceae with regular

anastomosis, for it seems to appear only in the large shade fronds.

The area where this Blechnum was found is fairly newly estab-

lished as the growth of this fern and the other vegetation suggests,

so perhaps there is another older established habitat here. In

Borneo it grows in fairly open but short mossy forest, a similar

kind being found on some of our higher mountains. On Batu

Brinchang, however, a search for a different habitat of this fern

is now practically impossible around this altitude. Below the road,

every accessible place is being used for depositing rubbish. It is a

great pity that such an interesting area is being ruined, as well as

being rendered unsightly.

Elaphoglossum melanostichum (Bl.) Moore, (Holttum p. 455).

Maxwell’s Hill, Perak; 24.1.1956, at 3,200 feet, no. 2711 was

sent to Kew where Dr. Holttum kindly checked the specimen.

Previously known from Penang Hill. This fern occurs locally

on Maxwell’s Hill between 3—3,200 feet, on trunks of tall trees,

usually on the forest edges in semi-shade. There were several

plants on a tree about 60 cm. from the ground and in November

1955, nearly all of these had new fertile fronds.

Elaphoglossum decurrens (Desv.) Moore, (Holttum p. 458).

Summit of Gunong Brinchang, Pahang; 31.7.1956, 6,666 feet,

no. 2911 sent to the British Museum and determined by the late

Mr. Alston.

In Malaya this has been collected twice previously in the low-

lands, but has however, been collected on Mt. Kinabalu in Borneo

at “5,000 feet and more” (see Holttum).

Dryopteris subarborea (Bak.) C.Chr., (Holttum p. 491).

At Cameron Highlands this fern is terrestial (see Holttum 492),

but at Fraser’s Hill I have never found it so, but always as an

epiphyte in tall primitive forest and usually hanging down from

large clumps of Asplenium nidus Linn., apparently as Kunstler

found it. At Maxwell’s Hill it grows on mossy logs and sloping

tree trunks in tall forest, but also on boulders on the exposed

ridge near The Box, and here has small fronds.

Dryopteris sparsa (Don) O. Ktze., (Holttum p. 492).

Near Taman Sedia, Cameron Highlands, Pahang, at about 4,700

feet. In 1956 this fern was Jocally common on mossy rocks in the

river and on moist earth banks in shady places. Plants were fertile.

267

Gardens Bulletin, S.

Tectaria melanocaulon (BI.) Copel. Ph. Journ. Sc. 2C, Bot., 6:

416. 1907; van Alderwerelt van Rosenburgh, Malayan Ferns

245. 1908 (Aspidium); Backer & Posthumus, Varenflora voor

Java, 74.1939 (Tectaria melanocaula Copel. ).

In tall forest near road to Cameron Highlands in Perak, at 26th

milestone; 3,200 feet, June 1958, specimen no. 3985.

Not previously recorded for Malaya. Plants were found in one

place but were very common here. They were on steeply sloping

ground near a large rock face. Only one plant was fertile; it is hoped

that later fuller field notes can be made.

The shining black stipes and rachises and very pale green fronds,

distinguish this fern from any other Malayan Tectaria. Distribu-

tion: China, Malaysia.

Since writing the above, Dr. Holttum, who kindly compared my

material at Kew, writes that the correct name should be Tectaria

melanocaulis (Bl.) Copel.

Pieocnemia conjugata (Bl.) Presl. (Holttum, p. 534).

This fern has been collected twice in Malaya, near Malacca and

at Baling in Kedah, the latter place suggesting that it may be a

limestone fern. In April 1956 I went to Baling hoping to find it on

the limestone hill of that name, but was unsuccessful. About four

miles on the road to Kroh, however, I found it growing in a small

quantity by the roadside, but not near any limestone. Twelve

plants were seen which were nearly all young, but some had

fronds to 170 cm. long, with stipes over 100 cm. long. There was

only one plant with fertile fronds, and these showed the indusia

clearly. Fronds were pale yellowish-green, but I have seen the

same colour in fronds of P. hemiteliiformis (Racib.) Holtt. (nor-

mally much darker) when growing in sunny places at Fraser’s

Hill. In this area the belukar is frequently cut back for emergency

reasons and possibly this has encouraged the growth of P. conju-

gata here. Apparently it requires only light shade, and was not

being smothered by taller growth. Specimen no. 2793.

Cystopteris tenuisecta (Bl.) Mett., (Holttum p. 540).

This fern is worthy of note, for when it was first found in Malaya

by Holttum in Cameron Highlands in 1936, it was apparently con-

fined to a small area on a newly cut earth bank. When I saw it 11

years later, it was growing on the forest floor in the same area and

was still rare, but the fronds were larger than those found by

Holttum. In August 1956 I was unable to go back to the original

place, but found that it was a common fern at about 5,000 feet

268

Vol. XVII. (1958).

altitude. The favourite habitat seems to be on loose earth overhang-

ing small streams in dark places, but here it was seldom fertile and

the fronds were small. On a flat area just above normal water level

above the stream, I found large plants which were fertile. The

largest frond measured was 130 cm. long by 51 cm. wide, but

were more common at about 80 by 30 cm. Fertile fronds were

either very new or just mature, and no old fronds were seen. Since

then I have found it at higher altitudes, up to 6,000 feet on Batu

Brinchang. It appears now to be one of the common streamside

ferns at 5—6,000 feet at Cameron Highlands.

Athyrium japonicum (Thbg.) Copeland, (Holttum p. 551).

Cameron Highlands, Pahang; July 1956; specimen no. 2904.

Previously collected from Maxwell’s Hill in Perak and from

Fraser’s Hill in Pahang, so links the two localities. In 1956 there

were several small plants of this fern at Robinson’s Falls, growing

in crevices on the large rocks in the river. Together with it were

Lindsaya cultrata and Dryopteris sparsa, all of which must often

be submerged. Athyrium japonicum had only one fertile frond. In

1958 I saw it in several places by and above streams in the vicinity

of Kemunting and Sultan Abu Bakar roads, at about 5,000 feet in

altitude. Large patches were on steep grassy banks just above the

roadside near Sungei Palas Estate bungalow. Here it is well away

from water, and is fairly exposed, and the fronds are uniformly

small (about 18 cm. long).

Athyrium curtisii Holttum, (Holtt. p. 560).

Ampang, Selangor; June 1957; specimen no. 3436.

Athyrium velutinum (Holttum) Holttum, (Holtt. p. 567).

Cameron Highlands, Pahang, in the forest behind Walkerburn

Cottage. Specimens no. 2936, 3937, 4022.

Only a few collections have been made of this fern. It appears

to be not uncommon locally where the forest is dark, the ground

sloping yet wet and rich in humus, but usually some distance above

streams. I have found it at about the same elevations as are given

by Holttum. In most plants that I have seen, the stock was not

short, but usually quite stout and erect and sometimes was built

up to 9 cm. in height, being supported by firm black roots. Living

fronds are very dark green except when very young, then they are

extremely pale. Fertile fronds always appear to be longer than the

sterile and more deeply cut. They also have free pinnules for nearly

half the length of each pinna. Pinnules are commonly 15 mm. long

by 5 mm. wide (occasionally to 17 mm. long), but the sterile

269

Gardens Bulletin, S.

fronds are about the same as given by Holttum. Indusia are a

medium brown colour in living specimens.

Athyrium boryanum (Willd.) Tawaga, (Holtt. p. 567).*

Near streams by road to Cameron Highlands, Perak, at 22%

milestone; 2,000 feet, August 1958; specimen no. 4006. A few

plants with large rootstocks were found just above a stream on

rocky ground at the forest edge. Although this fern has been col-

lected in Perak previously, it has not been found in this State in

recent years.

In February 1959 I came across a few plants of Athyrium bory-

anum near Trinkap, in the Cameron Highlands area, at c.4,000

feet altitude. The ferns were growing in the protection of a large

rock in a fairly open place on the roadside. Fronds were large and

fertile and all quite new.

This fern is certainly seasonal in its growth, fronds dying off at

certain periods, so this may be the reason that in Malaya, it escaped

the notice of collectors for about 50 years. In Perak there were no

plants showing in June, and in August they were all new. At Fra-

ser’s Hill where I first saw Athyrium boryanum in 1952, my notes

indicate that new fronds are produced earlier than in the Perak

plants. During April, June and August I have recorded new fronds,

but there were none at all in January when I searched for it in the

same place.

A note should be made of the rootstock of Athyrium boryanum.

Descriptions usually state that the rhizome is either erect or sub-

erect, whereas in the Malayan form it is most ecrtainly creeping,

with fronds borne closely together. The growing point occurs at

one end of the longitudinal rhizome, which is usual in truly creep-

ing rootstocks. The rhizome is thick and fleshy, and one which I

have recently brought from Fraser’s Hill for cultivation, measures

15 cm. long by 7 to 8 cm. through, and this is by no means the

largest. About 2 cm. of the old stipe bases remain attached in a

horizontal manner on the rhizome, each facing the growing-point.

They remain green and fleshy until they eventually fall off. The

oldest ends of large rhizomes are usually pitted with hollows where

the stipe bases have been.

Athyrium amplissimum (Baker) Holttum, (Holttum p. 569).

Cameron Highlands, Pahang, near Walkerburn Cottage; c.5,000

feet altitude. Specimen nos. 2935, 3936. On light forest edge in

fairly open places and in light shade. Previously found in Cameron

* Since preparing these notes, this species has been referred to a new

genus, Parathyrium Holttum, Kew Bulletin, 3, 1958, p. 448.

270

Vol. XVII. (1958).

Highlands, but only fronds with old sori had been seen and the

presence of an indusium had not been determined. In August 1956

I found several plants with fertile fronds. On these I found no

indusia, nor evidence that there had been any, and one of the fronds

had young sori which ought to have shown them, had they been

present, unless they fall long before maturity. Immature sporangia

were pale greenish in colour, but were dark and almost black when

mature.

No note, so far as I am aware, has been made of the mucilage

which completely covers new fronds at the top of the stout erect

stock. The mucilage was copious, and was present on all the plants

I saw; there were no scales visible and I seldom found any on the

new growth. There were, in fact very few broad scales present, the

stipes having none, but as Holttum points out, the scaliness varies

a great deal.

Athyrium dilatatum (Bl.) Milde, (Holttum p. 574).

Fraser’s Hill, Pahang, at c.4,000 feet; specimen nos. 3325, 3328,

4197. On two occasions, a few plants were found growing in

shady places in wet rocky ground, by pathsides in tall forest. One

was below Girdle Road near Peninjau, and the other was below

Lady Maxwell Road, both being near streams. In the third locality

plants of this fern were abundant in a narrow wet ravine, in tall

forest near the golf course, and so it is possible it is not uncom-

mon in similar places throughout this area. The plant sizes agree

with Holttum’s measurements, but the pinnules were cut half way

to the costae or less, never more. The colour of the living frond is

dark green.

Coniogramme fraxinea (Don) Diels, (Holttum p. 589).

Near the road to Cameron Highlands, Perak, between 2,000 and

3,000 feet. On sides of rocky streams in tall forest, it is not un-

common here. A persistently variegated form, which I have also

seen on the Taiping Hill, occurs frequently in the former locality.

Vittaria angustissima Holttum, (Holttum p. 610).

Cameron Highlands, Pahang; April 1958. On tree trunks; slopes

of Gunong Kemunting, c.5,000 feet, specimen no. 3941; slopes of

G. Jasar, c.5,400 feet, specimen no. 3928.

Vittaria flexuosa Fee, (Holttum p. 611).

In Malaya, the usual habitat for this fern is on trees in forest

where conditions are constantly humid (see Holttum). In the Gin-

ting Simpah forest on the Selangor side, there are many small to

very large granite outcrops. These usually contain plants quite

271

Gardens Bulletin, , s:

different from those in the forest surrounding these cliffs. Here,

between 700 and 2,000 feet, V. scolopendrina is usually to be

found with its rootstock wedged in small crevices in the vertical

rock. Usually the fronds are of normal size, others being fertile

when scarcely 20 cm. long. These outcrops are seldom near streams

and although sheltered by the tall forest trees, frequently become

very dry. |

Vittaria flexuosa Fee, (Holttum p. 611).

Penang Hill, Penang, c.2,500 feet; May 1956, specimen no.

2822. This plant agrees with the description of this species in

Holttum. It was growing out of an epiphytic spongy root mass

which was dead, but similar to that of an Asplenium nidus L. The

Vittaria was intermixed with V. ensiformis Sw., from which it was

easily distinguished. The fronds of V. flexuosa were pendulous, the

longest being 90 cm., but none was wider than 5 mm., often

narrower. The stipe was slender, the underside black and shiny and

the upper surface black at the very base. There were new fronds

with immature sori. When dry the frond edges become strongly

incurled.

The root mass was in the fork of a small tree in short, rather

sparse secondary growth on Western Hill and was fully exposed.

Only a small portion was collected and so it is hoped that the re-

mainder will survive in this locality.

Salvinia cucullata Roxb. ex Bory, (Holttum p. 621).

Kota Bharu, Kelantan, May 1957, specimen no. 3399. Previ-

ously collected from Kedah. I found that this little fern was not un-

common on the edges of the rice fields near the airfield at Kota

Bharu. They were in small patches, to about 13 cm. long and were

often mixed with Azolla pinnata R.Br. At this time of the year

nearly all the fields were dry, but in corners where moisture re-

mained, Salvinia was seen. I did not find any fertile plants.

I am grateful to Dr. R. E. Holttum for his continued help in

comparing and identifying my material at Kew, and to the Director

of the Singapore Botanic Gardens for checking nomenclature.

212

An interesting new record from the Malayan beach:

Spilanthes urens Jacq., its synonymy and distribution

By J. H. KERN & C. G. G. J. VAN STEENIS

Flora Malesiana Foundation, Leyden

IN 1934 Mr. Corner collected on Jason Bay, Johore, a remarkable

creeping Composite which at the time defeated identification. It

grows as most beach plants with a horizontal, more or less sub-

terranean stem, at the nodes giving off downward roots for anchor-

age and upward simple (lateral) stems (10-40 cm. high), each

with a few pairs of opposite narrow leaves and a terminal, rather

long-peduncled (5—15 cm.) head. The leaves are strictly opposite,

3-plinerved, 3-93 by 0-4-14 cm. The head consists entirely of

tubular, white flowers and measures c. 1 cm. diameter. The achene

is rather peculiar, being flattened elliptic, with one, less frequently

two, awns on top and 2(—3) yellowish, rounded ridges (obviously

corky), distinctly set off against the rather dark colour of the fruit

itself.

It has now appeared that it had been referred in Spilanthes, a

well-known genus of mostly erect, annual weeds. Bentham (1866)

had referred a North Bornean specimen collected by Barber to the

North Australian S. anactina F.v.M.; Martelli (1883) identified a

specimen collected by Beccari in Sarawak as belonging to the

American S. urens Jacq., and finally A. H. Moore (1907) had

described a North Bornean specimen as a new endemic species,

S. chamaecaula A. H. Moore.

The remarkable thing about the latter species appeared its rarity,

its exacting habitat, and the fact that it seems to lack immediate

relationships with any other Malaysian species in the infrageneric

system of the monographer A. H. Moore. He placed it with the

S. American S. urens Jacq. and some other species.

This circumstantial evidence raised our suspicion as it does not

fit with the generally wide distribution of plants restricted to the

tropical beach.

This led us to a closer examination of the characters A. H.

Moore used in his revision to discriminate it from its allied species.

It appeared that S. chamaecaula was keyed out against S. urens

on the strength of two characters, viz. the uniaristate achene (in

the other species there are 2 bristles) and the corky swollen ridges

which were said to be absent in the other species.

OA be

Gardens Bulletin, S.

The first character broke down at once in the Bornean material,

as in one head both 1- and 2-aristate achenes were found to occur.

The second character appeared at first sight satisfactory in our

scant material at Leyden. But in a bunch of S. American material

of S. urens, which we borrowed through the kindness of Dr. Lan-

jouw and Dr. Jonker from the Utrecht Herbarium, we found per-

tinent proof that in S. urens, which both in habit and in habitat

agrees so very well with S. chamaecaula, in all heads containing

mature achenes some of these are provided with the characteristic

swollen corky ridges.

This induced us to re-examine a complete head of the scant

Malaysian material and here we found to our satisfaction that be-

sides the corky achenes there were in one single head also achenes

without these ridges. Though it is possible that the ratio of achenes

with and those without corky ridges differs slightly in Malaysian

versus American material, it is useless for taxonomic distinction.

It is rather astonishing that the presence of ridged achenes has

hitherto been overlooked in S. urens, which is not a particularly

rare plant in America. This demonstrated to us that for a good

examination one must dissect an entire head; it is insufficient to

pick just a few random achenes. In doing so we did not succeed in

finding any regularity in the location of the corky achenes in the

head.

We have come to the conclusion that Martelli, who referred the

Bornean Beccari material to S. urens, was perfectly justified.

In the Old World a third species had been described from the

islands in the Gulf of Carpentaria, viz. S. anactina F.v.M. Already

Bentham, in the Flora Australiensis, had referred a Bornean col-

lection of Barber (obviously in the Kew Herbarium) to it. F. von

Mueller had described his species without having available ripe

achenes. As the Queensland specimens agree perfectly in habit and

habitat according to the description, we feel entirely at ease in

reducing S. anactina also to the synonymy of S. urens. We feel the

more confident as A. H. Moore merely distinguished S. urens from

S. anactina by saying that it was found in the New World while

S. anactina is from North Australia. There seems no doubt that

this also represents the same species.

There is a fourth species which we feel should be critically re-

examined, namely S. pusilla Hook. & Arn., from the New World.

We have a strong suspicion that this is also conspecific with S.

urens. The differences cited in Moore’s key to exist between these

two species appear very slight: the linear-spathulate leaves attri-

buted to S. pusilla are quite common in S. urens.

274

Vol. XVII. (1958).

It may seem remarkable that the localities in the Old World

tropical beaches are so scattered and distant, but this is by no

means uncommon in widely dispersed beach plants and is simi-

larly encountered in mapping localities of Scaevola plumieri,

Ipomoea pes-caprae ssp. pes-caprae, Launaea sarmentosa, Suriana

maritinia, Digitaria longissima, Triumfetta procumbens, etc.

The distribution of S. urens ranges from tropical and sub-tro-

pical Central and South America to North Australia (Gulf of

Carpentaria), and West Malaysia.

As far as we have ascertained within the scope of our examina-

tion the synonymy and the distribution in Malaysia run as fol-

lows :—

Spilanthes urens Jacquin, En. Syst. Pl. Carib. (1760) 28; Select.

Stirp. Am. Hist. (1763) 214, t. 126, fig. 1; Martelli, Nuov. Giorn.

Bot. Ital. 15 (1883) 296; Merrill, En. Born. (1921) 589.

?Spilanthes pusilla Hook. & Arn. in Journ. Bot., London 3 (1841)

B47.

Spilanthes anactina F.v. Mueller, Fragm. Phyt. Austr. 5 (1865)

63; Bentham, Fl. Austr. 3 (1866) 541; Bailey, Queens]. Fl. 3

(1900) 863.

Spilanthes chamaecaula A. H. Moore, Proc. Am. Ac. Arts &

Sc. 42 (1907) 528; Koster & Philipson, Blumea 6 (1950) 353.

Borneo. North Borneo: Burbidge (Gray); Barber (K). Sara-

wak: pr. Sibu, flowers white, leaves more or less fleshy, Beccari

P.B. 1750 (Fi, L), June 1866.

Malay Peninsula, Johore: Sg. Tuenseh, Jason Bay, Corner S.F.

28459 (Sing, Bo, K, L, A), June 1934.

We hope that this little note will draw attention to this curious

plant and stimulate collectors to find new localities.

275

A New Aroid from Sarawak

By C. X. FURTADO, Singapore.

AN ANOMALOUS AROID was noticed in the collections made three

years ago in Sarawak by Mr. J. W. Purseglove, then the Director

of the Botanic Gardens, Singapore. It belongs to the group of

aroids which shed off the upper portion of the spathes before the

fruits are formed on the enclosed spadices. Since the sheaths of the

petioles of its leaves are broad and produce long, tongue-shaped

free portions, the aroid could be a species of either Piptospatha or

Microcasia. In their general appearance and the way they bend, the

spathes resemble those of a Piptospatha, and the spadix is fertile

up to the apex, a character mentioned for Piptospatha only and

not for Microscasia which is described to have the spadix with a

sterile apex or with an apex with sterile flowers. But in Purse-

glove’s specimen the male flowers which occupy the upper two-

thirds of the spadix have each two horns through the apical pores

of which pollen is shed; and bicornulate stamens form a definitive

character of a species of Microcasia. Hence the new species is

assigned to this genus and named Microcasia purseglovei in honour

of the collector.

Hitherto only two species of this genus were known, M. elliptica

and M. pygmaea, both of which had helped to define the genus as

having, among other things, a sterile apex to the spadix; but in view

of the characters of the new species, the generic definition has to

be altered to include also the species with fully fertile spadices.

The two earlier described species were very tiny plants; but the

new species is a very much larger plant in regard to the vegetative

and reproductive parts.

Microcasia purseglovei Furtado spec. nov.

A duabus speciebus usque adhuc cognitis haec species stirpibus

omnino majoribus, spadicibus ad apicem usque fertilibus differt.

Caudiculus elongatus, repens, in parte erecta 25—30 cm. altus.

Folia: petioli tenues, laminis paulo breviores vel eis aequantes, 6-8

cm. longi; vagina papyracea, superne in partem liguliformiter

solutam 2-53 cm. longam producta; lamina supra saturate viridis,

276

Vol. XVII. (1958).

subtus albo-squamosa, ambitu elliptica, utrinque paulatim cur-

vato-attenuata, apice in pseudo-tubulum abrupte convoluta, margine

angustissime revoluta, 8-15 cm. longa, 2:5—3-5 cm. lata, nervis

Gardens Bulletin, S.

primariis paucis cum secundariis adscendentibus, dein margine

sub-parallelis, utrinque vix prominentibus praedita. Pedunculus

petiolos superans, 12—18 cm. longus, apice curvatus, spatham hori-

zontalem modo Piptospathae spp. ferens. Spatha circa 5 cm. longa,

leviter convoluta, cum tubo brevi viridescente persistente quam

lamina caduca albescens circa triplo breviore. Spadix quam spatha

brevior, breviter stipitatus, circa 2-5 cm. longus, omnino fertilis;

pars feminea basalis, parte mascula duplo brevior. Ovarium 1 mm.

altum, circa duplo diametiens, stigmate depresso. Stamina

bicornuta. Bacca ignota.

BoRNEO: Sarawak, Tau, juxta flumen Mayeng dictum (Purse-

glove 5344, Holotypus in SING, isoholotypi in K, L, SAR.).

Microcasia purseglovei Furtado (Holotypus)

A, Planta. Al, Folii apex. B, Spatha. C, Spadix cum parte spathae. D,

Ovarium. E, Ovarium longitudinaliter sectum. F, Stamen antice visum.

G, Stamen postice visum.

278

Some New or Noteworthy Species of Malaysia

By C. X. FURTADO

WHILE ARRANGING the material of the genus Ardisia in the Singa-

pore Herbarium, some new species or new records were noticed,

while others had to be excluded from the genus. The status of some

of these will have to be settled by comparing the types; but the

following notes are published here in order either to clarify some

confusions that existed in the herbarium and also in the recently

published literature on the subject, or to give a status to some of the

names that have been adopted in the Singapore herbarium where

the types and other specimens cited in this paper are preserved.

As originally published by Mez, the sections mentioned here were

subgenera, but they have been treated as Sections by Merrill and

others, an arrangement that seems to be more in accord with the

present concepts of nomenclature.

ARDISIA Swartz

Section PIMELANDRA Mez.

1. Ardisia ferox Furtado sp. nov.

A. pachysandra (Wall.) Mez sensu Ridl., Fl. Mal. Pen. II

(1923) 240 p.p.

Ab A. fuliginosa, cui affinissima, innovationibus deciduo hirsutis;

foliis utrinque glaberrimis, punctulis majoribus margines versus

sitis, vix reticulatis; inflorescentiis valde abbreviatis, omnino hir-

sutis; sepalis liguliformibus, hirsutis, basin versus fere +-connatis;

ovartis baccisque longe pilosis sat dissimillima.

Ramuli validiusculi, novelli dense ferrugineo pilosi, adulteriores

mox glabrati. Folia petiolis hirsutis vel glabris, 10-12 mm. longis,

in ramulos lineatim decurrentibus stipitata, coriacea, elliptica, 12—

25 cm. longa, 4-8-5 cm. lata, utrinque glaberrima, nitida, nervis

Jateralibus utrinsecus 14-16 patentibus, prope margines arcuatim

ascendentibus et inter se unitis praedita, apice obtusa, basin versus

oblique angustata, immo paulo decurrentia. Inflorescentia valde

abbreviata, flores hirsutos gerens. Sepala liguliformia, hirsuta,

basi fere 4-unita, haud imbricata. Petala sepalis longiora, extus

hirsuta, ovata, acuta. Stamina ovato-acuta, petalis breviora. Ova-

rium valde hirsutum; ovula glabra, verticaliter corrugata. Bacca

pisiformis, ferrugineo-pilosa, circa 5 mm. in diam.

279

Gardens Bulletin, S.

‘

5mm 2mm

4mm

Ardisia ferox Furtado (A-B1: Holotypus; C-H: Mat s.n.)

A, Ramulus cum foliis majoribus. B, Ramulus cum infructescentiis.

Bi, Fructus cum calyce. C, Flos ante anthesin. D, Petala duo postice

visa. E, Stamen antice visum. F, Ovarium. G, Placenta. H, Fructus

juvenilis.

280

Vol. XVII. (1958).

MALAYA: Johore, Gunong Pulai (Nur & Kiah 7,778—Holo-

typus; Ridley 12,189; Mat s.n. in 1892): Omnia specimina in her-

bario Singaporensi conservata.

2. Ardisia fuliginosa Bl., Bijdr. (1826) 692; Mez in Engl.

Pflanzenr. [IV 236 — Heft 9 (1902) 99.

A. tomentosa Presl., Rel. Haenk. II (1835) 66: Mez in Engl.

Pflanzenr. IV, 236 — Heft 9 (1902) 100; Merrill, Enum. Philipp

Pl. HI (1923) 265. syn. nov.

A. disticha A. DC., Prodr. VIII (1844) 129.

SUMATRA: Atjeh (v. Steenis 6,186); Takigeum (W. & C.

Bangham 726); loc. incert. (Forbes 2,142).

BORNEO: North Borneo, Marudu Kudat (Austin A 1,191);

Tawao (Elmer 20,462). Sarawak, Rejang (Haviland 3,029).

JAVA: Besuki (Steenis 10,826).

PHILIPPINES: loc. incert. (Cuming 849, isoholotypus of

A. disticha); Jamindan (Ramos & Edano 30,999); Mt. St. Isidoro

(Fenix 30,008); Angat (Ramos & Edano 34,062), etc.

There is a good deal of variation in the tomentum of the leaves

as well as in the branching of inflorescences.

3. Ardisia korthalsiana Scheff., Comm. Myrs. Arch. Ind. (1867)

57; Mez in Engl. Pflanzenr. IV 236 — Heft 9 (1902) 99.

A. teysmanniana Scheff. sensu Ridl., Fl. Mal. Pen. II (1923)

240 p.p.

MALAYA: Trengganu (Moysey & Kiah 33,384). Pahang,

Gunong Tapis (Symington & Kiah 28,825); Sungei Tahan (Kiah

s.n.); Sungei Teku (Kiah 31,709); Wray’s Camp on Tahan (Ridley

16,241). Malacca, loc. incert (Alvins 1755, as Kayu Segankan &

582 as Meddang Panjang); Bukit Bruang (Derry 456, as Tambang

Sisir); Lubok Kadondong (Ridley s.n. on 13—VI-1892); Bukit

Besar on Mt. Ophir (Ridley in Dec. 1898); Selandor (Alvins

673 — 251, as Pokok Lagan). Johore, Gunong Belumut (Holt-

tum 10,693 & 10,788).

SUMATRA: West Indragiri in Taluk region (Meijer 4,077 as

Balam Siram).

ANAMBA ISLANDS: North Jemaja near Padang (Henderson

203392).

There is a good deal of variation in the specimens. Some (e.g.

Moysey & Kiah 33,384 & Kiah 31,709) are practically glabrous

and shining in the lower surface with a few minute hairs on the

281

Gardens Bulletin, S.

midrib and with minutely dense reticulations and black glands on

the upper surface. The Johore specimens have glabrous and more

coriaceous leaves, and though the leaves are similar, black glands

are conspicuous on their upper surface in Holttum 10,788 but not

in Holttum 10,693. Many other specimens have leaves which are

more pilose beneath.

This is the first record in Malaya.

4. Ardisia pachysandra (Wall.) Mez in Engl. Pflanzenr IV 236

— Heit 9 (1902) 97 fig. 14 A-E, partim; King et Gamble, Mat.

Fl. Mal. Pen. IV (1905) 331; Ridl., Fl. Mal. Pen. II (1923)

240 p.p.

MALAYA: Kedah, Gunong Jerai—Kedah Peak (Ridley in VI

—1893); Gunong Bintang (Haniff 21,118). Perak, Piah (Penak

39,221); Pulau Rumbia (Seimund on 28—-III—1926); Tapah Rid-

ley 14,062). Penang, Telok Bahang (Curtis — 1156, as Limpa-

nan); Waterfall (Curtis 3,737; Nur 1,225); Government Hill

(Curtis = 1156; Ridley — Curtis 1156); Penara Bukit (Curtis

1156); loc. incert. (Curtis ? 3181 & 3164). Selangor, Weld Hill

(Rahman, Tree 334). Johore, Gunong Panti (Ridley in Dec.

1897); Sungei Kayu (Corner 29,197).

BANGKA: (Anta 212, & 685 & 966).

ANAMBA ISLANDS: near Terempak in Siantan (Henderson

20,186); Gunong Ranai in Bunguran (v. Steenis 1287); Padang

Leting.

JAVA: Tjibodas (Yates 2,924); Gunong Gendero (Kanta 236);

Besoeki (Buwalda 7,255 and 7,377). |

This species is listed here to show the range of distribution.

Usually non-Malayan specimens of this species are distributed un-

named or as A. fuliginosa.

There is a good deal of variation in this species, in regard to the

size of peduncle and the sepals and in the distribution of tomentum

and glands on the leaves; but the ovaries are glabrous. Some Javan-

ese specimens can be mistaken for A. fuliginosa if ovaries are not

examined. :

Ridley 176 & 2,800? which were cited here by Mez, have been

transferred to A. teysmanniana.

5. Ardisia teysmanniana Scheff in Tijdschr. Nederl. XX XI (1870)

368; Mez in Engl. Pflanzenr IV 236 — Heft 9 (1902) 97;

King et Gamble, Mat. Fl. Mal. Pen. IV (1905) 122; Ridl., Fi.

Mal. Pen. II (1923) 241 p.p.

A. pachysandra Mez op. cit. p. 97 partim.

282

Vol. XVII. (1958).

MALAYA: Perak? loc. incert (Ridley 462). Malacca, Sungei

Udang (Derry 1,143 as Medang Ketanak). Johore, Sungei Sedili

(Ngadiman 36,894). Singapore, Sembawang (Ridley 5,573);

Chan Chu Kang (Ridley 6,788); Changi (Ridley 2,800; 5652 &

176; Goodenough 2,800); Bukit Timah (Ridley 2,800; Corner

sn.; Chew 10).

Ridley 176 & 28004 were cited by Mez under A. pachysandra.

Section AKOSMOS Mez

6. Ardisia laevigata Bl., Bijdr. (1826) 690.

MALAYA: Pahang, Cameron Highlands, 1500-1700 m. alt.

(Henderson 18,022; Symington 21,016; Osman 20,810; Holttum

24,985).

JAVA: alt. 1500-2400 m.: Pasoeroea (N.-Ind. Flor. Serv.

No. 7, 143); Tjibodas (Sapun 239; Burkill 8,265).

This is a first record for this species outside Java. Burkill notes

that the flowers are downwardly directed and pinkish white, leaf-

stalks crimson and berries round greenish, all found on the same

plant at the same time. Henderson records as follows: “Small,

straggling tree in swampy places. Ripe fruit red.” Symington’s notes

suggest that the tree is 15 ft. tall and the fruit is green at first but

turns red later and finally black.

The Malayan specimens show more coriaceous leaves and

broader sepals which are somewhat ciliolate along margins. But no

material differences are noticed to separate these from the Javan

forms. Gland dots on sepals and petals may be found in the

Malayan as well as the Javan specimens.

7. Ardisia praetermissa Furtado sp. nov.

Inter species AKOSMOSIS insignis: innovationibus lepidotis; foliis

coriaceis, ellipticis, primo utrinque serius subtus tantum lepidotis,

margine integerrimis, laxe reticulatis; inflorescentiis quam petioli

1-5—2:5 cm. longi altioribus, in ramulos primarios apice floribus

pedicellatis congestis et bracteolis ciliolatis fere linearibus praeditos

divisis; sepalis punctulatis ciliolatis quam petala valde brevioribus,

basi 4—1/3-unitis; staminibus petala brevioribus apice subito acu-

minatis.

Frutex circa bimetralis, in innovationibus lepidotus. Folia ellip-

tica, coriacea, 12—18 cm. longa, 4—5-5 cm. lata, basin versus sensim

vel rotundato attenuata, in petiolos 1-2-5 cm. longos, lepidotos,

supra canaliculatos, margine scariosos paulo decurrentia, apice

283

Gardens Bulletin, S.

(2D

aw fla

EKA IE

ATT

TAN

3cm

6mm

Ardisia laevigata Bl. (Holttum 24,985)

A, Habitus. B, Floris alabastrum. C, Petala duo antice visa. D, Stamen

postice visum. E, Ibid antice visum. F, Ovarium. G, Placenta.

284

Vol. XVII. (1958).

3cm

Ardisia praetermissa Furtado (Alvins 342—Holotypus)

A, Habitus. B, Inflorescentiae ramulus cum bracteolis floreque. C, Flos

petalis desumptis, ut stylus apice bifidus (?) appareat. D, Petalum.

E, Stamen postice visum. F, Placenta nonnihil tabida.

285

Gardens Bulletin, S.

obtusa vel acuminata, primo utrinque dein subtus tantum lepidota,

supra veridescentia et in costa mediana valde sulcata, patente ner-

vosa, laxe reticulata, margine integerrima, punctulis multis minutis

subtus fere inconspicuis supra saepe manifestis aucta. Inflorescentia

axillaris, unica completa visa, circa 5 cm. longa, axi lepidota

in 7-8 ramulos ad 12 mm. usque longos, apice umbella-

tim vel subumbellatim floriferentes, bracteolis persistentibus, circa

1—2 mm. longis, linearibus vel lineari-lanceolatis, margine valde

ciliolatis praeditos divisa. Flores pedicellis 3-4 mm. longis deciduo

lepitodulis suffulti, ante anthesin stylis exsertis aucti, in alabastro

circa 5 mm. longi. Sepala extus minute lepidota, basi 4-unita, in

lobis ovata vel ovato-oblonga, apice acuta, dorso sparse punctulata,

margine albescentia ciliolataque, circa 2 mm. alta. Petala sepalis

subtriplo longiora, apice acuta vel obtusa, fere ad basin libera.

Ovarium glabrum; stylus tortuosus, apice bilobatus?

MALAYA: Malacca, Selandor (Alvins 679342, nom. vern.

Pokoh Pelandok Payah—-Holotypus in SING.).

This specimen was collected by Alvins on Jan. 30, 1885, and was

assigned in our herbarium first to A. crassa and then to A. colo-

rata. According to Alvins, this plant is common in swamps and

grows to a height of 6—7 ft. The label was originally numbered 679,

and subsequently it was numbered 342, but the original number

has not been cancelled.

The inflorescence is axillary bearing very short branches. In the

axis of the only one complete irfflorescence available, there are

marks of fallen off branches and below its base a scar of the axil-

lant leaf. Only 3 flowers were available, all badly preserved. The

style of the one examined bore a mark at the apex as if it was

bifid. The ovary and the placenta showed marks of disintegration.

The petals seem to be free nearly to base.

8. Ardisia singaporensis Rid]. in Journ. Roy. Asiat. Soc. Str. Br.

79 (1918), 92 et FE Nigieren Tl (1923) Zot

MALAYA: Singapore, Pulau Ubin (Ridley 2,816, Isosyntype

and s.n. in 1891; Goodenough on 3—III—1893); Changi Road

(Goodenough 2,833, Isosyntype); Changi (Goodenough 2,814):

All specimens cited are in Singapore herbarium.

The inflorescences are borne usually in the axils of the leaves

near the apex, but sometimes also of the leaves much below the

apex. The flowers are borne in umbels at the end of primary or

secondary branches of the inflorescence axis, rarely at the end of

the unbranched axis itself.

286

Vol. XVII. (1958).

This species should be placed in Section AKosMos and is in no

way related to A. miqueliana. Specimens of the species were dis-

tributed as A. ferruginea var. glabrata and some were named in

Singapore herb. as A. tuberculata:

Section STYLARDISIA Mez

9. Ardisia miqueliana Scheff. in Tijdschr. Ned. Ind. XXXI (1870)

367 et in Flora LIII (1870) 253; Mez in Engl. Pflanzenr. IV

236 — Heft 9 (1902) 115.

A. ferruginea Mez op. cit. (1902) 108; King et Gamble, Mat.

Fl. Mal. Pen. IX (1905) 325; Ridl., Fl. Mal. Pen. II (1923) 250,

non A. ferruginea H.B. & K. (1818)—Syn. nov.

A. rudis Sinclair in Gard. Bull. Sing. XV (1956) 24—Syn. nov.

MALAYA: Johore, Kampong Simpai (Lake & Kelsall, 4032,

Isoholotypus of A. ferruginea in Herb. Sing.); Kluang (Holttum.

9,356); Bukit Lenggor (Holttum 24,927); Sungei Kayu (Corner,

32,236; Kiah 32,076); Gunong Belumut (Holttum 10,621).

Singapore, Nee Soon (Sinclair 40,278).

Already King & Gamble had noted that the leaves were exactly

like those of A. miqueliana. The specimens collected by Lake &

Kelsall were apparently from a plant just beginning to flower and

so the fact that the inflorescence was actually terminal became

obscured due to the well developed lateral branches at the base of

the terminal panicle. I do not know the reasons that led Ridley to

transfer A. ferruginea Mez from Section AkosMos to Section

CRISPARDISIA. But there is no doubt that the species should be

placed in Section STYLARDISIA, the few well developed flowers

showing their styles well above the petals of the unopened flowers.

The holotype of A. miqueliana was from Bangka. Corner notes

that the plant is 20 ft. tall with monopodial side branches and with

silvery grey pustular bark with small lenticels: Twigs, petioles and

underside of leaves old hairy. Inflorescence hanging at the end of the

side branches. Pedicels purplish upwards and at top. Bracts green.

Sepals greenish white, but magenta at base. Petals white with a

magenta spot at base. Filaments magenta; connective greenish,

anther yellow.

10. Ardisia nurii Furtado spec. nov.

Ab A. crassa, cui valde affinis, foliis rigidioribus, nervis laterali-

bus reticulationibusque subtus validioribus, inflorescentiis omnino

minus tomentosis, floribus gracilioribus, sepalis fructibusque mino-

ribus haec species facile distinguenda. Ab A. colorata var. colorata,

287

Gardens Bulletin, S.

IO <7,

[Lae

Wz:

({l

IAS

(fie

(pl

Ey DX 8

if le ;

Wy)

Ardisia nurii Furtado (Nur 10,511—Holotypus)

A, Ramulus fructiferus. B, Fructus.

288

Vol. XVII. (1958).

quacum confusa, foliis coriaceis nitidisque, bracteolis flores axil-

lantibus persistentibus, floribus majoribus robustioribusque sat

distincta.

Arbusculus 3—4 metralis, ramulos robustos gerens. Folia ob-

longo-elliptica, apice acuminata, basi oblique acuta vel oblique

decurrentia, vel rotundato-acuta, 18-25 cm. longa, 6—7 cm. lata,

utrinque vel supra tantum nitida, subtus brunnescentia et lepidibus

minutis deciduis praedita, glandulis haud visibilibus, nervis laterali-

bus primariis pluribus subtus valde prominentibus supra immersis,

in nervum intramarginalem interruptum conjunctis, nervis secun-

dariis multo brevioribus, reticulationibus densis paulo prominulis

percursa, petiolis 2-3 cm. longis stipitata. Inflorescentia apicalis,

pyramidalis, in ramulos arcuatos bi- vel tri-pinnatim divisa, ferru-

gineo lepidota. Bracteolae minutae, linguiformes, ferrugineo-lepi-

dotae, margine ciliatae, pedunculi apice fructiferi persistentes.

Bacca subglobosa, circa 4 mm. longa, apice compressiuscula, nigra,

laevis; pedicelli umbellati, dense lepidoti, 6-10 mm. longi; sepala

ovato triangularia, extus dense lepidota, margine flavescentia

ciliata, dorso atro-pustulosa, basi breviter tubulosa.

MALAYA: Pahang, Fraser’s Hill, alt. 1300-1500 m. (Nur

10,551—Holotypus; Burkill & Holttum 8,514 and s.n. = 8,514;

Henderson 11,232). Omnia specimina in Herb. SING. conservata.

This appears to be localised to Fraser’s Hill. The duplicates of

the numbers were distributed to other herbaria under A. colorata

from which this species is distinguished by its coriaceous and shin-

ing leaves, persistent bracteoles at the base of the fruits and larger

and more robust fruiting calyces. The species appear more close to

A. crassa which, however, bears much larger flowers and fruits,

thicker tomentum on calyces, and less prominent veins and reti-

culations on the lower surface of the leaves.

11. Ardisia sessilis Scheff.. Comm. Myrs. Arch. Ind. (1867) 69;

Mez in Engl., Pflanzenr. IV. 236 — Heft 9 (1902) 115.

MALAYA: Singapore, Seletar (Ridley s.n. in 1894); Chan Chu

Kang (Ridley 3,653 & 3,844).

BORNEO: North Borneo, Tawao (Elmer 21,404 & 21,424).

East Borneo, Pelawan Besar (Act. 745).

This material has been named on the basis of the Fs stion

given by Mez, and requires verification by comparing with the type.

The Singapore specimens were quoted by Ridley under A. crassa

in Fl. Mal. Pen. II (1923) 249, while Elmer’s collections were

referred doubtfully to A. fortis Mez by Merrill in Univ. Calif. Publ.,

Bot. XV (1929) 234.

289

Gardens Bulletin, S.

ry Pbk

Ardisia breviramea Merr. (Elmer 20,425—Isotypus)

A, Ramulus floriferus. B, Flos. Bi, Ibid sepalo desumpto ut petala fere

libera appareant. C, Petala duo cum staminibus postice visa. D,

Stamina calyce corollaque desumptis. E, Stamen postice visum. F,

Calyx. F, Ovarium. H, Placenta.

290

Vol. XVII. (1958).

The flowers are practically sessile and bear ovate, entire, acute

sepals which are ciliate in the margins. The primary nerves tend to

join and form an intramarginal nerve on the leaves.

Section ACRARDISIA Mez

12. Ardisia breviramea Merr. in Univ. Calif. Publ. Bot. XV

(1929) 235.

NORTH BORNEO: Tawao (Elmer 20,425—AIsoholotype in

SING.).

Since the inflorescences are apical (a fact also noted by the

author of the species) and since the styles do not protrude just

before the flowers open, the species cannot be placed in the sec-

tions STYLARDISIA or AKOSMOs to which Merrill had referred

doubtfully. He had not seen the flowering specimens. Since the

Singapore specimen is the duplicate of the holotype and bears

flowers, it has been illustrated here as a help in understanding this

species which was described from a fruiting specimen only. As in

A. praetermissa, the petals are free almost to the base.

13. Ardisia goodenoughii Furtado sp. nov.

Ab A. divergente, cui valde affinis, haec species foliis majoribus

conspicue pustulatissimis, nervis lateralibus primariis pluribus,

sepalis valde glandulosis differt. Ab A. polyacte, cui similis, petio-

lis longioribus, foliorum glandulis pluribus conspicuioribus, flori-

bus minoribus paucioribusque in axi distanter sitis, petalis haud

acuminatissimis facile distinguitur.

Frutex circa 3 m. altus. Folia oblongo- vel lanceolato-elliptica,

basi in petiolos 6-10 mm. longos corrugato-decurrentia, 12-20

cm. longa, 5—9 cm. lata, interdum ad 30 cm. usque longa et 11

cm. lata, omnino praecipue secus margines valde glandulosa,

glandulis juvenilibus atribus vetustioribus pustulosis, densissime

reticulata. Inflorescentia apicalis, in ramos floribus pseudo-umbel-

latim praeditos divisa, foliis fere aequalis vel brevior. Flores pedi-

cellis 10-15 mm. longis suffulti, deciduo-puberuli. Sepala ovata, —

apice rotundata, basi imbricata, margine minute ciliata, dorso valde

pustulata. Petala extus parce pustulata, sepalis fere triplo longiora,

ovata, apice acuta vel breviter acuminata. Stamina petalis breviora.

Stylus ex alabastro ante anthesin haud exsertus.

MALAYA: Kemaman, Ulu Kajang (Corner 30, 432); Bukit

Kajang (Corner s.n. on 30-X—1935); Ulu Bendong (Corner

30,046 & 30,099). Selangor, Bukit Tangkol in Ulu Langat (Mil-

lard 1,380 as Kayu Bilal); Sungei Lalang in Kajang (Symington

291

Gardens Bulletin, S.

Ardisia goodenoughii Furtado (A, Strugnell 12,716; B—H, holotypus)

A, Ramulus fructiferus. B, Ramulus floriferus. C, Flos. D, Corolla.

E, Stamen postice visum. F, Stamen antice visum. G, Ovarium.

H, Placenta.

292

Vol. XVII. (1958).

22,681); Ampang (Strugnell 12,716); Kuala Lumpur (Good-

enough 41, Comm. sub numero Ridleyano 10,481, holotypus in

SING); Ginting Bidai (Ridley as Mata Ayam); Seminyih (Hume

8,371 & 8,401); Ginting Simpah (Hume 9,525). (Omnia speci-

mina in Herb. Sing. conservata).

BORNEO: Sandakan (Cuadra A2,221 SING.).

The Kemaman and the Bornean specimens represent either

material taken from young, vigorously growing shoots or a variety

of the type form. The leaves are somewhat thinner in structure

and the glands are largely not pustulose. In the Kemaman speci-

mens the inflorescence is very much more slender in its axis,

branches and pedicels, and the leaves are also much larger.

Section TINUS (Burm.) Mez

14. Ardisia gambleana Furtado sp. nov.

A. rhyncophyllae affinissima, sed petiolis longioribus, foliis

obovato-ellipticis, utrinque dense reticulatis, nervis lateralibus

prope marginem arcuatim porrectis paulatim evanescentibus, haud

in lineam conspicuam intramarginalem conjunctis, inflorescentiis

sueto tri- vel bifloris, sepalis ovato-rotundis margine plerumque

emarginatis nudisque sat distincta.

Frutex circa trimetralis. Folia chartacea, obovato-elliptica, apice

eleganter acuminata, basin versus cuneata, novella lepidibus minu-

tissimis obscuris deciduis tecta, dein glabra, glandulis atris dense

punctulosa, nervis primariis utrinsecus 7-10, prope marginem

arcuatim porrectis praedita. Inflorescentia axillaris, tri- vel biflora,

pedunculo 2-3 cm. longo, pedicellis fere aequilongo praedita,

omnino glandulosa. Sepala basi breviter connata, valde imbricata,

dense glandulosa, margine flavescentia nudaque, ovato-rotundata,

emarginata. Petala sepalis fere triplo longiora, ovata, apicem

versus sensim vel interdum abrupte attenuata, summo acuta, extus

dense punctulata. Stamina petalis breviora, ovata, acuta, postice

glandulosa. Ovarium globosum, glabrum, pauci-ovulatum. Bacca

globosa, juventute flavescens glandulosaque, maturitate atra,

corrugata.

MALAYA: Pahang, Tahan River (Ridley 2,670—Holotypus in

SING.): Jerantut (Holttum 24,746); Kota Glanggi (Henderson

22,438); Sungei Sat (Henderson 22,030); Raub (Strugnell

~ 23,456). Omnia specimina in herb. Singaporense conservata.

The holotype specimen was referred doubtfully to A. rhynco-

phylla Clarke by King & Gamble in observation of A. biflora K. &

G. in Materials IV (1905) 143, but it was not included as a

Malayan species because of the imperfectness of the specimen.

293

Gardens Bulletin, S.

2mm

5mm

fr §

Imm

a JAHRE EAR Poy

a ap

Ardisia gambleana Furtado (A, Ridley 2,670; B—F, Holttum 24,746)

A, Ramulus fructiferus. B, Flos. C, Stamen antice visum. D, Ovarium.

E, Placenta.

294

Vol. XVII. (1958).

’

Ardisia tiomanensis Furtado (Henderson 21,686—Holotypus)

A, Habitus. B, Floris alabastrum. C, Flos petalis desumptis. D, Sepa-

lum. E, Stamen postice visum. F, Placenta.

295

Gardens Bulletin, S.

15. Ardisia tiomanensis Furtado spec. nov.

A. monticolae affinissima, a qua foliis obovatis, inflorescentiis

robustioribus, florum alabastris majoribus apice obtusis, sepalis

semi-orbicularibus vel reniformibus valde imbricatis, extus deciduo

violascenteque pubescentibus, margine ciliatissimis sat distinctis-

sima.

Arbusculus circa 8 m. altus, saxatilis, cum innovationibus viola-

ceo-pubescentibus. Folia petiolis lepidotis supra canaliculatis circa

2 cm. longis stipitata, coriacea, obovato-elliptica, apice acuta vel

acuminata, basin versus sensim acuta, utrinque deciduo lepidota,

nervis multis reticulationibusque densis supra fere inconspicuis

subtus prominulis percursa, multi-punctulosa vel punctulis abs-

conditis inconspicuis diffusa, bicoloria. Inflorescentia axillaris,

omnino deciduo violaceo-tomentulosa, ad 2 cm. usque peduncu-

losa, 5-10 flores umbellatim gerens; alabastra circa 6-8 mm.

longa, 5-6 mm. in diam., apice rotundata, pedicellis 8-12 mm.

longis, violaceo-pubescentibus suffulta. Sepala fere reniformia,

medio multo dilata, valde imbricata, basi breviter connata, extus

violaceo-pubescentia, minute punctulosa, margine pallida optime-

que ciliata. Petala deciduo ciliolata vel scariosa. Ovarium glabrum;

stylus ante anthesin haud exsertus. Bacca ignota.

MALAYA: Pahang, Pulau Tioman, prope Ayer Surin, alt.

300-700 m. (Henderson 21,686—Holotypus in SING.).

The collection has been distributed as A. littoralis Andr. which

has very slender, acute flower buds, more united and glabrous

sepals, narrower petals and septate anthers. Though the leaves are

obovate-elliptic in A. littoralis, the violaceous pubescence is absent

from the young growth. Similarly A. tiomanensis also differs from

A. monticola which has elliptic-lanceolate leaves, no violaceous

tomentum in any parts, very slender, acute flower buds, less imbri-

cate sepals and acute petals.

The collector notes that this species is common in the rocky

jungle, growing to 25 ft. or so tall and that flowers are pink.

Section PyrGcus (Lour.) Mez

16. Ardisia calcicola Furtado sp. nov.

Inter species ad Sectionem PYRGUM pertinentes peculiarissima

facile distinguitur: foliis intergerrimis dense punctulatis, nervis

lateralibus interrupte absconditis, haud reticulatis; floribus in

pedunculo abbreviato, longe bracteolato, apice ramuli producto

valde congestis; pedicellis conspicue punctulatis; sepalis pro rata

longissimis, fere ad basin usque liberis, quam petala punctulata

296

Vol. XVII. (1958).

Ardisia calcicola Furtado (Endert 5,134, holotypus)

A, Habitus. B, Inflorescentia. C, Flos. D, Petala duo cum staminis

postice visa. E, Flos juvenilis sepalis desumptis ut stamina tubifor-

miter collata appareant. F, Stamen postice visum. G, Ovarium.

H, Placenta.

297

Gardens Bulletin, S.

circa 4-brevioribus, ovato-lanceolatis, utrinque lepidotis, atro

punctulatis.

Suffrutex circa 50-75 cm. altus, flores in ramulorum apicibus

lateralium racemoso-congestos gerens. Folia phyllotaxi in ramulis

incerto excepto, subverticillatim collata, obovato elliptica, 30—35

cm. longa, 4—5 cm. lata, basin versus sensim attenuata, apice ob-

tusa vel subacuta, eis ramuli 3—4-plo longiora, supra viridescentia,

subtus brunnescentia, opaca, margine revoluta, utrinque pustulis

minutis dense praedita, nervis primariis interrupte absconditis

prope marginem arcuato ascendentibus utrinsecus circa 12, nervis

secundariis fere parallelis inter primarios 1—2 fere inconspicuis per-

cursa, haud reticulata. Flores 5-6 mm. longi; pedicelli 5-6 mm.

longi, pustulati, bracteolis ovato-lanceolatis saepe acuminatis non-

nihil breviores vel eis subaequilongi. Sepala utrinque lepidota,

dextrorsum tegentia, atro-punctulata, petalis circa 4-breviora, fere

ad basin usque libera, ovato- vel oblongo-elliptica, apicem versus

sensim acuta. Petala sepalis paulo longiora, acuta, dorso punctu-

lata. Stamina breviter stipitata, petalis vix breviora. Ovarium glab-

rum ovulis biseriatim dispositis praeditum.

BORNEO: in regione Centrali in provincia malaice Kutai

dicitur, in saxis cretaceis, circa 30 m. alt. (Endert 5,134—Holo-

typus in SING.).

The collector notes that the shrub is 50-75 cm. tall, producing

dark red flowers and growing on the edge of limestone rockface.

The calyx is rather long and deeply divided, but such long calyces

are known to occur in the other sections of the genus. The flowers

are in a bad state to admit a further scrutiny. No fruits are avail-

able. Because the flowers are borne at the apices of special branches

and because the leaves are subverticillate, the species belongs

obviously to the Section PyrGus.

17. Ardisia calophylla Furtado sp. nov.

A. littoralis Andr. sensu Ridl., Fl. Mal. Pen. II (1923) 241 in

obs.

Ab A. kurzii, cui affinissima, foliis minoribus, coriaceis, obovatis

cum nervis reticulationibusque obscuris; inflorescentiis paniculatis,

quam folia paulo brevioribus; sepalis suborbicularibus margine

ciliolatis, sicut petala punctulatis; antheris acutis dorso glandulas

paucas distantes gerentibus sat distincta.

Ramuli glabri, in sicco sicut folia foenicolores. Folia glabra,

punctis manifestis destituta, obovato-elliptica, apice arcuatim acuta

vel breviter acuminata, 6-10 cm. longa, 3-4 cm. lata, basin versus

298

Vol. XVII. (1958).

sensim cuneata, imo paulo decurrentia, petiolis supra sulcatis 5-10

mm. longis stipitata, nervis reticulationibusque fere obscuris prae-

dita. Inflorescentia glabra, ramuli foliis subverticillatim praediti

apice sita, paniculata, in ramulos floriferos valde divisa, ad 6 cm. us-

que longa. Flores ante anthesin conoidei; apice rotundati 3-4 mm.

longi; pedicellis 3-5 mm. longis. Sepala basi 1/3 coalita, valde

dextrorsum imbricata, fere orbicularia, glabra, dorso glandulosa,

margine lutescentia, scarioso-ciliolata. Petala calyce subtripulo

longiora, glandulosa. Stamina petalis paulo breviora, acuta, dorso

pauciglandulosa. Ovarium ovoideum, glabrum.

MALAYA: Dindings, Bruas (Burn-Murdoch in 1909—Herb.

SING. ).

The collector notes that the flowers are white.

sem

4mm 7

Ardisia calophylla Furtado (Burn-Murdoch-Holoty pus)

A, Habitus. B, Flos per anthesin. C, Stamen postice visum. D, Ovarium.

E, Placenta.

299

Gardens Bulletin, S.

18. Ardisia purseglovei Furtado sp. nov.

A. candolleanae valde affinis, a qua foliis ovoideo oblongis basi

pseudo-cordulatis, subsessilibus, 2labris, margine obscure crenu-

latis, glandulas magnas mox deciduas unocuique sinu singulas

gerentibus, subtus minutissime punctulatis, cum nervis lateralibus

reticulationibusque obscuris; sepalis tetrameris vel pentameris,

extus tomentosis, intus glabris, margine ciliatis, eglandulosis; ova-

riis tomentellis facile distinguitur.

Frutex circa 3 m. altus, cum ramulis spiculiferentibus 5-10 cm.

longis. Folia coriacea vel chartacea, ovoideo oblonga, apicem ver-

sus arcuatim angustata, summo obtusa, basi rotundato truncata,

imo ob rugas pseudo-cordulata, petiolis brevissimis ad 3 mm. usque

longis stipitata, in ramulis spicas gerentibus subverticillata, mar-

gine obscure crenulata et revoluta, in unocuique sinu glandulas

conspicuas deciduas singulas ferentia, nervis reticulationibusque

obscurissimis praedita, subtus punctulis minutissimis dense aucta,

8—15 cm. longa, 2:5—4-5 cm. lata, utrinque glabra, novella subtus

deciduo rubro-lepidota. Inflorescentia apicalis omnino tomentella,

€X axi primario valde abbreviato, circa 2 cm. longo, in ramos circa

4, paulo ramulosos divisa. Flores in ramulis inflorescentiae pri-

mariis vel secundariis producti, tetrameri vel pentameri, pedicellis

puberulis circa 5 mm. longis suffulti. Sepala coriacea, basi 4—1/3-

coalita, ovato triangularia apice obtusa vel subacuta, intus glabra,

extus tomentosa, punctulis manifestioribus destituta, margine cilio-

lata. Petala calyce fere duplo longiora deciduo lepidotula, punctata.

Ovarium pubescens, ovoideum, cum stylo tortuoso.

BORNEO: Sarawak, Gunong Pueh, alt. + 150 m. (Purseglove

4670—holotypus in Herb. SING.). Specimina isoholotypica in K.

et L. distributa non vidi.

The species is readily recognised by the ovate oblong, subsessile,

pseudocordulate leaves having practically no reticulations and its

lateral nerves inconspicuous, the inflorescence being hairy (the

hairs are capitate), and the ovary puberulous. The leaf-margins

are somewhat crenulate, but as the margins are revolute, the crenu-

lations are not easily seen especially if the leaves are mounted with

their lower surface glued onto the sheet. A. brachythyrsa has also

subsessile leaves, but in that the leaves are much more rigid and

coriaceous, almost obovate, having their nerves and reticulations

quite conspicuous on both surfaces. The sepals are smaller there

and less tomentose and the ovary is quite glabrous.

300

Vol. XVII. (1958).

Ardisia purseglovei Furtado (Purseglove 4,670—Holotypus)

A, Habitus. B, Flos post anthesin. C, Flos petalis desumptis. D, Stamen

postice visum. E, Ovarium. F, Placenta.

301

Gardens Bulletin, S.

As the leaves are crenulate and bear a conspicuous gland in

each sinus, the species may be easily put in the Section CRISPAR-

DISIA as defined by Mez, but the mode of flowering and subverti-

cillate leaves have led me to place species in the Section PyRGus.

19. Ardisia vaughanii Ridl., Fl. Mal. Pen. II (1923) 248.

MALAYA: Perlis, Telor Jambu (Ridley 14,932, distributed as

A. sanguinolenta Wall.?). Kedah, Gunong Baling (Kiah 35,383

& sm. on 11—V—1938; Best 21,226; Nauen 38,022). Perak,

Temengoh (Schebesta 2). Pahang, Kuala Tembeling (Holttum

20,552 & 24,723; Henderson 21,780).

The type was from Kelantan. Ridley placed this species in the

Section TINOPSIS, but it should be placed in the Section PyrGus

(Lour.) Mez, the species being an ally of A. kurzii Clarke. Squires

848 from Anam, Indochina, distributed as A. rigida Kurz comes

very close to the Malayan species.

According to the notes of the collectors, this is a shrub 6—8 ft.

tall, bearing clusters of flowers having pale pink petals with dark

centre, deep pink peduncles and pedicels, and red fruits which

eventually turn black.

The following notes may be of use to supplement the original

description: | |

Leaves subverticillate on the main stem, with many small glands

inconspicuous in some leaves, .but generally quite conspicuous on

both surfaces, glabrous. Inflorescences terminal usually borne on

short lateral branches. Peduncles and pedicels deciduously puberul-

ous and lepidote, somewhat ferrugineous. Sepals ovate triangular

with rounded apex, 1/3 united at base, imbricate in buds, glan-

dular and minutely puberulous on the dorsum, ciliate in margins.

Petals with many glands, gradually acute. Stamens nearly as long as

the petals, acute or mucronulate at apex, provided on the dorsum

with no or few glands which are not black. Ovary glabrous. Fruit

black when dry, longitudinally ridged.

Section BLADHIA (Thunb.) Mez

20. Ardisia foliosa Furtado sp. nov.

A. linearifolia Ridl. in Journ. Roy. Asiatic. Soc. Str. Br. 61

(1912) 27 et Fl. Mal. Pen. II (1923) 255, non A. linearifolia

Miq. (1860); Syn. nov.

Suffrutex parte erecta 20-40 cm. longus, in parte basali repens

vel subterranea ramulos erectos gerens, cum innovationibus minute

puberulis. Folia subverticillatim congesta, membranacea, costis —

medianis exceptis minute puberulis utrinque glabra, lineari lanceo-

lata 10-12 cm. longa, 15-20 mm. lata, utrinque sensim acuta,

302

Vol. XVII. (1958).

; J

3mm @) , ©

3mm 3mm

Ardisia foliosa Furtado (A-E, Holotypus; F-J, Merotypus A. linearifoliae

Ridl.)

A, Habitus. B, Flos post anthesin apice pedunculi situs. C & H, Flos

petalis desumptis. D & I, Stamen postice visum. E & J, Placenta.

F, Folium. G, Flos post anthesin.

apice plerumque mucronulata, margine valde denticulata et revo-

luta, supra brunnea, subtus pallidiora, nervis secundariis utrinsecus

16-20, reticulationibus remotis subtus prominulis supra obscuris

praedita, glandulis novellis obscuris dein prominulis praecipue

secus marginem et apice punctata, petiolis 10-15 mm. longis sti-

pitata. Inflorescentiae axillares, simplices vel pauciramosae, axi

303

Gardens Bulletin, S.

minute puberulae, cum pedunculis gracilibus bracteolatis, apice

3—8 flores congestis, infra apicem flores solitarios vel etiam ramulos

florigerentes productis; pedicellis minute puberulis vel glabres-

centibus 8-15 mm. longis. Flores 3-4 mm. longi, ante anthesin

acuti. Sepala basi 1/3-unita, ovata, acuta, margine ciliolata,

albescentia, dorso pauciglandulosa vel non. Petala ovato acumi-

nata, interdum subito apiculata. Stamina petalis multo breviora.

Baccae non visae.

MALAYA: Kelantan (Henderson 29,656—Holotypus). Johore,

Sungei Kayu (Kiah 23,330). Perak, Gunong Bongsu (Haniff,

Merotypus A. linearifoliae). Omnia specimina in Herb. Sing.

conservata.

A. linearifolia Ridl. is an impriorable name since it is a later

homonym. This and the fact that the species was based on a frag-

mentary material cultivated in the Botanic Gardens, Singapore,

have induced me to establish a new species based on better material

the duplicates of which have been widely distributed. In accord-

ance with the definition given previously (Furtado in Gard. Bull.

Straits Settl. IX, 1937 p. 289) I have employed the term merotype

to designate the specimen taken from the type plant on which A.

linearifolia Ridl. was based. In the plate figs. F-J are from the

fragmentary merotype material preserved in herb. Singapore.

21. Ardisia tumida Furtado sp. nov.

Ab A. metallica, cui affinissima ut videtur, caule haud pubes-

centi, ramulis novellis deciduo ferrugineo-furfuraceis, foliis mar-

gine eserratis undulatisque, petiolis subtus valde tumidis haec

species sat distincta.

Suffrutex circa 60—100 cm. longus, lignosus, in vivo cortice valde

_ succosus, in sicco irregulariter angulatus, ramulis novellis axi deci-

duo ferrugineo-furfuraceis, 15-30 cm. longis praeditus. Folia

saepe subverticillatim collata, petiolis 1-2 cm. longis, supra paulo

alatis, sulcatis, subtus valde tumidis stipitata, obovato-elliptica

margine undulata, membranacea, pellucida, utrinque glabra, basin

versus paullatim angustata, imo sub-acuta, apice eleganter acumi-

nata, 15—20 cm. longa, 4:5—7 cm. lata, nervis primariis utrinsecus

14-20 praedita, prominulo reticulata, glandulis minutis pluribus

omnino dispersa. Inflorescentia stricte axillaris parva cum pedun-

culo 5—10 cm. longo, simplici vel bifurcato, 3—5 fructiferenti, pedi-

cellis fructiferis gracilibus, circa 10 mm. longis. Bacca globosa,

6—7 mm. in diam., purpureo-glandulosa; sepalis 1/3-coalitis, car-

nosis, glabris, obscure glandulosis vel non.

304

Vol. XVII. (1958).

Ardisia tumida Furtado (Corner 30,290, holotypus)

A, Fragmentum caulis cum fructu. B, Fragmentum caulis ut petiolus

subtus tumidus appareat.

305

Gardens Bulletin, S.

MALAYA: Kemaman in locis juxta flumina paludosis: Ulu

Bendong (Corner 30,290—Holotypus in SING.): Sungei Nipah

(Corner s.n. in Nov. 1935—SING.); Ulu Kajang (Corner s.n.

in Nov. 1935—SING.).

The species is readily distinguished by its peculiar, water-holding

cortex on the underside of the petioles, being very pronounced in

older leaves. Similar cortex is found all over the stem, a reason

why it dries to produce irregular ridges. Young twigs are covered

with fugaceous, rusty-furfur- Leaf margins are undulate, but not

serrate.

The collector notes that the plant is 3 ft. tall with pale red ber-

ries. No flowering specimen has been available.

HYMENANDRA A.D.C.

Hymenandra iteophylla (Ridl.) Furtado comb. nov.

Ardisia iteophylla Ridl. in Journ. Bot. LXII (1924) 298 et FI.

Malay Pen. V Suppl. (1924) 318: basinymus.

That the stamens are connate into a tube was noticed by Ridley

himself; they have also a small free tip. In Ardisia the stamens are

free.

This is a first record of the occurrence of the genus not only in

Malaya, but also outside Assam.

TETRARDISIA Mez

Tetrardisia corneri Furtado sp. nov.

A T. denticulata (Bl.) Mez, cui affinissima, sed foliis majoribus,

obovatis vel oblongis, inter venae nonnihil bullatis, inflorescentiis

longioribus facile distinguitur.

Suffrutex circa 60 cm. altus, lignosus. Folia obovato-elliptica vel

oblonga, basin versus sensim attenuata vel subito angustata, imo

acuta, apice acuminata, margine crenata, reticulationibus densis

utrinque prominulis praedita, nervis primariis utrinsecus 12—20

percursa, inter nervos bullata, 15—20 cm. longa, 45-8 cm. lata,

punctulis pluribus, atrescentibus praedita. Inflorescentiae termi-

nales vel axillares, 5-6 cm. longae, pedunculis gracilibus, apice

flores distantes gerentibus, vel etiam in 1—2 ramulos circa 1—2 cm.

longos florigerentos divisae; pedicellis 8-10 mm. longis, minute

puberulis. Flores tetrameri, ante anthesin acuti, 6-8 mm. longi.

Sepala imbricata, basi 1/3-unita, ovata apice rotundato-acuta,

dorso minute glandulosa, in fructu reflexa. Petala calyce fere

quadruplo longiora, apicem versus sensim acuta. Anthera petalis

vix breviora, apicem versus sensim attenuata, dorso atropunctata,

306

Vol. XVII. (1958).

ut tubus staminem appareat. F, Ovariu

307

Vol. XVII. (1958).

longitudinaliter imtrorsum et per foramina apicalia dehiscentia.

Ovarium ante anthesin unicum visum, imperfectum; stylus fili-

formis, paulo tortuosus, stigmate punctiformi praeditus. Bacca glo-

bosa, circa 6 mm. in diam., novella glandulis pluribus conspicuis-

que, in maturitate fere inconspicuis et sulcis verticalibus praedita.

MALAYA: Johore in locis paludosis: Sungei Berassau prope

viam Mawai—Jemaluang (Corner sn.—Holotypus); Mawai (Cor-

ner sn.); Gunong Panti ad basin (Ridley—Dec. 8, 1892 )—-Omnia

specimina in Herb. Sing. conservata.

The growth of inflorescence is somewhat similar to that in T.

tetrasepala and the anthers appear to dehisce through both the

apical pores and the longitudinal slits. Only one unopened flower

was available (besides one opened one) and this had decayed

somewhat. What looks like ovules in the ovary were really cavities.

One specimen (Mawai) had some inflorescences transformed

into small plants, though its apical inflorescence bore fruits.

Probably Alvins 2336 from Malacca is this species, but the label

does not belong here, for according to Alvins’ notes the plant is a

tree about 30 ft. tall with very hard, dark red wood. The leaves

are longer, lanceolate elliptic or ovate elliptic, gradually narrowed

to produce an acuminate apex and sub-rotund base. The specimen

has scars of fallen peduncles.

Ridley’s specimen from a herb grown in the Botanic Gardens,

Singapore, in 1892 and figured in colours by Alwis in a plate pre-

served in the Singapore herbarium should also be assigned here,

though being from a very young plant the leaves are smaller with

shorter petioles. The drawing shows different shades of pink or

mauve in the pedicels, calyx and corolla. Ridley suspected the

plant to have come from Pahang.

Tetrardisia porosa (Clarke) Furtado comb. nov.

Ardisia porosa Clarke in Hook. f., Fl. Brit. Ind. III (1882) 522;

Mez in Engl. Pflanzenr. IV 236 — Heft 9 (1902) 118; King &

Gamble, Mat. Fl. Malay. Pen. IV (1905) 132; Ridl., Fl. Mal.

Pen. II (1923) 248 fig. 97. Basinymus.

MALAYA: Perak, Taiping Hills (Henderson 10,454 & 10,438;

Ridley 14,262); Gunong Hijau (Ridley in February 1891); Batu

Togo (Wray 2,160); Larut (Scortechini 1167); Ayer Larut (Wray

39); Tupai (Wray 2,835). Malacca, Panchor (Goodenough)

1528). Negri Sembilan (Ridley in December 1890).

It is possible that the pollen shedding through apical pores of the

anthers is also a characteristic of the genus Tetrardisia.

309

Gardens Bulletin, S.

Tetrardisia tetrasepala (King et Gamble) Furtado comb. nov.

Ardisia tetrasepala K. & G. Mat. Fl. Malay. Pen. IV (1905)

352; Ridl. Fl. Mal. Pen. II (1923) 244—basinymus.

Ad descriptionem originalem adde:

Flores circa 10 mm. longi, ante anthesim valde acuti, tetrameri,

8—20 subumbellati, acropetales, in ramulo axillari vel apicali in

singulos annos pedunculiformiter 3-20 mm. aucto siti; pedicelli

graciles, 8-15 mm. longi. Sepala dextrorsum imbricata, ovata,

apice rotundata, margine albescente nuda vel ciliolata, dorso

minute punctulata, in fructu reflexa. Petala calyce fere triplo

longiora, lanceolata, apice sensim acuta, punctulis parvis multis

prope marginem picta. Anthera breviter stipitata, petalis vix bre-

viora, apice sensim acutissima, dorso bene punctulata, introrsum

longitudinaliter et. per foramina apicalia dehiscentia. Ovarium

glabrum vel minuti puberulum, cum ovulis paucis uniseriatim dis-

positis, stylo gracillimo, nonnihil spirali, petalis subaequante; stig-

mate punctiformi.

MALAYA: Johore, Gunong Pulai (Nur & Kiah 7,805—

Apotypus; Ridley sn. & 12,193: Omnia specimina in Herb. SING.).

The inflorescence may be termed sometimes axillary and some-

times terminal. But what appears to be peduncles are really the

current year growth of the axillary or terminal buds or twigs.

When flowers and fruits are shed, the “peduncles” will grow again

to produce flowers in the next season without producing any leaves.

Later the same will produce one or more leaves before flowering

again.

The tissues connected with the leaves produce a ridge on the

stem from node to node and as the petiole of the terminal leaf

grows much thicker than the main axis, the terminal inflorescence

appears nearly axillary. This mode of growth also tends to make

the stem zig-zag. Vestiges of fallen flowers may be seen on stem.

The anthers shed pollen both through apical pores and through

longitudinal slits. The leaf margins are undulate and sometimes

slightly crenate, and the petiole is often transversely wrinkled, the

wrinkles acquiring dark brown colour.

Tetrardisia was not recorded to occur outside Java. King &

Gamble had noted the tetramerous calyx, but they had no fiowers

to define the species generically. In the Singapore herbarium, of

the three collections from the type locality in Johore (Gunong

Pulai) only Nur and Kiah 7,805 have flowers which have served

to place the species definitely into this genus. Hence it may be

called the apotype. (cf. Furtado in Gard. Bull. Straits Settl. IX

(1937) 288).

310

Vol. XVII. (1958).

578 CG

¢ oe

Ut Rat g iy i, Wh Uy i

\ QO

VEZ “s

Tetrardisia tetrasepala (K. & G.) Furtado (A—G, Nur & Kiah 7,805, Apo-

typus; H, Ridley 12,193)

A, Habitus. A—a, Vestigia florum prioribus annis delapsorum. B, Flors

per anthesin. C, Calyx cum pedicello. D, Stamen postice visum. F,

Ovarium. G, Placenta. H, Umbella fructifera.

sid

The Genus Sphagnum in Malaysia

By ANNE JOHNSON

Department of Botany, University of Malaya,

Singapore

IN THE LARGE GENUS, Sphagnum, the degree of affinity and vari-

ability of the species occurring in South-East Asia has not been

fully worked out. Professor Le Roy Andrews has made very

valuable investigations and has indicated the synonymy for South

and North American species, and has made some reference to

species from other tropical regions. Fortunately he was able to

examine the types in the Warnstorf Moss Herbarium at Berlin-

Dahlem. This herbarium which was entirely destroyed in the last

war (Dalby, 1957), included the types described by Warnstorf in

Sphagnologia Universalis (1911). In the absence of types for re-

ference, the excellent critical work done by Le Roy Andrews before

the destruction of the herbarium must form the basis of present

day Sphagnum studies.

In this paper an account is given of the Malaysian Sphagna in

the Herbaria at Bogor, Indonesia, and at Singapore.

The classification of the genus in this paper closely follows that

of Andrews (1936). A complete synonymy is not given but new

synonyms are indicated. Other synonyms can be found in

Warnstorf (1911).

Key to the Species

la. Cortical cells of stem and branches without spiral thickenings;

some, at least retort-shaped

Subgen: LITOPHLOEA ................. 2:

2. Stem cortex cells all alike and all somewhat retort-shaped,

branch leaves lacking a distinct border but with a resorp-

tion furrow. fee ence’ Sect. MALACOSPHAGNUM

1. Sphagnum antarcticum.

2. Stem cortex cells of two kinds, with retort cells only in the

axils of leaves; branch leaves all bordered with one or

more rows of narrow cells

Sect. ACISPHAGNUDM ................... 3:

3. Small delicate plant, shining and iridescent when dry.

Branch leaves not fibrillose ...... 7. Sphagnum sericeum.

