CONTENTS

Pace

Hart I: 15th October, 1962 (pp. 1-186).

Prowse, G. A.: Diatoms of Malayan Freshwaters - - I

Prowse, G. A.: Further Malayan Freshwater Flagellata - 1.05

Gilliland, H. B.: A Checklist of Malayan Grasses - - 147

Fijr i ado, C. X.: The Species of Brackenridgea in the Singapore

Herbarium - - - - - - 181

Furtado, C. X.: Name Changes in Bornean Araliads - - 185

Hart II: 31st October, 1962 (pp. 187-372).

Corner, E. H.: The Classification of Moraceae - - 187

Ashton, P. S.: Some New Dipterocarpaceae from Borneo 253

Whitmore, T. C.: Studies in Systematic Bark Morphology III.

Bark Taxonomy in Dipterocarpaceae - - 321

Hart III: 10th December, 1962 (pp. 373-415).

Johnson, Anne: On a small collection of Bryophytes collected

by Professor H. B. Gilliland in Eastern New Guinea - 373

Johnson, Anne: A shori Note on some Soil Algae from New

Guinea - - - - - . ' 375

Johnson, Anne: Precursory Studies on the Epiterranean Soil

Algae of Singapore and Malaya - - - -379

Corner, E. J. H.: Taxonomic Notes on Ficus L., Asia and

Australasia. Addendum 11 - - - - - 385

Index..

CORRIGENDA

Volume 19

280

last line of legend to

13:

281

line 17:

309

2nd paragraph, line 3

311

line 10 from bottom:

for Kep 37183 read S.A. 0690

after Beluran add Sarawak: S.A.

0690, Nanga Mujan, Batang Ai.

for Kep 36027 read Kep 30627

for 17023 read 17025

GARDENS’ BULLETIN

SINGAPORE

15th October, 1962

CONTENTS

Page

Prowse: Diatoms of Malayan Freshwaters - - - - 1

Prowse: Further Malayan Freshwater Flagellata - - - 105

Gilliland: A Checklist of Malayan Grasses - - - - 147

Furtado: The Species of Brackenridgea in the Singapore Herbarium 181

Furtado: Name Changes in Bornean Araliads - - - 185

To be purchased at the Botanic Gardens, Singapore

Price $12 (Malayan)

Published by Authority

PRINTED BY LIM BUN H\N, ACTING GOVERNMENT PRINTER

1962

THE

Gardens' Bulletin

SINGAPORE

Vol. XIX, Part I 15th October, 1962

<*£^> <*£?»> <*£^> V2?*> <*2?Y<*2^ '*&*> <^V<*J*»

Diatoms of Malayan Freshwaters

By G. A. Prowse

Tropical Fish Culture Research Institute, Batu Berendam, Malacca

Diatoms or Bacillariophyceae are a class of algae which have

probably been studied more thoroughly taxonomically than any

other class, but whilst there are extensive accounts of species from

Indonesia and other parts of S.E. Asia, little seems to have been

published on those species occurring in Malaya itself (I here take

as my geographical boundaries the Federation of Malaya and

Singapore). Apart from a paper by Ruth Patrick (1936) describ¬

ing some 73 species and forms from Perak, virtually nothing has

been published on Malayan species. This present paper is an

attempt to rectify that position, but as it includes only those spe¬

cies which I have seen and drawn myself it can in no way be

regarded as a complete account of Malayan freshwater diatoms.

Several of the species described by Ruth Patrick are absent from

this paper, whilst many others are described which are not men¬

tioned in her paper. A few others which I have seen have been

excluded because material has been too scanty and imperfect for

accurate identification. A number of species described here are

usually regarded as marine forms; whilst it is quite possible that

they are of marine origin, they were found in quite non-saline

water, fully alive and often dividing, and in some cases occurring

together with healthy, dividing specimens of such notorious halo-

phobes as the larger desmids. This fact is not really surprising.

Malaya is surrounded by seas which are frequently subjected to

torrential rainfall and the coastal waters must become heavy

diluted. At times some of the marine organisms must be existing

in pockets of almost pure water. Such organisms must be highly

adaptable to survive, and if they are carried inland by tidal rivers

they might quite easily survive or even multiply in freshwaters

inland. Several species of Coscinodiscus, Stephanodiscus , Cyclo-

tdl a and Nitzschia fall within this category.

Gardens Bulletin, S.

Whilst most species mentioned have been recorded elsewhere,

several new taxa are described, but these are forms which have

unmistakable characteristics. Some authors may decide others of

the forms described should be separated as new taxa, but my own

opinion is that the differences from previously described forms

are too slight to warrant such separation. Despite the vast amount

of work which has been done on the morphology of diatoms, we

still know far too little about the physiological development, the

genetics and ecology of them. Culture work and statistical studies

of morphological forms have shown that in some cases at least

so-called varieties are merely nutritional forms, whilst in other

cases they have emphasized the differences between closely similar

forms. For this reason I have thought it better not to separate off

varieties and forms simply on the basis of slight differences in

shape and the number of striae.

Since the illustrations in this paper are all original, done under

phase contrast with the aid of a camera lucida, any errors or in¬

accuracies are therefore my own responsibility. Identifications

have been carried out with the aid of standard works of A. Cleve-

Euler, Hustedt, particularly in “Die Tropischebinnengewasser”,

De Toni’s “Sylloge Algarum” and others, as well as several papers,

all of which are quoted in the bibliography. Schmidt’s Atlas and

Van Heurck’s works are not yet available in Malaya, and it has

not been possible to refer to them; all identifications have been

made through the works of other authors.

The Bacillariophyceae or diatoms form a well-marked class of

algae with very distinctive features, and are unicellular, or form

simple colonies. Each has a silicified cell wall or jrustule com¬

posed of two distinguishable “halves”, one of which overlaps the

other rather like a lid on a date box. The cell wall is composed of

pectic compounds, usually heavily impregnated with silica; cellu¬

lose has been reported, but accounts are rather contradictory,

and it is certainly not a regular constituent of the wall. Each “half”

consists of a somewhat flattened valve attached at the margins to

a connecting band, the girdle or cingulum . The cingulum extends

almost completely round the periphery of the valve, but there is

always a gap between the two ends. The longer overlapping valve

is known as the epivalve or epitheca, and the shorter as the hypo-

valve or hypotheca. Thus there are two characteristic views of the

diatom, the valve view with the valve side uppermost, and the

girdle view with the cingulum lying uppermost. In addition to the

cingulum some genera exhibit one or more intercalary bands. The

valves, are variously ornamented with coarse or fine striae, punc-

tae or areolae which radiate from a central point in the centric

genera, or bilaterally with respect to an axial field in the pennate

genera. The axial field, which may be symmetric or asymmetric

with respect to the longitudinal axis, usually appears as a clear

VoL XIX . (1962).

Text fig. 1

Views of diatoms (diagrammatic).

(i) oblique view of a centric diatom Cyclotella

(ii) valve view of a pennate diatom Pinnularia

(iii) girdle view of Pinnularia.

e, epivalve; h, hypovalve; g, girdle or cingulum ; r, raphe; p.n., polar

nodules; c.n., central nodule.

band between the striae. It may be perforated by a complex longi¬

tudinal slot or raphe. The raphe usually has thickenings in the

walls at each end and at the centre, the polar and central nodules .

In the absence of a raphe the axial field is known as a pseudo¬

raphe. The ornamentations are usually thin places in the silicified

wall, although small pores may perforate the walls within the areo¬

lae of some centric diatoms, and in a few pennate species there are

longer median or polar perforations.

The chromatophores, which are rarely absent, are usually nu¬

merous and discoid or irregular in the Centrales, and single, lobed

and perforate, or two and laminate in the Pennales. The colour is

usually golden brown, but sometimes bright green, especially in

specimens from organic bottoms in ponds and lakes. Pyrenoids,

always without a starch sheath, may or may not be present. The

photosynthetic pigments consist of chlorophylls a and c, usually

obscured by p- and e- carotene and five xanthophylls; chlorophyll

b is absent. Reserve foods include fats in the form of oil droplets,

volutin and leucosin, but never starch.

Many pennate diatoms exhibit motility, moving forward and

backward in the direction of the long axis of the cell. Movement

is confined to those species possessing raphes, and is ascribed to

the streaming of cytoplasm between the polar and the central

nodules through fissures on the outer wall faces.

Gardens Bulletin , S.

Reproduction is by cell division, statospores (cysts or endos-

pores) and auxospores which may be formed without any sexual

union, or by fusion either of amoeboid gametes, or of autogam¬

ous nuclei, and uni- or biflagellate microspores which have been

variously interpreted as zoospores or gametes. In the species so

far investigated the nuclei of the vegetative cells are diploid.

Diatoms are widely distributed in fresh and salt waters, often

becoming very abundant in colder waters and in colder seasons.

They may be free-floating, often pelagic, or they may be sessile

forming an important part of the lithophytic and epiphytic flora.

In flowing waters many species form copious gelatinous masses on

rock surfaces. Although diatoms are mostly aquatic, many species

occur within the soil and on wet surfaces of terrestrial substrates.

KEY TO THE GENERA

Since this paper must be very incomplete, only a key to the

genera is provided, the species being described under each genus

in alphabetical order.

Valves orientated about a central point

with ornamentation radial or concen¬

tric; raphe or pseudoraphe absent . . I. CENTRALES.

Valves orientated about a longitudinal

axial field, with the ornamentation

bilateral to it and not radiating from

a central point; raphe or pseudoraphe

present . . .. . . II. PENNALES.

I. ORDER CENTRALES

Cells solitary or united variously into filaments, dendroid or zig¬

zag colonies, free-floating or sessile; cell-shape discoid, cylindric

or irregular, with valves round, elliptic, polygonal or irregular;

ornamentation radial or concentric about a central point; raphe

or pseudoraphe absent, cells immobile, often with spine-like,

mamillate or long horn like projections; chromatophores usually

numerous, small, discoid and often irregular and laminate, with

or without pyrenoids; reproduction by cell formed by rejuvenes¬

cence, and enlargement of cells without sexual reproduction; some

genera known to have diploid vegetative cells and autogamous

sexual reproduction. Largely a marine order, with a few fresh¬

water species.

Vol XIX. (1962).

Key to Centrales

(i) Cells united into long filaments.(ii)

(i) Cells not united into long filaments; at most short chains (iii)

(ii) Valves circular with short teeth or spines at the periphery

1. Melosira

(ii) Valves elliptic with two very long spines at the ends of the

longer axis.6. Chaetoceros

(iii) Cells a long cylinder with a long terminal spine at each end

1. Rhizosolenia

(iii) Cells disc-shaped or squat cylinders with no long terminal

spine.(* v )

(iv) Valves with coarse areolae.3. Coscinodiscus

(iv) Valves punctate, not areolate . (v)

(v) Valves with a peripheral ring of short spines. No well marked

outer zone of striae or punctae.(vi)

(v) Valves without a ring of short spines, but with a well marked

outer zone of striae or punctae.2. Cyclotella

(vi) Peripheral spines very small, striae very fine, hardly visible

4. Thalassiosira

(vi) Peripheral spines larger and conical, striae well-marked

5. Stephanodiscus

MELOSIRA C. A. Agardh 1824

Cells cylindrical, united into long filaments; valve view circular,

with or without marginal teeth, flat or convex, usually with orna¬

mentations in two distinct concentric areas; girdles with sulcus or

groove, pseudosulcus or without either; when sulcus or pseudo¬

sulcus absent central area of girdle usually ornamented, when pre¬

sent central area usually smooth; chromatophores numerous,

small, discoid, frequently crowded.

Auxospore formation asexual or autogamous; biflagellate

zoospores reported.

Text fig. 2

Girdle view of Melosira showing a single cell with a semicell attached

at either end. e, epivalve; h, hypovalve; s, sulcus; p, pseudosulcus; g,

girdle of parent cell.

Gardens Bulletin , S.

1. Melosira granulata (Ehrenberg) Ralfs. [Syn.: Gaillonella

granulata Ehr.; G. procera Ehr.; G. marchica Ehr.; Melosira

ordinata Kuetz.; Orthosira punctata W. Smith.] Cells 16 p. long

X 10/a diameter to 28/a long X 5/a diameter; sulcus shallow

and acute; valves with short marginal teeth, but end cells bear¬

ing also a few long marginal spines; punctae of valve view

large, scattered; girdle view coarsely punctate, in parallel stria-

tions which are straight or only very slightly spiral, 8-11 striae

in 10/a. (PI. I, figs, a & b). The slender form has been separated

as var. angustissima, but observations here and in the river Nile

suggest that it may only be a growth form resulting from dec¬

reasing mineral supplies (Prowse & Tailing 1958).

Malaya Perak: Taiping Lakes.

Selangor: Common in established fish-ponds.

Negri Sembilan: Common in established fish-ponds.

Malacca: Common in established fish-ponds.

Singapore: Common in established fish-ponds; Bota¬

nic Gardens Lake.

Distribution worldwide.

2. Melosira italica (Ehrenberg) Kuetzing. [Syn.: Gaillonella

italica Ehr.; Melosira crenulata Kuetz.; M. crenulata Kuetz. var.

italica Grun.; M. crenulata Kuetz. var. tenuis Grun.] Gells 16/a

long X 9-10/a diameter to 28/a long X 5/* diameter; sulcus

shallow, acute; valves with prominent marginal teeth, alike on

end and intercalary cells, with no long spines; valve view very

finely punctate; girdle view finely punctate in spiral striations

which are sometimes undulate or intersecting, 10-20 striae in

10/a (PI. I, figs, g & h). The narrow form is often separated as

var. tenuissima, but as in M . granulata it may be only a growth

form.

Malaya Perak: Taiping Lakes.

Selangor: Common in fish-ponds.

Negri Sembilan: Common in fish-ponds.

Malacca: Common in fish-ponds.

Singapore: Common in fish-ponds; Botanic Gardens

Lake.

Distribution worldwide.

3. Melosira roeseana Rabenhorst. [Syn.: Orthosira spinosa Grev.;

Melosira spinosa (Grev.) Breb.]. Cells 16-20/a long X 20-

30/a diameter; sulcus wide, concave and rounded; valve with

prominent marginal teeth; valve view with punctae in radial

striation with 1-^ central spots; girdle view finely punctate, the

punctae being finer and closer together in the central area of

Vol. XIX . (1962).

the girdle, the striae being parallel and straight, 15-20 in 10/a

in the central area of the girdle, 10-15 in 10/a nearer the valve

edges. (PL I, figs, c & i).

Malaya Selangor: Gombak River, rare.

Found in moorland bogs and waterfalls in Europe, Indonesia

and the Philippines, and probably elsewhere in Asia.

4. Melosira ruettneri Hustedt. Cells 34-38/x long X 25-

26/a diameter; sulcus absent, the wall being straight, but in the

middle forming a pseudosulcus; cell wall very thick, thinner

near the centre of the valve and at the pseudosulcus, and com¬

posed of two membranes, the outer irregularly areolate, with

the areolae smaller towards the centre of the girdle, and towards

the centre of the valve, the inner very finely punctate, the punc-

tae being in radial striae on the valve face, and spiral on the

girdle view; pseudosulcus smooth. (Pl. I, fig. d).

Malaya Selangor: Feeder stream to Klang Gates Reservoir,

rare.

Recorded from Indonesia.

CYCLOTELLA Kuetzing 1834

Cells discoid or drum shaped, solitary or colonial within a muci¬

laginous envelope; when filamentous the cells not tightly inter¬

locking as in Melosira; valve view circular, or sometimes elliptic;

ornamentation of the valve in two very distinct concentric regions,

the outer zone radially striate or punctate, the inner zone smooth,

or irregularly and finely punctate; girdle view straight or undulate;

chromatophores numerous, small and discoid.

Auxospores formed singly within a cell; some species known to

be autogamous.

5. Cyclotella kuctzingiana Thwaites. Cells 10-28/a in diameter,

outer zone with fine radial striae, 15-18 in 10 /a about i dia¬

meter in length; inner zone with a few fine, scattered punctae.

(PL II, figs, i & j).

Malaya Pahang: A stream near Kuantan.

Distribution widespread in freshwaters.

6. Cyclotella meneghiniana Kuetzing. Cells 10-25/a in diameter;

outer zone broad, with thick well-marked striae, 7-9 in 10/a;

central zone smooth, or with fine radial striae. (PL I, fig. e;

PL II, fig. h). The form with a smooth central zone is sometimes

separated as var. rectcmgulata Grun.

Malaya Selangor: Common in fish-ponds.

Negri Sembilan: Common in fish-ponds.

Malacca: Common in fish-ponds.

Singapore: Common in fish-ponds; Botanic Gardens

Lake.

Widespread in distribution.

Gardens Bulletin , S.

7. Cyclotella striata (Kuetzing) Grunow. [Syn. Coscinodiscus

striatus Kuetzing.]. Cells 25-40/x in diameter; outer zone broad

with well marked striae, 10-12 in 10^, every second or third

stria marked with a prominent pore; sometimes very short, fine,

marginal striae in between the long ones; at a deeper focus

submarginal curves often visible between the pores; inner zone

with irregular swellings, matt in texture. (PI. I, fig. f; PI. II,

figs, b & g).

Malaya Pahang: A stream near Kuantan.

Malacca: Non-saline water of Malacca River.

Widespread, marine in coastal waters and in estuaries in

Europe, Asia and America.

COSCINODISCUS Ehrenberg 1838

Cells discoid, solitary or sometimes forming short, very loose

mucilaginous chains, valve view circular, ornamented with distinct

areolae, which may be radial, tangential or somewhat irregularly

arranged, the ornamentation forming one zone only; marginal

teeth present or absent. Mainly a marine genus.

8. Coscinodiscus antiquus (Grunow) A. Cleve. Cells 60-80/x in

diameter; areolae large, polygonal of equal size except at mar¬

gin, arranged excentrically, on lines curving away from the

centre, sometimes almost tangential; marginal teeth absent;

marginal striae marked, 6-10 in 10/x. (PI. II, fig. f).

Malaya Malacca: Malacca River in non-saline water.

Marine in origin, widespread in coastal water in Europe and

America.

9. Coscinodiscus argus Ehrenberg. Cells 70-100/x in diameter;

areolae large, polygonal, each with a prominent central pore;

arranged in a close network, not markedly radial; areolae

smaller towards the centre and near the margin; some tendency

for the areolae to lie on lines spiralling towards the centre, mar¬

ginal region small, hardly distinguishable from the sub-marginal

areolae; marginal teeth absent. (PI. I, figs, j, 1 & m).

Malaya Pahang: A stream near Kuantan.

Marine in origin, widespread.

10. Coscinodiscus decipiens Grunow. [Syn. Thalassiosira decipiens

(Grun.) Joerg.]. Cells small, 8-20/x in diameter, forming a

short mucilaginous chain; valve flat, areolae roundish to poly¬

gonal, very excentric, in lines curving away from the centre;

inner areolae larger, decreasing in size towards the margin;

submarginal teeth prominent. (PI. Ill, fig. c).

Malaya Pahang: A stream near Kuantan, together with

desmids.

Marine in origin, widespread in coastal waters.

Vol XIX. (1962).

11. Coscinodiscus divisus Grunow. Cells 30-50/* in diameter;

valves flat; areolae small, polygonal, arranged in bundles, lines

in each bundle parallel to one side and crossed by lines curving

towards the centre; marginal zone with fine areolae, about 12-

18 in 10/*; a very small spine opposite each bundle. (PI. II, figs,

a & d). This species is very characteristic, with the radial lines

parallel to one side of each bundle.

Malaya Malacca: Malacca River in non-saline water.

Distribution widespread, marine or brackish water.

12. Coscinodiscus excentricus Ehrenberg. Cells 20-40/* in dia¬

meter, discoid; valve flat; areolae small of equal size, arranged

excentrically in lines curving away from the centre; teeth in a

marginal ring. (PI. Ill, fig. e). This is much smaller than the

typical form, but obviously belongs under this species.

Malaya Pahang: A stream near Kuantan, in non-saline water.

Marine in origin, widespread.

13. Coscinodiscus griseus Greville var gallopagensis Grunow.

Cells 30-40/* in diameter; valves concentrically undulate; areo¬

lae small, resembling pores, arranged in radiating rows some¬

times crossed by spiral rows; inner rows widely set apart about

6-9 in 10/*, outer ones much closer, or smaller areolae forming

marginal striations; marginal teeth absent. (PI. II, fig. e).

Malaya Pahang: A stream near Kuantan.

Marine in origin.

14. Coscinodiscus lineatus Ehrenberg. Cells large, 80-120/* in

diameter; areolae large, hexagonal, more or less equal in size,

arranged in linear rows parallel to the diameter 4-5 in 10/*,

except near the margin, where the areolae are smaller and irre¬

gular; margin wide, striae 10-12 in 10/*; smaller teeth on inner

side of margin, about 3 in 20/*. (PI. I, figs, k & n).

Malaya Pahang: A stream near Kuantan.

Malacca: Malacca River.

Marine, widespread.

15. Coscinodiscus lineatus Ehrenberg var. van heurcldi P. T.

Cleve. Very much smaller than the type, only 15-20/* in dia¬

meter; areolae arranged in linear rows, 10 in 10/*; marginal

teeth absent. (PI. HI, fig. d).

Malaya Pahang: A stream near Kuantan.

Marine in origin.

Gardens Bulletin, S.

16. Coscinodiscus symmetricus Kitton. Cells 80-90/a in diameter;

areolae hexagonal, alike in size, 4—5 in 10/a, arranged in wedge-

shaped bundles, the rays of each bundle being parallel sym¬

metrically about the middle line; rays crossed by secondary

rays curving towards the centre; margin striate, 9-10 striae in

10/a; teeth absent. (PI. II, fig. c).

Malaya Pahang: A stream near Kuantan.

Malacca: Malacca River.

Marine in origin.

THALASSIOSIRA P. T. Cleve 1873

Cells discoid or drum shaped, single or united into chains by a

central thread; valves circular very finely ornamented; teeth sub¬

marginal, usually very fine, often a single unpaired course tooth.

Largely a marine genus.

17. Thalassiosira fluviatilis Hustedt. Cells 15-20 /a in diameter,

drum-shaped, solitary; valve slightly convex, very finely punc¬

tate in radial striae, not in bundles, teeth submarginal, fine,

10-12 in 10/a, one much larger unpaired tooth or spine. (PI. II,

fig. k).

Malaya Pahang: A stream near Kuantan, together with des-

mids.

Widespread in brackish water, both inland and estuarine.

STEPHANODISCUS Ehrenberg 1845

Cells discoid, drum-shaped or cylindric, single and free-float¬

ing, rarely in chains; valves circular, radially punctate and with

a ring of short, but stout spines near the margin; girdle view

generally somewhat undulate, with smooth surfaces and without

intercalary bands, chromatophores several small discs or one or

two large irregular flat plates. Largely a marine genus.

18. Stephanodiscus biscrialis Prowse sp. nov. Cells drum-shaped,

valves circular, 40-50/x in diameter, hollowed in the middle,

striae in two series, the outer series finely punctate, 11-12 in

10/a, about one-third radius long, inner series prominent, highly

refractive, punctae not visible, each stria a continuation of every

alternate outer stria into the hollowed centre, about one-third

radius long, outer striae not joined in pairs to inner striae as in

S. astraea; central area apparently quite smooth; submarginal

spines short, prominent, 3i in 10/a (PI. Ill, fig. a).

Malaya Pahang: A stream near Kuantan. (Type locality).

Holotype No. Prowse 582a is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Vol XIX. (1962).

Frustula cylindrica, altitudine brevissima. Valvae orbiculares,

40-50/x in diam., medio concavae. Striae biseriatae, in serie

exteriore minute punctatae, 11-12 in 10/x circa tertiam radii par¬

tem longae, cum interioribus pariter ut in S. astraea non jugatae;

striae interiores singulae ex unisquibusque striis altematis exte-

rioribus orientes, circam tertiam radii partem longae. Area cen¬

tralis laevissima ut videtur; spinis submarginalibus brevibus, pro-

minentibus, 31 in 10/x..

Malaya Pahang, in fluvio proximo urbe Kuantan.

19. Stephanodiscus fenestralis Prowse sp. nov. Cells drum-shaped;

valves circular, 40-50/x in diameter, hollowed in the middle;

striae 12-14 in 10/x, about half the radius in length, reaching

to the edge of the central hollow where they meet the upper

edges of the sub-central rectangular refractive “windows” or

depressions, each about 5 striae wide and lying 3-4 striae

apart on the sloping sides of the central hollow; spaces between

the “windows” and the central area of the hollow apparently

quite smooth; submarginal spines short, prominent, 3-4 in 10/x.

(PI. ID, fig. b).

Malaya Pahang: A stream near Kuantan. (Type locality).

Holotype No. Prowse 582b is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Frustula cylindrica, longitudinae brevissima. Valvae orbiculares,

40-50/x in diam., medio late concavae. Striae 12-14 in 10/x, fere

semi-radiales, inter extremitates earum ac cavi marginem centralis

caveolulis rectangularibus, refractivis, singulis 5-striae latis, 3-4

striae inter se remotis, in modum fenestrarum dispositis; loculi in¬

ter caveolulas et cavus centrales laevissimi ut videtur. Spinae sub-

marginales breves, prominentes, 3-4 in 10/x.

Malaya Pahang, in fluvio proximo urbe Kuantan.

CHAETOCEROS Ehrenberg 1844

Cells cylindrical, elliptical or circular in valve view, in girdle

view rectangular with plane, concave or slightly convex ends; inter¬

calary bands present; cells united in chains, rarely single; each

valve bearing two long fine hair like spines which intersect with

those of the neighbouring cell; chromatophores small, discoid and

numerous. Auxospores formed singly in each cell. Mainly a marine

genus.

20. Chaetoceros amanita A. Cleve. Cells in girdle view rect¬

angular 8-12/x high X 15-20 /x in widest width, sides straight,

end walls curved, separated by lanceolate openings; valve view

elliptical; spines long and slender, not differing markedly on

Gardens Bulletin , S.

end cells; auxospore with a large primary valve and a smaller

secondary valve, both ornamented with short spines. (PI. Ill,

figs, g & h).

The characteristically shaped auxospore makes it certain that

the Malayan specimens belong to this species.

Malaya Pahang: A stream near Kuantan together with des-

mids.

Distributed in brackish waters in Europe and the Middle East.

RHIZOSOLENIA Ehrenberg 1843; emend. Brightwell 1858

Cells elongate cylinders, with many intercalary bands, and cir¬

cular or elliptical in cross-section, free-floating, solitary or some¬

times in straight or spirally twisted chains; end of valve calyptra-

like, terminating apically in a long centric or excentric spine; walls

very finely punctate or without markings, silicification slight;

chromatophores small, discoid and scattered. Reproduction by

auxospores, statopores and microspores. Mainly marine.

21. Rhizosolenia eriensis H. L. Smith. Cells cylindrical, slightly

flattened, 5-15 /a wide X 40-150/a long; intercalary bands scale¬

like and imbricated 2-4 in 10/a; valve end calyptra like, each

terminating asymmetrically in a single long spine, shorter in

length than the length of the cell body. (PI. Ill, fig. f).

Malaya Malacca: Batu Berendam, in fish-ponds at the Tro¬

pical Fish Culture Research Station.

Widespread in lakes in Europe, America and in Indonesia.

II. ORDER PENNALES

Cells solitary or variously united into ribbon like, stellate, zig¬

zag, dendroid or irregular colonies, free-floating, or attached by

gelatinous stalks or by other mucilaginous secretions; valves

elongate, rarely nearly round, bilaterally symmetric about a

median longitudinal line, not radiating from a central point; pseu¬

doraphe or raphe present, if raphe present, cells possess the pro¬

perty of mobility; valves without marginal spines or projections,

rarely winged; chromatophores variable, small and discoid, or

larger and laminate with pyrenoids; auxospores formed by the

conjugation of two cells, or autogamously; vegetative cells

diploid.

Vol. XIX. (1962).

Key to Pennales

(i) Cells with no true raphe; pseudoraphe on both valves (ii)

(i) Cells with a very short raphe at each end .(iii)

(i) Cells with one valve bearing a true raphe, the other with

only a pseudoraphe.(iv)

(i) Both valves bearing true raphes. (v)

(ii) Valves usually narrow and much elongated; cells solitary

or in tufted fan-shaped colonies . .,.9. Synedra

(ii) Valves usually broader; cells in ribbons, zig-zag bands or

stellate colonies, rarely solitary . 8. Fragilaria

(iii) Cells usually in ribbons; no denticulations on valve edges

10. Eunotia

(iii) Cells in branched dendroid colonies, joined end to end;

valves with fine denticulations at edge

11. Desmogonium

(iv) Valves elliptic to nearly circular; cells transversely curved

in girdle view. 12. Cocconeis

(iv) Valves more elongated; cells longitudinally curved in

girdle view. 13. Achnanthes

(v) Raphe in a distinct keel or wing.(vi)

(v) Raphe not in a keel or wing.(ix)

(vi) Keel usually attached to one lateral margin of each valve,

rarely central, and with a row of circular pores (carinal

dots) .. (vii)

(vi) Keel not attached to valve margin, but straight, sigmoid or

excentric to the median longitudinal line, no carinal dots

28. Tropidoneis

(vi) Keel usually wing-like attached to both margins of each

valve; valve face distinctly costate.(viii)

(vii) Both valves having the keel attached on the same adjacent

side . 29. Hantzschia

(vii) Valves having the keels attached on opposite sides

30. Nitzschia

(viii) Valve view nearly circular in outline, often bent

33. Campylodiscus

(viii) Valve view ovate, elliptic or nearly linear ... 32. Surirella

(viii) Valve view distinctly linear, sigmoid . . 31. Stenopterobia

(ix) Valve view more or less symmetrical about the median

longitudinal line . .. (x)

(ix) Valve view not symmetrical about the median longitudinal

^ ne . (xviii)

Gardens Bulletin , S.

(x) Valve view and girdle view usually symmetrical about the

median transverse line..(xi)

(x) Valve and girdle view not symmetrical about the median

transverse line; one pole usually wider than the other

26. Gomphonema

(xi) Raphe enclosed between parallel ribs or horns extending

from central nodule. (xii)

(xi) Raphe not enclosed between ribs.(xiii)

(xii) No longitudinal furrow outside ribs, transverse striations

finely punctate, not costate. 14. Frustulia

(xii) Longitudinal furrow on each side of ribs; transverse stria¬

tions costate or coarsely punctate.19. Diploneis

(xiii) Valve with transverse costae opening inwards; raphe com¬

plex, often curved.23. Pinnularia

(xiii) Valves punctate; if costate, costae showing punctae and

not opening inwards; raphe usually much simpler . (xiv)

(xiv) Striae crossed by longitudinal hyaline lines . (xvi)

(xiv) Striae not crossed by hyaline longitudinal lines.xv

(xv) Central nodule not reaching margins.22. Navicula

(xv) Central nodule reaching both margines ... 20. Stauroneis

(xvi) Longitudinal lines parallel to and near the lateral margins

(xvii)

(xvi) Longitudinal lines not parallel to lateral margins but zig¬

zagging and coalescing in places .... 21. Anomoeoneis

(xvii) Raphe bifurcate at poles and curving in opposite directions

at central nodule... 18. Neidium

(xvii) Raphe not bifurcate at poles and straight or only slightly

curved in same direction at central nodule . 17. Caloneis

(xviii) Valve view distinctly sigmoid.(xix)

(xviii) Valve view curved to one side, lunate or sickle-shaped . (xx)

(xix) Transverse striations crossed by longitudinal lines

15. Gyrosigma

(xix) Transverse striations cut by crossed lines at an angle

16. Pleurosigma

(xx) Cells in girdle view with parallel sides, more or less rect¬

angular ... 25. Cymbella

(xx) Cells in girdle view with outer margins convex .... (xxi)

(xxi) Transverse striations separated by costae . 27. Rhopalodia

(xxi) Transverse striations without costae.24. Amphora

Vol. XIX. (1962).

FRAGILARIA Lyngbye 1819, Rabenhorst 1864

Cells united into free-floating or sessile colonies, mostly zig-zag

chains or ribbons, sometimes flat, stellate colonies; cells rect¬

angular in girdle view, with one or two intercalary bands (some¬

times none), without septa or costae; valve view linear to fusiform,

bilaterally symmetrical, usually attenuated at the poles, sometimes

capitate and often medianly inflated, rarely constricted; transverse

striae usually fine, sometimes coarse; pseudoraphe narrow and

indistinct to broad and prominent; chromatophores numerous

small discoid bodies or one to four laminate plates with pyrenoids.

Auxospores formed singly within the cells.

22. Fragilaria lapponica Grunow var. tetragonalis Prowse var.

nov. Valves quadrangular, 45/x long, 40/x wide, with slightly in¬

curved margins. Striae short, thick, marginal, 6-10 in 10/x.

(PL IV, fig. f).

Malaya Malacca: Malacca River. (Type locality).

Holotype No. Prowse 183a is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Valvae quadrangulares, 45 ^ longae, 40/i latae, marginibus paulo

incurvis. Striae breves, crassae, marginales, 6-10 in 10^.

Malaya Malacca, in fluvio.

This is much larger than most forms of the species.

23. Fragilaria vaucheriae (Kuetzing) Boye Petersen. [Syn.:

Synedra vaucheriae Kuetz.; F. intermedia Grunow.]. Cells 50-90/x

long X 3-6/x wide, forming closely knit ribbons; girdle view

linear—rectangular; valve view elongate—lanceolate, narrowed

near the ends, capitate; pseudoraphe narrow; transverse striae

marked, 9-12 in 10/x, with a clear central area on one side. (PI.

IV, fig. a).

Malaya Pahang: Cameron Highlands.

Selangor: Gombak River.

Malacca: Malacca River.

Widely distributed in fresh water.

The Malayan specimens are larger than the. type, but it is doubt¬

ful if this is of varietal significance.

24. Fragilaria virescens Ralfs var. elliptica Hustedt. Cells 9-15/x

long X 5-7/x wide, rectangular in girdle view, elliptic-lanceolate

in valve view; pseudoraphe narrow, not very distinct; transverse

striae fine, parallel or slightly radiate, 15-20 in 10/x; united in

short ribbons. (PI. IV, fig. d).

Malaya Pahang: Stream near Kuantan, rare.

Widely distributed in freshwater in several countries, but rare.

Gardens Bulletin, S .

SYNEDRA Ehrenberg 1830

Cells narrow and much elongated, solitary or in fan-shaped tufts

or radiating colonies, free-floating or epiphytic, sessile or stalked;

valves linear to lanceolate, straight or sometimes curved, with

poles attenuated or not, often capitate; transverse striations lateral

to a conspicuous pseudoraphe, which is usually narrow but some¬

times broad; central smooth area present or not; girdle view

elongate with truncate ends, striated; chromatophores two large

plates, usually with 3 or more pyrenoids in each.

Auxospores 1-2 in each cell, formed without conjugation.

25. Synedra tabulata (C.A. Agardh) Kuetzing, var. acuminata

(Grunow) Hustedt. [Syn.: Diatoma tabulatum C.A. Ag., S. affi -

nis Kuetzing var. acuminata Grun.]. Cells 50-80 /a long X

3-5 /a wide, lanceolate, narrowed to slightly capitate ends; striae

13-15 in 10/a, shorter than the wide pseudoraphe. (PI. IV, figs,

b & c).

Malaya Pahang: Stream near Kuantan, together with desmids,

not common.

Widely spread in brackish waters in various countries.

26. Synedra ulna (Nitzsch) Ehrenberg var. amphirhynchus

(Ehrenberg) Grunow. [Syn.: Bacillaria ulna Nitzs., S. amphir¬

hynchus Ehr.]. Cells 150-240/a long X 5-7/a wide, nearly linear,

constricted near the poles, which are capitate; pseudoraphe nar¬

row, but well defined, widening in the middle to an elliptic to

rectangular, smooth middle area; striae well-marked, 8-10 in

10/A. (PI. IV, fig. h).

Malaya Selangor: Fish-ponds.

Negri Sembilan: Fish-ponds.

Malacca: Fish-ponds, Malacca River.

Singapore: Botanic Gardens Lake.

Widespread in freshwaters in various parts of the world.

27. Synedra ulna (Nitzsch) Ehrenberg var. danica (Kuetzing)

Grunow. [Syn.: Bacillaria ulna Nitzs., S . danica Kuetz., S. ulna

(Nitzs.) Ehr. var. chaseana B. W. Thomas, S. longissima W.

Smith var. acicularis Meister.]. Cells 150-200/j long X 5/x wide;

long, narrow linear-lanceolate, slightly narrowing to the poles

which are hardly capitate; pseudoraphe narrow but well defined,

widening at the centre into a rectangular, clear middle area;

striae well-marked, 9-10 in 10/a. (PL IV, fig. g).

Malaya Selangor: Klang River, fish-ponds.

Negri Sembilan: Fish-ponds.

Malacca: Fish-ponds.

Singapore: Botanic Gardens Lake.

Widespread in freshwater.

Vol. XIX. (1962).

EUNOTIA Ehrenberg 1837

Cells rectangular to linear in girdle view, both girdles and valves

strongly ornamented, usually with inter-calary bands, free floating

or epiphytic, solitary or united valve to valve into ribbons or

chains; valves arcuate with similar poles but dissimilar margins:

concave side regular, convex side regular, or inflated at the poles

or undulate, sometimes nearly straight; raphe very short, extend¬

ing from the fairly evident polar nodule diagonally to the concave

margin, no central nodule; neither costae nor septa present; inter¬

calary bands usually present; transverse striations or punctations

present, but without marginal denticulations or dots; chromato-

phores two, laminate, without pyrenoids.

Auxospores single from the conjugation of two cells.

28. Eunotia camelus Ehrenberg. Cells rectangular in girdle view,

united in ribbons; in valve view 25—30/a long X 5/a wide, dis¬

tinctly arcuate, with two prominent humps on the convex side;

ends narrowed on convex side, rounded—sub-capitate, reflexed

away from the concave side, nodules small, sub-terminal; stria¬

tions 11-14 in 10/a. (PI. V, figs, y & z).

The two forms figured are more arched and with more pro¬

minent humps than the type, and the ends are reflexed. They show

a marked resemblance to var. gibbosa Gandhi and var. ventricosa

Gandhi, but are regular and not gibbous on concave margin.

Malaya Malacca: Malacca River.

Distributed in South-east Asia.

29. Eunotia camelus Ehrenberg var. karveerensis Gandhi. Cells

rectangular in girdle view, united in ribbons; on valve view

35-40/a long X 5-7 wide, strongly arcuate, dorsal edge convex

with four strong rounded uniform humps; ends narrowed on

dorsal side, produced, rounded, sub-capitate; nodules small,

subterminal; striae 11-13 in 10/a. (PI. V, figs, c & d).

Malaya Malacca: Malacca River.

Described from India.

A. Berg (1939) has separated the two humped forms as forms of

E. pectinalis (Kuetz.) Rabh. but there seems no justification for

this as the two are unlike E. pectinalis , being much less arched,

very much less convex on the dorsal side, and with the ends

hardly reflexed. Gandhi (1957) has discussed the status of E.

camelus and I agree with him that both two and four-humped

forms should be included as forms of the same species, but un¬

like him I would take the two-humped forms as the type, since they

were so described by Ehrenberg—“E. minoribus, valvus striatis,

dorse elati gibbere duplici, ventre concave, apicibus productis

attenuatis obtusis.” (De Toni, 1889—1924)—and regard the four-

humped forms as being varieties.

Gardens Bulletin, S.

30. Eunotia cancellata A. Berg var. esseda A. Berg. Cells small,

rectangular in girdle view; in valve view 10-15/a long X

3-4 /x wide, only slightly arcuate, nearly straight on ventral

margin, except at the ends where it curves down slightly; dorsal

margin straight in the middle, curving away to the rounded,

sub-capitate ends, shoulders marked; nodules terminal with well

marked inner almost central secondary nodules; striae 15-20

in 10/a. (PL V, fig. n).

The Malayan form seems to come closest to the variety esseda

as described and figured by A. Cleve-Euler (1933).

Malaya Pahang, a stream near Kuantan.

Described from Europe.

31. Eunotia diodon Ehrenberg var. minor Grunow. Cells rect¬

angular in girdle view; in valve view 35-40/a long X 10-12/a

wide, very slightly curved on the ventral margin, much more

markedly convex on dorsal side, bi-undulate with a slight dep¬

ression between the two humps, curving down to the non-capi-

tate ends; nodules, prominent, terminal; striations 13-15 in 10/a,

slightly radial. (PI. V, figs, p & q).

Malaya Pahang: Cameron Highlands.

Malacca: Malacca River.

Widely distributed.

32. Eunotia exigua (Brebisson) Grunow. [Syn.: Himantidium

exiguum Brebisson]. Valves small, 15-20/a long X 2-3/a wide,

linear, slightly arched, both margins parallel in the middle, but

the ends rounded-capitate, slightly reflexed to the dorsal margin;

nodules terminal; striae fine, 20-24 in 10/a (PI. V, fig. j).

Malaya Pahang: Cameron Highlands.

Widely distributed.

33. Eunotia faba (Ehrenberg) Grunow var. denscstriata Oestrup.

[Syn.: Himantidium faba Ehr.]. Cells small, rectangular tabular

in girdle view; in valve view 13-15/a long X 3-4/a wide, slightly

arched, subconcave on the ventral margin, slightly more curved

on the dorsal margin; ends rounded, hardly capitate; nodules

terminal; striae 19-20 in 10/a. (PI. V, fig. m).

Malaya Pahang: Cameron Highlands.

Widely distributed.

34. Eunotia flexuosa Kuetzing. Cells linear in girdle view; in

valve view 120-270/a long X 3-5 /a wide, linear, straight or

slightly broadened; nodules terminal with a short pseudoraphe

extending inwards; transverse striations finely punctate, 12-18

in 10/a. (PL VI, figs, b & c).

Malaya Pahang: Cameron Highlands.

Malacca: Swamps.

Singapore.

Vol XIX . (1962).

35. Eunotia formica Ehrenberg. Cells rectangular in girdle view;

in valve view 40-130//. long X 10-14/a wide, with parallel mar¬

gins, ends cuneate making the whole valve resemble a bone in

appearance; nodules terminal, with a long pseudoraphe extend¬

ing inwards from it parallel to the sides; striations 9-14 in 10/x,

somewhat irregular, punctate. (PL VI, figs, g & h).

Malaya Pahang: Cameron Highlands.

Selangor: Gombak River.

Malacca: Malacca River.

Widely distributed.

36. Eunotia gracilis (Ehrenberg) Rabenhorst. [Syn.: Himantidium

gracilis Ehr.]. Cells in girdle view rectangular; in valve view,

80-110/x long X 4-6 /a wide, linear with parallel sides, distinctly

curved; ends rounded, slightly capitate, nodules terminal; trans¬

verse striae 12-16 in 10/a. (PL VII, fig. b).

Malaya Malacca: Malacca River.

Widespread in distribution.

37. Eunotia grunowi A. Berg var. uplandica A. Cleve f. sub-

undulata A. Cleve. [Syn.: E. pectinalis var. stricta (Rabenhorst)

Grunow forma]. Cells rectangular in girdle view; in valve view

50-70 [x long X 8-10/a wide, slightly curved, dorsal side more

convex than ventral side with marked shoulders, slightly in¬

curved in the middle; ends rounded, sub-capitate; nodules ter¬

minal; transverse striae 10-13 in 10/a. (Pl. V, fig. e).

Malaya Malacca: Malacca River.

Recorded from Scandinavia.

The form found in Malaya comes nearest to that described by

A. Cleve-Euler (1953).

38. Eunotia lunaris (Ehrenberg) Grunow. [Syn.: Synedra lunaris

Ehr.]. Cells linear in girdle view; in valve view 45-80/a long X

3-4/x wide, linear with parallel sides, curved forming a wide arc,

slightly thinner at the poles; nodules terminal, small; transverse

striae 14-18 in 10/a. (PL V, fig. f).

Malaya Pahang: Cameron Highlands.

Widespread in distribution.

39. Eunotia lunaris (Ehrenberg) Grunow var. capitata Grunow.

Differing from the type in that the ends are capitate and slightly

reflexed; in valve view 40-80/t long X 4-5 /a wide; transverse

striae 15-16 in 10/a. (PL V, fig. h).

Malaya Malacca: Malacca River.

Widespread in distribution.

Gardens Bulletin, S .

40. Eunotia major (W. Smith) Rabenhorst var. emarginata A.

Cleve. [Syn.: Himantidium majus W. Smith, E. major (W.

Smith) Rabh. var. bidens (Greg.) A. Cleve.]. Cells in girdle

view rectangular; in valve view 60- 100/a long X 10-12/a wide;

broadly curved, with slightly cuneate, rounded ends, the dorsal

side with two broad flattish humps; nodules terminal; trans¬

verse striae somewhat irregular, 10-12 in 10/a. (PI. VI, fig. f).

Malaya Perak: Taiping lakes.

Described from Europe.

41. Eunotia major (W. Smith) Rabenhorst var. indica Grunow.

[Syn.: Himantidium majus W. Smith, E. indica Grun.]. Cells

rectangular in girdle view; in valve view 70-90/a long X

10/a wide, curved, with both margins parallel in the middle but

curving down on the dorsal side to the sub-capitate, slightly

cuneate, ends; nodules terminal; transverse striae 8-12 in

10/x, irregularly distributed. (PI. V, fig. i).

Malaya Malacca: Malacca River.

Described from India and from Sweden, and is probably widely

distributed.

42. Eunotia major (W. Smith) Rabenhorst var. linearis A. Cleve.

[Syn.: Himantidium majus W. Smith, E. monodon Ehr. var.

major Hust.]. Cells rectangular in girdle view; in valve view

120-1 50/a long X 10-12/a wide, broadly curved with parallel

sides; ends rounded, slightly wider; nodules terminal, promi¬

nent; transverse striae 8-10 in 10/a. (PI. VII, fig. a).

Malaya Malacca: Malacca River.

Distribution widespread.

43. Eunotia monodon Ehrenberg var. alpina Kuetzing. Cells

small, 20-25/a long X 5/a wide in valve view, slightly curved,

concave on the ventral side, slightly more convex on the dorsal

side, narrowing gently to the rounded, hardly reflexed ends;

nodules terminal, small; striae 14 in 10/a. (PI. V, fig. s).

Malaya Pahang: Stream near Kuantan.

Distributed in Europe, possibly elsewhere.

44. Eunotia monodon Ehrenberg var. constricta A. Cleve-Euler.

[Syn.: E. major (W. Smith) Rabh. v. hybrida f. bidens A. Cleve].

Cells rectangular in girdle view; in valve view 65-70/a long X

lO-15/i wide, broadly arcuate, with two broad swellings on the

dorsal margin, slightly reflexed before the rounded, slightly cu¬

neate ends; nodules terminal, prominent; striae 10-12 in 10/a.

(PI. V, fig. a).

Malaya Malacca: Fish-ponds at Jasin and Malacca.

Probably widespread in distribution.

Vol. XIX. (1962).

45. Eunotia monodon Ehrenberg var. undulata A. Cleve-Euler.

[Syn.: E. major (W. Smith) Rabh. var. undulata A. Berg]. Cells

in girdle view rectangular; in valve view 65—70/a long X

8-1 0 /a wide, broadly arcuate, with the dorsal margin more con¬

vex and with four broad, flattish, equal humps; ends narrower,

rounded, not markedly capitate; nodules terminal, small; trans¬

verse striae 10-12 in 10/a. (PI. V, fig. g).

Malaya Malacca: Malacca River.

Distributed in Europe and probably elsewhere; not common.

This certainly comes nearest to Cleve-Euler’s figure and descrip¬

tion (1953), but it may possibly belong to another species.

46. Eunotia pectinalis (Kuetzing) Rabenhorst. [Syn.: Himantidium

pectinalis Kuetzing.]. Cells rectangular-tabular in girdle view, in

valve view 40-80 u long X 5-8/a wide, linear, slightly curved,

the dorsal margin parallel to the ventral margin, narrowing

slightly at the ends to the rounded slightly reflexed but non-

capitate poles; nodules terminal, small, transverse striae 9-11

in 10/a, well-marked. (PI. V, fig. b).

Malaya Pahang: Cameron Highlands.

Malacca: Widespread.

Widespread in distribution.

47. Eunotia pectinalis (Kuetzing) Rabenhorst var. minor (Kuetz¬

ing) Rabenhorst. [Syn.: Himantidium pectinalis Kuetz. var.

minor Kuetz.]. Much smaller than the type, 15-20/a long X

1.5-2/a wide. (PI. V, fig. w).

Malaya Pahang: Cameron Highlands.

Widespread in distribution.

48. Eunotia polydentula (Brun) emend. A. Berg var. perminuta

(Grunow) A. Berg. [Syn.: E. tridentula Ehr. var. perminuta

Grun.]. Cells small, 10-20/a long X 3-4/a wide, slightly ar¬

cuate with roughly parallel margins, the dorsal side with 3-4

shallow swellings; ends a little narrower, capitate and slightly

reflexed; nodules terminal; transverse striae 16-20 in 10/a. (PI.

V, figs, u & v).

Malaya Pahang: Cameron Highlands.

Johore: Ayer Hitam.

Described from Europe but probably more widely distributed.

49. Eunotia praemonos A. Cleve-Euler var. inflata (Grunow) A.

Cleve-Euler. [Syn.: E. praerupta Ehr. v. inflata Grun.]. Cells

small, 15-30/a long X 5-7/a wide, ventral side hardly con¬

cave, nearly straight, dorsal side much more convex; ends

broadly rounded, slightly reflexed; nodules terminal, well-mark¬

ed; transverse striae coarse, irregularly arranged, 7-13 in 10/x.

(PI. V, figs, r & 1).

Malaya Pahang: Stream near Kuantan.

Described from Europe but doubtless more widespread.

Gardens Bulletin , S.

50. Eunotia praemonos A. Cleve-Euler var. monodon (Oestrup)

A. Cleve-Euler. [Syn.: E. praerupta Ehr. var. curta Grun.

forma monodon Oest., E. praerupta Ehr. var. monodon Oest.].

Cells small, 20-25/a long X 7-8 /a wide; ventral side slightly

concave, dorsal side with a marked central wedge-shaped swell¬

ing; ends broadly rounded, only very slightly capitate; nodules

terminal, well marked; striae coarse, 10-12 in 10/a. (PI. V,

fig. k).

Malaya Pahang: Stream near Kuantan.

Described from Europe but probably occurs elsewhere.

51. Eunotia praemonos A. Cleve-Euler var. tibetica (Mereschko-

wski) A. Cleve-Euler. [Syn.: E. praemonos A. Cleve-Euler forma

tibetica A. Berg, E. praerupta var. tibetica Mereschkowski E.

praerupta-monos A. Berg forma undata A. Berg, E. praerupta

var. bidens Hust.]. Cells rectangular-tabular in girdle view; in

valve view 25-35/a long X 7-9/a wide, arcuate with two broad

humps on the dorsal side, the ends parallel, very bluntly round¬

ed poles, almost truncate; nodules small, terminal; transverse

striae irregular with some short ones on the dorsal margin,

10- 14 in 10/a. (PI. VI, fig. i).

Malaya Malacca: Malacca River.

Described from Tibet and Scandinavia but doubtless occurs

elsewhere.

Despite the difference in habitat the Malayan forms seem to fit

under no other variety, and any differences are too slight to justify

their separation under a new form.

52. Eunotia robusta Ralfs. Cells large, in girdle view rectangular-

tabular; in valve view 65-100/a long X 9-18/a wide, broadly li¬

near, arcuate, with 4-9 broad swellings on the dorsal side; poles

broadly rounded; nodules terminal, distinct; transverse striae

coarse, punctate, irregular, 9-12 in 10/a. (PI. VI, figs, a, d & e).

Malaya Malacca: Malacca River.

Widespread in distribution.

This species is unmistakable, despite the wide variation in the

number of dorsal humps.

53. Eunotia sarekensis A. Cleve-Euler var. minor A. Cleve-Euler.

[Syn.: E. papilio (Grunow) P. T. Cleve forma minor Hustedt.].

Cells small, rectangular in girdle view; in valve view 20-30/a

long X 9-12/a wide, arcuate with the dorsal side prominently

raised into two humps, shoulders sharp, sloping suddenly; ends

with almost parallel sides, poles bluntly rounded; nodules ter¬

minal; transverse striae very irregular with many short ones,

11- 14 in 10/a. (PI. V, fig. o).

Malaya Pahang: Cameron Highlands.

Described from Europe.

Vol XIX. (1962). )

54. Eunotia sudetica O. Mueller var. incisa (Mayer) A. Cleve-

Euler. [Syn.: E. pectinalis (Kuetz.) Rabh. var. incisa Mayer].

Cells in valve view 15-25 /a long X 7-10/* wide; ventral mar¬

gin almost straight, dorsal margin markedly convex, ends slightly

reflexed to the ventral side making a small depression or nick

near each end of the ventral margin; poles rounded; nodules

small, terminal; transverse striae slightly radial, 10-14 in 10/a.

(PI. V, figs, t & x).

Malaya Pahang: Stream near Kuantan.

Described from Europe.

DESMOGONIUM Ehrenberg 1848

Cells united end to end forming branching chains, or often den¬

droid colonies. In girdle view usually linear, in valve view linear.

Raphe short in terminal nodules; transverse striae distinct; margins

with small granules or denticulations, often difficult to see.

55. Desmogonium rabenhorstianum Grunow. Cells joined end to

end, often in pairs or fours, forming characteristic branched

chains or dendroid colonies; in girdle view linear with slightly

swollen ends; in valve view 100-200/a long X 5-8/a wide, li¬

near, slightly arcuate, with rounded, slightly capitate poles;

nodules terminal with a small pseudo-nodule in the middle of

each end; striae 13-16 in 10/a, each alternate one with a small

granule or denticulation at each end. (PL VII, figs, c & d). The

marginal granules are often difficult to see, showing up only

under the highest powers of phase contrast. The characteristic

habit of the colonies is enough to distinguish the species.

Malaya Perak: Taiping.

Pahang: Cameron Highlands, and in several coastal

streams.

Selangor: Gombak River.

Negri Sembilan: Port Dickson.

Malacca: Swamps.

Johore: Yong Peng.

Singapore.

Distributed in South-east Asia.

COCCONEIS Ehrenberg 1835; Grunow 1868

Cells transversely curved in girdle view, solitary, epiphytic upon

submerged aquatic plants; septa incomplete and intercalary bands

absent; valve view elliptic, epivalve with an axial pseudoraphe,

Gardens Bulletin, S .

hypovalve with a median true raphe which is straight or sigmoid,

with central and polar nodules; transverse striae often in distinct

punctate rows; chromatophores single, laminate, usually adjoining

the epivalve, with one or two pyrenoids.

Auxospores single from two conjugated cells, or formed par-

thenogenetically from a single gamete.

56. Cocconeis fcuerbornii Hustedt. Valves elliptic with broadly

rounded ends, 20-25/a long X 10—13/x wide; epivalve with a

narrow, linear pseudoraphe, striae radial, 17 in 10/x in the mid¬

dle, 20-25 in 10/a near the poles, punctate with the inner punc-

tae more widely spaced forming wavy longitudinal lines;

hypovalve with a linear raphe, axial area narrow linear widen¬

ing suddenly in the middle to form a stauros reaching half-way

to the lateral margins, striae radial, 20-23 in 10/a in the middle,

the punctae closer together and more evenly spaced than in

the epivalve. (PI. VIII, figs, k & 1).

Malaya Malacca: Malacca River,

Described from Indonesia.

57. Cocconeis thumensis Mayer. Valves elliptic with broadly

rounded ends, 13-20/a long X 5-8/a wide; epivalve with broad

lanceolate to nearly straight pseudoraphe, striae radial, 12-15

in 10/a coarsely punctate; hypovalve with linear raphe and nar¬

row linear axial area, striae radial 12-15 in 10/a, coarsely punc¬

tate. (PI. IV, figs, t & u).

Malaya Malacca: Malacca River.

Reported from Europe and Africa, probably widespread in

distribution.

There is some variation in this species, the two examples illus¬

trated being the extremes with many intermediates. There seems

no justification for separating the longer forms under a varietal

name.

ACHNANTHES Bory 1822

Cells in girdle view rectangular, bent or curved longitudinally,

usually attached by gelationous stalks, or sessile and united in

bundles at the valves, rarely into filaments, occasionally free-

floating and solitary; valve-view generally linear-lanceolate, or ellip¬

tic; hypovalve usually concave with raphe, a distinct central nodule

and less distinct polar nodules, the central area sometimes trans¬

versely widened into a stauros; epivalve generally convex, with a

pseudoraphe; transverse striations often somewhat radiate, in

some species costate; chromatophores one, two or numerous and

discoid. Two auxospores formed from two cells by the conjuga¬

tion of two pairs of gametes.

Vol. XIX. (1962).

58. Achnanthes brcvipes C. A. Agardh var. intermedia Kuetzing.

Cells in girdle view rectangular, 38-40/a long X 11-12/a wide,

distinctly bent longitudinally, with marginal rows of very coarse

punctae, costate in appearance, the concave margin broken in

the middle by a large hyaline space, four longitudinal rows of

coarse punctae down the middle, marginal striae 10-11 in 10/a;

valve view linear-lanceolate with a very slight constriction in

the middle, epivalve with a linear pseudoraphe excentrically

placed, striae costate, 10-11 in 10/a, very coarsely punctate;

hypovalve with a linear raphe, axial area linear, widening in the

middle to form a stauros, striae costate, 10-11 in 10/a, very

coarsely punctate. (PI. VIII, figs. h-j).

Malaya Selangor: Feeder stream to the Klang Gates reservoir.

Widespread in distribution.

59. Achnanthcs crenulata Grunow. Cells in girdle view rect¬

angular, longitudinally bent; in valve view linear-lanceolate, 50/a

long X 13-15/a wide, margin distinctly wavy; epivalve with a

linear pseudoraphe near one margin, striae 9-10 in 10/a,

coarsely punctate, punctae 9-10 in 10/a; hypovalve with raphe,

axial area narrow lanceolate widened in the middle to form a

stauros reaching the lateral margins, striae 9-10 in 10/a,

coarsely punctate, punctae 9-10 in 10/a. (PI. IV, fig. v).

Malaya Selangor: Gombak River.

Described also from Perak and Indonesia.

60. Achnanthes exigua Grunow. Valves quadratic-elliptic to

linear-elliptic, subrostrate, 8-1 0 /a long X 4-5 /a wide; epivalve

with median linear pseudoraphe, striae slightly radial, 22 in 10/a;

hypovalve with raphe, axial area narrow, linear, widened in the

middle to form a stauros, striae slightly radial, 25 in 10/a, with

one short stria opposite each limb of the stauros. (PI. IV, fig. 1).

Malaya Selangor: Feeder stream of the Klang Gates reservoir.

Widespread in distribution.

The small form described is much less rostrate than usual and

resembles closely the form figured by A. Cleve-Euler (1953) as

544 a & b and called “forma typica.” Ruth Patrick (1936) has re¬

corded the rostrate form from Perak.

61. Achnanthes hauckiana Grunow var. rostrata Schultz. Valves

lanceolate, 17-18/a long X 6-7/a wide, rostrate with rounded

poles; epivalve with a linear lanceolate pseudoraphe, striae 12-

14 in 10/a prominent, radial, punctae faint; hypovalve with

raphe, axial area narrow lanceolate slightly widened in the

middle, striae 12-14 in 10/a, prominent, radial punctae faint.

(PI. IV, fig. W ).

Malaya Pahang: Stream near Kuantan.

Widespread, usually in brackish waters.

Gardens Bulletin , S.

62. Achnanthes lanceolate (Br6bisson) Grunow. [Syn.: Achnan -

thidium lanceolatum Brebisson.]. Valves elliptic-lanceolate 15—

20 fx long X 7~8/x wide, with bluntly rounded ends, hardly nar¬

rowed below the poles; epivalve with a linear pseudoraphe and

marked, punctate, transverse striae, 12-14 in 10 /a, radial and

with a prominent refractive gap in the middle of one side; hypo-

valve with a straight raphe and linear to narrow lanceolate

central area, striae prominent, punctate, 12-14 in 10 /a, radial

and with a gap in one side corresponding to that of the epivalve,

but not refractive. (PI. IV, fig. r).

Malaya Pahang: Cameron Highlands.

Widespread in freshwaters.

63. Achnanthes Ianceolata (Brebisson) Grunow var. rostrata

Hustedt. Differs from the type in being distinctly rostrate to sub-

capitate at the poles, 15-20/* long X 7-9 /a wide; pseudoraphe

of epivalve linear, striae less marked, punctate, 12-14 in 10/a,

with refractive gap in middle of one side; axial area of hypo-

valve narrow linear with small lanceolate central area, striae

12-14 in 10/a, radial, without a gap on one side. (PI. IV, fig. s).

Malaya Pahang: Cameron Highlands.

Widespread in freshwaters.

64. Achnanthes stauroneiformis Prowse. sp. nov. Frustules

oblong-lanceolate, with linear-rostrate poles, truncate at both

ends, 30—32/^ long X 25-26/a wide; epivalve with a linear, cru¬

ciform pseudoraphe having the transverse arms reaching nearly

to the margins, striae 16 in 10/a, distinctly punctate, those op¬

posite the lobes of the stauros being short, marginal 3 on each

side; hypovalve similar but with a very distinct raphe, striae

closer together, 18-20 in 10/a, punctate, striae opposite the

lobes of the stauros short, marginal, 4-5 on each side. (PI. IV,

fig- q).

Malaya Malacca: Malacca River. (Type locality).

This species closely resembles Stauroneis except that only one

valve bears a raphe.

Holotype No. Prowse 124a is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Frustula oblongo-lanceolata, apice utrinque lineari-rostrata,

truncata, 30-32/a longa, 25-26 /a lata. Epivalva cum pseudoraphe

lineari cruciformi, lobis transversalibus fere ad margines utrinque

attingentibus; striae 16 in 10/x, conspicue punctatae, in stauri lobis

breves marginales utrinque 3. Hypovalva consimilis, sed raphen

clarissimam ferens; striae magis approximatae, 18-20 in 10/x,

punctatae, striae in stauri lobis breves marginales utrinque 4-5.

Malaya Malacca: in flumine principali frequens.

Vol. XIX. (1962).

65. Achnanthes subhudsonis Hustedt. Valves lanceolate, 10-15/a

long X 3-4 /a wide, with acute poles; epivalve with narrow

linear pseudoraphe, transverse striae fine, 16—17 in 10/a,

lightly radial; hypovalve with straight raphe and lanceolate

axial area, transverse striae fine, 18 in 10/a. (PI. IV, fig. e).

Malaya Pahang: Cameron Highlands.

Described from East Africa.

66. Achnanthes tenuissima Hustedt. Valves linear to rhombic-

lanceolate, 18-20/a long X 3/x wide, with narrow rounded

poles; epivalve with a broad lanceolate pseudoraphe, striae short,

marginal, very fine, 28-34 in 10/a, slightly radial; hypovalve with

a straight raphe and narrow linear axial area, striae very fine,

32-36 in 10/a, parallel. (PI. IV, fig. p).

Malaya Malacca: Fish-ponds near Malacca.

Described from Indonesia.

67. Achnanthes tropica Hustedt. Valves linear elliptic, 12—15/a

long X 2-3/a wide, with slightly convex to almost parallel sides,

and rounded poles; epivalve with a lanceolate pseudoraphe and

short marginal striae, 18-20 in 10/a, parallel to slightly radial;

hypovalve with a linear-lanceolate axial area, narrower than the

pseudoraphe, and a straight raphe, transverse striae slightly

closer together 20-22 in 10/a, parallel. (PI. IV, figs, m, n & o).

Malaya Malacca: Fish-ponds near Malacca.

Described from Indonesia.

68. Achnanthes weltereckii Hustedt. Valves linear to linear-

lanceolate, 10-15/a long X 3-4/a wide, with slightly convex

margins and rostrate to capitate poles; epivalve with a lanceo¬

late pseudoraphe, about 1/3 width of valve in the middle,

transverse striae 20-24 in 10/a, slightly radial or parallel; hypo¬

valve with a straight raphe and linear axial area, widening in

the middle almost to reach the margin, transverse striae radial,

28-30 in 10/a, two or three opposite the central area so that it

appears like a stauros. (PI. IV, figs, i, j & k).

Malaya Pahang: Cameron Highlands.

Described from Indonesia.

FRUSTULIA C. A. Agardh 1824; Grunow, 1865

Cells solitary and free-floating, or sessile and enclosed in a gela¬

tinous sheath or sometimes a gelatinous tube; in valve view

linear-elliptic to rhombo-lanceolate; in girdle view rectangular,

without intercalary bands; central nodule longitudinally elongated,

prolonged towards each pole as parallel ribs enclosing the raphe,

Gardens Bulletin, S,

and uniting apically with the polar nodule; transverse striations

crossed by fine longitudinal lines; chromatophores two, some¬

times incised longitudinally, one on each side of the girdle and

connected by a cytoplasmic bridge.

Auxospores formed in pairs from two cells.

69. Frustulia javanica Hustedt. Valves elliptic-lanceolate, 50-

60 fx long X 25-26/a wide, with a distinct swelling on each side

opposite the central nodule, and suddenly narrowed near the

poles, making the margin triundulate; transverse striae fine, 24-

27 in 10/a, parallel except at the poles and crossed by fine

longitudinal striae 22 in 10/i. (PI. VIII, fig. c).

Malaya Pahang: Stream near Kuantan.

Reported from Indonesia and S. Africa.

The Pahang specimens are slightly broader and more lanceolate

in shape than those described from Indonesia by Hustedt, but in

all other respects they agree.

70. Frustulia rhomboides (Ehrenberg) de Toni. [Syn.: Navicula

rhomboides Ehr.]. Valves rhombo-lanceolate, 50-80/x long X

12-15/a wide, hardly rostrate at the poles; transverse striae

marked 27-28 in 10/a, longitudinal striae fine, 30 in 10/a. (PL

VIII, figs, a & b).

Malaya Penang.

Perak: Taiping.

Pahang: Cameron Highlands, Fraser’s Hill, and in all

of many streams sampled.

Selangor: Kuala Lumpur, Klang Gates, Klang.

Negri Sembilan: Port Dickson.

Malacca: Various localities.

Johore: Muar, Yong Peng, Kota Tinggi.

Singapore.

Worldwide and common in distribution.

71. Frustulia saxonica Rabenhorst. [Syn.: Frustulia rhomboides

(Ehrenberg) de Toni v. saxonica (Rabh.) P. T. Cleve]. Valves

elliptic, 30-40/a long X 8-10/a wide, slightly rostrate at the

poles; transverse striae slightly radial, fine 35 in 10/a, longitudi¬

nal striae fine, 35 in 10/a. (PI. VIII, figs, f & g).

Malaya Pahang: Cameron Highlands.

Malacca: Fish-ponds, Malacca River.

Distribution worldwide.

Vol XIX. (1962).

GYROSIGMA Hassall 1845, emend. P. T. Cleve 1894

Cells solitary and free-floating, sometimes in gelatinous tubes;

girdle view linear-lanceolate, intercalary bands and septa absent;

valve view distinctly sigmoid, usually gradually attenuated to the

broadly rounded poles, sometimes sharply narrowed; rapne sig¬

moid with small central and polar nodules; axial area narrow,

with small, usually rounded, sometimes asymmetric, centra) area;

transverse striae crossed by longitudinal striae at right angles;

chromatophores two regularly or irregularly shaped plates, gene¬

rally with several pyrenoids.

72. Gyrosigma attenuatum (Kuetzing) P. T. Cleve. [Syn.: Pleuro -

sigma attenuatum W. Smith, Navicula attenuata Kuetz. var.

attenuata Kuetz., Pleurosigma attenuatum W. Smith var. caspia

Grun.]. Valves slightly sigmoid, 150-180/i long X 25-27 /a wide,

gradually attenuated to the rounded poles; raphe centrally

placed, central area asymmetric, transverse striae slightly radial,

12-14 in 10/a, longitudinal striae wider apart, 10-12 in 10/x,

(PI. X, fig. c).

Malaya Pahang: Stream near Kuantan.

Malacca: Malacca River.

Worldwide distribution.

The Malayan form is slightly wider than the type and might

possibly be referred to var. hippocampus W. Smith.

73. Gyrosigma distortum (W. Smith) P. T. Cleve. [Syn.: Pleuro¬

sigma distortum W. Smith]. Valves sigmoid, 100/a long X 20-

22/a wide, broad in the middle but sharply narrowed towards the

poles; raphe centrally placed, central area elliptic; transverse

striae marked, 15-16 in 10/a, longitudinal striae finer, 19-20

in 10/a. (PI. X, fig. b).

Malaya Pahang: Stream near Kuantan.

Widespread in distribution.

74. Gyrosigma distortum (W. Smith) P. T. Cleve var. parkeri

(Harrison) P. T. Cleve. [Syn.: Pleurosigma parkeri Harrison].

Differs from the type in the much more sharply narrowed ends;

valves distinctly sigmoid, 150-180/a long X 30-35/a wide, very

sharply narrowed towards the poles; raphe central, central area

elliptic; transverse striae marked, 15-17 in 10/a, longitudinal

striae finer, 19-22 in 10/a. (PI. IX, fig. e).

Malaya Pahang: Stream near Kuantan.

Widespread in distribution.

Gardens Bulletin , S.

75. Gyrosigma excentricum Prowse sp. nov. Valves linear sigmoid,

110-120/t long X 12-13/a wide, gradually narrowed towards

the rounded poles; raphe very oblique, sigmoid, disappearing in¬

to the convex margins below the poles, central area elliptic,

polar nodules very small, intra-marginal and almost invisible;

longitudinal striae very fine, 20-22 in 10/a, transverse striae

slightly wider apart, 18-20 in 10/a. (PL X, fig. d).

Malaya Pahang: Stream near Kuantan. (Type locality).

Holotype No. Prowse 582c is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Valvae lineari-sigmoidea, 110-120/a longae, 12-13/a latae, api-

cem versus utrinque gradatim attenuatae, summo rotundatae;

raphe conformis, ex nodulo centrali ellipsoideo infra polum in

marginem convexam utrinque evanescens; noduli polares minuti,

in margine fere inconspicui; striae longitudinales gracillimae, 20-

22 in 10/a, transversales distantiusculae 18-20 in 10/a.

Malaya Pahang, in fluvio proximo urbe Kuantan.

This species is very distinct with its extremely oblique raphe.

G. rectum (Donk.) P. T. Cleve, which has a similarly oblique

raphe, is not sigmoid but is quite straight.

76. Gyrosigma fasciolum (Ehrenberg) P. T. Cleve var arcuatum

(Donkin) P. T. Cleve. [Syn.: Pleurosigma arcuatum Donk.,

Pleurosigma fasciola W. Smith]. Valves markedly sigmoid, 90-

100/a long X 13-15/a wide, very sharply narrowed towards the

ends so that for a quarter of the length it is linear at each end

and only 2-3 /a wide; raphe central, central area almost circular;

transverse striae 24-26 in 10/a, longitudinal striae about equally

spaced. (PI. X, fig. a).

Malaya Pahang: Stream near Kuantan.

Described from N. Europe and usually regarded as marine, but

many other Baltic forms have been found in fresh water.

77. Gyrosigma scalproides (Rabenhorst) P. T. Cleve var eximium

(Thwaites) P. T. Cleve. [Syn.: Schizonema eximium Thwaites,

Colletonema eximium Thwaites, Endosigma eximium Brebis-

son, Pleurosigma eximium van Heurck.]. Valves linear sigmoid,

50-55/a long X 10-12/a wide, margins parallel except near the

poles where one margin is sharply bent to the rounded wedge-

shaped pole, the other margin only very slightly concave; raphe

central, curving at the poles towards the concave margin, central

area elliptic; transverse striae 22-24 in 10/a, longitudinal striae

26-28 in 10/a. (PI. IX, fig. 1).

Malaya Pahang: Stream near Kuantan.

Cosmopolitan; reported from Indonesia.

Vol XIX. (1962).

78. Gyrosigma spenceril (W. Smith) P. T. Cleve var. smithii

(Grunow) A. Cleve-Euler. [Syn.: Pleurosigma spencerii W.

Smith var. smithii Grunow.]. Valves slightly sigmoid, 100-120/a

long X 13-15/a wide, gradually narrowed to the rounded poles;

raphe central, central area elliptic; transverse striae prominent,

18—22 in 10/a, longitudinal striae closer, 22—24 in 10/a. (PI. IX,

fig. f).

Malaya Malacca: Malacca River.

Widespread in distribution.

PLEUROSIGMA W. Smith 1852, emend. P. T. Cleve 1894

Cells solitary, elliptic-lanceolate in girdle view, intercalary

bands and septa usually absent; valves sigmoid, gradually tapering

to rounded or sub-acute poles; raphe sigmoid, with small central

and polar nodules; axial area narrow, central area usually round;

striations consisting of transverse striae crossed obliquely in two

opposite directions by other striae thus making 3 series of stria¬

tions; chromatophores two, four, or many, regular or irregular

plates.

Two auxospores formed from the conjugation of two cells.

79. Pleurosigma elongatum W. Smith. [Syn. Pleurosigma angu-

latum W. Smith var. elongatum W. Smith.]. Valves long, slightly

sigmoid, uniformly narrowed to the rounded poles, 140-150/a

long X 18—20/a wide; raphe central, central area small, rhombo-

elliptic in shape; transverse striae 18-20 in 10/a, most clearly

visible at the margins, oblique striae wider apart 16-19 in 10/a.

(PI. X, fig. f).

Malaya Pahang: Stream near Kuantan.

Widespread, usually in brackish water.

80. Pleurosigma .salinarum (Grunow) P. T. Cleve. [Syn.: Pleuro¬

sigma delicatulum W. Smith var. salinarum Grunow.]. Valves

broad, slightly sigmoid, gradually narrowed to the rounded

poles, 130-150/a long X 23-26/a wide; raphe central, central

area small elliptic; transverse striae 20—22 in 10/a, oblique

striae finer and closer together, 22-24 in 10/a. (PI. X, fig. e).

Malaya Pahang: Stream near Kuantan.

Widespread in brackish water. Reported from Indonesia.

CALONEIS P. T. Cleve 1891

Cells solitary, free-floating, linear or lanceolate, usually with

convex sides and sometimes with marginal inflations in the valve

view; girdle view rectangular; raphe straight with rounded central

Gardens Bulletin , S.

and polar nodules; transverse striations crossed by one or more

longitudinal lines or hyaline areas parallel to the valve margin;

intercalary bands absent; chromatophores usually two, irregular,

each commonly with two pyrenoids.

81. Caloneis ladogensis P. T. Cleve var. cuneata A. Cleve. Valves

linear elliptic, small 20-25/a long X 8-10/a wide, margins al¬

most parallel in the middle, narrowing to almost cuneate poles;

axial area narrow, central area abruptly widened to form a rect¬

angle or stauros not reaching the margins; striae 16-19 in 10/a,

punctate and crossed near each margin by two hyaline longi¬

tudinal lines parallel to the margin. (PI. XVIII, fig. m).

Malaya Pahang: Cameron Highlands.

Reported from N. Europe.

This form can only come here, corresponding in all ways—

shape, rectangular central area and density of striations.

82. Caloneis silicula (Ehrenberg) P. T. Cleve var. minuta P. T.

Cleve. [Syn.: Navicula ventricosa Ehr. forma minuta Grunow,

Caloneis silicula (Ehr.) P. T. Cleve var. gibba Mayer.]. Valves

small, 28-30/a long X 7 -8/a wide, linear lanceolate with triun-

dulate margins and wedgeshaped poles; axial area narrow, cent¬

ral area a wide stauros reaching the margins; striae parallel,

19-21 in 10/a, finely punctate and crossed by a hardly visible

longitudinal line close to each margin. (PI. XIV, fig. n).

Malaya Malacca: Fish-ponds.

Described from freshwater of Europe, not common.

NEIDIUM Pfitzer 1871

Cells usually solitary and free-floating; girdle view rectangular

without intercalary bands; valve view linear, lanceolate, elliptic or

irregularly undulate with poles acute, rounded, nearly capitate or

rostrate, axial field usually narrow, with rounded or elliptic central

area; raphe straight, usually bifurcated at the poles, with central

ends facing in opposite directions; transverse striae punctate,

crossed near the margin by narrow longitudinal hyaline spaces or

lines; chromatophores two, longitudinally incised and thus often

appearing as four, each with a pyrenoid.

Auxospores two, formed by the conjugation of two approxi¬

mated cells.

83. Neidium hitchcockii (Ehrenberg) P. T. Cleve. [Syn.: Navicula

hitchcockii Ehr.]. Valves linear, 65-70/a long X 16—18/t wide,

with three prominent undulations on each margin, narrowing

to the cuneate, sub-acute poles; axial area narrow, widening at

Vol. XIX. (1962).

the poles, central area obliquely ellipsoid; striae punctate, ob¬

lique, 18-20 in 10/a, crossed near each margin by a longitudinal

hyaline line. (PL IX, fig. a).

Malaya Malacca: Fish-ponds in Malacca and Jasin.

Widespread in distribution.

Hustedt (1938) has described N. gracile from Indonesia, and

for geographical reasons the Malayan forms should be included

under that name. However Hustedt’s reasons for separating off

the species on the grounds of a narrow axial area and it being tro¬

pical, are so unconvincing that I would prefer to keep the species

as N. hitchcockii. Ehrenberg’s original description makes no re¬

ference to the width of the axial area, and other descriptions of

N. hitchcockii from north temperate areas (A. Cleve-Euler 1955,

Tiffany & Britton 1952) refer to a narrow axial area; so there

seems to be no difference between the forms found in temperate

waters and those found in the tropics.

84. Neidium iridis (Ehrenberg) P. T. Cleve var. ampliatum

(Ehrenberg) P. T. Cleve. [Syn.: Navicula iridis Ehr., Navicula

ampliata Ehr., Navicula affinis Greg., Neidium iridis (Ehr.)

P. T. Cleve var. intercedens Mayer.]. Valves elliptic lanceolate,

35-90/a long X 10-20/a wide, subrostrate at the poles, thus

differing from the type species; axial area narrow, widening at

the poles, central area obliquely elliptic; striae punctate, parallel

to oblique, 16—18 in 10/a, crossed near each margin by a longi¬

tudinal hyaline line. (PI. IX, figs, c & d).

Malaya Malacca: Malacca River, fish-ponds in Malacca town

area.

Widespread in distribution.

85. Neidium productum (W. Smith) P. T. Cleve [Syn.: Navicula

producta W. Smith, Neidium productum Hustedt, Neidium capi-

tatum A. Cleve, Neidium capitatum A. Cleve var. prolongata

A. Cleve.]. Valves linear-elliptic, with only slightly convex sides,

75-80/a long X 18-20/a wide; distinctly rostrate with rounded

poles; axial area narrow, wider at the poles, central area round;

striae punctate, parallel, 15-18 in 10/a, crossed near each margin

by a prominent, longitudinal, hyaline line. (PL IX, fig. b).

Malaya Malacca: Malacca River, fish-pond near Jasin.

Probably more widespread in Malaya.

Distribution worldwide.

86. Neidium productum (W. Smith) P. T. Cleve var. minor A.

Cleve. Smaller than the type, 40-45/a long X 8-10/a wide, with

practically straight valve margins and markedly rostrate to sub-

capitate poles; striae 19-20 in 10/x, crossed near each margin by

a longitudinal, hyaline line. (PL XII, fig. p).

Malaya Pahang: Cameron Highlands.

Widespread in distribution.

Gardens Bulletin , S.

DIPLONEIS Ehrenberg 1844

Cells solitary, rectangular in girdle view, valves usually elliptic,

sometimes linear, or with a median constriction; central nodule

quadrate, prolonged towards each pole as a pair of parallel horns

or ribs enclosing the raphe and with a longitudinal furrow on each

side of the horns and central nodule; transverse striations costate

or with coarse punctae or alveolae, often crossed by longitudinal

costae or rows of punctae; chromatophores two, sometimes longi¬

tudinally incised.

87. Diploneis bom bus Ehrenberg var. minor P. T. Cleve. [Syn.:

Navicula gemina Ehrenberg.]. Valves linear elliptic, 40-46/a

long X 9— lOju wide, markedly constricted in the middle to give

the shape of a broad figure eight, narrowing gradually at the

poles to a rounded wedge shape; central nodule large and the

paired ribs or horns straight or only slightly curved; transverse

costae radial, 7-9 in 10/a, each with coarse alveoli, a row of

punctate on each side of the central furrows; costae crossed by

longitudinal costae, 4 in the widest part and 2 at the poles.

(PI. XIII, fig. g).

Malaya Pahang: Stream near Kuan tan.

Described from N. Europe in brackish and marine waters, but in

Malaya found in non-saline water, although near the coast.

88. Diploneis decipiens A. Cleve var. parallel;* A. Cleve. [Syn.:

Diploneis elliptica (Kuetz.) P. T. Cleve var. linearis A. Cleve.].

Valves linear-elliptic, small, 18-20/a long X 10 p wide, ribs or

horns narrow, straight, not widened at the central nodule, longi¬

tudinal furrows straight; transverse costae nearly parallel in the

middle, radial near the poles, 12 in 10/a, each with a row of

coalescing punctae, a single row of punctae on either side of

the central furrows. (PI. IX, fig. i).

Malaya Pahang: Stream near Kuantan.

Described from brackish water in N. Europe.

89. Diploneis interrupta (Kuetzing) P. T. Cleve. [Syn.: Navicula

interrupta Kuetzing.]. Cells elliptic, constricted in the middle,

small, 27—30/a long X 10-12/a wide; ribs or horns almost

straight, central furrows straight; transverse costae radial, absent

opposite the central nodule, 10-12 in 10/a, each with 4 large

coarse punctae or alveolae in the widest part of the valve, 2 at

the poles, and a row of smaller punctae each side of the central

furrows. (PI. IX, fig. k).

Malaya Pahang: Stream near Kuantan.

Described from brackish water in N. Europe.

Vol. XIX. (1962).

90. Diploneis oblongella (Naegeli) P. T. Cleve var. baltica A.

Cleve. [Syn.: Navicula oblongella Naeg., Diploneis ovalis

(Hilse) P. T. Cleve var. oblongella Fontell, Diploneis oblon¬

gella (Naegeli) Cleve var. fossilis (Pantoscek) A. Cleve,

Navicula ovalis var. fossilis Pantoscek.]. Valves linear—elliptic,

30-40/a long X 15-17/a wide, margins only slightly convex,

nearly parallel, poles smoothly rounded; ribs or horns narrow,

straight, furrows narrow, slightly widened in the middle; costae

prominent, almost parallel in the middle, radial at the poles,

12-13 in 10/a, punctate with 12-14 punctae in 10/a short cos¬

tae of 2-3 punctae either side of the furrows. (PI. IX, fig. j).

Malaya Pahang: Stream near Kuantan.

Widespread in freshwaters.

91. Diploneis ovalis (Hilse) P. T. Cleve. [Syn.: Navicula ovalis

Hilse.]. Valves broadly elliptic, 20-25/x long X 12-15/a wide,

tapering gradually to the rounded poles; central nodule large,

ribs or horns narrow, furrows narrow, widening markedly oppo¬

site the central nodule; costae radial, 12-16 in 10/a, punctate,

with 16-20 punctae in 10/a, a row of costae with 1-4 punctae on

each side of the central furrows. (PI. IX, fig. h).

Malaya Pahang: Cameron Highlands, and a stream near

Kuantan.

Selangor: Gombak River.

Malacca: Malacca River.

Doubtless to be found elsewhere in Malaya.

Distribution worldwide in freshwaters.

92. Diploneis parina P. ' 1. Cleve. \^alves elliptic, 23—26/a long

X 14-15/a wide, tapering gradually to the rounded poles; cent¬

ral ribs narrow, slightly wider near the central nodule, central

nodule small; furrows expanded in the centre to enclose a lan¬

ceolate central area about 1/3 the width of the valve; costae

radial, 11-13 in I0p, each with two alternating rows of fine

punctae, a row of short prominent costae on each side of the

central furrows. (PI. XIII, fig. 1).

Malaya Pahang: Stream near Kuantan.

Distribution in European freshwaters.

This species closely resembles D. smithii (Breb.) P. T. Cleve,

but is much smaller, with a smaller central nodule and finer

punctae.

Gardens Bulletin, S.

STAURONEIS Ehrenberg 1843

Cells solitary or united valve to valve in short filaments; girdle

view rectangular, sometimes with intercalary bands; valve view

elliptic, lanceolate or rhombic, often capitate, sometimes with lon¬

gitudinal septa visible at the poles; raphe usually straight, axial

area usually narrow, occasionally wider, always widened sud¬

denly in the middle to form a stauros reaching the margins; striae

parallel or slightly radial, finely punctate; chromatophores two

with two to four pyrenoids.

Auxospores formed in pairs between two conjugating cells.

93. Stauroneis acuta W. Smith. Cells joined at the valves to form

short filaments; valves rhombo-lanceolate, 65-75/* long X

18—20/a wide, somewhat inflated in the middle and tapering to

rounded poles; raphe straight, axial area moderately wide, in

the middle expanded to the margins to form a stauros, central

area of stauros rhombic in shape, lateral limbs wider at the

margin; poles with visible septa; striae radiate, 14-18 in 10/x,

punctate. (PI. XIII, fig. f).

Malaya Malacca: Malacca River.

Widespread in freshwater, although not common.

This is a very characteristic species, especially in shape. The

Malayan specimens have a slightly finer structure than the type,

but hardly sufficiently so to warrant erecting a new variety.