312

Vol. XVII. (1958).

3. Not iridescent. Fibres present in branch leaves. ............ 4.

4. Stem leaves lingulate to triangular lingulate with

rounded fimbriate, laciniate or eroded apex ...... 5.

5. Stem leaves not fibrillose hyaline cells never

oe Ql op ae 3. Sphagnum cuspidatulum.

5. Stem leaves usually fibrillose in the centre, hya-

line cells septate

4. Sphagnum novo-guineense.

4. Stem leaves triangular to triangular-ovate with a

Se oli ARR coe 6.

6. Chlorophyll cells lying centrally, exposed on

both sides or exposed only on the dorsal

side. Branch leaves not squarrose ............... q.

7. Border of stem leaves broadened below

2. Sphagnum cuspidatum.

7. Border not broadened below ......0........-.:000+- 8.

8. Delicate aquatic plant, branch leaves

at least 3-0 mm. long

6. Sphagnum flaccidifolium.

8. Fairly robust species, branch leaves

less than 1-5 mm. long.

6. Sphagnum subrecurvum.

6. Chlorophyll cells lying exposed only on the

ventral side. Dry tip of branch leaves

squarrose ...... 8. Sphagnum junghuhnianum.

1b. Cortical cells of stem and branches with spiral thickenings, not

retort-shaped o......ssssscsee Subgen: INOPHLOEA ................. 2.

2. Stem leaves very small, ovate or ovoid, about 500,

oO CEE a ee 9. Sphagnum roseotinctum.

2. Stem leaves at least 1:2 mm. long, lingulate to trian-

Sahai angen! noe ah “eile alll Bee i ad set Mes ae 3.

3. Hyaline walls adjacent to chlorophyll cells in branch

leaves densely papillose

12. Sphagnum borneense.

3. Hyaline cells not papillose

4. Branch leaves with dorsal pseudopores

11. Sphagnum beccarii.

4. No dorsal pseudopores ......... 10. Sphagnum holttumii.

A. Subgenus Litophloea (Russ.) Andrews (1911)

313

Gardens Bulletin, S.

I. Section MALACOSPHAGNUM C. Mill.

1, Sphagnum antarcticum Mitt. in Jour. Linn. Soc. IV (1859)

106; Bartr., (1942) 246.

Robust plant. Main stem cortex 2—4 cells thick, outer cells all

more or less retort-shaped; wood cylinder distinct, solid, yellow to

brown. Stem leaves triangular-lingulate to lingulate, 1-0—2-0 mm.

long 0-7—1-:1 mm. broad, apex rounded, eroso-fimbriate. Branch

leaves broadly ovate, about 25 x 1-4 mm., strongly imbricate,

chlorophyll cells central, barrel-shaped in section, included between

hyaline cells or free, hyaline cell-walls adjacent to chlorophyll cells

smooth or papillose.

NEw GuINEA: Mount Wilhelmina, wet N. slope, burnt sub-alpine

forest 3,400 m. (Brass & Myer-Drees 9737/BO.).

DISTRIBUTION: Australia, New Zealand, Stewart Island, Campbell

Island.

This species, while typically a member of the small Malacos-

phagnum (Rigida) group differs from S. compactum D.C. in the

relatively large stem leaves with their broad rounded, constantly

fimbriate apex. The branch leaves have numerous dorsal pores at

the commissures while the pseudopores, typical of S. compactum,

are absent. The section of the branch leaves is similar to that of

S. compactum with the chlorophyll cells included on both sides, in

which feature it differs from S. strictum Sull. where the chlorophyll

cells are exposed on the outer surface of the leaf.

II. Section ACISPHAGNUM C. Miill. emend Andrews (1911).

Series Cuspidata Schlieph

Chlorophyll cells of branch leaves exposed dorsally

or central in position.

2. Sphagnum cuspidatum Ehrh. in Hoffm., Deutsch. Fl. 2 (1796)

22 Bartr., (1942) 246.

Delicate to fairly robust, often aquatic, pale green to pale brown.

Cortex 2—3 cells thick; wood cylinder distinct. Stem leaves tri-

angular-ovate; apex slightly truncate, dentate not lacerate; with a

strong border considerably broadened below; hyaline cells fibril-

lose. Branch leaves long lanceolate, involute; apex toothed; margin

entire with a border about three cells thick; hyaline cells fibrillose

with strongly ringed pores at ends and corners of cells (dorsal) ;

chlorophyll cells of branch leaves trapezoid in section with broader

exposure on outer side.

SUMATRA: Si Garang Garang, wet place in forest, 1,400 m. (Polak

154/BO.); Polok Magae (Polak 156/BO.); Si Gelapang, 1,400 m.

(Polak 157/BO.); Benkoelen Sikintjan Belirang, 1,400 m. (Rappard

314

Vol. XVII. (1958).

98/BO.); Gunong Koerintiju 1,800 m. (Brunnemeyer 9602/BO and

1079/BO.); Petanei Valley, 1,300 m. (Lorzing 8487/BO.); Tongging,

Toba Lake (Lorzing 8072/BO.); Tapianoeli, 1,400 m. (Polak 154/BO.);

Gunong Goh Lemboeh, Top Plateau, 3,000 m. (van Steenis 10171/BO).

Java: Gunong Dieng (Rant Bogor-1656/BO.); loc. incert. (Moh

Enoh 374/BO.).

BoRNEO: Western Division Anjongan, North of Pontianak, sandy

ground (Polak 157/BO.).

New Guinea: Mount Wilhelmina, very wet place in bog, 3,450 m.

(Brass & Myer-Drees 9758/BO.); Lake Habbema, common in bogs

and lining edges of pools, 3,225 m. (Brass 9448/BO.).

DISTRIBUTION: North America, Europe, Asia.

The following numbers labelled in the Bogor herbarium as

S. subrecurvum are definitely not this species. The stem leaves

have a border strongly broadened below. I find no reason for not

including them in S. cuspidatum:—

Brunnemeyer 9602; Rant (Bogor No. 1656); Toxopeus no number;

Lorzing 8487; Lorzing 8072; Brunnemeyer 10479; Polak 154.

3. Sphagnum cuspidatulum C. Mull. in Linnaea 38 (1874) 156.

Fairly robust. Main stem cortex 2—3 cells thick; wood cylinder

not distinct. Stem leaves triangular-lingulate to lingulate, not fibril-

lose; apex rounded laciniate; border narrow, not broadened below.

Branch leaves in five rows, ovate lanceolate with a very narrow

border; apex truncate dentate; hyaline cells with numerous pores

on the dorsal side; chlorophyll cells triangular in section, lying

dorsally.

CELEBES: Enrekang, Sawito, 1,800 m. (De Jong 108/BO.); Menado-

Possomeer, 2,000 m. (Boschproefstation 13/BO.).

DISTRIBUTION: India, Siam, Malaysia.

3 (a) var. malaccense (Warnst.) Warnst. (1911) 187; Dixon (1926),

2 (misprinted as S$. cuspidatum var.).

Sphagnum malaccense Warnst. in Hedwigia 31 (1892) 175.

Stricking pale brown colour when fresh. Stem leaves triangular-

lingulate to spathulate 1:3-1-6 x 1:0 mm., not fibrillose; apex

rounded, fimbriate; border indistinct. Branch leaves lanceolate

about 1:3 0:5 with a border two to five cells thick. Apex nar-

rowly truncate and split into four teeth.

CELEBES: Mamboeling by Mamasa (De Froideville 159/BO).

Matava: Kedah, Kedah Peak, common. + 914 m., apparently com-

monest species (Holttum 14881/SING.) Perak: Gunong Batu Puteh,

+ 2,100 m. (Wray 902, type, SING.). Pahang: Cameron Highlands

(Henderson 17877b/SING.); + 1368 m. (Holttum 23301/SING.);

Gunong Batu Brinchang, abundant in forest near summit (Holttum

23530/SING.. & BO.). 7

This variety is distinguished by its broad tongue-shaped or spa-

thulate stem leaves; which may be fimbriate, not only on the

315

: Gardens Bulletin, S.

rounded apex, but also on the side margin. However this last is not

a very constant feature.

3 (b) var. trengganuense Johnson var. nov.

Ab omnibus hujus speciei varietatibus hactenus cognitis caulibus

flavo-bruneis, foliis caulinis partim fibrosis hoc taxon facile distin-

guitur.

Planta robusta, cum caule flavo-bruneo. Folia caulina triangulo-

lingulata, partim fibrosa. Folia ramea plerumque 2 mm. longa, 0-6

mm. lata.

This variety differs from all the others hitherto described in the

golden-brown stem and partly fibrous stem leaves. The plant is

rather robust producing triangular-lingulate stem leaves. Its branch

leaves are 2 mm. long, 0-6 mm. broad.

MaLaya: Trengganu: Gunong Padang, 4,000 ft. (Moysey & Kiah

31023, Holotypus in SING).

4. Sphagnum novo-guineense Fleischer & Warnst. in Warnst.

(1914)5520.

Robust plant. Main stem cortex 2-3 cells thick distinct from the

wood cylinder. Stem leaves triangular lingulate, about 1:3 « 1-0

mm.; apex rounded, a little eroded or eaten away; narrow border

scarcely broadened below. Hyaline cells often septate and fibrillose

in the centre of the leaf. Branch leaves broadly ovate-lanceolate;

apex narrowly truncate, dentate, about 1-6 & 0-8 mm., with narrow

border, pseudopores present on dorsal side; chlorophyll cells tri-

angular in section exposed dorsally.

New GuINEA: Mt. Nellivig (Pulle 832/BO.); Mt. Goliath (De Koch

16/BO.).

5. Sphagnum flaccidifolium Dixon Ms. ex Johnson sp. nov.

A Sphagnum cuspidato, cui proxima, ramis flaccidis, foliis cau-

linis triangulis utrinque recto-marginatis (haud arcuatiin margi-

natis), foliis rameis multo longioribus, semper anguste limbatis,

cellulas chlorophyllosas in sectione eorum transversali liberas utrin-

que ferentibus haec species sat diversa.

Planta aquatilis. Caulis circa 1 mm. in diam., axi lignosus, epi-

derme 2-3 seriatim strata, manifeste diversa vestitus. Folia caulina

magna, triangularia, recto-marginata, multifibrosa, circa 1-6 mm.

longa, 1 mm. lata, apice cucullata et saepe bidentata, semper anguste

uniformiterque limbata. Folia ramea perlonga, anguste lanceolata,

3—5 mm. longa, circa 0-5 mm. lata, anguste limbata, integerrima,

apice truncata et 2-4 dentata, dorso poris interdum annulatis in

cellularum medio instructa, in sectione transversali cellulis chloro-

phyllosis rectangulis vel trapezoideis liberis utrinque disposita.

316

Vol. XVII. (1958).

An aquatic plant. Stem 1 mm. diameter, woody in the axis,

covered with 2-3 layered, well differentiated epidermis. Stem

leaves large, triangular with straight sides, multifibrous about 1-6

mm. long, 1 mm. wide, cucullate and often bidentate at the apex,

and always with a uniform narrow border. Branch leaves very long,

narrowly lanceolate 3-5 mm. long, about 0-6 mm. wide, entire,

having each a narrow border and a truncate 2—4 dentate apex, and

provided dorsally in the middle of the cells with pores which are

sometimes thickened with rings; rectangular to trapezoid chloro-

phyll cells being seen exposed equally on both sides in the trans-

verse section of the branch leaves.

Malaya: Selangor: Telok Forest Reserve, 7th Mile stone, Klang

(Carrick 500, Holotypus in SING.); Telok Panglima Garang (Holttum

28317/SING. & BO.).

SuMaATRA: Near Pakan Borae (Holttum 28127/SING.).

The specific epithet was proposed on herbarium sheets by H. N.

Dixon but, as far as I am aware, it has not been validly published

before.

This species is remarkable in its flaccid branches and exceed-

ingly long branch leaves. It is the only Malayan species found in

the lowlands.

6. Sphagnum subrecurvum Warnst. in Allgem. Bot. Zeitschr.

(1895) 134.

Fairly robust. Main stem cortex 2-3 cells thick with fairly distinct

wood cylinder. Stem leaves ovate-triangular, 1:00 & 05 mm. to

triangular 1:6 X 0-8 mm., all fibrillose, border present of equal

width above and below. Branch leaves lanceolate, about 1:3 « 0-5

mm., apex narrowly truncate, dentate; with equal narrow border;

pores minute, usually single; chlorophyll cells of branch leaves

triangular in section, exposed only on dorsal side.

SUMATRA: Tapianoeli, Lantong Nihoeda (Polak 155/BO.); Dolak

Margaoe (Polak 156/BO.); Si Gelapang (Polak 157/BO.); Danam

Biloeloek (Jacobson 110/BO.); Hoetagindjang, Toba Lake (Ruttner in

April 1929/BO.).

Java: Telaga Warna, 2,000 m. (van Steenis 4495 & 4545/BO.);

Telaga Loemoe, (Bogor 2747/BO.); Telaga Tjibong, + 2,000 m. (van

Steenis 4495/BO.); loc. incert. (Scroter, May 1927/BO.).

7, Sphagnum sericeum C. Mill. in Bot. Zeit. (1847) 481; Bartr.,

(1942) 246.

Small delicate plant, shining, iridescent when dry. Main stem

cortex 2-3 cells thick; wood cylinder distinct, yellow to brown

in colour. Stem leaves triangular, apex acute; narrow, finely

toothed border; 1:1 X 0:7 mm.; no fibres. Branch leaves ovate to

317

Gardens Bulletin, S.

oblong-lanceolate; 1:1 x 0-4 mm., acute apex, serrulate border,

no fibres, chlorophyll cells trapezoid in section, both sides exposed,

longest side ventral.

Java: Tjibeureum (Fleischer s.n./BO.); alt. 1,700 m. near Waterfalls

(Meyer B-3,447/BO. & B-1,099/BO.; van Steenis 2,136/BO.); Tiji-

kundul valley, alt. 1,700 m. on rocks (Meijer B—3,826/BO.); Gunong

Lalak 2,000—2,200 m. (van Steenis's.n. in VI-1941/BO.).

NEw GuInEa: Bernard Camp by Idenburg river, alt. 2,100 m. (Brass

12175/BO.).

BORNEO: West Koetai, Kemoel, alt, 1,700 m. (Endert 4530/BO.).

MataYa: Pahang: Gunong Tahan, alt. 1,500 m. by stream (Holttum

20911/SING.); Cameron Highlands (Holttum 23530/SING.).

DISTRIBUTION: Malaysia.

A very distinct species easily recognized by its delicate nature,

the shining iridescence of the leaves when dry, and the complete

absence of fibres. Andrew regards the loss of fibres as secondary;

their partial loss being exhibited in aquatic forms of the cuspidatum

and subsecundum type. There is no justification for the creation

of a separate section (Sericea Warnst., 1911) for the two diverse

species—S. sericeum and S. macrophyllum, merely because they

resemble one another in their complete absence of fibres.

Series Acutifolia

Chlorophyll cells of branch leaves exposed ventrally.

8. Sphagnum junghuhnianum Dz. et Molkenb., in Verhandel. d.

Kon. Akad Wetensch. Amsterdam (1854); Bryol, jav. 1 27.

Bartr. (1946) 246.

S. gedeanum Dz. et Molkenb. in op. cit. (1854); Bryol. jav.

i. 2s.

Most robust, usually not shining when dry, often tinged with

pink, pale red or violet. Main stem cortex 2-3 cells thick, distinct

from the woody cylinder. Stem leaves triangular about 1-8 x 0-7—

1:3 mm., auricles present or absent, not fibrillose or few to numer-

ous fibrils in hyaline cells, border indistinct or distinct, not broad-

ened below; tip truncate, toothed, somewhat squarrose. Branch

leaves lanceolate to ovate lanceolate, never in five rows, dry tips

squarrose; 1:5—2:0 x 0-4-0°8 mm., fibres few or many; border

very distinct; dorsal pores numerous near commissures; chlorophyll

cells triangular in section, exposed ventrally.

SUMATRA: Penghoeloe Bao, 2,000 m. (Frey-Wyssling 41/BO.); Gan-

jelanden, Gunong Goh Lemboeh, 2,700 m., in mossy forest (van Steenis

10223/BO.).

Java: Tjibeureum, 1,700 m. wet rocky walls and waterfall (Meijer

3448/BO.); 1,700 m., on andesite rocks (van Steenis 1156/BO.); 1,800

m. (Fleischer 501/BO.); Tjibodas (Massart 1262/BO.); Tjipanas

318

Vol. XVII. (1958).

(Fleischer in Aug. 1900/BO.); Gunong Patoeha, near Kawah Poetih,

under trees, wet semi-shady places. (Verdoorn, Series 1 in 1930/BO.);

2,200—2,300 m. (van Steenis 4429, 6920 & 6921/BO.); Gunong Papan-

dajan, Tegal, grassland 2,350 m., wet valley of small river (van Steenis

4291/BO.); rock valley of small river, 2,350 m. (van _ Steenis

4806/BO.); Priangan, Tjiparoegpoeg 2,350 m., valley of small river

(van Steenis 4806a/BO.); Gunong Oengaran, 2,050 m. (van Leeuwen

1248/BO.); Gunong Ardjoemo, 2,600 m. (Bremekamp/BO.); Air Panas

on slope by hot springs (Meijer B-3999/BO.).

CELEBES: Gunong Ngilataki, East of Lindoe Lake, 2,355 m. (Bloem-

bergen 3947/BO.); Gunong Bothian (Brunnemeyer 11968/BO.);

Porema, 1,500 m. (Kjellberg 79/BO.).

New GuINgEA: Mount Wilhelmina, wet slope, burnt subalpine forest,

3,400 m. (Brass & Myer-Drees 9738/BO.); Lake Habbema, 2,600 m.,

still water in stream (Brass 10897/BO.).

Mataya: Kedah: Kedah Peak, + 914 m., (Holttum 14882a/SING.).

Kelantan: Gunong Sitong, + 790 m., (Nur 12243/SING.). Selangor:

Ulu. Semangkok, (Ridley 277/SING.). Malacca: Gunong Ledang

(= Mt. Ophir) (Ridley 221/SING.). Pahang: Gunong Batu Brinchang,

+ 1,758 m. in open near summit, (Holttum 23529/SING. & BO.);

+ 948-1,370 m., (Holttum 20918/SING.); Pine Tree Hill,

+ 1,463 m. (Holttum 21561/SING.); Gunong Tahan; 1,066 m. on

ground on the ridge, (Holttum 20914 & 20915/SING.); 1,525 m. on

ground in wood, (Holttum 20905/SING.); 1,827 m. (Wray & Robinson

5435/SING.; (Ridley in 1908/SING.); Cameron Highlands, 1,553 m.

(Henderson 17877a/SING.).

DISTRIBUTION: Japan, Formosa, Malaysia, Himalaya.

After careful examination of all the specimens available to me,

I am convinced that S. gedeanum Dz. et Mb. is not a distinct

species. |

Warnstorf regarded S. gedeanum as a variety of S. junghuhnia-

num. I venture to suggest that there is no grounds even for varietal

distinction. The “gedeanum” features, i.e. presence of auricle, in-

distinctness of border and absence or paucity of spiral thickenings

in the stem leaves do not necessarily occur together in the speci-

mens and seem to vary independently. The presence of auricles is

very sporadic and found in specimens consigned to both “species”’.

In only five specimens did IJ find no trace of spiral thickenings of

the hyaline cells in the stem leaves. From this extreme there is a

complete range from a few cells with spiral thickenings near the

apex to complete thickening of all hyaline cells except for the

border (cf. Suzuki, 1956). The distinctness or otherwise of the

border seemed more probably to be a function of the age of the

specimen and possibly its habitat than a specific distinction. It

seems that S. junghuhnianum is a widespread and variable species

in Malaysia.

B. Subgenus Inophloea (Russ.) emend Andrews (1911)

319

Gardens Bulletin, S.

9. Sphagnum roseotinctum Johnson sp. nov.

A Sphagnum beccarii, cui affinis, plantis partim roseis, cortice

caulino multifibroso, foliis caulinis minoribus ovatisque haec spe-

cies sat distincta.

Planta partim roseo-tincta. Caulis crassus, 500u in diam., axi

lignosus, fusco-rubidus, cortice 3 strato, multi-fibroso, cum cellulis

superficialibus externe uniporosis vestitus. Folia caulina ovata,

parva, 500, longa, 250, lata, fibroso. Folia ramea ovata cucullata,

2 mm. longa, 0-7 mm. lata, cellulis chlorophyllosis anguste trape-

zoideis in sectione eorum transversali utrinque libere disposita.

Plant partly tinged rose. Stem thick, 500» in diameter, woody

and dark red in the axis, covered with 3 layered, multifibrous cor-

tex, which has the superficial cells provided externally with one

pore each. Stem leaves ovate, small, 500, long, 250, wide, fibrous.

Branch leaves ovate, cucullate, 2 mm. long, 0-7 mm. wide, with

chlorophyll cells narrow trapezoid in section and free on both sur-

faces, but with greater exposure on the ventral side.

MatayA: Kelantan: Gunong Sitong, on ridge 792 m. (Nur 12244/

SING., holotypus). Kedah: Kedah Peak (Spare 1439 & 1430/SING.).

This species is clearly distinct from S$. beccarii in the numerous

fibres in the stem cortex and in the very small stem leaves.

10. Sphagnum holttumii Johnson sp. nov.

Sphagnum magellanico similis, sed foliis caulinis brevissime lin-

gulatis, cellulis chlorophyllosis in sectione folii ramei transversali

rectangulis utrinque vel in latere ventrali tantum liberis haec species

sat diversa. :

Epidermis caulina 3-strata, multifibrosa, cum cellulis superficiali-

bus externe uniporosis. Folia caulina parva, lingulata, fibrosa, 2

mm. longa, 0:6 mm. lata, elliptico-ovata, apice cucullata, concava,

foraminibus magnia in cellularum angulis et prope commissuras

instructa, cellulis chlorophyllosis in transverse sectione rectangulis,

liberis ventre vel utrinque liberis praedita.

Stem epidermis three layered, multifibrous, with the superficial

cells provided each with a single pore. Stem leaves small, lingulate,

fibrous, 2 mm. long. 0-6 mm. wide, cucullate and concave at the

apex, and provided with large pores at the cell angles and junctions

chlorophyll cells in transverse section being exposed ventrally or

on both sides.

Ma.aya: Pahang: Gunong Tahan, alt. 1,500—2,000 m. in sandy and

other places, and in the open or shady localities (Holttum 20906, holo-

typus in SING.; 20908; 20909; 20910; 20916 and 20919—all in SING.);

Cameron Highlands, alt. 1,250 m. in swamp (Holttum 23300/SING.).

Kedah: Kedah Peak (= Gunong Jerai), alt. 1,250 m. (Abdul Kadir

19771/SING.). Johore: Gunong Pantai, alt. 500 m. (Corner 23207/

SING.).

320

Vol. XVII. (1958).

This species differs from S. magellanicum in the shape and posi-

tion of the chlorophyll cells and the shape of the stem leaves.

11. Sphagnum beccarii Hpe. in Nuovo Giorn. Bot. Ital. (1872)

278.

Sphagnum pauciporosum Warnst. in Hedwigia 39 (1900) 109;

cf. Andrews (1951).

Robust species. Stem with cortex about three cells thick, fibres

few. Outer cells with large single pore/cell. Wood cylinder distinct,

chocolate brown, pith pale yellow, thick walled cells. Branches

1-2 cm. long, blunt, turgid. Stem leaves lingulate, 1-75 0-75 mm.,

spiral thickenings near to apex, fimbriate tip; numerous large dor-

sal pores median in hyaline cells, often almost equal to cell dia-

meter, no border. Branch leaves wide imbricate, concave, about

1:72 « 15 mm., tip obtuse, cucullate, denticulate; ventral pores

near cell walls; pseudopores present dorsally, chlorophyll cells

trapezoid included dorsally.

SUMATRA: Gunong Kenieri, 3,200—3,300 m. (van Steenis 10256/BO.).

BoRNEO: Bei Kenepei (Fleischer, Serie II-1899/BO.).

Matava: Pahang: Cameron Highlands, 1,548 m. (Henderson 17877c/

SING.).

DISTRIBUTION: Malaysia.

In his paper Andrews (1951) “hesitates to suggest” that this

species is synonymous with S. erythrocalyx or S. leionotum but

points out several features of similarity with these South American

species. There is little material available of S. beccarii for the

detailed study needed to determine this point.

12. Sphagnum borneense Warnst. in Allgem. Bot. Zeitschr. (1895)

229.

Moderately robust. Main stem cortex fibrous, 3—4 cells thick,

outer cells with 1—4 pores. Stem leaves triangular with dentate tip,

about 1:3 0:5 mm., no fibres in hyaline cells. Branch leaves

large, concave, rounded, cucullate, ovate to oval, 1:5 & 0-8 mm.,

chlorophyll cells very small triangular in section, included on dor-

sal side; hyaline walls adjacent to chlorophyll cells densely

papillose.

Borneo: Sarawak: Mount Santubong (Native collector 2236/BO.).

The specimen examined by Andrews came to Europe as packing

material and Andrews doubts if it came from Borneo at all. The

specimen at Bogor seems a perfectly genuine collection from

Borneo. It is very like S. papillosum, but even more papillose.

321

Gardens Bulletin, S.

So) Os

Ag. Ob

Fig. 1. Folia. S. flaccidifolium: a = folium rameum; b = folium caulinum.

S. roseotinctum: c = folium caulinum; d = folium rameum;

e = folium caulinum valde ampliatum. S. cuspidatulum v. treng-

ganuense: f = folium caulinum; g = folium rameum. S. holt-

tumii; h = folium rameum; 1 = folium caulinum.

——)

'Ou

Fig. 2. Sections transversales. a, S. roseotinctum: t.s. folii ramei. b, S. holt-

tumii: t.s. folii ramei.

322

Vol. XVII. (1958).

Acknowledgements

This work was carried out in the Department of Botany, Uni-

versity of Malaya in Singapore, and I wish to acknowledge the help

and encouragement of Professor H. B. Gilliland. The specimens

were made available by the Directors of the Botanic Gardens in

Singapore and in Bogor. I would like to thank the staff of the

Kebun Raya, Indonesia, who made my trip to their herbarium a

most enjoyable one. Lastly I would like to thank Dr. C. X. Furtado

who gave considerable taxonomic help and translated the Latin

diagnoses.

References and other relevant papers

ANDREWS, A. LE Roy (1913). North American Flora, /5 part 1.

ANDREws, A. LE Roy (1936). Notes on the Warnstorf Sphagnum

Herbarium—Ann. Bryol. 9, 3—12.

ANDREWS, A. LE Roy (1951). Studies in the Warnstorf Sphagnum

Herbarium VI Bryologist 54, 83-90.

BaRTRAM, E. B. (1936). Bornean Mosses principally from Mt.

Kinapaie—Phil. Journ. Sci. 61, 235=—252.

BARTRAM, E. B. (1942). Third Archbold Expedition, Mosses

from the Snow Mountains, Netherlands New Guinea

—Lloydia, 5, 245-292.

BaRTRAM, E. B. (1945). Mosses of the Morobe District of N.E.

New Guinea—Bryologist, 48, 110-126.

DaLBy, D. H. (1957). The Warnstorf Moss Herbarium—-Trans.

Brit. Bryol. Soc. 3, 449.

Dixon, H. N. (1926). A list of the Mosses of the Malay Penin-

sula—Gard. Bull. Singapore 4, 1-46.

Dixon, H. N. (1932). Contributions to the Moss Flora of Su-

matra—Ann. Bryol. 5, 17—50.

Dixon, H. N. (1934). Mosses of Celebes—Ann. Bryol. 7, 19-

36.

Dixon, H. N. (1935). On the Moss Flora of Borneo—Journ.

UCing. Soc. Bot. 50, 57—140.

Drxon, H. N. (1939). High alpine mosses from Sumatra—Ann.

Bryol. 12, 48—56.

Drxon, H. N. (1941). New and Rare Bornean Mosses—Journ.

Bot. FORST «F2.

Dixon, H. N. (1942). Papuan Mosses—Journ. Bot. 53, 1-11.

az3

Gardens Bulletin, S.

Dixon, H. N. (1943). Alpine Mosses from New Guinea—Farlo-

wia I, 25—40.

FROELICH, J. (1955). Die von Prof. F. Ruttner 1928/29 auf Bali,

Java und Sumatra gesammelten Musci—Arch. Hy-

drobiol. Suppl. 21, 3/4 299-342.

Gipss, L. S. (1927). A Contribution to the Flora and Plant For-

mations of Mount Kinabalu and the highlands of

British North Borneo—Journ. Linn. Soc. Bot. 42,

1-240.

SUZUKI, Hyosrt (1956). Variations in Sphagnum junghuhnianum

var. pseudomolle Warnst and the Status of S. kieens

Warnst.—Jap. Journ. Bot. 15, 186-198.

WarnstorF, C. (1911). Sphagnologia Universalis—in Engl.

Pflanzenr. Heft 51].

324

Cleome ciliata Schum. et Thonn. in Singapore

By ANNE JOHNSON AND TAN KIAP SENG

Department of Botany,

University of Malaya, Singapore

AMONG THE many ecological problems awaiting solution in

Singapore Island, is the rapid spread of introduced weeds on soil

bared by land utilisation. In recent years Cleome ciliata Schum. et

Thonn. has shown wide distribution, occuring as a weed of waste-

ground, manure heaps and in exposed situations along the drains

of roadsides. This plant of tropical African origin (Hutchinson

and Dalziel, 1927) is widely distributed both in tropical Africa and

Jamaica. It was first collected in Singapore in 1927 and since then

has spread rapidly over Singapore Island and is locally abundant

in parts of the Federation of Malaya. In this paper some sugges-

tions will be made as the reasons for its rapid spread and its

successful colonisation of exposed sunny land. Other features of

ecological interest will also be mentioned.

The Good Colonising Power of Cleome ciliata in Singapore

Studies of the autecology of this species showed that particular

features of its morphology and life history gave it considerable

selective advantage in colonising bare exposed soil in Singapore.

(a) Vegetative development

The root system is a spreading one which consists of lateral

roots extending horizontally and forming a circular mat at a depth

of 5—10 cm. from the surface. For a well grown plant which sprawls

over an area of 120—160 cm. diameter, the root system will form

a mat some 60 cm. in diameter. Growing in this particular way, the

roots meet with little competition from other species of plants

growing in the same habitat. For instance Phyllanthus niuri,

Euphorbia hirta, Oldenlandia corymbosa and _ Alternanthera

speciosa have tap root systems in which the central main root

grows straight down to a depth well below that of the Cleome

ciliata root zone. Shallowly rooted sedges and grasses do not

penetrate below the first 6 cm. of soil. (Fig. 1).

The plant grows erect to a height of 30 cm. and then falls flat

on the ground where it continues to grow and flower. Such flat

sprawling plants have a good chance of survival in an exposed

situation.

325

Ground level

Gardens Bulletin, S.

G F D A 6 C

Fig. 1. Root systems of other herbaceous plants growing in association

with that of Clecme ciliata; (A) Cleome ciliata; (B) Alternan-

thera triandra; (C) Phyllanthus niuri; (D) Oldenlandia corymbosa;

(E) Cyperus spp.; (F) Borreria latifolia; (G) Euphorbia hirta.

(b) Reproduction

The flowers are distinctly dimorphous with respect to the ovaries

of the mature flower. In the normal ovary flower, the mature ovary

is almost as long as the filaments of the shortest pair of stamens

(8-0-10-0 mm. long), about 1:0 mm. in diameter and bears a pink,

papillose, sticky stigma. In the short ovary flower, the mature

ovary is only 2—3 mm. long and 0-5 mm. in diameter. The stigma

is green and undeveloped and no seed is set. (Fig. 2). Both types

of ovary occur on the same plant and the ratio of normal ovary

flower/short ovary flower varies from 1-1 to 2-0, i.e. there are

always more normal ovary flowers than sterile short ovary flowers.

Artificial pollination experiments showed that while the pollen

of both types of flowers was viable and successfully fertilised the

normal ovary flowers; the short ovary flowers would not set seed

either with pollen from flowers of its own type or with pollen from

the normal ovary flowers.

326

Vol. XVII. (1958).

Fig. 2. Flowers of Cleome ciliata

A, Short ovary flower with abortive ovules and undeveloped

stigma (b).

B. Normal ovary flower with fertile ovules and papillose

stigma (a).

In A and B, “honey guides” are indicated (c).

In the field Apis indica, a honey bee, is a regular visitor to both

types of flowers collecting nectar and pollen. Owing to the

proximity of the lower pair of stamens and the stigma of the nor-

mal ovary flowers, self-pollination may also take place to a limited

extent. The short ovary flowers seem simply to furnish additional

supplies of pollen, so that fertilisation of the normal ovary flowers

is assured. In the field it was observed that between 50 per cent

and 71 per cent flowers set seed and only a few of normal ovary

flowers failed to be fertilised.

The seeds are attached along the sutures formed by the two

carpels of the fruit and average 50 seeds per pod. They are small

and reniform with the embryo curled up with the sharp radicle

end touching the cotyledons to form a disc-like structure. The

seed coat is thick and transversely corrugated. On either side of

the tip of the seed coat are two float-like structures called here

“elaiosomes”. The elaiosomes which are full of fatty substances,

develop from the tips of the outer integuments from special cells

differentiated before fertilisation. Three days after fertilisation

these cells form a bunch-like structure which enlarge and appear

as glistening structure as the mature seed turns from brown to

black on ripening.

When the fruit is mature, it splits from below upwards forming

two valves. Some seeds are shot out immediately while others are

retained and later jerk out by movements of the plant in the wind,

327

Gardens Bulletin, S.

or are carried away by ants. The succulent elaiosomes are particu-

larly attractive to ants. They carry the seeds away and eat the

elaiosomes leaving the seed scattered abroad.

The seeds are lighter than water. This is partly due to the

corrugated seed coat which causes it to resist wetting; and partly

to the fat content of the elaiosomes which act as flotation devices.

Since one common habitat of Cleome ciliata is along drains and

canals on Singapore Island it is quite probable that the seed find

new habitats by dispersal for short distances in the water.

Experiments were done to test the viability and longevity of the

seed of Cleome ciliata. It was found that fresh seed would not

germinate in the dim light of a laboratory, but seeds, in similar

germinators placed on a windowsill receiving at least two hours

direct sunlight a day, germinated with a variable percentage

germination according to the seed sample used. Air drying of the

seed for 1-7 days does not apparently decrease the percentage

germination of samples, but air drying for four months decreases

it to 4 per cent.

This apparent marked effect of light is interesting since Cleome

ciliata is essentially a plant of sunny wastelands and is never found

deep in the forest, nor in shady situations. If seeds found their way

to such a habitat they would fail to germinate. Removal of

elaiosomes does not affect the percentage germination, therefore

ant-dispersal is quite feasible.

When seed of Cleome ciliata are sown thickly in a garden a good

percentage germination is obtained, but not all the seedlings will

survive. There is a marked effect of crowding resulting in the pro-

duction of etiolated thin plants, producing flowers somewhat

smaller than normal. In one experiment only 58 per cent plants had

survived after 52 days.

Such overcrowding by plants of the same species is unlikely to

occur in the field due to the diverse methods of seed dispersal

which ensures a wide range of destination for seed produced by

any one plant. Nevertheless Cleome ciliata meets competition from

taller life-forms, colonising bare ground in the next stage of the

succession. In the field it reacts rapidly showing etiolation of stems,

yellowing of leaves, smaller flowers and fewer seeds. If a locality

is completely covered with grasses such as Ischaemum; or with

Mimosa pudica or Cyperus species Cleome ciliata never becomes

established. Heavy shade from trees or scrub (e.g. Adinandra

dumosa/Wormia suffruticosa scrub) stops the colonisation by this

plant.

328

Vol. XVII. (1958).

DISCUSSION

Cleome ciliata is particularly successful as a pioneer weed colo-

nising bare ground. Its roots system is relatively shallow and does

not tap the same zones as other pioneer weeds growing in the same

vicinity. It has a very rapid growth and by its prostrate habit soon

covers extensive areas of bare soil. It has prolific production of

seeds which have a wide range of dormancy (from 10 to 126 days).

These are typical microbiotic seeds of a tropical climate

(Crocker & Barton 1953). About 50 per cent flowers set seed. The

other flowers are short ovary “pollen flowers” and have a sterile

ovary. The seeds are dispersed by splitting of the pod, by being

carried away in water currents of fioodtide, or by ant-dispersal.

In very exposed situations the stem and adaxial leaf surface

developed an erythrite red colour. The flower size was reduced

and the number of seeds per fruit decreased. These effects which

were not investigated further may well be due to a mineral

deficiency.

The tender leaves and shoots are the favourite food of the larvae

of a butterfly (Appis libytheae olferna). The Capparidaceae are

often the food plants of the Pieridae to which family Appis

libytheae olferna belongs (Corbet & Pendlebury, 1956). Although

damage may be done to individual plants, the plants are far too

numerous to be exterminated.

SUMMARY

1. Cleome ciliata Schum. and Thonn., 1828, a native of Tropical

Africa was first collected in Singapore in 1927 and since

then has spread throughout the Island and in the Federa-

tion of Malaya, growing in open waste-land and by road-

sides.

2. The plant produces two types of ovaries :—“‘normal” ovary

which is fertile; and “short”? ovary which is abortive. Both

types are produced on the same plant. All pollen produced

is viable.

3. The seeds bear two, float-like structures, called here “elaio-

somes” are not essential for germination of these seeds.

4. The elaiosomes play a double role in seed dispersal. They are

partly responsible for the flotation of seeds; and they

provide food for ants which scatter the seeds. The elaio-

somes are not essential for germination of these seeds.

329

Gardens Bulletin, S.

5. The percentage of germination of seed varies greatly but no

germination was observed unless they were exposed to

direct sunlight of at least 2 hours duration/day. This effect

may be due either to true light sensitivity or to the effect

of increased temperature.

60° THe morphological development of the plant from germina-

tion to fruiting is traced.

ACKNOWLEDGEMENTS

The experiments and observations described in this paper were

carried out by the junior author in connection with his course for

the degree of B.Sc., with Honours in Botany. For this work the

senior author acted as supervisor.

We wish to acknowledge the help and encouragement of Profes-

sor H. B. Gilliland, at whose suggestion this work was carried out.

We have received the help and co-operation of members of the

Departments of Chemistry, Physics, Geography and Zoology who

have made our investigation a very interesting one. The Botanic

Gardens’ staff have given their unfailing assistance and support.

We wish to thank Professor H. G. Baker for his helpful criticism.

REFERENCES

CorBET, A. S. & PENDLEBURY, H. M. (1956). The Butterflies of the

Malay Peninsula, page 125, Oliver & Boyd. 2nd

Edition.

CROCKER, W. (1950). Growth of Plants. Reinhold Publishing Cor-

poration, N.Y.

CROCKER, W. & BARTON, L. B. (1953). Physiology of Seeds. Chro-

nica Botanica Co., Waltham, Mass., U.S.A.

DE CANDOLLE, A. (1884). The Origin of Cultivated Plants, Kegan

Paul, Trench Co., London.

HUuTCHINSON, J. & DALZIEL, J. M. (1927). Flora of West Africa,

Vol, 1,

OLIVER, D. (1868). Flora of Tropical Africa 1, 78. Reeves & Co.,

Ltd., London.

Riwiey, H. N. (1922). Flora of the Malay Peninsula 1, 119.

Reeves & Co., Ltd., London.

330

OBITUARIES

Haji Mohamed Nur bin Mohamed Ghous, B.E.M.

MoHAMED Nur BIN MOHAMED GHOUS was born in Singapore

“during Bulan Puasa” in 1898. This was before registration of

births was a general practice and his birth appears to have been

undocumented. In that year Puasa occurred during March. Later

his birth was dated 22nd March, 1898.

His father was a Malacca Malay who travelled between Malacca

and Singapore trading in cloth. His mother, Yang Chik binte

Saman, was a Singapore Malay. They had four children, three boys

and one girl.

Mohamed Nur’s schooling was brief. He went first to the Kota

Raja Malay School in Singapore. After that he attended the Lorong

Pandan English School, a branch of the Government Victoria

School.

He joined the Gardens service in 1911 or 1912 at the age of 13

or 14 years during the close of H. N. Ridley’s directorship and was

employed on collecting seed for distribution in Malaya and over-

seas. Much of this work was with hevea seeds for which the

Gardens were the main source of supply.

After Ridley’s retirement, I. H. Burkill was appointed Director.

He arrived on 17th October, 1912, ten months after Ridley’s

departure. They had been months of less than standstill. The

printed literature which had come into the office was in a heap on

a table, thrown there as it arrived. In the herbarium the mounted

plants were in their cabinets, but the cabinets were in no systematic

arrangement. The Penang Gardens had been closed in mid-1910

and the land transferred to the Georgetown Municipality to make a

water reservoir. The Penang herbarium had been transferred to

Singapore. The sheets remained in the cabinets used in Penang, but

no start had been made to incorporating it with the Singapore her-

barium. Nor were the cabinets in any order, but simply stacked

wherever there was room.

To set about sorting out the confusion, Burkill picked out Ahmad

bin Hassan as Storekeeper and Mohamed Nur as Label Printer as

from Ist January, 1913. The duties of this latter post were more

varied than at present understood and Mohamed Nur had the run

of the herbarium for his ability to label plants in the garden fitted

551

Gardens Bulletin, S.

him for the same function in the herbarium. Thus began his asso-

ciation with the herbarium which was to continue for the rest of

his life.

For the herbarium to be at all usable it had to be got into order.

The cabinets were of miscellaneous sizes and did not block nicely,

but by sorting them out the best use of space was achieved. This

led of course to worse confusion in the sequence of families. Since

all the herbarium sheets were in labelled folders (according to the

Bentham and Hooker system), it was left to Mohamed Nur to

rearrange them. Commenting on the ability he displayed in doing

this, Burkill has said: “Nur means Light, and this is where he

shone it”. As science, whatever its context, is the pursuit of truth,

it seems wholely appropriate to quote R. J. Wilkinson [A Malay-

English Dictionary, 1955]: “Nur Ar. Light—whether the solar

light by which men see, or the primal light that illumined chaos

at the Creation, or the light of divine revelation that guides men to

Truth”. As Mohamed Nur was to give his life-work to the herba-

rium, and developed an innate perspicacity for botanical taxonomy,

how appropriated was he named!

Nur’s growing knowledge and an unusual habitual orderliness

clearly pointed his career to greater responsibilities. On Ist Septem-

ber, 1916 he was promoted to become Plant Collector. His collect-

ing was extensive and he held the post till the end of 1923, when

the post of Herbarium and Museum Assistant having fallen vacant,

he was appointed to it. I. H. Burkill, in recommending to Govern-

ment his promotion wrote: “The subordinate whose name I put

before you is singularly efficient and energetic, and is capable of

higher work than that of plant collector. He has acquired a surpris-

ing knowledge of the latin names of Malayan plants, and by his

methodical ways is calculated to keep the Museum and Herbarium

in due order’.

The post of Herbarium and Museum Assistant to which he came

on 1st January, 1924 was held by him throughout the rest of his

career. He retired on pension on 22nd March, 1953 on attaining

the age of 55 years after over 40 years service which he was yet to

extend by another 54 years. At the time of his retirement he had

acquired such a comprehensive knowledge of Malaysian plants, and

there being no-one available to fill his late post, that after a break

of one day as is required by regulations he was re-engaged in the

same post. This he held to the day of his death, 7th November,

1958. During this time he saw the herbarium grow to be the fourth

largest in Asia with a collection of about 400,000 specimens to

which he personally had in no small measure contributed.

332

Vol. XVII. (1958).

In the field, he visited most parts of Malaya; he also collected

in Thailand, Sumatra and Rhio. In his earlier days he accompanied

I. H. Burkill extensively over Malaya, and with Mohamed Hanifi,

Superintendent of the Penang Gardens, went to many places in the

northern half of the peninsula, the most extensive expedition being

made in January and February 1923 in Ulu Kelantan to the north-

ern side of Gunong Tahan. After Holttum and Henderson had

joined the Department in 1922 and 1924 respectively Mohamed

Nur accompanied them on many expeditions between then and the

war, from Langkawi and Perlis in the north to the southern end of

the peninsula, from the coastal plains and the mountains of the

west to the east coast and the off-shore islands of the Pahang/Johore

group. He also went in charge of independent assignments and in

October and November 1937 collected in Selangor for the Arnold

Arboretum, U.S.A., in conjunction with the F.M.S. Department of

Forests.

All his collecting is numbered in the field in the S.F.N. (Singa-

pore Field Number) series and his collections are laid in the

Singapore Herbarium with duplicates sent out to many other her-

baria. A list of his major collecting trips is given at the end of this

notice. On one of them he took the only known Malayan flowering

specimen of Bambusa pauciflora Ridl. which he got on the ridge

leading to the summit of Pine Tree Hill (4,780 ft.) on the Pahang-

Selangor border.

Ridley’s Flora of the Malay Peninsula was published between

1922 and 1925, the major part being out by 1924. With this as the

standard record of the Malayan flora, the sheets in the Singapore

Herbarium were rearranged:in accordance with its sequence. This

work naturally fell to Mohamed Nur and occupied him for much of

1924/25.

Immediately following on this Mohamed Nur undertook in 1926

the integration of the Perak Museum herbarium with the Singapore

herbarium. And for several years leading up to 1928 he carried

out various assignments for the F.M.S. Forest Department herba-

rium for which the F.M.S. Government gave him an honorarium.

These services were in fact regularly rendered by Mohamed Nur

right up to the time of his death and by the Gardens Department

generally then and now as a part of the Department’s fundamental

purpose of studying the Malayan flora.

Mohamed Nur’s contribution to the knowledge of the local flora

is not marked by any list of publications. His efforts have been

merged with and are contained in the publications of others to

whom he rendered ready and competent assistance in undertaking

333

Gardens Bulletin, S.

preliminary determinations of collected material. These determina-

tions were made with an unusually gifted knack, but entirely lack-

ing the substantial pedestal of scientific theory and precept, which

of course he had never received. His ability to “spot” sterile

material raised as much admiration as anxious doubts amongst the

conventional, but so often could it be shown subsequently that he

was usually right. Every generation produces a few men of genius,

and here in a humble way was one of them.

In the field of botany as in other sciences recognition of merit is

often acknowledged by co-workers by the use of personal names to

label some object or phenomenon. Bestowing of personal names

for new species or genera of plants is common and Mohamed Nur’s

name has been given to several species which are listed below. In

a similar but official manner Government in recognition of his long

and faithful service awarded him the British Empire Medal in 1956.

During his last few years of service he suffered increasingly from

breathlessness, but he refused steadfastly to take life more easily.

He maintained that if he stopped working he would quickly die.

Since there was no immediate replacement for him and since he

lived solely for his work it was to his and to the Gardens’ advantage

that he stayed in harness. Even during the Muslim fasting month

(Bulan Puasa) of 1958, which period to younger men in the best

of health is deemed a test of moral and physical endurance, he

refused to consider any adjustment of his working hours. His

objection was simple: he did not want to be different. But in fact

he was different in so many ways of punctilious honesty towards

his work. A single example will serve to show his faithfulness.

During the final fortnight of the Japanese military onslaught against

Singapore in 1942, when all was sudden death, chaos and confu-

sion, and again in 1945 at the time of the collapse of the Japanese

hegemony, the office attendance register shows that Mohamed Nur

reported with regularity for work, and at the height of these periods

of the subordinate staff only he and his life-long colleague ‘Che

Ahmad bin Hassan signed in.

In his private life there is little to record for his work was his

life. On 9th September, 1919 he married ‘Che Fatimah binte

Ariffin, in his only marriage by which there were six children,

three sons and three daughters, all of whom survive him. In 1949

he made the Hadj pilgrimage to Mecca and to other holy places of

Arabia.

His death was quite sudden and peaceful on the evening of 7th

November, 1958. He came to work as usual, but at midday col-

lapsed and was taken home. There he ate a little, then settled down

to sleep, and that was the end.

334

Vol. XVII. (1958).

In botanical history there are many instances of father-son suc-

cessions. This inheritance of similar interests and abilities would

make an interesting genetical study, and here indeed is yet another.

For the four years before Mohamed Nur’s death his son, Mohamed

Shah, had been working as a Plant Collector in the herbarium.

During this time Mohamed Shah showed and developed the family

trait of an untutored botanical taxonomic acumen. It must have

been a satisfaction to the old man that his son was destined to

follow his footsteps. Four months after his death Mohamed Shah

was confirmed in the post.

Plants named after Mohamed Nur

Dioscorea nurii .. Kunth in Engler’s Pflanzenreich, [V—

43, 1924 (Reduced to D. kingii

Kunth).

Begonia nurii .. Irmscher in Mitt. Inst. Bot. Hamburg

8, 1929.

Cryptocoryne nurii .. Furtado in Gardens’ Bull. S.S. 8, 1935.

Daemonorops nurii .. Furtado in Gardens’ Bull. Singapore 14,

1953.

Ardisia nurii .. Furtado sp. nov. in this issue.

Mohamed Nur’s major plant collecting expeditions

(No record is available of innumerable minor collecting trips and

single day forays)

Date Locality Accompanying

1913-24 (annually, Penang, Port Swettenham, occa- I. H. Burkill or

covering every sionally to Klang alone

month of the year

except January )

July 1914 Tampin, Negri Sembilan —

June 1915 Pulau Tinggi, P. Tioman, Johore/ I. H. Burkill

Pahang

November 1915 Pulau Terutau, S. Thailand Md. Haniff

February 1917 Taiping Hill, Perak Md. Haniff

December 1920 Salak & Sg. Siput, Perak Md. Haniff

August 1921 Sibolangit, Sumatra East Coast F. W. Foxworthy

J. A. Lorzing

November 1921 P. Langkawi group, Kedah Md. Haniff

June 1922 Tembeling, G. Tahan (south side)

Pahang Md. Haniff

s.d. 1922 G. Pulai, Johore Kiah

Date

Jan./Feb. 1923

Aug./Sept. 1923

November 1923

December 1923

Feb./ Mar. 1924

November 1924

February 1926

Apr./May 1927

Oct./Nov. 1927

December 1927

March 1928

Aug./Sept. 1928

April 1929

July/ Aug. 1929

November 1929

April 1930

May/June 1930

October 1930

November 1930

April 1931

May/June 1931

October 1931

November 1934

May 1936

April 1937

October 1937

Gardens Bulletin, S.

Locality

Sg. Kelantan, Sg. Labir, Gunong

Tahan (north side), Bukit Teman-

gan, Kuala Krai, Kelantan

Fraser’s Hill, Selangor/ Pahang

Gunong Angsi, Negri Sembilan

Bt. Tangga, Negri Sembilan

Sg. Buloh Forest Reserve, Selangor

K. Lipis, Gelas Valley, Gua Nineh,

G. Strong, Kelantan.

Serdang, Selangor

Raub, Kuala Lipis, Pahang

G. Panti, Johore

P. Tioman, Pahang

Pahang/ Kelantan Boundary

G. Panti, Johore

G. Muntahak, Johore

G. Tahan, Pahang/Kelantan

P. Tioman, Johore

K. Tembeling, Pahang, Kelantan/

Pahang Boundary & Kuantan,

Pahang

Perlis

Cameron Highlands, Pahang

Ipoh, Grik, Perak

Tasek Bera, Pahang

Rhio Islands

Mentakab, Tembeling, Pahang

Tembeling, Pahang/Kelantan

Bentong, Kuantan, Pahang

P. Langkawi, Kedah

Cameron Highlands, Pahang

Ringlet, Pahang

Telok Forest Reserve, Klang Sg.

Tinggi Forest Reserve K. Selan-

gor, Ginting Simpah and Bt.

Kanching, Selangor.

336

Accompanying

Md. Haniff

R. E. Holttum

M. R. Henderson

F. W. Foxworthy

Md. Haniff

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. Holttum

. Henderson

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F.M.S. Forest

Department

collectors.

H.M.B.

Haji Mohamed Nur bin Mohamed

Ghous, B.E.M.

Ngadiman bin Haji Ismail

NGADIMAN was born on 14th March, 1904 at 7 Scotts Road, Singa-

pore where his father was head gardener. He went to Tanglin

Tinggi Malay School and passed out at Standard V, the top grade.

He joined the Gardens Service on 1st January, 1921 as a label

printer and later worked as a gardener till 1929 when he was pro-

moted to the post of Plant Collector. He went on three major

collecting expeditions:

1930 to Tasek Bera, Pahang, with M. R. Henderson.

1931 to Tembeling, Pahang, and the Pahang/Kelantan

border with M. R. Henderson, E. J. H. Corner

and Mohamed. Nur.

and 1937 to Fraser’s Hill, Pahang, with E. J. H. Corner and

Kiah.

Most of his collecting was done however with Corner locally on

Singapore Island or in South Johore in single day expeditions

easily staged from Singapore.

In rural Malaya, and particularly in Kelantan, the “berok” or

pigtailed monkey (Macacus nemestrina) is often trained to pick

ripe coconuts. In April 1937 while in Kelantan Corner bought a

young berok and brought it back to Singapore with the object of

training it to pick botanical specimens from the tree tops. So pro-

mising was it that Ngadiman was sent to Kelantan later in 1937 to

find two more such monkeys, and to learn how to talk to them

and how to train them. In his quest he was successful. Thus began

an interesting and delightful incident in the history of Malayan

botanical exploration.

Such an exercise requiring co-operation between man and

monkey called for special gifts of patience and ingenuity. It was also

not without danger. Both Corner and Ngadiman were savagely

bitten by their “assistants”; on one occasion Ngadiman was off

work for 17 days on a medical certificate for “monkey bite on fore-

arms’. But the exercise was such a success and the result in terms

of cost as well as effort so relatively cheap that a fourth monkey

was acquired shortly before the war. But for the disruption which

the war brought, this aspect of botanical collecting would un-

doubtedly have been developed to its full potential.

RgO7

Gardens Bulletin, S.

After the war another attempt was made to train a berok and

Ngadiman went to Kelantan again bringing back a young one. It

grew however to be physically weak and with limitations of staff

and restrictions of collection in the Federation of Malaya owing

to the “emergency”, the attempt has not again been repeated.

With the acquisition of these monkeys, Ngadiman was tied to

their care and training. Soon after Corner had got his first one,

Ngadiman accompanied Corner with it to Fraser’s Hill with most

encouraging results. When two others were bought Ngadiman

would take one or the other out every day to the Bukit Timah

forest for training, and thereby the botany of the hill became very

considerably better known. In addition Ngadiman would take the

monkeys out to help Corner in studying the flora of the fresh water

swamp forest around Mawai and Sungei and Kuala Sedili.

One new species of plant has been called after him: Eugenia

ngadimaniana Henderson in Gardens’ Bulletin, Singapore 11,

1947.

As a result of Ngadiman’s association with Corner over the col-

lecting monkeys and Corner’s keen interest in wildlife protection,

it was natural that when botanical reserves, one of them the Bukit

Timah forest, were gazetted and put under charge of the Botanic

Gardens Department, Ngadiman should be connected with them.

Daily exercising the monkeys at Bukit Timah, Ngadiman was avail-

able for supervision. He was appointed Ranger in 1937 and re-

mained in charge of supervision of the reserves, firstly as Botanical

Reserves, then from 1951 onwards as Nature Reserves till the time

of his retirement when he held the rank of Head Ranger.

In mid-1956 advanced TB was diagnosed and the disease would

not respond to treatment. He died on 24th September, 1958, at the

age of 54 years.

Of his 11 children, two are employed in the Botanic Gardens to

maintain the family connexion with botany. H.M.B.

338

Kwan Koriba

Kwan KoriBa was born at Aomori, in the north of Honshu, in the

year 1882. As a boy he was educated in the Middle School of the

neighbouring city of Hirosaki, to which he finally returned in 1954,

and where he died, suddenly, on 15th December, 1957.

Koriba received his higher education at the University of Tokyo,

where he graduated in 1907, taking his doctor’s degree five years

later. He was Professor at the University of Tohoku from 1915 to

1920, and then at the University of Kyoto. In 1942, at the age of

60, he retired from the latter post, and was appointed Director of

the Botanic Gardens, Singapore, under the Japanese military ad-

ministration. Returning to Japan early in 1946, he lived in retire-

ment, continuing his botanical studies, until 1954, when he was

requested to undertake the office of President of the University, in

which office he continued until his death.

From his youth Koriba was interested in field botany, and in

observing the behaviour of plants in nature; on such subjects he

published several papers. He travelled extensively at various times,

to Europe, North and South America, and nearer home to Micro-

nesia and Java, taking the opportunities of such travel to widen

his acquaintance with plant life. While at Tokyo, he began to

devote himself to the study of environmental factors and their

effects on plants, and in this utilized his considerable knowledge of

physics; this work involved extensive research on transpiration and

evaporation. He continued his interest in the relation of plant-

behaviour to climate in Singapore, where he made records of the

periodicity of growth and leaf-change in many kinds of trees. A

summary of these observations has recently been published in Vol.

17 (1) of the Gardens Bulletin.

During his period of retirement from administrative duties, in

1946—54, he published two books, one on the forms of plants in

relation to organization and function, and one on plant physiology

and ecology. He planned a further book on the evolution of plants.

At Singapore he was given the rank of Brigadier-general, but

wore the uniform of that rank only on ceremonial occasions. He

devoted himself to his administrative duties and to his botanical

studies. He was dependent on the military organization for funds

and supplies of all kinds, and took every opportunity of securing

such amenities as were possible for the gardens staff. On several

occasions he took energetic action to prevent encroachment on the

339

Gardens Bulletin, S.

Nature Reserves of Singapore, in which he took great interest. He

was also greatly concerned that the herbarium and library at the

Gardens should be maintained intact. To his single-minded devo-

tion to botanical science the Singapore Botanic Gardens owes

much.

He was a man that commanded respect by his simplicity of life

and unfailing devotion to his duty, by his very broad scientific out-

look, and also by his sympathetic understanding of human nature,

an understanding that transcended racial boundaries, even in time

of war and of privation. R.E.H.

340

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OTHER PUBLICATIONS OF THE BOTANIC GARDENS

SINGAPORE

. Annual Reports 1875-—*

Reports for many years 1886 onward remain available.

Prices variable—will be quoted on application.

2. The Agricultural Bulletin of the Malay ah ora (Series I).

Nos. 1-9, 1891-1900.

Only Nos. 3, 5, 7, 9 and 10 remain available at 20 cents

each.

. The Agricultural Bulletin of the Straits and F.M.S. (Series II).

Vols. 1-10, 1901-12, monthly issues.

All available at 50 cents per issue or $5 per volume,

except Vol. 1, pts. 6 and 11.

4. The Gardens’ Bulletin, Straits Settlements (Series III).

Vols. 1, 1912-11 (3) 1941.

Price variable according to size.

All available except Vol. 2, pts. 2, 3, 6-11, and Vol. 6,

pts. 1 and 2.

Vol. 1 (1-5) 1912 issued under the title of Agricultural

Bulletin of the Straits and F.M.S.

5. The Gardens’ Bulletin, Singapore (Series I'V).

Vol. 11 (4) September 1947-*

All parts available. Price variable. Subscription rates will

be quoted on application. Issued irregularly.

6. An illustrated guide to the Botanic Gardens, Singapore.

Price $1.

. Materials for a Flora of the Malay Peninsula (Monocotyle-

dons). .

Parts I, II and III available.

. Malayan Garden Plants: a series of eleven booklets each illus-

trating 10 useful garden plants.

All available. Price 50 cents each.

9. A Revised Flora of Malaya.

(a) Vol. I Orchids by R. E. Holttum, 1957 (2nd edi-

tion). Price $20.

(b) Vol. II Ferns by R. E. Holttum 1955. Price $20.

. (a2) Malayan Orchid Hybrids by Mr. M. R. Henderson and

G. H. Addison. Price $21.

(b) Malayan Orchid Hybrids, Supplement I by G. H. Addison

(in preparation).

11. Wayside Trees of Malaya by E. J. H. Corner, 2 vols. Price $25.

Items 1-8 can be obtained from the Botanic Gardens,

Singapore, 10.

Items 8-11 can be obtained from the Government

Printer, Singapore, 13.

Prices quoted are in Malayan Dollars. In certain cases

postage is extra.

“IOWA

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BPUAYUAUAYVAY PD UA™"UWAU MAUI. MAUI. IV?

THE

SINGAPORE

Vol. XVII, Part III

To be purchased at the Botanic Gardens, Singapore

Price: $12.50

| (Volume XVII complete $25)

Published by Authority

PRINTED BY A. G. BANFIELD, GOVERNMENT PRINTER, SINGAPORE

——

- 2S

#? * RECEIVED .* |

GARDENS BULLETIN

pipea! ead a dic Se iat ie hat ll <PUPUUMV\PYVO UU VY VVE

18th May, 1960

VAM VUPUV OVW VU UV VOU UAV VOL AVAUVWAVVANE) &

CONTENTS

3 PAGE

SANTAPAU, H.: I. H. BuRKILL in India . 341

Furtapo, C. X: and R. E. Hottrum: I. H. Buin in Msi Peis ia)

VAN STEENIS, C. G. G. J.: Phellodendron, a genus of trees new to

the Malayan flora (Rutaceae) . 357

HoLttTuM, R. E.: Vegetative characters distihgeniithy the various

groups of ferns included in Dryopteris of Christensen’s Index

Filicum, and other ferns of similar habit and sori 361

Corner, E. J. H.: Taxonomic Notes on Ficus emg Asia aia

Australasia. Sections 1—4 368

Editor.: Posthumous Publication of iNew Di pterocarp Species from

North Borneo 486

B..E. S.: G. H. S. Woop, M.a., F.L.S. (A tribute) . 498

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I. H. Burkill in India

H. SANTAPAU, S.J., F.N.I.

FOR MANY YEARS I had been familiar with I. H. Burkill and the

splendid work he had done during his time in India as an officer

of the Botanical Survey of India; it was my privilege during an

extended visit to Kew Herbarium in 1946-1948 to come to know

him personally, and this almost daily contact deepened in me the

esteem that I had conceived from the perusal of his writings.

It is not an easy matter to give in short a sketch of the Burkill I

came to know and respect. In my opinion, and this seems to be the

common opinion of those who know him well, I. H. Burkill is a

charming gentleman of the good old school, always courteous and

obliging, ever ready to extend -.a helping hand to an honest strug-

gling botanist. In his work Burkill is one of the most meticulous

and accurate botanists that it has been my fortune to meet in my

life; in his papers and books every item is checked properly, every

minutest and most insignificant detail is tested. Such accuracy at

times might appear almost a waste of energy; but research workers

are grateful that he has had the patience to delve into such minute

details. As an example of his care, one of the first examples that

came to my notice in Kew Herbarium, I might cite but one case.

Burkill found that one of the Compositae of South America was

known under the Spanish-American name of “Coronilla de oro”

(meaning “The Little Golden Crown”); to Burkill it was not

enough to find out what the particular plant was, but he did his

best to ascertain why precisely such a plant was called by this

name.

341

Gardens Bulletin, S.

One of the reasons why he has been so accurate in his writings

is that he has spent a long life carefully taking down notes on any

point that at a later date might prove of interest. In a private

letter of Nov. 27th, 1951, he tells me, for instance: “Someone,

I forget who it was, put into print a statement that I was writing

up a history of botany in India. That was not exactly true; but I

was collecting material that would help towards a history . . . This

collecting began when I was arranging the collectors in Prain’s

and my volume of Dioscoreas; and the data went into slips . . .”

This thoroughness, in fact, explains the extraordinary amount of

information that one can derive from all Burkill’s books and

papers, even those written during his early years in Kew and India.

Biographical Data

1. H. Burkill was born on May 18th, 1870, near Leeds, York-

shire in England; he was trained in Cambridge, where he graduated

with B.A. Hons. in Natural Science from Caius College; I have

no details of his university career, except that he obtained the

Frank Smart Prize as a result of the performance at his final

examination. Shortly after his graduation, he was appointed As-

sistant Curator, University Herbarium, Cambridge, a post that he

held from 1891 to 1896; during that time he obtained his M.A.

from Cambridge University. From 1897 to 1899 he was on the

staff of the Royal Botanic Gardens, Kew, as Assistant in the

Herbarium; from 1899 to 1900 he was Principal Assistant in the

same Royal Botanic Gardens.

Burkill’s service in India began in 1901, when on the recom-

mendation of the authorities of the Royal Botanic Gardens, Kew,

he was appointed Assistant Reporter on Economic Products to the

Government of India, and was posted to the Indian Museum,

Calcutta, under George Watt as the Reporter. Throughout his

career in India, Burkill continued attached to the same office; in

1902 he was Officiating Reporter under Sir G. Watt; in 1907 he

was still mentioned as Officiating Reporter, although Sir G. Watt

had already retired. The Report of the Botanical Survey of India

for the years 1911-1912 says: “The Director was absent on pri-

vilege leave from 21st April to 2nd July 1911, during which time

Mr. I. H. Burkill officiated in addition to his substantive duties as

Economic Botanist of the Botanical Survey of India. Up to 31st

January 1912 Mr. Burkill was Assistant Reporter on Economic

Products officiating as Reporter, and thereafter on the abolition

of the Reportership became Economic Botanist to the Botanical

342

Vol. XVII. (1959).

Survey, which post he held for the remainder of the year.” This

would seem to indicate that Burkill was Officiating Reporter from

1902 to 1912, although in some official reports he is put down

as the Reporter from about 1908 onwards.

In the introduction to The Botany of the Abor Expedition

Burkill himself writes: “It fell to me, in the middle of the Hot

Weather of 1911, when acting as Director of the Botanical Survey

of India, during the leave of Colonel A. T.. Gage, to propose to

the Government of India that a botanist should be told off to

accompany the Abor Expedition; and to my gratification, after

Colonel Gage’s return, I was sent.” (Italics my own).

When in 1954 I informed Burkill of my appointment as Chief

Botanist to the Government of India, a new designation for the

post of Director, Botanical Survey of India, he wrote and gave

a number of details of the Calcutta he had known in his younger

days; one interesting detail, for instance, is the following: ‘Op-

posite to you in St. Xavier’s College is (or was) no. 47 Park

Street in which I lived from 1901 to about 1909. The house be-

longed to the Nawab of Murshedabad and the old Nawab did not

need a house in Calcutta; so some 5 of us chummed there. I used

to hear sometimes the austerity call at 4.30 a.m. which woke

St. Xavier’s household . . .”

Burkills Botanical Collecting Tours

During his time in the Indian Museum, Calcutta, as Officiating

Reporter or as Reporter on Economic Products, Burkill spent

much energy collecting information either in the field or through

correspondents on important economic products of India. From

various hints given by Burkill in his papers, I have been able to

deduce the dates of some of his major collecting expeditions.

Unfortunately due to pressure of work, I have not been able to

go through the publications of the Botanical Survey of India in

detail; I leave it to a future occasion to prepare a detailed list of

all Burkill’s exploration tours in India, as reported in the official

documents of the Calcutta Garden and Museum.

The Sikkim Expedition of 1906. “Mr. Burkill... paid an autumn

visit to Phallut in 1906 in search of Aconite tubers” (Kew Bull.

1907: 175). Burkill himself writes from Calcutta on October 15th,

1906: “I found my second trip to Phallut interesting and agree-

able...” What the first trip was, I cannot tell with the information

at my disposal. In the same account Burkill writes: “I amused

myself by the wayside making flower pollination notes and am

343

Gardens Bulletin, S.

now writing up a short paper . . .” Several papers, in fact, were

published by Burkill on the subject, and this would indicate that

he had made detailed excursions to many parts of India. His

published notes on flower pollination concern flowers of the fol-

lowing areas: Bengal and Assam, North-West Himalayas, Simla

Hills, Central Provinces and Berar, etc.

Journey into Nepal, 1907. An account of this trip is given by

Burkill in Rec. Bot. Surv. India 4 (4): 59-140, 1910. The first

paragraphs of the account state: “On November 28th, 1907, after

marching along nearly one hundred miles of the Nepalese frontier

between Jainagar and Raksal, I turned, with the permission of

the Nepalese Darbar, into the kingdom of Nepal and reached

Khatmandu by the usual route on December the second. Thence

... I visited the Trisuli valley, in the neighbourhood cf Naikot.

I returned from Nepal to the plains by a route through Pherphing,

which diversifies the first seventeen miles of the wav. My dates

almost coincide seasonally with the dates of Wailich’s march to

Khatmandu, and we seem, he and I, to have gathered at a period

87 years apart the same plants in the same spots .. . It is ill

gleaning for novelties after a botanist with the keenness of Wallich;

so I got no more than three species of Impatiens and apparently one

of Eriocaulon . . .”

The Notes from a Journey to Nepal, based on the observations

made in just a fortnight, cannot and do not claim to give an

account of the flora of Nepal; “what is written .. . is, indeed, but

a superficial account of the features of the vegetation between

Raksal and the Himalaya of Central Nepal as far back as 35

miles in a straight line from the skirts of the plains and not higher

than 7,0000 ft. There was but one excuse for writing it, ie., the

great want of knowledge of the Botany of that part of the chain.”

This account has always been of particular interest to me per-

sonally, and lately this interest has been enhanced by the fact that

one of my students has been doing research for several years in

an area of Nepal that includes the whole part visited by Burkill

and previously by Wallich. Burkill in his Notes lists about 400

genera of plants collected in his short trip, and this, to say the

least, is by no means a negligible result.

The Botany of the Abor Expedition. As stated above, it was at

Burkill’s suggestion that a botanist was assigned to the expedi-

tionary force that went into the Abor Hills; it was certainly lucky

for us, his successors, that Burkill was selected for the job. His

account is one of the most carefully complete papers that have

been published for any botanical expedition of the last two hundred

344

Vol. XVIT. (1959).

years in India with the exception of Hooker’s Himalayan journeys.

Burkill and his two collectors remained attached to the expedi-

tionary force from November 29th, 1911, till March 11th, 1912.

The methods of work followed by Burkill during this trip were

determined by the type of terrain and by the fact that for the

safety of the botanists they had to betake themselves to one of

the camps before nightfall. ‘““The plants low in the natural system

of classification could be of little use to me in my particular

object, because their geographic distribution is yet so little known.

I collected them as I was able . . . Material of tall trees was col-

lected by the aid of a shot gun. An anaeroid barometer and a

camera went with me everywhere; and for a time I had wet and

dry bulb thermometers in camp, though to little good effect...

To obtain as many plants in flower as possible, I moved back-

wards and forwards along the line, and I stationed my experienced

collector about two marches behind me, that I might have a pair

of eyes on the watch for that which had not been in flower when

I passed. Having a collector behind me gave another advantage,

for thereby there was an economy in transport, as it was easy

to send to him down the line by the food supply coolies half-

dried plants for finishing; but the carrying up the line of full sup-

plies of drying paper over two marches that could be avoided,

was a consideration. It was at all times necessary to keep the

weight of the collections down, and under these circumstances

my Diary was made to serve as much as possible. It became a

record of 8,000 observations supplementing the notes with the

specimens.” Much living material was sent directly from the field

to Calcutta and thence to Darjeeling; dried plants were also sent,

when ready, to Calcutta, where Col. Gage placed them in the

hands of assistants for a preliminary examination. The final study

of the materials collected by Burkill on this occasion was only

made prior to the publication of the report in 1924. The Botany of

the Abor Expedition is an excellent account, a model of how such

expeditions should be organised, collecting done and materials

studied; the presentation of the results is remarkable for concise-

ness and completeness.

Chapters on the History of Botany in India. Although this work

has only been published recently, long after the retirement of Mr.

Burkill from India, it forms, as it were, the crown of the work

that Burkill has done for Indian botany. In November 23rd, 1951,

I wrote to Burkill from Bombay: “Next August we are celebrating

the publication of the 50th volume of our Journal of the Bombay

Natural History Society. At a recent meeting of the Committee

345

Gardens Bulletin, S.

I was requested to ask you for a contribution for the August

number of the journal. The subject mentioned was a sort of Bio-

graphical Index of “Indian” botanists, i.e., of British and foreign

botanists that have worked in India from the beginning of the

XIXth Century onwards ... As for the length of the Index, no

limit has been set .. .” Burkill’s answer was generous in the ex-

treme: “You shall have all that I can give you” (27th Nov. 1951).

“I propose to separate from my slip index all the names of bota-

nists in India which belong to those deserving of mention in the

proposed account of Botany in India 1800 forward. When I have

done that I will see if I can find time to write an account and I

will communicate with you.” (16th Dec. 1951). On Sept. 18, 1952,

he writes again: “I assure you most appropriately the work I am

doing on the history of Indian Botany is absorbing my energies

. . . My undertaking is larger than I expected; but I am getting

on with it.” Finally on March 25th, 1953, he could write that the

MS of the first chapter had been posted to Bombay. Later on when

asking about chapter 2, I received the following answer: “TI shall

get down to part 2 as soon as I can. At the moment I am entwined

by African Dioscoreas. When last year something went wrong with

my inside-machinery and I could not get over to Kew or to the

libraries in London, I lost way. But I am able to get about again

—at any rate I get over to Kew; and I am in better fettle on the

eve of 84 than I was on the eve of 83.” Chapter 2 was finally ready

for the press in August 1956. At that time I wrote to Burkill:

“From all over India and beyond I have heard great appreciation

of your first chapter, with requests that when the series is over, it

may be published in book form.” Lately, at the beginning of this

year, 1960, I had a letter from Burkill’s son, Mr. H. M. Burkill

of Singapore, in which the son writes: “I would like to add that

I arrived back here from a visit to England but 4 days ago, and

[ left my father in good health. He was busily engaged in typing

the final text of Chapter 3 of his History of Botany in India.”

Burkill and Dioscorea. Burkill’s publications, as may be seen

from the list appended to this short paper, deal mainly with

economic products; only a few give accounts of botanical trips

in India. The genus Dioscorea has always held a special attraction

for him; the yams have been known in India for centuries as im-

portant food plants; it is only natural that the Reporter on Eco-

nomic Products would sooner or later devote his attention to Dio-

scorea. His first paper on the subject was published in Calcutta

in 1904; later on he became associated with Colonel, later Sir,

David Prain in a revision of the Asiatic species of this important

346

Vol. XVII. (1959).

genus; several papers were published jointly with Prain. The crown-

ing glory of the Dioscorea work is An Account of the Genus Dio-

scorea published in four large volumes in the Annals of the Royal

Botanic Gardens, Calcutta in 1936 and 1938. Burkill today stands

out as the greatest authority on the Dioscorea family; this is why

he has been chosen to write on the Dioscoreaceae for the Flora

Malesiana; he is now engaged in the last stages of the preparation

of an account of the Dioscoreas of Africa and Madagascar.

Conclusion

Burkill was appointed Director of the Botanic Gardens, Singa-

pore, on 10th September, 1912; he left Calcutta on leave in the

middle of April, 1912; his Indian service was officially terminated

at the end of his leave on the 16th October, 1912.

On the occasion of his 90th birthday, we, Indian botanists, wish

to associate ourselves with gratitude to the homage that the Singa-

pore Botanic Gardens are preparing for this happy day. May his

life and bright intellectual powers be spared for many more years

to come, so that he may continue to shed his light over the world

and be an example to us all.

Appendix

Some of Burkill’s Publications on Indian Botany

1904. A note upon the use of Marsdenia in the Rajmahal Hills.

Agric. Ledg. 8: 113-114, 1904.

Manihot utilissima.—The Tapioca Plant: its history, culti-

vation and uses. A review of existing information. [bid.

10: 123-148, illust. 1904.

Bixa orellana.—The Annatto-dye plant. [bid 12: 177-187,

1904.

On Dioscorea deltoidea, Wall., Dioscorea quinqueloba,

Thunb., and their allies. Journ. Asiat. Soc. Bengal n.s.

73 (II): Suppl. 1-11, 1904.

1906. A parasite upon a parasite,—A Viscum apparently V. arti-

culatum, Burm., on Loranthus vestitus, Wall., on Quer-

cus incana, Roxb. Journ. & Proc. Asiat. Soc. Bengal,

n.s. 2: 299-302, 1906.

Gentianacearum Species Asiaticas novas descripsit I. H.

Burkill sequentes. Ibid. 2: 309-327, 1906.

347

1907.

1908.

1909.

1910.

Gardens Bulletin, S.

On Swertia angustifolia, Ham., and its Allies. Ibid. 2: 363-—

381, 1906.

Notes on the pollination of flowers in India.

Note No. 1. The pollination of Thunbergia grandiflora,

Roxb., in Calcutta. Ibid. 2: 511-514. ff. 1-3, 1906.

Note No. 2. The pollination of Corchorus in Bengal

and Assam. Ibid. 2: 515-520, 1906.

Note No. 3. The mechanism of six flowers of the North-

West Himalaya. Ibid. 2: 521-525, t. 1, 1960.

Gossypium obtusifolium, Roxburgh. Mem. Agric. Dept.

India (Bot. ser.) 1 (4): 1-10, t. 1, 1906.

Psophocarpus tetragonolobus. (Goa Bean).—Goa beans

in India. Agric. Ledg. 4: 51-64, 1906.

Notes on the pollination of flowers in India.

Note No. 4. On cotton in Behar. Journ. & Proc. Asiat.

Soc. Bengal, n.s. 3: 517-526, 1907.

Alpine notes from Sikkim. Kew Bull. 1907: 92-94.

On Gentiana coronata, Royle. Journ. & Proc. Asiat. Soc.

Bengal. n.s. 3: 149-168, ff. 1-9, 1907.

Anguilicarpus—a new genus of the Cruciferae. Ibid. 3:

559-561, ff. 1-6, 1907.

A note on Impatiens balsamina, Linn., as a dye-plant.

Ibid. 3: 565-566, 1907.

Notes on the pollination of flowers in India.

Note No. 5. Some autumn observations in the Sikkim

Himalayas. Ibid. 4: 179-195, ff. 1-8, 1908.

Note No. 6. The Spring flora in the Simla Hills.

Ibid. 4: 197-231, ff. 1-16, 1908.

A Working List of the Flowering Plants of Baluchistan,

Calcutta.

On Coptis. Journ. & Proc. Asiat. Soc. Bengal n.s. 5: 73-—

88, ff. 1-20, 1909. :

First notes on Cymbopogon Martini, Stapf. Ibid. 5: 89-93,

ff. 1-6, 1909.

Notes on the pollination of flowers in India.

Note No. 7. A few observations made in the Central

Provinces and Behar. Ibid. 6: 101—107, ff. 1-3, 1910.

Notes from a journey to Nepal. Rec. Bot. Surv. India 4:

map, 59-140, 1910.

348

Volk, XV E1959).

1911. The polarity of the bulbils of Dioscorea bulbifera, Linn.

Journ. & Proc. Asiat. Soc. Bengal n.s. 7: 467-469, ff.

1-2, 1911.

Literature on the races of Rice in India. Compiled in the

Office of the Reporter on Economic Products to the

Government of India. Agric. Ledg. 16: 1-594, 1911.

Determination of the Prickly Pears now wild in India. Rec.

Bot. Surv. India 4: 287-322, map, 1911.

1916. Notes on the Pollination of flowers in India.

Note No. 8. Miscellanea. Journ. & Proc. Asiat. Soc.

Bengal n.s. 12: 239-265, ff. 1-10, 1916.

1924. The Botany of the Abor Expedition. Rec. Bot. Surv. India

10: 1-154, 1924; 155-420, tt. 1-10, 1925.

1954. Chapters on the History of Botany in India. I. From the

beginning to the middle of Wallich’s service. Journ.

Bombay Nat. Hist. Soc. 51: 846-878, 1954.

1956. Chapters ... II. The Advances, and in particular the plant

collecting, of the thirties and forties of the 19th century.

Ibid. 54: 42-86, 1956.

Burkill, I. H. & R. S. Finlow

1907. The Races of Jute. Agric. Ledg. 6: 41-137, f. 1, 1907.

On three varieties of Corchorus capsularis, Linn. which are

eaten. Journ. & Proc. Asiat. Soc. Bengal. n.s. 3: 633-—

638, ff. 1-7, 1907.

1911. Corchorus capsularis var. oocarpus—a new variety of the

common jute plant. Ibid. 7: 465-466, ff. 1-2, 1911.

Burkill, J. H. & D. Prain

1908. Dioscorearum novarum descriptiones quaedam. Ibid. 4:

447-457, 1908.

1914. A synopsis of the Dioscoreas of the Old World, Africa ex-

cluded, with descriptions of new species and varieties.

Ibid. 10: 5-41, 1914.

1919. Dioscorea sativa. Kew Bull. 1919: 339~375.

1926. Ad Dioscorearum orientalium Historias commentarii. Kew

Bull. 1926: 118-120.

1936. An Account of the genus Dioscorea. Ann. Roy. Bot. Gard.

Calcutta 14: 1-528, tt. 1-150, 1936 & 1938.

349

I. H. Burkill in Malaya

C. X. FURTADO AND R. E. HoLtTumM

ONLY A FEW years ago there was discussed in the Gardens’ Bulletin

the extensive pioneer work done by the late Mr. H. N. Ridley

not only in showing that Hevea brasiliensis could be grown in

Asia for the commercial production of rubber but also in laying

stable foundations for the development of the several scientific

departments for investigating the problems connected with fauna,

flora, agriculture and forestry of Malaya. In view of this great work

by Mr. Ridley, it might seem that his successor, Mr. Isaac Henry

Burkill, would have little to do but follow on established lines.

However, though we may not be able to do full justice to the sub-

ject, there are weighty reasons to consider the work of Mr. Burkill

quite unique in its way; and it was work carried out, like Ridley’s

under many difficulties.

When Ridley retired from the service in 1912 and even before

Burkill took the charge of the Gardens Department, the different

departments which Ridley had helped to found, dissociated their

connection with the Gardens’ Department in order to work entirely

as independent units in Malaya. Then the World War I (1914—

1918) rendered impossible the execution of several projects be-

cause of the difficulties of recruiting properly trained staff, also

because of lack of funds. Further, though the agricultural and the

forestry departments were mainly for the Malay States, they had

jurisdiction over the agricultural and forestry sections in the Straits

Settlements, while the authority of the Gardens’ Department was

restricted to the official gardens within the Straits Settlements, so

that the Gardens in the Malay States were placed under the

Malayan Agricultural Departments. Difficulties in the Straits

Settlements were mounting not only owing to lack of staff and

finance but also to lack of appreciation of the value of purely

botanical work; and perhaps this is the main reason why, soon

after World War I, the area in which the arboretum was estab-

lished and a good deal of acclimatisation work was being done on

fruit trees and other economic plants and where rubber industry

was born was taken away from the Gardens Department in order

to establish Raffles College—the precursor of the University of

Malaya, Faculties of Arts and Science.

350

Vol. XVII. (1959).

But Burkill came here with an experience in India, where at

the time, because of the limited availability of trained staff, one

had to use one’s skill to make a scientific undertaking a success.

A rapid survey must have shown to Burkill that, if the Gardens’

Department was to survive, it must become a centre for research

of systematic botany; and for this purpose the first thing needed

was to re-organise the whole herbarium and library so as to be-

come less dependent on the workers and herbaria in India and

England. Hitherto Ridley had worked in co-operation with the

different specialists in India, England, Italy and Germany. The

correspondence and his visits to Europe had helped him to des-

cribe his many new species and genera and write several papers.

This extensive work had not given him time to organise ade-

quately the herbarium and the library in Singapore; and perhaps

because of the danger of termites and other vermin important

specimens or types were sent to the workers in Kew, Calcutta and

elsewhere, so that Singapore lacked many of the types or authentic

material from Ridley’s own collections. Further, since many speci-

mens in the herbarium had no identifications written on their labels

even when sorted, they were liable to be misplaced in the course

of using the herbarium or mending the sheets. When working the

Flora of the Malay Peninsula IV (1922-25) Ridley used mainly

the duplicate specimens he had sent to Kew, so that most of the

specimens in the Singapore Herbarium still remained unlabelled.

Burkill therefore set himself immediately to the task of naming

the specimens and indicating appropriately those cited in the large

floristic works on Malaya, The Materials for a Flora of the

Malayan Peninsula: Dicotyledons Vols. I-V by King and later

assisted and continued by Gamble (1889-1915) and Monocoty-

ledons by Ridley, Vols. I-III (1907), and later in Ridley’s Flora of

the Malay Peninsula I-V (1922—25). Further he organised col-

lection trips to Malaya, Sumatra, Java and Sarawak in order to

enrich the Singapore herbarium and also to send duplicate speci-

mens to other institutions, so that such duplicates might be com-

pared with types or authentic material in other institutions. Thus

about 20,000 specimens were added by way of the collection in

the jungles, the duplicates of which were distributed to several

herbaria interested in Malaysian systematics and almost an equal

number was received in exchange or as gifts, in addition to acquir-

ing a set of Elmer’s collection in Borneo and the Philippines. Fur-

ther several hundreds of specimens of the plants cultivated in the

Gardens were made in order to form a herbarium of the Gardens’

plants.

351

Gardens Bulletin, S.

It was at this time that Burkill introduced the system of number-

ing the specimens in the field rather than in the herbarium; for

when numbered in the field, one can be sure that the different

specimens bearing the same number are from the same collection,

whereas, when numbered in the herbarium, specimens from dif-

ferent collections sometimes become mixed together so that the

duplicates bearing the same number may not be specifically or

varietally identical. The system of numbering the specimens in the

field seemed to be so interesting that many foreign botanists who

visited the Singapore herbarium often begged a specimen field label

to take away with them for adapting it to their conditions.

It was in the course of this re-organization of the herbarium and

the library that Burkill laid the foundation for his monumental

work, A Dictionary of the Economic Products of the Malay Penin-

sula 2 vols. 2402 pp., elaborated and published in 1935, after his

retirement in England. The work of re-organizing the herbarium

provided also vernacular names and some information which had

to be completed by making a further inquiry among the local peo-

ple and by consulting the reports of research workers in Malaya,

Indonesia, Philippines, India and elsewhere on the plants found in

Malaya. The results thus collated were no doubt later incorporated

in the Dictionary, but sometimes they formed the subject of special

papers in the Gardens’ Bulletin. Thus Burkill compiled a list of all

the persons who had contributed specimens to the Gardens’ Her-

barium with details of their travels, and this together with the

names of other persons who had contributed Malayan plants to

other herbaria in the world, formed the subject of a paper with

biographical sketches of the collectors published in the Gardens

Bulletin TV (1927) 113-202, with the following title: Botanical

Collectors, Collections and Collecting Places in the Malay Penin-

sula: A review of the work so far done towards a knowledge of

plantgeography of Malaya. An inquiry into the simples and crude

drugs of Malaya led Burkill to enlist the co-operation of the autho-

rities who could competently work out the collections or manus-

cripts studied by him. Thus the specimens of Chinese drugs col-

lected by Burkill were worked out by David Hooper and published

in Gardens’ Bulletin Vol. VI Part 1 (1929) 1-163. The prescrip-

tions collected from the Malays by Burkill and Mohammed Haniff

were issued as “Malay Village Medicine” the Malay names being

associated with the botanical names of the plants by Burkill him-

self, and issued in Gardens’ Bulletin VI (1930) 165-327. ‘The

manuscript book of “The Medical Book of Malayan Medicine’’,

translated by Inche Ismail Munshi, possibly in Penang circa 1886,

352

Volk. XVIT. (1959).

was edited by Burkill who gave the botanical determination of the

plants, while medical notes were supplied by J. D. Gimlette, who

had worked widely on the Malayan medical terminology. (Gard.

Bull. VI, 1930 pp. 323-474).

The undertaking to write an account about the Economic Pro-

ducts of Malaya led Burkill to make a survey of the efforts made

to acclimatize foreign plants in Malaya. This involved not only

culling information from the different published reports but also

noting and sorting out the old manuscript records of every plant

or seed that was introduced in the Botanic Gardens, which was

generally the place to which plants newly introduced into Malaya

were sent. The records were classified and made accessible in the

herbarium under their respective genus and thence were worked

out for the Dictionary.

The inquiry as to success or failure of the imtroduced plants

necessitated a census of all the plants that existed in the Botanic

Gardens and in its economic section which is now extinct. Thus

the herbarium of the plants in cultivation came into the existence

providing often valuable information of the success of plants that

were introduced and cultivated in Malaya. Since there were then

separate agricultural and forestry departments, the library ex-

changes and purchases had to be revised so as to make the institu-

tion more useful for its specialized purpose of the study of syste-

matic botany. At the same time the Gardens’ periodical which had

hitherto been The Agricultural Bulletin of the Straits and Fede-

rated Malay States, was continued with the changed title “The

Gardens’ Bulletin, Straits Settlements’. This change in the title

was also necessary since the Malayan Agricultural Department had

started a new periodical called “The Agricultural Bulletin of the

Federated Malay States’, causing thereby much confusion among

the foreign public who, at the time, knew very little of Malayan

geography and much less of Malayan political divisions.

The development of the Gardens’ Department as the systematic

centre for Malayan botany and the difficulties of enlisting the

services of the botanical institutions in India and England, then

suffering from shortage of staff, soon showed that several aspects

of research work in the Malayan departments concerned with

forestry, agriculture, rubber, malaria, food-stuffs of man and ani-

mals, and other subjects could not be carried on without making

frequent calls upon the services of the Gardens’ Department. An

attempt was therefore made to establish a separate systematic

research centre in Perak using Wray’s herbarium as the basis, but

353

Gardens Bulletin, S.

it proved a failure, because no systematist can provide the ser-

vices needed by research institutions without a properly organized

herbarium and library. Even police cases can be delayed or be

misdirected if plants suspected as poisons are not properly identi-

fied.

While continuing to co-operate, so far ‘as he was able, with the

Malayan Departments of Agriculture and Forestry, Burkill op-

posed proposals for union of the Botanic Gardens, S.S., with either

of these departments. He further insisted on the importance of a

herbarium being associated with a garden, where plants could

be studied in cultivation, as well as on the necessity of ample

opportunity for field study of native plants. When the Research

division of the Forestry Department was established, it became

evident that that Department was dependent on the herbarium

at the Botanic Gardens for a great amount of important informa-

tion concerning the forest plants of Malaya. Dr. F. W. Foxworthy,

the first Forest Research Officer, made frequent visits to Singapore,

and collaborated in a study of the seedlings of Dipterocarps which

Burkill had started: cf. Journ. Str. Br. Roy. Asiat. Soc. 75 (1917)

43-48; 76 (1917) 161-167; 79 (1918) 39-44; 81 (1920) 1-4

and 49-76; 86 (1922) 281-284 and 285-291; Journ. Malay. Br.

Roy. Asiat. Soc. I (1923) 218-222, and with Foxworthy in Journ.

Str. Br. Roy. Asiat. Soc. 86 (1922) 271-280.

A proposal was made for the moving of the headquarters of the

Gardens Department to Kuala Lumpur, in order that it might be

closely associated with the Forest Research Institute (with the

possibility of a common library of books and journals on plant

taxonomy), and about the time of Burkill’s retirement a committee

was appointed to make a recommendations to effect this change.

But, though there was agreement in principle as to the desirability

of the committee’s proposals, the Federal Government decided

that the cost of new buildings would be too great, and the matter

was shelved, later to be completely abandoned; and the Straits

Settlements continued to bear the whole cost of what was in effect

a Malayan botanical service. The effectiveness of this service was

increased during the last years of Burkill’s directorship by the

addition of two plant taxonomists and one graduate field officer

(Holttum, Henderson and Furtado respectively) and a library

assistant to the Gardens staff.

Though working under a great handicap, Burkill showed how

a trained botanist can use his time in making systematic and

biological observations on the many plants that are grown in a

354

Vol. XVII. (1959).

garden and also utilize the facilities available for growing plants

to clear certain doubts regarding their botanical affinities, dietetic

value, etc. Thus the first two or three volumes of the Gardens’

Bulletin contain a large number of notes regarding insect pests,

abnormalities, new varieties of orchids, flowering peculiarities of

many plants growing in the Gardens, digests of old records on

rubber and on the history of the Gardens in Malaya, and observa-

tions on the Yams (Dioscorea spp.) grown in the Gardens where

he had gathered them from different parts of the world in order to

prosecute his studies begun in Calcutta in the company of Dr.

(later Sir) David Prain. The following articles might be mentioned

since they show the keenness and the breadth of the studies: “The

Positions of the Agricultural Industries in the Straits Settlements

in 1913” (Gard. Bull. I, 1914 pp. 213-235); “The Treatmeni to

which the Para-Rubber Trees of the Botanic Gardens, Singapore,

have been subjected’ (Gard. Bull. I, 1915, pp. 248-295);

“Gordonia’ (Journ. Str. Br. Roy. Asiat. Soc. 76 (1917) 133-

159); “The Establishment of the Botanic Gardens, Singapore”

(Gard. Bull. I, 1918, pp. 55-72); “The Second Phase in the

History of the Botanic Gardens, Singapore” (Gard. Bull. II, 1918

pp. 93-108); “The Composition of a Piece of Well-drained Sin-

gapore Secondary Jungle Thirty Years Old’ (Gard. Bull. I, 1919

pp. 145-157); “Some Notes on the Pollination of Flowers in the

Botanic Gardens, Singapore, and in other Parts of the Peninsula”

(Gard. Bull. II, 1919 pp. 165-176); “The Genus Gordonia in

the Philippine Islands” (Philipp. Journ. Sci. XV, 1919 pp. 475-

478); “A List of Oriental Vernacular Names of the Genus Dios-

corea” (Gard. Bull. II, 1924 pp. 121-244).

However Burkill was only for a short space of 13 years in

Malaya; and during this period he was busy collecting material for

his monumental work of “A Dictionary of the Economic Products

of the Malay Peninsula’. Naturally these notes and observations

had to be worked out in England where he had an access to better

library and herbarium facilities and whence he helped, after his

retirement, the Gardens’ Herbarium to name many of the foreign

and other plants that were grown in the Gardens. Each of his

articles in the Dictionary is replete with information about the his-

tory of the plant, its products, vernacular names, uses, chemical

composition of the drug or poison if known, etc. In addition to

work on the Dictionary, other studies begun in Singapore or Cal-

cutta were completed in England. Thus “An Account of the genus

Dioscorea in the East’ was published by Prain and Burkill in four

355

Gardens Bulletin, S.

large volumes (two of which are plates), in the Annals of the

Botanic Gardens, Calcutta in 1937 and 1939. The following pa-

pers might also be mentioned: “The Chinese Mustards in the

Malay Peninsula’ (Gard. Bull. V, 1930 pp. 99-117); “An Enu-

meration of the Species of Paramignya, Atalantia and Citrus,

found in Malaya’ (Gard. Bull. V, 1931 pp. 212-223); “Notes on

Gluta”’ (Gard. Bull. V, 1931 pp. 224-230); “The rise and decline

of the Greater Yam in the service of man’ (Avance. Sci. Lond. VII

1950 pp. 443-448).

Also it may be noted that in addition to many species, at least

three genera have been named in honour of Mr. I. H. Burkill,

namely, Burkillia, West and West, Protococcaceae (1907), Bur-

killia Ridley, Leguminoseae (1925) and Burkillianthus Swingle,

Rutaceae (1938).

356

Phellodendron, a genus of trees new to the

Malayan flora (Rutaceae )

By C. G. G. J. vaN STEENIS

Flora Malesiana Foundation, Leyden

FoR SOME TIME I have been defeated in identifying a sheet from

the Malay Peninsula with decussate, pinnate, exstipular leaves and

choripetalous, 5—merous, male flowers. It would seem that the

choice was so much narrowed by these characters, that identifi-

cation would have been easy. And though it is now certain, that

it belongs to Phellodendron of the Rutaceae, it remains still re-

markable that the leaves of this genus do not show the distinct

pellucid glands which are so typical for rutaceous plants. There

are indeed minute, pin-like pellucid glands (crystal cells?) in

the parenchyma but of essentially. smaller size than is usual in

Rutaceae. It is true that Sprague (Kew Bull. 1920, 231) and

Engler (Pfl. Fam. ed. 2, 19a 1931, 298) mention the occurrence

of a large pellucid gland near each of the shallow leaf-crenations,

but this appears to be frequently indistinct in the sheets of the genus

preserved in the Rijksherbarium. See also Blenk, Flora 67

~ (1884) 278.

Phellodendron burkillii n.sp—Fig. 1.

Arbor, 18 m. alta, c. 35 cm. diam. Ramuli, foliorum raches,

petioluli et inflorescentiae axes laterales primarii appresse pube-

ruli. Folia 4—6-jugata, foliolis oblique ovato-oblongis ad sublanceo-

latis, acuminatis, acumine longo angusto saepe falcato, 6-10 x

2-3 cm., subtus in costae parte inferiore puberulis, ceterum glabris.

Corymbus axibus conspicue decussatis, multiflorus, c. 10-12 cm.

longus, 15—20 cm. latus. Flores subglabri. Calycis lobi. 1/3 mm.,

triangulares. Petala c. 23 * 1 mm., intus ad venam medianam

pilis paucis retrorsis suffulta. Filamenta in parte inferiore pilis lon-

gis patentibus munita. Ovarium glabrum. Styli pilis erectis parcis-

sime pubescentes.

Typus: KEP 78904, L.

Tree 18 m. tall, 35 cm. diam. Twigs with very few lenticels,

pithy, as the petioles, leaf-rachis, and inflorescential stalks puberu-

lous, terete, but flattened at the nodes. Leaves decussate, with

357

Gardens Bulletin, S.

4—6 pairs of leaflets, c. 20-30 cm. long; leaflets c. 4—6 pairs, dull,

+glaucous underneath, obliquely ovate-oblong to sublanceolate,

long- and narrow-, often falcate-acuminate, 6-10 by 2-3 cm,

base cuneate and on the acroscopical side often + rounded, ulti-

mate base short-decurrent into the petiolule; nerves c. 6—8 pairs,

inconspicuous, as the very dense venation not or obscurely pro-

minent; margin obsoletely crenate, in the crenations often an

obsolete, pellucid dot; petiole 3-6 cm; petiolules c. 3 mm. Termi-

nal corymb wide (c. 15-20 cm.), c. 10-12 cm. high, richly branch-

ed and with numerous flowers. Calyx c. 1 mm. diam., 4 mm. high,

the lobes acute-deltoid. Mature buds broad-elliptic almost globu-

lar, 2 mm. long. Petals oblong-elliptic, the margins very narrowly

imbricate, c. 24 times as long as wide, boat-shaped, c, 2% by 1

mm., tip thickened, retrorse, glabrous but for some appressed,

retrorse hairs on the midrib within. Stamens 5, erect in bud, the

filaments considerably lengthening during anthesis and becoming

longer than the anthers (c. 2-24 mm.), the lower half with scat-

tered, patent, long hairs; anthers free in the lower half and +

Sagittate, basifixed, acute-oblong, c. 1? mm. long. Ovary in 4

rudimentary, 5-angled, c. 4 mm. high, glabrous, with 5 erect, ap-

pressed styles 4 mm. high, with a few appressed-erect hairs.

MaLay PENINSULA. Kedah: Enggang Forest Reserve, Sik, KEP

78904 (L, type, KEP, K, SING, distributed as Micromelum pu-

bescens BI.), low-lying ground, bark smooth, grey-green, flowers

greenish-white, July 12, 1956.

The closest ally of this species seems to be P. macrophyllum

Dode from East Szechuan, China (Bull. Soc. Bot. Fr. 55, 1908,

648) which is, however, clearly specifically distinct, as appeared

from an examination of the type which was kindly loaned to me

by the Director of the Paris Herbarium. The differential characters

are that it has 2—4-jugate leaves (in the type sheet 2-jugate);

leaflets large, 10-15 (—20) by 5-8 (9) cm., broad-elliptic, ab-

ruptly acuminate, pubescent beneath on the main rib, venation

and intervenium, with a much coarser venation than in P.

burkillii, without marginal glandular dots, but with very numerous

dots of microscopic size and scattered large pellucid dots (as yet

unknown in the genus); inflorescence with very thick, lenticellate

dark-coloured peduncle and axis; flowers very dense; mature buds

3 mm.; calyx 1 mm. high. Petals 3 by 1 1/3 mm. Styles patent

long-villose. Fruit globular, 10 by 9 mm.

The genus is a truly East Asian—one of Sino-Japanese distri-

bution, north to Manchuria, south to Formosa.

358

WE oz

flower.

aes SD,

flower. f, stamen. g, pistil, all «

Flowering twig,

all x 6 (after KEP 78904, L.).

359

Vol. XVII. (1959).

ave: fil

as ark

‘ hes rol

ROO REA

6 (after Farges 1284, P.).

c, stamen. d, pistil,

~—Phellodendron macrophyllum Dode. e,

Phellodendron burkillii Steen. a,

Gardens Bulletin, S.

If such temperate to subtemperate ligneous genera occur in

the South-east Asian tropics they are generally found in the mon-

tane zone, as for example in Pentaphylax, Osmanthus, Camellia,

Anneslea, Distylium, Schoepfia, Pistacia (in Luzon), etc. This is

not always the case, as of Acer, which belongs in this group, two |

lowland localities have been recorded in addition to dozens in the

montane zone.

There is as far as I know one temperate to subtemperate ligne-

cus genus which only occurs in the lowland in Malaya and that is

Pistacia malayana Henderson; this shows an ecologically similar

behaviour as this new record of Phellodendron.

360

Vegetative characters distinguishing the various

groups of ferns included in Dryopteris of

Christensen’s Index Filicum, and other

| ferns of similar habit and sori

By R. E. HOLTTUM

IN A SERIES of papers begining in 1907, mainly on American spe-

cies, Christensen himself showed a way in which natural groups

within Dryopteris sens. lat. of his Index Filicum could be distin-

guished, the significant diagnostic features being provided by hairs

and scales on the fronds. Though he never published a formal con-

spectus of the whole group, he continued to develop his ideas

(see Christensen 1938). Ching’s series of papers on Dryopteris and

other genera of China, published from the year 1933 onwards,

carried the general investigation much further, dealing with the

ferns of Asia not considered in Christensen’s earlier work. Ching

also used the characters (both external and internal) shown by a

cross-section of the rachis, and the relation of rachis to pinna.

The use of this last kind of character was developed further by me,

and is most fully set forth, with some illustrations, in my book

on the ferns of Malaya (1954). In trying to understand more

clearly the limits of some of the genera concerned, I have been

much impressed by the significance of these characters, and to

clarify my thinking I have attempted a concise comparative pre-

sentation of them, as set forth at the end of the present paper.

It appears to me that Ching did not pursue effectively the line of

thought which he himself indicated when showing the distinction

between rachis-characters of Ctenitis and Dryopteris (sens. strict.) ;

if he had done so, he would not have suggested that Athyrium and

Thelypteris are closely related, nor that Rumohra adiantiformis

belongs to the same genus as the Dryopteroid ferns which he as-

sociated with it. |

I have been especially impressed by the importance of the

external shape of the rachis, and the way in which this shape is

modified when a pinna-rachis (or a simple pinna) is attached to it.

These characters are extremely uniform in genera which, as judged

by other characters, are seen to be natural ones, and I believe

that all the more highly evolved genera have each a particular

rachis-form. When however one looks at Dennstaedtia, Hypolepis

361

Gardens Bulletin, S.

and related genera. which have various primitive characters and

seem to be survivors of a very basic group, one finds less uni-

formity, and I believe that in this group one can see the beginnings

of both the Tectaria and Dryopteris rachis-characters, now so

strongly differentiated in these two more highly evolved genera.

I believe that the limits of the genera Dennstaedtia and Hypolepis

are not at present clearly defined, and that a further study of these

genera, with rachis-characters (as well as other factors) in mind,

may yield significant results.

There is a sharp distinction between the kind of grooved rachis

shown by Dryopteris proper (the groove opened to admit the

groove of a pinna-rachis) and on the other hand the kind of

rachis which, even if it has a median groove, does not open its

groove to admit a pinna-rachis groove. In ferns showing the

second of these alternatives, as for example the Thelypteroid ferns,

the main rachis and the costae of pinnae are grooved on the upper

surface, but the groove of the rachis does not open to admit the

groove the costa (the latter is in fact usually not grooved at its

base). In Tectaria and Ctenitis there is no median groove in the

upper surface of rachis or of costae of pinnae, though in Pleocnem-

ia (Holttum 1951) there is a slight median groove. Ctenitis and

Tectaria are characterized also by the short multicellular hairs

pointed out by Christensen (he called them Ctenitis-hairs). But

there are genera which have the rachis-form of Dryopteris and

hairs not much different from those of Ctenitis (e.g. Stenolepia

v.A.V.R.).

A quite distinct kind of rachis is found in Davallia and some

other genera. Here the middle of the upper surface of the rachis

is raised, and on either side of this raised median band, in all

smaller rachises, is a slight groove, the outer edge of the groove

forming a very firm narrow wing; the thickened edge of the lamina

of a leaflet is decurrent on this wing, the raised midrib of the

leaflet joining the side of the raised median band of the upper

surface of the rachis. An almost exactly similar arrangement

occurs in Pteridrys (Ching & C.Chr. 1934), but in rhizome this

genus is very different from Davallia. An indication of how this

arrangement of the rachis may develop in the Tectaria group of

genera is given by Lastreopsis Ching (united with Ctenitis by

Copeland; recently distinguished, with a list of species, by Miss

Tindale). In Lastreopsis the thickened lower edge of the lamina

of a leaflet is decurrent on the rachis-wing, but this wing is usually

much more lateral on the rachis than in Pteridrys.

The Davallia type of rachis is shown by Rumohra adiantiformis

(Forst.) Ching, and this same species also has a dorsiventral

362

Vol. XVII. (1959).

rhizome of Davallia type, so that a sterile plant of R. adiantiformis

might well be identified as Davallia. The spores of this species also

are like those of Davallia, not like those of Dryopteris and Polys-

tichum (Harris 1955, pl. 7, f. 2). It seems to me therefore proved

that Ching and Copeland were mistaken in associating species

allied to Dryopteris and Polystichum with Rumohra. For these

latter species, I have used the name Polystichopsis, but I am not

at all satisfied that they form a natural group, and I believe that

the matter needs further investigation. Furthermore, Mr. C. V.

Morton has pointed out to me that the generic name Polystichopsis

was first used in a different sense, and he is elsewhere proposing

another name.

In the following conspectus I have not attempted to distinguish

the genera of the Thelypteroid group, because I believe that more

study is needed to establish clearly their distinctive characters.

Everybody admits that a distinction between Thelypteris (or Las-

trea) and Cyclosorus on the characters of free or anastomosing

veins is a quite unnatural one. One could include all species in one

genus, but then subgenera would need to be distinguished. For ex-

ample, Lastrea oreopteris is surely very different from Thelypteris

palustris, and both are very different from the group of species to

which Thelypteris brunnea (Wall.) Ching belongs. Cytological

evidence (Manton, Manton & Sledge) indicates that some species

with free veins are more closely related to Cyclosorus than to other

free-veined species, and probably such species should be united to

Cyclosorus; but other morphological characters need to be

established.

As regards the genera of the Tectaria group, it seems to me

that Ctenitopsis Ching should be united to Tectaria, not to Ctenitis

as in Copeland’s Genera Filicum. Just as a division between

Thelypteris and Cyclosorus on whether veins are free or not in

unnatural, so also is a division between Tectaria and Ctenitopsis.

The series of forms included in Dryopteris dissecta (Forst.) O.

Ktze, which belong to Ctenitopsis, have exactly the frond-form of

some species of Tectaria. Perhaps Ctenitis itself needs further sub-

division, allowing some sections of it to be associated with Tec-

taria. I formerly attempted to distinguish Heterogonium from

Tectaria on frond-form (I see no other practicable distinction) but

now I am not sure whether this group should have generic separa-

tion. Aspidium boryanum Willd. (included by Copeland in Cteni-

tis) resembles Ctenitis rather than Dryopteris in external form of

the rachis, but differs from both in the vascular anatomy of the

stipe. I believe that this species (with some others which are closely

363

Gardens Bulletin, S.

related to it) is nearer to Athyrium than to Ctenitis, but quite dis-

tinct from Athyrium, in which genus I placed it in my book on the

ferns of Malaya (p. 567); in so doing I unfortunately made the

mistake of referring to it specimens which are in fact Athyrium.

Later (1959) I proposed a new generic name Parathyrium for

Aspidium boryanum Willd. and its allies, but this was antedated

by Dryoathyrium Ching (1941), based on the same type species.

In the following conspectus, Gymnocarpium (sensu Ching

1933) is shown as belonging to the Dryopteris group, not with

Thelypteris (Lastrea) where Copeland placed it (1947, p. 137),

and cytological evidence confirms this position. Dryopteris phegop-

teris (L.) Chr., on the other hand, belongs to the Thelypteris

group. I believe that Copeland overlooked both hairs and rachis-

characters when he wrote of these two that “their separation by

reasonable generic definition is hardly possible”. Similarly, I be-

lieve that the conspectus shows a clear separation between Athy-

rium and the Thelypteris group (on hairs, scales and rachis-

characters), though both Ching and Copeland held that they were

so closely allied as to be hardly separable in formal terms.

Cornopteris Nakai was based on a species like Diplazium and

Athyrium in rachis-characters but lacking an indusium. Hemionitis

opaca Don (in Dryopteris in the original Index Filicum of Chris-

tensen, transferred later to Diplazium by Christ) is very closely

related to the type species of Cornopteris, and other species have

been noted by Ching. Madame Tardieu-Blot however has added to

the genus Cornopteris some of the species of the alliance of Aspi-

dium boryanum Willd. (1956, 1958); the distinction between true

Cornopteris and these species is shown in the present conspectus.

The genus Hypodematium has abundant needle-like unicellular

hairs on its fronds, resembling in this the Thelypteroid ferns. Its

scales however are quite different from those of Thelypteris, and

I do not think that Hypodematium is of the Thelypteris alliance;

on the other hand, I am not sure that it really belongs with the

Tectaria group of genera.

Rhizome not dorsiventral (leaves and roots equally all round);

wing of pinna-rachis, if present, thin like the lamina, except

in Pteridrys and Lastreopsis

Main rachis lacking a median groove on upper surface; or if

grooved, the groove not open to admit grooves of pinna-

rachises,; rachis sometimes winged, the wing joining the

edge of the lamina of leaflets, not the edge of the costa of a

leaflet

364

= ee

Vol. XVII. (1959).

Hairs on upper surface of rachis multiseptate, usually very

short; scales lacking superficial hairs or glands

Several separate vascular bundles throughout the stipe

Tectaria group

A tooth present at the base of each sinus between lobes

of the lamina

Veins free; vascular bundles in stipe in a simple ring

Pteridrys

Veins anastomosing; vascular bundles in more complex

arrangement Pleocnemia

Arcypteris

No tooth present in sinus between two lobes of lamina

Basal basiscopic vein of a vein-group springing directly

from the costa, not from the costule which bears

the other veins of the group; or veins much anas-

tomosing Tectaria

(incl. Ctenitopsis )

Basal basiscopic vein of a vein-group springing from

the costule; veins always free

Indusium peltate; hairs on rachis few, mainly near

junction of rachis and pinnae on upper surface

Dryopolystichum

Indusium reniform; hairs many throughout rachis

Decurrent wing of lamina not thickened

Ctenitis

Decurrent wing of lamina thickened

, Lastreopsis

One U-shaped bundle in the stipe (2 bundles near base)

Dryoathyrium

Hairs on upper surface of rachis unicellular, more or less

needle-like; two vascular bundles in base of stipe, uniting

upwards to one of U-shape

Scales lacking superficial hairs or glands, confined to swol-

len bases of stipes Hypodematium

Scales bearing superficial glands or hairs, not confined to

base of stipe which is not swollen Thelypteris group

Main rachis with median groove in upper surface, this groove

open to admit grooves of pinna-rachises; raised edge of a

pinna-rachis groove joined to side of pinnule-costa groove,

edge of pinnule-lamina decurrent on pinna-rachis as a late-

ral wing

365

Gardens Bulletin, S.

Multicellular hairs, of a kind quite different from scales,

lacking; groove of rachis quite smooth within

Dryopteris-group

Several vascular bundles in stipe Dryopteris

Polystichum

“Polystichopsis”

Two vascular bundles in stipe Gymnocarpium

Multicellular hairs, quite different from scales, often present;

groove of rachis papillate within; some sori elongate and

asymmetric or unilateral; two vascular bundles at base

of stipe Athyrium group

Sori indusiate Athyrium

Sori not indusiate Cornopteris

Rhizome dorsiventral (roots from lower surface, leaves from upper

surface); pinna-rachis bearing a firm wing, edge of lamina

thickened at base and decurrent on the rachis-wing; midrib

of pinna raised, not grooved Rumohra (s. str.)

Literature Cited

CHING, R. C. 1933. On the nomenclature and systematic position

of Polypodium dryopteris L. and related species.

Contr. Biol. Lab. Sc. Soc. China 9: 30-42.

——-——-——— 1934. A revision of the compound-leaved Polysticha

and other related species in continental Asia includ-

ing Japan and Formosa. Sinensia 5: 23-71.

-—_——_—-— 1936. A revision of the Chinese and Sikkim-

Himalayan Dryopteris with reference to some species

of neighbouring regions, I. Bull. Fan Mem. Inst. Bot.

6: 237-332)

—-—----———— 1938. Ibid., Hl. lic. 8: 275-507.

—_—___—---—— 194]. The studios of Chinese Ferns XXXV. Lc. 11:

79-82.

CHING, R. C. & C. CHRISTENSEN, 1934. Pteridrys, a new fern genus

from tropical Asia. Bull. Fan Mem. Inst. Bot. 5:

125-148.

CHRISTENSEN, C. 1907. Revision of the American species of Dry-

opteris of the group of D. opposita. Vid. Selsk. Skr.

VII. 4.

————- 1911. On a natural classification of the species of

Dryopteris. Buiologiska Arbejder tilegnede Eug.

Warming. Copenhagen.

366

Vol. XVII. (1959).

—_——_———. 1913. A monograph of the genus Dryopteris, pt I.

Vid. Selsk. Skr. VII. 10.

1920. Ibid. pt II, lc. VIL. 6.

—_—_——_—_~—¥_1938. Filicinae (Ch. XX), in Verdoorn, Manual of

Pteridology. The Hague.

COPELAND, E. B. 1947. Genera Filicum. Waltham, Mass.

Harris, W. F. 1955. A manual of the spores of New Zealand Pte-

ridophyta. N.Z. Dept. of Sci. & Ind. Res. Bull. 116.

Hoittum, R. E. 1949. The fern-genus Heterogonium. Sarawak

Mus. Journ. 5: 156-166.

—_—_—_—-———- 1950. Further notes on the fern-genus Heterogo-

nium. Reinwardtia 1: 27-31.

1951. The fern-genus Pleocnemia. Reinwardtia 1:

171-189.

———— 1954. A Revised Flora of Malaya, II, Ferns of

Malaya, Singapore.

1955. Some additional species of Heterogonium.

Reinwardtia 3: 269-274.

1959. Notes on Malaysian Ferns. Kew Bull. no. 3,

1958, 447-455.

Manton, I. 1954. Appendix to Holttum, Ferns of Malaya.

MANTON, I. AND W. A. SLEDGE. 1954. Observations on the cyto-

logy and taxonomy of the Pteridophyte flora of Cey-

lon. Philos. Trans. Roy. Soc. London, ser. B, 238:

127-185.

TARDIEU-BLOT, M. L. 1956. Sur le genre Cornopteris Nakai et les

Cornopteris Malgaches. Mem. Inst. Sci. Madag. B,

7: 27-32.

TINDALE, M. L. 1957. A preliminary revision of the genus Las-

treopsis Ching. Vict. Nat. 73: 180-185.

367

Taxonomic Notes on Ficus Linn., Asia

and Australasia

I. SUBGEN. UROSTIGMA (GASP.) MIO.

By E. J. H. CORNER

Botany School, University of Cambridge

Summary

NEW SECTIONS, subsections, series and subseries.—sect. Urostigma

ser. Superbae Corner, ser. Caulobotryeae (Miq.) Corner, ser.

Orthoneurae Corner.—sect. Leucogyne Corner.—sect. Conosycea

(Miq.) Corner subsect. Conosycea ser. Drupaceae Corner, sub-

ser. Indicae Corner, subser. Zygotricheae Corner, subser. Crassira-

meae Corner; subsect. Dictyoneuron Corner ser. Glaberrimae

Corner, ser. Dubiae Corner, ser. Subvalidae (Miq.) Corner, ser.

Perforatae Corner; subsect. Benjamina (Miq.) Corner ser. Callo-

phylleae Corner; sect. Malvanthera Corner ser. Malvanthereae

Corner, subser. Eubracteatae Corner, subser. Platypodeae Corner,

subser. Hesperidiiformes Corner, ser. Cyclanthereae Corner.—sect.

Galoglychia (Gasp.) Endl. (subgen. Bibracteatae Mildbr. et

Burr. ).

Earlier specific epithets—F. virens Ait. (F. infectoria sensu

Roxb., F. lacur auct., non verae), F. costata Ait. (F. caudiculata

Trim.), F. depressa Bl. (F. pruniformis Bl.), F. drupacea Thunb.

(F. pilosa Reinw.), F. crassiramea Mig. (F. procera auct., non

Bl.), F. sundaica Bl. (F. indica sensu King, non Linn. which is

reduced to F. benghalensis Linn.), F. retusa Linn. (F. truncata

Miq.), F. pellucido-punctata Griff. (F. gelderi Miq.), F. piso-

carpa Bl. (F. microstoma Wall. ex King), F. subcordata Bl. (F.

garciniifolia Miq), F. microcarpa Linn. f. (F. retusa, F. nitida, F.

retusa var. nitida auct., non verae).

New names for later homonyms.—F. hookeriana Corner (F.

hookeri Miq), F. subtecta Corner (F. procera Bl.), F. maclel-

landi King var. rhododendrifolia Corner (F. rhododendrifolia

Miq.), F. curtipes Corner (F. obtusifolia Roxb.), F. subpuberula

Corner (F. puberula Miq.).

New species.—F. subgelderi Corner et v. rigida Corner, F. para-

camptophylla Corner, F. delosyce Corner et var. obtusa Corner,

F. spathulifolia Corner et var. annamensis Corner, v. substipitata

368

Vol. XVII. (1959).

Corner, F. calcicola Corner, F. episima Corner, F. tristaniifolia

Corner, F. palaquiifolia Corner, F. polygramma Corner, F. ben-

jaminoides Corner, F.. patellata Corner, F. triradiata Corner et var.

sessilicarpa Corner, F. augusta Corner.

New varieties.—F. superba Mig. v. alongensis (Gagnep.) Cor-

ner, v. henneana (Miq) Corner; F. concinna Mig. v. subsessilis

Corner; F. virens Ait. v. sublanceolata (Miq.) Corner, v. glabella

(Bl.) Corner; F. caulocarpa Migq. v. dasycarpa Corner; F. prolixa

Forst. f. v. subcordata Corner; F. arnottiana Mig. v. subcostata

Corner; F. drupacea Thunb. v. auranticarpa (Elm.) Corner, v.

glabrata Corner, v. pedicellata Corner, v. pubescens (Roth) Cor-

ner (F. mysorensis Heyne); F. cucurbitina King v. eubracteata

Corner; F. cordatula Merr. v. sericea (C. B. Robinson) Corner;

F. pubilimba Merr. v. ovata Corner; F. stupenda Miq. v. minor

Corner; F. crassiramea Miq. v. brevicupulata Corner, v. celebica

Corner v. clementis (Merr.) Corner, v. patellifera (Warb.) Cor-

ner; F. forstenii Miq. v. pacifica (Elm.) Corner, v. umbobracteata

(Elm.) Corner, v. villosa Corner; F. glaberrima Bl. v. bracteata

Corner, v. siamensis Corner; F. sundaica Bl. v. beccariana (King)

Corner; F. lowii King v. borneensis Corner, v. minor Corner; F.

sumatrana Miq. v. circumscissa Corner, v. microsyce Corner, V.

subsumatrana (Gagnep.) Corner; F. retusa Linn. v. borneensis

Corner; F. binnendykii Miq. v. coriacea Corner, v. cupulata Cor-

ner, v. latifolia Corner; F. subcordata Bl. v. malayana Corner;

F. benjamina Linn. v. bracteata Corner; F. callophylla Bl. v.

leytensis Corner, v. malayana Corner, v. minor Corner; F. mic-

rocarpa Linn. f. v. microcarpa f. pubescens Corner, v. eubracteata

Corner, v. hillii (Bailey) Corner, v. latifolia (Miq.) Corner, v.

naumanni (Engl.) Corner, v. saffordii (Merr.) Corner; F. glandi-

fera Summerh. v. brachysyce Corner; F. leucotricha Mig. v. mega-

carpa F.v.M. ex Corner, v. sessilis Corner; F. obliqua Forst. f. v.

petiolaris (Benth.) Corner, v. puberula (Benth.) Corner; F. hes-

peridiiformis King v. myrmekiocarpa (Summerh.) Corner; F.

xylosycia Diels v. cylindrocarpa (Diels) Corner.

Mixtae compositae.—F. caulobotrya Miq., F. onusta Wall. ex

Mig. (see under F. globosa Bl.).

‘In recent works on the classification of Ficus insufficient atten-

tion has been given to the names of subdivisions made by Miquel.

These must be typified and re-introduced with minimum of dis-

turbance, but a pre-requisite is a full understanding of all his

369

Gardens Bulletin, S.

species. The last to attempt this was King (1887-88), and he was

obliged to leave many uncertain. Since 1950 I have worked on the

revision of the Asiatic and Australasian species, using the large

collection of the Singapore Herbarium as the study-collection, and,

with the indispensable assistance of the herbaria of Leiden, Utrecht,

Bogor, and Kew (for many of Miquel’s types are in the Hooker

Herbarium), I have succeeded in recognising every taxon described

by Miquel from forma to genus. This series of papers gives the

bare taxonomic results: botanical appreciation must await full

publication. I should add that, until one can identify leaves in

particular from microscopic structure, it is commonly impossible

to identify many types. Hence I acknowledge with gratitude my

apprenticeship through the Singapore herbarium and my progress

through the courtesy of the Directors and staff of the herbaria of

Europe, Asia, New Guinea, and Australia, namely of the British

Museum, Kew, Edinburgh, Paris, Florence, Leiden, Utrecht,

Copenhagen, Berlin, Calcutta, Bogor, Manila, Lae, Brisbane, and

Adelaide. Thus I have studied over twenty thousand collections,

not counting the abundant duplicates. Further, in special matters,

I have received great help from the herbaria of Stockholm, Up-

psala, Geneva, Turin, Caen, the Arnold Arboretum, the New

York Botanic Garden, and the Smithsonian Institution. The funds

for this work have come mainly from the Cambridge Botany

School, but also from the Leverhulme Trust, the Nuffield Founda-

tion, and the Royal Society. To all these persons and institutions,

and to the many field-botanists who have joined the quest, I

express my gratitude. Taxonomy, of course, is a means to an end,

and what I have in mind is the evolution of tropical forest as seen

through one of its major constituents.

subgen. Urostigma (Gasp.) Mig.

Ann. Mus. Bot. Lugd. Bat. 3 (1867) 285.—Urostigma Gasp.

Parl. Giorn. Bot. 2 (1844) 214; Ann. Sci Nat. ser. 3, 3 (1845)

343.—sect. Urostigma (Gasp.) Endl. Gen. Pl. 4, 2 (1847) 35.

Lectotype—F. religiosa inn.

Gasparrini listed seven species without indication of a type.

Endlicher, in reducing the genus to a section, cited merely F. reli-

giosa Linn. and F. glaucophylla Desf., which had not been men-

tioned by Gasparrini. Hence I take F. religiosa as the lectotype

and it has always served as a typical member of the taxon

Urostigma.

370

Vol. XVII. (1959).

sect. Urostigma.—Figs with interleafing apical bracts. Male flow-

ers generally ostiolar. Ovaries entirely red-brown or in the upper

half. Cystoliths only on the lower side of the lamina. Petiole

often articulate to the lamina. Lectotype—F. religiosa Linn.

I divide this into four series.

1. ser. Religiosae Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867)

287.—Urostigma Gasp. ser. Religiosa Miq. Fl. Ind. Bat. 1, 2

(1859) 332.—Fig sessile (except F. verruculosa Warb., Africa):

basal bracts persistent: internal bristles none, or few and minute.

Lamina without hypodermis. Type—F. religiosa Linn.

2. ser. Superbae Corner ser. nov.—Receptacula pedunculata

(raro sessilia), bracteis basalibus caducis, setis internis nullis v.

paucis minutisque. Lamina hypodermide deficiens. Typus—F.

superba Miq.

3. ser. Caulobotryae (Mig.) Corner comb. nov.—Urostigama

Gasp. ser. Caulobotrya Mig. Fl. Ind. Bat. 1, 2 (1859) 334.—

Fig sessile or pedunculate: basal bracts persistent: internal bristles

abundant, chaffy-vesicular, white. Lamina without hypodermis.

Lectotype—F. caulocarpa Miq.

Miquel gave Urostigma superbum Mia., U. caulocarpum Miq.,

and U. stipulosum Miq. for this series. The last two are the same

and the first I exclude as ser. Superbae.

4. ser. Orthoneurae Corner ser. nov.—Receptacula sessilia v.

pedunculata, bracteis basalibus persistentibus, setis internis nullis.

Flores masculi dispersi. Petiolus non v. vix articulatus. Lamina

superne hypodermide unistratoso, cuticula inferiori gyroso-plicata,

praedita. Typus—F. orthoneura.Lévl. et Vant.

sect. Leucogyne Corner sect. nov.—Receptacula sessilia, bracteis

apicalibus interpositis. Flores masculi dispersi. Ovaria albida,

haud rubromaculata. Lamina superne hypodermide 1—2-stra-

toso, cystolithis solum pagina superiori, praedita. Petiolus haud

v. vix articulatus. Typus—F. rumphii BI.

This section includes F. rumphii, so often mistaken for F. reli-

giosa, and F. tsiela Roxb., which is close. Possibly F. menabeensis

H. Perr. (Madagascar) belongs here, but I am unable to place this

species satisfactorily. |

sect. Conosycea (Mig.) Corner comb. nov.—Urostigama Gasp.

sect. Conosycea Mig. Fl. Ind. Bat. 1, 2 (1859) 349.—Figs ses-

sile or pedunculate, apical bracts interleafing. Male flowers dis-

perse. Ovaries white with a red mark at the base, at least on the

stylar side, or the lower half red. Petiole not articulate. Lamina

af!

Gardens Bulletin, S.

generally with an upper hypodermis, cystoliths on both sides or

only on the upperside, rarely none. Lectotype—F. annulata BI.

This section was based on eleven species, namely U. altissimum

(BL) Miq., U. excelsum Miq. (= F. tinctoria Forst. f., subgen.

Ficus sect Sycidium Migq.), U. bicorne Mig. (= F. drupacea

Thunb.), U. conocarpum Mig. (= F. annulata Bl.), U. pilosum

(Reinw.) Mig. (= F. drupacea Thunb.), U. giganteum Mig. (=

F. stupenda Miq.), U. pruniforme (Bl.) Mig. (= F. depressa

Bl.), U. depressum (Bl.) Mig., U. benghalense (Linn.) Miq., U.

annulatum (Bl.) Migq., and U. xylophyllum Migq. I select F. annu-

lata Bl. as the lectotype because it is named twice, because its close

ally F. depressa Bl. is named twice, because it is well-known and a

good representative of this major section, and because it has the

conoid fig. Miquel did not use the section later. I recognise three

subsections with several series and subseries, as follows.

A. subsect. Conosycea.—Venation with intercostals. Type—

F. annulata Bl.

1. ser. Validae Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 285.—

Urostigma Gasp. ser. Valida Migq. FI. Ind. Bat. 1, 2 (1859) 334.—

Fig pedunculate, ripening green, brown, or brownish orange. Lec-

totype—Urostigma valida (Bl.) Miq. (= F. annulata Bl.).

I choose this type because Miquel obviously chose the name from

this species. It occurs, also, as U. flavescens (Bl.) Miq., and its

ally F. globosa occurs twice also, as U. globosum (Bl) Miq. and

U. manok Miq. Further, this choice gives the lectotype of sect.

Conosycea the position also of lectotype of a series.

2. ser. Drupaceae Corner ser. nov.—Receptacula sessilia v.

pedicellata, haud pedunculata, maturitae lutea, aurantiaca, dein

rubra. Typus—F. drupacea Thunb. (= F. pilosa Reinw.).

a. subser. Drupaceae.—Receptacula ellipsoidea, oblonga v.

pyriformi-pedicellata, bracteis basalibus plerumque conditis, api-

calibus plerumque gibbosis. Lamina costis basalibus haud elon-

gatis, cystolithis amphigenis. Typus—F. drupacea Thunb.

b. subser. Indicae Corner subser. nov.—Receptacula subglobosa

v. ellipsoidea, bracteis basalibus plerumque conspicuis, apicalibus

disco plano instructis. Lamina nervulis reticulatis ad paginas am-

bas leviter elevatis (sicco), cystolithis solum ad paginam superio-

rem v. nullis. Typus—F. benghalensis Linn. .

c. subser. Zygotricheae Corner subser. nov.—Receptacula sub-

globosa, ut in subser. Indicae. Cystolitha amphigena. Pili brunnei

copiosi, microscopice binati. Typus—F. consociata Bl.

aF2

Vol. XVI. (1959).

The microscopically twinned hairs are characteristic of F. con-

sociata and its close ally F. bracteata Wall.

d. subser. Crassirameae Corner subser. nov.—Receptacula sub-

globosa, oblonga, v. pyriformia, ut in subser. Indicae. Cystolitha

amphigena. Pili hyalini v. nulli, haud binati. Bah crassi-

ramea Miq.

B. subsect. Dictyoneuron Corner subsect. nov.—Lamina inter-

costis subparallelis deficiens, costis primariis multo majus evolutis

quam costis secondariis; cystolithis amphigenis, raro nullis. Typus

—F. sundaica Bl. (= F. indica sensu King, vide infra).

1. ser. Glaberrimae Corner ser. nov.—Receptacula pedunculata,

bracteis basalibus parvis v. nullis, ostiola non pertuso. Typus—F.

glaberrima Bl.

2. ser. Dubiae Corner ser. nov.—Receptacula pedicellata, haud

pedunculata, bracteis basalibus parvis v. minutis, ostiolo haud per-

tuso. Typus—F. dubia Wall. ex King.

3. ser. Subvalidae (Mig.) Corner comb. nov.—Urostigma Gasp.

ser. Subvalida Miq. Fl. Ind. Bat. 1, 2 (1859) 339. Fig sessile:

basal bracts well-developed: apical bracts in a disc, not perforate.

Typus—F’. sundaica Bl.

This group was described under Urostigma Gasp. in series-form

and 19 species were ascribed to it. I dispose these species as fol-

lows: —

a. subgen. Sycomorus:—U. lucescens B\. (= F. racemosa Linn.

var. elongata King);

b. subgen. Pharmacosycea:—U. hasseltii Miq. (= F. pubiner-

vis Bl.);

c. subgen. Ficus sect. Sycidium:—U. virgatum Reinw. ex BI.

(= F. virgata Reinw. ex Bl.);

d. subgen. Urostigma:—

(i) sect. Conosycea Mig. subsect. Conosycea ser. Drupaceae

Corner subser. Crassirameae Corner:—U. crassirameum Miq.;

(ii) sect. Conosycea Miq. subsect. Dictyoneuron Corner:—U.

sundaicum BL., U. glaberrimum Bl., U. sumatranum Migq., U. bin-

nendykii Miq., U. zollingerianum Miq. (== F. sumatrana Miq.),

U. peracutum Mig. (= F. binnendykii Miq.), U. pseudorubrum

Mig. (== F. sundaica Bl.);

(iii) sect. Conosycea Miq. subsect. Benjamina Mig. ser. Cailo-

phylleae Corner:—U. clusioides Mig. (= F. callophylla Bl.);

373

Gardens Bulletin, S.

(iv) sect. Urostigma:—vU. infectorium sensu Mig. (= F. virens

Ait.), U. canaliculatum Mig. (= F. virens Ait. var. glabella Bl.),

U. glabellum (Bl.) Mig. (= F. virens Ait. var. glabella Bl.), U.

moritzianum Mig. (= F. virens Ait. var. glabella Bl.), U. timo-

rense Mig. (= F. virens Ait.), U. concinnum Migq., U. parvi-

folium Miq. (= F. concinna Miq.):

Clearly the name should be used for one of the two larger groups

d (ii) or (iv), but d (iv) is already covered by the simultaneous

Urostigma Gasp. ser. Caulobotrya Miq. Therefore, I assign it to

d (11) as sect. Conosycea subsect. Dictyoneuron ser. Subvalidae

with F. sundaica Bl. as the type-species.

4. ser. Perforatae Corner ser. nov.—Receptacula sessilia,

bracteis basalibus bene evolutis, ostiolo pertuso, bracteis apicalibus

parvis non in discum planum instructis. Typus—F. pisocarpa BI.

(= F. microstoma Wall. ex Miq.).

C. subsect. Benjamina (Mig.) Corner comb. nov.—Urostigma

Gasp. ser. Benjamina Mig. FI. Ind. Bat. 1, 2 (1859) 344.—-Vena-

tion without intercostals, the secondary lateral nerves almost or

quite as strongly developed as the primary. Fig sessile, rarely

pedunculate or pedicellate. Type—F. benjamina Linn.

1. ser. Benjamineae Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867)

287.—Leaves subdistichous, thinly coriaceous, drying grey-green

to light brown: nerves distinctly raised on beth sides (dried): basal

nerves not elongate. Type—F. benjamina Linn.

2. ser. Callophylleae Corner ser. nov.—Folia spiraliter vel sub-

distiche disposita, sicco fusca v. fuscobrunmnea, saepe coriacea:

petiolo saepe nigro: costis basalibus saepe elongatis. Typus-—F.

callophylla Bl.

sect. Stilpnophyllum Endl. Gen. Pl. 4, 2 (1847) 35. Typus—

F.. elastica Roxb.

sect. Malvanthera Corner sect. nov.—Receptacula ostiolo bilabiato

vel triradiato, bracteis apicalibus internisque inflexis (haud in-

terpositis), bracteis basalibus 2—3 saepe caducis, praedita. Flo-

res masculi dispersi: antheris uniloculatis, saepe reniformibus,

aut longitudinaliter crescentiforme dehiscentibus aut transver-

saliter. Ovarium aut liberum basim versus rubromaculatum aut

plus minus in receptaculo immersum et apicem versus rubrum:

stigmate bifido v. simplo. Costae primariae et secundariae ut in

F. elastica quasi aequaliter evolutae, intercostis nullis. Typus—

F. macrophylla Desf.

This section resembles in the inflexed apical and internal bracts

the African sect. Galoglychia (— Bibracteatae), but differs in the

unilocular anther with two, not four, pollen-sacs. It consists of 19

374

Vol. XVIT. (1959).

species of Eastern Malaysia and Australia. I distinguish two series

and four subseries.

1. ser. Malvanthereae Corner ser. nov.—Anthera reniformia.

longitudinaliter crescentiforme dehiscentia. Typus—F. macro-

phylla Desf.

a. subser. Eubracteatae Corner subser. nov.—Bracteae basales

3, magnae, persistentes. Receptacula sessilia v. breviter pedun-

culata, ostiolo triradiato. Ovarium liberum, stigmate simplici.

Typus—F’. triradiata Corner sp. nov. (vide infra).

b. subser. Malvanthereae.—Bracteae basales 2 (? 3), mox

caducae, magnae. Receptacula pedunculata. Ovarium in recepta-

culo semi-immersum, stigmate simplici v. breviter bifido. Typus—

F. macrophylla Desf.

c. subser. Platypodeae Corner subser. nov.—Bracteae basales

2 v. 3, magnae, caducae. Receptacula pedunculata v. sessilia, 6—13

mm. lata (sicco). Ovarium liberum, stigmate simplici. Lamina

saepe parva (2-16 * 1.5—7.5 cm.), costis secundariis saepe minus

evolutis. Typus—F. platypoda A. Cunn. ex Mig.

d. subser. Hesperidiiformes Corner subser. nov.—Ut in subser.

Malvanthereae sed bracteis basalibus 3 parvis, persistentibus.

Typus—F.. hesperidiiformis King.

2. ser. Cyclanthereae Corner ser. nov.—Anthera depresso-

globosa, transversaliter dehiscentia. Ovarium in receptaculo semi-

immersum, stigmate bifido. Receptacula magna, oblonga, bracteis

3 parvis subcaducis praedita. Typus—F. sterrocarpa Diels.

The stamen resembles that of Brosimum alicastrum Sw. and 1s

unique in the genus.

sect. Galoglychia (Gasp.) Endl. Gen. Pl. 4, 2 (1847) 35.—

Galoglychia Gasp. Parl. Giorn. Bot. 2 (1844) 217; Ann. Sci.

Nat. ser. 3, 3 (1845) 346.—Ficus Linn. subgen. Bibracteatae

Mildbr. et Burr. Engl. Bot. Jahrb. 46 (1912) 175.

This section was based on F. galactophora Ten. and F. saus-

sureana DC., which are clearly African species of subgen. Bibrac-

teatae, and possibly identical. In view of the great diversity of the

Asiatic species of subgen. Urostigma and the parallel development

of sect. Malvanthera compared with the African Bibracteatae, 1

cannot recognise this as a subgenus, and I treat the African group

as a section of subgen. Urostigma.

sect. Americana Mig. Hook. Lond. J. Bot. 6 (1847) 525 (ut

Americanae); Mildbr. et Burr. Engl. Bot. Jahrb. 46 (1912)

169.

Gardens Bulletin, S.

This section is near both sect. Urostigma and sect. Conosycea,

but it differs in the two, generally connate, basal bracts. The ovary

has a red mark at the base and the cystoliths are generally amphi-

genous: thus the section is nearer to Conosycea. The species are

entirely American.

sect. Urostigma

F. superba Miq. var. alongensis (Gagn.) Corner comb. nov.—

F. alongensis Gagn. Lec. Notul. Syst. 4 (1927) 84.

This differs only in the small ovate-rotund lamina and short

petiole, and it appears to be a xerophytic form.

F. superba Miq. var. henneana (Mig.) Corner comb. nov.—F.

henneana Miq. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 216.—F.

gracilipes Bailey, Dept. Agr. Brisb. Bot. Bull. 3 (1891) 16.—F.

parkinsoni Hiern, J. Bot. (1901) 1.—F. pritzelii Warb. Fedde

Rep. 1 (1905) 74. Except for the axillary position of the fig and,

perhaps, the smaller leaves, I can find no distinction from F.

superba which extends from Indo-China and Malaya along the

Lesser Sunda [{slands to Timor. Hence this may well be its state

in Australia.

F. concinna Miq. var. subsessilis Corner var. nov.—F. subpedun-

culata Mig. Ann. Mus. Bot. Lugd. Bat..3 (1867) 217, 286

(haud 293).—Receptacula sessilia v. breviter pedunculata (0.5

mm.), bracteis basalibus caducis. N.E. India, Chekiang, Yun-

nan, Kwangtung.

Griffith 4589, 4590 (mixed with typical F. concinna); Henry

9122, 9965 (Yunnan); R. C. Ching 1917 (Chekiang, typus-herb.

Kew.); F. A. McClure 280 (C.C.C. 64, 1961, Kwangtung) ; Bodi-

nier 1027 (Macao).

At first perplexing, it is soon seen that, if there were a normal

peduncle, these collections would belong to F. concinna. Ching

1917, McClure 280, and Bodinier 1027 are var. concinna with

almost sessile fig. Griffith 4590 is the type of F. subpedunculata

Migq.: it has sessile figs at Leiden, but normally pedunculate figs

at Kew. Henry 9122 and 9965, as well as the Leiden specimen of

Griffith 4590, have more coriaceous leaves with the veins not or

scarcely raised above, and the lateral nerves at a wider angle.

Henry 9122 has, also, a rounded to subcordate base to the lamina

and approaches F’. cardiophylla Merr. (fewer costas, longer basal

nerves ).

F. virens Ait. Hort. Kew. 3 (1789) 451.—¥F. infectoria Willd.

sensu Roxb. Fl. Ind. 3 (1832) 551.—Urostigma infectorium

Mig. Zoll. Syst. Verz. (1854) 90.—F. infectoria (Miq.) Miq. Ann.

376

Vol. XVII. (1959).

Mus. Bot. Lugd. Bat. 3 (1867) 264, 286; King, Ann. R. Bot.

Gard. Calc. 1 (1887) 60, pl. 75.—F. lacur auct., non vera.—

This is the earliest name for the common banyan of India which

has the articulate lamina, and which extends through Malaysia

to Australia (F. cunninghamii) and to South Africa as the

complex of F. ingens Miq. F. virens was based on a plant grown

at Kew, of which there is a sterile specimen (the type) at the

British Museum. This specimen is labelled F. virens Ait. in

pencil and is one of a complete set of named specimens repre-

senting exactly those described by Aiton: hence there is no doubt

of its validity. But Aiton referred it to the West Indies, evidently

in the belief that it represented the plant figured as “Sloane Jam.

2 p. 140 t. 223” which is quoted after the description. The re-

ference to Sloane’s plant is, however, erroneous. Thus this name

has been overlooked by students of Asiatic botany.

F. virens Ait. var. sublanceolata (Miq.) Corner comb. nov.—

F. saxophila Bl. var. sublanceolata Miq. Ann. Mus. Bot. Lugd.

Bat. 3 (1867) 260.—F. caulobotrya (Miq.) Mia. var. fraseri Miq.

id. 287.—F. virens has numerous leaf-forms, to which varietal

names may be given, when they are better known, but I find so

much variation in herbarium-material, even on the same speci-

men, that I have been unable to recognise any such categories.

However, two varieties can usefully be distinguished on the fig,

whether pedunculate or sessile. As the type of F. virens is a

sterile sapling and there is no mention of the fig in the original

description, it is necessary to adopt the earliest varietal name,

but Miquel published simultaneously (1867) names for each

variety. F. infectoria (Mig.) Miq. var. aegeirophylla (Miq.)

Mig. refers to the pedunculate fig. while F. saxophila Bl. var. sub-

lanceolata Miq. and F. caulobotrya (Miq) Miq. var. fraseri

(Miq.) Mig. refer to the sessile fig. I propose to use the name

var. sublanceolata Migq. for the sessile fig, and thus to take var.

virens as representing the pedunculate fig. The decision is arbit-

rary, for both kinds occur in India and neighbouring countries,

from whence the Kew-plant must have come.

F. virens Ait. var. glabella (BI.) Corner comb. nov.—/. glabella

BI. Bijdr. (1825) 452.—This is the state with elliptic to obovate

lamina, cuneate leaf-base, shorter petiole, and more or less ses-

sile figs on short woody burrs on the twigs: behind the leaves.

It is the common state of the species in the rain-forests of Sun-

daland from lower Thailand and Sumatra to Borneo, where var.

377

Gardens Bulletin, S.

virens and var. sublanceolata rarely occur and, then, chiefly on

headlands and dry places. At the periphery of its range it grades

into var. sublanceolata. Hence I cannot maintain it as a species.

F. caulocarpa Miq. var. dasycarpa Corner var. nov.—F. argentea

Blanco sensu Merrill, Sp. Blanc. (1918) 129; Philip. J. Sci. 20

(1922) 368; Sata, Monogr. Ficus (1944) 216.—Receptacula et

pedunculi dense albo-villosi: bracteis basalibus 1—-1.5 mm.

longis, glabris. Philippines (Luzon, Mindoro).

Bur. Sci. 39732 (Mindoro, Paluan); PNH 4788 (Luzon, Zam-

bales prov., Mt. Pinatubo, typus—-Manila).

This was identified by Merrill as F. argentea Blanco, which is a

thoroughly dubious species, Merrill having first suggested that it

was F. ruficaulis Merr. Neither fits Blanco’s description.

F. prolixa Forst. f. var. subcordata Corner var. nov.—Lamina

ovato-elliptica, ad basin cordata, subcordata, v. late cuneata,

—15 X 8 cm.: nervis primariis utrinsecus 5-8, basalibus elon-

gatis 4-1/3 laminae: petiolo plus minus applanato. Ins. Caro-

line, Mariannas, Fanning.

Caroline Is]—Fosberg 25479, E.¥. Hosaka 3231, (Ulithi Atoll).

Mariannas—Fosberg 24972, 25004, 25112, (Rota); Fosberg

24731, 24758, Hosaka 2862, (Tinian); Fosberg 25221, 31278.

31791, Hosaka 2962, (Saipan); Anderson 123, 192, Fosberg

29302, 25331, ° 32620, 49355 (typus), 35515, “33516, ses,

35631, (Guam); Fosberg 31684 (Alamagan); Anderson 508, 521,

Fosberg 31392, (Pagan). Fanning—J.T. Arundel 14 (sterile, herb.

Kew.).

This has the scattered male flowers of F. prolixa, though the

more or less cordate leaf suggests F. virens. It is possible that the

two species intergrade in the Carolines, where some collections (as

Fosberg 25479) have so few scattered male flowers that the dis-

tinction seems trivial. Specimens with ovate-cordate leaves have

not been recorded from the main range of F. prolixa in Polynesia.

F. hookeriana Corner nom. nov.—F. hookeri Miq. Ann. Mus. Bot.

Lugd. Bat. 3 (1867) 215, 286 (non Sweet, 1827).—This be-

longs in ser. Orthoneurae.

F. caulobotrya Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 287.—

Urostigma caulobotryum Mig. Hook. Lond. J. Bot. 6 (1847)

568.—This is a mixture of leaves of F. tsjahela Burm. f. and figs

of F. arnottiana Mig. Such is the state of the type-number

(Wight 26) at Utrecht, Leiden, and Edinburgh (three sheets).

It is clearly a mixtum compositum of mounting the material,

and the name can be dismissed.

378

Vol. XVII. (1959).

sect. Conosycea (Miq.) Corner subsect.

Conosycea ser. Validae Miq.

F. arnottiana Mig. var. subcostata Corner v. nov.—Lamina sub-

acuta vel obtusa. Receptacula 10 mm. lata, maturitate rubra:

pedunculo 5S—8 mm. longo. Cuticula circum stomata plicato-

striata. Himalaya.

Duthie 23939 (Gahrwal); Gamble 23030 (Jehree); Polunin,

Sykes, et Williams 4145 (Nepal, Melcham, c. 7000 ped. alt.,

typus—herb. Br. Mus.).

This has the larger fig and more striate leaf-cuticle of F. costata

Ait. (v. infra), but the ovate-cordate lamina with few lateral ner-

ves, prominent basal nerves, and long petiole of F. arnottiana. It

lies outside the geographical range of both and, as an intermediate,

needs further study. Compare, also, F. glaberrima Bl. var. siamen-

sts Corner (v. infra).

F. costata Ait. Hort Kew. 3 (1789) 452.—-F. venusta Kth. et

Bouch. Ind. Sem. Hort. Berol. (1846) 16.—F. caudiculata

Trim. J. Bot. 23 (1885) 243.—F. mooniana King, Ann. R. Bot.

Gard. Calc. 1 (1887) 57, pl. 69.—Urostigma wightianum Mia.

var. majus Thw. En. Pl. Ceyl. (1864) 265.

F.. costata was said to come from the East Indies, and, according

to Index Kewensis, from the Nicobar Islands. No such plant has

since been recorded from the Nicobars, and I find that the type at

the British Museum agrees in leaf, venation, and microscopic

structure with F. caudiculata and F. mooniana of Ceylon. The type

is Clearly, a sapling with large leaves and is sterile. There is also a

specimen in the Herb. Martyn (Cambridge University Herbarium }

named F. costata which agrees so well with the type that it may

have come from the same plant cultivated at Kew. The cystoliths

only on the upperside of the lamina, the plicate cuticle, and the

upper hypodermal layer are distinctive microscopic characters in

the Asiatic species of this alliance. F. venusta was said to have

come from Cuba, but no such leaf has been found again in the

Cuban flora and the type-specimen at Berlin agrees exactly with

the specimen of F. costata at Cambridge.

There is some discrepancy, however, in the length of the petiole.

In the type of F. costata the petiole is long, up to —9 cm., and the

lamina is ovate-cordate to elliptic with subcordate base. The spe-

cimen in Herb. Martyn has the elliptic lamina with subcordate

base but shorter petiole 1.2—5.5 cm. long. In F. caudiculata the

lamina is elliptic with narrowly subcordate to cuneate base and

short petiole 1.3-3 cm. long. But a specimen from Indo-China

(Ch. d’Alleizette, environs de Tourane, 2.vi.09), which I cannot in

379

Gardens Bulletin, S.

any way distinguish from F. caudiculata, has the petiole —7 cm.

long. Further, King’s plate of F. mooniana shows a petiole inter-

mediate between the short petiole of F. caudiculata and the longer

petiole. Hence I assume that the long petiole and ovate-cordate

leaf must be the sapling form, which reduces to the elliptic leaf

with subcordate to cuneate base and short petiole on the upper

twigs of mature trees.

F. depressa BI. Cat. Gewass. Buitzg. (1823) 35; Bijdr. (1825)

450.—F. pruniformis Bl. Bijdr. (1825) 451.—Blume confused

this originally with specimens of F. magnoliifolia Bl., several

sheets of which are named F. depressa in Blume’s hand at Lei-

den. In 1825, he divided F. depressa into F.. pruniformis Bl. and

F. magnoliifolia Bl. without explanation. I reserve the name F.

depressa in the sense of F. pruniformis. The type-sheets of all

three species are well-preserved at Leiden, and that of F. priuni-

formis is identical with the lectotype of F. depressa.

F. globosa Bl}. Bijdr. (1825) 449.—Urostigma onustum Mig., Hook

Lond. J. Bot. 6 (1847) 575.—F. onusta Wall. ex Mig. Ann.

Mus. Bot. Lugd. Bat. 3 (1867) 285.—The type-sheet in Herb.

Hook at Kew consists of (a) two leaves and a twig with two

figs of F. pisocarpa Bl. (= F. microstoma Wall. ex King),

and (b) five leaves with two figs of F. globosa Bl. The sheet in

the Wallich-herbarium at Kew is entirely F. globosa Bl. W. B.

Hemsley wrote on the sheet in Herb. Hook. that Miquel’s des-

cription corresponded with the F. pisocarpa-element. On study-

ing Miquel’s description, however, it is clear to me that he

described the leaves of F. globosa and the figs of F. pisocarpa.

Thus the species is a mixture, and I assign it to F. globosa.

sect. Conosycea (Miq.) Corner subsect. Conosycea ser.