94. Stauroneis anceps Ehrenberg. Valves lanceolate, 90-110/x

long X 18-20/a wide, with distinctly capitate poles; raphe

straight, narrow, axial area narrow, widened in the middle to

form a stauros reaching the margins; striae radial, 16-18 in

10/x, punctate. (PI. XIII, fig. d).

Malaya Malacca: Malacca River.

Widespread in freshwater.

95. Stauroneis anceps Ehrenberg var. gothica A. Cleve-Euler.

Valves elliptic-lanceolate, 28-30//. long X 8-9/x wide, with

markedly rostrate to sub-capitate poles; axial area narrow, stau¬

ros wide, wider at the margins, each limb with 5 short marginal

striae; striae radial, 18-20 in 10/x, finely punctate. (PI. XIII,

% j>-

Malaya Pahang: Cameron Highlands.

Described from N. European freshwaters.

96. Stauroneis anceps Ehrenberg var. hyalina A. Brun and

Peragallo. Valves lanceolate, 45-50/* long X 8-9/x wide, with

poles hardly rostrate but almost acute; axial area very narrow,

limbs of stauros very narrow, only 2 or 3 striae wide; striae fine,

hardly visible, 25-30 in 10/x. (PI. XIII, fig. k).

Malaya Pahang: Stream near Kuantan.

Widespre ad in freshwaters.

Vol. XIX. (1962).

97. Stauroneis obtusa Lagcrstedt. Valves linear lanceolate, 40-

50/x long X 8-10/4 wide, narrowing gradually to the broadly

rounded almost truncate poles, with visible septa at the poles;

axial area narrow, in the middle widening to a prominent stau-

ros the limbs of which are wider at the margins; striae punctate,

radial, 19-20 in 10/4. (PL XIII, fig. i).

Malaya Pahang: Stream near Kuantan.

Widespread in fresh waters but not common.

98. Stauroneis phoenicenteron (Nitzsch) Ehrenberg var. elegans

Prowse var. nov. Valves linear-lanceolate, 90/4 long X

12/4 wide, slightly constricted in the middle, narrowed very

slightly below the rounded apices; axial area narrow, expanded

in the middle to form a stauros opposite each limb of which

are 4-5 short, marginal striae, striae punctate, distinctly radial

18-25 in 10/4. (PL XII, fig. d).

Malaya Malacca: Fish-ponds.

Holotype Prowse No. 504a is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Valvae lineari-lanceolate, 90/4 longae, 12/4 latae, medio leviter

constrictae, infra apices rotundatos paululo angustatae. Striae

punctatae, manifeste radiales, 18-25 in 10/4. Area centralis circum

raphen sita, angusta, medio in staurum dilatata, 4—5 striis brevibus

marginalibus adversus eum ornata.

Malaya Malacca, in stagnis piscosis.

99. Stauroneis phoenicenteron (Nitzsch) Ehrenberg var. halmei

Moelder forma marginestriata A. Cleve-Euler. Valves broadly

lanceolate, 29-30/x long X 9-10/4 wide, with broad, very

slightly rostrate poles; axial area narrow, widening in the middle

to a stauros with 3 short marginal striae opposite each limb of

the stauros; striae lightly radial, punctate, 17—20 in 10/4. (PL

XIII, fig. C ).

Malaya Pahang: Cameron Highlands.

Described from freshwaters in N. Europe.

The Malayan specimens are much smaller than the type. They

are however obviously a form of 5. phoenicenteron and apart from

size agree closely with var. halmei f. marginestriata.

100. Stauroneis phoenicenteron (Nitzsch) Ehrenberg var. inter¬

media (Dippel) A. Cleve-Euler. [Syn.: S. phoenicenteron var.

vulgaris Dippel forma intermedia Dippel, S. phoenicenteron var.

brevis A. Cleve.]. Valves broadly lanceolate, 70-120/x long X

15-25/4 wide, with rounded lightly rostrate poles; axial area

Gardens Bulletin , S.

linear, moderately wide, widening in the middle to form a stau-

ros reaching the lateral margins; striae radial, punctate, 13-16

in 10 M . (PI. XIII, figs, e & h).

Malaya Selangor: Gombak River.

Malacca: Fish-ponds.

Probably widespread in Malaya.

Cosmopolitan in distribution.

101. Stauroneis pusilla A. Cleve var. franconica (Mayer) A.

Cleve-Euler. [Syn.: S. anceps Ehrenberg var. pusilla A. Cleve,

S. franconica Mayer.]. Valves lanceolate, 30—45/x long X 7-

11 p wide, capitate at the poles, axial field narrow, widening in

the middle to form a stauros reaching the margins; striae diffi¬

cult to see, visible only with phase contrast under oil immersion,

more than 30 in 10/x. (PI. XIII, figs, a & b).

Malaya Pahang: Cameron Highlands.

Reported from European freshwaters.

The Malayan specimens are slightly larger than the type, but the

extremely fine striae, and the shape, make it reasonable to include

them under the above variety.

ANOMOEONEIS Pfitzer 1871

Cells generally solitary, rectangular in girdle view, in valve view

lanceolate to rhombic and elliptic, regularly with convex sides

and sometimes with a median inflation, poles acute, rounded or

capitate; axial area usually narrow, with a circular or rhombic

central area, but sometimes widening into a large lyrate area;

raphe straight; transverse striations crossed by longitudinal, zig¬

zag, hyaline lines; intercalary bands absent; chromatophore single,

laminate, longitudinally incised. Two auxospores formed from the

union of gametes in pairs from two sister cells enclosed in a gela¬

tinous envelope.

102. Anomoeoneis brachysira (Brebisson) Grunow. [Syn.: Navi -

cula brachysira Brebisson.]. Valves broadly rhombo-lanceolate,

20-25/x long X 7-8^ wide, poles broadly rounded; axial area

narrow, central area round-rhombic; transverse striae 25-27 in

10/x crossed by two zig-zag hyaline longitudinal lines on each

side of the raphe. (PI. XI, fig. j).

Malaya Pahang: Cameron Highlands.

Widespread in freshwaters.

Vol. XIX. (1962).

103. Anomoeoneis serians (Brebisson) P. T. Cleve var. acuta

Hustedt. [Syn.: Frustulia serians Breb., Navicula serians (Breb.)

Kuetz.]. Valves rhombo-lanceolate, 40-50/x long X 8-9/x wide,

uniformly narrowed to the acute poles; axial area narrow, cent¬

ral area round-rhombic; transverse striae 24-27 in 10/x, crossed

by 2-3 zigzag, hyaline longitudinal lines. (PI. XI, figs, b & c).

Malaya Malacca: Fish-ponds and swamps.

Described from Indonesia.

NAVICULA Bory 1822, emend. P. T. Cleve 1894

Cells generally solitary and free-floating, sometimes aggregated

into irregularly radiating clusters, occasionally joined valve to

valve to form ribbons; girdle view rectangular, with smooth gir¬

dles and without intercalary bands; valve view elongate, lanceo¬

late to elliptic, sometimes with undulate margins, frequently atte¬

nuated to capitate, rostrate or rounded poles; axial field usually

narrow, central area variable, occasionally expanded and fusing

to form large, lyrate, hyaline areas; raphe straight, simple, with

polar and central expansions; transverse striations medianly

radial to parallel, sometimes with regular or irregular cross lines,

in some species pseudocostate through the coalescence of coarse

punctae, but the costae without internal openings. Chromatophores

laminate, two, rarely four to eight, occasionally with one or more

pyrenoids.

Auxospores formed in pairs by the fusion of two gametes from

each of two approximated cells.

A very large and variable genus, probably the largest amongst

the freshwater diatoms and often divided into subgenera.

104. Navicula amphibola P. T. Cleve. Valves elliptic-lanceolate,

28-30^ long X 12—1 3/x wide, narrowing suddenly to rostrate

poles; axial area narrow, central area rectangular, almost stau-

roid; striae radial, punctate, 13-16 in 10^, shorter in the middle.

(PI. XI, figs, k & 1; PI. XII, fig. h).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

The Malayan form is distinctly smaller and more finely punc¬

tate than the type and perhaps it ought to be separated off as a

variety. However the shape, proportions and density of the striae

correspond closely to those of the species.

105. Navicula anglica Ralfs. [Syn.: Navicula tumida W. Smith

var. genuina Grunow, Navicula gastrum Ehrenberg var. anglica

Grunow.]. Valves elliptic-lanceolate, 25-28^ long X 7-9^ wide,

Gardens Bulletin , S.

poles rostrate; axial area narrow, central area small, round;

striae radial well marked, 10-14 in 10/a, shorter in the middle.

(PI. XI, fig. u).

Malaya Selangor: Stream near Klang Gates Reservoir.

Widespread in freshwaters.

106. Navicula antiqua A. Cleve. [Syn.: Navicula dicephala (Ehr.)

W. Smith var. subcapitata Mayer, Navicula pinnularioides Oes-

trup var. continua A. Cleve.]. Valves linear, 18—22 /a long X

7-8/a wide, with parallel or slightly concave sides and decidedly

wedge-shaped poles; axial area narrow, central area round;

striae slightly radial, 8-12 in 10/x. (PI. XI, fig. s).

Malaya Pahang: Cameron Highlands.

Found in N. European freshwaters.

The Malayan specimens fit best into this species, although no

forms with concave sides have been reported from Europe.

107. Navicula arenaria Donkin. [Syn.: Navicula lanceolata Kuetz.

var. arenaria (Donk.) P. T. Cleve.]. Cells in girdle view with

distinctly convex sides, slightly constricted in the middle oppo¬

site the central nodules; in valve view lanceolate with acute

poles, 45-50/a long X 9-10/x wide, often lying slightly tilted la¬

terally due to the convexity of the valve and the poles then

appearing to curve slightly to one side; raphe straight, following

the central line of the valve and curving in to the central nodule,

appearing to curve slightly to one side when the valve is tilted

laterally; axial area narrow, central area small; striae prominent,

coarsely punctate, radial, 8-9 in 10/a, broken by a short, hya¬

line longitudinal line on one side of the central area. (PI. XII,

fig. a).

Malaya Pahang: Stream near Kuantan.

Usually regarded as marine, but here described from non¬

saline water where it was in company with many healthy large

desmids. In some respects the Malayan forms resemble Navicula

lanceolata (Ag.) Kuetzing, but the valves in that species are

straight in girdle view, not convex as in the present forms. Thus

the Malayan forms more closely agree with Navicula arenaria

Donk. from which they differ only by the break in the striae on

one side of the central area. Whether this difference is sufficiently

consistent for varietal status remains to be seen.

108. Navicula brasiliensis Grunow var. minor Prowse var, nov.

Valves elliptic lanceolate, 35-40/a long X 17-18/a wide, slightly

rostrate at the poles; axial field narrow, central area small,

round; striae slightly radial, 9-12 in 10/a, punctate; the punc¬

tate closer together at the margins forming short transverse

Vol. XIX. (1962).

marginal lines, inner punctae wider apart on wavy longitudinal

lines. Differs from the type in being much smaller and with

more rostrate poles. (PI. XII, fig. g)*

Malaya Pahang: Stream near Kuantan. (Type locality).

Holotype Prowse No. 582d is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

This species is usually regarded as marine but is here found in

non-saline water.

A forma typica haec varietas differt natura minore et plure ros-

tratorum apicorum.

Valvae ellipticae-lanceolatae, apicibus rostratiusculis, 35-40/x

longae, 17-18 latae; area axialis angusta, area centralis minuta

rotunda; striae radialiusculae, 9-12 in 10/*, punctatae, puncta in

marginem propinquitatioria versus marginem lineis brevibus trans-

versalibus formatis, et puncta versus partem mediam distantius-

cula lineis longitudinalibus sinuosis formatis.

Malaya Pahang: in fluvio proximo urbe Kuantan.

109. Navicula cancellata Donkin var. apiculata Grunow. Cells in

girdle view with curved edges, concave at poles, slightly con¬

cave in the middle, valve view linear, 40-50/* long X 8-

lOfx wide, with acute wedge-shaped poles appearing one sided

when valve is slightly tilted laterally; axial area narrow, sud¬

denly widened at the central nodule where there is one short

marginal stria on each side; striae very prominent, 8-10/x,

parallel and formed by coalescing coarse punctae longitudinally

streaked. (PI. XI, fig. w).

Malaya Pahang: Stream near Kuantan.

Usually regarded as marine, but here found in non-saline water.

110. Navicula confervacea Kuetzing. Cells united valve face to

valve face to form short ribbons; valves lanceolate, 18-25/x

long X 7-9/x wide; axial area lanceolate, slightly widened at

the central area; transverse striae fine, radial, 18-20 in 10/*.

(PI. XII, figs, n & o).

Malaya Malacca: Fish-ponds, attached to the roots of

Eichornia.

Widespread, especially in the tropics.

This is most easily recognised by its ribbon like habit.

111. Navicula crucifera Grunow. [Syn.: Navicula apiculata

Brebisson, Navicula apiculata Brebisson var. maculifera Gru¬

now.]. Cells in girdle view with convex margins, sometimes

slightly concave in the middle; valves linear-lanceolate, 45-50/*

Gardens Bulletin , S.

Jong X 14-16/a wide, tapering to acute poles; axial area nar¬

row, central area wide, rectangular; striae somewhat radial 6-8

in 10/a very prominent, longitudinally streaked through the

coalescence of coarse punctae, striae opposite central area much

shorter and sometimes unequal in length. (PI. XI, fig. p).

Malaya Pahang: Stream near Kuantan.

Generally regarded as marine'but here found in freshwater in

the company of large desmids.

112. Navicula cryptocephala Kuetzing var. veneta (Kuetzing)

Grunow. [Syn.: Navicula veneta Kuetzing, Navicula lancettula

Schumann, Navicula cryptocephala Kuetz. var. lancettula van

Heurck, Navicula cryptocephala Kuetz. var. pumila (Grun.) P.

T. Cleve.]. Valves linear-lanceolate to rhombo-lanceolate,

23-25/a long X 5-7/a wide, evenly tapered to the roundish

poles; axial area narrow, central area very small; only slightly

widened; striae somewhat radial, fine, 15-16 in 10/a. (PI. XI,

fig. r).

Malaya Pahang: Cameron Highlands.

Widespread in distribution, often in brackish water, but here in

decidedly non-saline water far from the sea.

113. Navicula cuspidata Kuetzing var. ambigua (Ehrenberg) P.T.

Cleve. [Syn.: Navicula ambigua Ehr.]. Valves lanceolate,

95-100/a long X 25-30/a wide, tapering to markedly rostrate to

sub-capitate poles; axial area narrow, central area hardly wider,

but elongated; transverse striae parallel, 17-18 in 10 /i, distinctly

punctate and crossed bv longitudinal striae, 22-25 in 10 /j (PI

XII, fig. e).

Malaya Malacca: Malacca River.

Widespread but not common.

114. Navicula disparata Hustedt. Valves elliptic-lanceolate, 20-

25/a long X 8-10/a wide, tapering to slightly rostrate poles;

axial area lanceolate, widening in the middle to a roundish cent¬

ral area about 1/3 width of the valve; striae radial, fine 20-22

in 10/a at the poles, wider spaced in the middle. (PI. XII, fig. m).

Malaya Pahang: Stream near Kuantan.

Described from Indonesia.

The Malayan specimens are slightly larger than Hustedt’s, but in

all other respects agree.

115. Navicula elegantoides Hustedt. Valves elliptic-lanceolate,

75-80/a long X 30-35/a wide, tapering to the distinctly rostrate

poles; axial area widish, very much widened in the middle to

Vol XIX. (1962).

form a circular central area; striae slightly radial in the middle,

convergent at the poles, 5 in 10/a, thick but apparently smooth.

(PI. VIII, fig. d).

Malaya Malacca: Malacca River.

Described from the Philippines.

The species looks almost costate, but cannot be classed as a

Pinnularia because the pseudocostae have no internal openings.

116. Navicula feuerborni Hustedt. Valves narrow-lanceolate, 40-

48/a long X 6-8 /a wide, evenly tapered to the rounded poles;

axial area moderately narrow, asymmetrical, central area a long

ellipse not clearly set off and distinctly asymmetrical in shape;

raphe straight except at the poles where it is clearly hooked;

striae fine but distinct, radial in the middle, convergent at the

poles; 10-12 in 10/a in the middle, slightly closer together at the

poles; sometimes of uneven length opposite one side of the

central area. (PI. XI, fig. f).

Malaya Pahang: Cameron Highlands.

Described from Indonesia.

117. Navicula gastrum Ehrenberg. Valves broadly elliptic lanceo¬

late, 25-30/a long X 11-15/x wide, with short broadly ros¬

trate poles; axial area narrow, central area irregularly roundish

to rectangular, raphe straight; striae radial, 8-12 in 10/a, of

irregular lengths opposite the central area. (PI. XI, fig. v).

Malaya Malacca: Malacca River.

Cosmopolitan in freshwater.

118. Navicula glacialis P. T. Cleve var. septentrionalis P. T.

Cleve. Valve broadly elliptic-lanceolate, 45-50/a long X 25-

30/a wide; axial area narrow, central area rectangular; striae 9-

10 in 10/a, punctate with the punctae close together forming

short marginal transverse lines, and the inner punctae much

wider apart forming wavy longitudinal lines. (PI. XI, fig. h).

Malaya Pahang: Stream near Kuantan.

Generally described as marine, but here definitely in fresh¬

water.

119. Navicula grevillei (C. A. Agardh) P. T. Cleve. [Syn.:

Schizonema grevillei C. A. Agardh, Schizonema comoides van

Heurck, Navicula rhombica Donkin). Cells in girdle view with

curved margins, convex at the poles, slightly concave in the

middle; valve-view lanceolate to rhombo-lanceolate, 45-55/x

long X 12-15/a wide, tapered to the rounded poles; axial area

narrow, central area small, elliptic; striae fine, punctate, slightly

radial, 18-20 in 10/a, much wider apart opposite the central

area. (PI. XII, fig. b).

Malaya Pahang: Stream near Kuantan.

Generally regarded as marine, but here found in freshwater.

Gardens Bulletin, S.

120. Navicula halophila (Grunow) P. T. Cleve var. subcapitata

Oestrup. Valves lanceolate, 30-35/a long X 8-9/a wide, taper¬

ing to sub-capitate poles, axial area narrow, central area hardly

wider; transverse striae parallel, finely punctate, 19-20 in 10/a.

(PI. XI, fig. t).

Malaya Malacca: Malacca River.

Widespread in distribution, mainly in brackish water.

121. Navicula hungarica Grunow var. luneburgensis Grunow.

Valves lanceolate, 19-20/a long X 5-6/a wide, tapering to al¬

most acute poles; axial area narrow, central area slightly widen¬

ed; striae very prominent, 10 in 10/a, radial and longitudinally

striated through the coalescence of the coarse punctae. (PL XI,

fig. ci).

Malaya Pahang: Stream near Kuantan.

Widespread in distribution.

122. Navicula hustedtii Krasske. Valve elliptic-lanceolate, 15-

20/a long X 5-6/a wide, constricted beneath the distinctly capi¬

tate poles; axial area narrow, central area small, round; striae

radial, 26-30 in 10^, slightly wider apart opposite the central

area. (PI. XII, fig. k).

Malaya Malacca: Fish-ponds.

Widespread in distribution.

123. Navicula lagerheimi P. T. Cleve var. intermedia Hustedt.

Valves rhombic, 24-28/a long X 8-9/a wide, evenly tapered to

the broadly rounded poles; axial area narrow, central area rect¬

angular, stauroid; striae radial, 20-22 in 10/a distinctly punc¬

tate, very short opposite the central area and with a single pro¬

minent pore on one side of the central area. (PI. XI, fig. n).

Malaya Malacca: Malacca River.

Described from Indonesia.

Perhaps this species and its varieties ought to be merged with

Navicula mutica Kuetzing, from which they show little difference.

124. Navicula lyra Ehrenberg var. dilatata A. Schmidt. Valves

elliptic, 50-60/a long X 30-35/a wide, very slightly rostrate at

the rounded poles; axial area linear, widening at die centre to

join the lateral longitudinal hyaline spaces, the lateral hyaline

areas diverging from the middle and then slightly converging

again near the poles, thus forming a lyrate area; striae punctate,

slightly radial, 11 in 10^. (PI. XII, fig. c).

Malaya Pahang: Stream near Kuantan.

Generally regarded as marine, but here found in non-saline

water. A very characteristic form.

Vol XIX. (1962).

125. Navicula lyra Ehrenberg var. ehrenbergii P. T. Cleve. [Syn.:

Navicula lyra typica van Heurck.] Valves elliptic, 55—60/a long

X 23-25/a wide, markedly rostrate at the poles and slightly con¬

stricted in the middle of the sides; axial area linear, fusing in the

centre with the lateral longitudinal hyaline areas, lateral areas

diverging only very slightly in the mid&e and converging slightly

at the poles, forming a lyrate shape; striae punctate, slightly

radial, 12-14 in 10/a. (PI. XI, fig. o).

Malaya Pahang: Stream near Kuantan.

Usually regarded as marine.

126. Navicula microcephala Grunow. Valves narrow-lanceolate,

18-30/a long X 3/a wide, with sub-capitate poles; axial area

narrow, linear to linear lanceolate; transverse striae fine, 30 or

more in 10/a, strictly parallel. (PI. XII, fig. q).

Malaya Malacca: Fish-ponds.

Widespread in freshwaters, but not common.

127. Navicula minima Grunow. Valves linear-elliptic, 14—16/a

long X 4—5.5/a wide, with broad rounded poles; axial area

narrow, central area rectangular; transverse striae somewhat

radial, 26 in 10/a, shorter and wider spaced opposite the central

area. (PI. XI, fig. z).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

128. Navicula minuscula Grunow. Valves lanceolate, 12—15/a

long X 4-5/a wide, narrowing gradually to the slightly rostrate,

rounded poles; axial area very narrow, central area no wider

or even slightly constricted; striae hardly radial, fine, more than

30 in 10/a. (PI. XI, fig. y).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

129. Navicula ostcnfeldii Oestrup. Valves small, lanceolate, 25-

28/a long x 6-7 /a wide, with capitate poles; axial area nar¬

row, central area only slightly wider; striae fine, strictly parallel,

25-30 in 10/a. (PI. XII, fig. I).

Malaya Malacca: Malacca River.

Freshwater, widespread.

The Malayan forms are slightly larger than the type.

130. Navicula placenta Ehrenberg var. obtusa Meister. Valves

broadly elliptic-lanceolate, 50-55/a long X 25—30/a wide, gra¬

dually narrowing to the broadly rounded poles; axial area nar¬

row, central area small, round; striae punctate forming three

Gardens Bulletin, S.

series consisting of transverse lines crossed by two sets of

oblique striae and punctae widest apart in the middle and much

closer together near the margins and the poles. (PI. XI, fig. g).

Malaya Malacca: Malacca River.

So far only described from Asia, including Indonesia.

An unmistakable form.

131. Navicula platystoma Ehrenberg. [Syn.: Stauroneis platystoma

Ehrenberg.]. Valves broadly lanceolate, 20-25/x long X 10—14/jt

wide, distinctly rostrate at the poles; axial area narrow, central

area lanceolate, laterally elongated; striae radial, 15-19 in 10/x,

shorter opposite the central area. (PI. XI, fig. i).

Malaya Selangor: Mountain streams near Klang Gates Re¬

servoir.

This form closely resembles var. minor Schulz, from which it

differs only in being much smaller. Whether it should be classed

as a separate variety is a moot point.

132. Navicula punctulata F. Smith. Valves elliptic, 24-27/x long

X 14—15 /a wide, with broadly rounded poles; axial area nar¬

row, central area circular; striae radial, 12-14 in 10ft, punctate

with the punctae equidistant on the striae. (PI. XI, fig. di).

Malaya Pahang: Stream near Kuantan.

Most forms described from marine or brackish water habitats.

The present form is small and was found in non-saline water.

Possibly it should be separated as a variety.

133. Navicula pupula Kuetzing var. capitata Hustedt. Valves

linear or almost so, 40-45/x long X 10ft wide, with slightly

convex margins and slightly sub-capitate poles; axial area nar¬

row, but widening at the poles; central area somewhat rect¬

angular, but lateral “wings” wider near the margins; striae

radial, 13-15 in 10/* in the middle, 22-24 in 10 /* near the

poles, short opposite the central area, and distinctly curving out¬

wards near the pole leaving a wide polar area. (PI. XI, fig. a).

Malaya Malacca: Fish-ponds.

Widespread in distribution.

The present form is somewhat intermediate between var. capitata

Hustedt and var. rectangularis (Greg.) Grunow, but comes closer

to the former.

134. Navicula pusilla W. Smith. Valves elliptic-lanceolate, 35-40 /*

long X 18-20/t wide, distinctly rostrate, almost citriform in

appearance; axial area narrow, central area moderately large.

VoL XIX. (1962).

elliptic; striae punctate, radial 12-18 in 10/a, widest apart in the

middle with mixed long and short striae opposite die central

area, punctae closer together, 18-22 in 10/a. (PI. XII, fig. f).

Malaya Pahang: Stream near Kuantan.

Widespread in brackish water but also reported from freshwater.

The present form must come here, even though the central area is

of different shape.

135. Navicula radiosa Kuetzing var. minutissima (Grunow) P. T.

Cleve. [Syn.: Navicula tenella Brebisson var. minutissima Gru¬

now.]. Valves narrow-lanceolate, 27-30/a long X 5—6/x wide,

with rounded poles; axial area narrow, central area round; striae

radial in the middle, convergent at the poles, 18-19 in 10/x.

(PI. XI, fig. m).

Malaya Perak: Taiping lakes.

Widespread in freshwaters.

136. Navicula rhyncocephala Kuetzing. Valves lanceolate, 28-30 /a

long X 6-8/a wide, with slightly elongated, capitate poles; axial

area narrow, central area elliptic to circular; striae well-marked,

radial in the middle, somewhat convergent at the poles, 12-14

in 10/a, closer together at the poles. (PI. XI, figs, d & q).

Malaya Pahang: Cameron Highlands.

Widespread in freshwaters.

The Malayan form is a small form, but differs only in size.

137. Navicula rotacana (Rabenhorst) Grunow. [Syn.: Stauroneis

rotaeana Rabenhorst, Stauroneis ovalis Gregory.]. Valves

small, broadly elliptic, 14-16/* long X 8-9/a wide, with broadly

rounded poles; axial area moderately wide, central area stau-

roid, not reaching the margins; striae fine, somewhat radial,

25-28 in 10/a, shorter and slightly wider apart opposite the

central area. (PI. XI, fig. x).

Malaya Selangor: Mountain stream near Klang Gates reser¬

voir.

Widespread in distribution.

138. Navicula schroeteri Meister. Valves linear-elliptic, 40-50/a

long X 8-10/a wide, with broadly rounded poles; raphe straight

with a one-sided central pore; axial area narrow, central area

fairly large, elliptic; striae distinctly radial, 12-14 in 10/a, coar¬

sely punctate, but the punctae coalescing to give each stria a

longitudinal line effect. (PI. XI, fig. e).

Malaya Malacca: Fish-pond, Lorong Pandan, Malacca.

Described from Asia, including Indonesia.

Gardens Bulletin , S.

139. Navicula subtilissima P. T. Cleve. Valves linear, 20-25/a

long X 3-5 ix wide, with capitate poles; axial area narrow,

central area only slightly wider; striae fine, only slightly radial,

more than 40 in 10/a, hardly visible except in the middle where

they are slightly wider apart. (PI. XII, fig. i).

Malaya Negri Sembilan: Swamps at Port Dickson.

Malacca: Fish-ponds and swamps.

Johore: Swamps at Yong Peng.

Singapore: Swamps.

Probably widespread.

Cosmopolitan.

140. Navicula tenera Hustedt. [Syn.: Navicula uniseriata Hustedt.]

Valves elliptic, 13-18/a long X 6-8 /a wide, with broadly round¬

ed poles; raphe straight, central poles rather wide apart, axial

area lanceolate, asymmetric with a row of punctae on one side

of the raphe ending at the central pore; transverse striae radial,

16-18 in 10/a, broken on each side by a longitudinal hylaine

curved line parallel to the margin. (PI. IX, figs, g & m).

Malaya Pahang: Stream near Kuantan.

Described from Indonesia.

A very characteristic form with the asymmetrical row of

punctae.

141. Navicula toornensis P. T. Cleve. Valves small, 17-18/a long

X 7-9 /a wide, rhombic-elliptic with hardly rostrate, rounded

poles; axial area narrow, central area small, hardly wider than

the axial area; striae radial, 14-16 in 10/a, distinctly punctate,

punctae about 14 in 10/a. (PI. XI, fig. ai).

Malaya Pahang: Stream near Kuantan.

N. Europe in lightly brackish to freshwater.

142. Navicula tridentula Krasske. Valves small, 11-14/a long X

2-3/a wide, linear lanceolate with distinctly triundulate margins

narrowing to the rounded rostrate poles; axial area narrow

linear, widening in the middle and sometimes almost reaching

the margins; transverse striae extremely fine, not distinguishable

even under phase contrast. (PI. XII, fig. j).

Malaya Malacca: Swamps.

Widespread in boggy regions.

143. Navicula trituberculata Prowse sp. nov. Valves elliptic, 20/a

long X 10 /a wide, laterally sinuate, three-humped on each side,

attenuate and rounded at the poles; raphe straight; axial area

Vol. XIX. (1962).

narrow, in the middle expanded to form a stauros; striae 18-19

in 10[x, distinctly radial, the central ones being shorter. (PI. XI,

fig. bi).

Malaya Pahang: Stream near Kuantan. (Type locality).

Holotype Prowse No. 582e is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Valvae ellipticae, lateraliter sinuatae utrinque trituberculatae,

apice utrinque attenuatae, rotundatae, 20/i longae, 10 fx latae. Ra¬

phe recta; area axialis angusta medio stauro-formiter ampliata;

striae 18-19 in 10/a, manifeste radiales, in centro breviores.

Malaya Pahang: in fluvio proximo urbe Kuantan.

PINNULARIA Ehrenberg 1840

Cells solitary and free-floating, rarely in short filaments, sym¬

metric and rectangular in girdle view, girdle smooth, without in¬

tercalary bands; valves usually with straight or nearly straight

sides, sometimes medianly inflated or undulate, usually with

broadly rounded poles; axial field usually broad, expanded both

at the poles and in the middle, with a complex sigmoid or straight

raphe; face of valve crossed by distinct transverse costae with

internal openings; costae smooth, radial or parallel (usually con¬

vergent at the poles, and with two longitudinal lines often visible in

the costate part of the valve; chromatophores two, laminate,

usually with pyrenoids.

144. Pinnularia appendiculata (C. A. Agardh) P. T. Cleve var.

budcnsis Grunow. [Syn.: Frustulia appendiculata C.A. Ag.]

Valves lanceolate, 40/a long X 8-9/*, wide, slightly convex on the

margins, slightly constricted below the sub-capitate poles; axial

area narrow but broadening to a lanceolate central area, with a

wide stauros reaching the margins; raphe straight, curved

slightly only at the polar and central nodules; costae distinctly

radial, convergent at the poles, 18-20 in 10 /a. (PI. XIV, fig. k).

Malaya Pahang: Cameron Highlands.

Cosmopolitan.

145. Pinnularia biceps Gregory var. minor (Boye Petersen) A.

Cleve. [Syn. Pinnularia interrupta W. Smith f. minor Boye

Petersen, Pinnularia interrupta W. Smith v. stauroneiformis

Hustedt.]. Valves linear, 30-35 /a long X 7-8/a wide, with

straight margins sharply narrowed to the sub-capitate poles;

axial area narrow, widening to a lanceolate central area with a

very wide stauros reaching the margins; costae radial, conver¬

gent at the poles, 14-16 in 10/a. (PI. XIV, fig. m).

Malaya Malacca: Fish-ponds and the Malacca River.

Cosmopolitan.

Gardens Bulletin, S.

146. Pinnularia bogotensis (Grunow) P. T. Cleve var. continua

A. Cleve. [Syn.: Navicula bogotensis Grun.]. Valves linear,

90-100/a long X 13-15/a wide, with the middle slightly swollen,

ends straight, poles broadly rounded; raphe straight, doubly

forked at the polar nodules; axial area narrow, central area wide,

elliptic; costae radial, convergent at the poles, 14-16 in 10/a.

(PL XIV, fig. h).

Malaya Malacca: Fish-ponds.

Cosmopolitan.

147. Pinnularia borealis Ehrenberg. Valves linear-elliptic. 40-45 /a

long X 9-10/a wide, with broadly rounded poles; raphe slightly

curved, with one sided central pores and bifurcate at the poles;

axial area narrow, slightly widened in the middle; costae prac¬

tically parallel, shorter in the middle, very thick, 4—6 in 10/a.

(PL XIV, fig. q).

Malaya Malacca: Malacca River.

Widespread in fresh waters.

148. Pinnularia braunii (Grunow) P. T. Cleve. [Syn.: Navicula

braunii Grun.]. Valves elliptic-lanceolate, 65-70/a long X 10-

12/a wide, narrowing slightly below the sub-capitate poles;

raphe straight, axial area more or less wide lanceolate, widening

to the margin in the centre to form a wide stauros with very

short lateral arms; costae radial, convergent at the poles shorter

near the middle, 10-12 in 10/a. (PL XIV, fig. a).

Malaya Perak: Taiping lakes.

Pahang: Cameron Highlands.

Malacca: Fish-ponds.

Probably widespread in Malaya.

Cosmopolitan.

149. Pinnularia braunii (Grunow) P. T. Cleve var. amphicephala

(Mayer) Hustedt. [Syn. Pinnularia amphicephala Mayer.].

Valves linear-lanceolate, 35-75/a long X 8-1 I/a wide, with less

convex margins and more capitate poles than the type; axial

area wide, lanceolate, central area a wide stauros with short

lateral limbs; costae radial, convergent at the poles, shorter near

the middle, 10-14 in 10/a. (PL XIV, fig. 1; PL XV, fig. e).

Malaya Perak: Taiping lakes.

Pahang: Cameron Highlands.

Malacca: Fish-ponds.

Probably widespread in Malaya.

Cosmopolitan.

Vol XIX. (1962).

150. Pinnularia brebissonii (Kuetzing) P. T. Cleve van hybrida

(Grunow) A. Cleve. [Syn.: Navicula brebissonii Kuetz., Navi -

cula bicapitata (Lagerstedt) P. T. Cleve var. hybrida Grunow.].

Valves linear elliptic, 25-30 /a long X 7-9 /a wide, with slightly

convex margin narrowing sharply to the subrostate poles; axial

area narrow, widening in the middle to a stauros reaching the

margins; costae radial in the middle, convergent at the poles,

9-13 in 10/*. (PI. XIV, fig. o).

Malaya Perak: Taiping lakes.

Distribution worldwide.

151. Pinnularia brevicostata P. T. Cleve. Valves linear, 90-100 /a

long X 18-20/a wide, with broadly rounded poles; axial area

very wide, much more than 1/3 width of the valve, lanceolate;

costae short, marginal, parallel to verv slightly radial, 8-10 in

10/a. PI. XV, fig. g).

Malaya Perak: Taiping lakes.

Widespread in distribution.

152. Pinnularia gibba W. Smith var. interrupta A. Cleve-Euler.

Valves narrow linear-lanceolate, 65-70/* long X 8-10/a wide,

slightly swollen in the middle and tapering gradually to the

broadly sub-capitate poles; raphe linear, hooked at the poles;

axial area narrow, widening slightly towards the middle where

it reaches the margin over a wide area; transverse costae radial

in the middle, becoming parallel nearer the poles and conver¬

gent at the apices, 9-11 in 10/a. (PI. XIV, fig. b).

Malaya Perak: Taiping lakes.

Distribution worldwide.

153. Pinnularia hemiptera (Kuetzing) Rabenhorst. [Syn.: Navicula

hemiptera Kuetzing, Pinnularia hemiptera Mayer, Pinnularia

hybrida A. Cleve.]. Valves linear-lanceolate, 65-70/a long X

12-15/a wide, narrowing slightly to the rounded poles; raphe

linear, hooked at the poles; axial area lanceolate, about 1/3

width of the valve; costae parallel to very slightly radial, con¬

vergent at the poles, 8-12 in 10/a. (PI. XV, fig. c).

Malaya Perak: Taiping lakes.

Widespread in distribution.

154. Pinnularia legumen Ehrenberg. Valves linear, 70-75/*, long

X 12-15/a wide, with triundulate margins, narrowing below the

broadly capitate poles; raphe slightly curved, hooked at the

poles, curved to one side at central pores; axial area wide, about

1/3 width of the valve, central area large, rhombic, almost

reaching the margins; costae markedly radial in the middle, con¬

vergent at the poles, 8-12 in 10/a. (PI. VIII, fig. e).

Malaya Perak: Taiping lakes.

Worldwide in distribution.

Gardens Bulletin, S.

155. Pinnularia macilenta (Ehrenberg) P. T. Cleve. [Syn.: Navicula

macilenta Ehr.]. Valves linear, 130-170 /a long X 20-25/a wide,

with broadly rounded poles and parallel margins; raphe wide,

bifurcate at the poles, polar areas circular; axial area wide,

linear, about 1/3 the width of the valve; costae parallel, 7-9

in 10/Li, crossed on each side by a longitudinal band. (PI. XVIII,

fig. f).

Malaya Malacca: Malacca River.

Widespread in distribution.

156. Pinnularia mesolepta (Ehrenberg) W. Smith. [Syn.: Navicula

mesolepta Ehrenberg.]. Valves linear, 100-115/a long X 18-

20/a wide, with markedly triundulate margins, slightly wider in

the middle, and with broadly capitate poles; raphe straight, bi¬

furcate at the poles, polar areas circular; axial area narrow

linear, central area round rhombic, not reaching the margins;

transverse costae radial, shorter in the middle, convergent at

the poles, 10-13 in 10/a. (PI. XVI, fig. p).

Malaya Perak: Taiping lakes.

Widspread in distribution.

The Malayan forms are much larger than the type, and in the

specimens so far examined have alternating short and long

costae opposite the middle of the central area. It remains to be

seen whether these differences are sufficiently consistent to warrant

separation of these forms as a distinct variety.

157. Pinnularia microstauron (Ehrenberg) P. T. Cleve. [Syn.:

Stauroneis microstauron Ehr.]. Valves linear to linear lanceolate,

50-70/a long X 10-14/a wide, with parallel or slightly convex

margins narrowing to subrostrate rounded poles; axial area nar¬

row, linear, widening near the middle to a lanceolate area

reaching the margins, forming a stauros; costae radial in the

middle, convergent at the poles, 10-13 in 10/a. (PI. XIV, figs, c

& e).

Malaya Malacca: Fish-ponds and the Malacca River.

Widespread in distribution.

158. Pinnularia microstauron (Ehrenberg) P. T. Cleve var.

ambigua Meister. Differing from the type in that the margins are

slightly triundulate, and the poles more markedly rostrate; val¬

ves 75-80/a long X 14-16/a wide; transverse costae 10-13 in

1 0/a. (PI. XIV, fig. d).

Malaya Malacca: Malacca River.

Widespread in distribution.

Vol. XIX. (1962).

159. Pinnularia microstauron (Ehrenberg) P. T. Cleve var. brebis-

sonii (Kuetzing) Hustedt. [Syn. Navicula brebissonii Kuetz. pro

parte.]. Valves linear elliptic with only slightly convex sides,

tapering very smoothly to the non-rostrate poles, 60/a long X

10/a wide; axial and central area as in type; costae as in type,

10-13 in 10 ix. (PI. XIV, fig. f).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

160. Pinnularia parva (Gregory) P. T. Cleve var. parvula (Ralfs)

A. Cleve-Euler. [Syn.: Navicula parvula Ralfs, Pinnularia par¬

vula (Ralfs) A. Cleve, Pinnularia parva Gregory.]. Valves lan¬

ceolate, 50/a long X 10/a wide, with subrostrate poles; raphe nar¬

row, hooked at the poles; axial area wide, rhombic lanceolate;

costae slightly radial in the middle, convergent at the poles,

9-11 in 10/a. (PI. XV, fig. b).

Malaya Perak: Taiping lakes.

Widespread in freshwaters.

161. Pinnularia polyonca (Brebisson) P. T. Cleve. [Syn.: Navicula

polyonca Brebisson, Pinnularia mesolepta (Ehr.) W. Smith var.

polyonca (Breb.) Cleve, Navicula mesolepta Ehr.]. Valves

lanceolate, 85-90/a long X 12-15/a wide, a triundulate margin,

distinctly swollen in the middle, and slightly constricted beneath

the capitate poles; raphe narrow, bifurcate at the poles and with

distinct central pores; axial area wide, lanceolate, central area a

wide stauros reaching the margins; costae radial in the middle,

convergent at the poles, 10-12 in 10/a. (PI. XVI, fig. o).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

162. Pinnularia stauroptera (Grunow) P. T. Cleve var. subparal-

lela Mayer. [Syn.: Navicula stauroptera Grun.]. Valves linear,

95-100/a long X 11-14/a wide, with parallel margins slightly

constricted beneath the sub-capitate poles; raphe straight,

slightly forked at the poles, and curved at the central pores,

axial area fairly wide, 1/3 valve width, central area rhombic,

reaching the margins to form a stauros; costae radial in the

middle, convergent at the poles, 10-11 in 10/a. (PI. XV, fig. f).

Malaya Perak: Taiping lakes.

European freshwaters, not common.

163. Pinnularia stomatophoroides Mayer var. nuda A. Cleve-

Euler. Valves sub-linear, 90-95/Along X 14-15/a wide, with tri¬

undulate margins, slightly broader in the middle and slightly

constricted below the rounded sub-capitate poles; raphe straight,

Gardens Bulletin , 5.

forked at the poles, central pores conspicuous; axial area about

1/3 valve width, central area rhombic, reaching the lateral

margins over a narrow area; costae markedly radial in the

middle, convergent at the poles, 10-11 in 10/* (PI. XV, fig. a).

Malaya Perak: Taiping lakes.

European freshwaters, not common.

164. Pinnularia stricta Hustedt. Valves linear, 18—30/x long X

6-7/* wide, with almost straight margins, sometimes very

slightly concave, and broadly rounded poles; raphe straight,

curved to one side at the central pores; axial area narrow,

widening in a lanceolate manner to the very wide stauros reach¬

ing the lateral margins; costae strictly parallel, 12-14 in 10/*.

(PI. XIV, figs, i, j & p).

Malaya Pahang: Cameron Highlands.

Described from Indonesia.

165. Pinnularia tabellaria Ehrenberg. Valves linear, 85-90/* long

X 1(M 3/* wide, slightly swollen in the middle, with sub-capi¬

tate poles; raphe straight, hooked at the poles, curved to one

side at the central pores; axial area moderately wide, i-1/3

valve width, central area elliptic; costae slightly radial in the

middle, convergent at the poles, 12-14 in 10/*. (PI. XIV, fig. g).