Drupaceae Corner

F. drupacea Thunb. Diss. Ficus (1786) 6,1 1.—¥F. pilosa Reinw. ex

Bl. Bydr. (1825) 446—Examination of Thunberg’s type has

shown that this is the well-known F. pilosa from Java.

var. auranticarpa (Elm.) Corner comb. nov.—F. auranticarpa

Elm. Leafi. Philip. Bot. 9 (1937) 3454.—Bracteis basalibus glab-

ris 2-3 & 3—5 mm., non conditis. Luzon (Elmer 17186, typus),

‘Celebes (Beccari s.n., Kandari: R. Ist. Fir. 9347, 9347A).

var. glabrata Corner v. nov.—Stipulae et nodi ultimi brunneo-

pilosae, praeterea glabra. Bracteae basales 0.5—1 mm. longae, cres-

centiformes, margine ciliolato. Setae internae nullae v. sparsae.

Malaysia, Ins. Solomon.

380

Vol. XVII. (1959).

Kangeen Archipelago:—-Backer 167, 26956, 27147, 29766

(det. F. altissima). Sumbawa:—Herb. Lugd. Bat. 293 (det. F.

annulata). Soemba:—lIboet 103, 280 (det. F. altissima). Flo-

res:—Elbert 4320. Alor:—O. Jaag 767, 892. Timor Laut:—

Forbes 3371 (typus, herb. Br. Mus.). Solomon Isl.:—-Guppy 116

(Shortland Isl. ).

Glabrescent forms of var. drupacea have persistently brown

hairy basal bracts.

var. pedicellata Corner v. nov.—Ut var. glabrata sed receptacula

super bracteas basales pedicello 3-4 mm. longo praedita: setis

internis brunneis, —1 mm. longis, copiosis. Nova Guinea (NGF

8557, Western Highlands, Jimmi Valley, Tagan River, 2400 ped.

alt., typus herb. Lae).

F. cordatulae Merr. proximans.

var. pubescens (Roth) Corner comb. nov.—F’.. mysorensis Heyne

ex Roth, et var. pubescens Roth, in Roehm. et Schult. Syst. Veg.

1 (1817) 508.—I cannot distinguish F. mysorensis from the large

and varied complex of F. drupacea.

F. cucurbitina King var. eubracteata Corner v. nov.—Bracteae:

basales 5-7 x 10-13 mm., conspicuae, obtusae, glabrae v.

marginibus ciliolatis. Ins. Philippines (PNH 14352, Samar,,

Catarman, Mt. Cansayo, typus herb. Manila).

In v. cucurbitina bracteae basales pusillae, 0.3-1 mm. longae,,

sub receptaculo conditae.

F. cordatula Merr. var. sericea (C. B. Robinson) Corner comb.

nov.—F. sericea C. B. Robinson, Philip. J. Sci. Bot. 6 (1911)

319.—This is merely a form of F. cordatula with villous or to-

mentose twigs, stipules, and receptacles.

sect. Conosycea (Mig.) Corner subsect. Conosycea

ser. Indicae Corner

F. benghalensis Linn. Sp. Pl. (1753) 1059; C. Hegard, Amoen.

Acad. 1 (1749) 217; Miquel, Hook. Lond. J. Bot. 6 (1847)

571, 572; King, Ann. R. Bot. Gard. Calc. 1 (1887) 18, 19.—

F. indica Linn. Sp. Pl. (1753) 1060; emend. Lamarck, Enc. Bot.

2, pt. 2 (1788) 494; Smith, Rees Cycl. 14 (1810) n.41; Rox-

burgh, Hort. Beng. (1814) 65; B. Hamilton, Tr. Linn. Soc. 13:

(1822) 489; non sensu King, Ann. R. Bot. Gard. Calc. |

(1887) 338, 40.—F. cotoneaefolia Vahl, En. Pl. 2 (1806) 189;

non sensu Alston, The Kandy Flora (1938) 34.—F. cotonifolia

Stokes, Bot. Mat. Med. 4 (1812) 355.—I have reduced F.

indica Linn. to F. benghalensis Linn. because, as shown below.

381

Gardens Bulletin, S.

they are synonymous. Roxburgh and Buchanan Hamilton long

ago advised the converse, but their advice has never been ac-

cepted and to reinforce it now in view of the general agreement

about F. benghalensis and the Jong current misuse of F. indica

sensu King would be confusing. For this species, F. indica

sensu King, I have substituted F. sundaica Bl. (v. infra).

The name F. indica comes from the great Indian banyan of an-

cient and mediaeval reference, namely “Ficus indica Theophrasti”

(Tabern., Hist. p. 1370) and “Ficus indica foliis mali cotoneae

... (Bauhin, Pinax, p. 457). All this is explained by Cornelius

Hegard. It can refer only to the species known as F. benghalensis,

which is based on Rheede’s description and illustration (Hort.

Mal. 1, t.28, “Peralu”). Unfortunately, the poor wood-cuts of

Clusius, Tabernaemontanus, and Bauhin, which showed a pointed

and narrower leaf, lead the early botanists to suppose in their

ignorance that there were two species of such great banyan in

India, and the name F. indica Linn. was given to this fiction. Lin-

neus believed that the banyan figured as ““Katou-alou” by Rheede

(Hort. Mal. 3, t. 57) represented this F. indica and cited it as

well. King has shown, however, that Katou-alou is F. mysorensis

Heyne, which I call F. drupacea Thunb. var. pubescens (Roth

Comer (v. supra). Thus, the bad woodcut and the mis-reference

of Katou-alou fostered the belief in two great Indian banyans.

Then, as F. indica var: 6,” Ummneus (1753) paver” Ean

(Rheede, Hort. Mal. 3, t. 63) and references to American plants.

In 1763 (Sp. Pl. ed. 2, 1514-1515), he added to var. 6 the

“Varinga latifolia” of Rumphius (Herb. Amb. 3, t. 84). Lamarck

perceived the error of including var. @ under F. indica, and ex-

cluded it, thereby definitely emending F. indica as consisting of

(i) the references to Bauhin, Tournefort, Clusius, and Taber-

naemontanus (= F. benghalensis Linn.),

(ii) Katou-alou of Rheede (= F. mysorensis Heyne),

(ii1) Madagascan collections (? identification),

(iv) F. vasta Forsk.,

(v) A plant grown at Paris in the “Jardin du Roi” (which, ac-

cording to the specimen in the Lamarckian herbarium,

is F. tsiela Roxb.).

Of these five components, the first and second are elements of

the original circumscription, but the second has since been ex-

cluded as F. mysorensis; the others must be excluded as errone-

ous. Hence one is back at the original definition which is to be

identified with F. benghalensis. This was the opinion of King, but

382

Vol. XVIT. (1959).

he endeavoured to maintain the name for “Varinga latifolia” of

Rumphius, though this was not part of the original circumscription

and was definitely excluded by Lamarck. King’s use is further com-

plicated by his attempt to identify this Varinga latifolia.

The species, described and figured by King as F. indica, is a

common strangling fig spread from Assam to Borneo. It does not

occur in central India and it does not develop pillar-roots, as the

great Indian banyan does, and hence it cannot be the F. indica of

history. It does not occur, too, in the Moluccas, so it cannot be the

“Varinga latifolia”. Further, Rumphius described the fig as ellip-

soid, which Lamarck noted, and which lead Merrill (Interpr. Rum-

phius Herb. Amb., p. 194) to identify “Varinga latifolia” with F.

gelderi Miq. or F. altissima Bl., though neither of these occur in

the Moluccas. Personally, I think that “Varinga latifolia” is the

glabrescent state of F. drupacea Thunb. (F. pilosa Reinw.) which

is common in the Moluccas, where Rumphius could hardly have

missed it. Miquel had already given various names to the plant

intended by King as F. indica, but the earliest name that I can

find is F. sundaica Bl.

F. cotoneaefolia Vahl seems to have been intended as a new

name for F. indica Lam., which is really F. indica Linn. emend.

Lam., but the name was treated by King as a new species and so

used by Alston. I base my conclusion on these considerations :—

(i) Vahl saw no specimen,

(ii) He cited merely the Asiatic references given for F. indica

and var. 8 by Linneus in 1763, which are the Lamarc-

kian references,

(iu) He cited Lamarck’s emendation of F. indica,

(tv) He described F. indica. Linn. six pages later (p. 195),

basing his interpretation of it on the Linnean var. £,

as shown by the citation of “Tsiela” of Rheede,

“Varinga latifolia” of Rumphius,

(v) He seems to have been thoroughly confused for he also

cited “Varinga latifolia” under F. cotoneaefolia.

Smith (1810) promptly recognised F. cotoneaefolia as super-

fluous and reduced it to F. indica Linn. In contrast, King inter-

preted F’. cotoneaefolia by reference to “Katou-alou” of Rheede,

which he showed to be F. mysorensis, and Alston substituted F.

cotoneaefolia for F. mysorensis. There is, however, no evidence

that “Katou-alou” was an essential part of the original circums-

criptions of Linneus, Lamarck, or Vahl. Certainly they did not

describe F. mysorensis. Further, as the specific epithet indica came

from the early pre-Linnean authors, so cotoneaefolia clearly came

383

Gardens Bulletin, S.

from Bauhin’s ‘Ficus indica foliis mali cotoneae . . .” I conclude

the Smith was right and that Katou-alou was merely an erroneous

reference. A specimens in Vahl’s herbarium is named “F. ? coto-

neaefolia Vahl”, but it is F. tsiela Roxb., which was also cited by

Vahl under F. cotoneaefolia as ““Tsiela” of Rheede. Lamarck, also,

cited Tsiela under F. indica. This does not mean that they inter-

preted their species on Tsiela, but that they did not understand the

banyans of India and tried to fit the species known to Rheede into

the erroneous concept of F. indica.

F. pubilimba Merr. var. ovata Corner v. nov.—Glabra. Lamina

6-12 « 4-9.5 cm., ovata, breviter et obtuse subacuminata, basi

cordato vel rotundato-subtruncato; costis basalibus 1/3-—4

laminae elongatis. Malaya (Waterfall Gardens, Penang, leg.

Flippance Oct. 1932, typus herb. Singapore).

This is known only from a single tree, still thriving, in the

Penang Waterfall Gardens. The Province Wellesley Fig (Corner,

Wayside Trees of Malaya, p. 680) is typical F. publimba Merr.,

which has the more or less elliptic leaf with cuneate base.

sect. Conosycea (Miq.) Corner subsect. Conosycea

ser. Crassirameae Corner

F. stapenda Miq. var. minor Corner v. nov.—Ramuli 7-10 mm.

crassi. Lamina 8—24 x 6—10.5 cm., basim versus rotundata v.

late cuneata. Receptacula 30 « 22 mm., oblongo-ellipsoidea,

bracteis apicalibus in discum planum 5 mm. latum instructis:

bracteis basalibus 5-10 « 10-14 mm., obtusis. Flores minores,

masculi 2—6 mm. longi. Johore, Singapore, Brunei, North

Borneo.

Singapore:—Kelsall s.n., Chan Chu Kang, Nov. 1889; Good-

enough 3399, Bukit Timah, Nov. 1887 (typus, herb. Singapore).

Johore:—Sing. F.n. 37027, Mawai-Jemaluang Road, 17th mile,

May 1940. Brunei:—Brun. 5325, Ulu Belalong. North Borneo: —

Clemens 27132, Tenompok (ut Artocarpus); Elmer 21054, Tawao;

For. Dept. A4712, Kinabatangan, Bt. Garam.

F. crassirameae Miq. proximans sed bracteis basalibus breviori-

bus et vix induratis.

F. crassiramea Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 286.—

Urostigma crassirameum Miq. Pl. Jungh. (1851) 48.—¥F. pro-

cera Bl. var. crassiramea (Miq.) King, Ann. R. Bot. Gard. Calc.

1 (1887) 36, pl. 41.—F. procera auct., non vera.—As explained

under F. subtecta Corner, F. procera Bl. has been mistakenly

identified with F. crassiramea Miq., which is common and widely

distributed in Malaysia.

384

Vol. XVIT. (1959).

var. brevicupulata Corner v. nov.—Receptacula 16-19 x

13-15 mm., subcylindrica, bracteis apicalibus in discum planum

4—5 mm. latum instructis; bracteis basalibus 5-7 « 7—12 mm.,

crassis, subcupulatis. Celebes, Obi.

Celebes:—Kjellberg 1300, Poehara, Moeara Sampara; 2245,

Tabaramo. Obi:—Exp. v. Hulstyn 111 (typus, herb. Leiden).

var. celebica Corner v. nov.—Receptacula 27-28 & 20 mm.,

subcylindrica, bracteis basalibus 15-18 mm. longi. Celebes (Mina-

hassa, Koorders 19126, 19160, et 19334, typus herb. Leiden).

var. clementis (Merr.) Corner comb. nov.—F. clementis Merr.

Philip. J. Sci. Bot. 3 (1908) 130.—Receptacula 15-22 « 12-17

mm., bracteis basalibus 8-14 mm. longis. Ins. Philippine.

Clemens 421, 703 (typus); Elmer 10952; Merrill 1935; Bur.

Sci. 19495: PNH 1985; Coll. Agr. Laguna 513, 547.

var. patellifera (Warb.) Corner comb. nov.—F. patellifera Warb.

in K. Schum. et Laut. Nachtr. Fl. Schutzgeb. (1905) 241.—

Receptacula 12-14 mm. lata, subglobosa, bracteis apicalibus in

discum planum 3 mm. latum instructis; bracteis basalibus 5-8 x

10-15 mm. Nova Guinea.

Schlechter 14161, Ranu coast (typus, herb. Berlin); Kanehira

et Hatusima 12838, Nabire, Neth. New Guinea; Brass 3857, Terr.

N. Guinea, Central Div., Ononge Road, Dieni; Brass 24182, Milne

Bay. Peria Creek; Brass 28085, Sudest Island, Rambuso; NGF

hi23, Lie;

F.. forstenii Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 214, 285.—

F. palawanensis Merr. Govt. Lab. Publ. Philip. 29 (1905) 11.

var. pacifica (Elm.) Corner comb. nov.—F. pacifica Elm. Leafl.

Philip. Bot. 9 (1937) 3460. .

_ Lamina elliptico-obovata, breviter subacuminata, basi cuneato;

costis basalibus 1/3—4 laminae elongatis. Malaya, Ins. Philippine.

Malaya:—King’s Collector 6376 (Perak, Larut). Philippines:—

Elmer 7460, 9370, 14463, 16401; Merrill 4032; Wenzel 3022;

Bur. Sci. 26345, 76378; PNH 2050, 3673, 9066. (Elmer 14463,

16401, syntypes).

var. umbobracteata (Elm.) Corner comb. nov.—F. umbobracteata

Elm. Leafl. Philip. Bot. 4 (1911) 1247.—Receptacula 25-32

18-22 mm., bracteis apicalibus gibbosis, bracteis basalibus 7—9

mm. longis. Mindanao (Elmer 11956 typus, 14117).

var. villosa Corner v. nov.—Ramuli, stipulae, petioli, laminae

pagina inferior, bracteaeque basales pilis erectis (0.5 mm. longis)

villosi. Receptacula 20-22 x 18~—20 mm., bracteis basalibus 8-11

xX 9-12 mm. Malaya, Borneo.

385

Gardens Bulletin, S.

Malaya:—Corner s.n. Perak, Kroh, July 1936. Borneo:—

Endert 3344, W. Koetai, pr. L. Petah (typus, herb. Leiden); For.

Dept. B.N.B. A1073, Elopura, Sandakan; A3498, Lahad Datu,

Kertam Kechil River; Sing. F.n. 19235, Banguey Isl.

F. subtecta Corner nom. nov.—F. procera Bl. Bijdr. (1825) 445,

non Salisb. (1796).—F. procera Bl. has been consistently mis-

identified with F. crassiramea Mig. since King’s monograph.

Dried leaves of F. procera never show the characteristic shallow

pitting of the areolae on the underside of the lamina as occurs

in F. crassiramea, and there are constant differences in fig and

venation. Blume’s type 1s well-preserved at Leiden.

var. depressa Corner v. nov.—Receptacula 13-14 17-19

mm., depresso-globosa. Sumatra (Biinnemeyer 9089, G. Koerintji,

typus herb. Leiden).

F. subgelderi Corner sp. nov.—Corner, Wayside Trees, Malaya

(1940) pl. 208 (ut F. indica)—Arbor magna, epiphytica, suffo-

cans, radicibus columnaribus deficiens; cortice albido-griseo;

foliis subdistichis. Ramuli, petioli, stipulae, bracteae basales, et

costae paginae inferioris pilis erectis brevibus sparsim puberuli

v. subvillosi. Ramuli 3—5 mm. crassi. Lamina 4-16 x 1.5-7

cm., anguste obovata v. elliptica, acuminata, basi cuneato,

rigide coriacea: costis lateralibus utrinsecus 3—6 (—7), intercostis

2—3 vix bene evolutis, subtus elevatis: costis basalibus utrin-

secus 1, 1/3—4 laminae elongatis: petiole 5-32 mm. longo. Re-

ceptacula axillaria, binata, sessilia, maturitate e flava rubescen-

tia, 13-16 x 10.5-13 mm., ellipsoidea v. breviter oblonga,

bracteis apicalibus 2-3 in discum planum 3 mm. latum

instructis: bracteis basalibus 6—7.5 < 7—8.5 late ovatis, obtusis,

puberulis v. subtomentosis: setis internis nullis. Tepala 2

(mascula), 3 (feminea), libera, in floribus masculis et cecidio-

phoris valde indurata. Flores masculi sessilia v. breviter pedi-

cellata: anthero angusto, valde exserto. Stomata superficialia.

Malaya, Borneo.

var. subgelderi.

Malaya:—King’s Coll. s.n., Perak; For. Dept. 12712, Ampang

Reserve, Selangor; Derry 1211, Nyalas, Malacca; Kepong F. n.

71309, Kluang, Johore; Ridley s.n., Sept. 1890, et 5621, Bukit

Timah, Singapore; Sing. F. n. 29224 (typus, herb. Singapore) et

Ridley s.n., June 1898, Government Hill, Singapore.

Borneo:—Bur. Sci. (Manila) 1989, Sarawak; Brunei 226, S.

Belait, H. Low s.n., 1867, Labuan (herb. Beccari, R.Ist. Fir.

386

Vol. XVII. (1959).

9150A); For. Dept. B.N.B. 2323, Sandakan; Whitmore 583,

Kalabakan, W. Tawao; Kostermans 4179, Balikpapan, S. Wain,

et 6616, Samarinda, L. Djanan.

F. pellucido-punctatae Griff. (F. gelderi Miq) et F. sundaicae

Bl. (F. indica auct.) persimilis sed ostiolo non pertuso, intercostis

transversis satis distincta.

var. rigida Corner v. nov.—Urostigma rigidum Mig. Hook.

Lond. J. Bot. 6 (1847) 578.—F. rigida Miq. Ann. Mus. Bot. Lugd.

Bat. 3 (1867) 286, non Jack (1822).—Folia spiraliter disposita.

Glabra v. stipulae bracteaeque puberulae. Lamina 8-20 x 3-8

cm., basi subcordato v. cuneato: intercostis 3-8. Receptacula 10—

14 mm. lata, globosa, plus minus depressa, disco plano 5 mm.

lato praedita; bracteis basalibus 5-6 « 4—5 mm., v. 6-8 « 8-9

mm. Tepala vix indurata. Flores masculi bene pedicellati. Cam-

bodia, Cochinchina, Thailand, Malaya, Sumatra, Java, Borneo.

Cambodia:—Poilane 14769, Nord Kampat. Cochinchina:-—Fle-

ury 29911, Mt. de Nui Chua Chang, prov. Bienhou; Poilane

21309, km. 58 route col. n. 20, prov. Bienhou; Poilane 21513,

Dinh Quan, km. 46 route col. n. 20, prov. Bienhou. Thailand:—

Kerr 2743, Ban Pe, Ragawang: Kerr 16558, Kawchang, Ranaung;

Kerr 17440. Talang, Puket; J. Schmidt 140, Koh Chang (det. F.

consociata); RFD 6232, Kaw Kut, Trat. Malaya:—Hb. Hook,

Penang (typus F. rigidae Miq, et var. rigidae Corner); Philips s.n.,

1824, Penang; Scortechini 194b, Perak; Wray 2530, Perak; Sing.

F.N. 22344, Pahang; Sing. F.n. 23899, 30791, Sedili River,

Johore; Wallich 4570E, Singapore (ut F. longifolia Ham. ined.).

Sumatra:—Buwalda 6483, 6766, Indragiri; Grasshof 147, Lema-

tang Oeloe; NIFS bb 22450; ?Teysmann 727, Batang Baroes.

Java:—Koorders 9179, 9380, 20669, 23090, 24612, 38480,

38773, 38811, 39384; Forbes 530; Jacobs 4818; Bakh. v.d. Brink

6063; Korthals 39. Borneo:—Hewitt 6, Kuching.

F. paracamptophylla Corner sp. nov.—Valde scandens, foliis

spiraliter dispositis, radicibus gracilibus pendentibus subsparsis,

trunco —4 cm. crasso, haud suffocans. Glabra v. petiolo et

laminae pagina inferiori puberula. Ramuli 5—9 mm. crassi. Sti-

pulae 10-80 « 9—20 mm., brunnescentes, oblongo-lanceolatae,

persistentes. Lamina 10-26 « 3.5—9.5 cm., oblongo-elliptica,

subacuminata v. breviter et acute acuminata, ad basim sub-

cordata v. cuneato-rotundata, valde coriacea: costis lateralibus

utrinsecus 6-9, subtus vix elevatis, intercostis 1-4, saepe sub-

scalariformibus: costis basalibus utrinsecus 1 (—3), 4-4 laminae

elongatis: petiolo 15-30 « 3-6 mm., sicco fusco-brunneo.

Receptacula axillaria, sessilia, binata, 13-16 % 13-17 mm.

387

Gardens Bulletin, S.

(vivo, 20-25 mm.), ellipsoidea v. subconica, bracteis apicalibus

3 in discum planum 3—5 mm. latum, haud perforatum, instructis:

bracteis basalibus 4-7 *« 5-9 mm., obtusis v. subacutis, ap-

presse puberulis. Flores masculi copiosi, tepalis 2. Flores ceci-

diophori pedicellati, tepalis 2-3; feminei sessiles, tepalis 3.

Borneo, in silvis inter arbores insinuans, haud rara.

F. acamptophyllam Miq. simulans sed major, stipulis non cadu-

cis, receptaculi apice non pertuso.

Anderson 9092, Sarawak, Bau, Seburan, et 9103, Semengoh

Forest (typus, herb. Kuching); Corner s.n. Sarawak, Bako National

Park; Brunei 5323, Ulu Temburong; Kostermans 4027, Balik

Papan; Main (Exp. Polak) 2119, Indonesian Borneo, Pulow,

Kudala tas Djonkong; Nieuwenhuis 1425, Indonesian Borneo,

Bloe-oe; Sinclair 9352, Sandakan, Sepilok Forest.

sect. Conosycea (Mig.) Corner subsect. Dictyoneuron Corner

ser. Glaberrimae Corner

F. glaberrima Bl. var. bracteata Corner v. nov.—F. travancorica

King, Ann. R. Bot. Gard. Calc. 1 (1887) 28.—F. lawesii King,

J. As. Soc. Beng. 55 (1887) 403.—F. adamii Elm. Leafl. Philip.

Bot. 4 (1911) 1258 (ut F. adamsii laps. cal.).—F. villamilti

Merr. ex Sata, Monogr. (1944) 183.—Bracteae basales (ad

apicem pedunculi) 0.5—-1.5 mm. longae, crescentiformes: brac-

teae apicales saepe incrassatae. Flores cecidiophori plerumque

sessiles. India, Borneo, Philippines (Luzon, Mindanao), Nova

Guinea.

India:—Barber 6006, Anamalais; Beddome 7509, Travancore.

Borneo:—Kostermans 4918, E. Koetai, Sangkulirang Isl.; Purse-

glove 5183, Sarawak, S. Mayeng, Tau range. Philippines: —Bur.

Sci. 22821, 28100; Elmer 11177, 14029, 18284; For. Bur. 26361;

PNH 9480, 10623; Coll. Agr. Laguna 510. New Guinea:—Aet et

Idjan 738, 843, Jappen-Biak; Docters v. Leeuwen 10055, Rouf-

faer River; Forber 85, Sogeri; Hoogland 3748, Terr. N. Guinea,

Northern Div., Garara, Popondetta-Cape Killerton Road (typus,

herb Lae); Hoogland 4634, North Div., Tufi subdistr., Budi;

NGF 5216, 7423, Morobe, Bulolo: NGF 9067, Lae.

Var. glaberrima has no basal bracts. The so-called figs of F.

lawesii (Forbes 85) are abnormal galls.

var. Siamensis Corner v. nov.—Lamina 5-9.5 « 3-7.5 cm., late

‘elliptica, breviter acuminata, basi rotundato v. subcordato: costis

lateralibus utrinsecus 5—9, intercostis 1—4 vix distinctis: costis ba-

salibus haud elongatis: petiolo 10-30 mm. longo. Receptacula 8

388

Vol. XVII. (1959).

mm. lata, peduculo 10-15 mm. longo: bracteis basalibus 3, 1-2

mm. longis, obovatis, subacutis, caducis: bracteis apicalibus parvis,

gibbosis. Cuticula circum stomata et cystolitha plicato-striata.

Thailand, in collibus calcareis.

Kerr 2428, Hin Hap, Saraburi (typus, herb. Br. Mus.); Kerr

3021, Kao Sung, Paknampo; Kerr 9756, Krabin, Kao Sakan;

Nielsen 651, Koh Sichang.

This needs more collection because it may be a new species. It

relates F. arnottiana, F. costata, and F. glaberrima. It has the short

basal nerves, short petiole, and the fig of F. glaberrima, but the

apical bracts, the caducous basal bracts, the rounded subcordate

leaf-base, the tendency to intercostal veining, and the striate cuticle

of F. arnottiana. From F. costata it differs in the short lamina with

few lateral nerves and intercostals, the small fig with caducous

basal bracts, and the less plicate cuticle. The three species suggest

an ancient complex of the Asiatic mainland.

sect. Conosycea (Miq.) Corner subsect. Dictyoneuron Corner

ser. Validae (Mig.) Corner

F, sundaica Bl. Bijdr. (1825) 450.—F. indica sensu King, Ann.

R. Bot. Gard. Calc. 1 (1887) 39, pl. 45.—This is, as King

suggested, the same as the species for which he sought to main-

tain the name F. indica Linn. (see F. benghalensis, above).

Blume’s specimen is sterile but the venation and sunken stomata

leave no doubt: further, this common Ficus is not represented

among others of his from Java.

var. beccariana (King) Corner comb. nov.—F. korthalsii Miq.

Ann. Mus. Bot. Lugd. Bat. 3 (1867) 215, 286.—F. korthalsii

Mig. var. beccariana King, Ann. R. Bot. Gard. Calc. 1 (1887)

23, pl. 35A.—This variety is distinguished by the oblong or obo-

void fig with more unbonate disc of apical bracts. It often has a

shorter leaf and, when sterile, may be impossible to distinguish

from F. pellucido-punctata Griff (v. infra). Intermediates with

v. sundaica occur.

F. lowii King var. borneensis Corner v. nov.—Receptacula minora,

13-14 mm. lata, haud umbonata, bracteis apicalibus in discum

planum 3 mm. latum instructis. Lamina minus coriacea, costis

ad paginas ambas leviter elevatis, costis basalibus 4-1/3 laminae

elongatis. Glabra. Borneo (Kostermans 9758, G. Mentawir, Balik

Papan; typus, herb. Bogor).

var. minor Corner v. nov.—Receptacula 10 mm. lata, haud

umbonata, bracteis apicalibus in discum planum 3 mm. latum

instructis. Ramuli et stipulae pubescentes. Lamina rigide coriacea.

389

Gardens Bulletin, S.

costis basalibus 1/3—4 laminae elongatis. Malaya (Ridley 16176,

Wray’s Camp, G. Tahan, Pahang, typus herb. Singapore).

F. lowii is a rare species of Perak and these two collections indi-

cate that it must have a greater range and variability in the

mountains.

F. sumatrana Mig. var circumscissa Corner.—F. acamptophylla

sensu King, Ann. R. Bot. Gard. Calc. f* (1887) 40, pr. p.

Kunstler—Stipulae dense sericeae v. subtomentosae. Lamina

rigide coriacea, valde acuminata. Receptacula 8-10 mm. lata,

plus minus depresso-globosa, circum discum apicalem (2—3 mm.

latum) irregulariter circumscissa. Malaya.

Perak:—For. Dept. 39338, Piah For. Res.; Kunstler 4055

(Larut), 6106 (Gopeng), 10922 (Ulu Bubong); Ridley 10264,

Dindings; Wray 3788, Upper Perak. Trengganu:—Sing. F.n. 30357

(typus, herb. Singapore) et 30713, Bt. Kajang, Kemaman. Johore:

—Sing. F. n. 29937, Sedili River.

Haud F. acamptophylla Miq, forsan species nova.

var. microsyce Corner v. nov.—Receptacula parva, 4-6 mm.

lata (S—7 mm., vivo), disco apicali 2 mm. lato, bracteis basalibus

1.5-3 x 2-3 mm. Malaya, Sumatra, Java, Borneo, Celebes,

Philippines.

Malaya:—Curtis 3515, Pangkor, Perak; Ridley 13478, Sedenah,

Johore; Sing. F.n. 32241, Sedili River, Johore (typus, herb. Sin-

gapore). Sumatra:—Bangham 804, L. Tawar. Java:—Zollinger

1225. Borneo:—Endert 3216, W. Koetai; Clemens 26827, 26828,

Kinabalu; For. Dept. B.N.B. A4703, 10119, Kinabatangan. Cele-

bes:—Kjellberg 733, Kendari, Poehara. Philippines: Elmer

12386, Sibuyan (det. F. indica); Williams 226, Lamao River;

Coll. Agr. Laguna 611, Mt. Makiling.

F. binnendykii Migq. similis sed receptaculis non pertusis.

var. subsumatrama (Gagnep.) Corner comb. nov.—F. sub-

sumatrana Gagnep. Lec. Notul Syst. 4 (1927) 96.—Receptacula

7-10 mm. lata, bracteis basalibus 4-7 (-9) S-—8 mm. semi-

tecta, disco apicali 3—4 mm. lato. Indochina, Malaya, Sumatra,

Borneo. 3

Annam:—Poilane 1294 et 10786 (syntypes), 22415, 24034,

24159, 24475, 24487, 24605, 24670, 24683. Cochinchina:—

Poilane 19523. Malaya:—Burn Murdoch s.n., Feb. 1904, Se-

mangko Pass, Selangor. Sumatra:—Lorzing 5451, Sibolangit (det.

F. indica). Borneo:—Hallier s.n., Liang Gagang (Herb. Hort.

Bot. Bog. 3065); Kostermans 6994, Samarinda.

F. binnendykii Miq. var. cupulata Corner (v. infra) similis, sed

receptaculis non pertusis.

390

Vol. XVIT. (1959).

F. delosyce Corner sp. nov.—Arbor magna, epiphytica, suffocans,

cortice brunneo-carneo, foliis subdistichis, glabra v. receptaculis

puberulis. Ramuli 1-3 mm. crassi. Stipulae breves. Lamina 2.5—

8.5 x 1.2-4 cm., elliptica, raro obovata, subacuta v. breviter

acuminata, basi cuneato, tenue coriacea: costis lateralibus utrin-

secus 3-5, intercostis nullis, basalibus 1 et 1/3—4 laminae elon-

gatis: petiolo 6-13 mm. longo, gracili. Receptacula axillaria,

binata, sessilia, 7-14 « 6-8 mm., ellipsoideo-fusiformia, saepe

sparsim puberula, bracteis apicalibus subconicis: bracteis basali-

bus 2-3.5 2-2.5 mm., ovato-oblongis, subacutis, angustis:

setis internis nullis. Tepala 2 v. 3, libera. Flores sessiles v. pedi-

cellati.

var. delosyce

Malaya, Brunei, in silvis paludosis.

Singapore:—Sing. F.n. 37364, McRitchie Reservoir. Johore:——

Sing. F.n. 21337, 28178, 28607, (typus, herb. Singapore), 36800,

Sedili River; Ridley 13267, Tempayan River. Selangor:—Sing.

F.n. 30774, Pengkalan Kempas; 34033, Telok Reserve; 34143, S.

Tinggi; Burkill 1020, S. Buloh; Cons. For. 4870, Kuala Lumpur.

Negri Sembilan:—Ridley s.n., G. Angsi, Feb. 1904. Brunei:—

$5858, Seria; Brun. 5375, S. Belait.

var. obtusa Corner v. nov.—Lamina rotundato-obtusa. v.

subacuta. Borneo.

For. Dept. B.N.B. 2449, A3286, A3581, Sandakan; 4319,

Tawao; Kostermans 5681, Indonesian Borneo, E. Koetai, S.

Susuk (typus, herb. Bogor).

F. spathulifoliae Corner (v. infra) affinis sed costis paucis, re-

ceptaculis ellipsoideo-fusiformibus, bracteis basalibus parvis dis-

tincta.

F. spathulifolia Corner sp. nov.—Arbor mediocris, epiphytica, vix

suffocans, glabra v. stipulis puberulis, foliis spiraliter dispositis.

Ramuli 3—5 mm. crassi. Stipulae breves. Lamina 4-8 xk 2-4

cm., anguste. obovata v. spathulata, obtusa v. obtuse subacumi-

nata, basi cuneato, rigide coriacea, crassiuscula: costis lateralibus

utrinsecus 3—7, vix conspicuis, intercostis nullis: costis basalibus

utrinsecus 1, 3—3 laminae elongatis: petiolo 8-15 x 2-2.5 mm.,

applanato. Receptacula axillaria, binata, sessilia, 6-8 « 6-7

mm., subglobosa v. subobovoidea, umbonata, bracteis apicalibus

2 obtusis in discum convexum 3 mm. latum instructis: bracteis

basalibus 3—4 « 4—5 mm., ovatis, obtusis: setis internis nullis.

Tepala 2-3, libera. Flores masculi pedicellati, feminei et ceci-

diophori sessiles.

391

Gardens Bulletin, S.

var. spathulifolia

Malaya (Selangor), Sarawak, Brunei, in silvis paludosis. Selan-

gor:—Sing. F.n. 33970, Bt. Changgang, Klang (typus, herb.

Singapore); 34022, Telok Reserve, Klang. Sarawak:—-Anderson

9044, 9232, Binatang: W. M. A. Brooke 8778, Kelepu: Bur. Sci.

239; For. Dept. 7933, 8008, S. Kelepu, P: Bruit; Sing. F.n. 35686,

Manga Temulan. Brunei:—S5862, Seria.

var. annamensis Corner v. nov.—Ramuli 2—3 mm. crassi.

Lamina —10 X 5.2 cm., acuminata, apice subacuto: costis laterali-

bus utrinsecus 4—6. Bracteae basales minores, 1.5—2 « 2-3 mm.

(-3 xk 4mm., Poilane 23462). Tepala mascula praesertim 2. An-

nam, in silvis monticolis.

Poilane 22104, 22142 (typus, herb. Paris), 23462, 23749, prov.

Haut-Douaii, 1,000—1,200 m. alt.

var. substipitata Corner v. nov.—Ut v. spathulifolia sed recep-

taculis breviter pedicellatis, bracteis basalibus 1.5—2.5 mm. boneia

pedicellum obscurantibus. North Borneo.

For. Dept. B.N.B. A440, Sandakan (typus, herb. Kew); A1567,

Tawao.

Species lamina parva obtusa paucinervosa, receptaculis parvis

umbonatis, valde distincta, sed in v. annamense F. delosyce Cor-

ner approximans.

F. maclellandi King var. rhododendrifolia Corner v. nov.—F.

rhododendrifolia Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867)

286, non Kunth et Bouch. (1847).—Glabra, raro lamina in

juventute laxe pilosa. Bracteae basales 3, 2~3 mm. longae, vix

conditae. Himalayas, Khasia, Chittagong, North Burma, Pegu,

Yunnan, in silvis monticolis.

India: ——Brandis s.n., W. Duars; G. Mann s.n., Assam; Masters

s.n., Assam (typus, herb. Kew); Hooker s.n., Khasia et Chitta-

gong; Schlick s.n., Khasia. Burma:—-K. Ward 20427, Sumpra

Bum (det. F. benjamina).; Kurz 3135, Pegu. Yunnan:—Y. Tsiang

12332.

A var. maclellandi vix distincta sed adhuc raro collecta.

F. calcicola Corner sp. nov.—Arbor parva, epiphytica v. terricola,

foliis laxe spiraliter dispositis. Ramuli, petioli, stipulae, costarum

mediarum partes inferiores, bracteaeque basales pilis brunneis

piloso-tomentosi. Ramuli 2—3 mm. crassi. Stipulae —5 mm.

longae. Lamina 5-13 & 2.7-—5.5 cm., elliptica v. obovata, bre-

viter et obtuse v. subacute acuminata, basi cuneato, coriacea:

costis lateralibus utrinsecus 5~7 (—8), vix elevatis, intercostis

transversis 2—5: costis basalibus utrinsecus 1, 4-1/3 laminae

392

Vol. XVII. (1959).

elongatis: petiolo 7-22 mm. Receptacula axillaria, binata, ses-

silia, 7-10 mm. lata, subpyriformia v. subglobosa, glabra, brac-

teis apicalibus subprojicientibus in discum 2 mm. latum in-

structis: bracteis basalibus 1—2.5 mm. longis, ovatis, acutis, ap-

presse brunneo-pilosis: setis internis paucis, brunneis, v. nullis.

Tepala 3, raro 2 in floribus masculis vel 4 in femineis. Flores

masculi sparsi, breviter pedicellati, cecidiophoris breviores.

Burma, Thailand, Malaya, in collibus calcareis.

Malaya:—Burkill 6283, Tambun, Ipoh; Ridley 8518, 15th mile

Pahang track, Selangor; Kepong F. n. 37426, Kanching, Selangor;

Sing. F.n. 34388, Kanching (typus, herb. Singapore). Thailand:—

Kerr 130, 2394, Saraburi; 240, Prachuap; 2498, Sam Roi Yawt,

11376, Chumpawn; 19843, Krabin; Marcan 1738, Ratburi.

Burma:—Abdul Hak 22, Upper Burma, det. F. nitida.

F. microcarpae similis, sed pilis brunneis intercostisque differt:

forsan F. maclellandi affinis.

F. retusa Linn. Mant. (1767) 129; J. E. Smith, Rees Cycl. 14

(1810) n. 62.—Urostigma retusum (Linn.) Mig. Hook. Lond.

J. Bot. 6 (1847) 581.—U. truncatum Mig. Zoll. Syst. Verz.

(1854) 91, 97.—F. truncata Mig. Ann. Mus. Bot. Lugd. Bat.

3 (1867) 262, 286, non Vahl; King, Ann. R. Bot. Gard. Calc.

1 (1887) 41, pl. 48.—This name has been erroneously used for

F. microcarpa Linn. f. (v. infra). The type of F. retusa in the

Linnean herbarium, represented by a single good specimen, is

without doubt the species known as F. truncata Miq. Linneus

ascribed the specimen to India, where the species does not occur,

but there is no locality on the sheet, and the specimen agrees

exactly with the Javanese state of F. truncata; hence, Il assume

that it must have come from Java. As F. truncata is a later

homonym, a new name must be found for it and the only solu-

tion is to accept this disagreeable discovery.

var. borneensis Corner v. nov.—Lamina acute acuminata, tenue

v. valde coriacea, costis superne valde impressis, inferne prominen-

tibus: costis lateralibus utrinsecus 6-8, basalibus vix elongatis.

Bracteae basales 3—5 & 3—7 mm., 1/3—4 receptaculi obscurantes.

Tepala 2—3. Borneo, in silvis frequens.

Hallier 1020b, 1074, S. Sambas (1074, typus, herb. Leiden);

Kepong F.n. 80258, Beaufort, North Borneo; Main 1930, Danu

Pontoelok; Meijer 2463, 2582, Tarakan; Teysmann 7122, Blitong;

Corner s.n., 1959, Kuching.

A speciminibus javanicis varietatis retusae valde distincta, sed

malayana intermedia.

393

Gardens Bulletin, S.

sect. Conosycea (Mig.) Corner subsect. Dictyoneuron Corner

ser. Perforatae Corner

F. pellucido-punciata Griff. Notul. 4 (1854) 394; Ic. Pl. As. 4

(1854) t. 554. Syn.—F. gelderi Miq. Ann. Mus. Bot. Lugd.

Bat. 3 (1867) 216, 287.—F. indica Linn. var. gelderi (Miq.)

King, Ann. R. Bot. Gard. Calc. 1 (1887) 39, pl. 45.—Griffith’s

description and illustration of the ellipsoid fig with umbilicate

and half-open orifice shows that it is the same as F. gelderi. It

is represented at Kew by Griffith 4636, named F. pellucido-

punctata Griff., but of uncertain origin. Griffith gave Assam

(Gowahatti) in his description, but the species has not been

collected again from Assam. Herb. Helfer 4636 (Tenasserim

and Andamans), also named F. pellucido-punctata, is F. sun-

daica Bl. (== F. indica Linn. sensu King).

F. episima Corner sp. nov.—F. pellucido-punctatae affinis, sed

lamina disimillima. Ramuli 3—3.5 mm. crassi. Stipulae —20 mm.

longae. Lamina 6-10 X 3.2—6 cm., elliptica v. obovata, apice

obtuse rotundato v. subacuto, basi cuneato, rigide coriacea,

crassiuscula: costis lateralibus utrinsecus 3—5, basalibus 4—2/3

laminae elongatis: petiolo 12-20 mm. longo. Receptacula axil-

laria, binata, sessilia, -15 & 12 mm., late ellipsoidea, subtrun-

cata, ostiolo pertuso: bracteis basalibus 3—4.5 mm. longis,

ovatis, subacutis. Mindanao.

Elmer 11314, Mt. Apo (det. F. retusa); PNH 10045, Mt. Ka-

tanglad, Bukidnon; 10269, Surigao (typus, herb. Manila).

F. pisocarpa Bl. Bijdr. (1825) 454.—F. microstoma Wall. ex

King, Ann. R. Bot. Gard. Calc. 1 (1887) 38, pl. 44.—This un-

common species of Lower Thailand, Malaya, Java, and Borneo,

is close to F. pellucido-punctata. There is no specimen named

F. pisocarpa by Blume at Leiden, but there are three which he

must have seen, namely one collected by Reinwardt, another

by Blume himself, and the third labelled “H.L.B. 43 Gedeh”,

which is the type-locality of F. pisocarpa. These sheets were

un-named, but they are the same as F. microstoma, which fits

well with Blume’s description, particularly “fructibus pisiformi-

bus . . . sessilibus plano-umbilicatis”. No other Javanese banyan

fits this description and, as F. microstoma is the one species as

yet unaccounted for among Blume’s names, I have no hesitation,

after working through the Malaysian banyans, in identifying it

with F. microstoma.

King misinterpreted F. pisocarpa by identifying it with Kunstler

3555, which is F. caulocarpa Miq, having stipitate figs, not plano-

umbilicate, and different leaves. _

394

Vol. XVI. (1959).

F. binnendykii Mig. var. coriacea Corner v. nov.—Urostigma

tjiela Miq. Hook. Lond. J. Bot. 6 (1847) 580, pr. p. Cuming

1931.—Lamina 4-11 x 2-5 cm., anguste v. late elliptica, lan-

ceolata, rigide coriacea, crassiuscula, venis obscuris: costis late-

ralibus utrinsecus 3—5, basalibus 1/3-4 laminae elongatis.

Bracteae basales 2.5—-4 mm. longae. Annam, Malaya, Bangka,

Sarawak, Brunei, Luzon.

Annam:—Poilane 32798, Nui Dai Ding, prov. Koutum.

Malaya:—Singapore, Burkill 2031, Langlasse 313, 328, Ridley

3402, 5639, 5640, 6459, 9219, 10382, 10808, 12203, 14155,

Sinclair 4917, Sing. F.n. 33560; Johore, Ridley 12162, 13267,

Sing. F.n. 36562; Malacca, Cons. For. 2061; Perak, Cons. For.

2313, King’s coll. 6106, 10505, Scortechini 310, 322, 1051, Sing.

F.n. 31655 (typus, herb. Singapore). Bangka:—-Kostermans et Anta

172, 534, 1076. Sarawak:—Clemens 22318, Mt. Matang. Brunei:

—Brun. 542, Andulau. Luzon:—Bur. Sci. 22275, Cuming 1931.

F. sumatranae persimilis sed ostiolo pertuso, bracteis basalibus

parvis.

var. cupulata Corner v. nov.—Ut. var. binnendykii sed brac-

teae basales 4-6 « 5-8 mm., 1/3—% receptaculi obscurantes.

Lamina plus minus coriacea. Celebes.

Kjellberg 2769, Matano (typus, herb. Stockholm); 2823,

Sorvako.

var. latifolia Corner v. nov.—Ut var. binnendykii sed lamina

6-13 4—6.5 cm., late elliptica v. ovato-elliptica, basi rotundato

v. late cuneato: costis lateralibus utrinsecus 3—4. Borneo.

Beccari 3353, Pontianak, det. F. cycloneura (typus, herb. Flo-

rence); San. 17172, Tawao; Teysmann 7111, Kapoeas, det F.

cycloneura.

sect. Conosycea (Miq.) Corner subsect. Benjamina Miq.

ser. Benjamineae Mig.

F. subcordata Bl. Bijdr. (1825) 440.—¥F. garciniifolia Mig. Ann.

Mus. Bot. Lugd. Bat. 3 (1867) 218, 287.—F. calophylloides

Elm. Leafl. Philip. Bot. 4 (1911) 1246.—F. acrorrhyncha Sum-

merh. J. Arn. Arb. 13 (1932) 98.—F. fairchildii Backer, Blu-

mea 6 (1947-8) 306.—Urostigma balicum Miq., U. subcor-

datum (Bl.) Miq. Fl. Ind. Bat. I, 2 (1859) 348, 349.—This

is an uncommon, but wide-spread, species from Tonkin to New

Hebrides, yet I can see but slight differences in the size of leaf

and fig in the collections (35 in all), and I have compared the

_ types. Reinwardt 1338 (Timor) is the original F. subcordata

BL, labelled “arbor elegans ante domum in villa Koepang

395

Gardens Bulletin, S.

Timoriensium”. The specimen at Utrecht is labelled “F. subcor-

data. Timor” by Reinwardt; that at Leiden is labelled both by

Blume and Miquel and bears the note “Zipp. ?”. Both are sterile,

but the venation, cystoliths, and sunken stomata clinch the

identity with F. garciniifolia, the type of which also came from

Timor (de Vriese, Koepang). |

var. malayana Corner v. nov.—Receptacula majora, 35-50 x

20-25 mm., subcylindrica: bracteis basalibus annulato-conjunctis,

annulo 0.5—! mm. lato, integro vel obscure sublobato. Flores mas-

culi pauci, pedicellis 1.5—-5 mm. longis. Malaya, North Borneo.

Malaya:—-Curtis s.n., May 1902, road to Gap, Selangor (typus,

herb. Singapore); For. Dept. 6547, Kampong Bahru, Kuala Lum-

pur. Borneo:—San. 4365, Mt. Trus, Keningau distr.

F. benjamina Linn. var. bracteata Corner v. nov.—Bracteae

basales 2-3 « 2.5—-4 mm., haud conditae. Cochinchina, Cele-

bes, Morotai, Sumbawa, Timor, Philippines, New Guinea, New

Britain.

Cochinchina:—Poilane 17349, Budop. Celebes:—Kyjellberg

1386 (Eurekang), 1669 (Makule), 2290 (Timampoe). Morotat:

—Kostermans 780, 807, 1514. Sumbawa:—Elbert 3997, 4087.

Timor:—Forbes 3796. Philippines (Luzon, Negros, Batan, Min-

doro):—-Bur. Sci. 80168, 80300; Coll. Agr. Laguna 508, 557,

568; Cuming 1936; Elmer 10153 (det. F. indica), 16038; PNH

9482, 16897, 33342; Wenzel 1411, 1470. New Guinea:—Brass.

21616, 21902; Carr 16359 (typus, herb. Br. Mus.); Doctors v.

Leeuwen 9534; Hoogland 4167, 4269. New Britain: —NGF

7038.

sect. Conosycea (Miq.) Corner subsect. Benjamina Mig.

ser. Callophylleae Corner

F. callophylla Bl. var. leytensis Corner v. nov.—F. pachyphylla

Merr. Philip. J. Sci. Bot. 8 (1913) 365, non King (1887).—

Receptacula obovoidea, bracteis apicalibus in umbonem acutum

instructis, bracteis basalibus 7-9 x 7—8 mm. acutis. Leyte (Wen-

zel 209, typus varietatis etiam F. pachyphyllae Mertr.).

var. malayana Corner v. nov.—Lamina saepe major, valdius.

coriacea, venis quasi obscuratis. Receptacula majora, 14-17 x

15—20 mm., bracteis basalibus 8-10 * 7-12 mm. Tepala mascula

(3—)4. Thailand, Malaya, Sumatra.

Thailand:—J. Schmidt 537; G. Seidenfaden 2079. Malaya:—

Hullett 329; King’s coll. 4697. Sing. F.n. 21190 (typus, herb.

Singapore), 28634, 28657, 31662, 34144, 34460, 34464, 36955,

37987. Sumatra:—Forbes 1735, 2590.

396

Vol. XVIT. (1959).

var. minor Corner v. nov.—Receptacula minora 7-8 x 7-10

mm., bracteis basalibus 3—4 mm. longis. Celebes, Sumbawa.

Celebes:—Boschproefst. Cel/III-22, Cel/IV—114 (typus, leg.

27.VIII.32, herb. Leiden), Malili; Koorders 19199, 19329,

19337, Minahassa (det. F. retusa); Teysmann 12142, Pangkad-

jena. Sumbawa:— Elbert 3689.

F. cartipes Corner nom. nov.—F. obtusifolia Roxb. Fl. Ind. 3

(1832) 546, non HBK; King, Ann. R. Bot. Gard. Calc. |

(1887) 42, pl. 49.

F. tristaniifolia Corner sp. nov.—Arbor mediocris, epiphytica, vix

suffocans, foliis spiraliter dispositis, glabra. Ramuli 3—5 mm.

crassi. Stipulae —22 mm. longae. Lamina 4.5—10.5 & 1.5-5 cm.,

anguste obovata v. spathulata, apice obtuso subtruncato

v. subretuso, basi anguste cuneato, valdissime coriacea,

ccrassiuscula, marginibus recurvis, sicco ferruginea, venis quasi

obscuratis: costis lateralibus utrinsecus 7—9, obliquis, secundartis

vix minus evolutis: costis basalibus utrinsecus 1, 1/3—4 laminae

elongatis: petiolo 5-12 « 2 mm., brevi, applanato. Receptacula

axillaria, binata, sessilia, 8-10 mm. lata, subglobosa, bracteis

apicalibus 2 in discum planum 2 mm. latum instructis: bracteis

basalibus 3—5 mm. longis, 1/3—+ receptaculi obscurantibus; setis

internis nullis. Tepala 3, libera. Flores masculi et cecidiophori

pedicellati, feminei sessiles. Malaya, Sumatra, Borneo, in silvis

paludosis rara.

Malaya:—Ridley 1597, Malacca, S. Udang; Sing. F.n. 36606,

36698, Johore, Pontian, Penkalan Raja (36698 typus, herb. Singa-

pore). Sumatra:—Bruinier 306, East Coast, S. Rawa. Borneo:—

Buwalda 7852, Sampit; NIFS bb. 32411, Sampit; v. Romburgh

11, Paloh.

F. spathulifolia Corner (v. supra) facile confusa, sed foliis val-

dius coriaceis, venis ut in F. callophylla, receptaculis haud

umbonatis..

F. microcarpa Linn. f. Suppl. Spl. Pl. (1781) 442.—F. retusa

auct., F. nitida auct., F. retusa var. nitida auct., non verae.——

This is the plant generally, but wrongly, called F. retusa, F.

nitida, or F. retusa var. nitida. F. retusa Linn. is the species

generally called F. truncata Mig. (v. supra), while F. nitida

Thunb. is F. benjamina Linn., and F. benjamina sensu Thunb.

is this F. microcarpa. } have disentangled the confusion by study

of the types and original specimens. F. microcarpa, strangely

enough, never reached Carolus Linneus. The name-change is

397

Gardens Bulletin, S.

jarring, for this is one of the more widely cultivated species, but

in the absence of Nomina Specifica Conservanda aut Rejicienda

there is no way out. It ranges from Ceylon, India, South China,

and Ryu Kyu islands to Australia and New Caledonia.

var. microcarpa f. pubescens Corner f. nov.—Stipulae et

bracteae basales dense appresse puberulae. Ramuli et receptacula

subvillosa. Thailand, in collibus calcareis.

Kerr 2168, Chawng Ke, N. Sawan (typus, herb. Br. Mus.);

10649, Chawn Bung, Rotburi; Kostermans 681, Kwae Noi Basin

Exp., Brongkasi; 1441, Kin Sayok (120 km. n.w. Kwanburi).

_ var. eubracteata Corner v. nov.—Bracteae basales majores 4—5

mm. longae et latae, ovato-acutae. Lamina —11 x 5.5 cm., late

elliptica v. ovato-elliptica. Thailand.

RFD 16711, Chiengmai, Chiengdao (typus, herb, Bangkok);

11990, Lampang, Ngao. Mae Huat.

Ut. var. latifolia sed bracteis basalibus quasi duplice majoribus.

_ var. hillii (Bailey) Corner comb, nov.—F. hillii F.M. Bailey.

Contr. Queensl. Fl. 3 (1891) 16.—¥. schlechteri Warb. Tropenfi.

7 (1903) 582.—Ut v. mocrocarpa sed costis basalibus vix elon-

gatis, setis internis quasi nullis. Sunda Isl. (Alor Kechil), New

Guinea, Queensland, New Caledonia, Loyalty Islands.

Forsan a v. microcarpa vix distincta.

var. latifolia (Mig.) Corner comb. nov.—Urostigma accedens

Mig. v. latifolia Miq. Fl. Ind. Bat. 1, 2 (1859) 347.—F. rigo

F.M. Bailey, F. thynneana F.M. Bailey, Queensl. Agr. J. 1 (1897)

231, 235.—F. retusa Linn. v. rigo (Bailey) Diels, Engl. Bot.

Jahrb. 67 (1935) 183.—Ut v. microcarpa sed lamina -—12 « 9

cm., late elliptica, apici obtuso v. subacuminato, basi rotundato v.

late cuneato, venis subtus conspicuis, costis secundariis ut pri-

mariis quasi bene evolutis. Celebes. Moluccas, New Guinea,

Queensland, Caroline Islands.

Celebes:—Nielsen 804. Moluccas:—Reinwardt s.n., Halawahoea

(typus, herb. Leiden); de Vriese, Ternate; Alston 16649, Ternate.

New Guinea:—Brass 3592, Rona, Laloki River; Carr 11441,

Hisiu; Carr 11588, Veiya; NGF 8087, Port Moresby; Hellwig

187; Labillardiere, Waigeo Isl. Caroline Isl.:—Fosberg 25815,

25821; 25895, 31999,'32023.°E-Y. Hosaka 3351:

var. naumanni (Engl.) Corner comb. nov.—F. naumanni Engl.

Bot. Jahrb. 7 (1886) 451.—¥F. dahlii K. Schum. Notizbl. Bot.

Gart. Mus. Berlin 2 (1898) 111.—F. regnans Diels, Engl. Bot.

Jahr. 67 (1935) 182.—Receptacula pedunculo 1-5 mm. longo

praedita, bracteis basalibus caducis. New Guinea, New Britain,

New Ireland.

398

Vol. XV IGP).

New Guinea:—Brass 21926, Cape Vogel Pen., Menapi; 27781,

Sudest Isl., Joe Landing; Carr 12342, 12484, Rouna; Ledermann

7476, 9491, 10844, Sepik; Nyman 233, Stephansort; coll. cet. cit.

Engler, Segaar et MacCluer Bay. New Britain:—Dahl s.n., Ralum,

v. Hugel 572, Blanche Bay; Lauterbach 215, Ralum. New Ireland:

—Naumann s.n., Cartaret Bay (typus, herb. Berlin).

var. saftordii (Merr.) Corner comb. nov.—F. saffordii Merr.

Philip. J. Sci. Bot. 9 (1914) 73.—F. prolixa Forst. f. v. saffordii

(Merr.) Fosberg, Phytologia 5 (1955) 289.—Lamina -8.5 x

4.5 cm., ovato-cordata v. subcordata, apice acuminato, costis basa-

libus 4-1/3 laminae elongatis. Ins. Caroline, Marianas.

Marianas:—Anderson 235; Fosberg 25159, 31873, 35359;

McGregor 414 (typus); Nelson 150, det. F. tenuistipula Caroline

Is].:—Fosberg 32166; Kanehira 1907.

F. prolixa facile confusa sed cystolithis amphigenis, superne ut

verrucellis in lamina sicca, in F. prolixa modo in pagina inferior.

F. palaquiifolia Corner sp. nov.—Arbor magna. epiphytica, suf-

focans, foliis spiraliter dispositis, glabra. Ramuli 4-5 mm. crassi.

Stipulae —24 mm. longae. Lamina 7-12 & 3-6.5 cm., elliptica

v. subobovata, breviter et acute acuminata, basi cuneato, cori-

acea, sicco brunnea: costis lateralibus primariis utrinsecus 7—13,

vix elevatis, subcrenulatis, secundariis vix minus evolutis, inter-

costis nullis: costis basalibus utrinsecus 1, vix elongatis: petiolo

18-38 2-3 mm., sicco brunneo. Receptacula axillaria, binata,

sessilia, 9-10 mm. lata (vivo, 12-15 « 10-13 mm.), subpyri-

formia v. subglobosa, sicco subverrucosa (pariete pertenui ad

semina contracto), bracteis apicalibus 2—3 in discum convexum

v. subconicum 2—3 mm. latum instructis: bracteis basalibus 7-10

mm. longis, ovatis, obtusis, quasi + receptaculi obscurantibus:

setis internis nullis. Tepala 34, lineari-lanceolata, libera. Flores

masculi pedicellati, feminei et cecidiophori sessiles. Semina crasse

tunicata. Borneo.

Sing. F. n. 26703, 27413 (typus, herb. Singapore), Kinabalu, c.

1200 m. alt.; Hallier 3065, Liang Gagang.

F. polygramma, F. archboldiana, et F. benjaminoides affinis.

F. polygramma Corner sp. nov.—Arbor epiphytica, suffocans,

foliis spiraliter dispositis, glabra v. ramuli, stipulae, petioli, et

receptacula puberuli. Ramuli 3 mm. crassi. Stipulae 15—25 mm.

longae. Lamina 6-9 « 3—5.5 cm., elliptica, breviter acuminata,

basi late cuneato, coriacea, sicco brunnea: costis lateralibus

399

Gardens Bulletin, S.

primariis utrinsecus 9-13, ad paginas ambas leviter elevatis,

secundariis vix minus evolutis, intercostis nullis: costis basalibus

utrinsecus 1, haud elongatis: petiolo -25 « 2.5 mm., nigro.

Receptacula axillaria, binata, sessilia, 15 % 10-13 mm., pyri-

formia v. ellipsoidea, apice subtruncato v. obtuso, bracteis api-

calibus 3 in cono brevi 1.5 mm. lato instructis: bracteis basalibus

34.5 mm. longis, ovatis, obtusis v. subacutis, crassiusculis, ad

basim usque 1—2 mm. breviter et infundibuliforme connatis:

setis internis nullis. Tepala 3, spathulata, libera. Flores masculi

pedicellis subinduratis praediti, feminei et cecidiophori plus

minus sessiles. Celebes. |

Kjellberg 2702, Porema, c. 1200 m. alt. (typus, herb. Stock-

holm); Koorders 19429, 19275, Minahassa.

F. archboldianae Summerh. affinis sed bracteis basalibus con-

natis differt.

F. benjaminoides Corner sp. nov.—Arbor magna, epiphytica, suf-

focans, foliis spiraliter dispositis, glabra sed bracteis basalibus

puberulis. Ramuli 34.5 mm. crassi, suberecti. Stipulae -20 mm.

longae. Lamina 5-13 x 2.3—5.7 cm., anguste elliptica v. ovato-

elliptica, acuta v. subacuminata, basi cuneato, tenue coriacea,

sicco fusco-brunnea: costis lateralibus primariis utrinsecus 7—13,

leviter elevatis, secundariis vix minus evolutis, intercostis nullis:

costis basalibus utrinsecus 1—2, haud elongatis: petiolo 7—24

mm. longo, nigro. Receptacula axillaria, binata, sessilia, 5-6

mm. lata (6-7 x 7-8 mm., vivo), subglobosa, plus minus

depressa, subumbonata, bracteis apicalibus 2 in discum planum

v. subconvexum 2—2.5 mm. latum instructis: bracteis basalibus

2.54 * 3-6 mm., ovatis, obtusis, puberulis: setis internis nullis.

Flores masculi pedicellati, sparsi v. aggregati, tepalis 2—3 spa-

thulatis. Flores cecidiophori breviter pedicellati v. sessiles, tepalis

2—3; feminei sessiles, tepalis 3-4. Nova Guinea.

Brass 5433, Terr. N. Guinea, Central div., Mafulu, c. 1250 m.

alt; BW 6592, Neth. N. Guinea, Asmat region; Carr 12093,

Papua, Koitaki, c. 400 m. alt. (typus, herb. Br. Mus.).

F. benjaminae similis sed foliis spiraliter dispositis sicco fusco-

brunneis, ramulis suberectis, receptaculis minoribus, disco apicali

latiori.

F. patellata Corner sp. nov.—Arbor magna, epiphytica, sufiocans,

foliis spiraliter dispositis, glabra. Ramuli 2.5-3 mm. crassi. Sti-

pulae —11 mm. longae. Lamina S—-9 x 2.5-5 cm., elliptica,

400

Vol. XVII. (1959).

breviter acuminata, basi late cuneato, coriacea, sicco brunnea:

costis lateralibus primariis utrinsecus 13-16, gracilibus, vix

elevatis, secundariis vix minus evolutis, intercostis nullis: costis

basalibus utrinsecus 1, haud elongatis: petiolo 9-18 x 1.5-2

mm., nigro. Receptacula axillaria, binata, sessilia, 6-7 mm. lata,

subglobosa, bracteis apicalibus 3 in discum planum 2 mm. latum

instructis: bracteis basalibus in disco integro tumido patelli-

formi 6—7 mm. lato conjunctis: setis internis nullis. Flores mas-

culi breviter pedicellati, tepalis 2. Flores cecidiophori sessiles

v. subpedicellati, feminei sessiles, tepalis 3. Nova Guinea. (Exp.

Lundquist, Aet 141, McCluer Bay, Jakati, pr. Bobo, typus herb.

Leiden).

F. benjaminoides affinis sed receptaculo ad discum patellifor-

mem instructo facile distinguitur.

sect. Maivanthera Corner ser. Malvanthereae Corner

subser. Eubracteatae Corner

F. triradiata Corner sp. nov.—Arbor magna, epiphytica, suffocans,

glabra. Ramuli 3—6 mm. crassi. Stipulae —6 cm. longae, caducae.

Lamina 7-14 3.5-5.8 cm., elliptica v. obovato-elliptica, sub-

acuta, basi cuneato, rigide coriacea, margine plus minus recurvo:

costis lateralibus primariis utrinsecus 6-10, haud elevatis, secun-

dariis minus conspicuis: costis basalibus utrinsecus 1, haud v.

vix elongatis: petiolo 15-30 mm. longo, subapplanato. Recepta-

cula axillaria, binata: pedunculo 4-6 x 5-6 mm., apice —10

mm. lato incrassato: bracteis basalibus 3, 5-7 x 7-10 mm.,

ovatis, obtusis, persistentibus: receptaculi corpore 18-22 « 14—

17 mm., truncato-subconico, ostiolo profunde triradiato breviter

mammiullato, pariete 2—3.5 mm. crasso subligneo: setis internis

nullis. Flores masculi pedicellati, tepalis 4 spathulatis liberis,

anthero reniformi crescentiforme dehiscenti. Flores feminei et

cecidiophori sessiles v. breviter pedicellati, tepalis 4—5 lanceo-

latis: ovario libero rubro, stigmate simplici. Queensland.

C. T. White 10536, Mt. Spurgeon (typus, herb. Brisbane); H.

Flecher (N.Q. Naturalists Club n. 7260) Brooklyn-Mt. Lewis track.

var. sessilicarpa Corner v. nov.—Gracilis, laminis receptaculisque

minoribus. Ramuli 3-4 mm. crassi. Lamina 6-12 x 2-4 cm.,

anguste elliptica v. lanceolata, costis primariis lateralibus utrin-

secus 5~9. Receptacula 12-15 mm. (18—20 mm., vivo), subglobosa,

plus minus sessilia: bracteis basalibus S-7 8—10 mm. Queens-

land.

Brass 20052, Mt. Finnegan, 850 m. alt. (typus, herb. Brisbane).

401

Gardens Bulletin, S-

sect. Malvanthera Corner ser. Malvanthereae Corner

subser. Malvanthereae

F. glandifera Summerh. var. brachysyce Corner v. nov.—Recep-

tacula 10-13 mm. longa, subglobosa v. obovoidea, ad basim

cupula adnata 2.5—-5 mm. alta et 10-13 mm. lata inserta. La-

mina 4-14 x 2-6 cm., elliptica, costis lateralibus primariis

utrinsecus 11—15. Tepala feminea et cecidiophora 1—3. Celebes,

Nova Guinea.

Celebes:—Elbert 2945, Muna Isl., Lombai, S.E. Celebes. New

Guinea:—NGF 5218, (typus, herb. Lae), 5232, 7422, Bulolo,

Morobe distr., 3500 ped. alt.

sect. Malvanthera Corner ser. Malvanthereae Corner

subser. Platypodeae Corner

F. leucotricha Mig. var. megacarpa F.v.M. ex Corner v. nov.—

Ramuli crassior, 5-7 mm. Receptacula majora, 14 mm. lata.

pedunculo 8—14 mm. longo: bracteis basalibus multo majoribus,

~10 x 12 mm., apiculatis. Queensland (F. v. Mueller, Sea

Range, typus herb. Kew).

Specimina in herb. Kew et Utrecht det. “F. lanata sp. nov. OC

megacarpa”’ a F. v. Mueller, sed inedita. In v. leucotricha bracteae

basales 4-5 mm. longae.

var. sessilis Corner v. nov.—Receptacula sessilia: bracteis basa-

libus -10 & 7 mm., ovato-caudiculatis. New South Wales (speci-

men typicum in herb. Manchester “J. E. Smith N.S.W.” notatum).

F. obliqua Forst. ¢. var. petiolaris (Benth.) Corner comb. nov.—

F. platypoda A. Cunn. ex Mig. var. petiolaris Benth. Fl. Aus-

tral. 6 (1873) 169.—This has the larger figs of F. platypoda,

but the two basal bracts which distinguish F. obliqua. Some col-

lections of F. obliqua have figs of intermediate size. As a detail,

the leaves of var. petiolaris have the smooth lower epidermis of

F. obliqua, not the finely striate epidermis round the stomata

as in F. platypoda. The collection “Fraser, Brisbane and Has-

tings River’, cited by Bentham is to be excluded because it is

F. macrophylla Desf.

¥. obliqua Forst. f. var. puberula (Benth.) Corner comb. nov.—

F. eugenioides F.v.M. var. puberula Benth. Fl. Austral. 6

(1873) 167.—Urostigma brachypodum Miq. Hook. Lond. J.

Bot. 6 (1847) 562.—F. brachypoda Mig. Ann. Mus. Bot.

Lugd. Bat 3 (1867) 287.—Ramuli et petioli puberuli. Lamina

2.5-6 & 1.12.3 cm., lanceolato-elliptica, subacuta, basi. an-

guste et abrupte rotundato, costis lateralibus primariis utrinsecus

402

Vol. XVII. (1959).

8-9: petiolo 3-5 mm. longo. Receptacula 5—6 mm. lata, breviter

pedunculata. Queensland (A. Cunningham 304, York Sound,

typus herb. Kew).

F. platypodae vy. minoris similis sed puberula et receptaculum

minus.

F. subpuberula Corner nom. nov.—F. puberula A. Cunn. ex Miq.

Ann. Mus. Bot. Lugd. Bat. 3 (1867) 237, (non Kunth et

Bouch., 1847).—This little known species may be a variety of

F. platypoda.

F. platypoda A. Cunn. ex Mig. Ann. Mus. Bot. Lugd. Bat. 3

(1867) 287; Benth. Fl. Austral. 6 (1873) 169.—Urostigma

platypodum Miq. Hook. Lond. J. Bot. 6 (1847) 561.—Miquel

cited originally three collections :—

1. Cunningham, York Sound, (Herb. Hook), which is the

type;

2. Fraser, Brisbane and Hastings River, called “forma major”,

which Bentham transferred to F. platypoda var. petio-

laris Benth., but which is F. macrophylla Desf., and

must be excluded;

3. Bynoe (Herb. Hook), called ‘forma minor” which is F.

platypoda var. minor Benth.

sect. Malvanthera Corner ser. Malvanthereae Corner

subser. Hesperidiiformes Corner

F, augusta Corner sp. nov.—Arbor magna, epiphytica, suffocans.

Ramuli, stipulae, et saepe receptacula albido-puberuli, glabres-

centes. Ramuli 7-10 mm. crassi. Stipulae —-22 cm. longae, vivo

rubrae. Lamina —26 « 17 cm., ovato-subcordata v. late ellip-

tica, subacuta v. breviter acuminata, basi rotundato-cordato v.

late cuneato, coriacea: costis lateralibus primariis utrinsecus 12—

17, vix elevatis, secundariis minus conspicuis: costis basalibus

utrinsecus | (—2), 4-1/3 laminae elongatis: petiolo 30-100 x«

3-5 mm. Receptacula axillaria, binata, maturitate rubescentia;

pedunculo 4-6 « S5—7 mm. v. 20-25 « 8-10 mm., apice haud

v. vix dilatato: bracteis basalibus 3, 1-2.5 < 2-3.5 mm., cres-

centiformibus, denique caducis: receptaculi corpore 50-80 x

27-50 mm., oblongo-ellipsoideo, apice umbonato (5-7 x

10-17 mm.), bracteis apicalibus inflexis 2—3 gibbosis: setis in-

ternis nullis: pariete crasso, lignoso, processibus floriferis

praedito. Flores ut in F. xylosycia Diels, tepalis 3—4. Flores

403

Gardens Bulletin, S.

masculi pedicellati, anthero reniformi crescentiforme dehiscenti

sessili. Flores feminei sessiles, ovario in receptaculi pariete semi-

immerso, stigmate longe bifido. Nova Guinea.

Hoogland 5454, Terr. N. Guinea, Eastern Highlands, Goroka

subdistr., Asaro Mairi divide, 2400 m. alt. (typus, herb. Lae);

NGF 1020, Aiyura area.

F. hesperidiiformis King var. sisritek sea in (Summerh.) Corner

comb. nov.—F. myrmekiocarpa Summerh. J. Am. Arb. 22

(1941) 82.—This has somewhat smaller figs with more ex-

panded apex to the peduncle than typical F. hesperidiiformis,

but I can find no essential difference.

F. xylosycia Diels var. cylindrocarpa (Diels) Corner comb. nov.—

F. cylindrocarpa Diels, Engl. Bot. Jahrb. 67 (1935) 181—

This has a smaller, narrowly cylindric fig with slender peduncle,

but it grades into typical F. xylosycia, as in Carr 14050

(Papua).

404

Taxonomic Notes on Ficus Linn., Asia

and Australasia

If. SUBGEN. PHARMACOSYCEA MIQ.

By E. J. H. CORNER

Botany School, University of Cambridge

Summary

NEW SECTIONS, SERIES, and subseries:—sect. Oreosycea (Midq. )

Corner comb. nov., ser. Vasculosae Corner, subser. Vasculosae,

subser. Albipilae Corner, ser. Nervosae Comer, ser. Austrocale-

donicae Corner.

Earlier specific epithets: —F’. albipila (Miq.) King (F. colossea

F. v. Muell. ex Benth, F. microtricherinos Backer); F. habrophylla

G. Bennett (F. edulis Bur.).

New name for later homonym:—F. pachystemon Warb. (F.

mangiferifolia K. Schum. et Laut., haud Griffith).

New species:—F. gratiosa Corner, F. pachysycia Diels ex Cor-

ner, F. hombroniana Corner, F. subnervosa Corner, F. kjellbergii

Corner, F. madhucifolia Corner, F. pseudojaca Corner.

New varieties: —F. albipila (Miq.) King v. glabra Corner, F.

gratiosa Corner v. caudata Corner, F. pubinervis Bl. v. diandra

Corner et v. sibulanensis (Elm.) Corner, F. subtrinervia K.

Schum. et Laut. v. doormaniana (Diels) Corner, F. granatum

Forst. f. v. minor Corner, F. smithii v. robusta Corner, F. austro-

caledonica Bur. v. balansaeana Corner.

Notes on F. hadroneura Diels.

subgen. Pharmacosycea Migq.

Ann. Mus. Bot. Lugd. Bat. 3 (1867) 299.—Pharmacosycea

Mig. Hook. Lond. J. Bot. 6 (1847) 525; id. 7 (1848) 64.—

Monoecious, all three kinds of flower in the same receptacle.

Receptacles without lateral bracts on the body: interfloral bracts

often present. Male flowers ostiolar or disperse: stamens 1-3,

occasionally with pistillode. Gall- and female flowers similar,

but the female generally sessile with longer style: ovary white or

with a red spot at the base, sessile: style glabrous, stigma bifid.

405

Gardens Bulletin, S-

Seed smooth, often keeled over the apex. Trees, rarely shrubs (not

epiphytic, creeping, or climbing). Leaves generally entire and with

a gland in the axil of each main basal nerve. Lectotype:—F.

maxima P. Mill. (= F. radula Willd.).

sect. Pharmacosycea.—Ficus sect. Pharmacosycea (Miq.) Benth.

et Hook. Gen. Pl. 3 (1880) 369.—Fig solitary, orifice in some

species crateriform: interfloral bracts present: internal bristles

absent. Male flowers disperse: stamens 2—3, the filaments some-

times shortly joined. Ovary often with a red mark at the base on

the stylar side. Cystoliths amphigenous. Stomata superficial or

sunken. Microscopic gland-hairs with 4—6—12-celled head. Tro-

pical America, c. 12 species: lectotype F. maxima P. Mill.

sect. Oreosycea (Miq.) Corner comb. nov.—Urostigma Gasp. sect.

Oreosycea Mig. Hook. Lond. J. Bot. 6 (1847) 525, 585: Fl.

Ind. Bat. I, 2 (1859) 353.—Ficus Linn. subgen. Urostigma

(Gasp.) Miq. sect. Oreosycea Miq. Ann. Mus. Bot. Lugd. Bat.

3 (1867) 286.—Ficus Linn. sect. Leiosycea Miq. Hook. Lond.

J. Bot. 7 (1848) 454.—Ficus Linn. subgen. Urostigma (Gasp.)

Mig. sect. Stilpnophyllum Endl. subsect. Pedunculatae Sata,

Monogr. (1944) 179.—Figs typically paired, orifice not cra-~

teriform: interfloral bracts and internal bristles present or not.