Malaya Perak: Taiping lakes.

Tropical freshwaters.

166. Pinnularia trigonocephala P. T. Cleve. Valves large, linear,

130-140/* long X 25-28/* wide, slightly wider in the middle

and at the slightly wedgeshaped, sub-capitate poles; raphe

straight, forked at the poles, and with one sided central pores;

axial area less than i valve width, widening in the middle to the

elliptic central area; costae radial in the middle, convergent at the

poles, 7-8 in 10/*. (PI. XV, fig. d).

Malaya Malacca: Malacca River.

Worldwide in distribution, but not common.

AMPHORA Ehrenberg 1840

Cells usually sessile; girdle view broadly elliptic with truncate

ends, the concave faces of the valves being attached, girdles usually

separated by several punctate or striate intercalary bands; valve

view lunate, longitudinally asymmetric; striae transverse, punctate,

with a strongly excentric axial field nearer the concave margin;

raphe usually nearer the concave margin, with central nodule al¬

most adjacent to the margin; chromatophores single or 2-4.

Auxospores formed in pairs between two conjugating cells, or

formed singly by the union of gametes in a single cell.

Vol. XIX. (1962).

167. Amphora acutiuscula Kuetzing. [Syn.: Amphora lineata

Gregory, Amphora coffaeiformis C. A. Agardh var. salinarum

Grunow, Amphora coffaeiformis C. A. Agardh var. acutiuscula

(Kuetzing) Grunow.]. Cells in girdle view elliptic, narrowed

slightly before the truncate ends, girdle with several intercalary

bands which are striated, 18-20 striae in 10 /*; valve semi-lanceo¬

late, 30-60/x long X 6 - 8/1 wide, with only a slightly concave

to straight ventral margin, dorsal margin moderately convex,

narrowing slightly below the sub-capitate poles; raphe close to

the ventral margin, axial area narrow; dorsal striae finely punc¬

tate, 14-18 in 10/t, ventral striae short and fine, closer together.

(PI. XVII, figs, e, f, n, q, v & w; PI. XVIII, fig. b).

Malaya Pahang: Stream near Kuantan.

Widespread in brackish water, but here found in non-saline

water.

168. Amphora angusta (Gregory) P. T. Cleve emend. Cells in

girdle view elliptic-lanceolate, with narrowly rounded ends and

without intercalary bands; valve semi-lanceolate, 18 - 40/1 long

X 2-4/x wide, with acute poles, ventral face only slightly con¬

cave, valve resembling Cymbella; raphe straight, close to vent¬

ral face, axial area wider on the dorsal side of the raphe; dorsal

striae 15-17 in 10/x, ventral striae very short, more or less equal

in number to the dorsal striae. (PI. XVII, figs, g & i; PI. XVIII,

%• g)-

Malaya Pahang: Stream near Kuantan.

Usually regarded as marine but here found in non-saline water.

169. Amphora bitumida Prowse sp. nov. Frustules in girdle view

broadly elliptic, 18-23/x long X 11-12/x wide in the middle,

ends truncate, 3-4/x wide, under each pole slightly arcuate; gir¬

dle with many intercalary bands; valves lunate, 18-23/x long X

5-6/x wide in the middle, markedly convex on the dorsal side,

ventrally distinctly tumid and slightly concave opposite the cen¬

tral nodule, poles rostrate to almost subcapitate; raphe simple,

not sigmoid, axial area narrow on the dorsal side, sometimes

slightly wider near the central nodule, wider and semi-lanceo¬

late on the ventral side, widest in the middle; dorsal striae 17-18

in 10/x, radiate, punctate, ventral striae short, fine 22-25 in

10/x. (PI. XVII, figs, a, b & c).

Malaya Pahang: Stream near Kuantan. (Type locality).

Holotype Prowse No. 582f is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Frustula aspectu zonali lato-elliptica, apice utrinque truncata

3-4/x lata, infra apicem singulam arcuatiuscula, 18-23/x longa, me¬

dio 11-12/x lata, laciniae intercalares plures. Valvae lunate, 18-

Gardens Bulletin , S.

23 fi longae, medio amplissimae 5-6/a latae, dorso fortiter conve-

xae, ventre manifeste tumidae et nodulum centralem adversus

concaviusculae, apice utrinque rostratae, fere capitellatae.

Raphe simplex, haud sigmoidea; area centralis dorso angusta,

interdum apud nodulum centralem latiuscula, ventraliter latior,

semi-lanceolata, medio latissima. Striae dorsales 17-18 in 10/i,

radiatae, punctatae; eae ventrales breves, graciles 22-25 in 10/a.

Malaya Pahang: in fluvio proximo urbe Kuantan.

170. Amphora coffaciformis C. A. Agardh. Cells in girdle view

elliptic, slightly narrowed below the truncate ends, girdles with

several intercalary bands; valves lunate, 25-30/a long X 4—

6/a wide; with slightly concave ventral margin and convex dorsal

margin narrowed slightly below the rostrate poles; axial field

very narrow, raphe straight, near the ventral margin; dorsal

striae 20-24 in 10/x, ventral striae very short and fine. (PI. XVII,

fig. h & o).

Malaya Pahang: Stream near Kuantan.

Widespread, generally in brackish waters.

171. Amphora exigua Gregory. Cells in girdle view lanceolate,

slightly narrowed before the rounded-truncate ends, girdles with

several intercalary bands which are finely striate; valve view semi-

lanceolate, 25-30/a long X 6-7/a wide, with a straight to slightly

convex ventral margin and a convex dorsal margin, both sides

being slightly arcuate beneath the sub-capitate poles; axial area

narrow, raphe straight, close to the ventral margin; dorsal

striae slightly radial, 11-13 in 10/x, punctate, ventral striae very

short, fine and closer together, absent opposite the central

nodule. (PI. XVII, fig. d).

Malaya Pahang: Stream near Kuantan.

A so-called marine species here found in freshwater. It has

been recorded from brackish water in S. Africa (Cholnoky 1959).

172. Amphora holsatica Hustedt. Cells in girdle view elliptic-

lanceolate, slightly narrowing before the truncate ends, girdles

with several intercalary bands which are coarsely punctate;

valve semi-lanceolate, 40—45/a long X 7-9/a wide, ventral mar¬

gin straight or nearly so, dorsal side convex, both sides slightly

narrowing to the small sub-capitate poles; raphe close to the

ventral margin, straight; dorsal striae coarsely punctate, 12-13

in 10/a, punctate about 12 in 10/a, ventral striae very short and

fine, closer together, absent opposite the central nodule (PI

XVII, fig. m).

Malaya Pahang: Stream near Kuantan.

Widespread, usually in brackish water.

Vol. XIX . (1962).

173. Amphora holsatica Hustedt var. mala) ana Prowse var. nov.

Differing from the type by the punctae of the dorsal striae united

into lines nearer the raphe, elsewhere distinct, and below each

stria a dot separated by a hyaline gap. (PI. XVII, fig. 1).

Malaya Pahang: Stream near Kuantan. (Type locality).

Holotype Prowse No. 582g is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

A forma typica haec varietas differt punctis striarum dorsalium

alioqui distinctis, raphen versus in lineas unitis et infra unasquis-

que lineas punctis singulis per intervallos conspicous sejunctis.

174. Amphora fibica Ehrenberg. [Syn.: Amphora ovalis Kuetz.

var. libica P. T. Cleve.]. Cells in girdle view elliptic, with trun¬

cate ends; valve view lunate, 20-25/a long X 6-7/a wide, with

concave ventral margin, convex dorsal margin, tapering gra¬

dually to the rounded poles; raphe near the ventral margin,

curving very slightly towards the dorsal margin at the central

nodule; dorsal striae punctate, 15-18 in 10/a, much shorter

near the central nodule, ventral striae short, shortest at the mid¬

dle, fine and closer together. (PI. XVII, fig. p).

Malaya Pahang: Stream near Kuantan.

Widespread in distribution.

175. Amphora normani Rabenhorst. Cells in girdle view long-

elliptic, slightly narrowed below the truncate ends, girdles with

several fine intercalary bands; valve view lunate, 30-40/a long

X 8-10 /a wide, slightly constricted below the broad sub-capi¬

tate poles; raphe some distance away from the ventral margin,

leaving a wide space, central nodule curving dorsalwards, dorsal

axial area very narrow; dorsal striae radial, 17 in 10/a, shorter

opposite the central nodule, ventral striae very short. (PI. XVII,

fig. r).

Malaya Pahang: Stream near Kuantan.

Widespread in freshwaters.

176. Amphora pcrpusilla Grunow. Cell very small, in girdle view

elliptic, 8-12/a long X 4-5 /a wide, in die present form very

slightly constricted in the middle; valve view lunate, 8-12/a long

X H-2/a wide, with convex dorsal side, with a very shallow

kink in the middle, ventral side straight or hardly concave; raphe

straight, close to the ventral margin; axial area on dorsal side

very narrow, central area unmarked; dorsal striae 20-24 in 10/a

slightly radial, and slightly wider apart in the middle, ventral

margin smooth. (PL XVII, fig. j).

Malaya Pahang: Stream near Kuantan.

Gardens Bulletin , S.

The Malayan form differs slightly from the type, both in the

slight median constriction and in the slightly wider spacing of the

striae in the middle. Should these characters prove constant it

should be separated as a variety.

177. Amphora proteus Gregory. Cells in girdle view linear-elliptic

to almost rectangular with lightly convex margins and broad flat

truncate ends, girdles without intercalary bands; valve view lu¬

nate, 40—60/x long X 7—10/x wide, only slightly curved, with

broad rounded poles; raphe in the middle of the valve distinctly

curved ventralwards from the poles to the central nodule, and

hooked slightly dorsalwards at the central nodule; dorsal striae

slightly radial, 10-13 in 10/x, very coarsely punctate, punctae

arranged more or less in longitudinal lines as well as in trans¬

verse striae occasionally with a hyaline area opposite the central

nodule, ventral striae as long similarly punctate and similarly

spaced but absent opposite the central nodule. (PI. XVII, fig. u;

PI. XVIII, fig. a).

Malaya Pahang: Stream near Kuantan.

Generally regarded as marine, but here definitely in freshwater.

CYMBELLA C. A. Agardh 1830

Cells solitary and free-floating or attached at the ends of gela¬

tinous stalks, or confined in branched gelatinous tubes; girdle view

rectangular with smooth girdles, intercalary bands absent; valve

view asymmetric, sometimes only slightly so, lunate or more rarely

nearly elliptic, rhombic or naviculoid, dorsally convex, ventrally

concave, straight or somewhat convex; axial area wide or narrow,

nearer the ventral margin, central area with or without dots; raphe

curved, with well defined nodules; transverse striations radiate,

sometimes crossed by longitudinal lines; chromatophores a single

expanded plate.

Auxospores formed in pairs between the conjugating cells.

178. Cymbella cuspidata Kuetzing. Valves broadly lanceolate, 35-

40/x long X 15-16/a wide, asymmetrical, narrowing beneath the

subrostrate poles; raphe almost central, slightly curved, axial

area narrow, central area round-rhombic; striae radial, fine, 9-

11 in 10/x in the middle, 12-15 in 10/x near the poles. (PI.

XVII, fig. z).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

Vol. XIX. (1962).

179. CymbeHa javanica Hustedt. Valves lanceolate, 15-20/a long

X 4/x wide, asymmetric, with a markedly convex dorsal margin

and a slightly convex ventral margin, narrowing slightly beneath

the subrostrate poles; raphe hardly curved, axial area narrow,

widening in the middle to form a lanceolate area; striae promi¬

nent, radial, 11-12 in 10/a, slightly wider apart on the ventral

margin. (PI. XVII, fig. k).

Malaya Malacca: Malacca River.

Described from Indonesia.

180. CymbeHa kolbei Hustedt. Valves broadly lanceolate, asym¬

metric, 25-30/a long X 9-11/a wide, dorsal margin very con¬

vex, ventral margin less so, tapering smoothly to the rounded

poles; raphe central, slightly curved, axial area narrow, widen¬

ing near the middle to an elliptic central area; striae finely punc¬

tate, 11-12 in 10/a, slightly radial in the middle, more so at the

poles; a single puncta in the middle of the central area. (PI.

XVII, fig. y).

Malaya Malacca: Malacca River, fish-ponds and swamps.

Described from the Middle East and S. Africa, but probably

more widespread. The Malayan forms seem to fit best into this

species.

181. CymbeHa lanceolata (Ehrenberg) van Heurck. [Syn.: Coc -

conema lanceolata Ehrenberg.]. Valves distinctly asymmetrical,

naviculoid, 70-80/a long X 15-20/a wide, with a convex dorsal

margin, and concave ventral margin slightly tumid in the mid¬

dle, tapering smoothly to the rounded poles; raphe excentric,

slightly curved; axial area narrow, widening slightly towards

the middle; striae at right angles to the raphe in the middle,

radial at the poles, 9-11 in 10/x, markedly punctate, punctae

15-18 in 10/a. (PI. XVIII, fig. 1).

Malaya Malacca: Swampy water.

Widespread in distribution.

182. CymbeHa sumatrensis Hustedt. Valves broadly lanceolate,

35-40/a long X 9-10/a wide, distinctly asymmetric, with a very

convex dorsal margin and a less convex ventral margin, tapering

to the blunt, rounded poles; raphe nearly central, slightly convex

towards the dorsal margin; axial area narrow, widening in the

middle to form an elliptic central area; striae coarsely punctate,

radial, in the middle of the dorsal side 10 in 10/a, closer together

near the poles and on the ventral sides; a single isolated pore

opposite the middle ventral stria. (PI. XVIII, fig. h).

Malaya Malacca: Malacca River.

Described from Indonesia.

Gardens Bulletin , S.

183. Cymbella tumida (Br6bisson) van Heurck. [Syn.: Cocconema

tumidum BrSbisson, Cocconema stomatophorum Grunow.].

Valves broadly lunate, 50-70/x long X 18-20/a wide, with a

very convex dorsal margin, ventral margin straight or slightly

concave, swollen in the middle region, narrowing slightly at

the ends beneath the broad, blunt poles; raphe almost central,

a little nearer the ventral margin, slightly curved; axial area

linear, widening in the middle to the large, broadly rounded

central area; striae coarsely punctate, radial, 8-10 in 10/x in

the middle of the dorsal side, closer together elsewhere; a single

large pore in the middle of the ventral half of the central area.

(PI. XVII, fig. ai).

Malaya Pahang: Cameron Highlands.

Selangor: Gombak River.

Widespread in freshwaters.

A very characteristic species.

184. Cymbella turgida Gregory. [Syn.: Encyonema turgidum

(Greg.) Grunow.]. Valves lunate, 35-40/x long X 11-12/x wide,

with a very convex dorsal margin and a straight ventral margin,

with more or less sub-acute poles; raphe excentric, straight ex¬

cept at the poles, where it curves sharply towards the ventral

margin; axial area narrow, linear, only slightly wider in the

middle; striae on dorsal side radial, 8—10 in 10/x, coarsely punc¬

tate, punctae 17-20 in 10/x, on ventral side convergent and

closer together. (PI. XVII, fig. x).

Malaya Malacca: In a swamp.

Widespread in freshwaters.

185. Cymbella ventricosa Kuetzing. Valves lunate, 15-25/x long

X 5-7/a wide, with a distinctly convex margin and a straight

to slightly convex ventral margin, poles rounded acute; raphe

near the ventral margin, straight; axial area narrow, only slightly

wider in the middle; striae slightly radial, 14-16 in 10/x, finely

punctate. (PI. XVII, figs, s & t).

Malaya Pahang: Cameron Highlands.

Widespread in freshwaters.

GOMPHONEMA C. A. Agardh 1824

Cells usually epiphytic on the ends of dichotomously branched

gelationous stalks, sometimes sessile, more rarely solitary and free

floating; girdle view transversely asymmetrical, cuneate, the nar¬

rower end being the point of attachment; valve view also transver¬

sely asymmetrical, cuneate, clavate, lanceolate or nearly straight

with one pale broader than the other, and often capitate; axial field

narrow, straight, with a straight raphe and conspicuous central and

Vol. XIX. (1962).

polar nodules; striations transverse, parallel or radial; central area

sometimes extending to the margins, with an asymmetrically placed

dot or pore; chromatophore a single lobed plate, usually with a

single pyrenoid.

Auxospores formed in pairs between two conjugating cells.

186. Gomphonema gradle Ehrenberg. Valve view rhombic-

lanceolate, 40-50/a long X 8-10/x wide, almost isopolar with

narrow rounded poles; axial area narrow, linear, with a small

roundish central area, with a single isolated pore on one side;

striae slightly radial to almost parallel, 12-15 in 10/i, not mark¬

edly punctate. (PI. XVI, figs, a & e).

Malaya Malacca: Fish-pond.

Widespread in freshwaters.

187. Gomphonema Iongiceps Ehrenberg var. subclavata Grunow

forma gracilis Hustedt. Valves narrow linear rhombic, 65-70/1

long X 9-10/1 wide, margins slightly concave between the middle

and the narrow rounded poles; axial area narrow, central area

small, with a short marginal stria on each side and an isolated

pore on one side; striae slightly radial, 10-12 in 10/x, lightly

punctate. (PI. XVI, fig. m).

Malaya Pahang: Cameron Highlands.

Widespread in freshwaters, but not common.

188. Gomphonema parvulum (Kuetzing) van Heurck. [Syn.:

Sphenella parvula Kuetzing.]. Cells small, valves elliptic lanceo¬

late, 15-30// long X 4-7 /a wide, more or less rostrate to sub-

capitate at the free pole (i.e. not attached to the gelatinous stalk)

and slightly constricted above the attached pole; axial area nar¬

row, central area small, one-sided, with a single isolated pore on

one side; striae slightly radial to nearly parallel, 13-17 in 10//,

lightly punctate. (PI. XVI, figs, c, d, g, h & i).

Malaya Pahang: Cameron Highlands.

Selangor: Gombak River.

Malacca: Swamps.

Widespread in freshwaters. A very variable species of which

several varieties have been named. The Malayan material has too

many intermediate forms to justify separating any varieties.

189. Gomphonema subtile Ehrenberg. Valves slender, narrow-

lanceolate, 50-65/x long X 7-8/x wide, convex in the middle

but narrowing and slightly concave towards the poles; free pole

distinctly broadly capitate, the attached pole narrow, rounded;

axial area narrow, central area wider, lanceolate, with shorter

Gardens Bulletin , S.

striae on one side, and usually with a single isolated pore, some¬

times with more, sometimes with none; striae slightly radial,

12 in 10 fi 9 punctate. (PI. XVI, figs, b, j, k & 1).

Malaya Perak: Taiping lakes.

Widespread in freshwaters.

The shape is very characteristic, but the usual form has no iso¬

lated pore in the central area; in the Malayan material the other

forms seem equally common.

190. Gomphonema subventricosum Hustedt. Valves lanceolate,

32-40/a long X 7-8/a wide, broadest above the middle, taper¬

ing gently towards the subrostrate free pole, narrowing very

much below the middle towards the narrowly rounded pole of

attachment; axial area narrow, widening slightly in the middle,

where the striae are shorter and with a single isolated pore on

one side; striae slightly radial, 10-12 in 10/a, markedly punctate,

with the punctae drawn out longitudinally. (PI. XVI, figs, f &

n).

Malaya Selangor: Stream near the Klang Gates Reservoir.

Described from Indonesia.

RHOPALODIA O. Mueller 1895

Cells usually solitary and free-floating; girdle view with two

faces, a convex face much wider than the other straight or con¬

cave face; girdle view linear, nearly elliptic, clavate or almost

rounded, often medianly inflated and sometimes with a median

notch; valve view lunate to sickle-shaped, convex margin often

inflated, with acute ends, sometimes elongate, surface distinctly

ridged towards the convex margin; axial field in valve ridge, with

raphe bearing central and polar nodules; both valve and girdle

view transversely costate, with intermediate finer striae; intercalary

bands present or absent; chromatophores single, laminate, with

irregular margins.

Auxospores formed in pairs between two usually different sized

cells, elongating perpendicular to the long axis of the cells.

191. Rhopalodia gibberula (Ehrenberg) O. Mueller. [Syn.:

Epithemia gibberula Ehr.]. Girdle view elliptic, 40— 4-5 /<. long X

22-25/a wide, narrowing to the bluntly rounded ends; valves

sickle-shaped with a very convex dorsal margin having a notch

in the middle and a slightly concave ventral margin; costae

radial, 3-4 in 10/x, each pair with 4-6 striae between them,

striae 15-18 in 10/x on the ventral margin. (PI. XXII, fig. a).

Malaya Selangor: Stream near Klang Gates Reservoir.

Widespread usually in brackish water, but here in undoubtedly

freshwater.

Vol XIX. (1962).

192. Rhopalodia musculus (Kuetzing) O. Mueller. [Syn.: Epithemia

musculus Kuetzing.]. Girdle view broadly elliptic, almost cir¬

cular, 20 /a long X 18/a wide, with broadly rounded poles; val¬

ves broadly lunate, with a very convex dorsal margin, ventral

margin straight except at the poles where it curves sharply vent-

ralwards; costae 3-4 in 10/a, with 3-5 striae between each pair,

striae 16-18 in 10/x on the ventral margin. (PI. XXII, fig. f).

Malaya Pahang: Stream near Kuantan.

Widespread, usually in brackish water.

193. Rhopalodia parallela (Grunow) O. Mueller. [Syn.: Epithemia

gibba Kuetz. var. parallela Grunow.]. Girdle view linear, 65-80 /a

long X 14—15 jx wide, with sides nearly parallel and only

slightly narrowed at the truncate ends; valves half linear-lanceo¬

late, with a very slightly convex, almost straight dorsal margin,

slightly constricted in the middle, and straight ventral margin;

costae slightly radial, 6-8 in 10/a, with 2-4 striae between each

pair, striae 12-15 in 10/a (PI. XXII, fig. b).

Malaya Pahang: Stream near Kuantan.

Widespread, usually in brackish water.

TROPIDONEIS P. T. Cleve 1894

Cells usually solitary and free-floting; girdle view nearly rect¬

angular with a median constriction, to broad hour-glass shaped, a

keel curving in on each margin from the poles to the central no¬

dule, intercalary bands present or absent; valve view lanceolate,

usually acute at the poles, with the raphe in a wing or keel which

may be strictly central, or which may be sigmoid from one pole to

the other; axial field narrow, central area small, transverse striae

marked, sub-parallel to radial. Chromatophores 2, parallel to the

girdle. Mainly a marine genus.

194. Tropidoneis lepidoptera (Gregory) P. T. Cleve. [Syn.:

Amphiprora lepidoptera Gregory.]. Cell in girdle view long-

rectangular, 40-80/a long X 23-25/a wide, markedly constricted

in the middle, poles truncate, rounded; valves linear-lanceolate,

40-80/a long X 12-15/a wide, with narrowed sub-acute poles;

wing containing raphe central, usually seen obliquely slightly to

one side of the longitudinal axis; axial field very narrow, central

area small, round; transverse striae slightly radial, fine, 16-22

in 10/a. (PI. XVIII, figs, c, d, e & i).

Malaya Pahang: Stream near Kuantan.

Usually regarded as marine, but here found in freshwater.

Gardens Bulletin, S.

HANTZSCHIA Grunow 1877

Cells curved in both valve and girdle view, thus having a dorsal

and ventral margin; girdle view elongate-rectangular, somewhat

attenuate; valve view linear-lanceolate to linear, more or less at¬

tenuated to rostrate or capitate poles; raphe of each valve in a

keel near the ventral margin, so that both keels are on the same

side of the longitudinal axis, raphe with circular pores (“carinal

dots”) opening towards the interior of the cells; valves transversely

striate; chromatophore single near one side of the girdle.

195. Hantzschia amphioxys (Ehrenberg) Grunow var. capitata O.

Mueller. [Syn.: Eunotia amphioxys Ehr.]. Valves linear-lanceo¬

late, 35-40/1 long X 5/* wide, slightly curved, and narrowing to

the small capitate poles; carinal pores 8 in 10/*, much wider

apart in the middle and with a prominent central nodule; striae

22-25 in 10/*. (PI. XIX, fig. t).

Malaya Selangor: Stream near Klang Gates reservoir.

Widespread in freshwaters.

196. Hantzschia amphioxys (Ehrenberg) Grunow var. vivax

(Hantzsch) Grunow. [Syn.: Eunotia amphioxys Ehr., Nitzschia

vivax Hantzsch, Nitzschia elongata Hantz. var. minor Mayer.].

Valves linear, 180-225/x long X 10/x wide, slightly curved and

attenuated to the sub-acute rostrate poles, ventral margin con¬

stricted in the middle; carinal pores 6 in 10/*, central nodule

prominent, striae 18-20 in 10/*. (PI. XIX, fig. r).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

NITZSCHIA Hassall 1845

Cells solitary and free-floating or densely clustered in simple or

unbranched gelatinous tubes; girdle view elongate-rectangular, or

sigmoid, with somewhat attenuated poles; valves usually longitudi¬

nally asymmetric, rarely symmetric, very variable in shape, straight,

sigmoid, linear, elliptic, sometimes undulate, with or without

median constriction, poles acute, rostrate or capitate, often much

attenuate; raphe in keel which is usually near one margin, rarely

central with small nodules and a row of carinal dots opening to¬

wards the interior of the cell, keels of the two valves on opposite

sides of the longitudinal axis, except where the keel is central;

valve transversely striate or punctate and sometimes costate.

Two auxospores formed by the conjugation of two cells in some¬

what crossed opposition.

Vol. XIX. (1962).

197. Nitzschia acicularis (Kuetzing) W. Smith. [Syn.: Synedra

acicularis Kuetzing.]. Valves 60-65/a long X 5/a wide, spindle-

shaped with markedly attenuate straight ends; carinal pores

small, 12-16 in 10/a, transverse striae more than 30 in 10/a, very

fine, only just visible with phase contrast. (PI. XIX, fig. m).

Malaya Malacca: Swampy rice field.

Widespread in freshwater.

198. Nitzschia amphibia Grunow var. acutiuscula Grunow. Cells

small, valves 18-22/a long X 4/a wide, lanceolate with acute,

wedge-shaped poles; carinal pores small, 7-9 in 10/a, transverse

striae 17-20 in 10/a, punctate. (PI. XIX, fig. g).

Malaya Malacca: Fish-ponds.

Widespread in fresh to brackish water.

199. Nitzschia closterium (Ehrenberg) W. Smith. [Syn.: Ceratoneis

closterium Ehr.]. Valves 35-40/a long X 2-3 /a wide, spindle-

shaped in the middle, but curved in a sickle shape with very long

narrow attenuated ends; carinal pores 15-16 in 10/a, transverse

striae extremely fine and hardly visible. (PI. XX, fig. n).

Malaya Pahang: Stream near Kuantan.

Usually reported from brackish or sea water, but here in fresh¬

water.

A very characteristic form.

200. Nitzschia communis Rabenhorst var. abbreviata Grunow.

Cells very small, valves 6-8/a long X 2.5-3/a wide, elliptic with

narrow rounded poles; carinal pores 12-14 in 10/t, with a gap

in the middle, striae fine, 30 in 10/t. (PI. XIX, fig. j).

Malaya Pahang: Stream near Kuantan.

Widespread.

201. Nitzschia constricta (Gregory) Grunow. [Syn.: Tryblionella

constricta Greg.]. Valves linear-elliptic, 18-22/x long X 8-

9/j. wide, with slightly constricted margins and wedge-shaped

poles; carinal pores small, 10 in 10/x striae 16-20 in 10/i, dis¬

tinctly punctate, the punctae forming oblique lines sloping in

opposite directions to the longitudinal axis. (PI. XIX, fig. d).

Malaya Pahang: Stream near Kuantan.

Usually regarded as marine.

202. Nitzschia diducta Hustedt. Valves broadly linear, 38-40/t

long X 7-8/* wide, with slightly concave to parallel margins,

wedge-shaped ends, and narrow rounded, almost acute poles;

carinal pores 8-10 in 10/t, wider apart in the middle; transverse

striae well marked, 30 in 10/i. (PI. XIX, fig. x).

Malaya Pahang: Stream near Kuantan.

Described from Indonesia.

Gardens Bulletin, S .

203. Nitzschia fonticola Grunow. Cells small, valves 12-15/x long

X 2-4 /I wide, lanceolate, narrowing to the sub-rostrate to sub¬

acute poles; carinal pores 12-14 in 10/x, transverse striae fine

but clearly visible, 28-30 in 10/x. (PI. XIX, figs, a, b & c).

Malaya Negri Sembilan: fish-ponds.

Malacca: fish-ponds, Malacca River.

Widespread throughout the world.

204. Nitzschia gandersheimiensis Krasske. Valves linear, 90-100/x

long X 4/x wide, slightly narrowing to the rounded, hardly ros¬

trate poles; carinal pores very irregular, 8-9 in 10/x, transverse

striae very fine, just visible with phase contrast, more than 30

in 10/x. (PI. XIX, fig. u).

Malaya Malacca: Fish-ponds in Malacca and the Malacca

River.

Widespread.

205. Nitzschia granulata Grunow. Valve elliptic-lanceolate, 20/x

long X 11 /x wide, with rounded poles; carinal pores 7-8 in

10/x, transverse striae 7-8 in 10/x, with very coarse punctate, 6

in 10/x. (PI. XIX, fig. i).

Malaya Pahang: Stream near Kuantan.

Usually regarded as marine.

Cleve-Euler (1952) mentions very short marginal double rows

of fine punctae as well as the coarse ones. These were not visible

in the Malayan material, which agrees in all other respects. It

would however be very difficult to see the fine punctae in a com¬

plete specimen, since the coarse punctae of one valve would hide

the fine punctae in the others. I was not able to observe single

valves, only complete frustules.

206. Nitzschia hetifletiriana Grunow. Valves linear, 90-100/x long

X 4-5/x wide, often slightly curved and distinctly constricted

beneath the capitate poles; carinal pores 9-10 in 10/x, slightly

wider apart in the middle, transverse striae prominent, 15-20

in 10/x. (PI. XIX, fig. v).

Malaya Malacca: Swamp in Malacca.

Widespread.

207. Nitzschia ignorata Krasske. Cell in girdle view linear, sig¬

moid; valve view linear-lanceolate, 60-70/x long X 4/* wide,

distinctly sigmoid and narrowing evenly to the rounded poles;

keel very excentric, slightly kinked in the middle; carinal pores

7-10 in 10/x, wider in the middle, transverse striae fine, about

30 in 10/x. (PI. XIX, fig. s).

Malaya Malacca: Malacca River.

Widespread, usually in brackish water.

Vol. XIX . (1962).

208. Nitzschia littoralis Grunow var. tergestina Grunow. Valves

broad linear-elliptic, 28-30 /a long X 9-10 /a wide, with paral¬

lel to slightly concave margins and wedge shaped ends tapering

to rounded sub-rostrate poles; carinal pores 7-8 in 10/a with a

distinct central nodule; transverse striae 20 in 10/a, with a cent¬

ral narrow, lanceolate, clear longitudinal field less than 1/3

width of the valve. (PI. XX, fig. k).

Malaya Pahang: Stream near Kuantan.

Usually found in brackish water.

This may be identical with Nitzschia calida Grunow.

209. Nitzschia lorenziana Grunow var. subtilis Grunow. Valve

long, linear-lanceolate, 100-110/a long X 4-5 /a wide, distinctly

sigmoid with curved ends tapering to the narrow rounded poles;

keel excentric, carinal pores 8 in 10/a, transverse striae 20 in

10/a. (PI. XIX, fig. q).

Malaya Malacca: Malacca River.

Widespread in fresh to slightly brackish water.

210. Nitzschia marginulata Grunow. Valve broad linear, 100-

11 0/a long X 1 8-20/a wide, with wedge-shape ends tapering to

narrow, rounded poles; carinal pores 9-10 in 10/a, striae 19-22

in 10/a, punctate, valve face with a very broad, smooth, linear-

lanceolate central area stretching from pole to pole. (PI. XXI,

fig. b).

Malaya Pahang: Stream near Kuantan.

Usually found in brackish water.

21 1 . Nitzschia microcephala Grunow. Valves linear, 10-15/a long

X 2.5-3/a wide, with parallel or slightly convex margins, sharply

narrowed to slightly attenuate sub-capitate poles; carinal pores

12-14 in 10/a, transverse striae fine, 32-36 in 10/a. (PI. XIX,

% f).

Malaya Pahang: Stream near Kuantan.

Widespread in freshwaters, but not common, possibly overlooked

because of its small size.

212. Nitzschia navicularis (Brebisson) Grunow. [Syn.: Surirella

navicularis Breb.]. Valves elliptic, 30-35/a long X 15-16/a wide,

with smoothly rounded poles; carinal pores 7-8 in 10/a; valve

face with marginal transverse costae and a central longitudinal

clear area lanceolate in shape, two rows of alternating punctae

on each costae, costae 7-8 in 10/a. (PI. XX, fig. o).

Malaya Pahang: Stream near Kuantan.

Usually marine or in brackish water. The Malayan specimens

have slightly longer costae than the type, but there is no doubt

that they belong under this species.

Gardens Bulletin , S.

213. Nitzschia obtusa W. Smith. Cells long linear, slightly sig¬

moid; valves linear, 180-230/x long X 10-11/x wide, very

slightly constricted in the middle, ends rounded sometimes

slightly wedge-shaped, curving slightly to one side to make the

valve slightly sigmoid; keel excentric, notched in the middle

with a prominent central nodule, carinal pores 5-8 in 10/x,

transverse striae fine, 26 in 10/x. (PI. XX, fig. c).

Malaya Pahang: Stream near Kuantan.

Usually in brackish water or marine.

214. Nitzschia obtusa W. Smith var. scalpelliformis Grunow.

Much smaller than the type, and with a very sharp convex curve

on one side of each end so that the actual pole is narrow and

sometimes almost acute; valves 25-80 /x long X 4-5/x wide; ca¬

rinal pores 8-9 in 10/x, central nodule less well-marked, trans¬

verse striae 28-30 in 10/x. (PI. XIX, fig. o; PI. XX, fig. d).

Malaya Pahang: Stream near Kuantan.

Marine or in brackish water.

215. Nitzschia palea (Kuetzing) W. Smith. [Syn.: Synedra palea

Kuetz.]. Valves linear, 30-50/x long X 3-5/x wide, with ends

narrowing to the hardly sub-rostrate to sub-capitate poles; cari-

anl pores 10-14 in 10/x, transverse striae very fine 33-40 in

10/x. (PI. XIX, fig. k).

Malaya Perak: Taiping lakes.

Pahang: Cameron Highlands.

Malacca: Fish-ponds and Malacca River.

Probably widespread.

Common throughout the world in freshwaters.

216. Nitzschia parvula Lewis. [Syn.: Nitzschia obtusa W. Smith

var. brevissima Grunow.]. Cells small, linear-rectangular in gir¬

dle view, hardly sigmoid; valve view linear, 30-40/x long X

4-5/x wide, slightly constricted in the middle, narrowing at the

ends to the rostrate poles, which curve in opposite drections

making the valve slightly sigmoid; carinal pores 6-8 in 10/x,

transverse striae 29-30 in 10/x. (PI. XIX, fig. h).

Malaya Pahang: Cameron Highlands.

Europe in fresh to slightly brackish water.

217. Nitzschia paradoxa (Gmelin) Grunow. [Syn.: Bacillaria

paradoxa Gmelin.]. Cells united valve face to valve face to form

ribbons which zigzag by one cell gliding longitudinally on the

next; girdle view linear-rectangular, valve view linear, 60-90/x

Vol XIX. (1962).

long X 5-8/1 wide, narrowing in a wedge-shape to the sub¬

acute poles; keel central, carinal pores 8-9 in 10/*, transverse

striae 20-24 in 10/*. (PI. XX, fig. b; PI. XXI, fig. d).

Malaya Malacca: Malacca River.

Widespread, usually in brackish water but has been reported

from freshwater elsewhere.

This is better known as Bacillaria paradoxa Gmelin, the genus

Bacillaria having been erected on the basis of the central keel. In

Nitzschia there are several intergrades from the extreme excentric

position, and it seems rational to include this group under Nitzs¬

chia. The peculiar zigzag gliding motion makes this species easy

to identify and serves to distinguish it from all other species, in¬

cluding those with centrally placed keels.

218. Nitzschia plana W. Smith. Valves linear, 80-90/a long X

12/* wide, slightly narrowed in the middle and with wedge-

shaped poles; keel very excentric, carinal pores 4-6 in 10/*;

transverse striae 18-22 in 10/*, with a linear central area about

1/3 width of valve. (PI. XX, fig. e).

Malaya Pahang: Stream near Kuantan.

Found in brackish to freshwater as well as in the sea, but not

common.

219. Nitzschia punctata (W. Smith) Grunow var apiculata A.

Cleve-Euler. [Syn.: Tryblionella punctata W. Smith.]. Valves

elliptic, 20-25/* long X 12-13/* wide, narrowed at the ends to

the apiculate poles; keel excentric, carinal pores 8-9 in 10/*,

not marked; transverse striae marked, 9-10 in 10/*, curving

near the poles, and almost like costae in appearance. (PI. XIX,

fig. w).

Malaya Pahang: Stream near Kuantan.

Brackish water in Europe, not common.

220. Nitzschia punctata (W. Smith) Grunow var. malayana

Prowse var. nov. Differs from the type in being broad, with

concave margins and broad wedge-shaped ends.

Valves broadly lanceolate, 31/* long X 21/* wide, almost hexa¬

gonal, with slightly concave margins and broad wedge-shaped

ends; keel very excentric, carinal pores 8 in 10/*; transverse striae

prominent, punctate, slightly radial from central line of valve,

14-16 in 10/*. (PI. XX, fig. h).

Malaya Pahang: Stream near Kuantan. (Type locality).

Holotype Prowse No. 582h is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

A forma typica haec varieties differt natura latissima, cum mar-

ginibus concavis et apicibus latis cuneatis.

Gardens Bulletin, S.

Valvae late lanceolate, 31/x longae, 21/x latae, fere hexagonales,

marginibus concavulus et apicibus latis cuneatis. Carina valde ex-

centrica, dentibus 8 in 10/i. Striae transversales prominentes, punc-

tatae, ab axi centrale radiusculae, 14-16 in 10/x.

Malaya Pahang, in fluvio.

221. Nitzschia romana Grunow. Valves lanceolate to linear-

lanceolate, 20-35 /x long X 4-5/x wide, tapering evenly at the

ends to the acute poles; carinal pores 11—13 in 10/x, not very

prominent, transverse striae 24-26 in 10/t. (PI. XX, figs, i & j).

Malaya Pahang: Stream near Kuantan.

Widespread in freshwaters.

222. Nitzschia scalaris (Ehrenberg) W. Smith. [Syn.: Synedra

scalaris Ehr.]. Cells very long, valves linear, 450-500/x long X

16-17/x wide, with wedge-shaped poles, very slightly hooked to

one side; carinal pores prominent, irregular, drawn out trans¬

versely to irregular lengths, 3-5 in 10/x, transverse striae pro¬

minent, fine punctate, 9-11 in 10/x. (PI. XX, fig. g).

Malaya Pahang: Stream near Kuantan.

Usually marine or brackish.

This is a very characteristic species and one of the largest.

223. Nitzschia sigma (Kuetzing) W. Smith. [Syn.: Synedra sigma

Kuetz.]. Cell in girdle view sigmoid with somewhat narrowed

ends; valves linear, 60-100/x long X 3-5 /a wide, tapering to

the narrow poles; more or less sigmoid; keel excentric, carinal

pores 7-10 in 10/x, transverse striae 24-28 in 10/x. (PI. XX,

figs, a & m).

Malaya Pahang: Stream near Kuantan.

Widespread in both brackish and freshwater.

The smaller forms have often been separated off as var. rigidula

Grun., but the differences are not really sufficient to merit such a

separation.

224. Nitzschia sigma (Kuetzing) W. Smith var. clausii (Hantzsch)

Grunow. [Syn.: Nitzschia clausii Hantzsch.]. Cells much shorter;

valves linear, 40/x long X 4 /a wide, with parallel or slightly

concave margins narrowed at the ends to the short rostrate

poles, which point in opposite directions making the valve

slightly sigmoid; carinal pores 10-11 in 10/x, transverse striae

very fine, more than 30 in 10/x. (PI. XIX, fig. 1).

Malaya Malacca: Malacca River.

Widespread in both brackish and fresh water.

Vol XIX. (1962).

225. Nitzschia stagnorum Rabenhorst. Cells in girdle view linear

with parallel, slightly convex or slightly concave margins and

truncate ends; vfelve view linear, 40-45/x long X 6/x wide, ta¬

pering at the ends to the rostrate or sub-rostrate poles; keel nar¬

row, carinal pores 7-9 in 10/x, transverse striae 25-26 in 10/x.

(PI. XIX, fig. n).

Malaya Malacca: Swamps and the Malacca River.

Widespread in freshwaters.

226. Nitzschia subtilis (Kuetzing) Grunow. [Syn.: Synedra sub -

tills Kuetz.]. Valves linear-lanceolate, 90-130/x long X 3-5 ix

wide, evenly narrowed to the almost acute poles; keel excentric,

carinal pores 8-10 in 10/x, transverse striae 28-32 in 10/x. (PI.

XIX, fig. p; PI. XX, fig. f).

Malaya Malacca: Malacca River and fish-ponds.

Widespread in fresh water.

227. Nitzschia surirelloidea Prowse sp. nov. Valves broadly

linear, 270-280/4 long X 18/x wide, long sub-cuneate towards

both poles, slightly narrowed towards the middle; keel very ex-

centric; valve face with costae, striae prominent, solitary in each

costa, 5 in 10/t, reaching almost to the central axis; punctae

very weak, 9-10 in 10/x; carinal pores (teeth) as many as the

striae 5-10/x; central area narrow. (PI. XXI, fig. e).

Malaya Malacca: Malacca River (Type locality).

Holotype Prowse No. 198a is deposited at the Tropical Fish

Culture Research Institute, Malacca, Federation of Malaya.

Superficially this resembles Surirella, but is very long and nar¬

row. The one keel on each valve clearly distinguishes it as a

Nitzschia.

Valvae late linearcs, 270-280/x longae, 18/x latae, apicem versus

utrinque longae subcuneatae, medium versus paululo angustatae.

Carina valde excentrica. Striae prominentes, in unaquaque costa

solitariae, 5 in 10/x, fere axin centralem attingentes; punctis indis-

tinctis, 9-10 in 10/x; dentibus quot striae, 5 in 10/x. Area centra¬

lis angusta.

Superficialiter aspectum Surirellae accedens, sed haec species

longissima ac angustissima.

Malaya Malacca, in fluvio.