Male flowers disperse or ostiolar: stamens 1 or 2, rarely 3.

Ovary without a red spot. Cystoliths mostly hypogenous, in a few

species amphigenous. Stomata not sunken. Microscopic gland-

hairs with 2—4-celled head. Madagascar, Asia, Australasia, 45

species: lectotype—F. nervosa Heyne ex Roth.

ser. Vasculosae Corner ser. nov.—Folia exsiccata griseo-viridia

v. pallide brunnea, nervis reticulatis saepe ad paginam unam vel

ambas elevatis, glandulis basalibus saepe nullis v. indistinctis. Re-

ceptaculum pedunculo distincto praeditum, cellulis scleroticis in

pariete copiosis. Stamina plerumque 2 in floribus ostiolaribus (1

in F. gratiosa). Madagascar, Asia, New Guinea, Queensland, 7

species. Typus—F. vasculosa Wall ex Miq.

subser. Wasculosae.—sect. Leiocarpa Migq. l|.c. (typus—F. vas-

culosa Wall.)—-subsect. Pedunculatae Sata |.c. (lectotypus F. cal-

losa Willd., = F. malunuensis Warb.).—Folia nitida coriacea,

nervis intercostalibus numerosis (F. callosa) v. 0-1. Setae inter-

nae nullae (numerosae in F. gratiosa). Cystolitha hypogena.

Sempervirens. Asia, 4 species. Typus—F. vasculosa Wall.

subser. Albipilae Corner subser. nov.—Folia haud nitida, mem-

branacea v. chartaceo-corlacea, nervis intercostalibus 3-numerosis.

406

Vol. XVII. (1959).

Setae internae copiosae v. nullae. Arbores deciduae, saepe albi-

pilosae. Madagascar, Andaman Is]., Indochina, Thailand, Malay-

sia, Queensland, 3 species. Typus—F. albipila (Miq.) King.

ser. Nervosae Corner ser. nov.—Folia exsiccata saepissime fusco-

brunnea, leniter v. rigide coriacea, glandulis basalibus 2 conspi-

cuis, stipulis saepe elongatis. Receptaculum pedunculo saepe

brevi praeditum. Stamen 1, vel 2 (—3). Arbores grandes, ? sem-

pervirentes. Asia, New Guinea, Solomon Islands, 16 species.

Typus.—F. nervosa Heyne ex Roth.

ser. Austrocaledonicae Corner ser. nov.—Folia exsiccata ple-

rumque fusco-brunnea, membranacea v. coriacea, glandulis basali-

bus 2 plerumque conspicuis, stipulis haud elongatis. Stamina

1-2-3. Arbores mediocres v. suffrutices, raro grandes (F.

smithii), ? sempervirentes. New Caledonia, Loyalty Isl., New

Hebrides, Fiji, Solomon Isl., 23 species. Typus—F. austrocale-

donica Bur.

It has been assumed on geographical grounds that this subgenus

was confined to the American tropics but, as I have shown (Rein-

wardtia 4, 1958, 26), a block of Old World species, grouped about

F. vasculosa, F. nervosa, and F. austrocaledonica, has the same

essential characters and, indeed, is with difficulty distinguished.

‘These Old World species have been referred to subgen. Urostigma

where they are out of place, particularly in being independant trees

and not banyans or stranglers. The species of New Caledonia have

never been properly classified, and they are the closest in several

respects to the American. The subgenus is absent from Africa, ex-

cepting Madagascar, where F. assimilis J. Baker occurs: it is

almost the same as the wide-spread F. albipila (Thailand to

Queensland).

I divide the subgenus into two sections, maintaining the

geographical distinction for convenience, but redefinition will

be necessary when the American species are better known. Indeed,

nearly all distinctions, from varietal upwards, are unsatisfactory.

and the whole group gives the impression of one played out into

minor variation.

Typification. For the subgenus I take, on the concurrence of G.

DeWolf, who has been studying the American species, F. maxima

P. Mill.: he writes “It is the oldest species in point of description

and probably the commonest in point of occurrence, also the most

wide-spread”. For sect. Oreosycea, Miquel gave originally the two

407

Gardens Bulletin, S-

species Urostigma nervosum (Heyne) Mig. and U. modestum,

which is a synonym of the first. Sect. Leiosycea had only F. vas-

culosa Wall.

ser. Vasculosae Corner subser. Albipilae Corner

F. albipila (Miq.) King, Ann. R. Bot. Gard. Calc. 1 (1888) 179.—

Covellia ? albipila Miq. Fl. Ind. Bat. Suppl. (1860) 175, 434.

Morus leucophylla Miq. id. 415.—F. mollis Miq. var. albipila

Miq. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 283, 296. F. colos-

sea F, Muell. ex Benth. Fl. Austral. 6 (1873) 163.—F. mallo-

toides Val. ex Backer, Blumea 6 (1948) 304 (non Mildbr. et

Hutch., 1915).—F. microtricherinos Backer, Bekn. Fl. Java 6

(1948) 40.—F. cordifolia Bl. sensu Koord. et Val. Bijdr.

Booms. Java 11 (1906) 57, 60; Ic. Bog. XI, t.16: Atlas 4

(1916) f. 701.—This is the most widely distributed species of

the subgenus. Collections are few, possibly because it is a large

tree, but they indicate a range from North Thailand through

Malaya, Sumatra, Java, Timor, and New Guinea to Queensland,

where the species was common. F. assimilis Baker is a very

close ally in Madagascar. I have examined all types; sterile spe-

cimens can easily be recognised from the amphigenous cysto-

liths.

var. glabra Corner v. nov.—Praeter setas internas glabra. Borneo

(J. Motley 613, Banjermasin, typus herb. Cambridge).

This is the only collection of the species from Borneo.

ser. Vasculosae Corner subser. Vasculosae

F. gratiosa Corner sp. nov.—Arbor —10 m. alta, foliis spiraliter

dispositis. Ramuli, petioli, et costa (subtus) pilis cinnamomeis

v. fusco-brunneis 0.5~1 mm. longis villoso-tomentosis, etiam

saepe pilis minutis undulatis: nervi subtus villosi v. sparsim

puberuli. Ramuli 2—4 mm. crassi. Stipulae 10-25 mm. longae,

brunneo-sericeae. Lamina 12—20 « 3.5—5.7 cm., lanceolato-

elliptica v. lanceolato-obovata, subacuta v. obtuse subacuminata,

basi cuneata, subcoriacea, rigida, supra nitida, sicco griseo-

viridis v. subfusca: costis lateralibus utrinsecus 10—13, obliquis,

subtus valde elevatis, intercostis O—1, nervis reticulatis supra

leviter elevatis; nervis basalibus utrinsecus 1, brevibus, glandulis

basalibus 2: petiolo 12-25 mm. longo. Receptacula axillaria,

binata v. solitaria, brunneo-villosa glabrescentia: pedunculo 2-8

mm. longo: bracteis basalibus 1.5—-3 mm. longis, sericeis: pedi-

cello 0-2 mm. longo: corpore receptaculi 13—20 mm. lato, sub-

globoso, ostiolo 3—4 mm. lato bracteis parvis numerosis occluso:

408

Vol. XVII. (1959).

setis internis nullis: cellulis scleroticis in pariete copiosis, tenui-

tunicatis. Tepala (3—) 4, lanceolata, longa glabra, in floribus

masculis cecidiophorisque plus minus gamophylla. Flores mas-

culi ostiolares, raro dispersi, sessiles: stamen 1 (—2). Semina

- 1.2-1.5 mm. longa, anguste carinata. Cystolitha hypogena.

Celebes.

NIFS bb. 19584, Malili, 1,300 m. alt. (typus, herb. Leiden).

Koorders 19349, Minahassa (Ficus sp. “R”). Sarasin 438, Mina~

hassa, Tomaton (F. mollicosta Warb. ined.). Kjellberg 2688,

Porema, 1,200 m. alt.

var. caudata Corner v. nov.—Lamina —27 x 9 cm., elliptica,

apice caudato —30 mm. longo: intercostis 3—7: petiolo -40 mm.

Receptacula pedunculo 2—3 mm., pedicello -10 mm. praedita: setis

internis sparsis: pariete 2-3 mm. crasso. Celebes (Kjellberg 2886,

Todjamboe, 800 m. alt., typus—herb. Stockholm).

Species distinctissima, F. pubinervem Bl. revocans sed lamina

nitida, stipulis brevibus, F. vasculosae Wall. affinior.

ser. Nervosae Corner

F, pachysycia Diels ex Corner sp. nov.—Arbor, foliis laxe spiralitet

dispositis. Ramuli, stipulae, petioli, et costa subtus breviter ap-

presse brunneolo-pilosi, glabrescentes. Ramuli 4-5 mm. crassi.

Stipulae —20 mm. longae. Lamina 10-19 x 6.5—12 cm., ellip-

tica, obtusa, basi cuneata, crasse coriacea, rigida, sicco brunnea:

costis lateralibus utrinsecus 7—10, intercostis 3—5 ad ambas

paginas leviter elevatis: nervis basalibus utrinsecus 1 (—2), bre-

vibus: petiolo 12-25 « 4-5 mm., crasso. Receptacula (? axil-

laria, solitaria), 40-50 x 30-35 mm., oblongo-pyriformia,

attenuata in pedicellum brevem 2—4 mm. latum bracteis laterali-

bus 1—2, c. 2 mm. longis, praeditum, bracteis haud in collare

ternatis, ostiolo plano 4-5 mm. lato bracteis apicalibus 4—5 vix

projicientibus occluso: setis internis nullis: pariete crassissimo,

sicco 5~10 mm. crasso, valde sclerotico, lumine multo compresso

et in sectione elliptico. Tepala 4, libera, rubra. Flores masculi

dispersi copiosi: pedicellis 1.5—-2.5 mm. longis: tepalis spathula-

tis: staminibus 2 (—3), haud mucronatis, pistillodio nullo. Flores

cecidiophori pedicellis 0.5—2 mm. longis: tepalis ovato-lanceo-

latis: stigmate breviter bifido. Flores feminei similes, sessiles.

Semina ? Cystolitha hypogena. New Guinea (Clemens 3676,

-Morobe District; typus-herb. Berlin, det Diels ms.).

Lamina ut in F. subnervosa Corner, receptaculum F. habro-

phyllam Benn. (ser. Austrocaledonicae) revocans.

409

Gardens Bulletin, S.

F. hombroniana Corner sp. nov.—Arbor —27 m. alta, latice serifluo

albido, foliis spiraliter dispositis. Ramuli, stipulae petioli, et

nervi subtus dense v. sparsim pilis subfulvis 0.5—1 mm. longis

appresse vestiti, v. puberuli glabrescentes. Ramuli 2-4 mm.

crassi. Stipulae 20-75 mm. longae, subulatae, curvatae, sericeae

v. glabrae. Lamina 7-19 3.5-8.5 cm, elliptica, plerumque

obovata, obtusa v. subacuta (? in juventute subacuminata), basi

cuneata, subnitida, coriacea, sicco brunnea: costis lateralibus

utrinsecus 9-13, confertis, inferioribus ad costam quasi rectan-

gulatis, subparallelis, intercostis 0—4 indistinctis, vix elevatis:

costis basalibus utrinsecus 1 (—2), glandulis basalibus 2: petiolo

5-22 2-3 mm. Receptacula axillaria, binata, glabra, maturi-

tate rufa: pedunculo 0-2 mm. longo: bracteis basalibus 2-3,

1—-1.5 1.5-2 mm., ovato-acutis, appresse puberulis: pedicello

2-12 1 mm.: corpore receptaculi 10-12 mm. lato, subgloboso

v. ovoideo, ostiolo 1-1.5 mm. lato bracteis apicalibus 2—3 oc-

cluso: setis internis nullis: cellulis scleroticis copiosis per

parietem 1.2—2 mm. crassum. Perianthium gamophyllum, bre-

viter 2—3-lobatum, rubrum, glabrum. Flores masculi ostiolares

sessiles, et dispersi pedicellati: stamine 1. Flores feminei et ceci-

diophori sessiles v. pedicellati: stigmate bifido. Semina in loculis

in pariete ligneo receptaculi plus minus inserta. Cystolitha hy-

pogena. Moluccas, New Guinea, Solomon Isl.

Amboina:—de Fretes 5762; Hombron s.n. (herb. Paris); H.B.

1885 (herb. Utrecht 47537); de Vriese s.n. (herb. Leiden). Bat-

jan:—n. 5610 (Herb. Utrecht 47536). Boeroe:—Stresemann 367.

Saparoea:—Reinwardt s.n., 1821 (herb. Leiden). Netherlands New

Guinea:—BW 6506, Asmat region, Erma, ‘terkeen’ (Asmat).

Papua:—Brass 28598 (typus, Grey herb., Harvard), 28653, Wood-

lark Isl, Kalumadau. Solomon Isl.:—J.H.L. Waterhouse B286a,

Bougainville, Siwai, ‘tuparemu’, ‘tarimu’. Java:—cult. Hort. Bog.

VIII C 9.

A Hombron (Voyage de l’Astrolabe et de la Zélée, 1838-

1840) primo collecta sed in statu sterili. Hic F. pubinervem Bl.

pilis appressis stipulis elongatis similis, illic receptaculis floribusque

F. smithii Horne revocans; lamina etiam ut in F. pachystemon.

F. pubinervis Bl. var. diandra Corner v. nov.—Flores masculi

ostiolares sessiles, et dispersi pedicellati: staminibus 2, raro 1.

Receptacula 6—8 mm. lata, sessilia. Stipulae 10-28 mm. longae.

Celebes.

_ Kjellberg 2060, Waroe-waroe; Kjellberg 2100, Malili; NIFS

Cel/II-366, Malili, Oesoe; NIFS bb. 1831, Malili, La Roua; Teys-

mann 12423, Pangkadjena (typus, herb. Bogor); Teysmann 12773

(Maros), 13964 (Bonthain, Lokka).

410

Vol. XVI. (1959).

Receptaculis parvis diandris distinguenda; ad F. subtrinerviam

v. doormanianam approximans.

F. pubinervis Bl. v. sibulanensis (Elm.) Corner comb. nov.—

F. sibulanensis Elm. Leafl. Philip. Bot. 4 (1911) 1266.—F. pu-

binervis Bl. f. sibulanensis (Elm.) Sata, Monogr. (1944) 187.

—Lamina subtus pilis erectis molliter villosa. Philippines (Elmer

11154, Mindanao, Mt. Apo, typus; Bur Sci. 17530, Samar, det.

F. samarensis Merr. ined. ).

F. subnervosa Corner sp. nov.—Arbor ~25 m. alta, cortice griseo,

latice copioso, foliis laxe spiraliter dispositis, glabra. Ramuli

1—2.5 mm. crassi. Stipulae 7-26 mm. longae, in novellis pro-

minentes. Lamina 9-22 x 3.6-10 cm. elliptica v. obovata,

obtusa, subacuta v. obtuse subacuminata, basi cuneata, tenue

coriacea, sicco brunnea: costis lateralibus utrinsecus 5—9 (—10),

subtus vix elevatis, intercostis 1—3 indistinctis: costis basalibus

utrinsecus 1 (—2), brevibus, glandulis basalibus 2: petiolo 7—22

mm. Receptacula axillaria, binata v. solitaria: pedunculo 0-6

mm. longo: bracteis basalibus 3, 1 mm. longo: pedicello 2—4

mm. longo: corpore receptaculi 10-18 mm. lato (17-30 mm.,

vivo), subgloboso, ostiolo parvo plano bracteis apicalibus paucis

occluso: setis internis nullis: pariete 1-1.5 mm. crasso, carnoso,

cellulis scleroticis nullis v. sparsis. Perianthium breviter 3—4-

lobatum carnosum, rubrum. Flores masculi dispersi pauci pe-

dicellati, ostiolares sessiles: stamine 1. Flores cecidiophori pe-

dicellati, feminei sessiles: stigmate bifido. Semina 1 mm. longa,

vix carinata. Cystolitha hypogena. New Guinea.

Netherlands New Guinea:—Aet 582, Tarera, pr. Oeta; Aet et

Idjan 721, Jappen-Biak, pr. Seroei; Beccari s.n., Andai (R. Ist.

Fir. 9244); BW 1828, Hollandia, Skyline; Kostermans 374, 450

(typus, herb. Leiden), 2940, 2961, Manokwari, Warnapi, Warsui,

Momi; v. Royen 3060, Sorong.

Papua:—Brass 27288 (Fergusson Isl.), 28092 (Sudest Isl.),

28576 (Rossel Isl.). Terr. New Guinea:—Hoogland 4811, North-

er district, Tufi subdistr., Korea village; Ledermann 6819, Sepik

(det. F. subulata Bl.): J. C. Saunders 223, 449, Madang district,

Bismark Range, Ramu valley.

Species glabra paucinervosa, receptaculi pariete carnoso, peri-

anthio gamophyllo distincta. Cave F. casearioidem King et F.

pedunculosam Miq., species dioicas.

F, kjellbergii Corner sp. nov.—Arbor mediocris, —5 m. alta, glabra,

foliis dense spiraliter dispositis: Ramuli 1-2 mm. crassi. Stipulae

5-11 mm. longae. Lamina 4—7.5 2~—3.8 cm., obovata, obtusa

All

Gardens Bulletin, S.

v. subacuta, basi attenuato-cuneata, tenue coriacea, sicco fusco-

brunnea: costis lateralibus utrinsecus 7—9, haud v. vix elevatis,

intercostis nullis: costis basalibus utrinsecus 1, brevibus, glan-

dulis basalibus 2 in axillis costarum lateralium primarum:

petiolo 6-18 mm. longo. Receptacula axillaria, binata, maturi-

tate e flavo rubescentia; pedunculo nullo: bracteis basalibus 2-3,

0.5 mm. longis, ad basim pedicelli 6-9 mm. longi: corpore re-

ceptaculi 6-8 mm. lato, subgloboso, ostiolo plano 1 mm. lato

bracteis apicalibus parvis occluso: setis internis nullis: cellulis

‘ scleroticis in pariete copiosis. Perianthium breviter 3 (—4)-loba-

tum, rubrum. Flores masculi ostiolares sessiles (? paucissimi

dispersi): stamine 1. Flores cecidiophori pedicellati, feminei

sessiles: stigmate bifido. Cystolitha hypogena. Celebes (Kjell-

berg 2437, Tolala, typus herb. Stockholm).

Ut F. hombroniana Corner, sed in omnibus partibus minor et

paucinervosa, stipula haud prominenti.

F. subtrinervia Laut. et K. Schum. var. doormaniana (Diels) Corner

Nabire; NIFS bb. 30583, Jappen Isl., Seroei; Ledermann 8249,

comb. nov.—F. doormaniana Diels, Engl. Bot. Jahrb. 67

(1935) 192.—F. behrmanniana Diels, id.mLamina —24 8.5

cm., ovato-lanceolata v. elliptico-lanceolata, attenuata v. sub-

acuminata: costis lateralibus utrinsecus 8-16. Bracteae basales

acutae. Plus minus glabra. New Guinea.

H. J. Lam 1380, Mamberamo R., Doorman R. (typus); Docters

Leeuwen 10259, Rouffaer R.; Kanehira et Hatusima 12813, pr.

Sepik (typus, F. behrmanniana Diels); Brass 13222, 13433,

13679, Idenburg R.; Brass 28076, Papua, Sudest Isl.; Hinds s.n.,

(det F. pubinervis).

Ad F. pubinervem v. diandram Corner approximans, sed costis

lJateralibus pluribus.

F. pachystemon Warb. in Laut. et K. Schum. Nachtr. Fl. Schutzgeb.

(1905) 242.—F. mangiferifolia Laut. et K. Schum. Fl. Schutz-

geb. (1901) 275, haud Griffith 1854.—The type of F. pachys-

temon (Dahl s.n., Ralum, Gazelle Peninsula, Neu Pommern) at

Berlin has shorter leaves and smaller figs (10-11 x 8-9 mm.)

than in the later homonym F. mangiferifolia but I can see no

other difference. The species seems not uncommon as a river-

side shrub or tree in New Guinea and New Britain, and such

variability is likely.

F. madhucifolia Corner sp. nov.—Arbor, foltis fasciculato-

confertis, glabra. Ramuli 3 mm. crassi. Stipulae —30 mm. longae.

Lamina 8-20 x 3-9 cm., anguste obovata, subacuta, basi

412

_ oeror

Vol. XVIT. (1959).

cuneata, tenue subcoriacea, sicco brunnea: costis lateralibus ut-

rinsecus 7—9, subtus leviter elevatis, 0-2 (—3) intercostis: costis

basalibus utrinsecus 1, brevibus, glandulis basalibus 2: petiolo

7—20 mm. longo. Receptacula axillaria, binata, sessilia, 10—11

mm. lata, subglobosa, ostiolo plano bracteis apicalibus appla-

natis 3 occluso: bracteis basalibus 3, 2-3.5 « 2-—2.5 mm.,

ovatis, acutis: setis internis nullis: cellulis scleroticis in pariete

copiosis. Tepala 3—4, libera, rubra, oblonga. Flores masculi dis-

persi, copiosi, plus minus sessiles: stamine 1. Flores cecidio-

phori sessiles v. breviter pedicellati, feminei sessiles: stigmate

bifido. Cystolitha hypogena. Celebes (Buwalda 3719, Bantimoe-

rong, typus herb. Leiden; Teysmann 12621, 12637, Tjambu).

F. gratiosam Corner lamina floribusque sessilibus revocans, sed

glabra, receptacula sessilia, flores masculi dispersi copiosi.

F. hadroneura Diels, Engl. Bot. Jahrb. 67 (1935) 186.—This ex-

tremely interesting, large-leafed species, allied perhaps with F.

pseudojaca Corner, is represented by three sterile collections and

the type (Lane Poole and Brass 290), of which there is a leaf

only in the Berlin-herbarium, but a leafy twig with figs in the

Brisbane-herbarium. This Brisbane specimen is labelled by Diels

and must be taken as the type. Identical with it are: —

BW 390, 365, Manokwari, Prafi, ‘ateska’; J. C. Saunders 532,

Terr. New Guinea, Madang subdistr., Ramu valley, 5 miles S.E.

of the Faita airstrip, rainforest, c. 800 ft. alt., ‘uli’ (Bilia),

‘bambam’ (Amele), ‘sungung’ (Dumpu).

F. pseudojaca Corner sp. nov.—Arbor —35 m. alta, cortice brun-

neolo, latice cremeo, foliis distichis. Ramuli, petioli, et stipulae

pilis flexuosis cervicoloribus 0.5—1 mm. longis villosi, costae et

costulae subtus appresso-puberulae. Ramuli 2.5—4 mm. crassi.

Stipulae 12-18 mm. longae. Lamina 10-21.5 « 4.5-14 cm.,

elliptica v. ovato-elliptica, subacuta v. breviter acuminata, basi

rotundata v. late cuneata et minute auriculata, subasymmetrica,

chartacea v. tenue coriacea, sicco fusco-brunnea v. griseo-

viridis: costis lateralibus utrinsecus 10-15, supra impressis,

subtus prominentibus, intercostis 3—9, nervulis reticulatis supra

tenue, subtus valdius, elevatis: costis basalibus utrinsecus 3,

brevibus, glandulis basalibus 2: petiolo 6-20 2.5-3.5 mm.

Receptacula ? Cystolitha hypogena. New Guinea.

Netherlands New Guinea:—BW 1399, Salawati Isl. Terr. New

Guinea:—-NGF 1944, Lae; J. C. Saunders 289, 294, 301, 430

(typus, herb. Lae), 595, Madang district, Ramu Valley. Papua:—

L. E. Cheesman 40, Kokoda.

Specimina sterilia, nihilominus species facile distinguenda, Arto-

carpum simulans, forsan F. hadroneurae Diels affinis.

413

Gardens Bulletin, S.

ser. Austrocaledonicae Corner

F. habrophylla G. Bennett ex Seem. FI. Vit. (1868) 248.—

F. bennettii Seem. id., 250.—F. tanensis G. Bennett ex Seem.

id., 248.—F. edulis Bur. Ann. Sci. Nat. ser. 5, Bot. 14 (1872)

271.—The types of F. habrophylla and F. tanensis at Kew, both

from Tanna, show that they are identical with F. edulis, as

Bureau suspected. The type of F. bennettii, at the British Mu-

seum, is from a plant cultivated in the Sydney Botanic Garden.

This is an extremely variable species requiring field-examination.

¥. granatum Forst. f. var. minor Corner v. nov.—Arbor —25 m.

alta. Stipulae, pedunculi, bracteaeque basales tenue appresse

brunneo-puberuli. Ramuli 2-4 mm. crassi. Lamina 11-23

6—10 cm., elliptica v. elliptico-cbovata, obtusa v. subacuta, basi

cuneata: costis lateralibus utrinsecus 9-11, intercostis 1-3:

petiolo 20-80 x 1.5—2.5 mm. Receptacula maturitate rubra:

pedunculo O-5 mm. longo: bracteis basalibus 3, 1—-1.5 mm.

longis, caducis: pedicello 2-6 mm. longo: corpore receptaculi

10-11 & 8 mm., ellipsoideo, osticlo plano 1.5 mm. lato bracteis

apicalibus disciformibus occluso: setis internis copiosis: cellulis

scleroticis in pariete copiosis. New Hebrides (Aneityum, Ero-

mango: Kajewski 324, 732-typus herb. Kew, 956).

Ut v. granatum sed gracilior et receptacula multo minora, v. gra-

natum ipsa quamvis raro collecta nec satis cognita.

F. smithii Horne ex Baker var. robusta Corner vy. nov.—Arbor

—30 m. alta, stipulis ramulisque appresse brunneolo-pilosis gla-

brescentibus. Lamina 7-22 x 3.6—9.5 cm., elliptica v. elliptico-

obovata, obtusa v. subacuta: costis lateralibus utrinsecus 8-13:

petiolo 15-35 x 1.5—-3 mm. Receptacula 10-13 mm. lata,

pedunculo 7-15 x 1-1.5 mm, pedicello 0-5 mm.: cellulis

scleroticis sparsis in strato parietis interno. Flores masculi et

cecidiophori pedicellis induratis scleroticis. New Hebrides

(Aneityum), Fijt (Viti Levu, Vanua Levu, Taveuni), in silvis ad

700 m. alt.

New Hebrides:—Kajewski 862, 955, Aneityum, Anelgauhat

Bay. Vanua Levu:—A. C. Smith 605, 1728. Viti Levu:—A. C.

Smith 5357, 5879, 5952, 5995, 8497, 8709, 9162, 9273. Taveuni:

—J. W. Gillespie 4753, A. C. Smith 8339 (typus, herb. Cam-

bridge).

Var. smithii. gracilis, -15 m. alta: lamina -13 X 4.2 cm., an-

guste elliptica v. lanceolata, obtuse acuminata, costis lateralibus

414.

Vol. XVII. (1959).

utrinsecus 7-9, petiolo 4-14 mm.: receptacula 5—6 mm. lata,

pedunculo 6-12 x 0.5 mm., gracillimi, cellulis scleroticis in

pariete copiosis: Fiji, Solomon Isl. (Guadalcanal).

F. austrocaledonica Bur. var. balansaeana (Bur.) Corner comb.

nov.—F’. balansaeana Bur. Ann. Sci. Nat. ser. 5, Bot. 14 (1872).

262.—So far as I can see this differs from v. austrocaledonica

merely in the closely and appressedly hairy twigs, petioles, pedi-

cels, and fig-body. It is known only from the type, Balansa 138

(New Caledonia, Prony Bay).

415

Taxonomic Notes on Ficus Linn., Asia

and Australasia

Ii. SUBGEN. FICUS AND SECT. FICUS

By E. J. H. CORNER

Botany School, University of Cambridge

Summary

NEW SUBSECTIONS, series and subseries of sect. Ficus:—subsect.

Ficus ser. Rivulares, Pseudopalmeae, Sinosyceae, Cariceae, Eryth-

rogyneae, Podosyceae, subser. Basitepalae; subsect. Eriosyceae

(Miq.) Corner, ser. Eriosyceae, subser. Eriosyceae, Trichosyceae,

Dehiscentes, Cuneifoliae; ser. Auratae, subser. Auratae, Monan-

dreae.

Earlier specific epithets: —F. subincisa B. Ham. ex J.E. Sm. (F.

cunia B. Ham. ex Roxb.), F. neriifolia J.E. Sm. (F. nemoralis

Wall. ex Miq.), F. gasparriniana Miq. (F. silhetensis Miq.).

New species: —F. trivia, F. halmaherae, F. subfulva et var. villo-

sula, F. litseifolia, F. endospermifolia, F. bruneiensis, F. brunneo-

aurata, F. androchaete, F. eumorpha et v. subglabra, F. para-

morpha.

New varieties and forms:—F. deltoidea Jack v. angustifolia

(Mig.) Corner et f. angustissima, v. arenaria, v. bilobata, v. bor-

neensis et f. subhirsuta, v. intermedia, v. kunstleri (King) Corner,

v. lutescens (Desf.) Corner et f. longipedunculata et f. subsessilis

(Miq.) Corner, v. motleyana (Miq.) Corner, v. oligoneura (Miq.)

Corner, v. peltata, v. trengganuensis.—F.. oleaefolia King v. dodo-

naeiformis (Gagnep.) Corner, v. memecylifolia, v. myrsinoides, Vv.

riparia. F. neriifolia J.E. Sm. v. fieldingii (Miq.) Corner, v. nemo-

ralis (Wall. ex Mig.) Corner, v. trilepis (King) Corner. F. pedun-

culosa Miq. v. confertifolia (Merr.) Corner, v. imberbis (Elmer)

Corner, v. macropoda (Miq.) Corner, v. mearnsii (Merr.) Corner,

v. segaarensis (Engl.) Corner, v. velutina—F. erecta Thunb. v.

erecta f. sieboldii (Miq.) Corner, var. beecheyana (Hook. et Arm)

King f. koshunensis (Hayata) Cormer.—F. gasparriniana Mig. Vv.

esquirolii (Lévl. et Vant.) Corner, v. laceratifolia (Lévl. et Vant.)

Corner, v. viridescens (Lévl. et Vant.) Corner.—F. stenophylla

416

Vol. XVII. (1959).

Hemsl. v. macropodocarpa (Lévl. et Vant.) Corner, v. nhatrangen-

sis (Gagnep.) Corner.—F. ischnopoda Migq. v. subcylindrica.—-F.

pustulata Elmer v. lanceifolia (Sata) Corner.—F. grossularioides

Burm. f. v. robusta, v. stenoloba.—F. hirta Vahl v. appressa, Vv.

brevipila, v. dumosa (King) Corner, v. malayana, v. squamosa.—

F. simplicissima Lour. v. annamica (Gagnep.) Corner.—F. fulva

Reinw. v. timorensis—F. glandulifera Wall. v. camiguinensis

(Merr.) Corner, v. villosa.—F. chartacea Wall. v. lanceolata—F.

tuphapensis Drake v. annamensis (Gagnep.) Corner.—F. aurata

Mia. v. brevipilosa, v. longipilosa, v. palawanensis, v. pedunculata.

—F. macilenta King v. gibbsiae (Ridley) Corner, v. ilicifolia.—

F. setiflora Stapf v. adelpha, v. puberula.

subgen. Ficus

subgen. Eusyce Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867)

289.—subgen. Carica Mildbr. et Bur., Engl. Bot. Jahrb. 46

(1912) 174.—subgen. Metamorphe Sata J. Sci. Tr. Agr. Taiwan 6

(1934) 19.—subgen. Eumetamorphe Sata, Monogr. (1944) 244.

—subgen. Caricae Sata, Monogr. (1944) pt. 4, 179.—Dioecious.

Interfloral bracts absent. Style of gall-flowers much shorter than

that of the female; female stigma simple or bifid, gall-stigma simple

and, often, more or less infundibuliform. East Africa, Asia, Aus-

tralasia, 312 spp.

I distinguish eight sections: —

sect. Ficus, sect. Rhizocladus Endl., sect. Kalosyce Miq. (sect.

Synoecia Miq.), sect. Sinosycidium sect. nov., sect. Neomorphe

King, sect. Sycocarpus Mig. (sect. Covellia auct.), sect. Sycidium

Miq., sect. Adenosperma Corner.

sect. Ficus

This is the group with bistaminate male flowers which has F.

carica Linn. as its type. I divide it into two subsections, each with

several series. These subsections are so different in detail that they

appear worthy of sectional rank. There are, however, a few species

in South China and Indochina which come between them and in

some way bridge the differences. Hence I adopt the more conser-

vative classification. ;

subsect. Ficus——Semina laevia, non v. vix carinata. Tepala

plerumque glabra. Flores masculi ostiolares v. dispersi, antheris

non mucronatis. Lamina cystolithis praedita. Glandulae mic-

roscopicae capitatae, cellulis 2—4, v. flabelliformes. East Africa

usque ad Japan et New Guinea, 31 spp. Typus, F. carica Linn.

417

Gardens Bulletin, S.

ser. Rivulares Corner ser. nov.—Semina compressa, auriculi-

formia. Perianthium cupulare 2—4-lobatum, ovario albido plus

minus obtegens. Flores masculi ostiolares, staminibus (1—) 2.

Stigma simplex. Receptacula axillaria, solitaria. Cystolitha hypo-

gena. Ins. Philippines, | sp. F. rivularis Merr.

ser. Pseudopalmeae Corner ser. nov.—Ficus sect. Pseudopalma

Elmer, Leafl. Philip. Bot. 1 (1908) 283.—Semina lentiformia.

Flores sessiles, masculi ostiolares. Tepala libera. Ovarium cecidio-

phorum rufo-brunneum longistipitatum. Arbuscula pachycaulis

sparsim ramosa, foliis magnis brevipetiolatis grosse dentatis. Cys-

tolitha amphigena. Ins. Philippines, 1 sp F . pseudopalma Blanco.

ser. Sinosyceae Corner ser. nov.—Semina lentiformia. Tepala

libera, rubra v. albida. Flores masculi ostiolares v. dispersi, stami-

nibus 1—5. Receptacula plerumque solitaria, axillaria. Stipulae

binatae. Cystolitha amphigena. India, Burma, China, Indochina,

Thailand, 2 spp. F. henryi Diels, typus, et F. subincisa B. Ham. ex

J. E. Sm. (F. clavata Wall. ex Miq.).

ser. Cariceae Corner ser. nov.—Semina lentiformia. Tepala

libera, albida v. carnea, membranacea. Flores masculi ordinibus

numerosis ostiolares, raro etiam dispersi: staminibus 2—4. Recep-

tacula plerumque solitaria, axillaria. Folia palmato-lobata v. ovato-

cordata, plerumque dentata. Cystolitha hypogena. East Africa,

Asia Minor, Pakistan, India, Bonin Isl., 3 spp. F. carica Linn.

(typus), F. palmata Forsk., F. tidaiana Wilson.

ser. Erythrogyneae Corner ser. nov.—Erythrogyne Vis. ex Gasp.

Parl. Giorn. Bot. 2 (1844) 219; Ann. Sci. Nat. ser. 3, 3 (1845)

348.—Ficus sect. Erythrogynae (Vis.) Endl. Gen. Pl. Suppl. 4, 2

(1847) 34.—subgen. Erythrogyne (Vis.) Mig. Ann. Mus. Bot.

Lugd. Bat. 3 (1867) 289.—Semina magna reniformia, 2.5—6 mm.

longa, vix carinata, per receptaculum pauca: cotyledones condup-

licatae. Tepala libera rubra, ovario femineo semper multo breviora.

Flores masculi ostiolares. Ovarium cecidiophorum saepe crusta-

ceum et rugoso-angulatum. Frutices v. arbusculae, saepe epiphy-

ticae, haud sarmentosae. Folia acutilanceolata et penninervia, vel

obtusa costa dichotoma, integra. Cystolitha hypogena. Western

Malaysia, 2 spp. F. deltoidea Jack (typus), F. oleaefolia King.

ser. Podosyceae Corner ser. nov.—Ficus sect. Podosycea Miq.

Hook. Lond. J. Bot. 7 (1848) 442.—-sect. Didymophora Mig. et

sect. Thamnosycea Migq., id. 453, 454.—sect. Eusyce (Miq.) King

subsect. Frutescentiae Sata, Monogr. (1944), 332, 385 (lecto-

typus, F. pedunculosa Miq.).—Semina 1—3 mm. longa, lentifor-

mia Vv. reniformia, per receptaculum numerosa. Tepala libera, raro

418

Vol XVI: (ga).

conjuncta, rubra, ovario femineo equalia v. breviora. Flores mas-

culi dispersi, raro solum ostiolares. Frutices v. arbores, raro epi-

phyticae sed nonnullae repentes. Asia, Malaysia, 22 spp. Typus,

F, pedunculosa Miaq.

subser. Podosyceae.—Flores cecidiophori et sessiles et pedicellati,

tepalis ovario sessili v. substipitato plus minus equalibus. Flores

feminei tepalis ovario longioribus v. brevioribus. 10 spp. Typus,

F. pedunculosa Miq.

subser. Basitepalae Corner——Flores cecidiophori plus minus

sessiles, tepalis ovario longistipitato multo brevioribus, ut in flori-

bus femineis. Sinohimalaya, Philippines, 4 spp. Typus, F. abelii

Mig.

subsect. Eriosycea (Mig.) Corner subsect. nov.—Ficus sect.

Eriosycea Mig. Hook. Lond. J. Bot. 7 (1848) 455; emend.

Mig. FI. Ind. Bat. I, 2 (1859) 293; Ann. Mus. Bot. Lugd. Bat.

3 (1867) 290.—sect. Trichosycea Miq. Fl. Ind. Bat. I, 2

(1859) 296; Ann. Mus. Bot. Lugd. Bat. 3 (1867) 290.—Semina

plerumque tuberculata v. echinata, carinata et basi duplice cari-

nata, haud compressa. Tepala rubra v. albido-ochracea et setosa.

Flores masculi ostiolares: staminibus 2, 1—2, vel 1, antheris bre-

viter mucronatis v. modo acutis. Cystolitha nulla. Glandulae

microscopicae capitatae, cellulis plerumque numerosis. Frutices

v. arbores, haud epiphyticae, foliis saepe palmato-lobatis prae-

cipue juvenilibus. Asia, New Guinea, 28 spp. Typus, F. grossu-

larioides Burm. f.

ser. Eriosyceae Corner ser. nov.—Tepala rubra 3—7, plerumque

4—5, glabra v. apice marginibusque ciliolatis. Stamina et ovaria

saepe basi setulis circumdata.. 19 spp. Typus, F. grossularioides

Burm. f.

subser. Eriosyceae.—Folia subtus areolis obscuratis pilis un-

dulatis albidis griseis v. fulvis lanata, juvenilia plerumque palmato-

lobata. China, Indochina, Thailand, West Malaysia, 3 spp. Typus,

F. grossularioides Burm. f.

subser. Trichosycex Corner subser. nov.—sect. Trichosycea

Mig. l.c.—Folia subtus haud lanata, areolis haud obscuratis,

dentata v. denticulata, juvenilia saepe palmato-lobata. India, China,

usque ad ins. Moluccanas, 7 spp. Typus, F. hirta Vahl.

subser. Dehiscentes Corner subser. nov.—Folia subtus haud

lanata, integra, intercostis numerosis strictis, juvenilia raro pal-

mato-lobata. Receptacula cecidiophora irregulariter dehiscentia,

maturitate viridia v. flavescentia, feminea rubra indehiscentia. India

usque ad New Guinea, 3 spp. Typus, F. glandulifera Wall. ex Miq.

419

Gardens Bulletin, S-

subser. Cuneifoliae Corner subser. nov.—Folia subtus haud

lanata, parviuscula integra, mtercostis paucis et saepe laxis, basi

cuneata, brevipetiolata, juvenilia haud v. raro palmato-lobata.

Receptacula cecidiophora ut feminea rubescentia indehiscentia.

Stamina 1—2, vel 1, antheris haud mucronatis. India, China, Indo-

china, Thailand, Malaya, Sumatra, Borneo, 3 spp. Typus, F. char-

tacea Wall. ex King.

ser. Auratae Corner ser. nov.—Flores feminei et cecidiophori

tepalis 3-4 (—5) albidis v. ochraceis dense setosis. Setulae c. sta-

mina et ovaria plerumque deficientes. Setae internae copiosissimae

rigidae. Indochina, Thailand, Malaya, Sumatra, Borneo, Palawan,

9 spp. Typus, F. aurata Miq.

subser. Auratae.—Flores masculi ordinibus numerosis ostiolares,

sessiles et pedicellati, tepalis haud setosis, staminibus 2. Recep-

tacula 10-26 mm. lata, plerumque sessilia. Lamina majuscula. 4

spp. Typus, F. aurata Mia.

subser. Monandreae Corner subser. nov.—Flores masculi ordine

uno ostiolares sessiles, v. etiam ordine secundo pedicellati, tepalis

albido-setosis ut in floribus cecidiophoris, stamine uno. Recepta-

cula 10-16 mm. lata, sessilia v. pedunculata. Lamina saepe minus-

cula. Borneo, 5 spp. Typus, F. setiflora Stapf.

subsect. Ficus ser. Sinosyceae Corner

F. subincisa B. Ham. ex J.E. Sm. Rees Cyclop. 14 (1810) n. 91.—

F. chincha Roxb. FI. Ind. 3 (1832) 534.—F. clavata Wall. ex

Miq. Hook. Lond. J. Bot. 7 (1848) 431.—F. caudata Griff. Ic.

Pl. As. (1854) t. 557 (i).—The type of F. subincisa is in the

herbarium of the Linnean Society of London (n. 1610.42),

and there is a specimen of F. chincha named by Roxburgh in

the Martius herbarium at Brussels. They leave no doubt that

they are earlier names for the same species as F. clavata Wall.

subsect. Ficus ser. Erythrogyneae Corner

F. deltoidea Jack, Mal. Misc. 2 (1822) 71; Merrill, J. Arn. Arb.

33 (1952) 225.—This common shrub of Western Malaysia is

now represented in herbaria by more than 600 collections, and

I have myself studied many more living plants. Superficially

there is bewildering diversity, but attention to detail has lead me

to distinguish twelve varieties. Most of these appear to have geo-

graphical delimitation which it should not be difficult to check

because the plants are easy to recognise and to gather. It is,

420.

Vol. XVII. (1959).

however, necessary to collect gall-plants as well as the seed-

plants, because diagnostic characters lie in the gall-ovary and

tepals: in a few varieties it is impossible to classify seed-plants.

F. diversifolia Bl. is the common Javanese mountain shrub |

known, through cultivation in Europe, as var. lutescens Desf.,

specimens of which are preserved in the herbaria at Paris, Kew,

and Florence; Blume’s type is at Leiden. Of Jack’s two species,

F. ovoidea and F. deltoidea, the first is authenticated by Wallich

4526 from Penang, labelled F. ovoidea Jack: King made this

var. ovoidea which is antedated by var. angustifolia Miq. There

remains the identity of F. deltoidea. Jack did not mention penni-

nerved leaves, which he would surely have done if he had been

describing var. lutescens. Hence | identify his plant with that of

Sumatran and Malayan lowlands which lack the penninerved

leaves in the adult state: his description agrees, but lacks other

details diagnostic of particular varieties.

var. angustifolia (Mig.) Corner comb. nov.—Synoecia diversifolia

(Bl.) Mig. v. angustifolia Mig. Pl. Jungh. (1851) 67.—F. ovoi-

dea Jack. Mal. Misc. 2 (1822) 71.—F. diversifolia v. ovoidea

(Jack) King, Ann. R. Bot. Gard. Calc. 1, 2 (1888) 140, pl.

174D.—Miquel’s type, at Leiden, shows that it is the narrow

spathulate-leafed variety of F. deltoidea.

f. angustissima Corner f. nov.—Ramuli 1—1.5 mm. crassi.

Lamina 2—3.3 & 0.5—1 cm., anguste spathulata, costa sub mediam

laminae dichotoma. Receptacula 5 x 3-4 mm., pedunculo 2-5:

mm. longo. Semina 1—2 per receptaculum. Mentawei Isl.

H.H. B.B. 639 (Batoe), Iboet 259 (Siberut), Sing. F.n. 14753

(Sipora, typus herb. Singapore).

var. arenaria Corner v. nov.—Frutex —2 m. altus. Lamina 2.5-11

x 1.8-4 cm., plerumque elliptica v. anguste obovata, acuta v.

subacuminata, penninervia, costis lateralibus utrinsecus 3-5:

folia superiora pauca obovata, obtusa, costa ad v. super mediam

laminae dichotoma. Receptacula 10-12 < 7-9 mm., ellip-

soidea, v. 7-9 mm. lata subglobosa: pedunculo 1-9 mm. longo.

Flores cecidiophori tepalis ovario angulato tenuiter crustaceo

multo brevioribus. Borneo, ad terram arenosam.

Alston 13092 (Sampit); Brunei 5352 (Kuala Belait, typus herb.

Cambridge), 5367 (Badas); Bur. Sci. 290 (Sandakan); Dunsel-

man 103 (Singkawang); Hose 211 (Sarawak, Marudi, Baram) ;

Purseglove 5024 (Bako National Park); Richards 2644, 2645

(Marudi, Sarawak).

421

Gardens Bulletin, S.

This is like v. intermedia and v. motleyana, but differs from both

in the short gall-tepals. Thus it suggests v. deltoidea but its leaves

are mostly acute as in v. motleyana. I have seen many plants in the

Kerangas-vegetation in Sarawak and Brunei and consider that it is

usefully distinguished.

var. bilobata Corner v. nov.—Frutex v. arbuscula —5 m. altus.

Lamina 2-7 X 1-3.5 cm., spathulato-obovata v. obdeltoidea,

apice saepe breviter bilobata, costa ad v. infra mediam dicho-

toma: glandula ad furcam costae, basalibus 2 v. nullis, aliqu-

ando glandulis parvis ad furcas nervorum: petiolo 3—25 mm.

Receptacula 5-8 mm. lata, pedunculo 2-10 mm. longo. Tepala

cecidiophora ovario rugoso-angulato equalia. Malaya (Selangor,

Pahang, Perak, Kedah) in silvis montanis 1000-2000 m. alt. et

ad cacumina collium calcareorum.

Sing. F.n. 2368 (Taiping Hill, typus, herb. Simgapore), 8443.

11077, 19701, 29689, 32551, 38028, 38708; Ridley 7628; Haniff

3908; King’s Coll. 2430; Scortechini 308; Wray 1099; Yapp 488.

This approaches v. intermedia, v. kunstleri, v. lutescens, and v.

trengganuensis, but cannot be placed satisfactorily with any. It

requires field-study. Possibly all these varieties converge to this

effect in their high mountain forms.

var. borneensis Corner v. nov—Ramuli 2—5 mm. crassi. Lamina

4.5-13 < 3-9 cm., obovata, obtusa v. subrotundata, costa plus

minus ad mediam dichotoma, foliis ad basim caulis solo acutis

penninerviis: glandulis 2 basalibus, raro ad furcam costae: petiolo

4-25 mm. longo. 3-4 mm. crasso. Receptacula 8-12 mm. lata

(11-16 mm., viva), pedunculo 0.5—6 mm. longo: bracteis basali-

bus 2—2.5 x 3-4 mm. Flores cecidiophori sessiles v. breviter

pedicellati: tepalis ovario valde rugoso-angulato sessili v. longis-

tipitato multo brevioribus. Borneo.

Amdjak 492; Anderson 9754; Brunei 649; Dunselman 90; En-

dert 3920 (West Koetei, Mt. Kemoel, typus herb. Leiden) ; Hallier

1372, 2240; Sahib 16321; Sales 774; Sing. F.n. 27370; S. 4807,

5856, 9953.

This appears to be widely distributed in all parts of Borneo from

the lowlands to an altitude of 1,500 m. In the large leaf and fig it

approaches v. kunstleri (not known from Borneo), but differs in

the short peduncle, short gall-tepals, and the fewer leaf-glands.

Thus it is, in fact, an enlarged state of v. deltoidea.

f. subhirsuta Corner f. nov.—Receptacula tenuiter appresse

pilosa, aliquando etiam ramuli. Sarawak, S. Borneo. —

422.

Vol. XVII. (1959).

Beccari 274, 815, 1246; Clemens 21907 (Gat, Upper Rejang.

River, typus herb. Br. Mus.); Sarawak Mus. 4807; Teysmann

7118.

Varietates F. deltoideae praecipue glabrae.

var. intermedia Corner v. nov.—F. kinabaluensis Stapf, Tr. Linn.

Soc. Bot. ser. 2, 4 (1894) 226.—F. burkillii Ridl. Fl. Mal. Pen.

3 (1924) 330.—Frutex —4 m. alta. Folia plerumque dimorpha,

penninervia ut in v. motleyana, vel lamina obtusa spathulata 3—

10 x 1.5—5.5 cm., semper longiori quam lata, costa super

mediam, saepe prope apicem, dichotoma: glandulis 2 basalibus:

petiolo 2-15 mm. Receptacuia rubra dein nigro-purpurea, 6—10

mm. lata, subglobosa: pedunculo 1-11 (—15) mm.: bracteis

basalibus 1—2 mm. longis. Flores cecidiophori tepalis ligulatis

v. lanceolatis ovario laevi v. leviter subangulato longioribus.

Malaya, Borneo, (? Celebes), in silvis montanis 800—2,600 m.

alt. saepe epiphytica. Typus, Purseglove 4281 (Pahang, Fraser’s

Hill, herb. Singapore).

I distinguished this variety because it comes between v. motleyana

and v. lutescens and predominates in Malayan mountains where

neither of these varieties occurs. The differences from v. motleyana

are the more coriaceous leaf, the subglobose fig, the smooth gall-

ovary, and the greater production of obtuse leaves with dichotom-

ous midrib. On Mt. Kinabalu it seems that these obtuse leaves are

few, and specimens exactly like those of F. kinabaluensis have

been collected from G. Tahan in Malaya. Generally, in Malaya

abundant obovate obtuse leaves are formed and this is F. burkilii.

All the numerous collections: from Malaya identified as F. del-

toidea (or F. diversifolia) v. lutescens belong here and, also, the

following from Borneo:—

Clemens 10700, 10996, 20161, 28787, 28932, 29257, 29916,

30292, 30294, 30296, 30766, 30988, 31503, 31665, 32936,

32937, 32940, 33193, 33625, 33830, 40696, 50635, 51082.

Endert 3912, 3917. Haviland 1215, 1216. Purseglove 5024. San.

17580. Sing. F.n. 12209, 26400, 27524, 27695, 27714, 28007.

SK 29; 136.

var. kunstleri (King) Corner comb. nov.—F. diversifolia Bl. v.

kunstleri King, Ann. R. Bot. Gard. Calc. 1, 2 (1888) 140, pl.

175 A, B.

var. lutescens (Desf.) Corner comb. nov.—F. diversifolia Bl. Bijdr.

(1825) 456.—F. lutescens Desf. Cat. ed. 3 (1829) 413.—F.

diversifolia Bl. v. lutescens (Desf.) King, Ann R. Bot. Calc. 1,

423

Gardens Bulletin, S.

2 (1888) 140.—This is the Javanese and Sumatran plant with

the obtuse leaves broad and with the midrib dichotomous at or

below the middle.

f. longipedunculata Corner f. nov.—(F. diversifolia Bl. v. longi-

pedunculata Miq. ined.).—Pedunculi 14-30 mm. longi. Java,

Sumatra.

Sumatra:—Beccari 190 (Mt. Singalan); Bunnemeyer 5629 (G.

Gombah), 9187 (G. Koerintji, typus herb. Leiden); Meijer 5008

(Mt. Sago); v. Steenis 3669 (G. Pesagi).

Java:—Backer 8677, 25722, 25941; Blume s.n. (G. Salak);

Coert 689; Harreveld 1961; Herb. Junghuhn s.n. (G. Gedeh), n.

177, et s.n. (Hoch Ankola); Koorders 9404, 9406, 26491,

26812; Schiffner s.n. (1872); de Vriese s.n. (det. v. longipedun-

culata Miq. ms., ? Java, ? Sumatra).

f. subsessilis (Mig.) Corner comb. nov.—F. diversifolia Bl. v.

subsessilis Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 268, 289.—

Pedunculi 0-4 mm. longi. Java, Sumatra.

This is represented by many collections. The type (de Vriese,

Batjan, herb. Leiden, herb. Utrecht) was said to come from the

Moluccas, but these lie outside the range of the species and, I

think, this was an error for Java.

var. motleyana (Mig.) Corner comb. nov.—F. motleyana Miq.

Ann. Mus. Bot. Lugd. Bat. 3 (1867) 228, 294.—The typical

form of F. motleyana with oblong, shortly pedunculate fig and

only lanceolate, acute, penninerved leaves seems specifically dis-

tinct, but F. deltoidea v. intermedia provides so many inter-

mediates that it is almost impossible to draw a line between

them in the herbarium.

var. obligoneura (Mig.) Corner comb. nov.—F. oligoneura Mig.

Ann. Mus. Bot. Lugd. Bat. 3 (1867) 289.—Urostigma oligo-

neuron Mig. Fl. Ind. Bat. Suppl. (1860) 438.—This is close to

v. lutescens but has larger figs and long-pedicellate gall-flowers.

Its leaf is extremely variable.

y. peltata Corner v. nov.—Frutex repens gracilis, internodis elon-

gatis. Lamina 1.5—2.7 cm. lata, suborbicularis, anguste peltata,

costa infra mediam dichotoma: glandulis 3 (2 basalibus, 1 ad

furcam costae): petiolo 10-18 x 1 mm. Receptacula 8 X 5

mm., ellipsoidea: pedunculo 1-1.5 mm. longo: pedicello 2-3

mm. longo: Flores cecidiophori sessiles v. pedicellati: tepalis

4-5 ligulatis, ovario laevi equalibus v. longioribus. Sumatra (Mt.

Sago, Harau Canyon, ad scopulos).

424

ant 6.

~ Vol. XVIT. (1959). :

Meijer 3980 (typus, herb. Cambridge); Jacobsen 2194.

_ Habitu repenti foliis suborbicularibus anguste peltatis distinc-

tissima sed v. lutescentis affinis.

var. trengganuensis Corner v. nov.—Ramuli 2 mm. crassi. Lamina

2.3-8 & 1.8—-5.5 cm., elliptica v. rotundato-obovata, etiam sub-

bilobata, costa ad v. super mediam dichotoma: glandulis 3—4

(—5), 2 basalibus, ceteris ad furcas nervorum: petiolo 10-50 mm.

longo. Receptacula 9-12 mm. lata (14-18 « 11—14 mm., viva),

roseo-rubra dein nigro-purpurea: pedunculo 6-20 1.5 mm.:

bracteis basalibus 1—-1.5 mm. longis. Flores cecidiophori sessiles

v. pedicellis —1.5 mm. longis praediti: tepalis 3-4, lanceolatis v.

ligulatis ovario rugoso-angulato equalibus, saepe pilo apicali

praeditis. Malaya (Trengganu, Pahang costa orientali) in locis

arenosis et montanis, terrestris v. epiphytica.

Cons. For. 3112: Kep. F.n. 37354; Ridley 1401, 1444; Sing.

F.n. 15346, 17458, 17515, 25841, 30079 (typus, herb. Singa-

pore), 31039, 31075, 39914, 40405; Vesterdal 284.

Inter v. kunstleri et v. intermediam sed lamina elliptica obtusa.

F. oleaefolia King var. dodonaeiformis (Gagnep.) Corner comb.

nov.—F. dodonaeiformis Gagnep. Lec. Notul Syst. 4 (1927) 90.

var. epiphytica (Elmer) Corner comb. nov.—F. williamsti C. B.

Robinson Philip. J. Sci. Bot. 3 (1908) 177.—var. epiphytica

Elmer, Leafl. Philip. Bot. 7 (1914) 2396.

var. memecylifolia Corner v. nov.—Frutex v. arbuscula —5 m.

alta. Lamina 2-7 0.7—2.7 cm., lanceolato-elliptica, acuminata,

saepe obtusiuscule: petiolo 1-7 mm. longo. Receptacula cecidi-

ophora 4—5 mm. lata, subglobosa, feminea 4-7 & 3-6 mm. el-

lipsoidea: pedunculo 3-14 « 0.5 mm. Flores cecidiophori tepalis

ovario valde angulato plerumque brevioribus. Semina 4 mm. longa,

1—4 per receptaculum. Borneo, Celebes, in silvis montanis muscosis

900-2000 m. alt., saepe epiphytica.

A v. oleaefolia pedunculo longiori gracillimi et a v. myrsinoide

lamina acuminata differt.

Borneo:—Bur. Sci. 2027; Clemens 20696, 27939 (typus, herb.

Br. Mus.) 26708, 28758, 30570, 31276, 34010, 40741; Endert

3837, 4268, 4398; For. Dept. B.N.B. A4461; Hallier 2673, 3421,

Haviland 87; Pickles 3701, 3772; San. A4173; Sing. F.n. 26338.

27817. Celebes:—Kjellberg 1668 (Makale), 2310 (B. Wawoe

Meoesa), 2728 (Porema); NIFS 227 (0. Masamba Lemboeng).

var. myrsinoides Corner v. nov.—Frutex v. arbor ~—13 m. alta,

_ trunco —20 mm. crasso. Lamina 2.5—-9 > 0.8—3 cm., spathulata

v. anguste obovata, apice obtusa v. rotundata: petiolo 1-10

425

Gardens Bulletin, S-

mm. Receptacula cecidiophora 5-6 mm. lata subglobosa,

feminea 7 X 4-5 mm. ellipsoidea: pedunculo 1.5—-8 & 0.7 mm.

Flores cecidiophori tepalis ovario laevi v. subangulato equalibus

v. longioribus. Semina 3—4 mm. longa, 1-3 per receptaculum.

Borneo, in silvis montanis muscosis 500-2100 m. alt., saepe

epiphytica.

Kinabalu: —Clemens 11005, 29529, 29915, 29930, 31410,

31863, 32490, 32671, 32936, 40087, 40968; Sing. F.n. 26977,

27546, 27550 (typus, herb. Singapore); San. A4228; Sinclair

9058. Sarawak:—SK 87 (Merurong Plateau). Indonesian Bor-

neo:—Amdjah 490 (Batoe Lesseng).

Receptaculo parvo gracile pedunculato, foliis obtusis distincta-

var. riparia Corner v. nov.—Frutex riparius. Lamina 6-9 x

1.5—2 cm., lanceolata, acuta v. breviter acuminata: costis late-

ralibus utrinsecus 13—16: petiolo 2~S mm. Receptacula cecidio-

phora 5—6 mm. lata subglobosa: pedunculo 15-20 0.7 mm.

Flores cecidiophori tepalis ovario angulato brevioribus v.-

longioribus. Sarawak (Upper Baram River, 200 m. alt., Sing.

F.n. 6871, typus herb. Singapore).

sect. Ficus ser. Podosyceae Corner subser. Podosyceae

F. neriifolia J.E. Sm. Rees Cyclop. 14 (1810) n. 21.—

F. gemella Wall. ex Mig. Hook. Lond. J. Bot. 7 (1848) 454.—

F. nemoralis Wall. ex Miq. var. gemella (Wall. ex Miq.) King,

Ann. R. Bot. Gard. Calc. 1, 2 (1888) 162, pl. 206.—Smith’s

type (n. 1610. 39) in the herbarium of the Linnean Society of

London is this characteristic plant with thick-walled fig.

var. fieldingii (Miq.) Corner comb. nov.—F. nemoralis Wall. ex

Mia. var. fieldingii King, Ann. R. Bot. Gard. Calc. 1, 2 (1888)

163.—F. fieldingii Mig. Hook. Lond. J. Bot. 7 (1848) 439.

var. nemoralis (Wall. ex Miq.) Corner comb. nov.—F.

nemoralis Wall. ex Miq. Hook. Lond. J. Bot. 7 (1848) 453.

var. trilepis (King) Corner comb. nov.—F. nemoralis Wall.

var. trilepis King, Ann. R. Bot. Gard. Calc. 1, 2 (1888) 162.—F.

trilepis Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 228, 294.

F. pedunculosa Miq. var. confertifolia (Merr.) Corner comb. nov.—

F. confertifolia Merr. Philip. J. Sci. 18 (1921) 59.

var. imberbis (Elmer) Corner comb. nov.—F. luzonensis Merr.

var. imberbis Elmer, Leafl. Philip. Bot. 4 (1911) 1323.

var. macropoda (Miq.) Corner comb. nov.—F. macropoda Mig.

Hook. Lond. J. Bot. 7 (1848) 442.

426

~ Vol. XVII. (1959).

var. mearnsii (Merr.) Corner comb. nov.—F. mearnsii Merr.

Philip. J. Sci. Bot. 3 (1908) 402.

var. segaarensis (Engl.) Corner comb. nov.—F. segaarensis

Engl. Bot. Jahrb. 7 (1886) 453.

var. velutina Corner var. nov.—Hispido-villosa pilis brunneis

0.5 mm. longis mollibus, lamina supra tenuiter villosa. Lamina —13

x 6 cm., elliptico-obovata, breviter acuminata, acuta, basi an-

guste subcordata v. cuneata: costis lateralibus utrinsecus 6-8,

basalibus ad 4 laminae elongatis: petiolo —-20 mm. longo. Recep-

tacula ellipsoidea 12-15 mm. lata: pedunculo 30-40 mm.: pedi-

cello 4-5 mm.: setis internis copiosis. Flores masculi dispersi.

Burma (Toong Dong, Wallich 4528, typus herb. Kew).

In view of the numerous collections of F. pedunculosa, these

species can now be treated only as varieties. The species extends

from Formosa to New Guinea but in Malaysia it is not to the west

of Celebes. The sole collection of v. velutina from Burma is

therefore exceptional and needs further collecting.

F. erecta Thunb. var. erecta f. sieboldii (Mig.) Corner comb.

nov.—F’. sieboldii Miq. Ann. Mus. Bot. Lugd. Bat. 2 (1866)

199.—F. erecta Thunb. var. sieboldii (Miq.) King, Ann. R.

Bot. Gard. Calc. 1, 2 (1888) 142, pl. 178B.

var. beecheyana (Hook. et Arn.) King f. koshunensis (Hayata)

Corner comb. nov.—F. koshunensis Hayata, J. Coll. Sci. Imp.

Univ. Tokyo 30 (1911) 276.—F. beecheyana Hook. et Arn. var.

koshunensis (Hayata) Sata, J. Soc. Tr. Agr. 6 (1934) 22.

F. trivia Corner sp. nov.—F. cuneata Lévl. et Vant. Fedde’s Rep.

4 (1907) 84, pr. p. Bodinier 2363 (alt. p. = F. heteromorpha

Hemsl.).—Frutex v. arbor —7 m. alta, foliis longipetiolatis spira-

liter dispositis. Ramuli glabri v. puberuli: petioli et costae

(subtus) pilis fulvidulis patentibus 0.5—1.5 mm. longis sparsim

puberuli, v. subglabri. Ramuli 2.5—4 mm. crassi, brunnei. Sti-

pulae 6-15 mm. longae, appresse pilosae v. pilis patentibus.

Lamina 5-18 x 2.3—10 cm., elliptica v. obovata, angustior ad

ramulos superiores, apice acuta v. breviter acuminata —10 mm.

longo, basi cuneata, integra, laevis, membranacea v. subcoriacea,

sicco obscure viridis v. brunneola: costis lateralibus utrinsecus

4—7, v. 3-4 in foliis minoribus, obliquis, intercostis laxis 2—5,

subtus leviter elevatis: costis basalibus utrinsecus 1—2, ad 3 la-

minae elongatis, glandulis basalibus 2: petiolo 9-70 mm. longo.

Receptacula axillaria binata, maturitate rubra v. purpurea:

_pedunculo 3~8 mm. longo, glabro. v. puberulo: bracteis basali-

bus 3, 1—1.5 mm. longis, acutis, puberulis: corpore receptaculi

427

Gardens Bulletin, S.

8—10 mm. lato, subgloboso v. subellipsoideo, ostiolo bracteis 3

planis ut disco 1.5—2 mm. lato occluso: setis internis nullis v.

paucis minutis: cellulis scleroticis nullis. Flores masculi pedi-

cellati, ostiolares et dispersi, cecidiophoris equales v. longiores:

staminibus 2. Flores cecidiophori sessiles v. pedicellati: tepalis

ovario albido sessili v. substipitato longioribus. Flores feminei

sessiles, tepalis ovario brevioribus: stylo glabro, stigmate obliqua,

haud bifido. Semina 2 & 1.5 & 1.3 mm., subreniformia, vix

carinata, laevia. Cystolitha hypogena. China, Tonkin, in silvis

montanis.

F. chapaensis Gagnep. affinis sed stipula hirta, lamina laevi,

pedunculo longiori differt.

China:—Bodinier 2363 (Kweichou); Cavalerie 7809, 7810

(Kweichou); Esquirol 2053 (Kweichou); Ford 175, 177 (Kwang-

tung); A. N. Steward et H. C. Cheo 446 (Kwangsi, Ling Yuen

Hsien); Y. W. Taam 59 (Kwangsi, pr. Pai Shou); Y. Tsiang 6827

(Kweichou); T. M. Tsui 523, 716 (Kwangtung, Yang Shan, pr.

Linchow). Indochina:—Bon 2709 (Tonkin, Dong Ham valley,

typus herb. Paris).

F. gasparriniana Mig. Hook. Lond. J. Bot. 7 (1848) 436—F.

silhetensis Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 223, 291.

—The type at Kew, though poor, leaves no doubt that it is the

earlier name for F. silhetensis, a variable species.

var. esquirolii (Lévl. et Vant.) Corner comb. nov.—F.

esquirolii Lévl. et Vant. Fedde’s Rep. 4 (1907) 84.—This is a

variety with lanceolate multicostate lamina.

Cavalerie 125, 129; Decker 144; Esquirol 2588 (typus); Faber

446; Y. Tsiang 1471, 6053; T. M. Tsui 643; E. H. Wilson 2789.

var. laceratifolia (Lévl. et Vant.) Corner comb. nov.—

F. laceratifolia Lévl. et Vant. Fedde’s Rep. 4 (1907) 85.—F.

rhomboidalis Lévl. Fl. Kouy-Tcheou (1915) 433.—This is a form

with lobed dentate leaves and, to judge from herbarium material

of v. gasparriniana, it is not its sapling state.

Bodinier 1663 (typus); Cavalerie 420, 7813, 7814; Ducloux

2202; Esquirol 3267 (typus, F. rhomboidalis), 6365; Faber 444;

W. P. Fang 2415, 5645; Henry 12090; W. R. Price 1268; A. N-

Steward et H. C. Cheo 598.

var. viridescens (Lévl. et Vant.) Corner comb. nov.—

F. cyanus Lévl. et Vant., et v. viridescens Lévl. et Vant. Fedde’s

Rep. 4 (1907) 84.—This is the more villous state of F. gasparri-

niana, which seems commoner at higher altitudes. There are many

collections.

428

Vol. XVII, (1959).

F. stenophylla Hemsl. emend. Rehder, J. Arn. Arb. 17 (1936) 79,

80.—var. macropodocarpa (Lévl. et Vant.) Corner comb. nov.

—F. macropodocarpa Lévl. et Vant. Fedde’s Rep. 4 (1907) 66.

—This is a state with long peduncles (13-55 mm.).

China:—Cavalerie 3589, 7812; Esquirol 838, 2216; A. N. Ste-

ward et H. C. Cheo 833. Laos:—Poilane 1880. Annam:—Poilane

6584, 29168.

var. nhatrangensis (Gagnep.) Corner comb. nov.—¥F. nhatran-

gensis Gagnep. Lec. Notul. Syst. 4 (1927) 92.—This differs in the

broader lamina (4-16 & 1.5—5 cm.) with fewer pairs of lateral

nerves (5—10).

Poilane 3431 (typus, Nhatrang); Chevalier 38658 (Nhatrang)

(? Kostermans 390, Thailand).

F. ischnopoda Mia. var. subcylindrica Corner v. nov.—Receptacula

corpore 20-30 « 7-10 mm., subcylindrico: pedunculo 10-25

mm. longo. Annam (Tourane, Clemens 3442, typus herb. Kew,

et 4009). Receptacula matura, floribus masculis apertis, non

juvenilia.

subsect. Ficus ser. Podosyceae Corner subser. Basitepalae Corner

F. pustulata Elmer var. lanceifolia (Sata) Corner comb. nov.—

F. cardinalicarpa Elmer, Leafl. Philip. Bot. 4 (1912) 1391.—

var. lanceifolia Sata et v. linearifolia Sata, Monogr. (1944) 356.

This differs from v. pustulata only in the lanceolate multicostate

leaf.

subsect. Eriosycea (Mig.) Corner ser. Eriosyceae

Corner subser. Eriosyceae

F. grossularioides Burm. f. Fi. Ind. (1768) 227.—F. alba

Reinw. ex Bl. Bijdr. (1825) 467. var. robusta Corner v. nov.—

Ramuli, petioli, costae (subtus) pilis fulvis patentibus 1—2.5 mm.

longis hirsuti, glabrescentes. Ramuli 4-7 mm. crassi. Lamina

major, ad ramulos fertiles saepe trilobata v. ovato-cordata. Re-

ceptacula 15—20 mm. lata (20-30 mm., viva): pedunculo 4-6

3-4 mm.: bracteis basalibus 3—4 mm. longis: setis internis co-

piosis. Flores feminei pedicellis —~ 6mm. longis praediti, stylo

hirsuto. Malaya.

Pahang (Fraser’s Hill, 1,200 m. alt.):—Sing. F.n. 33187

(typus, herb. Singapore), Purseglove 4317, Corner s.n. 10.viii. 37.

Selangor:—Sing. F. N. 32970 (? Kuala Selangor).

Omnibus partibus robustior bene distincta.

429

Gardens Bulletin, S-

var. stenoloba Corner v. nov.—Ut v. grossularioides sed lamina

lanceolata- elliptica (-14 & 4 cm.), juvenilis anguste lanceolato-

lobata (lobo medio 1~—2.5 cm. lato). Sumatra, in silvis montanis

1,200 m. altis, praecipue ad clivis volcanicis.

Bartlett 8310 (Simeloengoen); Lorzing 4356, 6585, 8511 (typus,

herb. Leiden), 9471, 9890 (Sibolangit); Ouwehaud 249; Teys-

mann 714 (pr. Siboga); v. d. Vecht s.n. 25. xii. 54 (Berastagi).

subsect. Eriosycea (Miq.) Corner ser. Eriosyceae Corner

suber. Trichosyceae Corner

F. hirta Vahl. var. appressa Corner v. nov.—Ramuli pilis appres-

sis 1-2 mm. longis, costa subtus 2-3 mm. longis, praediti. Re-

ceptacula pilis appressis vel patentibus praedita. Laos, North

Thailand, in silvis montanis 500-1,300 m. alt.

Laos:—Poilane 20052, 20378, 26141 (typus, herb. Paris),

26347.—Thailand:—-RFD 9552 (Chiengrai).

var. brevipila Corner v. nov.—Ramuli, petioli, costae (subtus),

et receptacula pilis brunneolis patentibus —i mm. longis villosi-.

Lamina 8-16 x 4.7—8.5 cm., obovata, basi cuneata, integra,

supra subscabrida, subtus aspero-villosa: costis lateralibus utrinse-

cus 5—7, basalibus 1, ad 1/3—4 laminae elongatis: petiolo 11—23.

mm. longo. Receptacula 12-15 mm. lata sessilia, dense villosa:

bracteis basalibus 1.5~2 mm. longis: setis internis copiosis flavi-

dulis. Pedicelli florum glabri. Yunnan, Annam, in silvis montanis.

1,200—1,500 m. alt.

Kingdom Ward 12755 (Yunnan, Mong Hai); Poilane 4046.

(Annam, Nhatrang, typus herb. Paris).

var. dumosa (King) Corner comb. nov.—F. dumosa King, Ann.

R. Bot. Gard. Calc. 1, 2 (1888) 151, pl. 190.—This differs merely

in being thinly hispid-villous and in having for the most part pal-

mately lobed leaves, few being simple. All collections come from

the neighbourhood of G. Dempo and G. Raja in Sumatra. The

Indochinese records are F. hirta v. imberbis Gagnep.

var. malayana Corner v. nov.—F. cordata Ridley, J. R. As. Soc.

Str. Br. 57 (1911) 93.—Receptacula 12-25 « 10-16 mm. (18-—

33 & 15-22 mm., viva), conica, sessilia: bracteis basalibus 6—9

x 3-5 mm., caducis. Perianthium masculum et cecidiophorum

plus minus cupulare, 3—4-dentatum. Pili 2—4 mm. longi. Nepal,

Assam, Indochina, Thailand, Hainan, Malaya (Perak, Pahang),

in silvis montanis 500—1,700 m. alt.

Inter v. hirtam et v. roxburghii (Miq.) King intermedia.

430

“Vol. XVI. (1959).

Nepal:—-Stainton, Sykes, et Williams 5241 (Rupukot Tal.). As-

sam :—King’s coll. s.n. (Lushai Hills, 1890), Hook. f. et Thomson

sm. (Khasia). Hainan:—C. I. Lei 118 (det. F. fulva). Laos:—

Poilane 26246 (Huat Laos, inter M. Luong Nam Eh et M. Sing).

Annam:—Poilane s.n. (prov. Quang Tri, Dong Iam Pe, 22. ix. 39).

Thailand:—Kerr 701 (Chiengmai, Doi Sootep), 7614 (Pattani,

Betong), 20134 (Kao Krading, Toi), Sing. F.n. 4704 (Kantang).

Perak:—Ridley 14624 (type, F. cordata), Wray 3413. Pahang:—

Purseglove 4264, For. Dept. 7776, Sing. F. n. 32571, Corner s.n.

10.viii. 37 (typus, herb. Singapore), Fraser’s Hill: For. Dept.

28165, Kep. F.n. 30986, Ridley 13720, Sing. F.n. 10948, 33550,

Cameron Highlands: Sing. F. n. 31911, Sungei Tahan.

var. Squamosa Corner v. nov.—Ut. v. roxburghii (Miq.) King

sed receptacula minora 20—25 mm. lata, v. -28 « 18-20 mm.,

bracteis lateralibus nonnullis crassiusculis 3-5 mm. longis praedita:

bracteis basalibus 6-8 mm. longis, subcaducis. Ramuli et petioli

pilis brunneis 2-6 mm. longis praediti: receptacula pilis brunneis

1—2 mm. longis sparsim hispida et pilis brevioribus pallidis villosa.

Stylus glaber. Folia plerumque palmatolobata. Sumatra, in silvis

montanis 1,000—1,600 m. alt.

Alston 15134 (Sidikalang, inter Lae Pondor et Silalahi; typus,

herb. Br. Mus.); NIFS bb. 6851 (Karoland, Tongkok); O. Hage-

rup s.n. (Toba, 25.xi. 16); Ridley s.n. (Berastagi, 8.11. 21).

Bracteis lateralibus in sect. Fico singularis.

F. simplicissima Lour. var. annamica (Gagnep.) Corner comb.

nov.—F. silhetensis Mig. var. annamica Gagnep. Lec. Fl. Gen.

I.-C. 5 (1928) 788.—F. touranensis Gagnep. Lec. Notul. Syst.

4 (1927) 97.