228. Nitzschia towutensis Hustedt. Valves small, elliptic, 12-

15 /jl long X 4-6/4 wide, with distinctly convex sides and

short narrow rostrate poles; keel very excentric, carinal pores

small, 15-16 in 10/4, transverse striae fine, 28-30 in 10/x. (PI.

XIX, figs, y & z; PI. XX, fig. 1).

Malaya Pahang: Stream near Kuantan.

Described from Celebes.

Gardens Bulletin, S.

229. Nitzschia tryblionella Hantzsch var. victoriae Grunow.

Valves broad, linear-elliptic, 35—40/x long x 16—17/x wide,

with concave margins and broad wedge-shaped ends; face of

valve costate, costae slightly radial from central line of valve,

5-6 in 10/x; keel very excentric, carinal pores 7-9 in 10/x,

transverse striae very fine, hardly visible, even with phase con¬

trast. (PL XIX, fig. e).

Malaya Pahang: Stream near Kuantan.

Widespread in both brackish and fresh water, but not common.

STENOPTEROBIA Brebisson 1867

Cells long, linear, sigmoid; valves linear with a very narrow

wing on both margins containing raphes, each wing containing a

series of canals connecting to the interior (“wing canals”) sepa¬

rated by semi-circular membranes; thus each valve has two raphes;

face of valve with transverse striae and a central pseudo raphe.

230. Stenopterobia intermedia (Lewis) Fricke. [Syn.: Surirella

intermedia Lewis.]. Valve long, linear, 110-130/x long X 6-

8/a wide, sigmoid and tapering gently near the ends to the

rounded poles; wing canals 4-5 in 10/x, transverse striae 20-24

in 10/x. (PL XXI, fig. a).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

SURIRELLA Turpin 1828

Cells solitary and free-floating; valves elongate, lanceolate, ellip¬

tic or ovate, longitudinally symmetrical, transversely symmetrical

or asymmetrical, valve face flat, undulate or curved; girdle view

rectangular, sinuate or irregular; raphe in a prominent keel or wing

on each margin of the valve, connected by a series of canals

opening to the interior (“wing canals”) separated by semi-circular

membranes, valve face with costae corresponding to the wing

canals; chromatophore single, laminate or appearing as two ex¬

panded lobes.

231. Surirella angusta Kuetzing. Cells in girdle view rectangular

with nearly flat ends, valve view isopolar, linear, 30-50/x long

X 9-10/x wide, with parallel sides, and wedge-shaped ends;

valve wing narrow; costae 6-7 in 10/x, reaching almost to the

middle line, costae and intercostal regions about as wide as

each other. (PL XXIII, figs, b & i).

Malaya Pahang: Cameron Highlands.

Widespread in freshwaters.

Vol. XIX. (1962).

232. Surirella angusticostata Hustedt. Ceils isopolar, girdle view

linear rectangular with curved corners and flat ends; valve view

elliptic-lanceolate, 75-80/i long X 19-20/a wide, tapering to

more or less wedge-shaped poles; wing narrow, wing canals

small, costae very narrow, 4-5 in 10/a, much narrower than the

intercostal region and reaching the middle line. (PI. XXIII,

fig. c).

Malaya Pahang: Cameron Highlands.

Described from Sumatra.

233. Surirella biseriata BrSbisson. Cells very large, isopolar, in

girdle view rectangular with rounded ends; valve view linear to

elliptic-lanceolate, 200-370/a long X 50-80/a wide, with parallel

or slightly convex sides and somewhat wedge-shaped poles;

wings prominent, costae 10-18 in 100/a, almost reaching the

middle line, radial at the poles, costae and intercostal areas

equidistant. (PI. XXII, fig. d).

Malaya Malacca: Malacca River and swamps near Malacca

town.

Widespread in freshwater.

234. Surirella capronii Br6bisson. Cells large, heteropolar, in

girdle view broadly wedge-shaped; valve view ovate, 160-200/a

long X 60-80/a wide, with bluntly rounded poles; wing very

prominent, costae 10-15 in 100/a, nearly reaching the middle

line, radial at the ends, intercostal area narrower than the cos¬

tae; a prominent bulbous-based spine near each end on the

middle line. (PI. XXII, fig. e).

Malaya Malacca: Malacca River.

Widespread in freshwaters.

An unmistakable species.

235. Surirella lemmermanni Hustedt. Cells medium large, valve

view slightly heteropolar, lanceolate, 90-100/a X long 20/a

wide, with keel-shaped ends tapering to narrow poles; wing

narrow, costae 55-70 in 100/a, reaching the middle line and

radial at the ends, costae much wider than the intercostal

areas. (PI. XXII, fig. c).

Malaya Malacca: Malacca River.

Widespread in freshwaters, but not common.

236. Surirella linearis W. Smith. Cells isopolar; valves linear, 40-

100/a long X 12-20/x wide, with parallel or slightly convex

sides and bluntly rounded sub-cuneate ends; wing usually well-

marked, costae 20-40 in 100/a, wider than the intercostal dis¬

tances, almost reaching the middle line, radial at the poles. (PL

XXIII, figs, g & f; PL XXIV, figs, a & c).

Gardens Bulletin, S.

Malaya Penang: Fish-ponds.

Malacca: Malacca River.

Probably widespread in Malaya.

World-wide distribution in freshwaters.

There is some variation in the forms from all parts of the world,

and some of the Malayan forms have a higher ratio of costae than

usual, but there is so much intergrading that it seems unjustifiable

to separate them.

237. Surirella muelleri Hustedt. [Syn.: Surirella constricta O.

Mueller.]. Cells large, isopolar; valve view linear, 200-300/a

long X 45-55/a wide, with markedly concave sides and more or

less bluntly rounded, wedge-shaped ends; wings not very mark¬

ed; costae 16-18 in 100/a, about equal in width to the intercostal

distance, almost reaching the central line, radial at the poles.

(PI. XXIV, fig. d).

Malaya Malacca: Malacca River.

Described from Africa.

The Malayan forms agree so closely with those from Africa that

it is doubtful if they could be included under any other species.

238. Surirella robusta Ehrenberg. Cells large, distinctly hetero-

polar; in girdle view wedge-shaped with flat ends; valve-view

oval-elliptic, 150-250/a long X 50-80/a wide, distinctly hetero-

polar with blunt rounded ends; wings very marked, costae pro¬

minent, 8-15 in 10/a, slightly wider than the intercostal distance,

radial at the poles and nearly reaching the middle line. (PI.

XXI, fig. c).

Malaya Pahang: Cameron Highlands.

Malacca: Malacca River and swamps near Malacca

town.

Widespread in freshwaters.

A characteristic form.

239. Surirella robusta Ehrenberg var. splendida (Ehrenberg) van

Heurck. [Syn.: Navicula splendida Ehr., Surirella splendida W.

Smith.]. Cells smaller than the type, with the costae closer to¬

gether; valves oval-elliptic, 80-120/a long X 25-35/a wide,

distinctly heteropolar, with bluntly rounded ends; wings promi¬

nent, costae 20-25 in 100/a, nearly reaching the middle. (PI.

XXIII, fig. d, PI. XXIV, fig. b).

Malaya Pahang: Cameron Highlands.

Selangor: Gombak River.

Malacca: Malacca River, swamps near Malacca

Town.

Widespread in freshwaters.

Vol. XIX. (1962).

240. Surirella spiralis Kuetzing. Cells isopolar, spirally twisted

around the longitudinal axis, in girdle view shaped like a figure

8 with bluntly rounded ends; in valve view linear-elliptic, 50-

70/a long X 25-30/a wide, spirally twisted into a figure 8, with

with rounded wedge-shaped poles; wings prominent, costae 20-

30 in 100/a, wider than the intercostal distance, not reaching

the middle line, central area more or less linear-lanceolate. (PI.

XXIII, fig. 1).

Malaya Malacca: Malacca River, swamps and fish-ponds near

Malacca Town.

Probably widespread in Malaya.

Distribution worldwide in fresh waters.

241. Surirella tenera Gregory. Cells heteropolar, in girdle view

narrow wedge-shaped with truncate ends; valve view linear

ovate, 100-130/x long X 25-30/a wide, heteropolar, tapering at

the ends to the bluntly rounded poles; wings prominent, costae

20-30 in 100/a, wider than the intercostal distance, radial at the

poles, almost reaching the middle line. (PI. XXIII, fig. e).

Malaya Malacca: Malacca River.

Described from Africa and reported from Indonesia.

242. Surirella tenuissima Hustedt. Cells small, heteropolar, in

girdle view narrow wedge-shaped, with bluntly rounded ends;

valve view narrow ovate, 18-35/a long X 7.5-11/a wide, hete¬

ropolar with rounded truncate poles; wings narrow to moderately

marked, costae 40-70 in 100/a, wider than the intercostal dis¬

tance, reaching or nearly reaching the central line, transverse

striae sometimes visible on valve face. (PI. XXIII, figs, a, h, j &

k).

Malaya Pahang: Cameron Highlands.

Reported from Indonesia and S. America.

CAMPYLODISCUS Ehrenberg 1840

Cells solitary and free-floating, circular or nearly so in outline,

or saddle-shaped, bent in valve face, girdle view varying; raphe in a

narrow wing round valve margin, valves with radiating costae con¬

verging to a hyaline punctate or striate centre; a single broad lami¬

nate chromatophore.

243. Campylodiscus clypeus W. Smith. Valve medium large, a

slightly squarish circle in shape, 45/a in diameter; wing narrow,

costae wide, short, marginal, 15-35 in 100/a, radial and subtend¬

ed by radiating rows of coarse punctae, 7 rows in 10/a, with a

large squarish, smooth, central area; valve face undulate bulg¬

ing in from the costae and out again to the centre, so that some

of the punctae appear faint. (PI. XXII, fig. g).

Malaya Malacca: Malacca River.

Found in slightly brackish to freshwater in Europe.

Gardens Bulletin, S.

Addendum

MASTOGLOIA Thwaites 1856

Cells solitary or in gelatinous tubes, often in structureless gela¬

tinous envelopes; girdle view rectangular, septate with internal

part of septa fusiform and divided into chambers; valve view lan¬

ceolate, elliptic or rhombic, poles broadly rounded, acute or ros¬

trate, with marginal septa and inner septa divided into a series of

submarginal chambers lying in a longitudinal row; axial field

narrow, raphe straight, with small central and polar nodules, valve

face transversely striate; chromatophores two with extensive lateral

projections.

Auxospores formed in pairs by the conjugating of two cells.

244. Mastogloia exigua Lewis. Cells small, in girdle view rectan¬

gular with flat ends; valve view elliptic lanceolate 20-25//. long

X 7—8/a wide, with rounded poles; septum marginal except at

the poles, chambers 4 on each side, slightly rounded on the

inner side; axial field narrow, striae fine, sub-parallel 20-24 in

10/4. (PI. XVIII, figs, j & k).

Malaya Pahang: Stream near Kuantan.

Usually described from brackish water, but here in quite fresh¬

water.

Acknowledgements

To Dr. C. F. Hickling, Fisheries Adviser to the Secretary of

State for the Colonies of the United Kingdom Government, and

first Acting Director of the Tropical Fish Culture Research Insti¬

tute in Malacca, and to the Colonial Office, I wish to express my

thanks for all the encouragement given in this research, and for

permission to publish the results. To Mr. H. M. Burkill, Director

of the Botanic Gardens, Singapore, I am deeply grateful for all the

samples he has collected for me, for affording me facilities for

work in the Botanic Gardens in my early days in Singapore, and

for permitting this paper to be published in the Gardens’ Bulletin.

I am also very grateful to Dr. D. S. Johnson and students of the

Zoology Department in the University of Malaya in Singapore for

the algal samples they collected in the Cameron Highlands and

elsewhere. To Professor H. B. Gilliland and the Department of

Botany at the University of Malaya in Singapore I am extremely

grateful for all the encouragement and help in this research which

they have afforded me. To the University of Malaya library de¬

partment I am particularly grateful for the facilities they have given

me. I am also very grateful to all those people, too numerous to

list here, who have sent me reprints of various papers. To Dr. J.

Lund of the Freshwater Biological Association in England I am

Vol. XIX. (1962).

indeed grateful for the help he has given in identifications. Nor

must I forget Mr. S. F. Owen, of the Public Works Department of

the Federation of Malaya, who so kindly arranged facilities for me

to work in Malacca before the present laboratories were com¬

pleted, and who later sent me so many interesting algal samples.

To Mr. Ratnasabapathy, lately of King Edward VII School, Tai-

ping, and now in New Zealand, I am very grateful for the extremely

interesting collections he made in Taiping. To Mr. Frank Manley,

lately of the office of the Commander Royal Engineers, Malacca,

I owe special thanks for collecting so many samples on the east

side of Malaya. Thanks are also due to my various colleagues, in

particular Mr. George Tay, my laboratory assistant, who has

helped so much in collecting and culturing algae.

Lastly but not least, my thanks are due to Dr. C. X. R. Furtado

of the Botanic Gardens, Singapore, and his son, Mr. J. Furtado

of the Research Institute here, both of whom have so kindly

undertaken the translation into Latin of the diagnoses of new

species and varieties.

Gardens Bulletin, S.

BIBLIOGRAPHY

(this includes only literature actually consulted. )

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Cholnoky, B. J. 1957. Neue and seltene Diatomeen aus Afrika III Dia-

tomeen aus dem Tugeln-Flusssystem, hauptsachlich aus dem

Drakensbergen in Natal. Oest. Bot. Zeitschr. 104 (1-2):

25-99.

Cholnoky, B. J. 1957. Beitrage zur kenntnis der Suedafrikanischen Dia¬

tomeenflora I. Port. Act. Biol. (B) 6 (1): 53-93.

Cholnoky, B. J. 1957. Uber die Diatomeenflora einiger Gewaesser in den

Magalies-Bergcn nahe Rustenburg (Transvaal). Bot. Not. 110

(3): 325-362.

Cholnoky, B. J. 1958. Beitrage zur kenntnis der Sudafrikanischen Dia¬

tomeenflora II: Einige Gewasser in Waterberg—Gebiet,

Transvaal. Port. Act. Biol. (B) 6 (2): 99-160.

Cholnoky, B. J. 1958. Hydrobiologische Untersuchungen in Transvaal

II: Selbstreinigung in Jukskei-Crocodile Flusssystem. Hydro-

biologia 9 (3-4): 205-266.

Cholnoky, B. J. 1959. Neue und seltene Diatomeen aus Afrika IV. Dia¬

tomeen aus der Kaap-Provinz. Oest. Bot. Zeitschr. 106 (1-2):

1-69.

Cholnoky, B. J. 1960. Beitrage zur kenntnis der Diatomeenflora von

Natal (Sud Afrika). Nova Hedwigia 2: 1-128.

Cleve-Euler, A. 1951. Die Diatomeen von Schweden und Finnland Pt. I.

Kungl. Svenska. Vetenskap. Handl: Fjarde Series 2 (1) 163

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V—Schluss. Kungl. Svenska. Vetenskap. Handl: Fjarde se¬

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Cleve-Eijler, A. 1953. Die Diatomeen von Schweden und Finnland pt. II:

Arraphideae, Brachyraphideae. Kungl. Svenska Vetenskap.

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Ill: Monoraphideae, Biraphideae 1. Kungl. Svenska Vetens¬

kap. Handl: Fjarde series 4 (5) 153 pp. 41 pi.

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Gandhi, H. P. 1957. The Freshwater Diatoms from Radhanagari-Kolha-

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(3): 189-193.

Plate I.

(a & b) Melosira granulata (Ehrenberg) Ralfs; (c & i) Melosira roeseana RabenhorsU (d) Melosira ruettneri Hustedt; (e) Cyclotella meneghiniana

Kiietzing; (/) Cyclotella striata (Kiietzing) Grunow; (g & h) Melosira italica (Ehrenberg) Kiietzing; (/, m & l) Coscinodiscus argus Ehren¬

berg; (k & n) Coscinodiscus lineatus Ehrenberg.

Plate 11.

Cyclotella kuetzmg.ana Thwaites; { k) ThalassiosZ flu^

(Kuetzing) Grunow; (c) Coscinodiscus symmetries Kitton; ( e ) Coscinodiscus

antiquus (Grunow) A. Cleve; (/i) Cyclotella meneghiniana (Kuetzing); (i & /)

Hustedt.

Plate III.

(a) Stephanodiscus biserialis Prowse sp. nov.; (b) Stephanodiscus fenestralis Prowse S P- nov ‘> (?) Coscinodiscus decipiens Grunow; (d) Coscino-

discus lineatus Ehrenberg var. van heurckii P. T. Cleve; ( e ) Coscinodiscus excentricus Ehrenberg; (/) Rhizosolenia eriensis H. L. Smith;

(g & h) Chaetoceros amanita A. Cleve.

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1 =

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i —

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-i

i ~~

! i ll 1 1 ! i 1 \ *. '. ' 1 1 * M tTT rn i 1 111 n r 1 1 n n 11 n 11 n 1 1 n 111 i TTTn - n M 11 n i ii n TT i m 1 1 1111111 n m 1111 1 11 1 f!T! 11! I *.!! TTT. r ! TTTTTm

liu uin Min nniiiii.n l l , ,;;; i t l l ' l . ... 1

/^rmrrTTTT

rnMIlllllilTTIIlTn'IlhiiiiiiiiuiiiiT ll ll liiMU ii in i i i iHi i i i ni i i ii i i i TnTTF 1

hiuniliiintutiiiiiiiill nil \\\ .

. llll Hni l iiH 1 n lllll n lllll n lillll l l l H l | l miu ll ll i l lll lll l ll l l H 1 ILiiiUU^ UA ^

Plate IV.

IV.

(o) Fragilaria vaucheriae (Kiietzing) Boye Petersen; (b & c) Synedra tabulate (C. A. Ag.) Kiietzing ^ria^aZlTca^Slunow vif’fcX-

(d) Fragilaria vtrescens Ralfs. var. ellivtica Hn^dt; (e) Aclmanthes subhudsonis Hustedt; (/) Fragilaria lopponica ^unowvjv. tetra^

>g) Boye Petersen; (b & c) Synedra tabulate A lapponica Grunow var. tetra-

' ' - «>••«.«. var. elliptica Hustedt; (^) Aclmanthes subhudsonis Hustedt, (/) 8 Svnndra ulna fNitzscM Eh-

gonalis Prowse var. nov.; (g) Synedra'ulna (Nitzsch) Ehrenberg var. danica (Kiietzing) ^^Achrianthls exigua Grunow (m n &

renberg var . amphirhynchus (Ehrenberg) Grunow; rt / & A) Aclmanthes wolterecku Hustedt, (/) K •

*) Achnanthes tropica Hustedt; (p) Achnanthes tenuissima Hustedt; (?) Achnanthes ^auroneiiorms Prowse

lanceolata (Breb.) Grunow; ( 5 ) Achnanthes lanceolata (Breb.) Grunow var. rostrata Hustedt, ' & u) Cocconels lumens

A. Mayer; (v) Achnanthes crenulata Grunow; (tv) Achnanthes hauckiana Grunow var. rostrata Schulz.

Plate V.

(a) Eunotia monodon. Ehrenberg var. constricta A. Cleve-Euler; ( b ) Eunotia pectinalis (Kiietzing) Rabenhorst; (c & d) Eunotia camelus

Ehrenberg var. karveerensis Gandhi; ( e ) Eunotia grunowi A. Berg var. uplandica A. Cleve f. subundulata A. Cleve; (/) Eunotia luna -

ris (Ehrenberg) Grunow: ( g ) Eunotia monodon Ehrenberg var. undulata A- Cleve-Euler; (h ) Eunotia lunaris (Ehrenberg) Grunow

var. capitata Grunow; (/) Eunotia major (W. Smith) Rabenhorst var. indica (Grunow) A. Berg; (/) Eunotia exigua (Br6bisson)

Grunow; ( k) Eunotia praemonas A. Cleve-Euler var. monodon (Oestrup) A- Cleve-Euler; (/ & r) Eunotia praemonas A. Cleve-Euler

var. infiata (Grunow). A. Cleve-Euler; ( m ) Eunotia faba (Ehrenberg) Grunow var. densestriata Oestrup; («) Eunotia cancellata A.

Berg. var. esseda A. Berg; (o) Eunotia sarekensis A. Cleve-Euler var. minor A. Cleve-Euler; (p & q) Eunotia diodon Ehrenberg var.

minor Grunow; ( s ) Eunotia monodon Ehrenberg var. alpina Kiietzing; (/ & ■ r ) Eunotia sudetica O. Mueller var. incisa (May.) A.

Cleve-Euler; (u & v) Eunotia polydentula (Brun) A. Berg var. perminutn Grunow; (w) Eunotia pectinalis (Kiietzing) Rabenhorst

var. minor (Kiietzing) Rabenhorst; (y & z ) Eunotia camelus Ehrenberg.

Plate VI.

(a, d & e) Eunotia robusta RalfV <u s

(g & h) Eunotia formica Fh & c > Eunotia fiexuosa Kiietzing; (/) Eunotia major (W. Smith) Rabenhorst var. emarginata

nrenberg; (/) Eunotia praemonos A. Cleve-Euler var. tibetica (Mereschk.) A. Cleve-Euler.

A. Cleve;

Plate VII.

(a) Eunotia major (W. Smith) Rabenhorst var. linearis A. Cleve; ( b )

Eunotia gracilis (Ehrenberg) Rabenhorst; (c & d) Desmogonium raben-

horstianum Grunow; (c) general habit of colony.

Plate VIII.

(a & lZ S '“u“ rh ?F£ 0 ii% (Elenberg) de Toni; (c) Frustulia javanica Hustedt; (d) Navicula elegantoides Hustedt; (e) Pinnularia

Tone^feuerbornii HustedT^^ sax <>nica Rabenhorst; (h, i & j) Achnanthes brevipes C. A. Agardh var. intermedia Kuetzing; (k & /)

legu-

C OC¬

Plate IX.

(a) Neidium hitchcockii (Ehrenberg) P. T. Cleve; ( b) Neidium production (W. Smith) P. T. Cleve; (c & d) Neidium iridis (Ehren-

berg) P. T. Cleve var. ampliation (Ehrenberg) P. T. Cleve; ( e) Gyrosigrn a distortum (W. Smith) P. T. Cleve var. parkeri (Har¬

rison) P. T. Cleve; (/) Gyrosigrna spenceri (W. Smith) P. T. Cleve var. smithii (Grunow) A. Cleve-Euler; (g & m) Navicula

tenera Hustedt; ( h) Diploneis ovalis (Hilse) P. T. Cleve; (i) Diploneis decipiens A. Cleve var. parallela A. Cleve; (/) Diploneis

oblongella (Naegeli) P. T. Cleve var. baltica A. Cleve; ( k ) Diploneis interrupta (Kuetzing) P. T. Cleve; (/) Gyrosigrna scalproi-

des (Rabenhorst) P. T. Cleve.

Plate X.

(a)

Gyrosigma

Cleve; (/)

fasciolum (Ehrenberg)

attenuatum (Kuetzing)

Pleurosigma elongatum

P- T. Cleve var. arcuatum (Donkin) P. T. Cleve;

P. T. Cleve; ( d ) Gyrosigma excentricum Prowse

W. Smith.

(b) Gyrosigma distortum (W. Smith) P. T. Cleve; (c)

sp. nov.; ( e) Pleurosigma salinarum (Grunow) P. T.

Plate XI.

(a) Navicula pupula Kuetzing var. capitata Hustedt; ( b & c) Anomoeoneis serians (Br6bisson) P. T. Cleve var. acuta Hustedt;

( d & q) Navicula rhyncocephala Kuetzing; (e) Navicula schroeteri Meister; (/) Navicula feuerborni Hustedt; ( g) Navicula

placenta Ehrenberg var. obtusa Meister; ( h ) Navicula glacialis P. T. Cleve var. septentrionalis P. T. Cleve; (/) Navicula platy -

stoma Ehrenberg; (/) Anomoeoneis brachysira (Brebisson) Grunow; (k & /) forms of Navicula amphibola P. T. Cleve; (m)

Navicula radiosa Kuetzing var. minutissima (Grunow) P. T. Cleve; (n) Navicula lagerheimi P. T. Cleve var. intermedia Hus¬

tedt; (o) Navicula lyra Ehrenberg var. ehrenbergii P. T. Cleve; (p ) Navicula crucifera Grunow; (r) Navicula cryptocephala

Kuetzing var. veneta (Kuetzing) Grunow; ( s) Navicula antiqua A. Cleve; (t) Navicula halophila (Grunow) P. T. Cleve var.

subcapitata Oestrup; (//) Navicula anglica Ralfs; (v) Navicula gastrum Ehrenberg; (>v) Navicula cancellata Donkin var. api-

culata Grunow; ( x ) Navicula rotaeana (Rabenhorst) Grunow; ( y ) Navicula minuscula Grunow; ( z) Navicula minima Grunow;

(ai) Navicula toornensis P. T. Cleve; (bi) Navicula trituberculata Prowse sp. nov.; (ci) Navicula hungarica Grunow var. liine-

burgensis Grunow; (d i) Navicula punctulata W. Smith.

Plate XII.

(d) ,C s“aum^ n p£nSu^i (N£h/ph' 7/ l (C ' A ‘ Ae , ardh) T ' Cleve; ^ Navicula lyraEhrenberg var dilatata A. Schmidt;

(Fhrenberc) P T Cleve' (f\ ' Ehrenberg var. elegans Prowsc var. now.; (e) Navtcula cuspidata Kuetzmg var. ambigua

amphibola P T Cleve' (i)Navirui a p , usi , lla W ' Sm'th; (g) Navicula brasiliensis Grunow var. minor Prowse var. nov.; \h) Navicula

Mbnlissjma P. T. Cleve; (/) Navicula tridentula Krasske; (k) Navicula hustedtii Krasske; (/)

Smhh P T Cleve v^r A ) rw ,C "A &pfl ™' < ’ Husted,; (n & o) Navicula confervacea Kuetzing; (p) Neidium productum\ W

^ieve, ( q ) Navicula microcephala Grunow.

Plate XIII.

(a & b) Stauroneis pusilla A. Cl eve var. jranconica (May.) A. Cleve-Euler; (c) Stauroneis phoenicenteron (Nitzsch) Ehrenberg var. halmei M6el-

der f. marginestriata A. Cleve-Euler; ( d) Stauroneis anceps Ehrenberg; (e & h) Stauroneis phoenicenteron (Nitzsch) Ehrenberg var. inter¬

media (Dipp.) A. Cleve-Euler; (/) Stauroneis acuta W. Smith; ( g ) Diploneis bombus Ehrenberg var. minor P. T. Cleve; (i) Stauroneis

obtusa Lagerstedt; (/') Stauroneis anceps Ehrenberg var. gothica A. Cleve-Euler; ( k ) Stauroneis anceps var. hyalina Brun & Peragallo; (/)

Diploneis parma P. T. Cleve.

Plate XIV.

(a) Pinnularia braunii (Grunow) P. T Cleve* (b) Pinnularia gibba W. Smith var. interrupta A. Cleve-Euler; (c & e) Pinnularia micros¬

tauron (Ehrenberg) P. T. Cleve; id) Pinnularia microstauron var. ambigiui Meister; (/) Pinnularia microstauron var. brebissonii (Kuet-

zing) Hustedt; (g) Pinnularia tabellaria Ehrenberg- (h) Pinnularia bogotensis (Grunow) P. T. Cleve var. continua A. Cleve; (i, j &

p) Pinnularia stricta Hustedt; (k) Pinnularia avvendicula (C. A. Agardh) P. T. Cleve var. budensis Grunow; (/) Pinnularia braunii

(Grunow) P. T. Cleve. var. amphicephala (A. Mayer) Hustedt; (m) Pinnularia biceps Gregory var. minor (Boye Petersen) A.

Cleve; (n) Caloneis silicula (Ehrenberg) P T Cleve var. minuta P. T. Cleve; ( o ) Pinnularia brebissonii (Kuetzing) P. T. Cleve

var. hybrida (Grunow) A. Cleve; ( q) Pinnularia borealis Ehrenberg.

Plate XV.

(a) Pinnularia stornatophoroides Mayer var. nuda A. Cleve-Euler; ( b ) Pinnularia parva (Gregory) P. T. Cleve var. parvula (Ralfs.) A.

Cleve-Euler; (c) Pinnularia hemiptera (Kuetzing) Rabenhorst; ( d) Pinnularia trigonocephala P. T. Cleve; ( e ) Pinnularia braunii (Gru-

now) P. T. Cleve var. amphicephala (A. Mayer) Hustedt; (/) Pinnularia stauroptera (Grunow) P. T. Cleve var. subparallela Mayer;

(£) Pinnularia brevicostata P. T.. Cleve.

Plate XVI.

(a & e) Gomphonema gracile Ehrenberg; (b, j, k & /) Gomphonema subtile Ehrenberg; (c, d, g, h,

& i) Gomphonema parvulum (Kuetzing) van Heurck; (/ & n) Gomphonema subventricosum

Hustedt; (m) Gomphonema longiceps Ehrenberg var. subclayata Grunow f. gracilis Hustedt;

(o) Pinnularia polyonca (Brebisson) P. T. Cleve; ( p ) Pinnularia mesolepta (Ehrenberg) W.

Smith.

(0 ’ h i?jL1"l Ph0ra b !‘Tn da Pr Tn l P J OV -' <{) Amphora exigua Gregory; ( e , /, v & w) ^mp/tora acutiuscula Kuetzing; (, 4

Cv1l!h?nn?v 8 ‘ ta u G f e A? ty n\ P 'j T ' C lev e;(A&o) Amphora coffaeiformis C. A. Agardh; (/) Amphora perpusilla Grunow; \k)

C A y mnh!!r n ^ ’ A ™P ho ™ holsatica Hustedt var. malayana Prowse var. nov.; (m) Amphora holsatica Hustedt; (p)

rl a » ? nber i ; i r) " or 7""' Rabenhorst; & 0 Cymbella ventricosa Kuetzing; (u) Amphora proteus Gre¬

ta isson ) van C pfcurck tUr8tda ° re80ry; (y) Cym6e,/a kolbei Hustedt; ( z ) Cymbella cuspidata Kuetzing; (at) Cymbella tumida (Br6-

Plate XVIII.

(a) Amphora proteus Gregory; (b) Amphora acutiuscula Kiietzing; (c, d, e & i) Tropidoneis lepidoptera (Gregory) P. T. Cleve; (/) Pinnularia

macilenta (Ehrenberg) P. T. Cleve; (g) Amphora angusta (Gregory) P. T. Cleve; (/i) Cymbella sumatrensis Hustedt; (/ & k) Mastogloia exigua

Lewis; (/) Cymbella lanceolata (Ehrenberg) van Heurck; (m) Caloneis ladogensis P. T. Cleve var. cuneata A. Cleve.

Plate XIX.

(a, b & c) Nitzschia fonticola Grunow; ( d ) Nitzschia constricta Gregory) Grunow; ( e) Nitzschia tryblionella Hantzsch var. victoriae Grunow;

(/) Nitzschia microcephala Grunow; (g) Nitzschia amphibia Grunow var. acutiuscula Grunow; (/*) Nitzschia parvula Lewis; (!) Nitzschia

granulata Grunow; (/) Nitzschia communis Rabenhorst var. abbreviata Grunow; (k) Nitzschia palea (Kuetzing) W. Smith; (/) Nitzschia

sigma (Kuetzing) W. Smith var. clausii (Hantzsch) Grunow; (m) Nitzschia acicularis (Kuetzing) W. Smith; ( n) Nitzschia stagnorum Raben¬

horst; ( o) Nitzschia obtusa W. Smith var. scalp ellifor mis Grunow; (p) Nitzschia subtilis (Kuetzing) Grunow; (q) Nitzschia lorenziana Gm-

now var. subtilis Grunow; (r) Hantzschia amphioxys (Ehrenberg Grunow var. vivax (Hantzsch) Grunow; Nitzschia ignorata Krasske;

(0 Hantzschia amphioxys (Ehrenberg) Grunow var. capitata O. Mueller; (u) Nitzschia gandersheimiensis Krasske; (v) Nitzschia heufleuriana

Grunow; ( w) Nitzschia punctata (W. Smith) Grunow var. apiculata A. Cleve-Euler; (x) Nitzschia diducta Hustedt; (y & z) Nitzschia towu-

tensis Hustedt.

Plate XX.

(a f gr !}fl ( K V etzin g) w - Smith; ( b ) Nitzschia paradoxa (Gmelin) Grunow; (c) Nitzschia obtusa W. Smith; (d) Nitzschia

(Fhr(*nht>ro\ SC \l/‘Grunow; (e) Nitzschia plana W. Smith; (/) Nitzschia subtilis (Kiietzing) Grunow; (g) Nitzschia scalaris

n /.x M’t 'u-r* i- £ ltzsc hia punctata (W. Smith) Grunow var. malayana Prowse var. nov.; (/ & j ) Nitzschia romana Gru-

w irYxiti,. uttoralis Grunow var. tergestina Grunow; (/) Nitzschia towutensis Hustedt; (n) Nitzschia closterium (Ehrenberg)

W. Smith, (o) Nitzschia navwularis (Br6bisson) Grunow.

100

Plate XXL

I-1

Mwmniiiiiiui

ilflinMITIIITIITITlIIIIWFTTniTIIMITiriTIITIITTlIIlTIITrTTTITTTnT

ll!III!!lfl!ll7f777/n 2B

(a) Stenopterobia intermedia (Lewis) Fricke; (b) Nitzschia marginulata Grunow; (c) Surirella robusta Ehrenberg; (d) Nitzschia paradoxa

(Gmelin) Grunow, habit of colony; ( e) Nitzschia surirelloidea Prowse sp. nov.

101

Plate XXII.

<rt) ^fi^^^le^sliriren^ca^nifn 5J,V eller; (*) Rhopalodia parallela (Grunow) O. Mueller; (c) Surirella lemmermanni Hustedt; (d) Surirella biseriata

reoisson, (e) Surirella capronu Breb 1SS on; (/) Rhopalodia musculus (Kuetzing) O. Mueller; (g) Campylodiscus clypeus W. Smith.

102

Plate XXIII. /

(a, h, j & k) Surirella tenuissima Hustedt; (b & i) Surirella angusta Kiietzing; (c) Surirella angusticostata Hustedt; ( d ) Surirella robusta Ehrenberg var. splen -

dida (Ehrenberg) van Heurck; ( e) Surirella tenera Gregory; (/ & g) Surirella linearis W. Smith; (/) Surirella spiralis Kiietzing.

103

Plate XXIV.

(a & c ) Surirella linearis W. Smith; (b) Surirella robusta Ehrenberg var. splendida

(Ehrenberg) van Heurck; ( d ) Surirella muelleri Hustedt.

104

Further Malayan Freshwater Flagellata

By G. A. Prowse

Tropical Fish Culture Research Institute,

Batu Berendam, Malacca

Malaya has numerous rivers, swamps, paddy fields and ponds of

various kinds, and one would naturally expect to find an abund¬

ance of Flagellata in these waters. Despite this, very little is

known of the forms which occur in this area, largely because it

has been only in recent years that attention has been turned to

the freshwater microflora of Malaya. In addition, for accurate

determination the Flagellata must be studied in the living state, or

at least freshly killed with osmic acid vapour. Most fixatives, like

Formalin or alcohol, cause such distortion of the organism that

it is recognisable only with great difficulty. Thus, what knowledge

we have of the Flagellata is confined to those forms collected from

easily accessible places, and as more extensive areas are covered

we may expect the number of described species to increase con¬

siderably. References to the Malayan freshwater flagellates are

few and scanty in content, and with the exception of my own

longer paper on the Euglenineae (Prowse 1958), have all been

based on preserved specimens examined far from the native

habitat. The present paper cannot be regarded as in any way

complete, even in those groups discussed in it; the Volvocales,

Xanthophyceae and the Dinophyceae are omitted and will be des¬

cribed in later papers.

Most of the material on which this paper is based has been

examined immediately after collection, both living, and freshly

killed with osmic acid vapour. In some cases, especially with

those organisms inhabiting polluted waters, the material has been

examined again after keeping it in the laboratory for several days.

In a few cases the organisms have been studied in culture, al¬

though in no cases were the cultures absolutely free from other

organisms. Drawings were all carried out with the aid of a camera

lucida, using an Ortholux microscope; any inaccuracies are there¬

fore my own.

Since this paper is not intended to be in any way complete, I

have refrained from constructing keys beyond the level of genera.

Species are listed alphabetically under the genus. The Euglenineae

described here is additional to those mentioned in an earlier paper

(Prowse 1958), and as only one genus, Rhizaspis, has been added

to the earlier list, it has been felt not necessary to construct a new

key.

105

Gardens Bulletin, S .

The Flagellata are so called because of the presence of one or

more protoplasmic threads, the flagella, by means of which the

organism is propelled in the water. Originally the term was used

to distinguish naked flagellate forms from those forms such as the

Volvocales which possess a cell wall. This distinction is no longer

valid, since in the Chlorophyceae there are a few naked flagellate

members, particularly in the Polyblepharidaceae, whilst thalloid

and filamentous development has also been found in other groups

such as the Xanthophyceae, Chrysophyceae and even the Dino-

phyceae. As now used, the term Flagellata embraces a diverse

assemblage of forms belonging to well-defined and differing classes,

and includes as well a somewhat heterogeneous collection of

colourless forms whose affinities are uncertain. Apart from this

latter, which cannot be constituted a true class, each group in¬

cludes holophytic pigmented forms, and also colourless forms

amongst which the mode of nutrition may range from saprophytic

to completely holozoic. There are even species containing chro-

matophores which can display holozoic nutrition (e.g. Chrysa-

moeba) and these often show amoeboid characters, as do several

of the colourless forms. The distinguishing features of each group

will be dealt with under the class headings, although the key will

also include brief reference to them.

KEY TO THE FLAGELLATA

A. Cells with chromatophores .. B.

A. Cells colourless, without chromatophores.G.

B. Chromatophores grass green.C.

B. Chromatophores yellowish green to bright green, not grass

green .«.•. D.

B. Chromatophores yellowish to brownish.E.

B. Chromatophores variable in colour.F.

C. Cells usually with one chloroplast, rarely more; elaborate

vacuolar apparatus “gullet”, absent; flagella usually two or

four equal in length; storage material starch; complex

colonial forms occur .. VOLVOCALES (Chlorophyceae),

C. Cells usually with several chloroplasts; vacuolar system elab¬

orate, forming a “gullet”; storage material paramylum

EUGLENINEAE (I).

D. Cells bright green; chloroplasts numerous; elaborate vacuolar

system or “gullet” present; trichocysts present in the cell;

storage material fat ... CHLOROMONADINEAE (II).

106

Vol. XIX. (1962).

D. Cells more yellowish green; elaborate vacuolar system or

“gullet” absent; chloroplasts usually few; flagella usually

two, unequal in length; storage material oil, never starch

XANTHOPHYCEAE.

E. Chromatophores usually brownish in colour (but see F),

usually two in a number; cells markedly dorsiventral with

a groove or canal where the flagella are inserted; flagella

usually two, unequal; storage products solid carbohydrates-

starch and allied substances .. CRYPTOPHYCEAE (III).

E. Chromatophores brownish, usually numerous; membrane

usually consisting of sculptured plates; flagella two, one

tr an verse and undulatory, often in a groove, the other pro¬

jecting vertically and free; storage products starch and

oil.DINOPHYCEAE.

E. Chromatophores golden yellow, usually 1-2 in number;

flagella 1-3; cells often with an elaborate envelope; storage

products leucosin; colonial forms frequent

CHRYSOPHYCEAE (II).

F. Chromatophores blue, blue-green or red, usually two in num¬

ber; cells markedly dorsiventral with a groove or canal

where the flagella are inserted, flagella usually two, un¬

equal; storage products solid carbohydrates-starch and

allied substances_CRYPTOPHYCEAE in part (III).

G. Cells colourless, with an elaborate vacuolar apparatus or

“gullet”; storage product paramylum

EUGLENINEAE (in part) (I).

G. Cells colourless, distinctly dorsiventral with a groove or canal

where the flagella are attached, flagella usually two, un¬

equal; storage product starch or allied carbohydrate

CRYPTOPHYCEAE (in part) (III).

G. None of the above characters

RESIDUAL COLOURLESS FLAGELLATA (IV).

EUGLENINEAE

One hundred and twenty-five species of Euglenineae from

Malayan freshwaters have been described in an earlier paper

(Prowse 1958), and the following are species or forms additional

to that list. No key is provided here, but the genera are listed in

the same order as in the previous paper.

The most characteristic features distinguishing the Euglenineae

from the other classes, are the presence of a narrow canal or

cytostome leading back into an enlarged vacuolar swelling or

reservoir, and the fact that the storage product is paramylum, a

polysaccharide allied to starch, but not staining with iodine or

chlorzinc-iodide.

107

Gardens Bulletin, S .

Euglenaceae

Possessing green chloroplasts, ranging from discoid to band¬

shaped with or without pyrenoids. The green colour may in some

cases be obscured by the production of haematochrome. Colour¬

less forms have appeared which are so obviously related to pig¬

mented forms that there is no difficulty in identification.

EUGLENA Ehrenberg 1838

1. Euglena agilis Carter [syn. Euglena pisciformis Klebs]. CELLS

elongate-ovoid, 18-20/* long X 9-11/x wide, bluntly rounded

at the anterior end and tapered to a blunt point at the posterior

end; PERIPLAST very faintly striated, the striations not easily

visible. CHROMATOPHORES two parietal plates nearly filling

the lining of the periplast, but with the division between them

always visible; from the side they appear as elongated bands, but

in freshly killed specimens lightly stained with Gentian violet the

edges are seen to be lobed. PYRENOIDS one in each chro-

matophore, usually centrally placed and with a paramylum cup

on each side; additional paramylum in the form of a few very

short rods. EYESPOT faint, orange in colour. FLAGELLUM

a little over body length. The organism swims rapidly, with a

characteristic sideways flick of the tail end. (PI. I figs, h, i).

Malaya: Malacca—fish-ponds.

Worldwide in distribution.

2. Euglena exilis Gojdics. CELLS broadly fusiform 70-85/* long

X 18-22/* wide, rounded at the anterior end, tapering gently

to a point at the posterior end. PERIPLAST very prominently

striated, the striae seen clearly to consist of rows of dots. The

CYTOPLASM contains scattered spherical granules which stain

with neutral red, but are otherwise not easily visible. CHRO¬

MATOPHORES usually four elongated parietal plates, lobed

at the edges, the lobes best seen when the cell has swollen out

and is lightly stained with gentian violet. PYRENOIDS one in

the centre of each chromatophore, with a cup of paramylum

on each side; additional paramylum in the form of small rect¬

angular rods 2-3/a long. FLAGELLUM i-1 times body length.

EYESPOT large, orange red. Cell distinctly metabolic, bulging

in the middle and then narrowing down again. (PI. 1 fig. k).

Malaya: Malacca—fish-ponds.

Originally described from U.S.A.

Despite the larger size and fatter shape of the Malayan forms,

I have felt it better to place them under Gojdics species (Gojdics

measurements are 37-53/* X 8-10/*). They clearly agree in the

striated pellicle, shape and number of the chromatophores, and in

108

Vol. XIX. (1962).

the granules which stain neutral red. The flagellum in the Malayan

form is admittedly longer than in the original species from U.S.A.

There are distinct affinities with Euglena polymorpha Dangeard,

which is of a similar size and shape, but that species has 12-20

smaller chromatophores.

3. Euglena flava Dangeard. CELLS broadly fusiform, 55-70/x

long X 20-25/x wide, rounded at the anterior end and tapering

to a blunt tail at the posterior end. PERIPLAST faintly striated,

the striations difficult to see. CHROMATOPHORES 10-15

elongated discs with smooth edges, parietally placed parallel to

the long axis and appearing as elongated bands in side view.

PYRENOIDS one in each chromatophore with a cup of

paramylum on each side, additional paramylum not seen. The

cells are coloured red by numerous granules of haematochrome

which are either clumped in the middle, or dispersed throughout

the cell. FLAGELLUM about body length. EYESPOT a quite

large, orange-red concave disc. The cell is distinctly metabolic

stretching out and contracting to a round ball. (PI. I fig. g).

Malaya: Malacca—small experimental ponds at Batu Beren-

dam.

This species formed orange-red scums in small ponds manured

with buffalo manure. Why it should have occurred rather than

Euglena sanguinea Ehrenberg, to be found in almost any padi

field or carp pond, is hard to say. Perhaps the origin of the buffalo

manure might have explained it, but there are no records of this.

A bloom of Euglena flava Dangeard is quite different from the

brick-red blooms of Euglena sanguinea Ehrenberg, being much

more orange in colour. Chu (1946) has suggested that Euglena

flava Dangeard is synonymous with Euglena caudata Huebner,

which he found produced haematochrome in culture, but neither in

Dangeard’s original description, nor in the Malayan material of

Euglena flava was there any sign of lobing in the chloroplast as in

Euglena caudata.

4. Euglena gracilis Klebs. CELLS nearly cylindrical, 45-55^ long

X 8-12 /a wide, rounded at the anterior end and bluntly pointed

at the posterior end, although the tail may round off under

pressure from the cover slip. PERIPLAST with such faint stria¬

tions as to be hardly visible, even under oil immersion. CHRO¬

MATOPHORES 7-10 parietal yellowish-green circular to ovoid

discs, but the outlines may become irregular when they are

closely packed together. PYRENOIDS one in each chromato¬

phore, with a paramylum cup on each side; additional para¬

mylum ovoid bodies which may become so abundant as to bulge

out the periplast. FLAGELLUM half to body length, very

109

Gardens Bulletin, S.

active. EYESPOT crimson, prominent. The cell is very active,

swimming rapidly by a zig-zag motion which is characteristic,

and it can be very metabolic. (PI. I figs, e, f).

Malaya: Malacca—fish-ponds.

Selangor—Sungai Gombak, Kuala Lumpur.

Negri Sembilan—standing pools, Port Dickson.

Worldwide in distribution.

This species is easily recognised by its clear yellow-green colour

and its characteristic zig-zag swimming motion. It does not appear

to be abundant in any sample of water collected, although it is

probably wide-spread in distribution. Nevertheless it is probably

the easiest species to culture, being tolerant of the acid conditions

that often develop in cultures. Cultures of material from different

sources often show slight differences in the physiology of the organ¬

ism and in two cultures at Batu Berendam the cells have shown a

tendency to secrete haematochrome, although never in the quantity

to be found in the typical red species of Euglena.

5. Euglena intermedia (Klebs) Schmitz var. klebsii Lemmermann.

CELLS cylindrical 100-130/i. long X 15-18/x wide, only slight¬

narrowing to the blunt anterior end, and narrowing at the poste¬

rior end to a short blunt tail piece. PERIPLAST apparently

unstriated, even under oil-immersion. The CYTOPLASM is

granular and contains spherical bodies which stain with neutral

red, scattered throughout the cell except at the ends, where they

tend to be clumped. CHROMATOPHORES numerous, lenti¬

cular, 8-10/x long, without pyrenoids. PARAMYLUM in short

rods 2 in length, each with a slight central depression. EYE-

SPOT purplish red, conspicuous. FLAGELLUM rarely seen,

less than one-sixth body length, the cell usually moving by

means of a squirming motion. (PI. I fig. a).

Malaya: Selangor—Kuala Lumpur, a pond.

Negri Sembilan—Seremban and Port Dickson, stag¬

nant water.

Malacca—stagnant water.

Johore—Muar and Yong Peng, stagnant water.

Singapore—stagnant pools.

A very characteristic species, particular with its squirming habit.

The Malayan forms belong to the var. klebsii Lemmermann be¬

cause of the small paramylum rods. The species and its varieties

could be confused with Euglena deses Ehrenberg, and the type

was originally referred to that species as var. intermedia Klebs.

However Euglena intermedia and its forms, including the Malayan

material here described, have no pyrenoids as in Euglena deses .

110

Vol XIX. (1962).

6. Euglena mutabilis Schmitz [Syn. Euglena acus Ehrenberg var.

mutabilis Klebs.] CELLS cylindrical, 80-110/4 long X 5-9/4

wide, tapering very slightly to the blunt, often slightly oblique

anterior end and tapering to a point at the posterior end. The

PERIPLAST shows faint, steeply spiralling striae which are

visible with difficulty. The CYTOPLASM contains numerous

small spherical granules visible without staining and scattered

throughout the cell. CHROMATOPHORES yellowish-green,

usually four parietal plates, two on each side of the nucleus;

they are so closely appressed to each other and stretch nearly

round the cell that the division between them is difficult to see,

but will show up in certain positions of the cell, particularly

after staining. PYRENOIDS one in each chromatophore, usually

central and showing up as a bulge in edge view. PARAMYLUM

short rectangular bodies about 2/4 long and often so abundant

as to fill the cell. FLAGELLUM not seen in any of the Malayan

material examined, the cell being only slightly metabolic and

moving by a sideways lashing movement or looping from end to

end; eyespot crimson, prominent. (PI. I figs, b, c, d).

Malaya: Negeri Sembilan—ditches at Seremban and Port

Dickson.

Malacca—ditches at Batu Berendam and elsewhere.

Singapore—ditches near the Royal Island Club golf

course and Peirce reservoir.

Distribution worldwide.

LEPOCINCLIS Perty 1852

7. Lepocinclis fusiformis (Carter) Lemmermann (emend Conrad)

[syn. Euglena fusiformis Carter, Lepocinclis fusiformis Lem¬

mermann, Lepocinclis sphagnophila Lemmermann partem].

CELLS broadly fusiform, 50-60/4 long X 25-30/t wide, slightly

truncate at the anterior end, rounded acute at the posterior end.

PERIPLAST with distinct striations spiralling backwards from

left to right. CHROMATOPHORES numerous, discoid, parie¬

tal, each with a simple pyrenoid. PARAMYLUM in the form of

two large rings lying just inside the pellicle. FLAGELLUM

about H times body length. EYESPOT crimson, anterior. (PI.

I fig. 1).

Malaya: Singapore in a stagnant pool.

Worldwide in distribution.

PHACUS Dujardin 1841

8. Phacus meson Pochmann [syn. ? Phacus pleuronectes (O. F.

Mueller) Dujardin after Dangeard, Phacus longicauda (Ehren¬

berg) Dujardin var. brevicaudata Skvortzow, Phacus longicauda

var. indica Skvortzow]. CELL oval, 75-80/* long without tail

111

Gardens Bulletin , S.

X 50/* wide, rounded at the anterior end and tapering poste¬

riorly to a short, stout, straight, pointed tailpiece about 30/x

long. PERIPLAST distinctly striated longitudinally CHRO-

MATOPHORES numerous, discoid. PARAMYLUM two large

rings. FLAGELLUM about body length. EYESPOT crimson,

small. (PI. I. fig. n).

Malaya: Fish-ponds in Malacca.

Reported from various parts of S.E. Asia and possibly from

Europe.

This form resembles Phacus longicauda (Ehrenberg) Dujardin in

some respects, but differs in the much shorter, stouter, straight

tailpiece and the more oval shape of the body of the cell.

TRACHELOMONAS Ehrenberg 1833 emend Deflandre 1926

9. Trachelomonas allia Drezepolski forma. This form differs from

the type in being distinctly broader at the posterior end, tapering

gently to the rounded anterior end. As there are intermediate

forms there is no reason to separate the present one as a distinct

variety. (PI. I fig. q).

Malaya: Fish-ponds in Malacca.

10. Trachelomonas intermedia Dangeard. CELL subspherical to

elliptic, 18-20/* long X 15-17/* wide. LORICA golden brown,

finely but distinctly punctate. PORUS with a ring of thickening.

CHROMATOPHORES several, discoid. EYESPOT crimson,

clearly visible. FLAGELLUM H—2 times body length. (PI. 1

fig- j).

Malaya: A ditch in Malacca.

Widespread throughout the world.

11. Trachelomonas volvocinopsis Swirenko [syn. Trachelomonas

indica Skvortzow]. CELL spherical, 15/* in diameter. LORICA

smooth, brown without thickening to the porus. CHROMATO¬

PHORES 8-10, discoid, without pyrenoids. EYESPOT red.

FLAGELLUM 2-3 times body length. (PI. I fig. o).

Malaya: Selangor—standing water at Kuala Lumpur.

Negri Sembilan—standing water at Port Dickson and

Seremban.

Malacca—fish-ponds and ditches.

Johore—standing water at Muar.

Singapore—standing water near the reservoirs and the

Royal Island Club golf course.

This species might be confused with Trachelomonas volvocina

Ehrenberg, from which it differs by having more than two chro-

matophores and by completely lacking pyrenoids.

112

Vol. XIX. (1962).

STROMBOMONAS Deflandre 1930

12. Strombomonas verrucosa (Daday) Deflandre var. chinensts

(Skvortzow) Deflandre [syn. Trachelomonas chinensis Skvort-

zow and vars. ovata Skvortzow and assuriensis Skvortzow,

Trachelomonas conspersa Pascher emend. Gordon]. CELL

ovoid, 28-30/a long X 22-25/* wide, truncate at the anterior

end with a very short wide neck, subconical at the posterior end.

LORICA brown, thick and irregularly granular or verrucose on

the surface. CHROMATOPHORES numerous discoid. EYE-

SPOT crimson, clearly visible. FLAGELLUM about li times

body length. (PI. I fig. p).

Malaya: Fish-ponds in Malacca.

Described from China and Russia.

Peranemaceae

Cells metabolic or rigid, often crawling, but also free-swimming,

usually bilaterally asymmetrical, and dorsiventrally organised.

Flagella 1-2 or sometimes absent. Vacuolar system as in Euglena,

but usually with two rod-shaped bodies, or a tubular siphon the

former closely associated with the reservoir. Storage material para-

mylum granules and sometimes fat. The cells are complete without

a stigma or chloroplasts, although occasionally partly digested

chloroplast material may be seen inside the cell. Nutrition usually

holozoic.

PERANEMA Dujardin 1841

13. Peranema inflexum Skuja. CELL cylindrical fusiform, 30-40/a

long X 9-12/a wide, narrowed at the anterior end, bluntly

rounded at the posterior end, the whole cell usually slightly

bent or curved laterally. PERIPLAST with distinct striations

spiralling backwards from left to right. RESERVOIR and canal

about i body length with an associated rod-shaped organ

(“staborgan”). PARAMYLUM as small oval bodies scattered

throughout the cell. FLAGELLUM about body length. (PI. I

fig. m).

Malaya: A swamp in Kuala Lumpur.

Described from Northern Europe.

Gardens Bulletin, S.

Rhizaspidaceae Skuja

A single genus Rhizaspis Skuja

RHIZASPIS Skuja 1948

Cells without flagella, elliptic to ovate, slightly narrowed at the

anterior end, rounded to sub-cuneate at the posterior end, marked¬

ly flattened dorsiventrally, dorsal side convex, the ventral side

concave to slightly convex and often with a slight furrow in it.

Periplast smooth, colourless, firm except at the anterior end where

fine pseudopodia arise. Siphon or accessory rod-shaped organs

(“stab-organs”) absent, but there are several vacuoles at the

anterior end. The cytoplasm is colourless, with numerous spherical

granules of paramylum and sometimes 1-2 large refractive bodies

which are probably composed of leucosin.

This is a curious genus, very animal like in nature and nutrition.

On the other hand it shows distinct affinities with the Euglenineae

in the storage of paramylum, the colourless cytoplasm and the

firm periplast.

14. Rhizaspis granulata Skuja. CELLS broadly elliptic to ovate,

60-65/t long X 40—45/t wide X 18—20/* thick; at the narrowed

anterior end it is slightly protruded to form a rounded lip-like

structure, from which fine pseudopodia arise, at the posterior

end it is broadly cuneate; in side view the cell is curved with a

truncate anterior end and cuneate posterior end, the ventral side

concave with a broad shallow longitudinal furrow, the dorsal

side convex. PERIPLAST colourless, smooth and quite firm.

CYTOPLASM colourless, with several vacuoles at the anterior

end, numerous PARAMYLUM granules at the posterior end,

several food granules, and sometimes 1 large refractive granule

of leucosin, although this is not always present. NUCLEUS

large, subcentral. The cell moves by crawling over the sub¬

stratum by means of the pseudopodia, with the cell body erect,

or more usually held at an oblique angle. (PI. II figs, c, d).

Malaya: Experimental fish-ponds in Malacca.

Described from Northern Europe.

There is no doubt about the identity of the Malayan specimens.

They agree with Skuja’s description except in the number of

leucosin granules, for here they are often absent.

15. Rhizaspis simplex Skuja. CELLS oval, 50-55/c long X 30-

35/u. wide X 20-22/x thick, rounded at the posterior end, nar¬

rowed to the truncate anterior end, from which fine pseudopodia

arise, but without a lip-like protrusion; in side view the cell

convex on the dorsal side, less so on the ventral side; sometimes

114

Vol. XIX. (1962).

there are two low broadly rounded ridges on the dorsal side,

and a shallow furrow on the ventral side, but these are not

always very visible since the cell is a little metabolic and can

swell out its outlines. PERIPLAST colourless, smooth, firm but

elastic. CYTOPLASM colourless with several vacuoles at the

anterior end and numerous spherical granules of PARAMYLUM

at the posterior end, and often several food granules. Leucosin

appears to be absent. NUCLEUS large, sub-central. The cell

crawls along by means of the pseudopodia, keeping the body

erect, but occasionally this latter will shorten and fatten slightly.

(PI. II figs, a, b).

Malaya: Experimental fish-ponds in Malacca.

Described from Northern Europe.

The truncate anterior end places the Malayan forms under this

species, but they are a little fatter than the forms described by

Skuja, and the ridges and furrows are not always so easy to see.

CHLOROMONADINEAE

A class of flagellates with only a few forms known, and of

which the affinities are very uncertain. No coccoid or filamentous

forms have been recorded, all known forms being motile unicells,

usually oval or pyriform in shape. The cell is often dorsiventrally

flattened, with a ventral furrow. There are two flagella, one usually

trailing, and they are attached near the aperture of a complex

vacuolar system recalling that in the Euglenineae. In the pigmented

forms there are numerous discoid chloroplasts, bright green in

colour, but containing an excess of xanthophyll—they give a blue

colour with concentrated hydrochloric acid, and in this respect

the group resembles the Xanthophyceae. Neither starch nor para-

mylum occur, food reserves being fats and oils. In many forms

radially disposed trichocysts are found in the peripheral regions,

often more crowded at the anterior end, and these may be dis¬

charged on stimulus. Reproduction, as far as is known, is by

longitudinal fission. This class forms a somewhat isolated group,

but resembles the Euglenineae in the vacuolar system, but differs

in the nature of the pigments and food reserves, in which respect

it shows closer resemblance to the Xanthophyceae.

Two genera recorded from Malaya:—

1. Cells green, with trichocysts. Flagella apically attached

Gonyostomum.

2. Cells green, with trichocysts. Flagella laterally attached

Merotrichia.

115

Gardens Bulletin, S.

GONYOSTOMUM Diesing 1866

Cells motile, dorsiventrally flattened, obovate, to spherical in

front view, slightly metabolic, often narrowed posteriorly. Dorsal

surface convex, ventral surface flattened, with a shallow furrow

leading from the opening of the vacuolar system (“pharynx”) and

running longitudinally. Vacuolar system a triangular “pharynx”—

like structure. Chromatophores numerous, discoid, green. Eyespot

absent. Nucleus central. Trichocysts radial, abundant, more closely

packed at the anterior end. Flagella two, one trailing but both

attached at the mouth of the “pharynx”. Oil and fat droplets

occur as the food reserve.

16. Gonyostomum depressum Lemmermann [syn. Vacuolaria

depressa Lauterborn]. CELL round in face view, 30-40/a in

diameter, broad elliptic in side view, about 25-30/x thick.

CHROMATOPHORES numerous, green, discoid, lining the

periphery of the cell. VACUOLAR SYSTEM a small triangular

“pharynx”. TRICHOCYSTS numerous, peripheral, radial, but

crowded at the anterior end. FLAGELLA two, distinctly un¬

equal in length. Storage material oil and fat droplets. (PI. II fig.

o).

Malaya: Malacca—in acid swamps.

Described from Europe.

This species resembles Gonyostomum latum Iwanoff, but differs

in the unequal flagella.

17. Gonyostoinun semen Diesing [syn. Rhaphidomonas semen Eh-

renberg]. CELL elongate-obovate, 40-70/a long X 22-40/a wide,

rounded at the anterior end and narrowed to a subcuneate pos¬

terior end, very much flattened dorsiventrally, about 10-15/a

thick, and with a broad ventral furrow. CHROMATOPHORES

numerous, green, discoid, lining the periphery of the cell. VA¬

CUOLAR SYSTEM a prominent apical, triangular “pharynx”.

TRICHOCYSTS numerous, peripheral, radial, but very

crowded at the anterior end. FLAGELLA two, nearly equal,

but one trailing, both about body length. Storage material fat

and oil. (PI. II fig. g).

Malaya: Malacca—in acid swamps.

Worldwide in distribution.

MEROTRICHIA Mereschkowski 1877

Cells ellipsoidal to bean shaped, sometimes slightly metabolic,

rounded at the anterior end, occasionally slightly narrowed towards

the posterior end. Vacuolar system a rounded “pharynx” like

structure, laterally placed towards the anterior end. Chromato¬

phores numerous, green discoid. Eyespot absent. Nucleus central.

116

Vol. XIX. (1962).

Trichocysts scattered except at the anterior end, where they are

crowded. Flagella two (one only described for Merotrichia bacil-

lata Mereschkowski, but a second one is probably present), un¬

equal, attached laterally at the mouth of the pharynx. Food reserves

consists of oil and fat droplets. One species reported in Malaya.

18. Metrotrichia capitata Skuja. CELLS oval to bean-shaped, 30-

40 ix long X 22-25 ^ wide, rounded at the anterior end and nar¬

rowed towards the posterior end, but sometimes slightly meta¬

bolic; just below the anterior end is a lateral cleft indicating a

pharynx and the cell is often curved towards that side. CHRO-

MATOPHORES numerous, green, discoid, lining the periphery

of the cell, VACUOLAR SYSTEM a laterally placed “pharynx”

with a groove leading into it. TRICHOCYSTS crowded at the

anterior end, few and scattered elsewhere. FLAGELLA two, the

swimming one about i body length and pointing forward, the

other longer 1-1 i body length, trailing. Storage material fat and

oil. (PI. II. fig. h).

Malaya: Malacca—in acid swamps.

Described from Northern Europe.

CRYPTOPHYCEAE

Cells usually motile by means of two apical or lateral flagella,

although a few coccoid forms have been placed in this group.

The cells are dorsiventrally flattened, oval, slipper-shaped or reni-

form, and possess a longitudinal furrow. In many forms a gullet

extends inwards from the anterior end of the furrow. Chromato-

phores usually 2, occasionally many, sometimes one or absent,

the pigments varying from golden-brown, red, to distinctly blue.

Pyrenoids occur, apparently not embedded in the chromato-

phores, and occasionally they may be absent. Storage products

starch or allied carbohydrates, and sometimes oil as well. In some

forms trichocysts may occur closely surrounding the gullet. Re¬

production is usually by longitudinal division, although palmelloid

stages may be formed first.

Three genera have been found in Malaya:—

1. Cells pigmented, slipper shaped, with two flagella which

are attached slightly to one side of the apex

Cryptomonas.

2. Cells as above but without chromatophores Chilomonas.

3. Cells pigmented, reniform, with the two flagella attached

at the middle of the concave margin. Sennia.

117

Gardens Bulletin , S.

CRYPTOMONAS Ehrenberg 1838

Cells slipper-shaped, dorsiventrally flattened, convex on the

dorsal surface, flat or concave ventrally, with a broad longitudinal

furrow from which a gullet extends inwardly from the anterior

end. Flagella two, attached at the opening of the gullet. Chromato-

phores 1 or 2, lateral and parietal. Storage products starch grains.

Trichocysts often visible round the gullet. There are 1-3 con¬

tractile vacuoles. Stigma apparently absent.

19. Cryptomonas erosa. Ehrenberg var. reflexa Marsson. CELLS

slipper-shaped, 20-30/x long X 10—15/a wide, obliquely trun¬

cate at the anterior end, narrowed posteriorly to a blunt point

which is turned to one side, so that the whole cell is slightly sig¬

moid. CHROMATOPHORES two, lateral, golden brown to

yellowish green in colour. PYRENOIDS absent. STORAGE

PRODUCTS oval to polygonal starch grains lining the inner

sides of the chromatophores. FLAGELLA two, slightly un¬

equal, the longer being a little more than body length, the other

a little less, both inserted at the mouth of the gullet in the middle

of the oblique anterior end. (PI. II. figs, k, 1, n).

Malaya: Malacca — in experimental fish-ponds.

Worldwide in distribution.

This is very similar to Cryptomonas marssonii Skuja, which has

been separated off for reasons which are not very obvious, in parti¬

cular the more sharply truncate anterior end and the more dep¬

ressed mouth of the gullet.

20. Cryptomonas ovata Ehrenberg. CELLS almost ellipsoidal, 45-

50 ix long X 18-25/x wide, slightly obliquely truncate at the

anterior end, rounded at the posterior end, the whole cell curved

slightly to one side. CHROMATOPHORES two, lateral, large,

nearly filling the periphery of the cell but with the division

between them quite clear, golden brown to yellowish green in

colour, often appearing netted where the closely packed starch

grains press against them. PYRENOIDS absent. STORAGE

PRODUCTS starch grains, usually oval but appearing polygonal

when tightly packed, lining the inner sides of the chromato¬

phores. GULLET prominent, stretching 1/3-i length of the

cell, and surrounded by trichocysts. FLAGELLA two, nearly

equal, a little less than body length. (PI. II fig. m).

Malaya: Malacca—in small experimental fish-pond.

Worldwide in distribution.

21. Cryptomonas phaseolus Skuja. CELLS small, elliptic, 12-15/x

long X 8-9/a wide, rounded at both ends and only very slightly

oblique at the anterior end. CHROMATOPHORES two, lateral,

brownish in colour. PYRENOIDS absent. STORAGE PRO¬

DUCTS several comparatively large oval starch granules lining

118

Vol XIX. (1962).

the inner side of the chromatophores; occasionally 1-2 central,

oval refractive bodies. GULLET about 1/3—i body length,

with the opening at one side of the apex. FLAGELLA two

slightly unequal in length, a little shorter than body length. (PI.

II figs, i, j).

Malaya: Malacca—in experimental fish-pond.

Described from Northern Europe.

Skuja describes his specimens as violet brown and thiophile.

The Malacca specimens are more golden brown to reddish brown,

but they may well be thiophile, since the pond in which they were

found had a slight smell of hydrogen sulphide.

CfflLOMONAS Ehrenberg 1838

Cells resembling Cryptomonas but completely devoid of chro¬

matophores and pyrenoids. Cells more or less slipper-shaped, dor-

siventrally flattened, the opening of the gullet somewhat sunken

so that the surrounding margin has a lip-like appearance. One

lateral margin is distinctly more convex than the other, which may

be flattened. Trichocysts often surrounded the gullet, and storage

products are comparatively large starch granules. Flagella two,

unequal, attached at the mouth of the gullet.

Usually found in polluted water, often in abundance.

22. Chilomonas paramecium Ehrenberg. CELLS slipper-shaped,

25-30/a long X 10-15 /a wide, rounded at the posterior end,

lip-like at the anterior end, with a lateral kink indicating the

sunken opening of the gullet; lateral margin adjacent to the

gullet only slightly convex or even flattened, the other margin

distinctly convex. CHROMATOPHORES absent. PYRENOIDS

absent. GULLET prominent, about half the length of the cell,

surrounded by trichocysts. STORAGE PRODUCTS large round

starch granules. FLAGELLA two, unequal, the longer being

about body length. (PI. II fig. f).

Malaya: Selangor—polluted water from the Gombak river,

Kuala Lumpur.

Worldwide in distribution.

SENNIA Pascher 1913 emend. Skuja 1948

Cells bean-shaped, dorsiventrally flattened and one lateral margin

decidedly convex, the other slightly concave with an equatorial

groove running from it. Gullet in the middle of the concave side,

not very prominent. Trichocysts apparently absent. 1-2 contractile

vacuoles occur near the gullet. Chromatophores 1-2, golden yellow

to olive-green in colour. Pyrenoids present or absent. Eyespot

present or absent. Flagella two, unequal in length, laterally at¬

tached at the opening of the gullet.

119

Gardens Bulletin, S.

23. Sen ilia parvula Skuja. CELLS small, bean-shaped, 10/u long

X 5/j. wide, very convex on one lateral margin, flattened or

slightly concave on the other. CHROMATOPHORE one, curv¬

ing round the convex margin, olive-green in colour. STIGMA

absent in the Malayan specimens. PYRENOIDS absent. GUL¬

LET small, in the middle of the concave margin, without tri-

chocysts but with a single small contractile vacuole. FLAGELLA

two, divergent, unequal in length, the longer about body length.

(PI. II fig. e).

Malaya: Malacca—in an experimental fish-pond.

Reported from Northern Europe, America and Africa.

The Malayan specimens differ from the type only in the absence

of the stigma, a varying feature in this genus anyway.

CHRYSOPHYCEAE

The Chrysophyceae are characterised by possessing chromato-

phores which are golden-yellow to brown in colour, although in

waters rich in organic matter they may assume a greenish tint. The

main pigment giving the golden-yellow colour is phycochrysin,

but modified chlorophylls and xanthophylls may also occur.

Storage products consist of oil and whitish, highly refractive,

usually round lumps, leucosin, often forming a prominent mass

in the posterior portion of the cell. Leucosin appears to be a

carbohydrate, although some doubt has been thrown on this, and

it is readily soluble in most reagents and dots not react with

iodine. Starch is never formed.

All Chrysophyceae appear to be uninucleate and may range

from simple motile flagellate cells, and motile colonies to branched

palmelloid forms or even a few filamentous species. A characteris¬

tic feature of the class is the formation endogenously of cysts, the

walls of which are impregnated with silica.

The Chrysophyceae are frequently grouped together with the

Xanthophyceae and Bacillariophyceae in the phylum Chrysophyta.

In addition to the absence of chlorophyll b in all three classes, and

the absence of starch, there are certain other structural similarities.

In both the Xanthophyceae and Chrysophyceae endogenously

formed cysts have been reported. These are frequently impregnated

with silica, an obvious comparison with the Bacillariophyceae.

The cell envelope of some Chrysophyceae ( Hyalobryon) is lami¬

nated, recalling similar conditions in the Xanthophyceae ( Ophio-

cytium) and even the intercalary bands in the Diatoms. Leucosin

has been reported in certain Diatoms.

120

Vol XIX . (1962).

KEY TO THE ORDERS OF THE CHRYSOPHYCEAE

J. Cells normally motile and flagellate, sometimes amoeboid.

Colonial forms motile.CHRYSOMONADALES.

2. Cells normally amoeboid, holozoic, flagellate stages rare or

absent . [RHIZOCHRYSIDALES].

3. Cells in palmelloid colonies, often elaborately branched

[CHRYSOCAPSIDALES].

4. Cells coccoid, flagellate stages rare or absent

[CHRYSOPHAERALES].

5. Plants filamentous . [CHRYSOTRICHALES],

KEY TO THE CHRYSOMONADALES

1. Cells with 1 flagellum . CHROMULINEAE.

2. Cells with 2 equal flagella.ISOCHRYSIDEAE.

3. Cells with 2 unequal flagella.OCHROMONADEAE.

4. Cells with 3 flagella, one short and two long

[PRYMNESEAE].

CHROMULINEAE

(i) Cells naked....(ii).

(i) Cells not naked, possessing a wall or envelope.(iii).

(ii) Cells only rarely amoeboid, usually flagellate . . Chromulina.

(ii) Cells markedly amoeboid, with prominent pseudopodia

Chrysamoeba.

(iii) Cells with small lens-shaped siliceous scales embedded in the

wall, without regular arrangement and devoid of spines

(Microglena).

(iii) Cells with much larger siliceous scales, regularly arranged and

frequently bearing long spines . Mallomoncis .

(iii) Cells with siliceous material in complete transverse bands

(Conradiella).

(iii) Cells enclosed in a stalked envelope. Chrysopyxis.

(iii) Cells in spherical colonies, set with long needles

(Chrysosphaerella).

121

Gardens Bulletin , 5.

CHROMULINA Cienkowsky 1870

Cells naked, ovoid pyriform, elliptic or spherical unicells, often

very metabolic and each bearing a single apical flagellum. Periplast

of the cell often granular or set with small wart-like excrescences.

Chromatophores 1-2, clearly differentiated, curved plate or rings,

sometimes nearly filling the periphery of the cell, in most cases

without pyrenoids. In many species there is a stigma. At the

anterior end there are 1-2, sometimes more, contractile vacuoles.

Storage products leucosin, often as a large single sphere at the

posterior end of the cell, and oil droplets. The cysts are formed

endogenously, with the wall impregnated with silica and are

spherical with a symmetrically placed mouth, which may or may

not bear a prominent neck.

24. Chromulina sphaerica Bachman. CELLS spherical, 12-15/1 in

diameter. PERIPLAST smooth, not granular nor with excre¬

scences. CHROMATOPHORE a single larged curved plate

lying against the periplast but varying in position, without a

pyrenoid. STIGMA present, small crimson but conspicuous.

FLAGELLUM 1-1* body length. (PI. V fig. g).

Malaya: Malacca—in fish-ponds.

Described from Europe.

Bachman describes his species with the chromatophore in the

middle, whereas here it lines the periphery. It is therefore possible

that the Malayan forms should be regarded as a separate species,

although it is difficult to decide how much importance should be

given to the position of the chromatophore. In all other respects

the Malayan forms agree with Bachman’s description.

CHRYSAMOEBA Klebs. 1893

Cells motile with a single apical flagellum, often bearing fine

pseudopodia whilst still flagellate, occasionally losing the flagellum.

Chromatophores 1-2 curved plates or bands, central or parietal.

Stigma present or absent. Pyrenoids present or absent. Cysts, where

known, spherical with a centrally placed orifice. Leucosin granule

often present as storage material.

25. Chrysamoeba radians Klebs. FLAGELLATE STAGE ovoid,

15—25 /i long X 8-12/4 wide, often narrowed at the posterior

end. CHROMATOPHORE usually 1, bandshaped curving

round the cell even in amoeboid stages, occasionally two, with¬

out PYRENOIDS. STIGMA absent. FLAGELLUM 1-1* body

length rarely shorter. CYST spherical, 20-22 fx in diameter, with

a short tapering neck. (PI. V fig. i, j, k, 1).

Malaya: Malacca—in fish-ponds.

Worldwide in distribution.

122

Vol. XIX. (1962).

The Malayan forms are larger than most records for this species,

and the chromatophore is appreciably larger and curves just inside

the periplast rather than being central. For this reason they might

be regarded as a separate species. On the other hand they were

collected from very fertile ponds in a region of high light intensity.

It is possible that in less fertile waters and with lower light in¬

tensities the chromatophores may be smaller and contract to the

middie of the cell.

MALLOMONAS Perty 1852

Cells solitary, motile by means of a single flagellum, ovoid,

ellipsoid, cylindrical, spherical or pyriform in shape; periplast with

numerous circular, discoid or angular silicified imbricating scales;

scales usually regularly arranged, often in spirals, rarely truely

irregular, often bearing long siliceous spines or setae which may

be toothed, occasionally shorter ones or completely lacking;

chromatophores golden-yellow, 2, lateral and parietal; contractile

vacuoles present at the anterior end of the cell; eyespot often

lacking; nucleus ellipsoid, often large and distinct; storage material

leucosin, as large spherical granules at the posterior end.

26. Mallomonas acaroides Perty. CELLS ovoid to ellipsoid, 30-

50/x long X 15-22 p wide, rounded posteriorly and often

slightly narrowed at the anterior end. SCALES oval to elliptic,

arranged in spiral rows. SETAE long, somewhat curved, one

to each scale so that the whole cell is covered by spines.

CHROMATOPHORES 2, large, lateral and parietal, golden-

yellow in colour. FLAGELLUM about body length. (PI. Ill

figs, e, f, 1).

Malaya: Malacca—in experimental fish-ponds.

Distribution worldwide.

This is a characteristic if somewhat variable species. The

Malayan forms are larger than most of those recorded from else¬

where, but, not sufficiently so to warrant any separation as a

variety.

27. Mallomonas curta (Playfair) Conrad, [syn. Mallomonas lito -

mesa Stokes var. curta. Playfair]. CELLS elliptic 18-30/a long

X 11-15/a wide, with a very short broad papillum at the ante¬

rior end, rounded at the posterior end. SCALES circular,

arranged in shallow spirals. SETAE six only, about i body

length, attached at the papillum and curving backwards. FLA¬

GELLUM about body length. STIGMA small, crimson but

often lacking. (PI. Ill figs, g, i).

Malaya: Malacca—in swamp.

Described from Australia and reported from Europe.

123

Gardens Bulletin, S.

28. Mallomonas playfairii Conrad var. opisthiodonta Prowse. var.

nov. CELLS ellipsoid, 19-20/'. X 14— 15/x wide, broadly round¬

ed at both ends, but terminating anteriorly in a very short neck.

SCALES rhombic, spirally arranged at an angle 45°, without

spines except in the four rearmost scales, which each bear a

single short spine. CHROMATOPHORES two, large, parietal.

FLAGELLUM nearly body length. (PI. Ill fig. j).

Malaya: Malacca—fish-ponds. (Type locality).

Holotype: Prowse 398 deposited at Tropical Fish Culture

Research Institute, Malacca.

Cellulae ellipsoideae, 19-20/x longae, 14-15/4 latae, utrinque

late-rotundatae, unaquaeque antice in collum brevissimum ter-

minata. Squamae rhombicae, ad angulum 45° spiraliter dispo-

satae, spiris longissimis carentes, sed quattuor squamae postre-

mae tantum brevissime unispinosae. Flagellum corpori paene

aequilongum aequilongum. Chromatophora duo, magna, parie-

talia.

Malaya: Malacca—in stagnis piscosis. (Holotypus).

29. Mallomonas *pherica Prowse sp. nov. CELLS spherical, 8-9/i

in diameter. SCALES elliptic, arranged spirally at a slight

angle, only the anterior-most scales being spiny; SETAE 9-10

in all, 2—2i times longer than the body, strongly reflexed.

CHROMATOPHORES 2, discoid, parietal. FLAGELLUM

nearly as long as the body. (PI. Ill fig. m).

Malaya: Malacca—in fish-ponds. (Type locality).

Holotype: Prowse 395 deposited at Tropical Fish Culture

Research Institute, Malacca.

Cellulae sphericae, 8-9/4 in diam. Squamae ellipticae, ad angu¬

lum paululum spiraliter dispositae, eae antice sitae tantum spinasae;

spinis 9-10, quam corpus 2-2i-plo longioribus, valide reflexis.

Flagellum corpori paene aequilongum. Chromatophora duo, dis-

coidea, parietalia.

Malaya: Malacca—in stagnis piscosis. (Holotypus).

30. Mallomonas splendens (G. S. West) Playfair [syn. Lagerheimia

splendens G. S. West]. CELLS cylindrical, 30-50/c long X 15—

17/x wide, slightly narrowed at each end to the rounded poles.

SCALES nearly rhombic, finely punctate and spirally arranged.

SETAE confined to the poles, 4 at each end, about i body

length, divergent. CHROMATOPHORE one, large, lining one

side. FLAGELLUM about i body length. (PI. Ill fig. c).

Malaya: Selangor—acid swamp in Kuala Lumpur.

Negri Sembilan—acid swamps at Port Dickson and

Seremban.

Malacca—acid swamps in various areas.

124

Vol. XIX. (1962).

Singapore—swampy pools near the Royal Island Club

golf course and Peirce reservoir.

Worldwide in distribution, including Indonesia.

A very characteristic species.

31. Mallomonas tcilingioides Prowse sp. nov. CELLS elongate-

elliptic, 50-55/a long, 18-20 /a wide, equally rounded at both

ends, or slightly narrower at the anterior end. SCALES dis¬

tinctly quadrate, spirally arranged, only the ones near the poles

bearing setae, the others apparently smooth 5-9 in the largest

spirals (median ones). SETAE slender, simple, 15-20 at each

end of the cell, 30-32/a long. CHROMATOPHORES two,

large, parietal, golden yellow STIGMA prominent, crimson, at

the anterior end. FLAGELLUM i body length. LEUCOSIN

a large single grain at the posterior end. (PI. Ill figs, d, k).

Superficially resembling Mallomonas teilingii (Teiling) Conrad,

from which it differs in scales being quadrate (not discoid) and in

having slightly more numerous setae.

Malaya: Malacca—in fish-ponds. (Type locality).

Holotype: Prowse 414a deposited at Tropical Fish Culture Re¬

search Institute, Malacca.

Superficialiter Mallomonas teilingii (Teiling) Conrad similis, a

qua squamis quadratis, non discoideis, et setis paulo numerosiori-

bus differt.

Cellulae elongato-ellipticae, 50-55/a longae, 18-20/a latae, utrin-

que pariter rotundatae vel apice anteriori angustiusculae. Squamae

conspicue quadratae, spiraliter dispositae, apicales tantum setosae,

alterae apparenter leves, in singulis spiris maximis 5-9. Setae

graciles simplices, utraque cellulae apice 15-20 sitae, 30-32/a

longae. Flagellum longitudine semi-cellulare. Chromatophora dua,

magna, parietalia, fulva; ocellum prominens, coccineum, prope

apicem anteriorem situm. Leucosin unigranulare, magnum ad ex-

tremitatem posteriorem situm.

Malaya: Malacca—in fossis piscicolis. (Holotypus).

Philipose (1953) has described a species which shows some

resemblances to the present one, but he was not able to determine

the exact nature of the scales. He suggests that they were discoid

and circular, and this fact and his report of fewer setae at the ante¬

rior end, suggests that he was dealing with a different species.

32. Mallomonas tonsurata Teiling var. alpina (Pascher) Krieger.

[syn. Mallomonas alpina Pascher, Mallomonas tonsurata var.

megalepis Schiller]. CELLS small, ovoid, 15-20/a long X 10-

12/a wide, rounded at the posterior end, narrowing at the ante¬

rior end. SCALES small, elliptic, with a V-shaped mark on the

125

Gardens Bulletin, 5.

base of each, and arranged in very shallow spirals. SETAE con¬

fined to the anterior end, usually in three rows of six, compara¬

tively stout, toothed, longer than body length. CHROMATO-

PHORES 2, lateral and parietal, golden-yellow. STIGMA small,

crimson, situated at the anterior end. FLAGELLUM about body

length. (PI. Ill figs, a, b).

Malaya: Malacca—in fish-ponds.

Described from Europe, but reported also from Java.

This variety differs from the type mainly in its smaller size and

the toothed setae.

33. Mallomonas tonsurata Teiling var. dorsidentata Prowse var.

nov. Cells elongate-ovoid, 23-25/* long X 8-9/* wide, slightly

narrowed at the anterior end. SCALES small, elliptic, arranged

at a slightly acute angle, the rearmost scales very shortly un-

identate. SETAE very long, 20-30 in number, confined to the

anterior half of the cell, the apical ones 15/* long, shorter than

the infra-apical ones which are 20/* long. CHROMATO-

PHORES two large, discoid, parietal. FLAGELLUM nearly

body length. (PI. Ill fig. h).

Malaya: Malacca—in fish-ponds. (Type locality).

Holotype: Prowse 406 deposited at Tropical Fish Culture Re¬

search Institute, Malacca.

Cellulae elongato-ovoideae, apice antico paulo angustiores, 23-

25/* longae, 8-9/* latae. Squamae parvae, ellipticae, ad angulum

acutiusculum dispositae; squamae postremae brevissime uniden-

tatae. Setae longissimae in dimidia cellulae antica circumscriptae,

20-30, eae apicales 15/* longae breviores quam infra-apicales

20/* longae. Flagellum corpori fere aequilongum. Chromatophora

duo, magna, discoidea, parietalia.

Malaya: Malacca—in stagnis piscosis. (Holotypus).

CHRYSOPYXIS Stein 1878

Cells each enclosed in a cellulose envelope which is flask-shaped

in face view, with a slender thread like attachment extension at

the base; in side view the base is saddle-shaped, sitting on an algal

filament, with a thread like attachment extension each side, which

may completely surround the filament from one side to the other.

Chromatophores one or two, golden yellow, usually parietal.

Flagellum one, but often replaced by a slender protoplasmic ex¬

tension (rhizopodium) which may or may not be slightly branched,

in both cases protruding from the orifice of the envelope. Con¬

tractile vacuoles occur, apical when a flagellum is present, ap¬

parently basal when a rhizopodium is present; in the latter case

it seems likely that the cell has inverted within the envelope, so

that the rhizopodium arises from the morphologically posterior

end of the cell.

126

Vol. XIX. (1962).

34. Chrysopyxis sp. ENVELOPE flask-shaped, 10-12/a high X

6-8/a wide, sharply drawn out basally to a fine peduncle,

apically tapering to a narrow neck. CELL round, three quarters

filling the envelope. CHROMATOPHORES two, parietal, or

possibly one, divided deeply into two lobes. ‘FLAGELLUM’

long, twice body length, probably rhizopodial in nature since it

does not arise from the anterior end of the cell, but sometimes

from the side and sometimes from the morphological posterior

end, the cell lying at all angles in the envelope and may even

be inverted. CONTRACTILE VACUOLES variable in posi¬

tion. (PI. IV fig. e).

Malaya: Malacca—in fish-ponds.

This shows very great resemblances to Chrysopyxis iwanoffi

Lauterbom, but differs in that the rhizopodium is unbranched and

closely resembles a flagellum.