F. halmaherae Corner sp. nov.—Arbor —30 m. alta, foliis longi-

petiolatis spiraliter dispositis. Ramuli, petioli, costae (subtus)

pilis rigidis fulvidulis (? irritantibus) 1-2 mm. longis hispidi,

nervuli subtus pilis mollibus —0.5 mm. longis villosi, areolae

glabrae. Ramuli 10-12 mm. crassi. Stipulae — 25 mm. longae,

appresse pilosae. Lamina —20 cm. longa et lata (? multo major),

Ovata-cordata, acuminata, denticulata, membranacea: costis

lateralibus utrinsecus 4—6, intercostis —12 subtus elevatis; costis

basalibus utrinsecus 3—4, ad 4 laminae elongatis, glandulis basa-

libus 2: petiolo 100-160 mm. longo. Receptacula axillaria,

binata, pedunculata, primo ut ramuli hispida dein pilis rigidis

caducis molliter et breviter villosa: pedunculo 2.5-4 « 2.5-3

431

Gardens Bulletin, S.

mm.: bracteis basalibus 1.5—2 mm. longis, appresse puberulis:

corpore receptaculi -22 & 17 mm., ellipsoideo, ostiolo subde-

presso bracteis apicalibus numerosis parvis appresse pilosis

occluso; setis internis —1.5 mm. longis, flavidulis rigidis copiosis:

cellulis in pariete receptaculi omnibus fibriformibus elongatis

crasse tunicatis. Flores masculi ordinibus duobus ostiolares

sessiles: staminibus 2, filamentis rubris. Flores cecidiophori ses-

siles v. pedicellis glabris albidis -2 mm. longis praediti. Flores

feminei et semina ? Cystolitha nulla. Halmaheira (Gunong

Sembilan, c. 600 m. alt., “common along path in forest”; Pleyte

369, typus herb. Leiden).

Ut arbor Hibiscus, forsan F. mollissimae Ridley (Malaya)

affinis, sed cellulis fibriformibus in pariete receptaculi singularis.

F. fulva Reinw. ex BI. var. timorensis Corner v. nov.—Semina

laevia, vix carinata. Stylus glaber v. subpuberula. Setae recep-

taculi internae paucissimae, breves. Pedicelli florum femineorum

glabri. Timor (Forbes 3444 et 3579, typus herb. Br. Mus.).

F. subfulva Corner sp. nov.—Arbor —7 m. alta, foliis spiraliter

dispositis. Ramuli, petioli, et costae (subtus) pilis fulvidulis —0.5

mm. longis sparsim puberuli. Ramuli 4-6 mm. crassi. Stipulae

—15 mm. longae, appresse pilosae. Lamina 10-25 x 5-—11.5

cm., elliptica, apice acuminata —10 mm. longo, basi rotundato-

subcordata v. cuneata, integra, margine saepe recurvata, plus

minus rigide coriacea, utrimque scabrida, sicco brunnea: costis.

lateralibus utrinsecus 4—6, intercostis —10 subtus elevatis: costis.

basalibus utrinsecus 2—3, ad 3-2/3 laminae elongatis, glandulis

basalibus 2: petiolo 12-50 mm. longo. Receptacula axillaria

binata pedunculata, sparsim appresse puberula, glabrescentia,

maturitate rubra (cecidiophora et feminea): pedunculo 4-10

mm.: bracteis basalibus 2—4 mm. longis, ovato-acutis, appresse

pilosis: pedicello nullo: corpore receptaculi 10-12 mm. lato

(14-16 16-20 mm., viva), globoso v. ellipsoideo, ostiolo

bracteis apicalibus 3—S planis occluso sed maturitate hiascenti:

setis internis 2-4 mm. longis, flavidis rigidis copiosis: cellulis.

scleroticis copiosis. Pedicelli florum albido-setosi: tepala glabra

rubra spathulata. Stamina 2. Stylus glaber. Semina 1.5 mm.

longa, tuberculata, valde carinata. Cystolitha nulla. Borneo.

var. subfulva.—Kinabalu, Sing. F. n. 26612, 26642; G. Klam,

Hallier 2349, typus herb. Leiden.

var. villosula Corner v. nov.—Costae, intercostae et nervuli

reticulati subtus pilis fulvidulis —-0.5 mm. longis dense villosuli-

Setae internae 1—1.5 mm. longae: cellulis scleroticis nullis.

432

Vol. XVII. (1959).

Hallier 2869 (typus, herb. Leiden) et 2977, Lianggagang; Havi-

land 85 (Braang); Beccari 962 (Kuching, det. F. leucoptera Miq.);

Clemens 20698 (Sarawak, Bidi Cave).

F. auratae Migq. persimilis sed vero inter F’. fulvam et F. glan-

dulosam Wall., fere ut hybrida, referenda.

subsect. Eriosycea (Miq.) Corner ser. Eriosyceae Corner

subser. Dehiscentes Corner

F. glandulifera (Wall. ex Mig.) King var. camiguinensis (Merr.)

Corner comb. nov.—F. banahaensis Elmer, Leafl. Philip. Bot.

1 (1907) 252.—var. typica et var. camiguinensis (Merr.) Sata

Monogr. (1944) 267, 370.—F. camiguinensis Merr. Philip. J.

Sci. Bot. 9 (1914) 276.—This differs merely in details of hairi-

ness.

var. villosa Corner v. nov.—Ut v. glandulifera sed costis et

nervulis reticulatis subtus villosulis, pilis saepe undulosis, haud

glabrescentibus. Borneo, Celebes, Morotai, New Guinea.

Borneo:—Bur. Sci. 193 (B.N.B.); For. Dept. B.N.B. 2930;

Kostermans 6187 (pr. Sangkalang), 9546 (Mahakam River), East

Borneo. Celebes:—NIFS bb 24166 (O. Masamba, Biridondok).

Morotai:—Kostermans 1003, 1083. New Guinea:—BW 755

(Cyclops Mt.); Carr 12763 (Papua, Koitaki); Doctors v. Leeuwen

10537 (Nassau Mt.); Kanihera et Hatusima 13219 (Waren); NGF

3753 Sepik district, typus herb. Lae); NIFS bb 30745 (Biak),

31116 (Membramo, Hollandia).

subsect. Eriosycea (Migq.) Corner ser. Eriosyceae Corner

subsect. Cuneifoliae Corner

F. chartacea Wall. ex King var. lanceolata Corner var. nov.—

Lamina 7-23 x 1-3 cm., lanceolata: costis lateralibus utrin-

secus 9-16, angulo lato exeuntibus, intercostis O—1: petiolo 5—20

mm. Receptacula breviter stipitata. Frutex riparius, in silvis.

Annam, Thailand.

Annam:—Poilane 10574, 12233, 18313, 19770, 32287 (det.

F. macilenta King). Thailand:—Kerr 773 (Kaw Pa-ngan) 11724

(Nam Chut, Ranawng, typus herb. Br. Mus.); RFD 14834 (Nak-

hawn Srithamarat, Chawang).

F. litseifolia Corner sp. nov.—Frutex v. arbor —8 m. alta, trunco

—20 cm. crasso, cortice rufobrunneo, latice albo: foliis brevipe-

tiolatis spiraliter dispositis. Ramuli, petioli, stipulae, costae

(subtus) et receptacula pilis albidis v. brunneis —O.5 mm. longis.

433

Gardens Bulletin, S.

appressis tenuiter vestiti, v. costae subtus pilis brevibus patenti-

bus subscabrae. Ramuli 1.5-3 mm. crassi. Stipulae —7 mm.

longae. Lamina 4-14 x 1.5—5 cm., lanceolata, anguste elliptica

v. obovata, apice acuminata 7-18 mm. longo, basi cuneata v.

basi cuneata v. subrotundata, integra v. distanter denticulata,

raro sublobata, coriacea, laevis v. subtus subscabridula, sicco

viridis v. subdtus brunneola: costis lateralibus utrinsecus 4-8,

angulo lato exeuntibus, intercostis 0O—2 laxis, subtus leviter ele-

vatis: costis basalibus utrinsecus 1, brevibus v. ad 4 laminae

elongatis, glandulis basalibus nullis: petiolo 4-12 & 1 mm.

Receptacula axillaria binata glabrescentia, maturitate rubra:

pedunculo 2—7 mm. longo: bracteis basalibus 1 mm. longis: cor-

pore receptaculi 5—8 mm. lato, ostiolo plano bracteis apicalibus

parvis numerosis occluso: setis internis copiosis: cellulis sclero-

ticis nullis. Tepala 4—5, rubra libera glabra v. apice ciliata.

Flores masculi ordine uno sessiles ostiolares; staminibus 2.

Flores feminei et cecidiophori sessiles v. pedicellis pilosis —2 mm.

longis praediti: stylo femineo sparsim piloso. Semina 0.8—1 mm.

longa, tuberculata, carinata. Cystolitha nulla, lamina pilis mi-

croscopicis cystolithiformibus praedita. Malaya, Sumatra, in

silvis montanis 1,000—1,600 m. alt.

F. oreophilae Ridley affinis sed brevipilosa, receptaculis et

seminibus minoribus, florum pedicellis pilosis, petiolis brevioribus

differt.

Malaya: —Sing. F.n. 31094 (Trengganu, G. Padang). Sumatra : —

Altson 14740 (Atjeh, Blangkedjeren) ; Bartlett 11128 (Tapianoeli,

Toba); Forbes 2241 (Mt. Dempo); Jacobs 4672 et Meijer 3670.

5290 (Pajakumbah, Mt. Sago); v. Steenis 3548 (Palembang, G.

Raja, typus herb. Leiden).

F. tuphapensis Drake var. annamensis (Gagnep.) Corner comb.

nov. F. annamensis Gagnep. Lec. Notul. Syst. 4 (1927) 85.—

F. cambodica Gagnep. id. 87.—This differs from v. tuphapensis

in the larger fig, the two stamens rather than one in the male

flower, and the puberulous female style. It relates F. tuphapensis

closely with F. chartacea.

subsect. Eriosycea (Migq.) Corner ser. Auratae Corner

subser. Auratae

F. endospermifolia Corner sp. nov.—Arbor —25 m. alta, foliis

longipetiolatis spiraliter dispositis, omnibus partibus pilis brun-

neis rigidis patentibus 0.5—1.5 mm. longis hispido-villosis, ner-

vulis subtus villosulis v. minute puberulis. Ramuli 4-7 mm.

crassi, lente glabrescentes. Stipulae 10-30 mm. longae, appresse

434

Vol. XVIT. (1959).

pilosae. Lamina 13-25 cm. lata, tam longa quam lata v. paulum

latior, ovata, apice acuminata —12 mm. longo, basi cordata v.

subtruncata, denticulata, membranacea, supra hispidula, subtus

villosa, sicco brunnea: costis lateralibus utrinsecus 4—5, glandulis

axillaribus praeditis, intercostis strictis —16, subtus elevatis,

areolis glabris: costis basalibus utrinsecus 3—4, ad 4—2/3 laminae

elongatis, glandulis basalibus 2: petiolo 35-100 mm. longo,

glandula subnodali praedito. Receptacula axillaria binata ses-

silia, hispido-villosa v. tomentosa, 18-25 « 15~—20 mm. (23-

30 mm. longa et lata, viva), ovoidea v. subglobosa, ostiolo parvo

subdepresso: bracteis basalibus 3—5 mm. longis, ovatis, obtusis

v. breviter acuminatis, appresse pilosis, caducis: setis internis

—2.5 mm. longis, flavidis, copiosis: cellulis scleroticis in pariete

receptaculi cecidiophori copiosis, feminei sparsis v. nullis.

Florum pedicelli pilosi. Flores masculi 2.5—7 mm. alti, ordinibus

numerosis ostiolares, sessiles v. pedicellis -4 mm. longis praediti:

tepalis 4, rubris, margine ciliolatis, haud setosis: staminibus 2,

antheris breviter mucronatis. Flores neutri (in rec. fem.) ordini-

bus 2-3 ostiolares. Flores feminei et cecidiophori sessiles v.

pedicellis —4 mm. longis praediti: tepalis 3-4, lanceolatis v.

anguste spathulatis, flavidis, apicem versus dense setosis: stylo

femineo piloso. Semina 1—1.3 mm. longa, breviter oblonga, cari-

nata, tuberculata. Cysotolitha nulla. Borneo (Mt. Kinabalu), in

silvis montanis 1,000—1,800 m. alt.

Clemens 28223 (Tenompok), 34381 (Colombon River): For.

Dept. B.N.B. 3621, A4492 (Tenompok, typus herb. Kew); San.

15444 et Sinclair 8974 (Kota Belud. Kandasan); Sing. F.n. 26902.

27842 (Tenompok).

F. bruneiensis, F. eumorpha, et haec species arbores spectabiles

latifoliatae hispidissimae ad regionem North Borneo restrictae.

F. auratae affinis sed F. bruneiensis et F. endospermifolia inter

alia praecipue florum masculorum tepalis rubris, non setosis, ut

in ser. Eriosyceae distinctae.

F. bruneiensis Corner sp. nov.—Arbor —13 m. alta, cortice brun-

neo, latice albo copioso, foliis longipetiolatis spiraliter dispositis:

stipulis, petiolis costisque primo rubris (vivis). Petioli et costae

(subtus) pilis aureis v. brunneis rigidis patentibus 2-4 mm. lon-

gis strigosi, nervuli pilis brevioribus: nervuli reticulati et areolae

subtus toto minute velutinati. Ramuli 6-9 (—12) mm. crassi,

minute puberuli mox glabri, brunnei, cavi. Stipulae 20-60 mm.

longae, lanceolato-ellipticae caducae, appresse sericeo-puberuli,

pilis brunneis longis paucis ad carinam praediti. Lamina 28-45

x 16-28 cm., elliptica v. ovata, apice acuminata -15 mm-

435

Gardens Bulletin, S.

longo, basi cordata v. rotundato-cuneata, denticulata, membra-

nacae, scabrida, sicco viride-brunnea: costis lateralibus utrin-

secus 5—6, glandulis axillaribus praeditis, intercostis strictis —12,

subtus elevatis, areolis ut nervulis reticulatis minute velutinosis:

costis basalibus utrinsecus 3—4, ad 4 laminae elongatis, glandulis

basalibus 2: petiolo 80-180 m. longo, glandula subnodali prae-

dito. Receptacula axillaria binata sessilia, pilis pallidis 1-2 mm.

longis dense hispido-villosa, maturitate rubra, 24-30 « 20-26

mm. (pilis exclusis: -30 mm. lata, viva), subglobosa v. ovoideo-

conica, ostiolo bracteis apicalibus 3—5 occluso: bracteis basalibus

6-10 & 5-7 mm., ovatis v. oblongo-ellipticis, acutis, minute ap-

presse puberulis, caducis: setis internis —2.5 mm. longis, copiosis,

flavidis: cellulis scleroticis sparsis in pariete receptaculi cecidio-

phori. Flores masculi et cecidiophori ut in F. endospermifolia,

sed majusculi: aliquando staminibus 3. Cystolitha nulla. Brunei

(Ulu Belalong, prope flumen; Brunei 5338, typus herb. Cam-

bridge).

Inter F. endospermifoliam et F. brunneo-auratam, sed ramulis

‘subglabris, receptaculis majoribus. Forsan arbor montana secundo

flumine descendens.

F. brunneo-aurata Corner sp. nov.—Arbor —15 m. alta, foliis

longipetiolatis spiraliter dispositis, juvenilibus 3—5S palmato-loba-

tis. Indumentum duplex: ramuli et petioli pilis aureis v. fusco-

brunneis 1—4 mm. longis patentibus strigosi, costae subtus bre-

vioribus, et omnes partes (praeter paginam laminae superiorem)

pilis brevibus pallidis minute velutinosi. Ramuli 3-6 (—10) mm.

crassi. Stipulae 10-35 mm. longae (ad folia juvenilia —70 x

18 mm.) sericeae, pilis longis brunneis paucis ad carinam prae-

ditae. Lamina 15-30 * 6-14 cm., (-50 x 45 cm., juvenilia

lobata), ovato-elliptica v. elliptica, apice acuminata —10 mm.

longo, basi anguste subcordata v. cuneata, serrulata v. subintegra,

chartaceo-coriacae, sicco brunnea: costis lateralibus utrinsecus

(3—) 4-5 (-8), glandulis axillaribus praeditis, intercostis nu-

merosis strictis, subtus elevatis, areolis ut nervis et nervulis

minute velutinosis: costis basalibus utrinsecus 2—3, ad 1/3-2/3

laminae elongatis, glandulis basalibus 2: petiolo 15-60 mm.

longo, glandula subnodali praedito. Receptacula axillaria binata

sessilia velutinata glabrescentia, maturitate rubra, 11-16 mm.

lata (12-22 mm., viva) subglobosa, subsulcata, ostiolo bracteis

apicalibus 3 planis occluso: bracteis basalibus 1—1.5 mm. longis,

pusillis ovatis acutis: setis internis 1.5-3 mm. longis brunneolis

copiosis: cellulis scleroticis numerosis. Flore masculi ordinibus

436

Vol. XVII. (1959).

numerosis ostiolares, sessiles v. pedicellis subrubris sparsim

pilosis praediti: tepalis 4 rubris spathulatis, marginibus ciliolatis:

staminibus 2. Flores neutri (in rec. fem.) ostiolares, tepalis rub-

ris. Flores feminei et cecidiophori sessiles v. pedicellis glabris v.

sparsim pilosis, aliquando cecidiophoris induratis, praediti: tepa-

lis 3-4, dense setosis ut in F. aurata, albidis: stylo femineo rubro,

puberulo v. glabro. Semina ut in F. aurata. Cystolitha nulla. Bor-

neo, in silvis secundariis ad 1,100 m. alt., forsan communis.

F. hirta et F. fulva confusa et praecipue F. subfulva. Plantas

numerosas vivas in Sarawak et Brunei studui, semper distinctas.

Sarawak:—Beccari 1360 (det. F. hirta v. roxburghii); Corner

s.n. Jan. 1959; Haviland 153; Brunei:—Brunei 5307, 5308. Brit.

North Borneo, Kinabalu : —Sing. F.n. 25125 (Dallas), 26608 (Mene-

tendok Gorge, typus herb. Singapore). Indonesian Borneo: -—~

Hallier 1519 et 2525 (G. Klam), 2909 (Lianggagang), det. F. fulva.

F. aurata Mig. var. brevipilosa Corner v. nov.—F. in-

aequipetiolata Merr. Philip. J. Sci. 21 (1922) 517.—Ramuli,

petioli, et receptacula pilis brevioribus 0.5—1 mm. longis spar-

sim hirsuti, nervuli subglabri. Lamina 12—26 « 2.5—8.5 cm.,

oblonga, lanceolata, v. subovata, utrimque scabriuscula: costis

lateralibus utrinsecus 4—9, v. 16—20 in laminis oblongis v. lan-

ceolatis. British North Borneo.

Bur. Sci. 178, 485, 602 (typus), 1141, 1885; Creagh s.n. (1895);

For. Dept. B.N.B. 2459, 9368, 9371: Ridley s.n. (1887).

var. longipilosa Corner v. nov.—Ramuli, petioli, et costae

(subtus) pilis 2-4 mm. longis villosi (1-2 mm. in v. aurata). Lamina

elliptico-obovata v. oblonga. Malaya, Sumatra, Riouw Archipelago,

Borneo.

Malaya (Singapore to Perak): —For. Dept. 12110, 15076; Hullett

381; Hume 9203; King’s coll. 8553; Sing. F.n. 29387, 32313 (typus.

herb. Singapore). Sumatra :—Bartlett 7885 (Asahan); Curtis 3548

(Indragiri); Postumus 563, 674, 685 (Djambi). Riouw :—Bunne-

meyer 6952 (Lingga); Buwalda 6229 (Riouw). Borneo:—Brune1

575; Hallier 2238 (G. Klam); Motley 600 (Banjermasim); Teys-

mann 7899 (det. F. hirta).

var. palawanensis Corner v. nov.—Quisumbing, Philip. J. Sci.

41 (1930) 317 (ut F. chrysocarpa).—Folia sicca fusco-brunnea

(aureo-brunnea in v. aurata): juvenilia breviter trilobata, basi cor-

dato-subtruncata, petiolo —16 cm. longo: lamina superiora 15-28

< 8-17 cm., elliptica v. obovata, basi cordata v. late cuneata; cos-

tis lateralibus utrinsecus 4—6. Pili ut in v. aurata: nervuli subtus

437

Gardens Bulletin, S.

villosuli, areolae glabrae. Receptacula 14-18 & 12-14 mm. Pala-

wan, Balabac. Bur. Sci. 49913, 77382 (typus, herb. Singapore),

77425.

Forsan species nova, F. brunneo-auratae affinis. Folia juvenilia

palmato-lobata in F. aurata mcognita.

var. pedunculata Corner v. nov.—Ut v. aurata sed receptacula

pedunculis 2-5 mm. longis, cellulis scleroticis in pariete praedita.

Malaya. ;

Perak : King’s coll. 3738, 4328, 5834, 7776. 10106, 10225 (det.

F. hirta et F. chrysocarpa); Scortechini 365 (typus, herb. Singapore),

529: Wray 493. Trengganu:—Kep. F.n. 26990 (S. Kemaman);

Selangor :—-H. M. Burkill 1021 (S. Buloh), For. Dept. 12900 (Gin-

ting Simpak). Java: —-HHBB 135866, 135867, 135868 (cult.).

subsect. Eriosycea (Miq.) Corner ser. Auratae Corner

subser. Monandreae Corner

F. andrechaete Corner sp. nov.—Arbor —10 m. alta, foliis spirali-

ter dispositis. Ramuli, stipulae, petioli, et receptacula pilis rigidis

appressis flavidulis 0.5-1 mm. longis tenuiter vestiti, glabre-

scentes: costae subtus pilis patentibus sparsis, nervuli subglabri.

Ramuli 3—4 mm. crassi. Stipulae —7 mm. longae. Lamina 11-20

x 4-9 cm., elliptica v. subobovata, saepe angusta, apice acumi-

nata -—10 mm. longo, basi cuneata, integra, subcoriacea,

utrimque subscabrida, sicco brunnea: costis lateralibus utrin-

secus 4—6, obliquis, intercostis —4, subtus leviter elevatis: costis

basalibus utrinsecus 1, ad 1/3-4 laminae elongatis, glandulis

basalibus 2: petiolo 12-50 mm. longo: lamina juvenilia —33

13 cm., oblongo-elliptica, apice -35 mm. longo acuminata, haud

lobata, costis lateralibus utrinsecus 5—8, petiolo —-110 mm. longo.

Receptacula axillaria binata: pedunculo 3-5 x 2 mm.: bracteis

basalibus 1-1.5 & 2-3 mm., ovatis, subacutis: corpore recep-

taculi 12-15 mm. lato, subgloboso, ostiolo parvo plano: setis

internis —1.5 mm. longis, copiosis: cellulis scleroticis nullis.

Florum pedicelli setosi. Flores masculi ordinibus duobus ostio-

lares, secundo pedicellati: tepalis 3, setosis, albidis: stamine uno.

Flores feminei et cecidiophori sessiles v. pedicellis -5 mm. longis

praediti: tepalis 3—4, setosis, albidis: stylo glabro. Semina 1.3

mm. longa, carinata, tuberculata. Cystolitha nulla. North

Borneo, in silvis prope flumina.

F. macilentae affinis, sed omnibus partibus major.

Brunei 389, 5309, 5310, 5320 (Ulu Belalong, typus herb. Cam-

bridge); For. Dept. B.N.B. San. 16285, 17397; Nieuwenhuis

(Taheri) 952 (Tepoese, Lilit boelan).

438

Vol. XVI. (1959).

F. macilenta King var. gibbsiae (Ridley) Corner comb. nov.—

F. gibbsiae Ridley, J. Linn. Soc. Bot. 42 (1915) 137.—This differs

from v. macilenta in the pedunculate fig, and the less hairy leaves

and fig.

var. ilicifolia Corner v. nov.—Ramuli, petioli, stipulae, costae

(subtus), et receptacula sparsim et minute puberuli, glabrescentes.

Lamina 2.5-11 & 1-4 cm., lanceolato-elliptica, apice acuminata

-10 mm. longo, basi cuneata, grosse et distanter dentata dein

integra, coriacea, utrimque scabriuscula, sicco flavo-viridis et

subtus aureo-brunnea: costis lateralibus utrinsecus 4—8, intercostis

0-1, nervulis reticulatis subtus leviter elevatis: costis basalibus bre-

vibus: petiolo 8-40 mm. Receptacula solitaria, 7-11 mm. lata:

pedunculo 3-10 mm. longo: cellulis scleroticis copiosis. Kinabalu,

1500-2000 m. alt. Clemens 50712, Gurulau spur; Sing. F. n.

26952, path to Ranau, typus herb. Singapore.

A var. gibbsiae pedunculo longiori, lamina subscabrida, saepe

dentata differt.

F. eumorpha Corner sp. nov.—Arbor —18 m. alta, Ramuli, folia,

et receptacula pilis fusco-brunneis patentibus 0.5—1.5 mm. longis

dense hispidi, nervuli subtus et areolae pilis albidis v. fulvidulis

—0.5 mm. longis minute velutinosi. Ramuli 4—S mm. crassi.

Stipulae —15 mm. longae, strigoso-hispidae. Lamina 8-20 x

6.5—-14 cm., ovato-cordata v. subtriangularis, longior quam lata,

breviter acuminata, dentata, membranacea, supra _hispido-

pustulata, sicco griseo et subtus aureo-brunnea: costis lateralibus

utrinsecus 5—7 (-—8), intercostis —6, subtus elevatis, areolis im-

pressis velutinosis: costis basalibus utrinsecus 3, ad 1/3-4

laminae elongatis, glandulis basalibus 2: petiolo 40-85 mm.

longo. Receptacula axillaria binata sessilia, aureo-hispida, 16—17

< 14-15 mm. (pilis exclusis; 18—20 mm., viva), subglobosa v.

ellipsoidea, ostiolo bracteis apicalibus 3-4 subumbonatis

occluso: bracteis basalibus 3—3.5 mm. longis, ovato-acutis, ap-

presse pilosis: setis internis 1-2 mm. longis, copiosis. cellulis

scleroticis sparsis. Flores masculi et cecidiophori ? Flores neutri

ordine uno ostiolares sessiles. Flores feminei sessiles v. pedicellis

albido-setosis —2.5 mm. longis praediti: tepalis 3, albidis, dense

setosis: stylo puberulo vy. glabro. stigmate bifido. Semina 1—1.2

mm. longa, subcarinata, subtuberculata (? immatura). Cystolitha

nulla. Borneo.

var. eumorpha.—Sing. F. n. 27566, Tibobah River (¢ypus herb.

Singapore); Clemens 32524, Marai Parai “perhaps the tallest Ficus

in this vicinity”.

439

Gardens Bulletin, S.

var. subglabra Corner v. nov.—Areolae subtus puberula v.

subglabra. Florum femineorum tepala glabra v. ad apicem pilis

1—3 praedita. Receptacula, minora, 8-9 « 10—11 mm., bracteis

basalibus obtusis, cellulis scleroticis copiosis. Central East Bor-

neo (W. Koetai, Mt. Kemoel, 1,800 m. alt.; Endert 4416, typus

herb. Leiden).

Inter F. endospermifoliam et F. paramorpham sed. floribus

masculis incognitis, incertae sedis.

F. paramorpha Corner sp. nov.—Frutex v. arbor parva. Ramuli,

petioli, et costae (subtus) pilis fusco-brunneis patentibus —0.5

mm. longis villosi, nervuli subtus pilis brevioribus, aliquando

ramuli et petioli appresse pilosi. Ramuli 2—3 mm. crassi. Sti-

pulae —7 mm. longae, appresse brunneo-pilosae. Lamina 6-14

< 3-5.5 cm, elliptica, breviter acuminata, basi cuneata, serrato-

denticulata, plus minus chartacea, supra hispido-scabra, sicco

grisea et subtus aureo-brunnea: costis lateralibus utrinsecus 5—6,

intercostis —5, subtus valde elevatis, areolis quasi glabris: costis

basalibus utrinsecus 1 (—2), ad 3-1/3 laminae elongatis, glan-

dulis basalibus 2: petiolo 12-25 mm. Receptacula axillaria

binata sessilia, pilis fusco-brunneis brevibus appressis tenuiter

vestita, maturitate rubra, 9-10 mm. lata, subglobosa: bracteis

basalibus 1.5 mm. lJongis, ovato-subacutis: setis internis —2.5

mm. longis, copiosis, aureis: cellulis scleroticis copiosis. Florum

pedicelli setosi: tepala 3-4 albido-setosa. Flores masculi ordine

uno ostiolares: stamine uno. Flores feminei et cecidiophori ses-

siles v. pedicellati: stylo glabro. Semina 1—1.2 mm. longa, sub-

carinata, subtuberculata (? immatura). Cystolitha nulla. Borneo

Kinabalu, 1,600—2,200 m. alt.).

Clemens 31777, Upper Kinataki River, typus herb. Leiden:

32523, Marai Parai.

F. eumorphae affinis sed lamina elliptica, receptaculis minoribus

inter alia differt.

F. setifiora Stapf var. adelpha Corner v. nov.—Ramuli, petioli,

costae (subtus), et receptacula pilis albidis v. aureo-brunneis

obliquis 0.5—1 mm. longis villosi. Ramuli 2—2.5 mm. crassi.

Lamina 4.5-12 « 2-4.5 cm., anguste elliptica v. lanceolata,

breviter acuminata, basi anguste cuneata, supra hispidulo-

scabra, subtus villosa: petiolo 3-18 mm. Receptacula 6-7 mm.

lata, pedunculo 1-2.5 mm. longo. Stylus femineus_pilosus.

Semina obtuse carinata, valde tuberculata. North Borneo

Kinabalu, 1,100—2,500 m. alt.).

Ut speciminibus exsiccatis videtur, decidua. A var. Setiflora

indumento villoso, lamina anguste elliptica, pedunculo distincto

differt.

440

Vol. XVII. (1959).

Sing. F.n. 26930, 27876 (Tenompok, typus herb. Singapore):

Sinclair 8975 (Bt. Kinasaruban); Clemens (Tenompok, Colombon

River, et Mt. Nunok) 28402, 28433, 29353, 29482, 30295, 32684.

32719, 33905, 34453, 40123.

var. puberula Corner v. nov.—Ramuli et petioli pilis —-O.5 mm.

longis subvillosuli, costae subtus sparsim puberulae. Receptacula

sparsim appresse puberula, pedunculo 0.5-1 mm. longo. Stylus

femineus glaber. Semina obtuse carinata, valde tuberculata. Central

East Borneo (W. Koetai, Mt. Kemoel, 1.700 m. alt.: Endert 4275,

typus herb. Leiden).

Forsan v. adelphae forma.

441

Taxonomic Notes on Ficus Linn., Asia

and Australasia

IV. SUBGEN. FICUS SECT. SYCIDIUM MIQ.

By E. J. H. CORNER

Botany School, University of Cambridge

Summary

New sections, subsections, and series:—-subsect. Sycidium ser.

Prostratae, Pungentes, Phaeopilosae, and Copiosae ser. nov.;

subsect. Varinga (Miq.) Corner comb. nov., ser. Heterophylleae,

Cyrtophylleae, and Exasperatae ser. noy.; subsect. Palaeomorphe

(King) Corner comb. nov., ser. Subulatae and Fibrosifoliae ser.

nov.

Earlier specific epithets: —F. semicordata B. Ham. ex J.E. Sm.

(F. cunia B. Ham. ex Roxb.), F. pungens Reinw. ex BI. (F. myrio-

carpa Miq.), F. wassa Roxb. (F. duriuscula King), F. fraseri Miq.

(F. proteus Bur.), F. coronata Spin. (F. stephanocarpa Warb.), F.

asperiuscula Kunth et Bouché (F. leptorhyncha Koord. et Val.),

F. virgata Reinw. ex BI. (F. decaisneana Migq.), F. heteropleura

BI. (F. urophylla Wall. ex Mig.), F. sinuata Thunb. (F. rostrata

Lam.).

New names for later homonyms: —F. gul Laut. et K. Schum.

(F. rudis Mig., non Pers.), F. andamanica Corner (F. macropoda

Kurz, non Miq.), F. elmeri Merr. (F. semicordata Migq., non J.E.

Sm.), F. celebensis Corner (F. irregularis Miq., non Steud.).

New species: —F. koutumensis, F. complexa, F. porphyrochaete,

F. subsidens, F. griseifolia, F. primaria, F. cauta, F. goniophylla,

F. tenuicuspidata et v. major, F. oleracea et v. pugnans, F. chryso-

chaete, F. imbricata, F. gryllus, F. erinobotrya et v. solomonensis

et f. glabrior, F. leptogramma, F. praetermissa, F. midotis, F.

leptocalama, F. rubrocuspidata, F. rubromidotis.

New subspecies and varieties: —-F. gul Laut. et K. Schum. v.

lasiocarpa Corner, v. solomonensis Corner; F. montana Burm. f.

v. purpurascens (Bl.) Corner; F. madurensis Mig. v. angustifolia

Corner; F. copiosa Steud. v. pubescens Corner; F. wassa Roxb.

v. nubigena (Diels) Corner, v. obversifolia (Miq.) Corner; F.

balica Miq. v. colffsii Corner; F. cumingii Mig. v. androbrota

(Summerh.) Corner v. angustissima (Merr.) Corner, v. worcesteri

(Merr.) Corner; F. fiskei Elmer v. multinervia Corner; F. riedelii

442

Vol. XVIT. (1959).

Mig. v. minor Corner; F. ampelas Burm. f. v. linearis Corner, v.

soronensis (King) Corner; F. guyeri Elmer v. sibuyanensis (El-

mer) Corner; F. melinocarpa Bl. v. hololampra (Diels) Corner;

F. trachypison Laut. et K. Schum. v. pallida Corner; F. tonsa Mia.

v. aspera Corner, v. leptodictya (Diels) Corner, v. subcordata Cor-

ner; F. irisana Elmer v. validicaudata (Merr.) Corner; F. oppo-

sita Mig. v. micracantha (Miqg.) Corner, v. indecora (A. Cunn.

ex Miq.) Corner; F. storckii Seem. v. kajewskii (Summerh,) Cor-

ner; F. tinctoria Forst. f. ssp. gibbosa (Bl.) Corner et v. rigida

Miq., ssp. parasitica (Willd.) Corner et v. anastomosans (Wall.

ex Kurz) Corner, ssp. swinhoei (King) Corner; F. virgata Reinw.

ex Bl. v. philippinensis (Mig.) Corner, v. sessilis (Bur.) Corner;

F. subulata Bl. v. gracillima (Diels) Corner; F. sinuata Thunb.

v. oblonga Corner, ssp. cuspidata (Reinw. ex Bl.) Corner; F. hete-

ropleura Bl. v. hirta Corner, v. mindanaensis (Warb.) Corner; F.

aurita Reinw. ex Bl. v. auriculifera (Merr.) Corner, v. celebica

(Reinw. ex Bl.) Corner; F. obscura Bl. v. angustata (Mig.) Cor-

ner, v. borneensis (Miq.) Corner, v. kunstleri (King) Corner.

Notes on F. madurensis Mig., F. ampelas Burm. f., F. scobina

Benth., F. aspera Forst. f., F. cyrtophylla Wall. ex Miq., F. obs-

cura Bl.

sect. Sycidium Mig.

Hook. Lond. J. Bot. 7 (1848) 228; Ann. Mus. Bot. Lugd. Bat.

3 (1867) 291; King, Ann. R. Bot. Gard. Calc. 1, 2 (1888) 73;

OK in T. Post Lex. Gen. Phan. (1904) 236; Sata, Monogr. (1944)

252.—subgen. Sycidium (Miq.) Mildbr. et Burr. Engl. Bot. Jahrb.

46 (1912) 175—Dioecious. Figs pedunculate or pedicellate with-

out a collar of basal bracts, rarely sessile: body often with scatter-

ed lateral bracts. Tepals free or shortly joined, red, pink, or white.

Male flowers ostiolar: stamen 1, or 1—2: anther not mucronate.

Style subterminal: stigma simple, subclavate. Seed small, lenticular

or shortly oblong, generally with a single keel, smooth or, in a

few species, reticulate: hilum rarely prominent. Cystoliths various-

ly present. Africa, Asia, Australasia, c.100 spp. Lectotypus:—F.

aspera Forst. f.

In 1848 Miquel made three series of this section. King (1887)

extrated ten species along with numerous synonyms, and made of

them sect. Palaeomorphe King. These comprise most of the two se-

ries Cuspidate and Pallidae, which I refer accordingly to subsect.

Palaeomorphe. The species of the third series Scabrae Miq. were

mostly retain by King in sect. Sycidium, the typification of which

should, therefore, come from this series. Choice of type might

favour F. montana Burm f., which appears with six other species:

443

Gardens Bulletin, S.

now known to be synonyms, in ser. Scabrae, but F. montana has

generally a gall-ovary in the male flower and should have been

placed by King in sect. Palaeomorphe. Second choice might be F.

scabra Forst. f., but Miquel does not appear to have seen a spe-

cimen of it in 1848. Hence I choose the related F. aspera Forst.

f. which makes the largest group of species representative of the

section and subsect. Sycidium.

subsect. Sycidium.—sect. Covellia (Gasp.) Miq. subsect. Pandani-

florae Sata, Monogr. (1944) 293 (ut Pandanusiiflorae; lecto-

typus, F. minahassae Teysm. et de Vr.).—subsect. Pseudosy-

cidium Sata, id. 253, 258, 381 (lectotypus, F. fiskei Elmer) .—

Seed lenticular, as long as wide, slightly keeled all round or in

the upper half, rarely not at all. Male flowers without a normal

gall-ovary, sometimes with a rudiment. Fig often pedunculate

with a collar of three basal bracts. Cystoliths often papillate.

Trees or shrubs, not epiphytic or climbing. Trop. Asia, Austra-

lasia, 65 spp. Lectotype, F. aspera Forst. f.

ser. Prostratae Corner ser. nov.—Albido- v. brunneo-pilosae.

Folia brevipetiolata disticha. Receptacula geocarpica ad ramos

efoliatos internodis elongatis e trunco basim versus orientes, brac-

teis basalibus ternatis pedunculata, corpore saepe bracteis laterali-

bus paucis, setis internis praedito, cellulis scleroticis nullis. Tepala

rubra glabra. Stamine 1—2. Ovarium rubrum v. albidum, stylo

glabro. Semina !—1.5 mm. longa, hilo non prominenti. Cystolitha

plerumque hypogena. Continens Asiatica, 3 spp. Typus, F. semi-

cordata B. Ham. ex J. E. Sm. (F. cunia B. Ham.).

This series relates with sect. Ficus, but habit and convenience

place it here. F. semicordata has two stamens in the male flower, F.

prostrata has one, while in F. koutumensis the male flowers are

not known. The first two species have usually been placed in sect.

Sycocarpus (Covellia) because of their cauliflorous habit, but the

free tepals at once exclude them, and relate them with the following

two series.

ser. Pungentes Corner ser. nov.—Bosscheria Teysm. et de Vr.

Nat Tijd. N.I. 23 (1861) 212.—sect. Covellia (Gasp.) Miq. sub-

sect. Pandaniflorae Sata, Monogr. (1944) 293 (ut Pandanusii-

florae; lectotypus, F. minahassae Teysm. et de Vr.).—Brunneo-

setosa. Folia longipetiolata spiraliter disposita, aut brevipetiolata et

disticha. Receptacula parva pedunculata, bracteis basalibus terna-

tis, lateralibus deficientibus, rami- et cauliflora ad ramos longos

efoliatos pendentes sed haud geocarpicos: setis internis copiosis

longis albidis: cellulis scleroticis nullis. Tepala 3—4 rubra glabra.

445

Vol. XVII. (1959).

Ovarium brunneo-rufum, femineum stylo hirsutissimo. Semina

0.5—0.7 mm. longa, hilo parvo prominenti. Cystolitha hypogena.

Borneo, Philippines, Malaysia Orientalis, 3 spp. Typus, F. pungens

Reinw. ex BI.

As the preceding series, this has also been wrongly assigned to

sect. Sycocarpus. F. minahassae and F. pungens are closely allied,

but the third, F. petrotica Diels, has distichous, short-petiolate,

leaves.

ser. Phaeopilosae Corner ser. nov.—Purpureo- v. brunneo-

setosae v. hirtae. Folia longipetiolata symmetrica spiraliter

disposita. Receptacula axillaria v. cauliflora, raro geocarpica,

sessilia v. pedicellata, bracteis lateralibus dispersis, ad corpus re-

ceptaculi saepe bene evolutis: cellulis scleroticis nullis. Tepala

rubra glabra. Ovarium albidum, stylo glabro. Borneo, Philippines,

Malaysia Orientalis, 7 spp. Typus, F. complexa Corner.

This is the most generalised series and it relates to the more

generalised species of sect. Adenospermak and sect. Sycocarpus.

Except for F. gul Laut. et K. Schum., the species occur in New

Guinea.

ser. Copiosae Corner ser. nov.—Albido-pilosae v. glabrae, saepe

scabridae. Folia plerumque longipetiolata et symmetrica, spira-

liter disposita v. decussata. Receptacula axillaria v. rami- et cauli-

flora, pedicellata bracteis lateralibus paucis parvis dispersis: cellu-

lis scleroticis nullis v. paucis. Tepala purpureo-rubra pallescentia

v. albida, saepe puberula v. hispidula. Ovarium albidum. India

usque ad insulas Solomon, 11 spp. Typus, F. copiosa Steud.

ser. Scabrae Mig. Hook. Lond. J. Bot. 7 (1848) 228—subgen

Palaeomorphe (King) Sata sect. Palaeomorphe ser. Nonminutuli-

florae Sata (lectotype, F. ulmifolia Lam.), subser. Fulvobrunnei-

foliae Sata (lectotypus, F. ulmifolia Lam.) et subser. Metallici-

foliae Sata (lectotypus, F’. blepharistoma Warb.), Monogr. (1944)

223, 224, 378.—subgen. Eumetamorphe Sata sect Sycidium Miq.

subsect. Sycidium ser. Viridifoliae Sata (lectotypus, F. worcesteri

Merr.) et ser. Lineariangustifoliae Sata (lectotypus, F. angustissima

Merr.), Monogr. (1944) 253, 254, 380.—subsect. Pseudosyci-

dium Sata (lectotype, F. fiskei Elmer )et ser. Scabricordatogib-

bosiifoliae Sata (lectotype, F. fiskei Elmer), ser. Lanceifolicae Sata

(lectotype, F. celtoides Elmer), ser. Subscabririgidiifoliae Sata

(lectotype, F. ampelas Burm. f.), Monogr. (1944) 253, 258, 260,

262, 381, 382.—Albido-, luteo- v. brunneo-pilosae. Folia brevi-

petiolata decussata, spiraliter disposita, v. disticha, saepe asym-

metrica et scabra. Receptacula axillaria v. cauliflora, pedunculata

445

Gardens Bulletin, S.

v. pedicellata: cellulis scleroticis nullis v. copiosis. Ovarium albi-

dum: stylo puberulo v. glabro. Seminis hilum non prominens.

Madagascar, Ryu Kyu, Formosa, Malay Archipelago, Australasia,

c. 50 spp. Lectotypus, F. aspera Forst. f.

This is the short petiolate remainder, comprising the majority of

the species. Let us hope that some of these synonyms may never

have to be used.

subsect. Varinga (Miq.) Corner comb. nov.—Ficus sect. Carica

Mig. subsect. Varinga Miq. Ann. Sci. Nat. ser. 3, 1 (1844) 33

(haud sect. Varinga Mig. sensu OK in T. Post Lex. Gen. Phan.

1904, 236).—Ut subsect. Sycidium sed semina breviter oblonga:

tepala et ovarium albida. Arbores v. frutices, raro sarmentosae

(Africa), haud epiphytica. Africa, Ceylon, Asia continentalis,

Sumatra, Java, Borneo, c. 13 spp. Lectotypus, F. scabrella Roxb.

(F. heterophylla Linn. f.).

As the character of subsect. Varinga, Miquel gave the unista-

minate flower. This excludes all the species which he cited except

F. scabreilla Roxb., which is a synonym of F. heterophylla Linn.

f., and F. radicans Roxb. which, as a synonym of F. heteropleura

Bl., is provided for in subsect. Palaeomorphe (King) Corner.

Hence my choice of F. scabrella as lectotype. The species much

resemble those of ser. Scabrae (subsect. Sycidium) but cannot be

fitted in with them, and the group permeates a different geogra-

phical region.

ser. Heterophylleae Corner ser. nov.—Receptacula axillaria

solitaria, 10-20 mm. lata v. latiora, plerumque pedunculata brac-

teis basalibus ternatis, lateralibus et setis internis deficientibus.

Tepala plus minus glabra. Semina laevia. Lamina plus minus sym-

metrica. Africa, Asia, 6 spp. Typus, F. heterophylla Linn. f.

ser. Cyrtophylleae Corner ser. nov.—Receptacula binata, axil-

laria v. ad truncum ramosque fasciculata, pedicellata, bracteis late-

ralibus dispersis. Tepala plus minus glabra. Semina laevia. Folia

disticha brevipetiolata, saepe asymmetrica. Africa, Asia, 6 spp.

Typus, F. cyrtophylla Wall. ex Miq.

ser. Exasperatae Corner ser. nov.—Receptacula axillaria, ple-

rumque solitaria, pedicellata, bracteis lateralibus dispersis. Tepala

hispida. Semina reticulata. Africa usque ad Indiam, Ceylon, 1 sp.

F. exasperata Vahl.

subsect. Palaeomorphe (King) Corner comb. nov.—Ficus sect.

Paleomorphe King, Ann. R. Bot. Gard. Calc. 1 (1887) 1, 3.—

subgen. Palaeomorphe (King) Sata, J. Soc. Tr. Agr. 6 (1934)

446

Vol. XVII. (1959).

26; Monogr. (1944) 217.—Ficus sect. Grossularia OK in T.

Post Lex. Gen. Phan. (1904) 236.—Semina oblonga, apice

carinata v. gibbosa, laevia. Flores masculi plerumque ovario

cecidiophori praediti. Folia disticha brevipetiolata. Receptacula

pedicellata v. sessilia, bracteis haud in collare ternatis. Cysto-

litha haud papillata. Arbores, frutices, saepe epiphytica etiam

suffocantes, v. sarmentosae: ramulorum radicumque cortice

saepe flavescenti. Asia, Australasia, 19 spp. Lectotypus, F. gib-

bosa Bl. = F. tinctoria Forst. f. ssp. gibbosa (Bl.) Corner.

The tendancy is to discard this group because of the variable

development of the gall-ovary in the male flower in the whole

section. When, however, the shape of the seed, the leaf-arrange-

ment, and the habit of the plants are taken into account, the

well-developed, insect-inhabited, gall-ovary in these species is, as

King perceived, a good character. Thus the group is certainly dis-

tinguishable as a natural entity from subsect. Sycidium, and should,

perhaps, be given its original sectional rank. However, I prefer to

await the elucidation of the African and Madagascan species and

their relation with ser. Copiosae (subsect. Sycidium), for they im-

pinge on the species of subsect. Palaeomorphe.

ser. Pallidae Mig. Hook. Lond. J. Bot. 7 (1848) 433.—ser.

Euglabrifoliae Sata (lectotypus, F. tinctoria Forst. f.) Monogr.

(1944) 217, 218, 377.—ser. Glabrifoliae Sata (lectotypus, F.

philippinensis Miq.), id. 262, 382.—Arbores mediocres v. gran-

des, saepe epiphyticae et suffocantes. Stipula terminalis plerumque

conspicua, recta, rigida. Lamina coriacea, haud caudata, plerum-

que integra, sicco pallida v. subtus brunneo-areolata. Receptacula

sicca laevia, bracteis lateralibus nullis: setis internis minutis.: cel-

lulis scleroticis in pariete copiosis. Tepala puberula v. hispidula.

Cystolitha amphigena. Asia, Australasia, 3 spp. Lectotypus, F.

tinctoria Forst. f.

Of the species assigned first to this series by Miquel, F. hede-

racea Roxb. belongs to sect. Rhizocladus Endl., F. septica Burm

f. to sect. Sycocarpus Miq., and F. undulata B. Ham. (F. nervosa

Heyne) to subgen. Pharmacosycea Miq. These excluded, the re-

maining eleven are F. tinctoria and F. virgata Reinw. ex Bh. or

their synonyms. I choose F. tinctoria as type because it is the better

known.

ser. Subulatae Corner ser. nov.—Arbores parvae v. frutices,

saepe epiphyticae v. sarmentosae, haud suffocantes. Stipula ter-

minalis conspicua curvata gracilis. Lamina subcoriacea v. mem-

branacea, acuminata v. caudata, integra. Receptaculis sicca con-

tracta et rugosa, plerumque bracteis lateralibus paucis parvis

447

Gardens Bulletin, S.

praedita: setis internis et cellulis scleroticis nullis. Tepala albida.

Flores feminei et cecidiophori pedicellis induratis, tepalis linea-

ribus v. subulatis. Cystolitha hypogena, raro amphigena. India

usque ad insulas Solomon, 2 spp. F. subulata Bl. (typus), F.

armiti King. |

ser. Cuspidatae Mig. Hook. Lond. J. Bot. 7 (1848) 428.—ser.

Glabriusculifoliae Sata (lectotype, F. caudatifolia Warb.), Monogr.

(1944) 217, 220, 378.—Arbores parvae, frutices, v. sarmentosae.

Stipula inconspicula. Lamina caudata, dentata v. integra, sicco

nervis reticulatis supra leviter elevatis, saepe fusco-brunnea. Recep-

tacula raro bracteis lateralibus praedita: cellulis scleroticis nullis.

Tepala albida membranacea. Semina valde carinato-gibbosa. Cys-

tolitha hypogena. India usque ad Malaysiam Occidentalem, 4 spp.

Lectotypus, F. cuspidata Reinw. ex Bl. = F. sinuata Thunb. ssp.

cuspidata (Reinw. ex Bl.) Corner.

I have taken F. cuspidata as lectotype because it agrees with

Miquel’s diagnosis of the series and obviously provided the name.

ser. Minutuliflorae Sata, Monogr. (1944) 217, 222, 378.—

Arbores parvae v. frutices, saepe epiphyticae, plerumque luteo-

v. brunneo-pilosae. Stipula inconspicua. Lamina caudata, basi

saepe auriculata, dentata v. integra, membranacea. Receptacula

saepe bracteis lateralibus parvis 1—2 praedita: cellulis scleroticis

nullis. Tepala albida membranacea. Cystolitha hypogena, raro

amphigena. Sumatra, Borneo, Philippines, Celebes, Moluccas, New

Guinea, 3 spp. Lectotypus, F. hispidulosa Elmer (F. aurita Bl.).

ser Fibrosifoliae Corner ser. nov.—Ut ser. Minutuliflorae sed

lamina fibris microscopicis sclereidiformibus in mesophylla copio-

sis praedita. Stipulae saepe conspicuae. Flores feminei et cecidio-

phori sessiles. Tepala rubra v. albida. Cystolitha amphigena v.

hypogena. Burma, Thailand, Malaysia Occidentalis usque ad in-

sulas Philippinenses et Moluccanas, 7 spp. Typus, F. obscura Bl.

This group is abundantly distinct in the microscopic feature of

sclereid-like fibres excurrent from the vascular bundle-sheaths and

permeating the whole mesophyll between upper and lower epi-

dermis. It is a feature which can be observed very easily and

quickly in a minute fragment of dried leaf-tissue cleared by plac-

ing in a drop of dilute potash on a slide and warming for a

minute: the fibres appear like white worms under a low power of

the microscope. The species are closely related and show, as any

other natural group of Ficus, the common evolutionary trends in

leaf and receptacle.

448

Vol. XVII. (1959).

subsect. Sycidium ser. Prostratae Corner

F. semicordata B. Ham. ex J. E. Smith, Rees Cyclop. !4 (1810)

i /1.—F. cape Clam. ex Roxb. Fil. Ind. 3 (1832): 561;

Wight Ic. t. 648: (F. cunea B. Ham. ms.).—This is represented

by two sheets in the Smith herbarium of the Linnean Society of

London, namely n. 1610. 26 (narrow leaf) and 1610.27 (broad

leaf, the lectotype). The name antedates F. semicordata Miq.

for which F.. elmeri Merr. must be used. The common name for

the species has been F. cunia, of uncertain etymology, but it is

spelled F. cunea on the labels of B. Hamilton’s specimens.

var. conglomerata (Roxb.) Corner comb. nov.—F. conglome-

rata Roxb. Fl. Ind. 3 (1832) 559; Wight Ic. t. 669.—F. cunia B.

Ham ex Roxb. var. conglomerata (Roxb.) Kurz, For. Fl. Br.

Burma 2 (1877) 461.

F. koutumensis Corner sp. nov.—Arbor —9 m. alta, foliis sym-

metricis distichis. Ramuli, petioli, et costae (subtus) pilis albidis

v. brunneis —1.5 mm. longis hispidi, nervuli pilis brevioribus:

lamina supra pilis albidis appressis scabrida, glabrescens. Ra-

muli 4-6 mm. crassi, ulteriores 2-3 mm. Stipulae —17 mm.

longae, puberulo-glabrescentes v. glabrae, subcaducae. Lamina

10-21 « 3-9.5 cm., oblongo-elliptica, apice acuminata —15

mm. longo, basi late cuneata v. rotundata symmetrica, denti-

culata v. serrulata, subcoriacea, supra scabrida, subtus hispido-

scabrida, sicco brunnea: costis lateralibus utrinsecus 6-8,

obliquis, intercostis -10, subtus leviter elevatis: costis basalibus

utrinsecus 2—3 (—4), ad 3—4 laminae elongatis, glandulis basali-

bus 2: petiolo 12-55 mm. -longo. Receptacula ad ramulos

efoliatos geocarpicos ut in F. semicordata J. E. Sm., stipulis plus

minus persistentibus; pedunculo 3-10 mm. longo, albido-villoso

dein glabro: bracteis basalibus in collare ternatis, 2-3 mm.

longis, glabris v. appresse puberulis: corpore receptaculi 15—20

mm. lato, albido-villoso dein glabro, bracteis lateralibus nullis v.

unico, ostiolo plano bracteis apicalibus planis 4—S occluso: setis

internis paucis albidis: cellulis scleroticis nullis. Flores feminei

ut in F’. semicordata, plus minus sessiles: tepala 3—4 libera rubra

lanceolata, ovario equalia: ovario sessili v. breviter stipitato,

rubro: stylo valde laterali, glabro, stigmate simplici. Semina

laevia, vix carinata et vix gibbosa. Cystolitha hypogena v. supra

vix evoluta. Annam, pr. Koutum, in silvis montium 1,000-—

1,800 m. alt.

Poilane 18220, mont. Mam Ray: 32027, pr. Moi de Tu-inh

(typus, herb. Paris).

449

Gardens Bulletin, S.

Lamina symmetrica ut in F. prostrata Wall. sed denticulata,

scabrida, et hispida ut in F. semicordata J. E. Sm.: stipulis et

seminibus intermedia.

subsect. Sycidium ser. Pungentes Corner

F. pungens Reinw. ex BI. Bijdr. (1825) 478.—F. myriocarpa Mig.

Ann. Mus. Bot. Lugd. Bat. 3 (1867) 230, 296.—The type of

F. pungens (Reinwardt 1486, Ternate) is sterile. I know of no

way to distinguish sterile material surely of F. minahassae

Teysm. et Binn. and F. myriocarpa Miq., but there are later col-

lections of F. myriocarpa from Ternate and Morotai, whereas

F. minanassae occurs to the westward (Talaud, Philippines,

Celebes, North Borneo) and does not stretch into the Moluccas

proper or to New Guinea, which is the domain of F. myrio-

carpa. Hence I identify F. pungens with F. myriocarpa. King

mistook a Moluccan species of sect. Sycocarpus (F. calcarata

Corner sp. nov.) for F. pungens Reinw. in which he was fol-

lowed by Diels.

subsect. Sycidium ser. Phaeopilosae Corner

F. complexa Corner sp. nov.—Arbor —7 in. alta, latice albido,

foliis longipetiolatis spiraliter dispositis. Ramuli et petioli pilis

brunneis v. purpureis 1-3 mm. longis erectis v. curvatis rigidis

strigosi, glabrescentes: costae subtus pilis brunneolis 0.5—1 mm.

longis appressis v. patentibus, nervuli puberulo-scabridi: lamina

supra pilis appressis rigidis sparsis, glabrescens. Ramuli 4-6 mm.

crassi, glandulis subnodalibus deficientibus. Stipulae 15-25

9-15 mm., ovato-lanceolatae, persistentes, appresso-puberulae

v. subglabrae v. ad carinam pilis erectis brunneis strigosae.

Lamina 18-32 x 12-19 cm., late elliptica v. ovata, apice

gracili acuminata —25 mm. longo, basi cordata, symmetrica,

denticulata, utrimque scabrida, membranacea, sicco fusca et

subtus griseo-viridis: costis lateralibus utrinsecus 5—7, plerumque

glandula axillari praeditis, intercostis 5-10, subtus elevatis: cos-

tis basalibus utrinsecus 3—4, ad 1/3—4 laminae elongatis, glandu-

lo axillari praeditis: petiolo 4-15 cm. longo. Receptacula axil-

laria, solitaria v. binata, etiam rami— et cauliflora ad ramulos

congestos, internodis vix evolutis, -5\0 «x 5 mm., efoliatos,

maturitate brunneo-rubra: pedicello 4-15 « 3—4 mm., bracteis

lateralibus dispersis 2-5 < 2-4 mm., appressis v. patentibus,

haud ternatis, praedito: corpore receptaculi 10-13 mm. lato

(16-20 mm., vivo), subgloboso, interne ob flores breves cavo,

pilis albidis —1 mm. longis laxe hispido, glabrescenti scabrido,

bracteis lateralibus appressis numerosis ovato-lanceolatis acutis

450

Vol. XVII. (1959).

5-16 & 4-9 mm, receptaculis juvenilibus toto obscurantibus,

ostiolo bracteis similibus erectis numerosis 4-8 mm. longis oc-

cluso: setis internis minutis sparsis: pariete crasso (4-5 mm.,

vivo), cellulis scleroticis nullis. Tepala 4—5 fusco-rubra, spathu-

lata v. in floribus sessilibus ovato-lanceolata, libera, glabra. Flores

masculi ostiolares sessiles, ordinibus 1—2 instructi: stamine 1,

anthero minute mucronato, pistillodio nullo. Flores cecidiophori

sessiles v. pedicellis glabris —1 mm. longis praediti, feminei sub-

sessiles: ovario sessili albido: stylo glabro, stigmate subinfundi-

buliforme puberulo. Semina 1 mm. longa, lentiformia, laevia,

vix compressa, subcarinata. Cystolitha hypogena: pili papillati.

Papua, in silvis montium 1,000—1,600 m. alt.

Brass 23213, Milne Bay district, Maneau Range, Mt. Dayman;

Carr 14023 (Lalu River), 14660 (Boridi), 15753 (Isuarava,

typus herb. Br. Mus.).

F. conocephalifoliae Ridley affinis sed lamina latiori, plus ovato-

cordata, receptaculi bracteis lateralibus longioribus floribus sessi-

libus v. breviter pedicellatis. An F. eustephana Diels, e specimine

miserabili descripta ?.

F. porphyrochaete Corner sp. nov.—Arbor --12 m. alta, latice

albido subseriflua, foliis longipetiolatis spiraliter dispositis.

Ramuli, petioli, et receptacula setis purpureo-brunneis 1-4 mm.

longis (ad costas et receptacula 1—2 mm.) irritantibus erectis

hispidi, costae subtus pilis brunneolis plus minus appressis:

lamina supra sparsim et appresse albido-hirsuta, glabrescens

scabra. Ramuli 5—7 mm. crassi. Stipulae 15-35 x 6-11 mm.,

ovato-lanceolatae, persistentes, appresso-hirsutae. Lamina 23—40

x 11-20 cm., elliptico-obovata, apice acuminata —25 mm.

longo, basi rotundato-cuneata v. subcordata, denticulata, mem-

branacea, utrimque scabrida, sicco griseo-viridis, cyanea, Vv.

brunneola: costis lateralibus utrinsecus 6—8, curvato-ascendenti-

bus, glandula axillari praeditis, intercostis 6-12, subtus elevatis:

costis basalibus utrinsecus 3, ad 4-1/3 laminae elongatis,

glandulis basalibus 2: petiolo 25-75 mm. longo. Receptacula

axillaria et cauliflora in cumulis magnis (—-60 receptaculis ag-

gregatis) ad ramulos efoliatos congestos -50 & 5-10 mm., in-

ternodis vix evolutis, maturitate e brunneo rubescentia; pedicello

3-20 mm. longo, in receptaculis axillaribus brevi, bracteis

lateralibus 1—2 (—3) dispersis, haud ternatis, ovato-lanceolatis

puberulis, 1-2 mm. longis, praedito: corpore receptaculi 8-11

mm. lato (setis exclusis: 17-19 mm., vivo), subgloboso hispido

glabrescenti, bracteis lateralibus paucis 1-2.5 %& 3-5 mm.,

ostiolo bracteis numerosis erectis, externis ad apicem recurvatis,

1-1.5 mm. longis occluso: setis internis brevibus, brunneolis,

451

Gardens Bulletin, S.

sparsis v. copiosis: cellulis scleroticis nullis. Tepala 4—6 fusco-

rubra albomarginata oblonga libera glabra. Flores masculi ostio-

lares sessiles, ordine uno instructi: stamine 1, non mucronato,

pistillodio nullo. Flores feminei et cecidiophori sessiles v. pedi-

cellis albidis glabris —2.5 mm. longis praediti: ovario sessili

albido-luteo: stylo glabro. Semina 0.8—1 mm. longa, lentifor-

mia, subcarinata, laevia, hilo vix prominenti. Cystolitha amphi-

gena: pili papillati. Papua, Solomon Islands, in silvis —5O0 m.

alt.

Brass 24162, Papua, Milne Bay district, Kwagira River, Peria

Creek (typus, Gray herb. Harvard): Carr 16381, Papua, Kokoda;

J. H. L. Waterhouse B146, B146a (Bougainville Isl, Siwai), 236

(New Georgia).

F. complexae Corner affinis sed receptaculis minoribus setosis

parvibracteatis, cystolithis amphigenis.

F. gul Laut. et K. Schum. Fl. Deutsch. Schutzgeb. Suds. (1901)

278.—-F. rudis Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 222,

291 (non Pers.).—F. keyensis Laut. et K. Schum. id. 270.—F.

manilensis Warb., Perkins Fragm. Fl. Philip. 3 (1905) 199.—

This is a widespread, common, and variable species extending

from North Borneo and the Philippines to New Guinea and New

Britain.

var. lasiocarpa Corner v. nov.—Receptacula pilis albidis, lute-

olis, v. brunneolis 1-2 mm. longis dense villosa, bracteis latera-

libus obscuratis. Morotai, Halmaheira, Netherlands New Guinea.

Aet 218 (Exp. Lundequist) et BW 6889, SW New Guinea,

Kaimana; A. W. Herre 636, Manokwari; Idjan 183, Halmaheira,

Goal; Kanehira et Hatusima 13353, Momi, 60 miles south of

Manokwari (typus, herb. Bogor); Kostermans 1348 (Morotai).

2619 (Manokwari, Warnapi).

var. solomonensis Corner v. nov.—Ramuli. petioli, et costae

pilis 1-2 mm. intensius brunneis villosi. Receptacula pilis brunneis

—0.5 mm. longis hispida, 9-12 mm. lata, bracteis lateralibus 1.5—3

mm. longis, bracteis apicalibus 1-1.5 mm. longis projicientibus:

pedicello —12 mm. longo. Tepala ad apicem pilosa. Solomon

Islands.

Brass 2578, San Cristoval, Waimamuru; Kajewski 1651, 1712

(Bougainville, Kupei Gold Field; typus 1651, herb. Kew) 2118

(Bougainville, Buin, Koniguru), 2796 (San Cristoval, Puepue

River).

A v. gul receptaculis majoribus et bracteis apicalibus projicienti-

bus differt: ad F. porphyrochaete Corner tendit.

452

Vol: XVET 2459).

subsect. Sycidium ser. Copiosae Corner

F. montana Burm. f., Fl. Ind. (1768) 226.—F. quercifolia Roxb.

Fl. Ind. 3 (1832) 534; Wight Ic. t. 646; King, Ann. R. Bot.

Gard. Calc. 1 (1888) 77, pl. 95.—This is undoubtedly the cor-

rect name for F. queércifolia Roxb., though Burmann’s type

(herb. Delessert, Geneva) had been misnamed F. ampelas.

var. purpurascens (Bl.) Corner comb. nov.—F. purpurascens BI.

Bijdr. (1825) 471; King, Ann. R. Bot. Gard. Calc. 1 (1888)

75, pl. 93.—Leaves purple or violet beneath. Java.

A recent collection (Meijer 3981, Tjibodas) proves that this is

merely a variety of F. montana. Blume’s type is at Leiden.

F. madurensis Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 222,

291.—F. smaragdina S. Moore, J. Bot. (1925) 171.—F. co-

piosa Steud. sensu Ridley, Fl. Mal. Pen. 3 (1924) 340.—This

has a more tree-like habit than F. montana, and is truly cauli-

florous, with more persistent stipules, larger leaves, and broader

tepals; and it is a plant of high forest, unlike F. montana. How-

ever, it requires field-study and it would be desirable to grow it

along with F. montana, F. andamanica, F. wassa, and F. co-

piosa, which may be almost impossible to distinguish from sterile

material. Because it has been misunderstood, I cite the collec-

tions which I refer to F. madurensis:—

Tenasserim:—Beddome 7489 (F. smaragdina, type). Thailand:

—Kostermans 176, Kwae Noi Exped., pr. Wangka. Perak:—

Wray 1732, Sing. F.n. 12728, Curtis s.n. May 1890, (Maxwell’s

Hill). Pahang:—Ridley 13719 (Telom), Sing. F.n. 31335, 32941

(Cameron Highlands). Selangor:—Ridley 8496, Robinson and

Kloss s.n. 7 Sept., 1917, (Ginting Simpah). Negri Sembilan:—

Ridley s.n. 23 Dec., 1920 (Bt. Tangga), Sing. F.n. 9856 and Fur-

tado s.n. 16 June, 1937 (Ulu Bendol). Sumatra:—Beccari 772

(Padang; referred to F. copiosa Steud. by King); Borssum-Waal-

kes 1060 (Krakatau); Lorzing 4632 (Sibolangit), v. Steenis 9850

(Atjeh, Gajolanden). Java:—Zollinger 3803.4 (It. sec., Br.

Mus.); Horsfield s.n. (Br. Mus.); Reinwardt s.n. (Utrecht, det. F.

subcrenata Miq. ined.) Madura:—de Vriese s.n. (type, Leiden),;

Zollinger 2449.

It looks as if the species has become extinct in Java.

var. angustifolia Corner v. nov.—Lamina 20-25 x 5-6.5 cm.

lanceolata. Sumatra.

Krukoff 4178, Asahan, Masihi For. Res. (typus: herb. Leiden);

Lorzing 5662 Sibolansi).

453

Gardens Bulletin, S.

F, andamanica Corner nom. nov.—F. macropoda Kurz, Prelim.

Rep. For. Veg. Pegu App. A (1875) p. cxxiii; For. Fl. Br.

Burma 2 (1877) 459, (non Miquel).—This is close to F. madu-

rensis but has a longer fig-pedicel, internal bristles, puberulous

tepals, and no ovary in the male flower. These are small points

but they show that the two are not identical, and both require

further field-study. In having the reticulate seed, it may connect

F. exasperata Vahl (Peninsular India, Ceylon, East Africa)

with F. madurensis and F. montana of Western Malaysia. I have

seen the following collections: —

South Andaman Isl.:—Kurz s.n. and n.2 (syntypes, but with

n.2, at Kew, are mixed figs of F. variegata Bl.); King’s collector

s.n. and n. 326; Parkinson 411. Barren Isl.; Prain s.n. 1891, Nar-

condam, common. Gt. Cocos Isl.:—Prain s.n. (det. F. brevicuspis

Miq.).

F. subsidens Corner sp. nov.—Frutex laxe ramosa, ramulis de-

cumbentibus v. repentibus, internodis elongatis, foliis longipetio-

latis laxe spiraliter dispositis, scabridula. Ramuli 3-4 mm.

crassi. Stipulae caducae. Lamina 17-22 « 9-11.5 cm., ellip-

tica, apice breviter acuminata -—10 mm. longo, basi late

cuneata, denticulata, utrimque scabrida, membranacea, sicco

brunneo-viridis: costis lateralibus utrinsecus 8—10, intercostis

strictis —11, subtus leviter elevatis: costis basalibus utrinsecus

1 (2), brevibus, glandulis basalibus 2: petiolo 30-70 x 3-4

mm. Receptacula binata, axillaria ad ramulos frondosos et ad

caules repentes defoliatos, scabridula, maturitate externe et in-

terne roseo-rubra: pedicello 5-8 mm. longo, bracteis lateralibus

2-3 parvis dispersis, haud ternatis: corpore receptaculi 6—8 mm.

lato (10-11 mm., vivo), subgloboso, bracteis lateralibus parvis

paucis, ostiolo subdepresso bracteis apicalibus umbonatis 4—5

occluso: setis internis et cellulis scleroticis nullis. Flores masculi

et cecidiophori? Flores feminei sessiles v. pedicellis glabris —lmm.

longis praediti: tepala 3—5 lineari-lanceolata, leniter gamophylla,

glabra, albida: ovario sessili v. substipitato, parte inferiori car-

noso: stylo glabro. Semina 1—1.2 mm. longa, lentiformia, vix

compressa, subcarinata, laevia. Cystolitha ut pili microscopic

amphigena. North Borneo (Kinabalu), inter lapides riparia, c.

1,500 m. alt.

Sing. F.n. 26443, Kinataki River, leg. C. E. Carr (typus, herb.

Singapore); Clemens 32402, pr. mont. Nunkok.

Species notabilis, in distributione singularis, F. montanam Burm.

f. revocans sed semina haud reticulata et lamina majoribus, inter-

costis pluribus.

454

Vol. XVII. (1959).

F. copiosa Steud. var. pubescens Corner v. nov.—F.

krausseana Rechinger, Fedde’s Rep. 11 (1912) 180.—F. acan-

thophylla Summerh. J. Arn. Arb. 10 (1929) 142.—Ramuli,

petioli, lamina (subtus), et receptacula juvenilia pilis albidis

-1 mm. longis villosi v. hispiduli, plus minus muriculati. Moluc-

cas, New Guinea, New Britain, New Ireland, Solomon Islands,

Queensland.

Moluccas:—Beguin 1827, Boger 9 (Halmaheira); Kostermans

1339 (Morotai); Labillardiere s.n. (Boeroe et Amboina). New

Guinea:—Brass 1388, 8749, 24127; Carr 11728, 11732, 16445;

Hoogland 4209; Kostermans 310 (typus, herb. Leiden), 493,

2853; NGF 861. New Britain:—-Waterhouse 247. New Ireland :—

Labillardiére s.n. Solomon Isl.:—Brass 2580 (San Cristoval),

3193 et 3196 (Ysabel); Kajewski 1836, Rechinger 4713, Water-

house 156, 226, B324, 801 (Bougainville). Queensland:—S. T.

Blake 14976; Kajewski 1178 et C. T. White 8021 (det. F. mag-

nifolia F.v.M.).

Ut species nullo modo distinguenda.

F. wassa Roxb. Fl. Ind. 3 (1832) 539; Wight Ic. t. 666.—F.

duriuscula King, Ann. R. Bot. Gard. Calc. 1 (1888) 155, pl.

195.—F. eulampra K. Sch. Fl. Deutsch. Schutzgeb. Suds.

(1901) 279.—F. lamprophylla Laut. et K. Schum. id. 271.—

F. portus-finschii Warb. in Laut. et K. Schum. Nachtr. FI.

Schuztgeb. (1905) 250.--—F. rhodocarpa Summerh. J. Arn. Arb.

10 (1929) 150.—F. anggica Diels, Engl. Bot. Jahrb. 67 (1935)

198.—F. wassa is identifiable from Roxburgh’s specimen (herb.

Martii, Brussels), his description, and Wight’s figure. F. copiosa

Steud. is identifiable from’ Roxburgh’s description of F. poly-

carpa Roxb. (non Jacqu.) and Wight’s figure (t. 632), but I

have found no specimen of Roxburgh’s. I treat F. copiosa in the

customary sense, as the species with large leaves and figs, but

Buch. Hamilton 2426 (at Edinburgh), labelled “F. polycarpa

Roxb. Hort. Beng. 66: colitur in horto prope Calcuttam botanico

e Moluccis”, resembles F. wassa rather than F. copiosa. Wassa

is a common name in Eastern Malaysia for any rough-leafed fig,

comparable with ampelas in Western Malaysia, and may signify

F. ampelas, F. melinocarpa, F. heteropoda, F. copiosa, F. wassa

or any other.

var. nubigena (Diels) Corner comb. nov.—F. nubigena Diels,

Engl. Bot. Jahrb. 67 (1935) 209.—F. caroli Diels id. 200.—

Diels indicated that F. caroli might have to be reduced to F. nubi-

gena. Many specimens are now at hand for the common F. wassa

455

Gardens Bulletin, S.

and several of these are intermediate to F. nubigena which, in fact,

differs merely in the following, rather vague, and, in some respects,

probably phenotypic characters :—

Leaves mostly decussate; lamina elliptic, often dentate; petioles

2-22 mm., rather short. Tepals dark red, fading pinkish white.

New Guinea, mountain and mossy forest, 1,300—2,400 m. alt.

Brass 5098; BW 3087, 3101; Carr 13304, 13630;/513656,

14211, 14342, 14866, 15979, 16095, 16402; Eyma 4913, 5319,

5424; Kanehira et Hatusima 13637; Ledermann 11826 (F. caroli,

type), 11970, 12535 (F. nubigena, type), 12901; Nielsen 808;

NGF 7847.

var. obversifolia (Miq.) Corner comb. nov.—F. ampelas Burm.

{. var. obversifolia Mig. Ann. Mus. Bot. Lugd, Bat. 3 (1867)

272.—F. reticulatissima S. Moore, J. Bot. (1925) Suppl. 108.—

Scabridissima. Ramuli, petioli, et costae (subtus) pilis pustulatis

~0.5 mm. longis muriculati. Flores, Timor, Halmaheira, Morotai.

Timor: Forbes 3551, 3704, 3294. Flores:—Elbert 4321. Hal-

maheira:—Nedi 221, Teysmann 5649 (typus, herb. Leiden).

Morotai:—Kostermans 1487.

Muriculata ut F. copiosa Steud. sed in omnibus partibus minor

ut in F. wassa.

F. balica Mig. var. colfsii Corner v. nov.—Ramuli, petioli, et

receptacula albido-villosi, haud glabrescentes: lamina subtus

molliter villosa. Ramuli 2—4 mm. crassi. Lamina basi cordata:

costis lateralibus 6-8 v. 8-11. Receptacula pedicellis 8-30 mm.;

corpore 12-15 mm. lato. Flores sessiles v. pedicellis rubris —1

mm. longis praediti; tepala 4-5 rubra libera oblonga, glabra v.

puberula. Flores masculi ordinibus duobus instructi: stamine 1

(—2), pistillodio nullo. Lombok, Sumbawa, Flores, in silvis ad

1200 m., alt.

Colfs 295, Sumbawa (typus, herb. Leiden); Elbert 4220, Flores;

Rensch 645, Sumbawa; Elbert 895, 896, 1651, Lombok.