35. Chrysopyxis sp. ENVELOPE ovoid, 18-20/a long X 8-

1 0/a wide, narrowing apically to a widish neck, tapering basally

to a short peduncle. CELL round, two thirds filling the en¬

velope, slightly flattened anteriorly. CHROMATOPHERE one,

large, parietal, curving transversely round the cell. FLAGEL¬

LUM one, arising from the anterior end of the cell, li body

length. CONTRACTILE VACUOLES two, apical. (PI. IV fig.

f).

Malaya: Malacca—in fish-ponds.

This species resembles Chrysopyxis urna Korshikov in the enve¬

lope, but differs in retaining the flagellum and in the apical contrac¬

tile vacuoles.

ISOCHRYSIDEAE

2 equal flagella

1. Cells apparently naked, long stalked, epiphytic

(Stylochrysallis).

2. Cells within an envelope.ii.

ii. Envelope with a prominent orifice, no siliceous scales .... iii.

ii. Envelope with siliceous scales, completely surrounding the

protoplast.v.

iii. Envelope long stalked.( Stylotheca ).

iii. Envelope very shortly stalked or sessile.( Derepyxis ).

iv. Cells in pairs, never more, with broad bases attached to each

other. Chrysodidymus.

iv. Cells in spherical colonies, attached bases tapering .. Synura.

iv. Cells in bands usually of two rows, basal attachments broad

( Catenochrysis )

=* ( Chlorodesmus ).

127

Gardens Bulletin, S.

CHRYSODIDYMUS Prowse gen. nov.

Somewhat similar to Synura, this genus is readily distinguished

by its colonies being always of two cells, wider posteriorly. The

motion is forward and backwards in a straight line, occasionally

making a turn of 180° and moving on as before, never effecting its

movement as in Synura by rolling in complete revolutions. Cells

trapezoidal in shape, united directly and firmly at the posterior

ends to form bicellular colonies, the anterior end free and narrower

than the posterior ones. Involucre similar to that in Synura , with

siliceous, shortly spinose scales. Flagella two to each cell, apical,

apparently equal in length, in motion one pointing forward, the

other trailing behind. Chromatophores two, large, discoid and pa¬

rietal.

Synurae aliquatenus simile sed coloniis nunquam non bicel-

lularibus, protinus vel retrorsum recte progredientibus interdum

per 180° vertentibus et dein ut supra moventibus, nunquam ut in

Synura per gyrationes vel circumvolutiones proficientibus, cellulis

postice lateoribus hoc genus facile distinguendum.

Cellulae ambitu trapezoideae, apicibus posticis extemplo firmi-

terque in colonias bicellulares unitae; apicibus anticibus liberis

quam postices angustioribus; involucrum eo Synurae simile, squa-

mis siliceis, spinulosis praeditum. Flagella per cellulam singulam

bina, apicalia, simulate aequilonga, in motu unum porrectum, alte-

rum a tergo trahens. Chrornatophora bina, magna, discoidea, parie-

talia. Species holotypica: Chrysodidymus synuroideus Prowse.

36. Chrysodidymus synuroideus Prowse sp. nov. CELLS 14-15/x

long, posterior end 10-11 /a wide, anterior end 7-8/u. wide,

trapezoid in shape. CHROMATOPHORES. two large, parietal

golden yellow. FLAGELLA body length or slightly longer, ap¬

parently equal in length. (PI. IV fig. n.).

Malaya: Malacca— in acid swamps. (Type locality).

Holotype: Prowse 248a deposited at Tropical Fish Culture Re¬

search Institute, Malacca.

Cellulae ambitu trapezoideae, 14-15/x longae, apicibus poste-

rioribus 10—11 /a. latis, apicibus aiterioribus 7-8/x latis. Chromato-

phora dua, magna, parietalia, fulva. Flagella corpori aequilonga

vel paululo longiora, inter se aequilonga ut videtur.

Malaya: Malacca— in locis paludosis acidis (Holotypus).

37. Chrysodidymus gracilis Prowse sp. nov. CELLS longer and

narrower than those in the preceding species 22-24/x long,

posterior end 8.5-9/t wide, anterior end 6-6.5 jutwide. FLAGEL¬

LA nearly body length. (PI. IV fig. m).

Malaya: Malacca—in acid swamps.

Holotype: Prowse 248b deposited at Tropical Fish Culture Re¬

search Institute, Malacca.

128

Vol. XIX. (1962).

Cellulae eis prioris speciei longiores angustioresque, 22-24/a

longae, apicibus posterioribus 8.5-9 jx latis, apicibus anterioribus

6-6.5 fi latis. Flagella corpori fere aequilonga.

Malaya: Malacca in locis paludosis acidis. (Holotypus).

SYNURA Ehrenberg 1838

Cells ovoid, pyriform or clavate, united in spherical to sausage¬

shaped colonies, with the cells attached to each other by the atte¬

nuated posterior ends, even in two-celled colonies, thus differing

markedly from Chrysodidymus and Catenochrysis. Cell membrane

bearing siliceous scales which are frequently shortly spinose, one

spine to each scale. Chromatophores two, large, parietal, golden

yellow in colour. Flagella two to each cell, apparently equal in

length, but one projects forwards, the other trailing sideways; the

forward projecting flagellum is of the “tinsel” type, with a row

of very short cilia, the trailing flagellum being smooth. The colony

moves by rolling over and over, revolving through 360° continu¬

ously in one direction.

38. Synura uvella Ehrenberg. CELLS elongate-ovoid to pyriform,

20-25/a long X 12-15 /a wide, broadly rounded at the anterior

end, narrowing markedly at the posterior end, often to a thread¬

like attachment organ. CHROMATOPHORES two, large,

parietal, golden yellow. FLAGELLA two, apical, apparently

equal, more than body length. COLONIES of few to many

cells, spherical to sausage shaped. (PI. IV figs, d, k, 1).

Malaya: Selangor—in acid swamps near Kuala Lumpur.

Negri Sembilan—in various acid swamps.

Malacca—in acid swamps.

Singapore—in acid swamps near the Royal Island

Club golf course.

Probably to be found in all parts of Malaya.

Worldwide in distribution.

OCHROMONADEAE

2 unequal flagella

(i) Cells without cellulose envelopes, naked.ii.

(i) Cells with cellulose envelopes.iii-

(ii) Cells solitary . Ochromonas .

(ii) Cells colonial, usually in spherical colonies .... ( Uroglena ,

Volvochrysis, Synochromonas, Synuropsis).

129

Gardens Bulletin , S.

(iii) Cells in dendroid colonies, or epiphytic and solitary. Proto¬

plasts stalked inside envelope.iv.

(iii) Ceils solitary. Protoplasts not stalked inside envelope

{Pseudokephyrion, Kephyriopsis, Bitricha, Styloceras ).

(iv) Envelope markedly laminate, with ends of lamina projecting

Hyalobryon.

(iv) Envelope not laminate or if lamina present visible with diffi¬

culty and not projecting. Dinobryon .

OCHROMONAS Wyssotzki 1887

Cells usually solitary, naked, free swimming or occasionally

attached by a posterior threadlike extension. Shape very variable,

occasionally amoeboid. Periplast usually fine and smooth, rarely

thick and verrucose. Chromatophores 1-2, golden-yellow, often

very much reduced. Flagella two, unequal, the longer one pointing

forwards in swimming, the shorter trailing sideways. Palmelloid

stages often found.

39. Ochromonas hinterzartensis Doflein. CELL elongated ovoid,

14-15 fx long X 7-8/x wide, rounded at the posterior end,

slightly narrower anteriorly. PERIPLAST beset with peripheral

granules, but not verrucose. CHROMATOPHORE single, band¬

shaped, curving round lower half of cell, golden yellow in

colour. STIGMA absent. FLAGELLA two, unequal, the longer

body length, the shorter about 1/3 body length. LEUCOSIN

as four large oval granules. (PI. V fig. h).

Malaya: Malacca—in fish-ponds.

Described from Northern Europe.

This Malayan form seems to fit best into this species, especially

with its peripheral granules and several large leucosin granules.

The shorter flagellum is a little longer than that in the holotype

as described from Germany, but it is difficult to be accurate in

estimates of the lengths of flagella.

DINOBRYON Ehrenberg 1835

(includes Epipyxis (Ehrenberg) Lauterborn

and Dinobryopsis Lemmermann).

Cells free-floating, sessile or epiphytic, solitary or in colonies,

each cell enclosed in a conical, campanulate or cylindric cellulose

envelope, which is open at the top, pointed at the base, and may

have smooth or undulate sides, colonies dendroid, divergent or

compact. Protoplast oval to fusiform, attached to the base or side

of the envelope by a stalk. Chromatophores 1-2, elongate, parietal,

golden yellow in colour. Eyespot crimson, apical. Spherical

siliceous cysts are quite common in some species.

130

Vol. XIX . (1962).

40. Dinobryon bavaricum Imhof. [syn. Dinobryon stipitatum

Stein, D. stipitatum var. bavaricum (Imhof) Zacharias, D. stipi¬

tatum var. undulatum Lemmermann, D. stipitatum subspec. eus-

tipitatum Pascher, D. stipitatum subspec. bavaricum Pascher, D.

stipitatum var. affine (Lemmermann) Pascher, Dinobryon elon-

gatum Imhof, D. elongatum var. undulatum Lemmermann, D.

elongatum var. affine Lemmermann, Dinobryon bavaricum

Imhor. var. affine Lemmermann, Dinobryon sociale Ehrenberg

var. bavaricum (Imhof.) Bachman, Dinobryon cylindricum Imhof

var. ceylonicum Lemmermann]. CELLS in rather narrow den¬

droid colonies. ENVELOPE ( lorica ) 40-60/a long X 8-10 /a

wide, cylindric in the upper part with undulate walls, and drawn

out to a long pointed conical basal region. CHROMATOPHO-

RES two, parietal, golden yellow. FLAGELLA two, very un¬

equal, the longer H length of cell protoplast. (PI. IV fig. a).

Malaya: Malacca—in unlimed experimental fish-ponds and

acid swamps.

Distribution generally in colder waters in various parts of the

world.

The occurrence of this very striking species in fish ponds is

surprising, since it has generally been regarded as a cold-water

form. Even in India it has only been reported from high mountain

waters, whereas in Malacca it was quite frequent in unlimed fish¬

ponds with temperatures up to 31 ° C. It is probable that tempera¬

ture is not a determining factor for its occurrence, and some

chemical factor is of greater importance, particularly pH.

41. Dinobryon sertularia Ehrenberg [syn. Dinobryon thyrsoideum

Chodat, Dinobryon sertularia var. thyrsoideum (Chodat) Lem¬

mermann]. COLONIES dendroid, often large and thick and

rather bushy. ENVELOPE 25-30/ilong X 8-10/a wide, cylin¬

dric to slightly campanulate, swollen in the central portion,

flaring at the mouth, and tapering to a short bluntly pointed basal

portion. CHROMATOPHORES two, parietal, golden yellow.

FLAGELLA two, unequal, the longer about body length. CYSTS

quite common, spherical, with a curved apical neck to the pore,

the whole enclosed in a gelatinous envelope at the mouth of the

lorica. (PI. IV figs, b, c).

Malaya: Penang—acid swamps.

Perak—acid swamps near Ipoh.

Pahang—acid swamps near Kuantan.

Negri Sembilan—acid swamps near Seremban and

Port Dickson.

Malacca—acid fish-ponds and acid swamps.

Johore—acid swamps near Muar and Yong Peng.

Singapore—acid swamps near the Royal Island Club

golf course.

Worldwide in distribution.

131

Gardens Bulletin, S*

42. Dinobryon inflatum Lemmermann. CELLS solitary, epiphytic.

ENVELOPE broadly oval, 30-35/a long X 15-20/a wide,,

tapering to a short stalk basally and narrowing to a short but

distinct cylindrical neck apically. PROTOPLAST round to fusi¬

form, attached near the base of the envelope by a fine stalk.

CHROMATOPHORE one, apparently near the base, golden

yellow in colour. FLAGELLA two, unequal, the longer the

length of the protoplast, the other about i this length. (PI. IV

fig- g).

Malaya: Malacca—in acid swamp epiphytic on Mougeotia.

Originally described from Northern Europe.

I have tentatively placed the Malayan material under this species,

by virtue of the general shape of the envelope and the distinct neck

apically, but details of the original description are somewhat lack¬

ing.

HYALOBRYON Lauterborn 1896

Cells epiphytic, usually solitary but occasionally forming den¬

droid colonies, each cell in a cellulose envelope which is distinctly

laminate, formed of overlapping thimble-like segments. Envelope

(lorica) usually cylindric, sometimes narrower towards the anterior

end, occasionally spreading at the mouth, tapering at the posterior

end to the attachment point or even a stalk. Protoplast oval to

spindle-shaped with usually 2 unequal chromatophores, sometimes

apparently only one; chromatophores golden yellow in colour.

Flagella two, unequal. Cysts where known ellipsoid with a distinct

porus and plug.

43. Hyalobryon lauterbornii Lemmermann. CELLS solitary,

epiphytic. ENVELOPE cylindric, 35-40/a long X 8-10/a

wide, tapering to a pointed conical base, hardly flaring at the

mouth; laminations of the wall close together, starting from

the base and hardly projecting, most easily seen in empty en¬

velopes. PROTOPLAST ovoid, stalked, attached near base of

the envelope. CHROMATOPHORE apparently one, golden

yellow in colour. FLAGELLA two, unequal, the longer about

body length, the shorter i-1/3 as long. (PI. IV figs, h, i & j).

Malaya: Malacca—in experimental fish-ponds, epiphytic on

filamentous algae.

Worldwide in distribution. This is the commonest of the species

and is very variable.

The Colourless Flagellata

This is a heterogenous assemblage of genera and species with

divers affinities, which in many cases are somewhat obscure. They

all have in common the complete absence of a chromatophore

132

Vol. XIX. (1962).

and the method of nutrition may range from saprobic to com¬

pletely holozoic, some forms displaying distinctly animal-like

tendencies. The colourless Euglenineae, Volvocales and Dino-

flagellata, all of which show well marked affinities to pigmented

forms, are excluded from the account.

There are numerous ways of classifying the colourless flagel¬

lates, but since the number to be described below is small, the

simplest way is to separate them on the basis of flagellation.

KEY TO THE COLOURLESS FLAGELLATA

(i) Cells with flagella attached apically.ii.

(i) Cells with flagella attached to each lateral margin.v.

(ii) Cells with a single flagellum.iii.

(ii) Cells with two unequal flagella . ..iv.

(iii) Cells in stalked campanulate envelopes (lorica) arranged in a

dendroid fashion . Poteriodendron.

(iii) Cells arranged in a granular gelatinous sheaths

Phalansterium.

(iii) Cells with a flaring collar and enclosed in a thin lorica, usually

solitary. Salpingoeca .

(iv) Cells naked, colonial, at end of granular stalk . . Anthophysa.

(v) One flagellum at each margin.. . Turbomonas.

(v) Four flagella at each margin, two long and two short

Trepomonas.

POTERIODENDRON Stein 1878

Cells enclosed in campanulate, stalked cellulose lorica which

are arranged in a dendroid fashion, with each lorica attached by

means of its stalk to the inside of the lorica below. The proto¬

plasts are usually single in a lorica, uniflagellate, the flagellum

attached slightly subapically with a prominent lip on one side of

the apex; this lip is retractile and can be withdrawn. A contractile

vacuole is present in the basal part of the protoplast. One species

is known.

44. Poteriodendron petiolatum Stein. LORICA campanulate, 20-

25/z long X 8-10/x wide, slightly widened at the mouth, with

thickened walls basally and with an internal basal knob, each

lorica attached inside a lower one by a thin stalk 15-30/x long,

the lowest lorica with a long stalk 60-70/1 long, with a slight

basal holdfast. PROTOPLAST one in each lorica attached by

133

Gardens Bulletin, S .

a short stalk to the base, ovoid, 8-10/a long X 4-5 /a wide, with

a distinct apical lip which can be retracted. FLAGELLUM

single, apical, slightly longer than body length. (PI. V. fig. a).

Malaya: Malacca—in a ditch both attached and as free-swim¬

ming colonies.

Distribution world-wide.

PHALANSTERIUM Lienkowski 1865

Cells enclosed in a granular gelatinous mass, impregnated with

iron; the mass may be spherical (P. consociatum (Fresenius) Cien-

kowski) with the cells peripheral, or dendroid (P. digitatum Stein)

with the cells at the ends of the branches. Each cell is uniflagellate

with the long flagellum protruding through the surface of the gela¬

tinous mass, and the individual protoplasts each have a long narrow

apical collar. One or two contractile occur at the posterior end.

45. Phallansterium digitatum Stein. SHEATH dendroid, with

rounded very slightly concave ends, apparently hollow in the

greater part, terminal branches about 10 /a wide. CELL ovoid,

6-8/1 long X 3-4/a wide, with a long apical colour, not easily

visible and through which the flagellum protrudes; cell often

appearing slightly beaked sub-apically on living material. FLA¬

GELLUM single, 1 0/a or more long, protruding through the

ends of the gelationous sheath. (PI. V fig. f).

Malaya: Malacca—in acid swamps.

Worldwide in distribution.

The Malayan material shows a number of differences from the

type description, being somewhat smaller. The narrow apical col¬

lars are often difficult to see in the Malayan material, and the cells

often show a slightly beaked appearance not mentioned in the

original. This beaked appearance may be due to contraction of the

cell longitudinally, as often happens with cells under coverslips

under conditions of decreased aeration. The whole organism, des¬

pite its smaller size, so closely resembles Phalansterium digitatum

Stein that it seems logical to include it under that species, at least

until further investigation proves otherwise.

SALPINGOECA J. Clark 1867

Cells enclosed in a thin envelope or lorica, which is usually

constricted at the neck or mouth, and is basally attached, either

sessile or stalked. The protoplast may occupy part or nearly all

of the lorica. being attached to it by a thin stalk which may be

long to extremely short. A characteristic feature of the proto¬

plast is the slightly flaring funnel-like collar just below the apex

and protruding beyond the mouth of the lorica. There is a single

apical flagellum, and a single contractile vacuole.

134

Vol. XIX. (1962).

46. Salpingoeca frequentissima (Zacharias) Lemmermann [syn.

Diplosiga frequentissima Zacharias, Diplosigopsis frequentissima

(Zacharias) Lemmermann]. LORICA very thin, vase shaped,

10/a long X 5 wide, constricted at the neck and widely flaring

at the mouth, sharply pointed to very shortly stalked at the base.

Protoplast nearly filling the lorica, with a flaring collar 3-5 /a

long X 3-5/a wide projecting beyond the mouth of the lorica.

FLAGELLUM single, apical, about body length or slightly

longer. CONTRACTILE vacuole single, nearly central. (PI. V.

fig. b).

Malaya: Malacca—in a fish pond, attached to Mougeotia and

to phytoplankton.

Worldwide in distribution.

This is very common on planktonic diatoms in temperate waters,

but in Malaya is more frequent on filamentous algae, where it can

be very abundant indeed.

STOMATOCHONE Pascher 1942

Cells single, epiphytic on other algae, attached by a short stalk,

and obliquely truncate; deeply incurved at the apex so as to form

a rim or lip. Flagella two, distinctly unequal in length. There is

a single basal contractile vacuole.

There are reasons for thinking that these are colourless members

of the Chrysophyceae (Bourelly 1957), but in the absence of a

cyst such an affinity still remains uncertain. The organisms are

often as dense clusters on planktonic algae.

47. Stomatochone infundibuliformis Pascher. CELLS single,

small, 5-7/a long X 4—5/a wide, with an obliquely truncated

apex, rounding posteriorly and then narrowing to a very short

stalk by which it is attached. FLAGELLA two, very unequal,

the longer li body length, the shorter about i body length,

both arising from a prominent apical depression. CONTRAC¬

TILE vacuole basal, NUCLEUS median. (PI. V fig. d).

Malaya: Malacca—in an acid experimental fish-pond, and in

acid swamps, abundant as an epiphyte on Pleuro-

taenium kayei.

Worldwide in distribution.

The Malayan specimens seem much smaller than the size usually

given for this species, but it is doubtful if such size differences are

of real taxonomic significance.

ANTHOPHYSA Bory de St. Vincent 1822

Cells colonial, forming dense clumps at the ends of a pro¬

minent branched, brownish, somewhat granular, rigid gelatinous

stalk. Individual cells roughly conical, with two unequal apical

flagella. Stigma present or absent.

135

Gardens Bulletin, S.

48. Anthophysa vegetans (O. F. Mueller) Stein, [syn. Volvox vege¬

tans O. F. Mueller]. CELLS conical, 10-12/a long X 5-6 wide,

obliquely truncate at the anterior end with a slight central dep¬

ression, tapering posteriorly to a blunt, narrow, attachment end.

FLAGELLA two, apical, unequal, the longer li body length,

the shorter i to i body length. STIGMA absent in the Malayan

material. CONTRACTILE VACUOLE sub-apical. STALK

variable in length, rigid, gelatinous, thickly impregnated with

iron, sparsely branched. (PL V. fig. e).

Malaya: Malacca—in experimental fish-ponds, often attached

to decaying debris.

Worldwide in distribution.

TURBOMONAS Prowse gen, nov.

Cells small, hyaline, flattened, obovate or sub-cordate, spirally

twisted lengthwise. Flagella two, each attached midway on each

lateral margin. Motion forward, swiftly gyrational in nature, the

flagella whirling at right-angles to the longitudinal axis. Nucleus

one, central.

Cellulae parvae, hyalinae, applanatae, obovatae vel subcordatae,

longitudinaliter obtortae. Flagella bina, unumquodque in margine

utriusque lateris medio insertum. Motus cellulae porro ex natura

gyrationis, flagellis ad axin longitudinalem per angulum rectum

gyrantibus effectus. Nuclea uno, centrali.

Species holotypica: T. gyrans Prowse.

49. Turbomonas gyrans Prowse sp. nov. CELLS 12-13/a long,

7-9/a wide, 3-3.5/t thick, narrowed at the anterior end, wider and

almost flattened posteriorly, twisted lengthwise through 90°,

propelled forward both by the gyrations of the body and the

whirling of the flagella. EYESPOT absent. FLAGELLA nearly

as long as the body, pointing in opposite directions. CYTO¬

PLASM hyaline, with a few refractive granules. (PI. V fig. c).

Malaya: Selangor—Sg. Gombak, Kuala Lumpur Holotype

Prowse 388).

And occurring elsewhere in water containing putrid material.

Holotype is deposited at Tropical Fish Culture Research Insti¬

tute, Malacca.

Cellulae 12-13/t longae, 7-9/a latae, 3-3.5 ^ crassae, antice

angustatae, postice latiores et fere applanatae, per 90° longitudi¬

naliter obtortae, gyrationibus corporis flagello-rumque conjunctim

propulsae; ocellis nullis. Flagella corpori paene aequilonga, oppo¬

site directa. Cytoplasms hyalinum, cum paucis granulis refractivis.

Malaya: Selangor, Sg. Gombak, Kuala Lumpur (Holotypus).

In regionibus diversis: in aqua cum materiis putridis inquinata.

136

Vol. XIX. (1962).

This organism is reminiscent of the Distomataceae in the way it

rotates with the flagella whirling out at right angles, but it differs

in having a single central nucleus. Staining with Gentian Violet

suggests that although the flagella appear to be lateral in attach¬

ment, they are actually apical in origin, having fused with the

margin of the cell until midway. This species should be compared

with the next species, Trepomonas rotans, in which there are two

lateral nuclei and the flagella are truly laterally attached.

TREPOMONAS Dujardin 1841

Cells mainly broadly oval, flattened in cross-section, slightly

twisted, with two posterior lobe-like inflations. Nuclei two, one

near each margin. Flagella two long and two short on each margin,

opposite the nuclei, near the anterior end of each lobe. Cells swim

by sweeping the flagella at right angles to the long axis, the body

rotating and moving forward, all with a rather deliberate motion.

50. Trepomonas rotans Klebs. CELLS broadly oval, 1 8-20/a long

X 1 1-12/a wide, rounded at the anterior end, the main body

of the cell narrowing to the posterior end, but the lobes widen¬

ing posteriorly and projecting slightly beyond the end of the main

body, so that the posterior end of the cell is truncate and slightly

concave. FLAGELLA 2 long, more than body length, 2 short

about 1/3 body length, on each margin opposite each nucleus

which is about 1/3 from the anterior end. VACUOLES fairly

large and numerous. (PI. V. fig. m).

Malaya: Johore—stagnant water from Kota Tinggi.

Worldwide in distribution.

The slower deliberate rotating of the cell is very different from

the fast whirling of Turbomonas gyrans.

I am grateful to the Colonial Office and the Tropical Fish

culture Research Institute, Malacca for facilities to carry out these

studies. I am also very grateful to all those people who have kindly

collected material for me, among them Mr. H. M. Burkill, Director

of the Botanic Gardens, Dr. D. S. Johnson of the Department of

Zoology, University of Malaya in Singapore, Dr. Anne Johnson

of the Department of Botany in the same University, Miss Joan

Floyd (now Mrs. Charlton) late of Victoria Institution, Kuala

Lumpur, and Mr. S. F. Owen, P.W.D. Headquarters in Kuala

Lumpur. I am deeply grateful for all the encouragement 1 have

been given by the Department of Botany, University of Malaya,

and by the Botanic Gardens, Singapore. I am equally grateful to

all those generous people, too numerous to name, who have sent

me reprints, which have so greatly assisted this work. To the

Director of the Botanic Gardens 1 am especially grateful for per¬

mission to publish in the Gardens Bulletin. Finally 1 owe a deep

sense of gratitude to Dr. Furtado of the Botanic Gardens, Singa¬

pore, for the Latin translations.

137

Gardens Bulletin , S’,

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Biology 2nd Edition. New York; 117-170 & 190-231.

140

(a) Euglena intermedia (Klebs.) Schmitz var. klebsii Lemmermann; (b, c, d) Euglena muta-

bilis Schmitz; (e , /) Euglena gracilis Klebs; (g) Euglena flava Dangeard; (h, i) Euglena

agilis Carter; (/) Trachelomonas intermedia Dangeard; (it) Euglena exilis Gojdics; (/)

Lepocinclis fusiformis (Carter) Lemmermann; (m) Peranema inflexum Skuja; (n) Phacus

meson Pochmann; (o) Trachelomonas volvocinopsis Swirenko; (p) Strombomonas verrucosa

(Daday) Deflandre var. chinensis (Skvortzow) Deflandre; (q) Trachelomonas allia

Drezepolski fa.

Gardens Bulletin , 5.

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Vol. XIX. (1962)

143

(a, b ) Mallomonas tonsurata Teiling var. alpina (Pascher) Krieger; (c) Mallomonas splendens (G. S. West) Playfair; (d, k) Mallo¬

monas teilingioides Prowse sp. nov.; (e, f, l) Mallomonas acaroides Perty; (g, /) Mallomonas curta (Playfair) Conrad; (/*)

Mallomonas tonsurata var. dorsidentata Prowse var. nov.; (/) Mallomonas playfairii Conrad var. opisthiodonta Prowse var. nov.;

(m) Mallomonas spherica Prowse sp. nov .

Gardens Bulletin, S.

Plate IV iia

(a) Dinobryon bavaricum Imhof; (b , c) Dinobryon sertularia Ehrenberg; (d, k) Synura uve

Ehrenberg; ( e ) Chrysopxis sp.; (/) Chrysopxis sp.; (g) Dinobryon inflatum Lemmerm<' d

{h, i, i) Hyalobryon lauterbornii Lemmermann; (/) Synura uvella Ehrenberg, two-cei

colony; ( m ) Chrysodidymus gracilis Prowse sp. nov.; (n) Chrysodidymus synurow

Prowse sp. nov.

144

Vol. XIX. (1962)

145

(a) Poteriodendron petiolatum Stein; ( b ) Salpingoeca frequentissima (Zacharias) Lemmermann; (c) Turbomonas gyrans Prowse sp

nov.; (d) Stomatochone sp.; ( e ) Anthophysa vegetans (O. F. Muller) Stein; (/) Phalansterium digitatum Stein; '(g) Chromulina

sphaerica Bachman; (h) Ochromonas hinterzartensis Doflein: (/-/) Chrysamoeba radians Klebs; (m) Trepomonas rotans Klebs.

A Check List of Malayan Grasses

By H. B. Gilliland

Department of Botany, University of Singapore

The publication of Dr. N. L. Bor’s “The Grasses of Burma,

Ceylon, India and Pakistan” in 1960 makes it possible to draw

up a check list of the grasses of the Malay Peninsula with reason¬

ably up to date names.

In point of time, the first synthesis of our knowledge of the

grasses of the Malay Peninsula was prepared by Sir Joseph Hooker

in his “Flora of British India” Vol. VII which was published in

1897. That great graminologist, Dr. Otto Stapf, had just recently

joined the staff of the Kew Herbarium and assisted with this ac¬

count. Since Burma and the Malay Peninsula were at that time

administered from India, the plants of Burma and the Malay

Peninsula were included in that monumental work.

Simultaneously an independent synthesis of plant records for the

Malay Peninsula was begun by Dr. King—then Curator of the

Calcutta Herbarium in association with Mr. J. S. Gamble and

many other botanists. This is commonly referred to as King &

Gamble’s “Materials for a Flora of the Malay Peninsula”, and was

published in the Journal of the Asiatic Society of Bengal between

1889 and 1908. This, however, did not include the Monocoty¬

ledons which were specially dealt with by H. N. Ridley, then

Director of the Botanic Gardens of Singapore and the Straits Set¬

tlements, under the title of “Materials for a Flora of the Malay

Peninsula—Monocotyledons” in 1907/1908. Volume III of his

work, published in 1907, contains the first complete account of

the grasses of Malaya. On retirement Mr. Ridley devoted his time

to a further study of the flora of Malaya at Kew, and in 1925

published Volume V of “The Flora of the Malay Peninsula” which

carried our knowledge of Malayan grasses a stage further.

Mr. I. H. Burkill, who from 1912 to 1925 had succeeded Ridley

as Director of the Botanic Gardens of Singapore and the Straits

Settlements, published in 1935 a “Dictionary of the Economic

Products of the Malay Peninsula” in 2 volumes. This contains

further references to and discussions of many Malayan grasses.

The first work to merit discussion thereafter was a paper by

P. Jansen, a botanist on the staff of the Flora Malesiana Founda¬

tion at Leyden, who published “Notes on Malaysian Grasses” in

Reinwardtia, Vol. II, part 2, in 1953. Many references to herba¬

ceous grasses of the Malay Peninsula occur in this paper. Mean¬

time M. R. Henderson, by now Director of the Botanic Gardens,

147

Gardens Bulletin, S.

Singapore, produced the volume entitled “Malayan Wild Flowers

—Monocotyledons” published by the Malayan Nature Society in

Kuala Lumpur in 1954. This contains a useful account of the

commoner Malayan herbaceous grasses.

It will be noted that the last two works referred to have dis¬

cussed the grasses without reference to the aborescent bamboos.

However, Dr. R. E. Holttum, former Director of the Botanic

Gardens, Singapore and first Professor of Botany at the University

of Malaya, had been making a study of the bamboos and was able

to bring our knowledge of this interesting and important group

in Malaya up to date in a paper published in 1958 in the Gardens’

Bulletin, Singapore. Actually, Professor Holttum had prepared a

manuscript account of the grasses of the Malay Peninsula which

he very kindly passed on to the present writer and which has been

of great help in preparing the present list. Comments on the signi¬

ficance of the geographical distribution of the present records will

appear elsewhere.

1. Acroceras munroanum (Balansa) Henr. Blumea 3. 445,

1940; Bor, Grasses Burma, Ceylon, India & Pakistan,

275, 1960.

Panicum munroanum Balansa in Morot, J. de Bot. 4, 140;

1890.

Acroceras crassi-apiculatum (Merr.) Alston in Trimen

Handb. FI. Ceyl. 6 Suppl. 324, 1931; Burkill Kew Bull.

317, 1935; Diet. Econ. Prod. Mai. Pen. 1, 39, 1935.

Panicum ridleyi Hack. nom. nttd. ex Ridl. in Trans. Linn.

Soc. ser. 2, 3, 400, 1893.

Acroceras ridleyi (Hack.) Stapj ex Ridley, Flor. Mai. 5,

229, 1925.

A creeping grass of coastal sands.

2. Acroceras zizanioides (H.B.K.) Dandy, J. Bot. 69, 54, 1931;

Bor, l.c. 275, 1960.

Panicum zizanioides H.B.K. Nov. Gen. & Sp. 1, 110,

1816.

Acroceras sparsum Stapf. ex Ridl. Flor. Mai. 5, 229, 1925;

Burkill, l.c. I, 39, 1935.

Common in disturbed places with good light.

3. Alloteropsis cimicina (Linn.) Stapf. Flor. Trop. Afr, 9, 487.

1919; Ridl. Flor. Mai. 5, 223, 1925; Burkill, l.c. /, 105,

1935; Bor, l.c. 276, 1960.

Milium cimicinum Linn. Mant. Alt. 184, 1771.

A ruderal probably introduced through disturbance.

4. Apluda mutica Linn. Sp. PI. 82, 1753; Bor, l.c. 93, 1960.

Apluda varia var. intermedia Ridl. Mat. 3, 164, 1907; Flor.

Mai. 5, 207, 1925; Burkill, l.c. 1, 193, 1935.

A ruderal found only in the northern States.

148

Vol. XIX. (1962).

5. Aristida culionensis Pilg. in Perk. Fragm. FI. Philip 7, 145,

1904.

Has been recorded from Perlis.

6. Aristida setacea Retz. Obs. Bot. 4 , 22, 1786; Jansen in

Reinwardtia 2, 230, 1953; Bor, l.c. 412, 1960.

Aristida adscensionis Ridl. non Linn. Flor. Mai. 5, 242,

1925.

Rare in the northwest.

7. Arthraxon lanceolatus (Roxb.) Hochst. in Flora 59, 188,

1856; Bor, l.c. 100, 1960.

Andropogon lanceolatus Roxb. Flor. Ind. 7, 257, 1820.

Rare on limestone in Kedah.

8. Arthraxon nudus (Nees) Hochst. in Flora 59, 188, 1856;

Bor, l.c. 101, 1960.

Batratherum nudum Nees ex Steud . Syn. PI. Glum. 583,

1853.

Lankawi islands, rare.

9. Arundinefla setosa Trin. Gram. Panic. 63, 1826; Ridl. Flor.

Mai. 5, 239, 1925; Burkill, l.c. 7, 259, 1935; Bor, l.c.

424, 1960.

Occurs only in the northwest.

10. Arundo donax Linn. Sp. PI. 81, 1753; Agric. Bull. Straits

& F.M.S. 10, 260, 1911; Burkill, l.c. 7, 260, 1953; Bor,

l.c. 41, 1960.

“Teberau gading” Introduced.

11. Axonopus affinis Chase, J. Wash. Acad. Sci. 29, 180, 1938;

Bor, l.c. 227, 1960.

A. compressus var. affinis Henderson, Malay, Wild FIs.

Monocots. 339, 1954.

12. Axonopus compressus (Sw.) P. Beauv. Ess. Agrost. 12, 154,

167, 1812; Ridl. Flor. Mai. 5, 216, 1925; Burkill, l.c.

7, 276, 1953; Jagoe, Gard. Bull. S.S. 109, 1940; Hen¬

derson, l.c. 337, 1954; Gilliland, Comm. Mai. Pits. 26,

1958; Bor, l.c. 278, 1960.

Milium compressus Swartz Prodr. Veg. Ind. Occ. 24, 1788.

Paspalum platicaulon Poir. in Lamk. Encycl. Meth. Bot.

5, 34, 1804.

P. “platycaule” of Ridl. Mat. 3, 125, 1907.

Introduced “carpet grass” now abundant in lawns, etc.

13. Bambusa arundinacea (Retz.) Willd. Sp. PI. 2, 245, 1799;

Holttum, Gard. Bull. Sing. 16, 59, 1958.

Bambos arundinacea Retz . Obs. Bot. 5, 24, 1789; Holttum

in Taxon, 5, 65, 1956.

Planted in Singapore and Penang.

149

Gardens Bulletin, S.

14. Bambusa blumeana Schult. Syst. Veg. 7, 1343; Ridl. Flor..

Mai. 5, 256; 1925; Burkill, l.c. /, 298, 1935; Holttum,

l.c. 57, 1958.

Bambusa spinosa Blutne non Roxb. ex Nees. in Bot. Zeit.

580, 1825; Ridl. Mat. 3, 183, 1907.

The “buloh duri” is probably introduced.

15. Bambusa burmanica Gamble in Ann. R. Bot. Gard. Calcutta

7, 35, pi. 33, 1896; Holttum, l.c. 62, 1958.

Known only from northern Kedah.

16. Bambusa glaucescens (Willd.) Sieb. ex Munro Trans. Linn.

Soc. 26, 89, 1868; Holttum, Kew Bull. 2, 207, 1956;

l.c. 67, 1958; Gilliland. Comm. Mai. Pits. 29, 1958.

Ludolfia glaucescens Willd. in Ges. Nat. Freund. Berl.

Mag. 2, 320, 1808.

Bambusa multiplex Raeusch. non Louv. ex Burkill, l.c. 1,

299, 1935.

Bambusa nana Roxb. ex Munro l.c. 89, 1868; Ridl. Mat.

3, 184, 1907; Flor. Mai. 5, 258, 1925.

The “buloh pagar” makes an excellent hedge. Not native in

Malaya.

17. Bambusa heterostachva (Munro) Holttum in Jour. Arn. Arb.

27, 341, 1946; l.c. 65, 1958.

Gigantochloa heterostachya Munro in Trans. Linn. Soc.

26, 125, 1868; Ridl. Mat. 3, 188, 1907; Flor. Mai. 5, 262,

1925; Burkill, l.c. 1, 1069, 1935.

Gigantochloa latispiculata Gamble in Ann. R. Bot. Gard.

Calcutta 7, 67, pi. 59, 1896; Ridl. Mat. 3, 189, 1907r

Flor. Mai. 5, 263, 1935; Burkill, l.c. 1069, 1935.

Bambusa latispiculata (Gamble) Holttum, Jour. Arn. Arb.

27, 341, 1946.

Type from Malacca; also recorded Negri Sembilan, Johore,

Singapore & Perak.

18. Bambusa magica Ridl. J.S.B.R.A. Soc. 44, 208, 1904; Mat.

3, 184, 1907; FlSr. Mai. 5, 258, 1952; Burkill, l.c. 1,

299, 1935; Holttum, l.c. 75, 1958.

Bambusa elegans Ridl. J.S.B.R.A. Soc. 44, 208, 1904; Mat.

3, 185, 1907; Flor. Mai. 5, 258, 1925.

The “buloh perindu” is endemic on the higher more ex¬

posed ridges of the Main Range.

19. Bambusa montana (Ridl.) Holttum Kew Bull. 2, 206, 1956;

l.c. 77, 1958.

Dinochloa montana Ridl. J.S.B.R.A. Soc. 44, 2/0, 1905;

Mat. 3, 193, 1907; Flor. Mai. 5, 267, 1925.

Endemic to Penang Hill.

150

voi. xix. (mi).

20. Bambusa pauciflora Ridl. Flor. Mai. 5, 529, 1925; Burkill,

l.c. 7, 299, 1935; Holttum, l.c. 76, 1958.

“Buloh padi” is a rare endemic only known from Fraser’s

Hill.

21. Bambusa ridleyi Gamble, Ann. R. Bot. Gard. Calc. 7, 34,

pi. 32, 1896; Ridl. Mat. 3, 184, 1907; Flor. Mai. J,

257, f. 225, 1925; Holttum, l.c. 71, 1958.

Endemic from forest in Singapore and Pahang.

22. Bambusa vulgaris Schrad. in Wendl. Coll. PI. 2, 26, t. 47,

1810; Ridl. Mat. 3, 185, 1907; Flor. Mai. 5, 256, 1952;

Burkill, l.c. 7, 300, 1935; Holttum l.c. 63, 1958.

‘Buloh minyak" is the commonest cultivated bamboo of

Malaya.

23. Bambusa wrayi Stapf; Kew Bull. 14, 1893; Ridl. Mat. 3,

183, 1907; Flor. Mai. 5, 259, 1925; Burkill, l.c. 7, 301,

1935; Holttum, l.c. 72, 1958.

The “buloh bersumpitan” is an endemic species from the

mountains in Perak.

24. Bothriochloa intermedia (R. Br.) A. Camus, Ann. Soc. Linn.

Lyon. N.S. 76, 164, 1931; Bor. l.c. 108, 1960.

Andropogon intermedium R. Br. FI. N. Holt. 202, 1810;

Ridl. Mat. 3, 166, 1907; Henderson, l.c. 346, 1956.

Amphilophis glabra var. paupera Stapf in Ridl. Flor. Mai.

5, 209, 1925.

A rare roadside weed north of Johore.

25. Bothriochloa pertusa (Linn.) A. Camus l.c. 1931; Bor, l.c.

109, 1960.

Holcus pertusus Linn. Mant. Alt. 301, 1771.

Amphilophis pertusus (Linn.) Stapf in Flor. Trop. Afr. 9,

175, 1917; Flor. Mai. 5, 209, 1925.

Rare, recorded from Malacca.

26. Brachiaria distachya (Linn.) Stapf in Flor. Trop. Afr. 9, 565,

1919; Ridl. Flor. Mai. 5, 219, 1925; Burkill, l.c. 7, 356,

1935; Jansen, l.c. 238, 1953; Henderson, l.c. 342,

1954; Bor, l.c. 281, 1960.

Panicum distachyum Linn. Mant. 1, 138, 1767; Ridl. Mat.

3, 133, 1907.

Rather rare from Malacca & Port Swettenham.

27. Brachiaria holotricha Ohwi in Bull. Tokyo Sci. Mus. 18, 4,

1947; Jansen l.c. 238, 1953.

Rare, recorded from Penang.

151

Gardens Bulletin, 5.

28. Brachiaria mutica (Forsk.) Stapf l.c. 526, 1919; Ridl. Flor„

Mai. 5, 219, 1952; Burkill, l.c. /, 356, 1935.

Panicum muticum Forsk. Flor. Aegypt.Arab. 20, 1775;

Ridl. Mat. 3, 133, 1907.

A stoloniferous prolific grass introduced in Malaya.

29. Brachiaria paspaloides (Presl.) Hubb. in Hk. Ic. PI. T3363*

1938; Henderson, l.c. 342, 1954; Bor, l.c. 284, 1960.

Urochloa paspaloides Presl. Rel. Haenk. 1, 318, 1830.

A weed of cultivated and waste land.

30. Brachiaria reptans (Linn.) Gardn. & Hubb., Hk. Ic. PI. sub

t.3363, 1938; Bor, l.c. 285, 1960.

Panicum reptans Linn. Syst. Nat. ed. 10, 2, 870, 1759.

Urochloa reptans (Linn.) Stapf. l.c. 601, 1920; Ridl. Flor .