F. griseifolia Corner sp. nov.—Arbor —8 m. alta, latice seriflua,

foliis laxe spiraliter dispositis. Petioli, laminae pagina superior.

et costae (subtus) pilis rigidis conicis erectis —0.5 mm. longi

sparism muriculati, nervuli subtus scabriduli; ramuli stipu-

laeque glabri. Ramuli 2-3 mm. crassi, fusco-brunnei. Stipulae

5—9 mm. longae, graciles, liberae, caducae. Lamina 8.5—17.5

< 5-12 cm., ovato-elliptica, apice acuminata 12—22 mm.

longo, basi cordata v. rotundata, saepe subasymmetrica, serru-

lato-denticulata, rigide membranacea, utrimque scabrida, sicco

griseo-olivacea v. plumbea: costis lateralibus utrinsecus 5—8, in-

tercostis strictis —8, subtus elevatis: costis basalibus utrinsecus

456

Vol. XVIT. (1959).

2—3, ad 1/3 laminae elongatis, glandulis nullis v. indistinctis:

petiolo 20-55 & 2 mm., glandula subnodali praedito. Recep-

tacula rami- et cauliflora ad ramulos efoliatos congestos —30

< 3-4 mm., basi —10 mm. latos, internodis vix evolutis: pedi-

cello 15-20 & 1 mm., gracili, sparsim muriculato v. glabro,

bracteis lateralibus glabris subacutis 1-3, 0.5-1 mm. longis,

dispersis, haud ternatis: corpore receptaculi 7 & 6 mm. (imma-

turo), ellipsoideo, subscabrido, bracteis lateralibus 1—2 parvis

appressis, ostiolo bracteis apicalibus erectis numerosis acutis

glabris 1-1.5 mm. longis occluso: setis internis —1 mm. longis,

flexuosis, copiosis, albidis: cellulis sclerotocis nullis. Tepala 4—5

fusco-rubra ovata-lanceolata libera glabra: florum pedicelli al-

bidi glabri. Flores masculi ostiolares, sessiles v. breviter pedi-

cellati, ordinibus duobus instructi: stamine 1, non mucronato,

pistillodio plus minus bene evoluto. Flores cecidiophori sessiles

v. pedicellati: ovario sessili, albido-luteo, ad basim lateris sty-

laris rubro-marginato. Cystolitha hypogena. Papua (Hoogland

3993, Alola, Iora valley. pr. Kokoda, 1200 m., alt.; typus, herb.

Lae).

Ut F. gui Laut. et K. Schum. sed plus minus glabra; muriculata

ut F. copiosa Steud; bracteis apicalibus erectis distinguenda.

F,. primaria Corner sp. nov.—Arbor —20 m. alta: cortice griseo-

brunneo pustulato, 12 mm. crasso: latice albido serifluo; foltis

spiraliter dispositis. Ramuli, petioli, et costae (subtus) pilis rigi-

dis patentibus albidis 1-2 mm. longis hispidi, nervuli pilis bre-

vioribus v. puberuli: lamina supra hispido-scabrida. Ramuli 5—6

mm. crassi, brunnei. Stipulae 5-10 mm. longae, puberulae,

caducae. Lamina 17—24 x 8-11 cm., elliptico-obovata, breviter

attenuato-acuminata, basi rotundato-subcordata, denticulata,

membranacea, utrimque scabrido-hispidula, sicco brunneo-viri-

dis: costis lateralibus utrinsecus 9-11, intercostis strictis —11,

subtus valde elevatis: costis basalibus utrinsecus 2—3, ad 4 lami-

nae elongatis, glandulis basalibus 2: petiolo 36-65 mm. longo.

Receptacula cauliflora ad ramulos breves congestos 8-14 mm.

crassos, internodis vix evolutis: pedicello -55 mm. longo, spar-

sim hispidulo, bracteis lateralibus 3, 1.5 mm. longis, circum

medium pedicelli dispersis, haud ternatis: corpore receptaculi

26 & 22 mm. (immaturo), pyriformi, bracteis lateralibus nullis,

scabrido-hispidulo pilis —-0.3 mm. longis, ostiolo c. 5 mm. lato

bracteis apicalibus parvis numerosis erectis v. recurvatis, occ-

luso: setis internis brevibus copiosis albidis: cellulis scleroticis

nullis. Flores feminei (juveniles) sessiles v. pedicellati: tepala

457

Gardens Bulletin, S.

4-5 libera glabra oblongo-spathulata, pallide carnea: ovario ses-

sili albido: stylo valde laterali, glabro v. sub stigmate clavato

piloso. Cystolitha hypogena. Terr. New Guinea (Hoogland 4958,

Madang district, Kokun River, pr. Jal, Gogol River Valley:

typus: herb. Lae).

Species nobilis ut F. montana gigantea, sed ob flores masculos

incognitos sedi incertis, forsan F. griseifoliae Corner affinis.

subsect. Sycidium ser. Scabrae Miq.

F. cumingii Miq. var. androbrota (Summerh.) Corner comb. nov.-—

F.. androbrota Summerth. J. Arn. Arb. 10 (1929) 143.—F. dich-

roa Summerh. id. 147.—This New Guinea variety differs from

v. cumingii merely in the broadly elliptic lamina, 4-8 cm. wide,

the numerous intercostals (3-11), the glabrous flower-pedicels,

and the presence of a gall-ovary in the stalked male flowers ad-

joining the gall-flowers (not in the sessile male flowers round

the orifice). Thus it approaches v. worcesteri with larger lamina.

Some Philippine collections of v. terminalifolia (Elm.) Sata

(=F. terminalifolia Elm). match those of v. androbrota in leaf

and glabrous flower-pedicels: thus, it may not be possible to

distinguish seed-plants.

Aet (Exp. Lundquist) 520; Branderhorst 53, 128 (det. F.

dichora); Brass 800 (F. androbrota, type), 1190 (F. dichroa,

type), 8298, 8548; Carr 11208; Koch 4.

var. angustissima (Merr.) Corner v. nov.—F. angustissima

Merr. Govt. Lab. Publ. Philip. 29 (1905) 11.—This differs from

Vv. cumingii in the linear acuminate lamina (14—32 x 0.5-—1.5

cm.) with 35-60 pairs of lateral nerves. It appears to be a shrub

—3 m. high, but whether a sapling-form, a coppiced form, or a

distinct variety is uncertain: a few collections are intermediate

with v. cumingii. It is recorded from Luzon and Mindanao.

Merrill 2696 (type); Elmer 9357 (not seen), 13920; PNH

11067; Herb. Univ. Philip. 4907; Vidal 3818.

var. worcesteri (Merr.) Corner comb. nov.—F. worcester

Merr. Philip. J. Sci. Bot. 9 (1914) 274.—This differs from v.

terminalifolia (Elm.) Sata in the larger lamina (13-37 x 5-17

cm.; 4-17 « 1.5—6 cm. in v. terminalifolia) and the glabrous

flower-pedicels. It lacks the gall-ovary in all the male flowers.

That it is not a large-leafed sapling is shown by its sporadic distri-

bution and the fact that small-leafed saplings of v. terminalifolia

are known. Some collections, including the type, have pale pink

tepals, which suggests that v. worcesteri links with F. tonsa Miq.

458

Vol. XVI. (1959).

Bur. Sci. 7178 (Cavilli Isl. Sulu Sea; type); 24309, 24513

(Samar, Catubig River); 31029, 31067, 31369 (Panay, Capiz

province, Jamindan), PNH 14315 (Samar, Mt. Mahagua).

F. fiskei Elmer var. multinervia Corner v. nov.—Sparsim muri-

culata. Lamina scabrida, basi utrimque cordata sed asymmetrica:

costis lateralibus utrinsecus 8—11, costis basalibus haud elongatis.

Tepala subrubra. Mindanao (Bur. Sci. 49506, Davao province,

Mt. Mayo, det. F. hemicardia Merr., typus, herb. Bogor).

In v. fiskei, v. cebuensis Merr., et v. laevifolia Merr. costae late-

rales utrinsecus 5—8, basales ad 1/3-—4 laminae elongatae. Var.

multinervia foliis F. oleraceae Corner (Solomon Isl.) simillima,

sed pilis muriculatis et setis internis deficientibus F. fiskei constat.

F. riedelii Mig. var. minor Corner v. nov.—Receptacula minora

6—8 mm. lata, pilis conicis —0.5 mm. longis vestita, bracteis late-

ralibus et apicalibus minoribus 1—1.5 mm. longis, pedicello 0—2

mm. longo: setis internis paucis brevibus. Lamina minor v. an-

gustior. Frutex v. arbor parva. Celebes.

Beccari s.n., Kandari (R.Ist. Fir. 9346, det. F. obscura Bl. by

King); Eyma 413, subdiv. Enrekang, inter Pasoei et Rante Lomo,

600—1,300 m. alt. Eyma 3408, Menado, subdiv. Kolone. Dale, inter

Koroworo et Tompira (typus, herb. Leiden); Elbert 3329 (Ka-

baena Isl.), 3048 (distr. Rumbia, pr. Zankaya), S.E. Celebes;

Kjellberg 357, 361 743, Kendari; 1385, Enrekang; 3779, Malili.

F. ampelas Burm. f. Fl. Ind. (1768) 226; emend. Mig. Hook. Lond.

J. Bot. 7 (1848) 428 (excl. syn. Rheede).—There are two

sheets in herb. Delessert (Geneva) named in Burman’s hand F.

ampelas. One, which I will call A, is the usual interpretation as

given by Miquel, King, and Valeton. The other, which I will

call B, is F. montana Burm. f. (= F. quercifolia Roxb.),

though a leaf of F. ampelas is also attached to B. Both have

been labelled “F. ampelas Burm. type” by Hochreutiner.

A. This sheet I have selected as the type because it maintains

current usage and because it has written on it the full citation given

by Burman, that is:—

Folium politorium.

Varinga Rumph. Herb. lib. 6 cap. 69 T63.

Terega H. Mal. tom. 3 p. 79 tab. 60.

The reference to Terega is to F. montana Burm. f. and was ex-

cluded by Miquel (1848).

459

Gardens Bulletin, S.

B. This sheet bears the single erroneous citation “Rumph 4 p.

128 tab. 61”, which should read “tab. 63”. The specimen is typical

F. montana with reticulate seed, and is to be excluded from F.

ampelas.

The name was published as F. ampelos, but this is clearly a

misprint for Burman wrote F. ampelas unmistakably on both

sheets, and it is etymologically wrong. Miquel and King used the

correct F. ampelas.

var. linearis Corner v. nov.—Frutex —2 m. altus. Lamina 3—-6.5

x 0.5—0.9 cm., lineari-oblonga, subacuta, rigide coriacea, scabri-

dissima: costis lateralibus utrinsecus 10-20, intercostis nullis: cos-

tis basalibus brevibus: petiolo 2-5 mm. longo. Receptacula 4 mm.

lata: pedicello 2-3 mm. longo, bracteis lateralibus 2—3 dispersis.

Stomata immersa. Celebes (Kjellberg) 3773, Waroe; typus, herb.

Stockholm ).

F. fallax Mig. (Celebes) revocans, sed setis internis, tepalis

puberulis F. ampelas constat.

var. soronensis (King) Corner comb. nov.—F. exasperata Roxb.

Fl. Ind. 3 (1832) 555.—F. biglandulosa Mig. Hook. Lond. J.

Bot. 7 (1848) 229 (haud FI. Ind. Bat. 1, 2, 1859, 298).—F.

asperior Miq. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 291.—F. soro-

nensis King, J. As. Soc. Beng. 55, 2 (1887) 411.—F. blephari-

sepala Warb. in Laut. et K. Schum. Nachtr. Fl. Schutzgeb. (1905)

246.—Tepala ciliato-puberula: florum pedicelli saepe hispiduli.

Corpus receptaculi saepe bracteis lateralibus 1—2 praeditum.

Lamina scabrida v. sublaevis, saepe subsymmetrica: costis laterali-

bus utrinsecus 6-9. Moluccas, Key Isl., Aru, New Guinea, New

Britain.

Varietas orientalis plurinervia, in Moluccis specimina inter-

media.

F. guyeri Elmer var. sibuyanensis (Elmer) Corner comb. nov.—

F. sibuyanensis Elmer, Leafl. Philip. Bot. 4 (1911) 1319.—

Lamina 5-15 x 3-8 cm., elliptica v. ovato-elliptica, apice acu-

minata 8—24 mm. longo v. subcaudata, laevis v. scabrida: costis

lateralibus utrinsecus 3—6: petiolo 3-10 mm. longo. Pedicelli

receptaculi 3—5 mm. longi, v. 15-18 mm. Luzon, Sibuyan,

Panay, Samar.

Elmer 12236 (typus); Bur. Sci. 14509, 22811, 30846, 32284,

47345.

Ob setas internas longas F. guyeri ascripta, sed lamina ut in F.

ampelas Burm. f.

460

Vol. XVII. (1959). '

F. cauta Corner sp. nov.—Arbor —15 m. alta, ? latice subnullo,

foliis distichis, glabra. Ramuli 1.5—2 mm. crassi, straminei dein

brunneoli. Stipulae 6-15 mm. longae, graciles. Lamina 8—20

3.5—7.5 cm., obovato-elliptica, apice acuminata —12 mm. longo,

basi cuneata, integra v. dentato-angulata, membranacea v. sub-

coriacea, laevis v. subtus subscabrida, sicco griseo-viridis: costis

lateralibus utrinsecus 3—5 (—6), curvato-ascendentibus, intercos-

tis 1-4 laxis: costis basalibus utrinsecus 1, ad 1/3—4 laminae

elongatis, glandulis basalibus 2 exiguis: petiolo 6-20 mm. longo.

Receptacula axillaria solitaria: pedunculo 1-8 mm. longo: brac-

teis basalibus in collare ternatis, parvis, caducis: pedicello 2-15

mm. longo: corpore receptaculi 13-17 mm. lato (immaturo),

bracteis lateralibus nullis, ostiolo plano bracteis apicalibus parvis

numerosis occluso: setis internis et cellulis scleroticis nullis.

Tepala 3—4 albida libera glabra, lanceolata v. oblonga. Flores

masculi ostiolares, ordinibus duobus instructi, sessiles v. pedi-

cellati: stamine 1, pistillodio minuto v. nullo. Flores feminei et

cecidiophori sessiles v. pedicellis —1.5 mm. longis praediti:

ovario sessili, pallido, stylo glabro. Semina 1—1.2 mm. longa,

lentiformia, valde carinata, laevia. Cystolitha hypogena. Celebes,

in silvis —1,200 m. alt.

Bunnemeyer 11477, Lombasang (typus, herb. Leiden); NIFS

bb. 13517, Manado, Kakaskassen (det. F. rostrata Lam.); Forman

350, G. Lokon, Tetepamgan, Minahassa; Kjellberg 2602 (Ko-

sali Porema), 2691 (Porema); Koorders 19255, 19348, Minahassa,

Menado; Reinwardt 1563, Minahassa; Sarasin 603 (det. F. pau-

cinervosa Warb. ms., non Merrill); Teysmann 14022, Bonthain.

Lamina ut in F. uniglandulosa Wall. sed cellulis fibrosiformis

deficientibus, habitu et receptaculis toto dissimilis. Affinitas incerta,

forsan F. ulmifoliae Lam.

F. goniophylla Corner sp. nov.—Arbor —10 m. alta, foliis brevi-

petiolatis distichis. Ramuli, petioli, et nervi pilis conicis —0.3

mm. longis sparsim hispiduli. Ramuli 1.5 mm. crassi, fusco-brun-

nei. Stipulae 2-3 mm. longae, perbreves, caducae. Lamina 3—9

< 1.7—4 cm., anguste elliptica, inequilateralis, utroque lobis bre-

vibus 1—2 subangulatis, apice attenuata, basi cuneata, mem-

branacea, utrimque scabrida, sicco griseo-viridis: costis laterali-

bus utrinsecus 3-4(—5), rectangulatis, subtus _ elevatis,

intercostis nullis: costis basalibus utrinsecus 1, ad 4-1/3

laminae elongatis, glandulis basalibus 2: petiolo 2-4 mm. longo.

Receptacula cauliflora ad ramulos breves congestos efoliatos,

pilis erectis 0.5 mm. longis hispidula, maturitate brunnea intus

461

Gardens Bulletin, S.

rubescentia: pedicello 20-30 mm. longo, sursum gradatim dila-

tato, bracteis lateralibus 2—3 parvis dispersis: corpore recepta-

culi 20 mm. lato, obconico, bracteis lateralibus parvis 1-2 dis-

persis, ad apicem receptaculi applanatum numerosis umbonatis

confertis 2—~3 mm. longis, appressis v. incurvatis, ostiolo parvo

plano bracteis apicalibus minoribus erectis occluso: setis internis

brevibus sparsis albidis: cellulis scleroticis nullis. Tepala 4 ob-

longa, raro bifida, libera, glabra, albida. Florum pedicelli longi,

glabri, in cecidiophoris 1.5—4.5 mm. Flores masculi ordinibus

2—3 instructi: stamine 1, pistillodio nullo. Ovarium sessile albi-

dum. Cystolitha hypogena. Celebes (Kjellberg 1925, Todjam-

boe, c. 1,200 m. alt., in silvis: typus, herb. Stockholm).

Species insignis, foliis F. tinctoriam Forst. f. subsp. gibbosam

(BI.) Corner revocantibus, receptaculis F. gul Laut. et K. Schum.,

sed tepala albida. Planta feminea inquirenda.

F. melinocarpa BI. var. hololampra (Diels) Corner comb. nov.—

F. hololampra Diels, Engl. Bot. Jahrb. 67 (1935) 201.—F. col-

linsii Elmer, Leafl. Philip. Bot. 9 (1937) 3468.—Glabra sed

scabridula v. foliis supra nitidis et laevibus. Receptacula setis

internis praedita.

This is merely the glabrous state of F. melinocarpa, co-extensive

from Sumatra to the Solomon Islands, but more abundant, ap-

parently, in New Guinea. Intermediates with varying indumentum

Occur.

F. trachypison Laut. et K. Schum. var. pallida Corner v. nov.—

Frutex v. arbor —15 m, alta, foliis distichis. Ramuli, petioli, et

receptacula scabridi, haud v. vix hispiduli. Lamina tenuior,

sicco pallidior griseo-viridis, intercostis subtus vix elevatis. Te-

pala albida, apice hirta. Terr. New Guinea, Papua, Solomon

Isl. (Bougainville).

Ramuli 1-2 mm. crassi, brunneo-badii. Stipulae parvae caducae.

Lamina 4-18 * 2.5—8.5 cm., elliptica, apice acuto attenuata Vv.

subacuminata, basi late cuneata v. uno latere subcordata, sub-

symmetrica, denticulata v. integra, utrimque scabrida: costis late-

ralibus utrinsecus 5—9, intercostis —9: costis basalibus utrinsecus

1-2, ad 3-1/3 laminae elongatis, glandulis basalibus 2: petiolo

4-12 mm. longo. Receptacula axillaria et ad ramulos efoliatos

fasciculata: pedunculo 2—5 mm. longo: bracteis basalibus ternatis

0.5 —1 mm. longis: corpore receptaculi 6-9 mm. lato, globoso,

bracteis lateralibus nullis: setis internis albidis copiosis brevibus:

cellulis scleroticis nullis. Semina reticulata.

Brass 25346, Normanby Isl., Waikaiuna (typus, Gray herb.

Harvard); A. W. Herre 223, Huon Gulf; Hoogland 3828, Terr.

462

Vol. XVIT. (1959).

New Guinea, north division, pr. Dobodura; Hoogland 4240, Terr.

N.G., north division, Tufi subdistrict, pr. Guragura; NGF 5636,

Lae; J. R. Saunders 39, Terr. N.G., north division, Tufi subdis-

trict, pr. Oi-ai on Penari-track; J. H. L. Waterhouse 32, Bougain-

ville, Siwai.

Varietas incertae sedis propter affinitatem diversam, vix species

ipsa. Facies ut F. ampelas Burm. f. var. soronensis (King) Corner

sed differt in seminibus reticulatis, bracteis pedunculi basalibus

ternatis, cellulis scleroticis in pariete receptaculi nullis, intercostis

numerosis, qui sunt F. melinocarpae Bl. et F. trachypison proprii.

Lamina parva et receptaculum parvum ut F. trachypison, sed eo

F. storckii Seem. (Fiji) revocans. Plantae vivae juvenes et in statu

cauliflori inspiciendae.

F. tonsa Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 234, 297.—

Celebes, Talaud Isl.

Riedel s.n., Menado (HB 5705, typus); Boschpr. st. Cel/II—

405, Malili, Kawata; Koorders 19057, 19063, 19193, 19247, 19274,

Minahassa; Sarasin 588, Minahassa, Tomohon; Teysmann 11976

(Pangkadjena), 12648, 12737 (Balek Angin).

Lam 2627, Talaud, Karakelang, east of Beo; 3125, Talaud,

Salibabu, east slope of G. Ajambana. -

This is a large, but little known, tree relating F. todayensis

Elmer and F. irisana Elmer of the Philippines with F. leptoclada

Benth. of Queensland, through F. tonsa v. leptodictya of New

Guinea. They may, indeed, be varieties of one species, but v. tonsa

is characterised by sunken stomate.

var. aspera Corner v. nov.—Lamina 5-16 2-5.5 cm.,

utrimque scaberrima, anguste elliptica, breviter acuminata, basi

subsymmetrica, uno latere cuneata altero subrotundata, apicem

versus denticulata: costis lateralibus utrinsecus 5—8, intercostis

strictis —8: costis basalibus 1 + 2, ad 4-24 laminae elongatis,

glandulis basalibus 2: petiolo 3-8 mm. longo. Receptacula minora,

8—10 mm. lata, scabra, bracteis lateralibus nullis: pedunculo 1.5—

2.5 mm. longo: setis internis —1 mm. longis, copiosis albidis: cellulis

scleroticis in pariete receptaculi interno copiosis. Tepala 4—5 rubra

glabra. Cystolitha amphigena: stomata superficialia. Terr. New

Guinea (NGF 4863, East Highlands, pr. Nondugl, arbor parva in

querceto: typus, herb. Lae).

Ut forma F. trachypison Laut. et K. Schum. angustifolia, sed

setis internis longis et cellulis scleroticis copiosis, bracteis lateralibus

nullis ad F. tonsam pertinens.

var. leptodictya (Diels) Corner comb. nov.—F. leptodictya

Diels, Engl. Bot. Jahrb. 67 (1935) 196.—Costae basales ad 1

laminae elongatae. Stomata superficialia. New Guinea.

463

Gardens Bulletin, S.

Brass 27291, Fergusson Isl., Agamoia; Carr 14799 (Boridi).

16059 (Isuarava): Eyma 4355, 4534, Wissel Lake: Hoogland 3914,

Kokoda; Ledermann 12826, Sepik (typus); NGF 4183. Sogeri.

A var. tonsa vix distincta sed in regione novoguineensi et ad

F. wassa Roxb. v. nubigena (Diels) Corner approximans.

var. subcordata Corner v. nov.—Folia applanata disticha.

Lamina 5-13 « 1.5—4.5 cm., anguste elliptica v. ovato-lanceolata,

acuta v. subacuminata, basi rotundata v. subcordata, symmetrica,

subscabrida: costis lateralibus utrinsecus 7—10, intercostis 2-4:

costis basalibus utrinsecus 2, brevibus: petiolo 3-10 mm. Recep-

tacula axillaria, solitaria, glabra, 9-10 mm. lata, bracteis laterali-

bus nullis: pedunculo 7-15 1 mm.: bracteis basalibus ternatis,

1 mm. longis: pedicello 1.5—-8 mm. longo: setis internis brevibus

copiosis: cellulis scleroticis nullis. Flores feminei sessiles v. pedi-

cellis rubris glabris praediti: tepalis 4-5 fusco-rubris, apice his-

pidulis: stylo puberulo. Semina | mm. longa, lentiformia, sub-

carinata, laevia. Stomata superficialia. Solomon Isl. (Kajewski

1657, Bougainville, Kupei Gold Field, in silvis 950 m. alt., arbor

communis —10 m. alt.; typus, herb. Br. Mus.).

A var. leptodictya lamina anguste elliptica, basi symmetrica

subcordata, receptaculo minori, cellulis scleroticis deficentibus

differt. Ad F. leptocladam Benth. (Queensland) approximans.

F. irisana Elmer var. validicaudata (Merr.) Corner comb. nov.—

F. guyeri Elmer v. validicaudata (Merr.) Sata, Monogr. (1944)

290.—? v. minimaefolia Sata id.—F. fastigiata Elmer, leafl.

Philip. Bot. 1 (1906, April 10) 44.—F. validicaudata Mert.

Philip. J. Sci. 1 (1906, April 15) Suppl. 45.—This is a dwarf

state of F. irisana with small leaves and figs, occurring on high

mountains and exposed places: it may well be phenotypic. The

red tepals and lack of internal bristles show that it does not

belong with F. guyeri Elm.

Clemens 17386; Bur. Sci. 2571, 12644, 27032, 37487, 38099,

42157; Elmer 6072. 8001 (F. fastigiata, typus), 8555, 8577, 8710.

14284, 21944, 21999; For. Bur. 936, 5084, 18140, 20125; PNH

1879, 7692; Vanoverbergh 1033; Whitford 1201 (F. validicaudata,

typus).

F. tenuicuspidata Corner sp. nov.—Arbor tenuiramosa, foliis brevi-

petiolatis distichis, glabra, praeter receptacula non scabrida.

Ramuli 1—-1.5 mm. crassi, brunnei. Stipulae —8 mm. longae,

binatae, caducae. Lamina 7.5—11 « 2-3.5 cm., elliptica, apice

valde caudato-acuminata 25—38 mm. longo, (1/3—4 laminae ef-

ficienti) basi cuneata, integra, laevis, sicco griseo-viridis: costis

lateralibus utrinsecus 6—8, intercostis —3, subtus leviter elevatis:

464

Vol. XVI. (1959).

costis basalibus utrinsecus 1, brevibus, glandulis basalibus 2:

petiolo 1.5—4 mm. longo. Receptacula axillaria, ? solitaria, sca-

bridula: pedunculo 2—3 mm. longo: bracteis basalibus ternatis,

0.5 mm. longis: pedicello 2-3 mm. longo: corpore receptaculi

8—11 mm. lato, bracteis lateralibus nullis, ostiolo parum dep-

resso: setis internis copiosis minutis: cellulis scleroticis copiosis.

Flores feminei sessiles v. pedicellis rubris, glabris v. hispidulis,

—1 mm. longis praediti: tepalis 4-5 fusco-rubris liberis hispidu-

lis: ovario sessili rubro, lateribus pallidis: stylo glabro. Semina

1 mm. longa, lentiformia subcarinata, laevia. Cystolitha amphi-

gena, papillata.

v. tenuicuspidata.—Celebes.

. Boschpr. st. Cel 1/21, Bonthain; Biinnemeyer 12641, Bonthain,

1400 m. alt.: Elbert 2730, Buton Isl.; Eyma 1013, subdiv. Enre-

kang pr. Latimodjong (typus, herb, Leiden).

F. virgatae Reinw. simillima sed bracteis basalibus ternatis,

seminibus lentiformibus, cystolithis papillatis, rectius prope F.

tonsam Miq. et F. irisanam Elmer referenda.

v. major Corner var. nov.—Lamina —22 8 cm., apice 25-35

mm. longo, intercostis 4-7. Receptacula axillaria, solitaria v. fasci-

culato-ramiflora, glabra, majora, 12—15 mm. lata: pedunculo lon-

giori 10-17 mm.; pedicello nullo. Tepala 3—4 glabra. Flores masculi

sessiles v. rubro-pedicellati: stamine 1, haud mucronato, ad basim

setulis circumdato, pistillodio nullo. Flores cecidiophori sessiles v.

pedicellis glabris rubris —2.5 mm. longis praediti: ovario sessili

albido. Flores feminei sessiles v. breviter pedicellati: ovario albido:

stylo glabro rubro. Semina 1 mm. longa, lentiformia, subcarinata,

laevia. Mindanao (Bukidnon province, Mahilucot River; Bur. Sci.

38635, typus, herb. Leiden; 38683).

F. cumingii Miq. v. worcesteri (Merr.) Corner similis sed tepalis

rubris rectius inter F. irisanam et F. tonsam ascribenda.

F. elmeri Merr. Govt. Lab. Publ. Philip. 29 (1905) 9.—

F.. semicordata Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 226,

293 (haud J. E. Smith).—This synonym must be re-instated as

the correct name for F. semicordata Mia. in view of the oversight

of F. semicordata B. Ham ex J. E. Sm. (= F. cunia B. Ham. ex

‘Roxb. ).

F. oleracea Corner sp. nov.—Arbor —20 m. alta, foliis distichis,

glabra v. ramulis petiolisque pilis erectis pallidis mollibus, 2—3

mm. longis, laxe pubescentibus. Ramuli 3—5 mm. crassi, brun-

neoli. Stipulae 6-15 mm. longae, binatae, lanceolatae, glabrae,

novellis subpersistentes. Lamina 16-35 « 5-12 cm. (-47 X

465

Gardens Bulletin, S.

15 cm., juvenilis), oblonga v. ovato-oblonga, apice breviter acu-

minata 10-20 mm. longo, basi plus minus valde cordata, sym-

metrica v. asymmetrica, uno latere late lobata et petiolum obte-

genti, integra, coriacea, laevis v. subtus subscabrida, sicco

griseo-viridis v. brunnea: costis lateralibus utrinsecus (7—)

9-12, (—16, foliis juvenilibus), divaricatis, intercostis 5—10,

subtus leviter elevatis: costis basalibus utrinsecus 3—4, brevibus,

glandulis basalibus 2: petiolo 5-12 « 2—4 mm., brevi, crasso.

Receptacula axillaria, ramiflora et cauliflora, ad truncum in

cumulis grandibus evoluta, maturitate e cremeo rubescentia:

pedunculo 3—10 mm. longo: bracteis basalibus in collare ter-

natis, 1 mm. longis: pedicello O-8 mm. longo: corpore recepta-

culi 15-32 mm. lato (25—48 mm., vivo), subgloboso, bracteis

lateralibus nullis, ostiolo 2—4 mm. lato bracteis apicalibus 3—4

Vix projicientibus occluso: setis internis —0.6 mm. longis, albidis,

copiosis: pariete 1-2 mm. crasso, cellulis scleroticis nullis.

Tepala 4—6, rubro-carnea dein pallida, libera, glabra v. pube-

rula, lineari-oblonga, ovarium superantia. Flores masculi ostio-

lares, ordinibus 1—2 instructi, pedicellati: stamine 1, pistillodio

nullo. Flores feminei et cecidiophori sessiles v. pedicellis glabris

albidis —4.5 mm. longis praediti: ovario sessili, pallido: stylo

femineo valde laterali, glabro v. puberulo, stigmate simplici.

Semina 1.2 mm. longa, lentiformia, vix carinata, minute reticu-

lata. Cystolitha amphigena. Solomon Islands.

var. oOleracea.--Kajewski 1721 (typus, herb. Kew), 1912,

Bougainville, Kupei Gold Field et Buin; J. H. L. Waterhouse 133

(New Georgia), 124, B203, 779 (Bougainville, Siwai), 203, 847

(Bougainville, Maisua); ? R. B. Comins 174 (San Cristoval).

var. pugnans Corner y. nov.—Ramiuli, petioli, costa (subtus), et

receptacula pilis erectis pallidis, flavidis v. brunneolis, 0.5 mm.

longis villosuli, ad nervulos brevioribus. Lamina supra scaberrima,

subtus villosula. Kajewski 1941, Bougainville, Buin, Kugumaru

(typus, herb. Kew).

Species F. fiskei Elm. revocans sed arbor altior, receptaculis

majoribus, setis internis praeditis, costis curvato-ascendentibus, in-

tercostis numerosis. F. tonsae Miq. et F. elmeri Merr. affinis sed

foliis majoribus brevipetiolatis, receptaculis majoribus, habitu cau-

liflori spectabilis.

F. chrysochaete Corner sp. nov—Arbor mediocris —10 m. alta,

v. suffruticosa, ramis horizontalibus, foliis distichis et ad ramu-

los flabelliforme dispositis. Ramuli et petioli pilis albidis, aureis,

v. brunneolis mollibus erectis, 1-2 mm. longis, villosi: costae

466

Vol. XVII. (1959).

subtus sparsim et breviter puberulae, nervuli glabri v. scabriduli:

lamina supra sparsim et appresse pilosa, glabrescens, costis

subvillosis. Ramuli 1-2 mm. crassi, brunneoli, saepe alternis an-

fracti. Stipulae -12 mm. longae, binatae, novellis subpersistenti-

bus, albido-pilosae. Lamina 11-25 »< 3-8 cm., anguste

oblongo-elliptica v. subovata, apice acuminata 7-20 mm. longo,

basi leniter cordata symmetrica v. subasymmetrica, latere acros-

copico latiori, denticulata v. integra, chartaceo-subcoriacea,

supra nitida et scaberrima, subtus hispidulo-velutinosa, sicco

griseo-viridis v. olivacea: costis lateralibus utrinsecus 6-9

(—10), obliquis, intercostis 5-14, subtus elevatis: costis basali-

bus utrinsecus 3—4, ad 4—+ laminae elongatis, glandulis basali-

bus 2: petiolo 5-10 & 1.5—2 mm. brevi. Receptacula axillaria,

solitaria, pilis erectis 0.5 mm. longis hispidula, glabrescentia,

scabridula, maturitate rubra: pedunculo 5—1G mm. longo, gra-

cili: bracteis basalibus in collare ternatis, 1-1.5 mm. longis,

ovato-acutis, appresse puberulis: pedicello O—-7 mm. longo:

corpore receptaculi 8-10 mm. lato (12-15 mm., vivo), subglo-

boso, bracteis lateralibus nullis, ostiolo bracteis apicalibus

parvis numerosis occluso: setis internis —1 mm. longis, albidis

v. aureis, copiosis: cellulis scleroticis nullis. Tepala 5—6 albida

libera, spathulato-oblonga, ovarium superantia, appresse hispi-

dissima, stamine et ovario setulis circumdatis. Flores masculi

ostiolares, ordinibus duobus instructi, sessiles v. pedicellati:

stamine 1 (—2), pistillodio nullo. Flores feminei et cecidiophori

sessiles vel pedicellis glabris -2 mm. longis praediti: ovario ses-

sili, albido: stylo femineo puberulo. Semina 0.8 mm. longa, lenti-

formia, subcarinata, tenuiter reticulata. Cystolitha amphigena:

pili laeves. Solomon Islands.

L. J. Brass 2960 (Ulawa). 3199 (Ysabel, Maringe, typus herb.

Kew); Kajewski 1983, 2148 (Bougainville, Buin, Koniguru et

Kugumaru), 2317 (Malaita); J. H. L. Waterhouse 201 (New Geor-

gia), 232, 232a (Bougainville, Siwai).

Species lamina oblonga brevipetiolata, costis obliquis, intercostis

numerosis strictis, aureopilosa facile distinguenda, inter F. melino-

carpam Bl. et F. fulvopilosam Summerh. sed habitu humili fron-

doso-flabellitormi dissimilis——F. imbricatae Corner comparanda.

F. imbricata Corner sp. nov.—Arbor —20 m. alta, ramulis divari-

catis, foliis distichis. Ramuli, petioli, et costae (subtus) pilis erec-

tis albidis v. flavidis 1-2 mm. longis plus minus villosi, nervuli

subtus sparsim et breviter pilosi v. subglabri: lamina supra spar-

sim et breviter pilosa, glabrescens. Ramuli 1-2 mm. crassi,

467

Gardens Bulletin, S.

brunneoli. Stipulae —10 mm. longae, binatae, lanceolatae, spar-

sim pilosae, caducae. Lamina 10-18 x 3.5—7.7 cm., elliptica,

inaequilateralis, apice acuminata —17 mm. longo, basi rotun-

data v. late cuneata et asymmetrica, saepe latere acroscopico

auriculato-lobulato, denticulata v. integra, supra scabrida, sub-

tus scabrido-pilosa, rigide membranacea, sicco griseo-viridis:

costis lateralibus utrinsecus 4—7, obliquis curvatis, —10 inter-

costis, subtus elevatis: costis basalibus 2 + 3, ad 4-1/3 laminae

elongatis, glandulis basalibus 2: petiolo 3-8 mm., brevi. Recep-

tacula axillaria solitaria (? etiam ramiflora), pilis brunneolis

rigidis —O0.5 mm. longis hispidula, subglabrescentia: pedunculo

5—18 1 mm.: bracteis basalibus in collare ternatis, 1.5—2 mm.

longis, lanceolatis: pedicello 4-15 mm. longo, gracili: corpore

receptaculi 10-12 mm. lato (-25 mm., vivo, ficiformi, apice

applanato), subgloboso, bracteis lateralibus nullis, apicem ver-

sus plus minus contracto, ostiolo bracteis apicalibus parvis

numerosis subprojicientibus occluso: setis internis —0.5 mm.

longis, albidis, sparsis: cellulis scleroticis nullis. Tepala 4—6 fus-

co-rubra, albescentia in receptaculis cecidiophoris, libera, lan-

ceolato-oblonga, ovarium multo superantia, apicem versus

hispidula. Flores masculi ostiolares, ordinibus 2-3 instructi,

sessiles v. pedicellati: stamine 1, pistillodio nullo. Flores feminei

et cecidiophori sessiles v. breviter pedicellati: ovario sessili, pal-

lido: stylo glabro. Semina 0.7—0.8 mm. longa, lentiformia, sub-

carinata, subreticulata. Cystolitha hypogena: pili laeves. Solomon

Islands (Bougainville).

Kajewski 1724, Kupoi village; J. H. L. Waterhouse B37 (Buka,

Skotolan), 176, 818 (Siwai, typus 818 herb. Kew).

F. chrysochaete Corner affinis sed arbor major, receptaculo ma-

jori, setis internis paucis, tepalis rubris, laminae basi plus minus

asymmetrico, costis lateralibus paucioribus. Auricula ad basim

laminae acroscopica forsan ad plantas juveniles pertinens quod

foliis in Kajewski 1724 (arbor alta) deficiens.

F. gryllus Corner sp. nov.—Arbor parva, foliis asymmetricis dis-

tichis. Ramuli, petioli, stipulae, receptacula, et costae (subtus )

pilis albidis v. brunneis rigidis erectis 1-2 mm. longis hispido-

scabridi: nervuli subtus pilis brevioribus: lamina supra pilis

albidis erectis —1 mm. longis basi pustulato hispido-scabrida.

Ramuli 5—6 mm. crassi, brunneoli, cavi. Stipulae 20-30 mm.

longae, caducae. Lamina -35 16 cm., ovata, asymmetrica,

apice acuminata 30-40 mm. longo, base valde asymmetrica, uno

468

Vol. XVII. (1959).

latere late et profunde cordata, altero cuneato-subcordata, den-

tata, membranacea, utrimque scabrida, sicco griseo-viridis: cos-

tis lateralibus utrinsecus 7—8, intercostis —12, subtus elevatis:

costis basalibus 2 +4, ad 4 laminae elongatis, glandulis basali-

bus 2: petiolo 10-40 3 mm., sub auricula laminae condito.

Receptacula axillaria solitaria hispidissima, maturitate auran-

tiaca: pedunculo 4-6 2.5 mm.: bracteis basalibus in collare

ternatis, 3—4 mm. longis, ovato-lanceolatis, acutis, hispidis:

corpore receptaculi 15 & 12 mm. (pilis exclusis), ellipsoideo,

bracteis lateralibus 2—4 mm. longis sparsis, circum ostiolum

pluribus confertis erectis hispidis 2-3 mm. longis: setis internis

albidis —1.5 mm. longis, copiosis: cellulis scleroticis nullis (re-

ceptaculo femineo). Flores feminei sessiles vel pedicellis glabris

—1.5 mm. longis praediti: tepala 5—6 albida spathulata libera,

apicem versus hispida: ovario sessili albido, setulis paucis cir-

cumdato: stylo sparsim puberulo. Semina 0.8-0.9 mm. longa,

lentiformia v. breviter oblonga, subcarinata, laevia. Cystolitha

hypogena: pili papillati. Solomon Islands (L. J. Brass 3407,

Ysabel, Tiratona, in silvis c. 600 m. alt., typus herb. Kew).

Receptaculo horrido, lamina valde asymmetrica, species hispi-

dissima facile distinguenda, F. erinobotryae Corner affinis.

F. erinobotrya Corner sp. nov.—Suffrutex v. arbor parva, foliis

asymmetricis distichis. Ramuli, petioli, laminae, et receptacula

pilis albidis v. brunneis erectis rigidis 1~2 mm. longis hispidi:

nervuli subtus pilis brevioribus. Ramuli 2—3 mm. crassi, fusco-

brunnei. Stipulae —12 mm. longae, pilosae caducae. Lamina 13

—32 < 5.5-14 cm., ovato-elliptica, apice acuminata —17 mm.

longo, basi valde asymmetrica, uno latere cordata petiolum ob-

tegenti, altero anguste subcuneato, denticulata v. subintegra,

membranacea, supra scaberrima, subtus hispido-villosula, sicco

griseo-viridis: costis lateralibus utrinsecus 7-11, inferioribus

saepe glandulo axillari praeditis, -11 (—14) intercostis regula-

riter instructis, subtus elevatis: costis basalibus 3 + 5, ad 1/3-4

laminae elongatis, glandulis basalibus 2. petiolo 7-10 mm.

longo. Receptacula axillaria solitaria, dein cauliflora ad ramu-

los efoliatos -12 cm. longis paniculatos, basi 5-20 mm., apice

2~3 mm. crassos, internodis vix evolutis, hispidissima, maturi-

tate rubra: pedicello 7~9 mm. longo, bracteis lateralibus 1-2

dispersis parvis, haud in collare confertis: corpore receptaculi

7-10 mm. lato, subgloboso, bracteis lateralibus 0-2, apicalibus

numerosis parvis vix projicientibus: setis internis -1 mm. longis,

albidis v. brunneolis, copiosis: cellulis scleroticis nullis. Tepala

5—7 (vel 3—5 in floribus masculis) subrubra v. albida, lanceolata

469

Gardens Bulletin, S.

v. spathulata, libera, glabra v. apicem versus sparsim hispidula.

Flores masculi ostiolares, ordinibus duobus instructi, sessiles v.

breviter pedicellati, pedicellis hispidulis: stamine 1, pistillodio

nullo. Flores feminei et cecidiophori sessiles v. pedicellis —1.3

mm. longis, hispidulis praediti: ovario sessili albido: stylo glabro

v. subpuberulo. Semina 0.7—-8 mm. longa, lentiformia, subca-

rinata, laevia. Cystolitha amphigena, supra saepe ut pili mic-

roscopici: pili laeves.

v. erinobotrya.—New Britain: G. Bateson 148, between Baining

Mts. and Toma; Dept. Agr. T.N.G. A23, Malabunga Mission;

NGF 3510, Galilo village (typus, herb. Lae); Warburg 20824,

20825, Ralum (ut F. semicordata Miq.); J. H. L. Waterhouse

268, Gazelle Peninsula.

Inter F. elmeri Merrill (== F. semicordata Miq.) et F. fulvo-

pilosam Summerh., sed F. gryllus Corner affinis.

var. solomonensis Corner v. nov.—Pili ramulorum petiolorum-

que —1 mm. longi. Basis laminae minus cordatus, petiolum haud

obtegens: costae laterales utrinsecus 5—8: petiolus 5-27 mm.

Receptacula pedunculis 2—9 mm. longis, bracteis basalibus in

collare ternatis, pedicellis O-9 mm. longis praedita: corpore sine

bracteis lateralibus. Tepala 4-5, florum pedicelli glabri. Cysto-

litha plerumque ut pili microscopici. Arbor -10 m, alta: ramulis

1-2 mm. crassis: lamina 6-26 & 3-13 cm., Solomon Islands.

L. J. Brass 3294, (Ysabel, Tasia), 3483 (Florida, Olevuga);

Kajewski 2364, Malaita, (typus, herb. Kew); J. H. L. Waterhouse

B99, B99a, 148, 794, (Bougainville, Siwai), 165 (New Georgia).

F. fulvopilosae Summerh. affinis et forsan subspecies sed foltis

acute acuminatis, intercostis numerosis strictis, receptaculis mino-

ribus, et habitu cauliflori dissimilis.

f. glabrior Corner f. nov.—Sparsim brunneo-hispidula vel glabra.

Setae internae —0.5 mm. longae, copiosae. Cystolitha hypogena

bene evoluta. Solomon Islands.

L. J. Brass 2557, San Cristoval, Waimamura: E. S. Brown 5054

(Guadalcanal, Aruligo), 5762, 5763 (Nggela, Soso; typus 5763,

herb. Brit. Mus.), W/265 (San Cristoval, Aronopo); J. H. L.

Waterhouse B260, B315 (Bougainville, Siwai).

F. storckii Seem. revocans sed lamina plus asymmetrica et acute

acuminata, intercostis numerosis strictis, et cauliflora. F. trachy-

pison Laut. et K. Schum var. pallidae Corner et F. melinocarpae

Bl. var. hololamprae (Diels) Corner (species ? non caulifiora)

comparanda.

470

a ee NT mae ey EN wry TW um mrt cere Ny grt ae emt hE Te op ae

Vol. XVII. (1959).

F. opposita Mig. Hook. Lond. J. Bot. 7 (1848) 426.—F. con-

jugata Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 222, 291.—

F. opposita occurs in Australia and along the southern coasi of

New Guinea, extending to the Eastern Highlands in the Terri-

tory of New Guinea. F. conjugata was collected in Pulau Sang-

rang (Java), by Teysmann and de Vriese (type at Leiden), but

has never been found thereabouts again. Nevertheless I can

detect no difference from F. opposita, which is itself variable.

The species of sect. Sycidium in Madagascar, however, are clo-

sest to the Australian alliance of F. opposita and this lone sta-

tion of F. conjugata becomes significant.

var. micracantha (Mig.) Corner comb. nov.—F. aculeata Mig.

var. micracantha (Miq.) Benth. Fl. Austral. 6 (1873) 175.—F.

aculeata A. Cunn. ex Mig. Hook. Lond. J. Bot. 7 (1848) 426.—

F. orbicularis A. Cunn. ex Miq. id.—F. micracantha Mig. Ann.

Mus. Bot. Lugd. Bat. 3 (1867) 221, 291.—That this is merely a

small-leafed variety with aculeate hairs and slightly smaller figs is

shown by intermediate collections, particularly from New Guinea.

var. indecora (A. Cunn. ex Mig.) Corner comb. nov.—

F. indecora A. Cunn. ex Mig. Hook Lond. J. Bot 7 (1848) 426.—

This has been confused with F. orbicularis (v. micracantha,

above), but is distinguished by the foveolate areolae (stomatal

pits) on the underside of the lamina and by the lack of the aculeate

hairs. The few collections indicate that it is restricted to Western

Australia, where it is close to F. scobina Benth.

Broadbent 774a, Yarru, Grey Creek; A. Cunningham 65 and

306 (1820), Careening Bay (type); Gardner 66, E. Kimberley.

Moogerooga Creek; F. v. Mueller s.n.. Victoria River (1855):

Ostenfeld 1176, 1177, Derby; Perry 2578, pr. Carlton Station:

Schultz 407, Port Darwin.

F. scobina Benth. Fl. Austral. 6 (1873) 176.—1I take Schultz 410

as the type (lectotype) because Cunningham 124, also cited by

Bentham, is F. opposita Mig. v. micracantha (Miq.) Corner.

F. scobina is very near to F. cumingii Miq. of the Philippines,

extending to New Guinea as v. androbrota (Summerh.) Corner.

so that it may be an Australian variety, but the collections are

too few.

- Schultz 160, 410, 499, Port Darwin; Schomburgk, s.n. Port

Darwin; Specht 873, Arnhem Land, Yirrkala.

F. fraseri Mig. Hook. Lond. J. Bot. 7 (1848) 235.—F. proteus

Bur. Ann. Sc. Nat. ser 5, 14 (1872) 250.—F. stenocarpa F.v.

Muell. ex Benth. Fl. Austral. 6 (1873) 174.—This has been con-

fused with F. fraseri (Mig.) Miq., based on Urostigma fraseri

47]

Gardens Bulletin, S.

Miq., and which is a synonym of F. virens Ait. The type of F.

fraser’ Mig. (Fraser 154, Brisbane River) shows that it is

identical with the well-known F. proteus.

F. coronata Spin. Cat. H. St. Sebast. (1818) 29; Roehm. et Schult.

Mant. 1 (1822) 327; Mant. 3, add. 2 (1827) 403.—F. muntia

Link, En. Pl. 2 (1822) 450.—F. stephanocarpa Warb. Fedde’s

Rep. 1 (1905) 75.—F. aspera hort. et F. scabra hort., non

verae.—I have examined the type of F. coronata Spin., which is

well-preserved in the Turin-herbarium, and it is exactly the spe-

cies known as F. stephanocarpa Warb.; indeed, the description

is unmistakable, as is that, also, of F. muntia. It appears to

have been one of the early species introduced to cultivation in

European hot-houses.

F. aspera Forst. f. Prodr. Fl. Austral. (1786) 76, n. 404—

F. parcelli Veitch ex Cogn. et Marchal, Pl. Ornem. 2 (1874) t.

47.—F. parcelli is merely the variegated state of F. aspera, but

there are few collections of this species, and none that is good.

Indeed, it is difficult to know how it differs from F. scabra

Forst. f., except in the larger villous figs and the more copious

veining.

F. storckii Seem. var. kajewskii (Summerh.) Corner comb. nov.—

F. kajewskii Summerh. J. Arn. Arb. 13 (1932) 103; Bull. B.

Bish. Mus. 141 (1936) 53.—A character of F. storkii is the

presence of abundant cystoliths on both sides of the lamina, and

this is the surest distinction, in the herbarium, from glabrous

forms of F. scabra, which has cystoliths only on the underside,

Amphigenous cystoliths also distinguish F. kajewskii, which dif-

fers from F. storkii merely in the smaller leaves and figs. Thus,

it may be the state of F. storckii in exposed situations, or it may

represent twigs of old trees (10-15 m. high) with reduced size

of leaf and fig; several more or less intermediate collections come

from Fiji. However, var. kajewskii strongly resembles the wide-

spread F’. ampelas Burm. f. which differs mainly in the narrower

and more asymmetric leaf-base, the basal bracts not ternate but

scattered on the fig-stalk, and the presence of sclerotic cells in

the fig-wall. I consider that they are parallel reductions, Malay-

sian and Polynesian, of the F. melinocarpa -stock.

New Hebrides.—Aubert de Ia Rue s.n. (Aug. 1937); E. S.

Brown s.n., Espirito Santo; Kajewski 216, 233, 343, 737 (typus).

Fiji—Parham 316; Degener 14517; A. C. Smith 460, 532,

1323, 5272, 607 1;,.6606;.7 72843403:

472

Vol., XVI. (1959).

subsect. Varinga (Miq.) Corner, ser. Cyrtophylleae Corner

F. asperiuscula Kunth et Bouché, Ind. Sem. Hort. Berol. (1846)

21; Ann. Sci. Nat. ser. 3, 7 (1847) 253; Miq. Fl. Ind. Bat. 1,

2 (1859) 300; Ann. Mus. Bot. Lugd. Bat. 3 (1867) 292.—F.

coronata Reinw. ex Bl. Bijdr. (1825) 470 (non Spin.).—F.

grewiaefolia Kunth et Bouché id. (non Bl.)—Covellia zollinge-

riana Miq. Hook. Lond. J. Bot. 7 (1848) 460.—F. leptorhyncha

Koord. et Val. Bijdr. Booms Java 11 (1906) 156.—F. inaequi-

latera Ridl. J. Str. Br. R. As. Soc. 1 (1923) 93.—I have not

found the type of F. asperiuscula but there are several collections

from Java at Leiden, identified as F. coronata Reinw., on which

Miquel has written “F. asperiuscula Kth. et Bouche’. As this

species agrees with the description, I take Miquel’s authority for

the interpretation of F. asperiuscula. It occurs only in Sumatra

and Java, the Malayan records being F. obscura Bl., which has

been much confused with it. A diagnostic character is the hooked

apex of nearly all the hairs.

F. cyrtophylla Wall. ex Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867)

282, 296.—Covellia cyrtophylla Wall. ex Mig. Hook. Lond. J.

Bot. 7 (1848) 460.—F. asymmetrica Lévl. et Vant. Fedde’s

Rep. 4 (1907) 82.—This species was erroneously reduced to

F. obscura Bl. by King. It is a species of northern India, upper

Burma, Tonkin, and South China, allied with F. praetermissa

Corner and with the East African species centred about F. stor-

tophylla Warb. It parallels in a remarkable way F. fulvopilosa

Summerh. of Fiji.

India (Assam, Bhutan, Khasia, Sikkim).—T. Anderson 222;

C. B. Clarke 11961, 13571, 25411, 25508; Cooper and Bulley

1060; Gamble 1152a, 9654; Griffith 4643; Hooker s.n., Khasia

and Sikkim; King 2370, 5066, 5132; N.E. Parry 784; Thomson

s.n. Sikkim; Wallich 4532 (typus).

Burma.—E. M. Buchanan 18, Myitkyina; Kingdon Ward 171

(Ngawchang Valley, Htawgaw), 9112 and 12932 (Nam Tamai

valley), 20576 (Sumpra Bum); J. H. Lace 5345, Kadu; S. Toppin

4085, Kachin.

China (Kweichow, Yunnan).—Bodinier 2577; Cavalerie 2716,

3596, 3598; Esquirol 2692, 3108, 3549, 5238; Forrest 9419,

111535429995: Henry 11462, 12180; 'T. T) Yuuswsse, 17465,

20488.

Tonkin.—Balansa 747. 2943; Poilane 25473, 25492, 25960.

F. leptogramma Corner sp. nov.—Arbor —7 m, alta, glabra, follis

brevipetiolatis distichis. Ramuli 2-3 mm. crassi, pallide griseo-

brunnei, sicco sulcato-striati, vetustiores frustis corticis elongatis

473

Gardens Bulletin, S-

angustis vestiti. Stipulae —15 mm. longae, binatae, subpersis-

tentes, striatae. Lamina 10-35 2.7-14 cm., elliptica v. subo-

vata, apice caudata —60 mm. longo, basi rotundata v. cuneata,

breviter dentato-repanda, utrimque subscabrida, membranacea,

sicco supra grisea, subtus brunneola; costis lateralibus utrinsecus.

8-12, saepe glandula axillari praeditis, intercostis strictis —12,.

subtus leviter elevatis; costis basalibus utrinsecus 1—2, brevibus,

glandulis basalibus 2: petiolo 3-12 (—20) mm. longo, sicco

striato. Receptacula in cumulis —12 cm. latis ad ramos breves.

efoliatos congestos cauliflora: pedicellis 17-25 > 2-2.5 mm.,

bracteis lateralibus 1-3, 1-2 mm. longis, dispersis; corpore

recptaculi 10-15 & 10-12 mm. (16—20 « 15-17 mm., vivo),

subellipsoideo, lenticellis asperato, bracteis lateralibus numerosis

2—3 mm. longis et latis, dispersis, apicem receptaculi versus

confertis, ostiolo bracteis apicalibus planis parvis occluso: setis.

internis et cellulis scleroticis nullis. Tepala albida, libera v. levi-

ter connata, glabra, anguste oblonga v. subspathulata, ovario

equalia. Flores masculi 2—3.5 mm. alti, ostiolares, ordinibus

2-3 instructi, sessiles vy. subpedicellati: tepala 3-4: stamine

1 (—2), submucronato, pistillodio nullo v. raro evoluto. Flores

cecidiophori 2.5—6.5 mm. alti, sessiles v. pedicellis glabris —3.5

mm. longi praediti;: tepala 5—6: ovario sessili v. substipitato,

albido. Flores feminei similes sed pedicellis —2.5 mm., tepalis

5—8, ovario ad 1.5 mm. stipitato, stylo glabro, stigmate clavato.

Semina 1 mm. longa, lentiformia, subcompressa, laevia, haud

carinata. Cystolitha amphigena, supra sparsa. North Borneo,

Sarawak, in silvis.

Kinabalu.—Sing. F.n. 26754 (Menetendok River, 1,000 m. alt.),

27328 (Koung, 500 m, alt,; typus, herb. Singapore). Sarawak.—

Corner s.n. (1959), Kuching, Bt. Kuap.

F.. praetermissae Corner affinis sed inter alia cauliflora, recep-

taculis majoribus. Ramulis stipulis petiolisque siccis striatis facile

recognoscenda. .

F, praetermissa Corner sp. nov.—Gagnep. Fl. Gen. I-.C. 5 (1928)

812 (ut F. rostrata Lam.).—Frutex, ? sarmentosus, v. arbor —6

m. alta, minute scabrido-puberula v. subglabra, foliis brevipe-

tiolatis distichis. Ramuli 1.5—3 mm. crassi, brunnei. Stipulae

parvae caducae. Lamina 7-25 x 3.5—9.5 cm., elliptica v. an-

guste obovata, apice caudata 12-35 mm. longo, basi cuneata

et subasymmetrica, dentato-repanda v. denticulata, membra-

nacea, scabrida, sicco pallide brunnea: costis lateralibus utrin-

secus 5—8 (—9), obliquis, intercostis strictis 3-7, subtus leviter

474

Vol. XVII. (1959).

elevatis: costis basalibus utrinsecus |, brevibus v. ad 1/3 lami-

nae elongatis, glandulis basalibus 1-2 v. nullis: petiolo 2—15

mm. longo. Receptacula axillaria, maturitate rubra dein nigra:

pedicellis 2-5 mm. longis, bracteis lateralibus 1—2 parvis dis-

persis: corpore receptaculi 6-10 mm. lato, bracteis lateralibus

0-2 dispersis, ostiolo plano v. leviter depresso bracteis apica-

libus parvis occluso: setis internis et cellulis scleroticis nullis.

Tepala 3-5 albida glabra libera, anguste oblonga, ovario lon-

giora. Flores masculi ostiolares, ordinibus 2—3 instructi: stamine

1, pistillodio nullo v. parvo. Flores feminei et cecidiophori sub-

sessiles: ovario sessili albido: stylo glabro v. sparsim puberulo.

Semina sublenticularia v. breviter oblonga, vix carinata, laevia.

Cystolitha amphigena copiosa. Tonkin, Laos, Thailand.

Tonkin.—Balansa 2970 (Mt. Bavi, typus, herb. Paris), 4457

(Moe Ha; subsarmentosa). Laos.—Poilane 1904 (prov. Sau Nena,

Muong Pun), 26434 (Haut Laos, inter Tafu et Bettown-Sai); Spire

199 (Bai Som Pho); Thorel (Bang Mue, pr. Lakon). Thailand. —

Kostermans 771 (prov. Kolug chada, Rau ti River valley); Winit

1763 (Nau, Nam Ken).

Species insignis praetermissa. A F. sinuata Thunb. (F. rostrata

Lam.) intercostis numerosis strictis, cystolithis amphigenis, brac-

teis corporis receptaculi lateralibus, pistillodio florum masculorum

deficienti profunde differt, et speciebus africanis, ut F. urceolari

Welw. ex Hiern, vero affinis.

subsect. Palaeomorphe (King) Corner ser. Pallidae Mig.

F. tinctoria Forst. f. Prodr. Fl. Austral. (1786) 76.—I distinguish

four subspecies, three of which fill large geographical regions

but with intermixture or overlapping at the boundaries. There

is no specific difference in fig, flower, seed, or leaf, and it is

possible to arrange specimens to form a continuous series. F.

tinctoria and F. virens Ait. (subgen. Urostigma) are the most

widespread species of Ficus and both are epiphytic stranglers,

but this subsect. Palaeomorphe does not extend to Africa, as

does sect. Urostigma to which F. virens belongs.

subsp. tinctoria.—Arbor, saepe epiphytica et suffocans. Lamina

4-13 cm. lata, obtusa, subacuta, v. subacuminata, oblonga, ellip-

tica v. ovato-elliptica, nec dentata nec angulata, basi sub-

cordata, rotundata, v. late cuneata, laevis v. subscabrida,

subtus brunneo-areolata: stomata immersa. Receptacula pedi-

cellata. Hainan, Formosa, Philippines (incl. Palawan), Celebes,

475

Gardens Bulletin, S.

Sumbawa, Moluccas, Alor, New Guinea, North Australia,

Micronesia, Polynesia to Tahiti and Austral Isl.

This is the insular state of the species as opposed to the conti-

nental.

subsp. swinhoei (King) Corner comb. nov.—F. swinhoei King, Ann.

R. Bot. Gard. Calc. 1, 2 (1888) 81, p. 101 C.—F. fenicis Merr.

Philip. J. Sci. 18 (1921) 66.—Subscandens ad saxa scopulosque,

scaberrima, hispidula. Lamina -10 6 cm., minor, elliptica,

obtusa v. acuta, subsymmetrica, basi subcordata v. late cuneata,

rigide coriacea, subtus haud brunneo-areolata: stomata immersa.

Receptacula 8 mm. lata: pedunculo brevi, —1.5 mm. longo, brac-

teis basalibus ternatis: pedicello 2—4 mm. longo. Formosa,

Mindanao.

subsp. gibbosa (Bl.) Corner comb. nov.—F. reticulata Thunb. Diss.

Ficus (1786) 6, 12.—F. gibbosa Bl. Bijdr. (1825) 466.—Arbor

epiphytica suffocans, etiam littoralis ad saxa repens. Lamina 4—

13 & 2.5-6 cm., anguste elliptica v. lanceolata saepe plus minus

obovata, saepe subrhomboidalis, angulata v. utrimque 1-2 an-

gulato-dentata, apice acuta v. valde acuminata, basi cuneata,

coriacea, haud v. vix scabrida, subtus brunneo-areolata. Recep-

tacula 7-12 mm. lata.

var. gibbosa.—Lamina flexibilis, stomatibus immersis. Recep-

tacula pedicellata. South Andaman Isl., south China, Hainan, Indo-

china, Thailand, Malaya, Sumatra, Java, Borneo, Palawan, Cele-

bes, Soembawa, Banda, Halmaheira.

var. rigida (Mig.) Corner comb. nov.—F. rigida Bl. Bijdr.

(1825) 465, (haud Jack).—F. pereng Steud. Nomencl. ed. 2 (1840)

637.—F. gibbosa BI. var. rigida Mig. Ann. Mus. Bot. Lugd. Bat 3

(1867) 276.—var. rigida (Bl.) Val. Bijdr. Booms. Java 11 (1906)

171 :—Lamina rigide coriacea, acute acuminata: stomatibus haud

immersis. Receptacula sessilia v. breviter pedicellata: bracteis

basalibus ternatis, 1-2 mm. longis. Sumatra, Java, Soembawa, Alor.

South Borneo. ;

This is the state of the species in South East Asia and Western

Malaysia, but it intergrades with ssp. tinctoria in Celebes, Hal-

maheira, and Palawan, and with ssp. parasitica in Hainan, south

China, Indochina, and the Andaman Islands. How the species

is to be classified in these boundary-regions will be a problem

for local botanists to investigate.

subsp. parasitica (Willd.) Corner comb. nov.—F. parasitica Willd.

Mem. Acad. R. Sc. Bell. Lett. Fr. Guill..2 (1798) 102, t. 3.—

F. gibbosa Bl. var. parasitica (Willd.) King, Ann. R. Bot. Gard.

Calc. 1 (1887) 6, pl. 2 a-b.—Lamina ut in ssp. gibbosa sed saepius

476

Vol. XVH. (1959).

subrhomboidalis et angulata, tenuiuscule coriacea, scabrida v.

laevis, saepe subtus puberula v. subvillosa, haud brunneo-

areolata: stomatibus haud immersis. Receptacula pedicellata.

var. parasitica.—Arbor epiphytica suffocans. Receptacula 7-11

mm. lata. Ceylon, India, Burma, south China, Indochina, Andaman

Islands, Nicobar Islands.

var. anastomosans (Wall. ex Kurz) Corner comb. nov.—

F. anastomosans Wall. ex Kurz, For. Fl. Br. Burma 2 (1877) 455.

—Frutex v. ad rupes calcareos repens. Glabra v. ramulis foliis

receptaculisque albido-puberulis. Lamina 3-9 x 1-5 cm., ellip-

tica v. subrhomboidalis, dentata v. pinnato-lobata, subscabrida:

petiolo —5 mm. longo. Receptacula 6-8 * 5-6 mm., pedicello

—15 mm. longo. Burma, Thailand.

Lobb s.n. Moulmein; Wallich 4513, Kogun, typus; A.F.G. Kerr

282 (Prachuap), 2510 (Sam Roi Yawt), 5239 (Mé Poi, Ching Dao),

6089 (Mé Lamung, Kampeng Pet), 12854 (Wang Kanai, Kanburi).

Varietas inspicienda. Forsan forma juvenilis fertilis. Folia ut

in Streblus asper Lour.

F. virgata Reinw. ex BI. Bijdr. (1825) 454.—F. decaisneana Miq.

Fl. Ind. Bat. 1, 2 (1859) 312.—F. firmula Migq. et F. trymato-

carpa Mig. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 224, 292.—

F. pinkiana F.v.M. in Wing’s South Sci. Rec. 2 (1882) 273.-—-

F. esmeralda Bailey, Queensl. Agr. J. 1 (1897) 452.—F. inaequi-

folia Elmer, Leafl. Philip. Bot. | (1907) 242.—As this species

was named by Reinwardt and published by Blume, the type

is Reinwardt 1376 (Banda Major, Moluccas) in the Leiden her-

barium, where it is named by both authors. Blume, however,

added Java as a locality. No Javanese specimen exists and the

species, as defined on Reinwardt 1376, does not occur there.

Possibly, as others have suggested, Blume mistook a specimen

of F. subulata for Reinwardt’s species, but that does not mean

that it should be reduced to F. subulata. | have carefully studied

Reinwardt 1376 and am sure from its amphigenous cystoliths

that it is the wide-spread species known as F. philippinensis

and F. decaisneana. lt is closely allied with F. tinctoria and with

F. subulata, and most specimens have been wrongly referred

to these species.

var. philippinensis (Miq.) Corner comb. nov.—F. philippinensis

Miq. et F. insularis Mig. Hook. Lond. J. Bot. 7 (1848) 435.—-

F. magnifica Elmer, Leafl. Philip. Bot. 1 (1906) 51.—F. setibrac-

teata Elmer, Leafl. Philip. Bot. 7 (1914) 2411, 2413.—Receptacula

interne et tepala rubra, pallescentia: setis internis copiosis; pedi-

cellis florum saepe hispidulis. Philippines, Formosa, Ryu Kyu.

477

Gardens Bulletin, S.

var. sessilis (Bur.) Corner comb. nov.—F. philippinensis Mia.

var. sessilis Bur. Ann., Sci. Nat. ser. 5, 14 (1872) 253. Ut v. virgata,

tepalis albidis, sed receptaculis plus minus sessilibus: bracteis

basalibus 3-6, 1-3 mm. longis. Philippines, Moluccas, New Guinea,

New Britain, Solomon Isl., New Caledonia, Loyalty Isl., New

Hebrides. |

F. celebensis Corner nom. nov.—F. irregularis Mig. Ann. Mus.

Bot. Lugd. Bat. 3 (1867) 224, 292 (non Steud.).—This willow-

like tree is now cultivated in several parts of the tropics and

subtropics, but its identity is generally not known. Hence this

inevitable name-change will not be serious: it will indicate the

origin of the plant and how little is yet known of it in the

wild state.

subsect. Paiaeomorphe (King) Corner ser. Subulatae Corner

F. subulata Bl. var. gracillima (Diels) Corner comb. nov.—F.

gracillima Diels, Engl. Bot. Jahrb. 67 (1935) 194.—? F. otario-

phylla Diels, id. 209.—Over its great range from the Himalayas

to the Solomon Islands and New Hebrides, var. subulata is re-

markably uniform. In New Guinea there occurs this state, var.

gracillima, with small lanceolate, caudate-acuminate lamina (3—

13 x 0.9—3 cm.) without intercostal veining, generally with one

basal gland in the axil of one of the lower lateral nerves (not

of a basal nerve), and with small figs 4-5 mm. wide (dried):

further, the male flower lacks the gall-ovary. These plants are

said to be slender epiphytes or scandent bushes in the mountains

of New Guinea. Var. subulata is normally an epiphyte rooting

to the ground, but not strangling, yet it, too, sends out slender

creeping runners with reduced leaves and short adventitious

roots whereby it attaches itself to other trunks. Several coilec-

tions, also, from New Guinea bridge the difference in leaf-size.

Therefore, though var. gracillima in the extreme looks a different

species, it clearly grades into typical F. subulata. In the Hima-

layas F. sikkimensis, which I regard as synonymous with v.

subulata, is a less reduced state, also lacking the gall-ovary in the

male flower.

F, otariophylla, from New Guinea, has the short stipules and

small figs of var. gracillima, but the figs are cauliflorous. Its main

twigs bear large leaves (-30 & 9 cm., lamina) as in var. subulata,

but the side-branches bear abruptly the small leaves of var. gracil-

lima, Such a rapid change from large to small leaf on branching

478

FO Be SSeS se SE PRA S

Vol. XVIT. (1959).

occurs also in F. virgata Reinw. ex Bl. Hence, I suspect that F.

otariophylla may be the sapling, or bathyphyllous, form of var.

gracillima.

I refer the following to var. gracillima:

Brass 11391 (Neth. New Guinea, Bele River, 2,200 m. alt.),

13865 (Idenburg River, Bernhard Camp), 23280 (Milne Bay

district, Mt. Dayman, 1,370 m. alt.); Carr 13261, 13353, 13449

(Boridi, 1,500 m. alt.), 15387 and 15430 (Isuarava, 1,500 m.

alt.), 15980 (Lala River, 1,600 m. alt.); Clemens 5045 Morobe

district, Ogeramnang, 5,900 ft. alt.); Docters v. Leeuwen 10121,

11110 (Rouffaer River); Ledermann 8174 (Sepik, type); Meyer-

Drees 603 (Neth, New Guinea, Bernhard Biv.); NGF 6161 (East-

ern Highlands pr. Goroka, 7,300 ft. alt.), 8477 (Morobe district,

pr. Skindewai, 5,400 ft. alt.), 9433 and 9455 (Western Highlands,

Mt. Hagen, 7,000 ft. alt.), 9565 (Eastern Highlands, 6,500 ft.

alt.).

F. otariophylla Diels —Ledermann 8572 (Sepik, type); Carr

15925, 16110a (Isuarava, 1,100 m. alt.).

subsect. Palaeomorphe (King) Corner ser. Cuspidatae Mig.

F. sinuata Thunb. Diss. Ficus (1786) 6, 12.—F. rostrata Lam.

Encycl. 2 (1788) 498.—Examination of Thunberg’s type

(Uppsala), shows that it is F. rostrata Lam. (type, Paris). Thun-

berg ascribed it to Ceylon, where this species does not occur,

and he must have made a mistake for Java, where it is common.

ssp. sinuata var. oblonga Corner nom. nov.—F. cuspidata

Reinw. ex BI. var. sinuata King, Ann. R. Bot. Gard. Calc. 1,

2 (1888) 89, pl. 112 C.—This differs from v. simuata in the

narrowly oblong lamina with more numerous lateral nerves

at right angles to the midrib. With the change in specific epithet.

so King’s varietal name must be changed.

Malaya.—King’s coll. 7256 (Perak); Sing. F.n. 21338, 29023,

32502 (Johore). Sumatra.—O. Hagerup s.n. (1916-17, Lake Toba),

Yates 1414 (East Coast). Java—dZollinger 1674.

ssp. cuspidata (Reinw. ex BI.) Corner _comb. nov.—F’.

cuspidata Reinw. ex Bl. Bijdr. (1825) 464 (incl. a and B).—I

cannot separate this from the variable F. sinuata except as a

montane subspecies with smaller, subsessile, fig and smaller, more

or less lanceolate, leaf. Small-leafed states of ssp. sinuata v.

sinuata seem to merge into ssp. cuspidata, but its greater fe-

semblance is with v. oblonga.

479

Gardens Bulletin, S.

F. heteropleura Bl. Bijdr. (1825) 466.—F. urophylla Wall. ex

Miq. Hook. Lond. J. Bot. 7 (1848) 429.—F. caudatifolia

Warb. Perk. Fragm. Fl. Philip. 3 (1905) 194.—F. eucaudata

Elmer, Leafl. Philip. Bot. 1 (1906) 40.—F. rostrata Lam. var.

urophylla (Wall.) Koord. et Val. Bijdr. Booms. Java 11 (1906)

175.—Blume’s type at Leiden is well-preserved and accompanied

by a fine painting, which leaves no doubt of its identity.

var. hirta Corner var. nov.—Ut v. heteropleura sed ramulis,

petiolis, costisque (subtus) pilis albidis dein brunneis 0.5—1 mm.

longis hispido-villosis. Receptacula minora 5—10 mm. lata, pedi-

cello breviori —3 mm. longo. Sarawak.

J. A. R. Anderson 9147 (Lundu); Beccari s.n. et 780 (Ku-

ching); Bur. Sci. Pl. Saraw. 1938 (typus, herb. Leiden); Kep.

F.n. 80038 (B.6 For. Res.).

var. mindanaensis (Warb.) Corner comb. nov. F. mindanaensis

Warb. Perk. Fragm. Fl. Philip. 3 (1905) 195.—This is distin-

guished by its larger leaf and fig, but there are several intermediate

collections from the Philippines so that it is with difficulty separ-

able.

subsect. Palaeomorphe (King) Corner ser. Minutuliflorae Sata

F. aurita Reinw. ex Bl. var. auriculifera (Merr.) Corner comb.

nov.—F’. quriculifera Merr. Univ. Calif. Publ. Bot. 15 (1929)

46.—Stipuiae caducae. Lamina auricula basali minori —8 X

2 mm. praedita: costis lateralibus utrinsecus 4—7. Borneo, Cele-

bes, Philippines, Amboina.

Borneo.—Clemens 10467, 26194, 26611, 30288 (Kinabalu);

Elmer 20468 (Tawao, type); Endert 4667 (W. Koetai), Celebes.

—Bloembergan 4106 (Menado, G. Ngila-laki); Kjellberg 2351

(Malili), NIFS Cel/V-284 (Malili, Kawata). Bohol.—Bur. Sci.

42744. Amboina.—Binnendyk 6820.

var. celebica (Reinw. ex BI.) Corner comb. nov.—F.

celebica Reinw. ex Bl. Bijdr. (1825) 461.—Stipulae persistentes.

Lamina haud auriculata, basi cuneata: costis lateralibus utrinsecus

3—7. Celebes.

Reinwardt 1626, Kamanga, type; Koorders 19257, Menado,

Riedel s.n., Gorontalo; Teysmann et de Vriese s.n., s.l.

subsect. Palaeomorphe (King) Corner ser. Fibrosifoliae Corner

F. obscura BI. Bijdr. (1825) 474.—This common and variable

species of Western Malaysia has been much confused through

King’s error in identifying with it F. asperiuscula Kunth et

480

Vol. XVI. (1959):

Bouché and F. cyrtophylla Wall. ex Miq. In subordinating the

swarm of unnecessary species created by Miquel, King mistook

F. obscura and used F. pisifera Wall. ex Miq. in its place. Four

sheets named by Blume at Leiden leave no doubt what he in-

tended, and on this basis I have re-constructed. In recognising

five varieties I realise that intermediates occur which will require

investigation, but most of the five hundred collections which have

now been made fall into these categories without much difficulty.

They seem to have little geographical significance.

var. obscura.—Receptacula 9-12 mm. lata (12—20 mm., viva),

plus minus brunneo-hispida v. villosa (ut ramuli). Lamina supra

scabrida. Java, Sumatra, Borneo, Celebes, Philippines, Talaud.

The large-fruited, hairy, and scabrid state, not known from

Malaya, except for one plant of doubtful origin in the Penang

Waterfall Gardens.

var. angustata (Mig.) Corner comb. nov.—F. hypsophila Miq.

var. angustata Miq. Pl. Jungh. (1851) 60.—Plus minus glabra,

laevis v. vix scabrida. Lamina basi anguste cuneata, vix asym-

metrica, saepe coriacea et bullata, denticulata v. integra, stipulis

caducis. Receptacula 5-8 mm. lata. Java, Sumatra, Lingga, Bor-

neo, Celebes, Moluccas (Soela).

F’. uniglandulosae Wall. ex Miq. similis, sed nervis ut in F. obs-

cura et tepalis albidis. Forsan species distincta. F. uniglandulosa

nondum in Java inventa.

var. borneensis (Miq.) Corner comb. nov.—F. microtus Miq.

var. borneensis Miq. Ann. Mus. Bot. Lugd. Bat. 3 (1867) 273.—

F. pisifera Wall. ex Mig. Hook. Lond. J.-Bot. 7 (1848) 427;

King, Ann. R. Bot. Gard. Calc. 1 (1887) 3, pl. 1.—Ramuli et

petioli brunneo-villosi glabrescentes. Lamina plus minus scabrida.

Receptacula 5—8 mm. lata, scabrida, brunneo-puberula glabres-

centia: pedicello 2-6 mm. longo. Lower Thailand, Malaya, Riouw,

Sumatra, Java, Borneo, Celebes, Philippines.

The commonest variety, as a small-fruited state of v. obscura.

var. kunstleri (King) Corner comb. nov.—F. celebica Reinw.

ex Bl. var. kunstleri King, Ann. R. Bot. Gard. Calc. 1 (1887) 12,

pl. 10 A.—F. acuminatissima Miq. Hook. Lond. J. Bot. 7 (1848)

233.—F. lancifolia Miq. id. 452.—Ut. var. scaberrima sed lamina

lanceolata acuminata. Malaya (Perak), Sumatra (Fort de Kock),

Bangka, Borneo, Celebes, Philippines.

Forsan v. scaberrima statu vetuso sed in Java non inventa. King’s

coll. 3927 (Perak, typus) in errore sub F. celebica (F. aurita var.

celebica) disposita.

481

Gardens Bulletin, S.

var. scaberrima (BI.) Miq. Ann. Mus. Bot. Lugd. Bat. 3 (1867)

273.—F. scaberrima Bl. Bijdr. (1825) 474.—Ut in v. obscura sed

receptaculis 6-7 mm. latis, sessilibus v. breviter pedicellatis —2

mm., brunneo-setosis, tarde glabrescentibus. Sumatra, Java, Bor-

neo, Celebes, Philippines.

F. midotis Corner sp. nov.—Frutex v. arbor —11 m. alta, saepe

epipytica sed non suffocans, glabra, sparsim puberula, v. scab-

ridula, foliis distichis. Ramuli 1-2.5 mm. crassi, brunnei dein

lutescens. Stipulae -12 mm. longae, caducae. Lamina 11-33

3.5-13 cm., ovato-elliptica, oblonga, v. ad ramulos ultimos

anguste elliptica, symmetrica, apice caudata 12-45 mm. longo,

basi late v. anguste cuneata, symmetrica v. uno latere breviter

decurrens et subauriculata, crenato-dentata v. integra, membra-

nacea v. subcoriacea, utrimque subscabrida v. laevis, sicco

brunnea: costis lateralibus utrinsecus (6—) 8—13, obliquis, saepe

glandula axillari praeditis, intercostis 3—7, subtus leviter ele-

vatis: costis basalibus utrinsecus 1—2, v. latere latiori —4, glan-

dulis basalibus 1-2: petiolo 3-20 x 2-4 mm., Receptacula

axillaria binata et ad ramos et truncum usque ad basim fasci-

culata, scabridula, maturitate e luteo rubescentia: pedicellis 3-

25 mm. longis, brevioribus ad receptacula axillaria pertinen-

tibus, bracteis lateralibus 1-2 minutis dispersis: corpore recep-

taculi 6-10 mm. lato (10-12 * 9-10 mm., vivo), raro bractea

una laterali praedito, apice subumbonato, ostiolo haud v. vix

depresso bracteis apicalibus parvis occluso: setis internis nullis,

Vv. paucis minutis: cellulis scleroticis nullis. Tepala albida, ob-

longa, acuta v. obtusa, puberula v. apicem versus denticulata,

ad basim leviter connata. Flores masculi ostiolares sessiles, or-

dine uno instructi: tepala S—7: stamine 1, pistillodio bene evo-

luto. Flores feminei et cecidiophori sessiles: tepala 4—5: ovario

sessili, vel cecidiophoro stipitato: stylo glabro. Semina 1.5 mm.

longa, subcompressa, oblonga, apice carinata. Cystolitha hypo-

gena: fibris sclereidiformibus in mesophyllo copiosis. Borneo in

silvis planis et montanis —2,800 m. alt.

Br. North Borneo (in regionibus omnibus ).—Amdjah 651 (det.

F. subulata); Clemens (Kinabalu) 10366, 10469, 27017, 27685,

28667, 29650 bis, 30294, 30624, 36057, 31350, 31551, 31539,

32013, 32222, 32617, 33294, 33643, 33997, 40392, 40624;

Elmer 21908; For. Dept. 1360, 1518, 1555, 7276, A 2936, A

3242, A 4714; Ramos 1441, 1478 (det. F. pisifera); Sing. F.n.

18987, 19061, 26324 (typus, herb. Singapore). 27331, 27380,

27854; D. Wood 1982 (det. F. pisifera). Koetai—Endert 4774;

482

Vol. XVIU. (1959).

Kostermans 5563, 5626. S.E. Borneo.—v. Slooten 2132, Tanah

boemboe, Batu Litjin (det. F. pisifera). Sarawak —J. A. R. An-

derson 8972; Beccari 843; Haviland 1612; Purseglove 4717.

F. obscurae Bl. affinis sed foliis quasi laevibus symmettricis pluri-

nervis, stipulis caducis, receptaculis longipedicellatis, cystolithis

hypogenis differt, et speciminibus jam numerosis certe distin-

guenda.

F. leptocalama Corner sp. nov.—Arbor —5 m. alta, glabra, foliis

brevipetiolatis distichis. Ramuli 1 mm. crassi, graciles, pallide

ochracei. Stipulae —-8 mm. longae, plus minus persistentes, ap-

pressae. Lamina 8-15 x 2.5-4.5 cm., elliptica, subasymmet-

rica, apice caudata —30 mm. longo, basi cuneata uno latere

minute auriculata, integra, membranacea, supra subscabrida,

siccO griseo-viridis: costis lateralibus utrinsecus 7-10, intercostis

~—3, subtus leviter elevatis: costis basalibus utrinsecus 1, brevi-

bus, glandulis basalibus 1-2: petiolo 1-2 mm. longo, brevis-

simo. Receptacula geocarpica ad ramulos graciles repentes efo-

liatos -1.8 m. longos, basi —10 mm. crassos, e basi trunco cres-

centes, binata, maturitate rubescentia: pedicello 1-2 mm. longo,

bracteis lateralibus 2—3 minutis dispersis: corpore receptaculi

6-8 mm. lato, subgloboso, bracteis lateralibus 1-3 parvis dis-

persis, scabriusculo, ostiolo bracteis apicalibus parvis numerosis

occluso: setis internis et cellulis scleroticis nullis. Tepala 4 al-

bida oblongo-lanceolata glabra libera v. vix connata. Flores

feminei sessiles v. breviter pedicellati: ovario sessili: stylo glabro.

Semina 1.2 mm. longa, oblonga, laevia, subcompressa. haud

carinata. Cystolitha hypogena: fibris sclereidiformibus in meso-

phyllo copiosis. North Borneo (Sing. F. n. 26829, Kinabalu,

Menetendok River, Kuala Serab, 1000 m, alt.; typus herb.

Singapore).

F. midotis Corner affinis sed gracilior, geocarpica, et seminibus

haud carinatis.

F. rubrocuspidata Corner sp. nov.—Frutex v. arbuscula, saepe

epiphytica, glabra, haud scabra, foliis brevipetiolatis distichis.

Ramuli 1—1.5 mm. crassi, cremei v. pallide ochracei v. brun-

neoli. Stipulae —8 mm. longae, lanceolatae, caducae. Lamina 5—

23 x 1.2-4.5 cm., elliptico-obovata v. lanceolato-elliptica.

symmetrica, apice caudato-acuminata -45 mm. longo, basi

cuneata, symmetrica, integra, membranacea, laevis, sicco griseo-

viridis: costis lateralibus utrinsecus 5—10 (—14 in foliis juveni-

libus), quasi rectangulatis, intercostis 0—1 (—2): costis basalibus

utrinsecus 1, glandula basali 1: petiolo 2-6 mm. longo, gracili.

483

Gardens Bulletin, S.

Receptacula axillaria binata, etiam ad ramulos defoliatos fas-

ciculata, laevia, maturitate e luteo rubra: pedicello 0-1.5 mm.

longo: corpore receptaculi 3-5 mm. lato, bracteis lateralibus nul-

lis: setis internis et cellulis scleroticis nullis. Tepala 4 rubra, saepe

aetate pallescentia, libera acuta glabra. Flores masculi ostio-

lares, ordine uno instructi: stamine 1, pistillodio nullo v. minuto.

Flores feminei et cecidiophori sessiles: ovario sessili albido:

stylo glabro. Semina 0.8 mm. longa, breviter oblonga, vix cari-

nata, laevia. Cystolitha hypogena: fibris sclereidiformibus in

mesophyllo copiosis. Borneo, in regionibus omnibus, in silvis

—1,500 m. alt.

Amdjah 315; Brunei 5328; Clemens 20192, 25254, 26254,

26986, 30572, 30729, 30824, 31351, 31892, 40365, 40365 A;

Endert 3066, 4677; For. Dept. B.N.B. 1808, 2977, 3117, A

4528, A 4647; Motley 1226; Purseglove 4671, 5470; Sing. F. n.

26339, 27323 (Kinabalu, Koung, 400 m, alt.; typus herb. Singa-

pore).

F. microsphaerae Warb. (ins. philipp.) persimilis sed tepalis

rubris, fibris sclereidiformibus in mescphyllo, costis quasi rectan-

gulatis, lamina haud scabrida differt, etiam in insulis philippinensi-

bus nondum inventa. F. sinuata Thunb. ssp. cuspidata (Reinw. )

Corner similis et eisdem modis differt, etiam seminibus vix cari-

natis. Vero F. uniglandulosae Wall. ex Miq. affinis.

F. rubromidotis Corner sp. nov.—Frutex vel arbor parva, saepe

epiphytica, glabra v. ramulis stipulisque puberulis, foliis bre-

vipetiolatis distichis. Ramuli 2-3 mm. crassi, griseo-ochracei.

Stipulae 10-20 mm. longae, late lanceolatae, binatae, scario-

sae, striatae, brunneolae, saepe ramulos investientes, persis-

tentes. Lamina 15-36 5-12 cm. oblongo-elliptica, apice

attenuato-acuminata v. caudata 12-50 mm. longo, basi rotun-

dato-cuneata uno latere breviter auriculata, integra, rigidiuscule

subcoriacea, laevis v. subtus subscabrida, subbullata, sicco fusco-

olivacea: costis lateralibus utrinsecus (9—) 12—20, subtus valde

elevatis, saepe glandula axillari praeditis, intercostis 6-10, sub-

tus elevatis: costis basalibus utrinsecus 1, brevibus, glandulis

basalibus 1-2: petiolo 1-10 * 3-4 mm., crasso, sub auricula

plus minus condito. Receptacula axillaria binata et ad truncum

ramosque fasciculata, maturitate e luteo rubra: pedicello 1—2

mm. longo vel receptaculo cauliflori 10—15 mm., gracili, bracteis

lateralibus 2—3 minutis dispersis: corpore receptaculi S—7 mm.

lato, subgloboso, bracteis lateralibus nullis, ostiolo plano parvo:

setis internis et cellulis scleroticis nullis. Tepala 4 rubra v. brun-

neorubra oblonga obtusa glabra libra. Flores masculi ostiolares

484

Vol. XVII. CEGs9).

sessiles, ordine uno instructi: stamine 1, pistillodio bene evoluto.

Flores cecidiophori sessiles v. pedicellati, feminei sessiles: ovario

sessili albido: stylo glabro. Semina 1 x 0.5—0.6 mm., oblongo,

subcarinata, laevia. Cystolitha hypogena, fibris sclereidiformibus

in mesophyllo copiosis. Sarawak, Brunei, in silvis humidis, prae-

_sertim ripariis.

W. M. A. Brooke 9125 (Kuching); Brunei 5331, 5340 (Ulu

Belalong); Clemens 20539 (Kuching, G. Tiang), 20699 (Bidi

Cave Mountain; typus, herb. Kew), 22117 (Gat, Upper Rejang

River).

Stipulis scariosis striatis persistentibus, costis numerosis valde

distincta, F. midotis Corner affinis.

Dietdy age ce

485

Posthumous Publication of New Dipterocarp

Species from North Borneo

Footnote:—These notes are put together from material supplied by the

Forest Department of North Borneo and the Forest Research Institute of

the Federation of Malaya, and are now published to validate the new spe-

cies, descriptions of which the authors had left in manuscript. 2

Editor.

1. Dipterocarpus exalatus van Slooten sp.nov. (§ Angulati).

A D. kunstleri, cui simillima, ramulis magis applanatis, inter-

nodiis virgularum glabris, foliis adultis angustioribus, fructibus

validius angulatis, laciniis accrescentibus perbrevioribus vel caren-

tibus haec species sat distincta.

Arbor elata circa 40 m. alta, cum trunco 90 cm. in diam. Folia

chartacea vel coriacea, lanceolata, interdum elliptico- vel oblongo-

lanceolata, apice abrupte caudato-acuminata, basin versus cuneata,

margine crenato-sinuata, 12—22 cm. longa, 5.5—9 cm. lata, costis

nervisque subtus minute lepidotis exceptis utrinque glabra, nervis.

lateralibus utrinsecus 15—20 praedita, petiolis 2-3 cm. longis sti-

pitata. Inflorescentia in ramulis producta, axillaris vel apicalis, soli-

taria, ad 22 cm. usque longa, in ramulos sympodiales 4—8 floribus.

praeditos 1—3-plo divisa. Flores alternati, vel fere secundi; calyx

infundibuliformis, coriaceus vel ligneus, in pedicellum 10-15 mm.

longum attenuatus, prominenter S-angulatus, apice in lobos duos.

majores ad 10 mm. usque longos et in tres alteros fere dimidios.

divisus; petala oblique spathulata, basi connata, in alabastro torta

et extus minute stellato tomentosa. Fructus valde 5-angulatus, 4—5

cm. longus, 2.3—3.5 cm. in diam. glaber, cum laciniis 2 reductis-

vel interdum accrescentibus ad 12 cm. usque longis, ad 3 cm.

usque latis, alteris 3 excrescentibus auriculatis. Nux ovoideus, 4

cm. longus, 2 cm. in diam., superne pubescens, in stylopodium

exsertum, angulato-costatum, paullatim attenuatus.

BoRNEO: Parte orientale indonesiana, apud Balikpapan (bb.

13,913, holotypus in BO). Parte septentrionale britanica, Sanda-

kan, (SAN 1721, paratypi in SAN & BO).

This species is very like D. kunstleri but differs from that in

more flattened twigs, glabrous internodes of the young shoots, nar-

rower leaves, more prominently-angled fruits, much shorter accres--

cent lobes of the fruit.

[The name has been used before as a nomen nudum. The draw-—

ings with notes were prepared by Dr. van Slooten.]

486

Vol. XVII. (1959).

Dipterocarpus exalatus v. Sl. (bb. 13,913, holotypus)

a, Ramulus floriferus. b, Alabastrum ante anthesin. c, Flos post anthesin.

d, Ibid. apertus ut stamina ovariumque appareant. c, Calyx longitudinaliter

dissectus ut ovarium appareat. f, Stamina. |

487

Gardens Bulletin, S.

dk.

Dipterocarpus exalatus v. Sl. (SAN 1721: paratypus)

a, Ramulus fructiferus. b, Fructus juvenilis cum calyce dissecto.

488

Vol. XVII. (1959).

2. Shorea argentifolia Sym. sp.nov. (§ Grex “Red Meranti’’)

Ponenda inter species staminibus 15 biseriatim dispositis; an-

theris quadriloculatis ad anthesin interdum reflexis; appendiculis

haud ciliatis, interdum reflexis in serie interiore quam anthera

plerumque brevioribus. S. leprosulae Miq. similis, sed foliis mino-

ribus, nervis eorum lateralibus pluribus, floribus conspicue pedi-

cellatis, stylis brevioribus haec species sat dissimilis.

Arbor elata, circa 50 m. alta, cum ramulis apice applanatis, gri-

seis, stellato-tomentosis, et gemmis latentibus fulvo-tomentosis.

Stipula conspicua, oblongo-elliptica, acuta, 1-2 cm. longa, circa

9-nervia, extus tomentosa, intus lepidoto-puberula, diu persistens,

dein decidua. Folia oblonga vel obovato-oblonga, interdum ellip-

tico-oblonga, 5—9 cm. longa, 2—4 cm. lata, basi abrupte cuneata

vel rotundiuscula, apice attenuata, acuta vel subito acuminata,

supra glabra, subtus dense tomento-stellata fulvo- vel argenteo-

brunnea, utrinsecus 20—25—nervia, petiolis 0.75-1.25 cm. longis

eodemmodo stellato-tomentosis stipitata. Jnflorescentia paniculata,

ad 14 cm. usque longa, eodemmodo tomentosa, in ramulis secun-

dariis flores 4-8 congestos producta, cum bracteis bracteolisque

pubescentibus, diu persistentibus dein deciduis. Flores ante anthe-

sin 3.5 mm. longi, breviter pedicellati, extus argenteo-tomentosi;

sepala majora 3, ovato-oblonga, extus tomentosa, intus glabra, al-

tera 2 multo minora, ovata, extus partim tomentosa; petala albes-

centia, in parte ante anthesin exposita tomentosa excepta, glabra;

stamina 15 biseriatim disposita, 5 interiora exterioribus 10 fere

duplo altiora, basi dilatata. Fructus 5-angulatus, luteo-brunneus;

calycis lobi stellato-puberuli, concavi, cum laciniis accrescentibus

3, obovato-lanceolatis, 8—11 cm. longis, 1-2 cm. latis, laciniis al-

teris 2 excrescentibus, linearibus, 0.8—1.5 cm. longis, ad 2 mm.

usque latis. Nux ovoideus, in stylopodium conicum attenuatus,

minute tomentosus 0.75—1.5 cm. altus.

BoRNEO: Brunei, Batu Apoi (KEP 39,617, paratypus in KEP;

KEP 40.471, paratypus in KEP). North Borneo: Tawau (KEP

38,838—SAN 4462, holotypus in KEP).

This tall tree has a bole with regular fissures with brown outer

bark and purple brown inner one. The leaves are very narrow.

The tomentum which varies in colour from grey or silvery grey

to yellowish brown is found on the underside of the leaves, branch-

lets, inflorescence axis, bracts and the flower parts exposed in the

buds. Similar to S$. leprosula Miq., but leaves smaller having more

489

Gardens Bulletin, S.

Lig

"yy he

uh,

hi ny

aes

SIRS

OS.

Shorea argentifolia Sym. (fig. 1-17; KEP 38,838; fig. 18; KEP 39,617; fig. 19:

KEP 40,471)

1, Ramulus floriferus. 2, Paniculae fragmentum. 3, Bractea. 4, Alabastrum.

5-9, Sepala. 10, Alabastrum sepalis remotis. 11 & 12, Petala. 13, Flos

sepalis desumptis. 14 & 15, Stamina. 16, Flos sepalis, petalis staminibus

remotis ut pistillum appareat. 17, Ovarium longitudinaliter discissum.

18, Ramulus fructiferus. 19, Fructus (Amussis cum fig. 1, 18 et 19 tantum

congruens; alterae figurae ampliatae).

Vol. XVII. (1959).

lateral nerves, flowers more distinctly pedicelled and style shorter.

It belongs to the S. parvifolia group of the Red Meranti (cf. Sy-

mington, Malay For. Rec. 16, 1943, p. 4 & 58-59).

[The drawing was made by Mr. Wong Sze Moy under the direc-

tion of Symington whose notes have been also incorporated here.

The name has been used in literature as a nomen nudum even in

A. L. Howard’s “A Manual of the Timbers of the World” (1948)}.

3. Shorea superba Sym. sp.nov. (§ Eushorea subsect. ciliata)

Inter omnes hujus subsectionis species truncis fere levigatis

griseis, basi longitudinaliter paulo-tabularibus; foliis multinervatis,

supra griseo rubris, subtus brunneis vel argenteo-griseis, fructibus

parvis cum lacintis 3 majoribus et 2 multo minoribus haec species

sat distincta.

Arbor elata, circa 60-75 m. alta, cum ramulis apice brunneis,

minute stellato-tomentosis. Stipula ovato elliptica, mox decidua,

roseo-viridis, 1—-1.5 cm. longa, 4—5 mm. lata, eodemmodo stellata.

Folia oblongo-lanceolata, 5—9 cm. longa, 2—3 cm. lata, acuminata

vel caudato-acuminata, basi cuneata vel rotundata, supra costis

deciduo-tomentosis exceptis glabra, nitida, brunnea, subtus minute

stellato-tomentosa, vetustiore fere glabra glaucescentia, utrinsecus

15-25 nervia. Inflorescentia in ramulis apicem versus oriens, ter-

minalis axillarisque, axi stellato-tomentosa, in ramulos primarios

vel secundarios floriferos divisa, bracteolis deciduis, multinerviis

tomentosis praedita. Flores per ramulum floriferum 2—8, ante an-

thesin 2—4 mm. longi, pedicellis brevissimis circa 1 mm. longis

suffulti; sepala 3 majora quam altera 2, omnia ovata, apice acuta

vel fere, tomentosa; petala linearia, undulata, in alabastro torta,

puberula, extus grisea, intus brunnea; stamina 28—32 altitudine

variabilia, in appendiculas apice 1-5 ciliatas terminata. Fructus

breviter stipitatus, laciniis puberulis praeditus, 3 accrescentibus

spathuloideis 5-6 cm. longis 1.25 cm. latis, alteris 2, minoribus.

angustioribus, linearibus, apice acutis vel obtusis, 3 cm. longis,

3—4 mm. latis. Nux ovoideus, sericeo-tomentosus, in stylopodium

abrupte angustatus, ad 1.2 cm. longus, 0.75 cm. in diam.

BORNEO: North Borneo: Sandakan: Bettotan (KEP 38,853=—

Puasa SAN 4,477, holotypus in KEP, isotypi in SAN & SING):

Tawao (Elmer 21,714, paratypus in SING).

This species produces relatively smooth, grey bole with irregu-

larly scaling or cracked bark, small buttresses, many werved

49]

Gardens Bulletin, S.

Shorea superba Sym. (fig. 1-14: Holotypus; fig. 15: Elmer 21,714)

1, Ramulus floriferus. 2, Paniculae fragmentum. 3, Alabastrum. 4—8, Sepala.

9-10, Petala. 11-12, Stamina. 13, Flos sepalis, petalis staminibusque

desumptis ut pistillum appareat. 14, Ovarium longitudinaliter discissum.

15, Fructus. (Amussis cum fig. 1 et 15 congruens; alterae fig. ampliatae).

492

— es eee

Vol. XVII. (1959).

leaves which are shining brown above and pale fawn or silvery

grey with minute stellate hairs below, silky tomentose flowers and

small fruits with minutely pubescent wings.

[The drawing was made by Mr. Wong Sze Moy under the direc-

tion of Symington, whose notes have been incorporated here. The

species has been listed as Shorea sp. ‘B’ (not to be confused with

Symington’s Shorea sp. “B” of Malayan Forest Records, No. 16,

1943, p. 95) in the herbarium of the Forest Research Institute,

Kepong. |

4. Shorea symingtonii Wood sp. nov. (§ Anthoshorea)

A S. virescente Parijs, cui affinis haec species differt nervis folio-

rum lateralibus pluribus, petalis latioribus, fructibus basi valde at-

tenuatis, lacintis eorum longioribus.

Arbor elata, circa 60 m. alta, cum ramulis juventute applanatis,

dense fulvo-tomentosis, dein fuscentibus parce tomentosis. Stipula

1-1.5 cm. longa, 2—5 mm. lata, tomentosa decidua. Folia oblonga

vel oblanceolata, 12-16 cm. longa 6—7 cm. lata, obtuso-apiculata

vel acuminata, interdum bullatula, basi rotundata vel saepe cordu-

lata, utrinque fulva vel brunnea, supra glabra, subtus secus costas

et nervos parce puberula, margine revoluta, utrinsecus 16—24 ner-

via, petiolis 2 cm. longis, brunneis, minute stellato-tomentosis suf-

fulta. Inflorescentia terminalis vel etiam in parte ramuli infra-ter-

minale axillaris, circa 20 cm. longa, laxe multiramosa, deciduo

bracteata, florum bracteolis deciduis, puberulis. Flores remoti,

ante anthesin oblongi vel lanceolati 10-13 mm. longi, 3-4 mm. in

diam., pubescentes, fusci, pedicellis ad 5 mm. usque altis, apicem

versus crassioribus, concoloribus, pubescentibus suffulti; sepala

lineario-lanceolata, coriacea, margine incrassata, leviter cymbifor-

mia, utrinque fusco-pubescentia, inaequalia, tria alteris duobus

maiora, omnia conformia; petala oblonga, apice obtusa vel trun-

cata multinervosa, chartacea, in partibus alabastri expositis albo-

tomentosa; stamina 15, per series duas disposita, interiora 5 altiora

quam exteriora 10. Fructus 5-laciniatus, basi valde angustatus,

pedicello 2-3 mm. longo suffultus, interdum glandulosus, laciniis 3

longioribus lineario-oblongis, apice sensim arcuato-acutis, 12—17

cm. longis, 1.6—2 cm. latis; alteris laciniis 2, linearibus, apicem

versus angustatis, acutis, 8-10 cm. longis 7-10 mm. latis. Nux

ovoideus, 2—2.5 cm. longus, minute pubescente rugosus.

BorNEO: North Borneo: Sandakan, Sepilok (KEP 35,608 —SAN

16,522, holotypus in SAN, isotypi in KEP, SING et al; SAN

15481, paratypus in SAN, isoparatypi in KEP, SING, etc.).

493

Gardens Bulletin, §-

t / (

. AN SA Hy

aN \

\ Vie

Shorea symingtonii Wood (A—C: KEP 35,608; D-H: SAN 15481)

A, Ramulus floriferus. B, Flos. C, Ibid. petalis partim remotis ut dispositio

staminum appareat. D, Surculus cum bracteolis. E, Folium. F, Fructus.

G & H, Glandula ampliata.

494

Vol. XVII. (1959).

This large tree with shallow fissured and large buttressed bole,

has large leaves 12-16 cm. long, 6—7 cm. wide, often cordulate

at base, having 16-24 pairs of lateral nerves, the midrib and

lateral veins being minutely hairy below; the fruit 5-winged (3

large and 2 shorter ones) narrowed at base, shortly pedicelled. It

is an ally of S. virescens Parijs, but has more lateral nerves to the

leaves, flowers more apart, petals wider, fruits strongly attenuate at

base and bearing longer lobes.

[The drawing with fruit was prepared by Miss P. W. Wood at

Kew, the one with flowers has been made from the isotype speci-

men in Singapore by the Singapore artist.]

Originally this species was separated as new by Symington but

was not named. Some specimens have been distributed as S. sy-

mingtoniana which is a pre-occupied name. It has been listed as

Shorea sp. ‘L’ in the herbarium of the Forest Research Institute,

Kepong.

5. Hopea pentanervia Sym. ex Wood sp. nov. (§ Euhopea)

H. nutanti Ridl. similis, sed foliis minoribus glabrisque, nervis

eorum lateralibus paucioribus utrinsecus plerumque 5, floribus

minoribus, laciniis fructus angustioribus sat dissimilis.

Arbor circa 30 m. alta. Folia ovata vel elliptica, olivaceo-viridia,

6-9.5 cm. longa, 2.5—5 cm. lata, caudato-acuminata, basi paulo

oblique rotundata vel breviter cuneata, utrinque glabra, supra

nitida, utrinsecus nervis lateralibus 4—7 plerumque 5, porrectis

subtus prominenter, supra dibiliter percursa, petiolis circa 1—1.3

cm. longis stipitata. Inflorescentia axillaris, solitaria vel interdum

binaria, in ramos secundarios floriferos ad 1.5 cm. usque longos

divisa, bracteis deciduis, axi‘omnino glabra. Flores in alabastro

ovoidei, cum pedicellis 4 mm. altis circa 2 cm. longi; sepala utrin-

que glabra, 2 majora petalis paulo longiora, lingulata, externe

longitudinaliter reflexa, olivaceo-brunnea, altera 3 minora ovata

vel fere orbicularia, albescentia, margine fimbriata; petala linearia,

apicem versus paulo angustata, obtusa, margine fimbriata, extus

albo-tomentosa, intus fere glabra vel minute puberula, flavescentia;

stamina 15, per series 2 disposita, inaequalia. Fructus bi-laciniatus,

omnino glaber, fusco-brunneus, pedicello 1.5—2 mm. longo stipi-

tatus; laciniis majoribus 2 spathulatis, 4.5—-5.5 cm. longis, 1 cm.

Jatis; minoribus 3, circa 5 mm. longis, quam nux cum stylopodio

brevioribus. Nux ovato-conicalis, breviter apiculatus, 6 mm. lon-

gus, 2.5 mm. in diam.

BorNEO: Sarawak, Baram (SAR 1257, holotypus in KEP,

isoholotypus in SAR). North Borneo, Bongawan—Paper (Mail,

SAN A. 1753, paratypus in SAN, isoparatypus in KEP & SING).

495

Gardens Bulletin, S.

AMM

HUY)

Hopea pentanervia Wood (A—D: A—1753 in SING; E-I: SAR 1257 in KEP)

A, Ramulus fructiferus. B, Folii fragmentum cum glandula (ampliatum).

C, Fructus. D, Nux. E, Ramulus floriferus. F, Flos. G, Ovarium cum sta-

minibus. H & I: Stamina.

496

Vol. XVI. (1959).

The leaves are glabrous, provided usually with 5 pairs of nerves

and often with domatial glands in the axils of basal nerves. Bole

often fluted, somewhat cracked and flaking. The species is similar

to H. nutans Ridl., but differs in smaller glabrous leaves having

fewer lateral nerves, smaller flowers, and in the fruit wings being

narrow.

[The drawing is made by the artist of the Botanic Gardens, Sin-

gapore, from the specimens in Kepong Herbarium.]

497

G. H. S. Wood, M. A., F. L. S.

(A tribute)

[Footnote:—Mr. G. H. S. Wood died on 6th May, 1957 in the Kuala Belait

hospital, Brunei State, as a result of burns received in an accident at his

camp. A short obituary notice will be found in the Malayan Forester Vol.

XX, p. 121. The following account of his last collecting trip was written.

shortly after his death. Apart from his numerous collections he left some

work nearly ready for publication, including field notes on all and com-

pleted descriptions of two of the new species published in this number. His.

contribution to the botany of Borneo was far greater than his published

work can ever indicate, and it is fitting that it should not be forgotten.

Since then Mr. P. S. Ashton has been collecting continuously in the forests

of Brunei until his departure in February 1960, and Dr. W. Meijer in 1959

joined the Forest Department in North Borneo as successor to Mr. Wood.

Thus the work of botanical collecting, which Wood did so much to further,

is being faithfully carried on after his departure.

B.E.S.]

“For Lycidas is dead, dead ere his prime

Young Lycidas, and hath not left his peer.”

IN 1957 the late Geoffrey Wood, then Forest Botanist, North

Borneo, had only recently returned from his first long leave in

England (where I had visited him working in the Kew Herbarium).

He wished to widen his field knowledge of North Borneo trees by

studying them in their Brunei habitat, and a tour was arranged

for that purpose. Peter Ashton, Forest Botanist, Brunei, (with

whom Geofirey travelled out in the Canton) and myself were pri-

vileged to be with him almost continuously for what were destined

to be the last few weeks of his life. You cannot, for long, live

under the same tarpaulin or share the same longhouse verandah

with a man without obtaining a clear idea of his character and

qualities; this brief tribute is an attempt to give some idea of

how he lived and worked and appeared to us during these last

days.

He arrived in Brunei on 13th March, bringing with him three

of the North Borneo forest staff. Two of these, Kapis and Kalukut

by name, were Dusuns from the upper reaches of the Kinabatangan

River; the third was a Christian forest guard from Jesselton. The

two Dusuns were expert tree climbers; given time they could climb

any tree, by building a bamboo ladder up it if necessary. Kapis,

the elder man, was frequently and proudly referred to by Geoffrey

as the finest jungle man in North Borneo and it was an exceedingly

498

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es te

SEE Ce |

enti: Seale

Vol. XVIT. (1959).

odd tree if neither Kapis, with his partly inherited and partly self-

taught native jungle lore, nor Geoffrey, with his patiently acquired

scientific knowledge, could “place” it.

The Dusuns were the most taciturn and inscrutable of men; they

hardly ever spoke, except in Dusun amongst themselves, and it

was impossible to guess most of the time what was in their minds.

Yet Geoffrey had won their confidence and their loyalty in a

remarkable way. He used to work very long hours, out in the

forest from 7 a.m. to 3 p.m., often 4 p.m., sometimes 5 or even

6 p.m.; a short break on return to camp; and then the laborious

work of trimming and putting away the day’s collections, usually

three to four hours’ work ending up by the light of the battered

old Tilley lamp that he carried. Yet they never groused or com-

plained. As a party they were perhaps the most efficient collecting

unit the forests of Borneo had seen up till then. Not since the

days of Beccari, nearly a century ago, had any botanist collected

the trees month after month over a period of years, as Geoffrey

did. And when it was all over, perhaps the most moving moment

in the funeral service was when these two Dusuns came forward

to the graveside, side by side, to sprinkle earth on the coffin.

Sporadic collecting had been done over the past 23 years in

Brunei by successive State Forest Officers and by the forest staff,

but only twice before had a qualified botanist toured in Brunei.

The late C. F. Symington came over from Malaya for two weeks

in 1938, and Mr. J. Wyatt-Smith spent a week here during his

visit to Borneo in 1954. This previous work had given us a general

picture of the Brunei forests. The plan for Geoffrey’s strong party

was intensive collecting at a Epenper of localities known to be of

special interest.

The first month was spent in the Temburong district, with camps.

at Kuala Belalong (submontane country in the upper Temburong),

Bangar (exploring the Biang ridge), Labu Estate (Bukit Patoi and

Bukit Peradayan and Labu swamp forest). The second month

(interrupted by Easter week, during which Geoffrey sailed for

Sandakan at the Kuala Belait interport regatta) was spent in the

Shorea albida forests behind Seria, and in the Andulau Forest

Reserve. Priority was given to the Dipterocarps, of which many

interesting collections (mostly sterile) were made, but trees of

other species were collected if in flower or fruit. The collections.

amount to some hundreds of species, which will take a long time

to classify and name. Several Dipterocarps new to Brunei (and

to science ?) were obtained, and by running through the sheets.

in the Brunei herbarium Geoffrey was able to clear up a number

of puzzles.

499

Gardens Bulletin, S.

The fates were against him on this trip. On the day we left

Brunei to start the tour his ankle swelled up alarmingly and turned

blue—delayed reaction from a blow from a hockey stick in a

game just before he left Sandakan. For over a week he could only

hobble a few steps, and had to confine his collecting to what he

could see from a dug-out. In botanically virgin country this must

have been terribly disappointing, but he never complained, and

derived some satisfaction from poring over the collections Peter

and J brought back daily from the surrounding ridges; his presence

in camp was invaluable, because his knowledge enabled him to

direct our efforts to best advantage. Any kind of dead leaf that

looked as if it might be a dipterocarp was picked up and brought

in. Geoffrey could tell infallibly (a) if it was a dipterocarp or not;

(5) if it was, to which group it belonged, and whether it was worth

collecting or not. Rest and hot fomentations eventually reduced

the swelling, and for the last day or two at that camp he was able

to climb the hills. For the rest of that month in Temburong dis-

trict the fates relented, and he was on top of his form; but during

the sailing regatta he contracted a chill, and was troubled with

a severe cold and fever during the last week in the Andulau Forest

Reserve.

Looking back over those two months, what impressions remain?

First, I think, must come his intense interest in his work, and his

already deep and extensive knowledge of Bornean plants. He was

extremely conscientious and thorough; however hot the day, how-

ever trying the leeches, painted flies, mosquitoes, sandflies, or

other pests, he resisted the temptation to scamp the work, and the

notes written on each collection were always full and detailed.

This attribute alone marked him out as a great field collector, in

the tradition of Beccari and Symington.

Next, perhaps comes his endless patience. His Dusun climbers

were sure but often slow, yet never did he betray the slightest

impatience with them, but was wont to jolly them along with

cheerful cries and imitations of the Helmeted Hornbill’s weird

notes. How inappropriate therefore that the fates should have

exacted so severe a penalty for a moment’s impatience with a

refractory camp fire.

Never again shall we see him, clad in jungle green, gazing up

at the tree-tops in search of fertile material, with the binoculars

he always carried—sometimes standing, but as often as not lying

flat on his back on the leechy forest floor. Never again shall we

see him sitting on the ground, puffing away at his pipe, writing

those endless notes in his field book on some specimen which his

climbers were busy collecting. On a long trip such as this he let

500

Vol. XVIT. (1959).

his beard grow, I suspect chiefly because the few minutes saved

by not shaving every day could be devoted to writing some extra

notes. Never again shall we see him snipping away with a large

pair of clippers, as he used to do hour after hour trimming his

material to give at least a dozen specimens; for he collected not

only for Sandakan Herbarium but for the great herbaria of the

world, Singapore, Kepong, Kew, Leiden, Arnold Arboretum, and

the rest, to which duplicates were distributed.

It is difficult to write of his other interests; chiefly because we

were so enveloped and caught up by his enthusiasm on this trip,

and usually so tired out by the time the last specimen was put to

bed, that I cannot remember any subject other than botany being

discussed at length. We shared a common interest in mountains

(when in England he had asked me to propose him for member-

ship of the Himalayan Club) and sometimes talked of the Cuillins.

or the Himalayas, or the Silvretta. He was interested in photo-

graphy and carried two cameras, one for monochrome and one for

colour, but these were chiefly used to picture the barks of trees

or colourful figs.

“Lycidas is dead, dead ere his prime.” Had he been spared and

devoted enough years to the work he might have become a great

botanist; but he was cut off when he was only just beginning to

tackle taxonomic work and describe new species. I have already

suggested that he was a great collector, and he had the luck to

be in the field during 1955, one of the great dipterocarp flowering

and fruiting years, when he obtained many species in flower or

fruit for the first time in botanical history; the magnificent

material he collected and distributed round the world will be his

memorial. According to Milton,

“Fame is the spur that the clear spirit doth raise

To scorn delights and live laborious days.”

{ doubt whether botanist explorers follow their arduous calling in

the hope of achieving fame so much as from sheer curiosity and

love of travel. However Geoffrey might have answered the question

we do not know, for he has now joined the select few, including

George Forrest and Reginald Farrer. who died in harness, or (in

the words inscribed on Farrer’s lonely tombstone in far Northern

Burma), who “died for love of beauty, and in search of rare plants.”

B.E:S.

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OTHER PUBLICATIONS OF THE BOTANIC GARDENS

SINGAPORE

1. Annual Report 1875*

Reports for many years 1886 onward remain available.

Price variable.

2. The Agricultural Bulletin of the Malay Peninsula (Series I).

Nos. 1-9, 1891-1900.

Only Nos. 3, 5, 7, 8, 9 available at 20 cents each.

3. The Agricultural Bulletin of the Straits and F.MLS. (Series I).

Vols. 1-10, 1901-1912, monthly issues.

All available except Vol. 1, pts. 6 and 11.

At 50 cents each or $5 per volume.

4. The Gardens’ Bulletin, Straits Settlements (Series III).

Vol. 1-11 (3) January 1912—August 1941.

Vol. 1 (1-5) January-May 1912 is issued under title of

Agricultural Bulletin of the Straits and F.M.S.

All parts available, except vol. 2 (2, 3, 6-11) and 6

(reprinting in press).

Prices variable according to size.

5. The Gardens’ Bulletin, Singapore (Series IV).

Vol. 11 (4) September 1947*.

All parts available. Prices variable. Issues are irregular

and volumes are not related to calendar time. Subscrip-

tion will be entered against deposit.

Available on exchange.

6. Malayan Garden Plants: a series of eleven booklets cach

illustrating 10 useful garden plants.

Price 50 cents each.

7. An illustrated guide to the Botanic Gardens, me 20 3

Price $1.

8. Materials for a Flora of the Malay Peninsula, Monocotyle-

dons.

. Vols. I, I, Ill remain available.

9. 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.

10. (a) Malayan Orchid Hybrids by M. R. Henderson and G. Hi.

Addison. Price $21.

(b) Malayan Orchid Hybrids, Supplement I by G. H. Addison

(In press).

11. Wayside Trees of Malaya by E. J. H. Corner, 2 vols.

Price $25.

items 1—8 can be obtained from the Botanic Gardens,

Singapore.

Items 6 and 9-11 can be obtained from the Government

Printer, Singapore.

Prices are quoted in Malayan Dollars.

Postage overseas is extra.

7832—55C—6/60

GARDENS BULLETIN

SINGAPORE

JV OV OV OV YU OVUM UMVIM MIVA APYAMUAOI OVO

Vol. XVII 1958 — 1960

OMY VIVO OV VOW VMOU DOU UYU UMM PU PUP MPU?

Part 1 — Ist November, 1958.

Part 2 — 5th December, 1959.

Part 3 — 18th May, 1960.

To be purchased at the Botanic Gardens, Singapore

Part 1, $12.50, Part 2, $12.50, Part 3, $12. 50.

| Complete Volume $25 —

(Overseas postage extra)

Prices quoted in Malayan currency

Published by Authority

PRINTED BY LEE Kim HENG, ACTING GOVERNMENT PRINTER, SINGAPORE.

1960

AMANAAAAADOAAD 5 da! . cae ion

DB ADAVBABAAA AAPA AVAPAAAA. PAA A AAPAAAAAAAAPAUAAAAAAA, PUPUPAMNAI

CORRIGENDA—Volume 17

Note: Part 2 page headings—for Vol. XVII (1958) read Vol. XVII (1959)

Part 3 page headings—for Vol. XVII (1959) read Vol. XVII (1960)

line 37: for Vittaria flexuosa Fee read Vittaria scolopendrin

(Bory) Thw.

Section heading: for Ser. Validae (Miq.) read Ser. Subvalidae

Corner (Miq.) Corner

Section heading: for Subsect Benjamina read Subsect Benjamina

Mig. (Miq.) Corner

penultimate line: for Smithii read Smithii

last line: for Wall. ex. Mig. read (Wall. ex. Miq.)

King

line 17: for Adenospermal read Adenosperma

line 9: for inconspicula read _inconspicua

ne’ TY; for minanassae read _minahassae

line 3 for retuso read retusto

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INDEX - VOLUME 17

References are given

(a) Names of authors of papers in capitals:

(b) New taxa and binomials in bold print:

Abarema nediana Kosterm. 8

Aglaomorpha heraclea (Kze.) Copeland 256

Agriculture in Malaya 187

ALLEN, B. M.: 251, 253

Alseodaphne magnifica Kosterm. 3

Ampelopteris prolifera (Retz.) Copeland 261

Ararocarpus Scheffer 93

Arcypteris 365

Ardisia 279

A. breviramea Merr. 291

calcicola Furtado 298

calophylla Furtado 298

disticha A.D.C. 281

ferox Furtado 279

— ferruginea Mez 287

foliosa Furtado 302

fuliginosa Bl. 281

gambeleana Furtado 293

goodenoughii Furtado 291

iteophylla Ridl. 306

korthalsiana Scheff. 281

laevigata Bl. 283 —

littoralis Andr. 298

miqueliana Scheff. 287

nurii Furtado 287

pachysandra (Wall.) Mez 282

pachysandra (Wall.) Mez sensu Ridley

. 279

porosa Clarke 309

purseglovei Furtado 300

praetermissa Furtado 283

rudis Sinclair 287

A. sessilis Scheff. 289

singaporensis Ridl. 286

tetrasepala King & Gamble 310

teysmanniana Scheff. 282

teysmanniana Scheff. sensu Ridley 281

tiomanensis Furtado 296

tomentosa Pres]. 281

tumida Furtado 304

vaughanii Rid]. 302

Athyrium 366

A. amplissimum (Baker) Holttum 270

boryanum (Willd.) Tawaga 270

curtisii Holttum 269

dilatum (BI.) Milde 271

japonicum (Thunb.) Copeland 269

velutinum (Holtt.) Holttum 269

Bamboos: pioneer work on 193

Banyan 382

Blechnum fraseri (A. Cunn.) Luerssen

var. philippinense Christ. 263

B. orientale Linn. 263

Botrychium daucifolium Wall. 251

Brownlowia purseglovei Kosterm. |

BURKILL, H. M.: 201

BURKILE, i? fie 73

Burkill, I. H.: Biographic notes on 341, 35(

Christiopteris tricuspis (Hook.) Christ. 259

Colysis acuminata (Baker) Holttum 256

Coniogramme fraxinea (Don) Diels 271

503

Conservation 151, 201

CORNER, E. J. H.: 209, 368, 405, 416, 442

Cornopteris 366

Covellia albipila Miq. 408

C. cyrtophylla Wall. 473 .

zollingeriana Mig. 473

Ctenitis 365

Ctenitopsis 365

Ctenopteris celebica (Bl.) Copeland 260

Cyathea excavata Holttum 255

Cystopteris tenuisecta (Bl.) Mett. 268

Dipterocarpaceae, New Species from Borneo

486

Dipterocarpus exaltus van Slooten 486

Dryoathyrium 365

Dryopolystichum 365

Dryopteris 361, 366

D. subarborea (Bak.) C. Chr. 267

sparsa (Don) O. Ktze. 267

Elaphoglossum decurrens (Desv.) Moore 267

E. melanostichum (Bl.) Moore 267

Erythrophloeum densiflorum (Elm.) Merr. 5

E. unijugum Airy Shaw 5

Ficus 368

generic subdivisions

Albipilae Corner (Subseries) 406, 408

Americana Miq. (Section) 375

Auratae Corner (Series) 420, 434

Auratae (Subseries) 420,434

Austrocaledonicae Corner (Series) 407, 414

Basitepalae Corner (Subseries) 419, 429

Benjamina (Mig.) Corner (Subsection) 374,

395

Benjamineae Mig. (Series) 374, 395

Callophylleae Corner (Series) 374, 396

Cariceae Corner (Series) 418

Caulobotryae (Mig.) Corner (Series) 371

Conosycea (Mig.) Corner (Section) 371,

379

Conosycea (Subsection) 372, 379

Copiocseae Corner (Series) 445, 453

Crassirameae Corner (Subseries) 373, 284

Cuneifoliae Corner (Subseries) 420, 433

504

Gardens Bulletin, S.

Cuspidatae Miq. (Series) 448, 479

Cyclanthereae Corner (Series) 375

Cyrtophylieae Corner (Series) 446, 473

Dehiscentes Corner (Subseries) 419, 433

Dictyneuron Corner (Subsection) 373, 388

Drupaceae Corner (Series) 372, 380

Drupaceae (Subseries) 372

Dubiae Corner (Series) 373

Eriosycea (Miq.) Corner (Subsection) 419,

429

Eriosyceae Corner (Series) 419, 429

Eriosyceae (Subseries) 419, 429

Krythrogyneae Corner (Series) 418, 420

Exasperatae Corner (Series) 446

Ficus (Subgenus) 416, 417, 442

Ficus (Section) 416, 417

Ficus (Subsection) 417, 420

Fibrosifoliae Corner (Series) 448, 480

Galoglychia (Gasp.) Endl. (Section) 375 |

Glaberrimae Corner (Series) 373, 388

Hesperidiiformes Corner (Subseries) 375,

403

Heterophylleae Corner (Series) 446

Indicae Corner (Subseries) 372, 381

Leucogynae Corner (Section) 371

Malvanthera Corner (Section) 375, 401

Malvanthereae Corner (Series) 375, 401

Malvanthereae (Subseries) 375, 402

Minutuliflorae Sata (Series) 448, 480

Monandreae Corner (Subseries) 420, 438

Nervosae Corner (Series) 407, 409

Oreasycea (Mig.) Corner (Section) 406

Orthoneurae Corner (Series) 371

Palaeomorphe (King) Corner (Subsection) —

446, 475 ;

Pallidae Miq. (Series) 447, 475

Perforatae Corner (Series) 374, 394

Phaeopilosae Corner (Series) 445, 450 ~

Pharmacosycea Mig. (Subgenus) 405

Pharmacosycea (Section) 406

Platypodeae Corner (Subseries) 375, 402 |

Podosyceae Corner (Series) 418, 426

Podosyceae (Subseries) 419, 426

Prostratae Corner (Series) 444, 449

Pseudopalmae Corner (Series) 418

Pungentes Corner (Series) 444,450

Religiosae Mig. (Series) 371

Vol. XVI. (1958-1960)

Rivulares Corner (Series) 418

Scabrae Miq. (Series) 445, 458

Sinosyceae Corner (Series) 418, 420

Subulatae Corner (Series) 447, 478

Subvalidae (Mig.) Corner (Series) 373, 389

Superbae Corner (Series) 371

Stilpnophyllum Endl. (Section) 374

Sycidium Mig. (Section) 442, 443

Sycidium (Subsection) 444, 449

Trichosyceae Corner (Subseries) 419, 430

Urostigma (Gasp.) Mig. (Subgenus) 368,

370

Urostigma (Section) 371, 376

Vasculosae Corner (Series) 406, 408

Vasculosae (Subseries) 406, 408

Validae Migq. (Series) 372, 379

Varinga (Mig.) Corner (Subsection) 446,

473

Zygotricheae Corner (Subseries) 372

Ficus: specific divisions

F. abelii Mig. 419

acamptophylla King 390

acanthophylla Summerhayes 455

acrorrhyncha Summerhayes 395

aculeata A. Cunn. 471

var. micrantha (Mig.) Benth 471

acuminatissina Mig. 481

adami Elm. 388

alba Reinw. 429

albipila (Miq.) King 407, 408

var. glabra Corner 408

alongensis Gagn. 376

ampelas Burm. f. 459

var. linearis Corner 460

obversifolia Miq. 456

sonorensis (King) Corner 460

anastomosans- Wall. 477

andamica Corner 454

androbrota Summerhayes 458

androchaete Corner 438

anggica Diels 455

angustissima Merr. 458

annamensis Gagnep. 434

annulata Bl. 372

argentea Bl. 378

armiti King 448

F. arnottiana Migq., var. subcostata Corner

379

aspera Forst. f. 444, 446, 472

asperior Mig. 460

asperiuscula Kunth & Bouché 473

asymmetrica Lévl. & Vant. 473

augusta Corner 403

auranticarpa Elm. 380

aurata Miq. 420

var. brevipilosa Corner 437

longipilosa Corner 437

palawensis Corner 437

pedunculata Corner 438

auriculifera Merr. 480

aurita Reinw.

var. auriculifera (Merr.) Corner 480

celebica (Reinw.) Corner 480

austrocaledonica Bur. 407

var. balansaeana (Bur.) Corner 415

balansaeana Bur. 415

balica Mig. var. colfsii Corner 456

banahaensis Elmer 433

beecheyana Hook. & Arn.

var. koshunensis (Hayata) Sata 427

behrmanniana Diels 412

bengalensis Linn. 372, 381

benjamina Linn. 374

var. bracteata Corner 396

benjaminoides Corner 400

bennettii Seem 414

biglandulosa Mig. 460

blepharisepala Warb. 460

binnendykii Miq.

var. coriacea Corner 395

cupulata Corner 395

latifolia Corner 395

brachypoda Miq. 402

bruneiensis Corner 435

brunneo-aurata Corner 436

burkillii Ridl. 423

ealcicola Corner 392

callophylla Bl. 374

var. leytensis Corner 396

malayana Corner 396

minor Corner 397

calophylloides Elm. 395

505

F. cambodica Gagnep. 434

camiguinensis Merr. 433

cardinalicarpa Elmer 429

carica Linn. 417, 418

caroli Diels 455

cartacea Wall.

var. lanceolata Corner 433

caudata Griff. 420

caudatifolia Warb. 480

caudiculata Trim. 379

caulobotrya Mig. 378

caulobotrya (Miq.) Miq. var. fraseri Mia.

377

caulocarpa Miq. 371

var. dasycarpa Corner 378

cauta Corner 461

celebensis Corner 478

celebica Reinw. var. kunstleri King 481

chartacea Wall. 420

chincha Roxb. 420

chrysochaete Corner 466

clavata Wall. 420

clementis Merr. 385

collinsii Elmer 462

colossea F. Muell. 408

complexa Corner 445, 450

concinna Mig. var. subsessilis Corner 376

confertifolia Merr. 426

conglomerata Roxb. 449

conjugata Miq. 471

consociata Bl. 372

copiosa Steud. 445

var. pubescens Corner 455

copiosa Steud. sensu Ridley 453

cordata Ridley 430

cordatula Merr. var. sericea (C. B. Rob.)

Corner 381

cordifolia Bl. 408

coronata Reinw. 473

coronata Spin. 472

costata Ait. 379

cotoneaefolia Vahl. 381

cotonifolia Stokes 381

crassiramea Miq. 373, 384

var. brevicupulata Corner 385

celebica Corne 385

clementis (Merr.) Corner 385

patellifera (Warb.) Corner 385

506

-Gardens Bulletin, S.

crinobotrya Corner 469

var. solomonensis Corner 470

f. glabrior Corner 470

cucurbitina King, var. eubracteata Corne1

381

cumingii Miq.

var. androbrota (Summerh.) Corner 458

angustissima (Merr.) Corner 458

worcesteri (Merr.) Corner 458

cuneata Lévl. & Vant. 427

cunia B. Ham. 449

var. conglomerata (Roxb.) Kurz 449

curtipes Corner 397

cuspidata Reinw. 448, 479

var. sinuata King 479

cyanus Lévl. & Vant. 428

cylindrocarpa Diels 404

cyrtophylla Wall. 446, 473

decaineana Miq. 477

delosyce Corner 391

var. cbtusa Corner 391

deltoidea Jack 418, 420

var. angustifolia (Mig.) Corner 420

f. angustissima Corner 421

arenaria Corner 421

bilobata Corner 422

borneensis Corner 422

f. subhirsuta Corner 422

intermedia Corner 423

kunstleri (King) Corner 423

lutescens (Desf.) Corner 423

f. longipedunculata Corner 424

subsessilis (Mig.) Corner 424

motleyana (Miq.) Corner 424

oblongoneura (Miq.) Corner 424

peltata Corner 424

trengganuensis Corner 425

depressa Bl. 380

dichroa Summerh. 458

diversifolia Bl. 423

var. kunstleri King 423

longipedunculata Mig. 424

lutescens (Desf.) King 423

ovoidea (Jack) King 421

dodonaeiformis Gagnep. 425

doormaniana Diels 412

Vol. XVII. (1958—1960)

F. drupacea Thunb. 372, 380

var. auranticarpa (Elm.) Corner 380

glabrata Corner 380

pedicellata Corner 381

pubescens (Roth.) Corner 381

dubia Wall. 373

dumosa King 430

duriuscula King 455

elastica Roxb. 374

edulis Bur. 414

elmeri Merr. 465

endospermifolia Corner 434

episima Corner 394

erecta Thunb.

var. beecheyana (Hook. & Arn.) King

f. koshunensis (Hayata) Corner

427

erecta,

f. sieboldii (Miq.) Corner 427

sieboldii (Miq.) King 427

esmeralda Bailey 477

esquirolli Lévl. & Arn. 428

eucaudata Elmer 480

eugenioides F.v. Mueller, var. puberula

Benth. 402

eulampra K. Schum. 455

eumorpha Corner 439

var. subglabra Corner 440

exasperata Vahl 440

exasperata Roxb. 460

fairchildii Backer 395

fastigiata Elmer 464

fenicis Merr. 476

fieldingii Miq. 426

fiskei Elmer, var. multinervia Corner 459

firmula Miq. 477

forstenii Miq. 385°

var. pacifica (Elm.) Corner 385

umbobracteata (Elm.) Corner 385

villosa Corner 385

fraseri Miq. 471

fulva Reinw. var. timorensis Corner 432

garciniifolia Miq. 395

gaspatriniana Mig. 428

var. esquirolii (Lévl. & Vant.) Corner428

laceratifolia (Lévl. & Vant.) Cor-

ner 428

viridescens (Lévl. & Vant.) Cor-

ner 428

507

F. gelderi Mig. 394

gemella Wall. 426

gibbosa Bl. 447, 476

var. parasitica (Willd.) King 476

rigida Mig. 476

rigida (Bl.) Val. 476

gibbsiae Ridley 439

glabella Bl. 377

glaberrima Bl. 373

var. bracteata Corner 388

siamensis Corner 388

glandifera Summerhayes, var. brachysyce

Corner 402

glandulifera (Wall.) King 419

var. camiguinensis (Merr.) Corner 433

villosa Corner 433

globosa Bl. 380

goniophylla Corner 461

gracilipes Bailey 376

gracillima Diels 478

granatum Forst. f., var. minor Corner 414

gratiosa Corner 408

var. caudata Corner 409

grewiaefolia Kunth. & Bouché 473

griseifolia Corner 456

grossularioides Burm. f. 419

var. robusta Corner 429

stenoloba Corner 430

eryllus Corner 468

gul Laut. & K. Schum. 452

var. lasiocarpa Corner 452

solomonensis Corner 452

guyeri Elmer

var. minimaefolia Sata 464

sibuyanensis (Elmer) Corner 460

validicaudata (Merr.) Sata 464

habrophylla G. Bennett 414

hadroneura Diels 413

halmaherae Corner 431

henneana Miq. 376

henryi Diels 418

hesperidiiformis King 375

var. myrmekiocarpa (Summerh.) Corner

403

heterophylla Linn. f. 446

F. heteropleura Bl. 480

var. hirta Corner 480

mindanaensis (Warb.) Corner 480

hillii F. M. Bailey 398

hirta Vahl. 419

var. appressa Corner 430

brevipila Corner 430

dumosa (King) Corner 430

malayana Corner 430

squamosa Corner 431

hispidulosa Elmer 448

hololampra Diels 462

hombroniana Corner 410

hookeri Miq. 378

hookeriana Corner 378

hypsophila Miq. var. angustata Miq. 481

Ficus indica foliis mali cotoneae 382

Ficus indica Theophrasti 382

iidaiana Wilson 418

imbricata Corner 467

inaequifolia Elmer 477

inaequilatera Ridl. 473

inaequipetiolata Merr. 437

indecora A. Cunn. 471

indica Linn. 381, 389

var. gelderi (Mig.) King 394

infectoria (Miq.) Miq. 376

infectoria Willd. 376

insularis Miq. 477

irisana Elmer, var. validicaudata (Merr.)

Corner 464

irregularis Miq. 478

ischnopoda Miq., var. subcylindrica Cor-

ner 429

kajewskii Summerhayes 472

keyensis Laut. & K. Schum. 452

kinabaluensis Stapf. 423

kiellbereii Corner 411

koutumensis Corner 449

korthalsii Mig. 389

var. beccariana King 389

koshunensis Hayata 427

krausseana Rechinger 455

laceratifolia Lévl. & Arn. 428

lamprophylla Laut. & K. Schum. 455

lancifolia Miq. 481

lawesii King 388

leptocalama Corner 483

FP:

508

Gardens Bulletin, S.

leptodictya Diels 463

leptogramma Corner 473

leptorhyncha Koord. & Val. 473

leuchotricha Miq.

var. megacarpa F.v. M. 402

sessilis Corner 402

litseifolia Corner 433

lowit King

var. borneensis Corner 389

minor Corner 389

lutescens Desf. 423

luzonensis Merr. var. imberbis Elmer 426

macilenta King

var. gibbsiae (Ridl.) Corner 439

ilicifelia Corner 439

maclellandi King, var. rhododendrifolia

Corner 392

macrophylla Desf. 374, 375

macropoda Miq. 426

macropoda Kurz. 454

macropodocarpa Lévl. & Vant. 429

madhueifolia Corner 412

madhurensis Mig. 453

var. angustifolia Corner 453

magnifica Elmer 477

mallotoides Vai. 408

mangiferifolia Laut. & K. Schum. 412

manilensis Warb. 452

maxima P. Mill. 406

mearnsii Merr. 427

melinocarpa Bl., var. hoelolampra (Diels)

Corner 462

micrantha Miq. 471

microcarpa Linn. f. 397

var. eubracteata Corner 398

hillii (Bailey) Corner 398

latifolia (Miq.) Corner 398

f. pubescens Corner 398

naumannii (Engl.) Corner 398

saffordii (Merr.) Corner 399

microstoma Wall. 394

microtricherinos Backer 408

microtus Miq., var. borneensis Mig. 481

midotis Corner 482

mindanaensis Warb. 480

mollis Migq., var. albipila Mig. 408

motleyana Miq. 424 ~

muntia Link 472

Vol. XVII. (1958-1960)

F. montana Burm. f. 453

var. purpurascens (Bl.) Corner 453

mooniana King 379

myriocarpa Miq. 450

myrmekiocarpa Summerh. 404

mysorensis Heyne 381

var. pubescens Roth 381

naumannii Engl. 398

nemoralis Wall. 426

var. gemella (Wall.) King 426

fieldingii King 426

trilepis King 426

neriifolia J. E. Sm. 426

var. fieldingii (Miq.) Corner 426

nemoralis (Wall.) Corner 426

trilepis (King) Corner 426

nervosa Heyne 406, 407

nhatrangensis Gagnep 429

nitida auct. 397

nubigena Diels 455

oblongoneura Mig. 424

obliqua Forst. f.

var. petiolaris (Benth.) Corner 402

puberula (Benth.) Corner 402

obscura Bl. 448, 480

var. angustata (Miq.) Corner 481

borneensis (Miq.) Corner 481

kunstleri (King) Corner 481

scaberrima (Bl.) Miq. 482

obtusifolia Roxb. 397

oleaefolia King 418

var. dodonaeiformis (Gagnep.) Corner

425

epiphvtica (Elmer) Corner 425

memaecylifolia Corner 425

myrsinoides Corner 425

riparia Corner 426

oleracea Corner 465

var. pugmans Corner 466

onusta Wall. 380

opposita Mig. 471

var. indecora (A. Cunn.) Corner 471

micrantha (Miq.) Corner 471

orbicularis A. Cunn. 471

orthoneura Lévl. & Vant. 371

otariophylla Diels 478

ovoidea Jack 421

paramorpha Corner 440

F. pachyphylla Merr. 396

parasitica Willd. 476

parcelli Veitch 472

pachystemon Warb. 412

pachysycia Diels 409

pacifica Elm. 385

palaquiifolia Corner 399

palawensis Merr. 385

palmata Forsk 418

paracamptophylla Corner 387

parkinsonii Hiern 376

pateliata Corner 400

patellifera Warb. 385

pedunculosa Miq. 419

var. confertifolia (Merr.) Corner 426

imberis (Elmer) Corner 426

inacropoda (Miq.) Corner 426

mearnsii (Merr.) Corner 427

segaarensis (Engl.) Corner 427

velutina Corner 427

pellucida-punctata Griff. 394

pereng Steud. 474

philippinensis Miq. 477

var. sessilis Bur. 478

pilosa Reinw. 380

pinkiana F. v. M. 477

pisifera Wall. 481

pisocarpa Bi. 374, 394

platypoda A. Cunn. 375, 403

var. petiolaris Benth 402

polygramma Corner 399

porphyrochaete Corner 451

portus-finschii Warb. 455

praetermissa Corner 474

primaria Corner 457

pritzelii Warb. 376

procera Bl. 386

var. crassiramea (Miq.) King 384

prolixa Forst. f.

var. subcordata Corner 378

proteus Bur. 471

pruniformis Bl. 380

pseudojaca Corner 413

pseudopalma Blanco 418

puberula A. Cunn. 403

pubilimba Merr. var. evata Corner 384

pubinervis Bi.

var. diandra Corner 410

509

Gardens Bulletin, S.

F. pubinervis BI. F. sibuyanensis Elmer 460

sibulanensis (Elm.) Corner 411

f. sibulanensis (Elm.) Sata 411

pungens Reinw. 445, 450

purpurascens Bl. 453

pustulata Elm. var. lanceifolia (Sata)

Corner 429

quercifolia Roxb. 453

regnans Diels 398

religiosa Linn. 370, 371

reticulata Thunb. 476

reticulatissima S. Moore 456

retusa Linn. 393

var. borneensis Corner 393

rigo (Bailey) Diels 398

rhododendrifolia Miq. 392

rhodocarpa Summerhayes 455

rhomboidalis Lévl. 428

riedelii Miq., var. minor Corner 459

rigida Bl. 476

rigida Miq. 387

rigo F. M. Bailey 398

rivularis Merr. 418

rostrata Lam. 479

var. urophylla (Wall.) Koord. & Val.

480

rubrocuspidata Corner 483

rubromidotis Corner 483

rudis Miq. 452

rumphii Bl. 371

saffordii Merr. 399

saxophila Bl., var. sublanceolata Miq. 377

scaberrima Bl. 482

scabrella Roxb. 446

schlecteri Warb. 398

scobina Benth. 471

segaarensis Engl. 427

semicordata B. Ham. 444, 449

var. conglomerata (Roxb.) Corner 449

semicordata Miq. 465

sericea C. B. Robinson 381

setibracteata Elmer 477

setiflora Stapf. 420

var. adelpha Corner 440

puberula Corner 441

simplicissima Lour.,

var. annamica (Gagnep.) Corner 431

sibulanensis Elmer 411

510

sieboldii Miq. 427

silhetensis Mig. 428

var. annamica Gagnep. 431

sinuata Thunb. 479

subsp. cuspidata (Reinw.) Corner 479

var. oblonga Corner 479

smaragdina §. Moore 453

smithii Horne

var. robusta Corner 414

smithii 414

sororensis King 460

spathulifclia Corner 391

var. annamensis Corner 392

substipitata 392

stenocarpa F. v. Muell. 471

stenophylla Hemsl.

var. macropodocarpa (Lévl. & Vant.)

Corner 429

nhatrangensis (Gagnep.) Corner 429

stephanocarpa Warb. 472

sterrocarpa Diels 375

storckii Semm.

var. kajewskii (Summerh.) Corner 472

stupenda Miq.

var. minor Corner 384

subcordata Bl. 395

var. malayana Corner 396

subfulva Corner 432

var. villosula Corner 432

subgelderi Corner 386

var. subgelderi 380

rigida Corner 387

subincisa B. Ham. 418, 420

subnervosa Corner 411

subpedunculata Miq. 376

subpuberula Corner 403

subsidens Corner 454

subsumatrana Gagnep. 390

subtecta Corner 386

var. depressa Corner 386

subtrinervia Laut. & K. Schum.

var. doormaniana (Diels) Corner 412

subulata BI.

var. gracillima (Diels) Corner 478

sumatrana Miq.

var. circumscissa Corner 390

microsyce Corner 390

Vol. XVII. (1958—1960)

subsumatrana (Gagnep.) Corner | F. virgata Reinw. 477

390 var. philippinensis (Miq.) Corner 477

F. sundaica Bl. 373, 389 sessilis (Bur.) Corner. 478

var. beccariana (King) Corner 389 wassa Roxb. 455

superba Miq. var. nubigena (Diels) Corner 455

var. alongensis (Gagnep.) Corner 376 obversifolia (Mig.) Corner 456

_ enneana (Miq.) Corner 376 williamsii C. B. Robinson 425

swinhoei King 476 worcesteri Merr. 458

tanensis G. Bennett 414 pac ycianel

tenuicuspidata Corner 464

Bi aagee-Cormer 465 var. cylindrocarpa (Diels) Corner 404

thynneana F. M. Bailey 398 Flora Malesiana 161

tinctoria Forst. f. 447, 475 Forestry in Malaya 172

sub. sp. gibbosa (Bl.) Corner 476

var. gibbosa 476

rigida (Miq.) Corner 476 Founding and work in the early tropical

sub. sp. parasitica (Willd.) Corner 476 Gardens 126

var. anastomosans (Wall.) Corner 477 FURTADO, C. X.: 195; 276; 279, 350

parasitica 477 GILLILAND, H. B.: 82, 228

sub. sp. swinhoei (King) Corner 746

Gactoria 475 GILMOUR, J.-S. L215

tonsa Miq. 463 Gingers, Pioneer work on 192

var. aspera Corner 463

leptodictya (Diels) Corner 463

FOSBERG, F. R.: 219

Grammitis crispatula Holtt. 259

subcordata Corner 464 Gymnacranthera (Myristicaceae) 93

touranensis Gagnep. 431 Index to 120

trachypison Laut. & K. Schum. Gymnocarpium 366

var. pallida Corner 462 History of Botanic Gardens 125

travancorica King 388

trilepis Mia. 426 HOLTITUM, R..E-: 190, 330,-361

triradiata Corner 375, 401 Hymenandra iteophylla (Ridl.) Furtado 306

var. sessilicarpa Corner 401

Hopea pentanervia Sym. 495

tristaniifolia Corner 397

trivia Corner 427 Hypodermatium 365

truncata Miq. 393 Indonesia: Central Institute for Nature Re-

trymatocarpa Miq. 477 search 206

tuphapensis Drake. JOHNSON, A.: 312, 325

var. annamensis (Gapnep.) Corner 434 J

umbobracteata Elmer 385 Juncus prismatocarpus R. Br. 91

urophylla Wall. 480 Katou-alou 382

vahlii K. Schum. 398 KERN. J. H: 91. 273

validicaudata Merr. 464 4 “

vasceulasa Waill--406 Kew, Royal Botanic Gardens 155

venusta Kth. & Bouch. 379 KORIBA, KWAN: 11

villamilii Merr. 388 Koriba, Kwan, Obituary of 339

virens Ait. 376 ;

var. glabyella (BI.) Corner 377 KOSTERMANS, A. F. G. H.: 1

sublanceolata (Miq.) Corner 377 KUSNOTO SETYODIWIRYO: 206

511

LAM) Tek: 166

Lastreopsis 365

Leiden, Rijksherbarium 166

Leucostegia pallida (Meitt.) Copeland 263

Lindsaya cultrata (Willd.) Sw. 262

Loxogramme scolopendrina (Bory) Presl. 256

MALAYAN AGRICULTURIST: 187

Manila, National Museum of the Philippines

199

MATTSSON, L.: 225

Meiogyne virgata (Bl.) Mig. 93

Merinthosonus drynarioides (Hook.) Cope-

land 257

Micrecasia purseglovei Furtado 271

Microlepia puberula a.V. v.R. 262

Mohamed Nur bin Mohamed Ghous, Obi-

tuary of 331

Morus leucophylla Miq. 408

Nature Reserves 151

Ngadiman bin Haji Ismail, Obituary of 337

Nomenclature, International Code of 215

Obituaries 331

Orchids: Pioneer work on 190

Pacific Region, Importance of Biological Re-

search in the 219

Palms, Study of 195

Parathyrium Holttum 270

Penang Botanic Gardens 128

Peralu 382

Phellodendron burkillii van Steenis 357

Pleocnemia 365

P. conjugata (BI.) Pres]. 267

Polypodium prainii (Bedd.) C. Chr. 259

Polystichopsis 366

Polystichum 366

Pteridys 365

PURSEGLOVE, J. W.: 125

QUISUMBING, E.: 199

Gardens Bulletin, S.

Rainwater, pH of 244

Regenerating Forest 228

Research, Importance of 219

Reserves,

Aboriginee 202

Forest 1515°202

Nature 151, 203

Water Catchment 202

ROBINSON, R. A.: 244

Rubber, Founding of Malaya’s Rubber In-

dustry 134, 175

Rumhora 366

RUSSELL, T:\Aeeess

Salvinia cucullata Roxb. 272

SANTAPAU, H.: 341

Serianthes gigalobium Kosterm. 5

Shorea argentifolia Sym. 489

S. superba Sym. 491

symingtonii Wood 493

SINCLAIR, J.: 93, 96

Singapore Botanic Gardens 129

Sphagnum antarticum Mitt. 314

S. beccarii Hpe. 321

borneense Warnst. 321

cuspidatum Ehrh. 314

cuspidatulum C. Mull. 315

var. malaccense (Warnst.) Warnst. 315

trengganuense Johnson 316

flaccidifolium Dixon 316

holttumii Johnson 320

malaccense Warnst. 315

novo-guineense Fleis. & Warnst. 316

roseotinctum Johnson 320

subrecurvum Warnst. 317

sericum C. Mull. 317

Spilanthes anactina F. v. M. 273

S. chamaecaula A. H. Moore 273

pusilla Hook. & Arn. 273

urens Jacq. 273

Sympetalandra borneensis Stapf. 5

Synoecia diversifolia (Bl.) Miq. var. augusti-

folia Miq. 421

512

Vol. XVII. (1958—1960)

TAN KIAP SENG: 325

Taxonomy, Importance of Tropical 209

Tectaria 365

T. melanocauJon (Bl.) Copeland 268

Teijsmamicdendren sinclairii Kosterm. 6

T. holophyllum Kosterm. 8

_ Tetrardisia corneri Furtado 305

JT. porosa (Clarke) Furtado 309

T. tetrasepala (K. & G.) Furtado 310

Thelypteris beddomei (Bak.) Ching 261

T. brunnea (Wall.) Ching 260

Trichomanes motleyi v. d. Bosch 254

Tsiela 382

LN.Eos C0. 225

Urostigma accedens Miq., var. latifolia Miq.

398

U. balicum Miq. 395

binnendykii Mia. 373

brachypodum Mig. 402

canaliculatum Miq. 374

clusioides Mig. 373

concinnum Miq. 374

caulobotryum Miq. 378

crassirameum Miq. 373, 384

glabellum (Bl.) Mig. 374

glaberrima Bl. 373

U. hasseltii Mig. 373

513

infectorium Mig. 374, 376

lutescens Bl. 373

moritziana Mig. 374

obligoneuron Mig. 424

onustum Mig. 380

parvifolium Miq. 374

peracutum Mig. 373

platypodum Miq. 403

pseudorubrum Mig. 373

retusum (Linn.) Mig. 393

rigidum Mig. 387

subcordatum (Bl.) Mig. 395

sumatranum Migq. 373

sundaicum Bl. 373

timorense Miq. 374

tjiela Mig. 395

truncatum Mig. 393

virgatum Reinw. 373

wightianum Miq., var. majus Thw. 379

zollingerianum Miq. 373

VAN STEENIS: C. Guth 16h, 273.397

Varinga latifolia 382

Vittaria angustissima Holtt. 271

V. flexuosa Fee 271

WANTMAN, M. J.: 228

Wood, G. H. S.: A tribute 498

WYATT-SMITH, J.:

WYCHERLEY, P. R175

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