Mai. 5, 220, 1925; Burkill, l.c. 2, 2211, 1935 ,

A creeping perennial introduced at Port Swettenham.

31. Briza minor Linn, Sp. PI. 70, 1753; Bor, l.c. 528, 1960.

A European weed now established on hill stations.

32. Capillipeditim parviflorum (R. Br.) Stapf l.c. 169, 1917;

Bor, l.c. 112, 1960.

A swamp grass in Malaya.

33. Centotheca lappacea (Linn.) Desv. Nuov. Bull. Soc. Phil.

Paris 2, 189, 1810; Ridl. Mat. 3, 181, 1907; Flor. Mai.

5, 253, 1952; Burkill, l.c. /, 508, 1935; Henderson,

l.c. 309, 1954; Bor, l.c. 459, 1960.

Cenchrus lappaceus Linn. Sp. PI. ed. 2, 2, 1488, 1763.

Centotheca latifolia (Osbeck) Trin. Fund-Agrost. 141,

1820; Jansen, l.c. 253, 1953.

“Rumput lilit kain” is common associated with forest

paths, clearings, etc.

34. Centotheca longilamina Ohwi, Bull. Tokyo Sci. Mus. 18 ,

10, 1947; Jansen, l.c. 253, 1953.

C. lappacea var. longilamina (Ohwi) Bor, l.c. 459, 1960.

Also a forest grass.

35. Chloris harbata Sw. FI. Ind. Occ. /, 200, 1797; Ridl. Mat.

3, 173, 1907; Flor. Mai. 5, 250, 1925; Burkill, l.c. /,

529, 1935; Henderson, l.c. 315, 1954; Bor, l.c. 465,

1960.

An introduced ruderal now well established.

36. Chloris dolichostachya Lagasca Gen. et Spec. PI. 5, 1816;

Bor, l.c. 466, 1960.

Known only from the Lankawi islands in Malaya.

152

Vol. XIX. (1962).

37. Chrysopogon aciculatus (Retz.) Trin. Fund. Agrost. 188,.

1820; Ridl. Flor. Mai. 5, 207, 1925; Burkill, l.c. 7,

535, 1935; Henderson, l.c. 344, 1954; Gilliland, Comm.

Mai. Pits. 50, 1958; Bor, l.c. 115, 1960.

Andropogon aciculatus Retz . Obs. Bot. 5, 22, 1789; Ridl.

Mat. 3, 169, 1907.

The “Love-grass” of Malayan lawns and all inhabited land.

38. Chrysopogon collinus Ridl. Flor. Mai. 5, 208, 1925.

Jansen in Reinwardtia 2, 254, 1953 suggests that this may

turn out to be a form of C. montanus Trin. which Bor.

1. c. 116, 1960 equates to C. fulvus (Spreng.) Chiov.

A rare grass of the northwest.

39. Chrysopogon orientalis (Desv.) A. Camus, FI. Gen. Indoch.

7, 332, 1922.

Raphis orientalis Desv. Opusc. 69, 1831.

Andropogon wightianus Steud. Syn. PL Glum. 1, 395,

1854; Ridl. Mat. 3, 167, 1907.

Chrysopogon wightianus (Steud.) Thw. Enum. PI. Zeyl.

366, 1864; Ridl. Flor. Mai 5, 208, 1925.

Pahang, Kelantan & Trengganu and the northwest.

40. Coelachne simpliciuscula (Wight & Arn.) Munro ex Benth.

in J. Linn. Soc. Bot. 79, 93, 1881; Bor, l.c. 576, 1960.

Panicum simpliciusculum Wight & Arn. ex Steud. Syn. PI.

Glum. 1, 96, 1854.

A swamp grass from Kedah.

41. Coelorachis foveolata (Holtt.) Jansen l.c. 256, 1953.

Rottboellia foveolata Holttum, Gard. Bull. Sing. 9, 247,

1947 .

Rare recorded only from the northwest.

42. Coelorachis glandulosa (Trin.) Stapf ex Ridl. Flor. Mai. 5 ,

204, 1925; Burkill, l.c. 7, 618, 1935; Jansen, l.c. 255,

1953; Henderson, l.c. 347, 1954; Bor, l.c. 120, 1960.

Rottboellia glandulosa Trin. Mem. Acad. Petersb. ser. 6,

2, 250, 1833; Ridl. Mat. 3, 161, 1907.

A coarse grass of river banks widely distributed in Malaya.

43. Coelorachis helferi (Hk. f.) Henr. Blumea 4 , 518, 1941;

Jansen, l.c. 256, 1953; Bor, l.c. 120, 1960.

Rottboellia helferi Hk. f. Flor. Brit. Ind. 7, 158, 1896.

Mnesithea rupincola Ridl. J.S.B.R.A. Soc. 57, 116, 1910;

Flor. Mai. 5, 206, 1925.

Rare on riverbed rocks in Perak.

153

Gardens Bulletin, S.

44. Coix gigantea Koenig ex Roxb. FI. Ind. ed. 2, 3, 570, 1832;

Ridl. Flor. Mai. 5, 191, 1925; Bor, l.c. 264, 1960.

Coix lachryma-jobi var. gigantea (Roxb.) of Ridl. Mat. 3,

151, 1907.

From the Kinta river in Perak.

45. Coix lachryma-jobi Linn. Sp. PI. 972, 1753; Ridl. Mat. 3,

151, 1907; Flor. Mai. 5, 191, 1925; Burkill, l.c. /, 629,

1935; Henderson, l.c. 6, 354, 1954; Bor, l.c. 264, 1960.

Cultivated and wild races of “Job’s Tears” or “Jelai” are

found.

46. Cymbopogon calcicola C. E. Hubb. Kew Bull. 24, 1941;

Henderson, l.c. 354, 1954.

On limestone in Lankawi, Kedah and Pahang.

47. Cymbopogon flexuosus (Nees) Wats, in Atkins, Gaz. N.W.

Prov. Ind. 392, 1882; Bor. l.c. 127, 1960.

Andropogon flexuosus Nees ex Steud. Syn. PI. Glum. 1,

388, 1854. This is probably Ridley's “C. citratus Stapf'

(A. citratus DC) of the Flor. Mai. 5, 211, 1925; Burkill,

l.c. 1, 724, 1935; Henderson, l.c. 354, 1954.

“Lemon Grass.”

48. Cymbopogon nardus (Linn.) Rendle, Cat. Welw. Afr. PI. 2,

155, 1899; Ridl. Flor. Mai. 5, 211, 1925; Burkill, l.c.

/, 727, 1935; Henderson, l.c. 353, 1954; Bor, l.c. 130,

1960.

Andropogon nardus Linn. Sp. PI. 1046, 1753; Ridl. Mat.

3, 168, 1907.

“Citronella grass.”

49. Cynodon arcuatus Presl. Rel. Haenk. /, 290, 1830; Bor, l.c.

469, 1960.

Cynodon dactylon Ridl. non Pers. Mat. 3, 173, 1907; Flor.

Mai. 5, 249, 1925; Burkill, l.c. 1, 729, 1935; Henderson,

l.c. 314, 1954.

A stoloniferous but not rhizomatous species, widespread,

flowering only in dry spells.

50. Cyrtococcum accrescens (Trin.) Stapf in Hook. Ic. PI. t3096,

1922; Ridl. Flor. Mai. 5, 234, 1925; Burkill, l.c. /, 744,

1935; Jansen, l.c. 257, 1953; Bor, l.c. 291, 1960.

Panicum accrescens Trin. Sp. Gram. Ic. 1, pi. 88, 1827.

Panicum patens Ridl. non Linn. Mat. 3, 142, 1907.

A grass tolerant of some shade, widely distributed.

51. Cyrtococcum oxyphyllum (Steud.) Stapf in Hk. Ic. PI. t3096,

1922; Ridl. Flor. Mai. 5, 233, 1925; Bor, l.c. 291,

1960.

Panicum oxyphyllum Steud. Syn. PI. Glum. 1, 68, 1854.

154

Vol. XIX. (1962).

:2\ Panicum pilipes Nees et Arn. ex Buese in Miq. PI. Jun-

ghn. 3, 376, 1854; Ridl. Mat. 3, 141, 1907; Henderson ,

l.c. 331, 1954.

Panicum trigonum Ridl. non Retz. J.S.B.R.A. Soc. 23, 21,

1891.

A creeping grass of shady places, widespread.

52. Cyrtococcuni patens (Linn.) A. Camus, Bull. Mus. Nat. Hist.

Paris 27, 118, 1921; Bor, l.c. 292, 1960.

Panicum patens Linn. Sp. PI. 58, 1753.

Cyrtococcuni carinatum (Presl.) Stapf in Ridl. Flor. Mai.

5, 233, 1925.

Cyrtococcum trigonum (Retz.) A. Camus, Burkill, l.c. 1,

744, 1935.

Panicum trigonum Henderson l.c. 330, 1954.

From Malacca, Negri Sembilan & Selangor.

53. Dactyloctenium aegyptium (Linn.) Beauv. Ess. Agrost. ExpL

PI. 15, 159, 1812; Henderson, l.c. 316, 1954; Bor. l.c.

489, I960.

Cynosurus aegyptius Linn. Sp. PI. 72, 1753.

Dactyloctenium aegyptiacum Willd. Enum. Hort. Berol.

1029, 1809; Ridl. Flor. Mai. 5, 251, 1925; Burkill, l.c .

I, 747, 1953.

Eleusine aegyptiaca Ridl. non Desf. Mat. 3, 174, 1907.

An introduced weed of sandy places.

54. Dendrocalamus asper (Schult.) Backer ex Heyne Nutt. PI.

Ned. Ind. ed. 2, /, 301, 1927; Burkill, l.c. /, 781, 1935;.

Holttum Gard. Bull. Sing. 16, 100, 1958.

Bambusa aspera Schult. Syst. Nat. 7, 1352, 1830.

Dendrocalamus flagellifer Munro Trans. Linn. Soc. 26,.

150, 1866; Ridl. Flor. Mai. 5, 265, 1925.

The “buloh betong” or “Rebong China” is a favourite

“bamboo shoot”. Widespread.

55. Dendrocalamus dumosus (Ridl.) Holttum. Gard. Bull. Sing.

II, 296, 1947 & l.c. 96, 1958.

Schizostachym dumosum Ridl. J.S.B.R.A. Soc. 61, 64,.

1912; Flor. Mai. 5, 271, 1925.

A rare bamboo from limestone in Kedah and the Lankawi

islands.

56. Dendrocalamus elegans (Ridl.) Holttum, Gard. Bull. Sing.

11, 296, 1947 & l.c. 95, 1958.

Schizostachyum elegans Ridl. J.S.B.R.A. Soc. 73, 146,

1916; Flor. Mai. 5, 271, 1925.

Another rare bamboo from limestone in the Lankawi

islands.

155

Gardens Bulletin, S.

:57. Dendrocalamus giganteus Munro, Trans. Linn. Soc. 26, 150,

1868; Ridl. Mat. 5, 191, 1907; Flor. Mai. 5 , 265, 1925;

Burkill, l.c. 7, 781, 1935; Holttum, l.c. 103, 1958.

“Buloh semiliang” is sporadic probably introduced.

58. Dendrocalamus hirtellus Ridl. J.S.B.R.A. Soc. 75, 146, 1916;

Flor. Mai. 5, 266, 1925; Holttum, l.c. 93, 1958.

“Buloh kapor” is recorded from open places in forest in

Johore, Kedah & Kelantan.

59. Dendrocalamus pendulus Ridl. J.S.B.R.A. Soc 44, 210, 1905;

Mat. 5, 192, 1907; Flor. Mai. 5, 266, 1925; Burkill,

l.c. 1, 782, 1935; Holttum, l.c. 90, 1958.

Cephalostachyum malayanum Ridl. J.S.B.R.A. Soc. 57,

118, 1910; Flor. Mai. 5, 267, 1925.

Schizostachym subcordatum Ridl. J.S.B.R.A. Soc. 82, 204,

1920; Flor. Mai. 5, 269, 1925.

“Buloh akar” is abundant in the valleys of the Main

Range.

60. Dendrocalamus sinuatus (Gamble) Holttum, l.c 11, 296,

1947; 16, 97, 1958.

Oxytenanthera sinuata Gamble, Ann. R. Bot. Gard. Calc.

7, 71. pi. 62, 1896; Ridl. Flor. Mai. 5, 264, 1925; Burkill,

l.c. 2, 1617; 1935.

This “buloh akar”, whose locus classicus is Seremban,

Negri Sembilan, is uncommon.

61. Dendrocalamus strictus (Roxb.) Nees, Linnaea 9, 476, 1834,

Burkill, l.c. 7, 782, 1935; Holttum, l.c. 98, 1958.

Bambo stricta Roxb. Corom. PI. 7, 58, t80, 1798.

The “male bamboo” of India is known in cultivation in

Penang, Kuala Lumpur & Singapore.

62. Dichanthium annulatum (Forsk.) Stapf. Flor. Trop. Afr. 9,

178, 1917; Bor, l.c. 133, 1960.

Andropogon annulatus Forsk, FI. Aeg.-Arab. 173, 1795.

This and the next species are hay grasses which do best in

the north.

63. Dichanthium caricosum (Linn.) A. Camus, Bull. Mus. Nat.

Hist. Paris 27, 549, 1921; Ridl. Flor. Mai. 5, 210,

1925; Burkill, l.c. 7, 802, 1935; Bor, l.c. 134, 1960.

Andropogon caricosus Linn. Sp. PI. ed. 2, 1480, 1763.

Recorded from Kedah.

64. Digitaria adsccndens (H.B.K:) Henr., Blumea, 7, 92, 1934;

Bor, l.c. 298, 1960

Panicum adscendens H.B.K. Nov. Gen. el Sp. PI. 1, 97,

1816.

156

Vol. XIX. (1962).

Digitaria marginata Link., Hort. Berol. 1, 229, 1821; Ridl.

Flor. Mol. 5, 214, 1925; Burkill, & c. 1, 808, 1935; Hen¬

derson, l.c. 340, 1954.

Digitaria chinensis Hornem. Suppl. Hort. Bot. Hafn. 8,

1819; Ridl. Flor. Mai. 5, 215, 1925.

65. Digitaria bicornis (Lamk.) Roem. & Schultz ex Lond. Hort.

Brit. 24, n. 1578, 1830; Bor, l.c. 299, 1960.

Paspalum bicorne Lamk. Tab. Encycl. Meth. Bot. 1, 176,

1791.

Paspalum heteranthum Hk. ). non Link, nec Nees et

Mayen Flor. Brit. Ind. 7, 16, 1896; Ridl. Mat. 3, 126,

1907.

Digitaria barbata Willd. Enum. Hort. Berol. 91, 1809;

Ridl. Flor. Mai. 5, 215, 1925.

Recorded from the eastern coastal areas.

66. Digitaria didactyla Willd. Enum. PI. Hort. Berol. 91, 1809;

Burkill, l.c. /, 809, 1935; Gilliland, Comm. Mai. Pits.

80, 1958; Bor. l.c. 300, 1960.

“Serangoon grass” is widely used for lawns but is not

native.

67. Digitaria longiflora (Retz.) Pers. Syn. 1, 85, 1805; Ridl. Flor.

Mai. 5, 214, 1925; Burkill, l.c. /, 808, 1935; Hender¬

son, l.c. 342, 1954; Bor, l.c. 302, 1960.

Paspalum longiflorum Retz. Obs. 4, 15, 1786; Ridl. Mat.

3, 126,

A roadside ruderal widespread in Malaya.

68. Digitaria pertenuis Buse in Miq. PI. Junehn. 381, 1854; Ridl.

Flor. Mai. 5, 214, 1925.

A garden weed from Penang and Singapore.

69. Digitaria pusilia Ridl. Flor. Mai. 5, 215, 1925.

A rare grass recorded from N. Kedah.

70. Dimeria ciliata Merr. Philip. J. Sci. 9, 262, 1914; Jansen,

l.c. 267, 1953.

Dimeria fuscescens var. barbata Ridl. Flor. Mai. 5, 191,

1925.

Rare in the far north.

71. Dimeria glabra Ridl. Flor. Mai. 5, 192, 1925.

Dimeria ornithopoda var. glabra (Ridl.) Jansen, l.c. 266,

1953.

A rare grass of drying swamps.

72. Dimeria kurzii Hk. f. FI. Brit. Ind. 7, 103, 1897; Ridl. Flor.

Mai. 5, 192, 1925; Bor, l.c., 142, 1960.

Rare in the far north.

157

Gardens Bulletin , S.

73. Dimeria leptorachis Hack. Monogr. Androp. 89, 1889;

Jansen, l.c. 267, 1953.

Once recorded from Kedah.

74. Dimeria ornithopoda Trin. Fund. Agrost. 167, tl4, 1820;

Ridl. Mat. 3, 151, 1907; Flor. Mai. 5, 192, 1925;

Burkill, l.c. /, 810, 1935; Jansen, l.c. 265, 1953; Hen¬

derson, l.c. 346, 1954; Bor, l.c. 142, 1960.

A grass of lawns & dry grassy spots.

75. Dinebra retroflexa (Vahl.) Panz., Denkschr. Acad. Moench.

274, tl2, 1814; Bor, l.c. 491, 1960.

Cynosurus retro flex us Vahl. Symb. Bot. 2, 20, 1791.

Dinebra arabica Jacq . Fragm. 77, tl21, 1809; Ridl. Flor.

Mai. 5, 249, 1925.

Seems to have come and gone from Malaya at Port Swet-

tenham.

76. Dinochloa seandens (Bl.) O. Ktze. Rev. Gen. PI. 773, 1891;

Merrill, J.S.B.R.A. Soc. 86 , 53, 1921; Holttum, Gard.

Bull. Sing. 16, 84, 1958.

Bambusa seandens Bl. ex Nees in Flora 7, 291, 1824.

Dinochloa tjankorreh Buse ex Miq. PI. Jungh. 388, 1854;

Ridl. Mat. 3, 192, 1907; Flor. Mai. 5, 267, 1925.

A “buloh akar” from the Dindings & Perak.

77. Dinochloa sp. Holttum, Gard. Bull. Sing. 16, 85, 1958.

From Gunong Panti & G. Pulai in Johore.

78. Diplachne fusca (Linn.) P. Beauv. Agrost. 80, 163, 1812;

Burkill, l.c. /, 834, 1935; Bor, l.c. 492, 1960.

Festuca fusca Linn. Sp. PI. ed. 2, 109, 1762.

Widespread in the north.

79. Diplachne malayana C. E. Hubb., Kew Bull. 106, 1934.

Recorded from North Kedah.

80. Echinochloa colonum (Linn.) Link. Hort. Berol. 209, 1827;

Ridl. Flor. Mai. 5, 222, 1925; Burkill, l.c. /, 888, 1935;

Henderson, l.c. 336, 1954; Gilliland, Comm. Mai. Pits.

89, 1958; Bor, l.c. 308, 1960.

Panicum colonum Linn. Syst. Nat. ed. 10, 2, 870, 1759;

Ridl. Mat. 3, 132, 1907.

“Padi burong” is associated with padi as a weed of cul¬

tivation.

81. Echinochloa crus-galli (Linn.) Beauv. Ess. Agrost. 53, 161,

1812; Ridl. Flor. Mai. 5, 222, 1925; Burkill, l.c. /,

889, 1935; Henderson, l.c. 337, 1954; Bor, l.c. 310,

1960.

Panicum crus-galli Linn. Sp. PI. 56, 1753; Ridl. Mat. 3,

132, 1907.

158

voi xix. (m2).

A sporadic weed like the last and also called “padi

burong”.

82. Echinochloa fcrumentacea Link. Hort. Berol. /, 214, 1827;

Bor, l.c. 311, 1960.

Echinochloa crus-galli var. frumentacea Ridl. Flor. Mai.

5, 223, 1925.

Panicum crus-galli var. frumentaceum Ridl. Mat. 3, 132,

133, 1907.

Occasionally cultivated.

83. Echinochloa stagnina (Retz.) P. Beauv. Ess. Agrost. 53, 161,

171, 1812; Burkill, l.c. /, 888, 1935; Bor, l.c. 311,

1960.

Panicum crus-galli var. stagninum Ridl. Mat. 3, 132, 1907.

Occasional escape from cultivation.

84. Eleusine corocana (Linn.) Gaertn. Fruct. & Sem. /, 8, t.l.fll,

1788; Ridl. Mat. J, 174, 1907; Burkill, l.c. /, 916,

1935; Bor, l.c. 492, 1960.

Cynosurus corocanus Linn. Syst. Nat. ed. 10, 2, 875, 1759.

A cultivated plant sometimes found in Malaya, “Ragi”.

85. Eleusine indica (Linn.) Gaertn. Frust. & Sem. /, 8, 1789;

Ridl. Mat. J, 173, 1907; Flor. Mai. 5, 250, 1925;

Burkill, l.c. /, 917, 1935; Bor, l.c. 493, 1960.

Cynosurus indicus Linn. Sp. PI. 72, 1753.

“Rumput sambau” is a weed of waste land throughout

Malaya.

86. Eragrostis atrovirens (Desf.) Trin. ex Steud. Nom. Bot. ed.

2, /, 562, 1840; Bor, l.c. 503, 1960.

Poa atrovirens Desf. FI. Atlant. 1, 73, tl4, 1798.

Eragrostis elegantula (Kunth) Nees in Steud. Syn. PI.

Glum. 1, 271, 1854 non E. elegantula Nees 1851; Ridl.

Mat. 3, 179, 1907; Flor. Mai. 5, 248, 1925; Burkill, l.c.

1, 933, 1935.

Eragrostis chariis Auct. now (Schult.) Hitchc.; Hubb. in

Kew Bull. 345, 1949; Jansen, l.c. 269; 1953.

A glaucous perennial which survives periodic inundation.

87. Eragrostis japonica (Thunb.) Trin., Mem. Acad. Petersb. ser.

6, /, 405, 1831; Bor, l.c. 509, 1960.

Poa japonica Thunb. Flor. Jap. 51, 1784.

Diandrochloa japonica (Thunb.) de Winter in Bothalia 7,

2, 388, 1960.

Eragrostis interrupta Ridl. Mat. 3, 178, 1907; Flor. Mai.

5, 246, 1925; Burkill, l.c. 1, 934, 1935.

A tall grass rare in the north of Malaya.

159

Gardens Bulletin, S .

88. Eragrostis inalavana Stapf in Hk. f. Flor. Brit. Ind. 7, 317„

1897; Ridl. Mat. 3, 178, 1907; Flor. Mai. 5, 247,

1925; Jansen, l.c. 272, 1953; Bor, l.c. 510, 1960.

Resembling E. unioloides but with more numerous florets,

widespread in sandy sites.

89. Eragrostis pilosa (Linn.) Beauv. Ess. Agrost. 71, 162, 175,.

1812; Ridl. Mat. 3, 179, 1917; Flor. Mai. 5, 247, 1925;

Burkill, l.c. I, 934, 1935; Bor, l.c. 512, 1960.

Poa pilosa Linn. Sp. PI. 68, 1753.

A sporadic ruderal of cultivation.

90. Eragrostis tenella (Linn.) Beauv. ex Roem. & Schult. Syst.

Veg. 2, 576, 1817; Ridl. Mat. 3, 177, 1907; Flor. Mai.

5, 245; Jansen, l.c. 274, 1953; Bor, l.c. 513, 1960.

Poa tenella Linn. Sp. PI. 69, 1753.

Eragrostis amabilis Kuntze of Burkill, l.c. I, 933, 1935,

A common weed throughout Malaya.

91. Eragrostis unioloides (Retz.) Nees in Steud. Syn. PI. Glum.

/, 264, 1854; Burkill, l.c. /, 934, 1935; Bor, l.c. 515,

1960.

Eragrostis amabilis Ridl., non W. & A. in Mat. 3, 178 ,

1907; Flor. Mai. 5, 246, 1925.

“Rumput kolam padang” is widespread as a weed.

92. Eremochloa ciliaris (Linn.) Merr. Philip. J. Sci. /, Suppl. 5,

331, 1906; Bor, l.c. 146, 1960.

Nardus ciliaris Linn. Sp. PI. 53, 1753.

Eremochloa malayana Ridl. Mat. 3, 155, 1907; Flor. Mai,

5, 196, 1925; Burkill, l.c. 1, 935, 1935.

A rare grass of the north of Malaya.

93. Eriachne pallcsccns R. Br. Prodr. FI. Nov. Holl. 184, 1810;

Ridl. Mat. 3, 172, 1907; Flor. Mai. 5, 240, 1925;

Burkill, l.c. 1, 936, 1935; Henderson, l.c. 316, 1954;

Bor, l.c. 479, 1960.

Common in dry exposed places.

94. Eriochloa procera (Retz.) Hubb., Kew Bull. 256, 1930;

Jansen, l.c. 276, 1953; Bor, l.c. 312, 1960.

Agrostis procera Retz. Obs. Bot. 4, 19, 1786.

Eriochloa polvstachya Ridl. non H.B.K. in Mat. 3 127

1907.

Eriochloa annulata (Fluegge) Kunth, Rev. Gram. 1, 30,

1829; Ridl. Flor. Mai. 5, 223, 1925; Henderson, l.c. 325,

1954.

Eriochloa ramosa (Retz.) Ktze; Burkill, l.c. 1, 938, 1935.

Waste ground throughout Malaya.

160

Vol. XIX. (1962).

95. Eulalia leschenaultiana (Dene) Ohwi, Bull. Tokyo Sci. Mus.

18, 2, 1947; Jansen, l.c. 277, 1953; Bor, l.c. 155, 1960.

Andropogon leschenaultianus Dene, Herb. Timor. Descr.

29, 1835.

Recorded from the Peninsula fide Jansen.

*96. Eulalia milsumi Ridl. Flor. Mai. 5, 196, t222, 1925; Jansen,

l.c. 277, 1953.

Local from Klang Gates, Selangor.

*97. Eulalia ridleyi (Hack.) Stapf. ex Ridl. Flor. Mai. 5, 197,

1925; Jansen, l.c. 277, 1953.

Pollinia ridleyi Hack. Oestr. Bot. Zeitsch. 40, 1, 7, 1891;

Ridl. Mat. 3, 155, 1907.

Sand dunes of the East Coast of Malaya.

'98. Eulalia speciosa (Debeaux) O. Ktze. Rev. Gen. PI. 2, 775,

1891; Bor, l.c. 157, 1960.

Erianthus speciosus Debeaux, Act. Soc. Linn. Bordeaux

32, 53, 1878.

Eulalia lanipes Ridl. J. F.M.S. Mus. 7, 56, 1916; Flor.

Mai. 5, 196, 1925; Jansen, l.c. 277, 1953.

Recorded from Kedah Peak.

■99. Eustachys tenera (Presl.) C. E. Hubb. Kew Bull. 25, 1941.

Cynodon tener Presl. Rel. Haenk. 1, 291, 1830.

Chloris ridleyi Hack. Oestr. Bot. Zeitsch. 52, 237, 1902.

Penang, Kelantan, Pahang, Singapore.

100. Gamotia stricta Brogn. in Duperrey, Voy. Coq. Bot. 133,

t21, 1831; Ridl. Flor. Mai. 5, 242, 1925; Burkill, l.c.

/, 1061, 1935; Bor, l.c. 569, 1960.

Usually at high altitudes in the mountains.

101. Gigantochloa apus (Schult.) Kurz, Tijds. Ned. Ind. 27, 226,

1864; Holttum, Gard. Bull. Sing. 16, 112, 1958.

Bambusa apus Schult. Syst. Veg. 7, 1353, 1830.

Cultivated in Singapore and at Serdang, Selangor.

102. Gigantochloa hasskarliana (Kurz.) Backer ex Heyne Nutt.

Plant. Ned. Ind. ed. 2, /, 299, 1927; Holttum, l.c. 118,

1958.

Schizostachyum hasskarlianum Kurz, Indian Forester 1,

353, 1876.

Oxytenanthera nigrociliata Munro non Buse, Trans. Linn.

Soc. 26, 128, 1868; Ridl. Mat. 3, 189, 1917; Flor. Mai.

5, 263 , 7925; Burkill, l.c. 2, 1617, 1935.

Cultivated for tall hedges in Singapore and Penang.

103. Gigantochloa latifolia Ridl. Flor. Mai. 5, 262, 1925; Holt¬

tum, l.c. 132, 1958.

161

Gardens Bulletin, S.

Oxytenanthera auriculata Prain ex Ridl. Flor. Mai. 5, 264,

1925.

var. alba Holttum, l.c. 133, 1958.

var. efimbriata Holttum, l.c. 133, 1958.

Recorded from Kedah & Pahang & cultivated in Singapore.

104. Gigantochloa levis (Blanco) Merr. Am. J. Bot. 3 , 1916; Holt¬

tum, l.c. 119, 1958.

Bambusa levis Blanco, Flor. Filip, ed. 1, 272, 1837.

Gigantochloa verticillata Munro non Willd. ex Ridl. Mat.

3, 186, 1907; Flor. Mai. 5, 260, 1925. p.p.; Burkill, l.c.

1, 1019, 1935.

Bamboo shoots are prepared from this widely cultivated

species.

105. Gigantochloa tigulata Gamble, Ann. R. Bot. Gard. Calc.

7, 67, pi. 58, 1896; Ridl. Mat. 3, 188, 1907; Flor. Mai.

5, 262, 1925; Burkill, l.c. /, 1069, 1935; Holttum, l.c.

129, 1958.

A species with many forms from the northern States of

Malaya.

106. Gigantochloa maxima Kurz, Ind. Flor. 1, 343, 1876; Holt¬

tum, 114, 1958.

Gigantochloa verticillata Munro non Willd. in Ridl. Mat.

3, 1907\ Flor. Mai. 5, 260, 1925. p.p.; Burkill, l.c. 1,

1069, 1935.

var. minor Holttum, l.c. 116, 1958.

var. viridis Holttum, l.c. 115, 1958.

A bamboo valued for its strength as a structural material.

107. Gigantochloa ridleyi Holttum, Gard. Bull. Sing. 15, 275,

1956; l.c. 127, 1958.

Cultivated in Province Wellesley and Singapore.

108. Gigantochloa scortechinii Gamble, Ann. R. Bot. Gard. Calc,

7, 62, pi. 53, 1896; Ridl. Mat. 3, 186, 1907; Flor. Mai.

5, 261, 1925; Burkill, l.c. 1, 1069, 1935; Holttum, l.c.

122, 1958.

var. albovestita Holttum l.c. 124, 1958.

The “buloh aur” grows in the north and along the Main

Range.

109. Gigantochloa wrayi Gamble, l.c. 64, pi. 55, 1896; RidL

Mat. 3, 187, 1907; Flor. Mai. 5, 261, 1925; Burkill,

l.c. /, 1070, 1935; Holttum, l.c. 124, 1958.

Gigantochloa kurzii Gamble, l.c. 65, 1896; Ridl. Mat. 3 „

187, 1907; Flor. Mai. 5, 261, 1925 quoad spec, malay.

“Buloh beti” is native to the northern States.

162

VoL XIX. (1962).

110. Haekelochloa granulans (Linn.) O. Ktze Rev. Gen. PI. 2,

776, 1891; Bor, l.c. 159, 1960.

Cenchrus granulans Linn. Mant. 2, 575, 1771.

Manisuris granularis (Linn.) Linn. f. Nov. Gram. Gen.

40, 1779; Ridl. Mat. 3, 163, 1907; Flor. Mai. 5, 205,

1925; Bur kill, l.c. 2, 1422, 1935.

A rare weed of sandy places.

3 11. Hemarthria vaginata Buse in Miq. Plant. Jungh. 354, 1854.

Rottboellia protensa Ridl. non Hack, Mat. 3, 163, 1907.

Hemarthria protensa Ridl. non Steud., Flor. Mai. 5, 205,

1925.

A rare grass recorded in Kedah and Kelantan.

112. Heteropogon contortus (Linn.) Beauv. ex Roem. & Schult.

Syst. Veg. 2, 836, 1817; Ridl. Flor. Mai. 5, 213, 1925;

Burkill, l.c. 7, 1143, 1935; Bor, l.c. 163, 1960.

Andropogon contortus Linn. Sp. PI. 1045, 1753; Ridl. Mat.

3, 167, 1907.

Open sandy places of coastal areas.

1 13. Hymenachne aurita (Presl.) Backer in Heyne, Nutt. PI. Ned.

Ind. ed. 2, 7, 197, 1922.

Panicum auritum Presl. ex Nees Agrost. Bras. 1/6, 1829;

Ridl. Mat. 3, 135, 1907; Flor. Mal. 5, 226, 1925; Burkill,

l.c. 2, 1655, 1935; Henderson, l.c. 382, 1954.

A tall grass of riverbanks.

114. Hymenachne pseudointerrupta C, Muell., Bot. Zeit. 79, 333,

1861; Bor, l.c. 313, 1960.

Panicum amplexicaule Henderson, l.c. 529, 1954.

Panicum myuros Ridl. non H.B.K., Mat. 3, 135, 1907.

Hymenachne myuros Ridl. non Beauv., Flor. Mal. 5, 230,

1925; Burkill, l.c. 1, 1214, 1935.

“Rumput kumpai M is a widespread grass of open swamps.

115. Ichnanthus vicinus (F. M. Bailey) Merr. Enum. Philip. 7, 70,

1923; Burkill, l.c. 2, 1221, 1935; Bor, l.c. 314, 1960.

Panicum vicinum F. M. Bailey, Syn. Queens. PI. Suppl. 3,

82. 1890 .

Ichnanthus pollens Munro non Sw. in Ridl. Mat. 3, 142,

1907; Flor. Mal. 5, 231, 1925.

A widespread forest grass.

116. Imperata cylindrica (Linn.) Beauv. Ess. Agrost. 165, 1812;

Burkill, l.c. 2, 1228, 1935; Henderson, l.c. 34, 1954;

Gilliland, Comm. Mal. Pits. 121, 1958; Bor, l.c. 169,

1960.

Lagurus cylindricus Linn. Syst. Nat. ed. 10, 878, 1759.

163

Gardens Bulletin, S.

Imperata arundinacea Cyrillo, PI. Rar. Neap, 2, 27, 1788;

Ridl. Mat. 3, 152, 1907; Flor. Mai. 5, 193, 1925.

Lalang occurs wherever Man misuses land. Its valuable

role in holding soil against erosion should not be over¬

looked however.

117. Imperata conferta (J. S. Presl.) Ohwi, Bot. Mag. Tokyo, 55.

549 , 1941; Bor, l.c. 169, 1960.

Saccharum confertum J.S. Presl. ex C.B. Presl. Rel. Haenk

1, 346, 1830.

Imperata exaltata Ridl. non Brogn. Mat. 3, 152, 1907;

Flor. Mai. 5, 193, 1925.

Imperata contractu Hitchc.; Burkill, l.c. 2, 1228. 1935;

Henderson, l.c. 343, 1954.

Larger than Lalang and not so common.

118. Isachne albens Trin. Sp. Gram. / t. 85, 1826; Ridl. Mat.

3, 128, 1907; Flor. Mai. J, 237, 1925; Burkill, l.c. 2,

1252, 1935; Henderson, l.c. 326, 1954; Jansen, l.c.

280, 1953; Bor, l.c. 579, 1960.

A grass of the mountains & hill stations.

119. Isachne conftisa Ohwi. Bull. Tokyo Sci., Mus. 18, 14. 1947;

Jansen, l.c. 282, 1953; Bor, l.c. 578, 1960.

Isachne rigida Nees, Ridl. Flor. Mai. 5, 238, 1925; Burkill.

l.c. 2, 1253, 1935; Henderson, l.c. 326, 1954.

Occurs in the northern States.

120. Isachne globosa (Thunb.) O. Ktze Rev. Gen. PI. 2, 778,

1891; Burkill, l.c. 2, 1252, 1935; Jansen, l.c. 282, 1953;

Bor, l.c. 580, 1960.

Milium globosum Thunb. Flor. Jap. 49, 1784.

Isachne australis R. Br. Prodr. FI. N. Holl. 196, 1810;

Ridl. Mat. 3, 129, 1907; Flor. Mai. 5, 239, 1925.

Widespread on the margins of swamps.

121. Isachne kinabaluensis Merr. J.S.B.R.A. Soc. 76, 1917; Jan¬

sen, l.c. 283, 1953.

Isachne javana Ridl. non Nees, Mat. 3, 128, 1907; Flor.

Mai. 5, 238, 1925.

A mountain species from Gunong Ledang in Johore

northwards.

122. Isachne kunthiana (Wight & Arn. ex Steud.) Miq. FI. lnd„

Bot. 3, 460, 1857; Jansen, l.c. 285, 1953; Bor, l.c. 581,

1960.

Panicum kunthianum Wight & Arn. ex Steud., Syn. PL

Glum. 1, 96, 1854.

Isachne semitalis Ridl. Flor. Mai. 5, 237, 1925.

From Singapore northwards.

164

Vol. XIX. (1962).

123. Isachne lankawiensis Jansen, Reinwardtia 2, 284, 1953.

From limestone in the Lankawi islands.

124. Isachne vniliacea Roth. Nov. Sp. PI. 58, 1821; Ridl. Mat. 3 r

129, 1907; Flor. Mai. 5, 239, 1925; Burkill, l.c. 2,

1253, 1935; Jansen, l.c. 285, 1953; Bor, l.c. 522, 1960.

From Singapore northwards.

125. Isachne saxicola Ridl. Flor. Mai. 5, 237, 1925; Jansen, l.c..

288, 1953.

Isachne javana var. saxicola Ridl. J. F.M.S. Mus. 6, 196,.

1915.

var. denticulata Ridl. Flor. Mai. 5, 237, 1925.

Recorded from Gunong Tahan.

126. Isachne sylvestris Ridl. J.S.B.R.A. Soc. 44, 206, 1905; Mat.

5, 129, 1907; Flor. Mai. 5, 238, 1925.

Isachne albens var. sylvestris (Ridl.) Jansen, l.c. 280, 1953 .

Rare, recorded from the Dindings.

127. Ischaemum apricum Ridl. Flor. Mai. 5, 203, 1925; Jansen,

l.c. 292, 1953.

Recorded from Penang.

128. Ischaemum fieldingianum Rendle, J. Bot. 22, 101, 1894;

Ridl. Mat. 3, 159, 1907; Flor. Mai. 5, 202, 1925; Jan¬

sen, l.c. 298, 1953.

Ischaemum commelynifolium Stapf ex Ridl. Flor. Mai. 5,

202, 1925.

Ischaemum maculatum Stapf ex Ridl. l.c. 202 , 1925.

Widespread from Johore swamps to Kedah Peak.

129. Ischaemum fluviatile Ridl. Flor. Mai. 5, 202, 1925.

Recorded from Kelantan.

130. Ischaemum goebelii Hack., Oest. Bot. Zeit. 51, 149, 1901;

Bor, l.c. 179, 1960.

Ischaemum imbricatum (Munro) Stapf ex Ridl. Flor. Mai.

5, 200, 1925.

Ischaemum barbatum var. imbricatum (Munro) Jansen,

l.c. 294, 1953.

Kedah & Perlis & recorded as a weed in Singapore.

131. Ischaemum indicum (Houtt.) Merr., J. Am. Arb. 19, 320,.

1938; Bor, l.c. 180, 1960.

Phleum indicum Houtt. Nat. Hist. 11, 13, 198, t.90 f.2,

1782.

Ischaemum ciliare Retz. Obs. Bot. 6, 36, 1791; Ridl. Mat.

3, 160, 1907.

165

Gardens Bulletin, S.

lschaemum aristatum Ridl. non Linn. Flor. Mai. 5, 203,

1925; Burkill, l.c. 2, 1253; Henderson, l.c. 351, 1954.

lschaemum barbatum Retz. Obs. Bot. 6, 35, 1791; Ohm,

Act. Phyt. Geobot. 11, 175, 1942; Jansen, l.c. 293, 1953.

Common, especially in the north.

132. lschaemum laeve Ridl. J.S.B.R.A. Soc. 14. 217, 1905; Mat.

3, 158, 1907; Flor. Mai. 5, 201, 1925.

Recorded from Singapore.

133. lschaemum macrunim Stapf ex Ridl. Flor. Mai. 5, 203,

1925; Jansen, l.c. 298, 1953.

Singapore and Negri Sembilan.

134. lschaemum magnum Rendle, J. Bot. 32, 102, 1894; Ridl.

Mat. 3, 158, 1907; Flor. Mai. 5, 200, 1925; Jansen,

l.c. 299, 1953; Bor, l.c. 182, 1960.

The locus classicus is Singapore and it belongs to the

south.

135. lschaemum inuticum Linn. Sp. PI. 1049, 1753; Ridl. Mat.

3, 159, 1907; Flor. Mai. 5, 201, 1925; Henderson, l.c.

350, 1954; Bor, l.c. 183, 1960.

Common throughout Malaya especially near the sea.

136. lschaemum polystachyum Presl. Rel. Haenk. 1, 328, 1830;

Jansen, l.c. 300, 1953.

lschaemum plumosulum Stapf ex Ridl. Flor. Mai. 5, 201,

1925.

Pollinia rupestris Ridl. J.S.B.R.A. Soc. 44, 208, 1905; Mat.

3, 156, 1907; Flor. Mai. 5, 199, 1925.

Recorded from Penang & Johore west coast.

137. lschaemum rugosunt Salisb. Ic. Stirp. Rar. 1, t. 1. 1791;

Ridl. Mat. 3, 157, 1907; Flor. Mai. 5, 200, 1925;

Henderson, l.c. 350, 1954; Bor, l.c. 184, 1960.

From Singapore northwards.

138. lschaemum trimorense Kunth. Rev. Gram. 369, t.98, 1829;

Ridl; Mat. 3, 160, 1907; Flor. Mai. 5, 203, 1925;

Burkill, l.c. 2, 1253, 1935; Jansen, l.c. 301, 1953;

Henderson, l.c. 351, 1954; Bor, l.c. 185, 1960.

Common in open places.

139. Leersia hexandra Swartz, Prod. Veg. Ind. Occ. 21, 1778,

Ridl. Mat. 3, 148, 1907; Flor. Mai. 5, 253, 1925;

Burkill, l.c. 2, 1327, 1935; Henderson, l.c. 320, 1954;

Bor, l.c. 599, 1960.

Rice-grass occurs in swamps everywhere.

166

Vol. XIX. (1962).

140. Leptaspis urceolata (Roxb.) R. Br., Benn. PI. Jav. Rar. 23*

t.6, 1838; Ridl. Mat. 3, 150, 1907; Flor. Mai. 5, 255,

1925; Burkill, l.c. 2, 1333, 1935; Jansen, l.c. 304,

1953; Henderson, l.c. 307, 1954; Bor, l.c. 617, 1960.

Phams urceolatus Roxb. FI. lnd. ed. 2, 3, 611, 1832.

A widespread forest grass.

141. Leptochloa chinensis (Linn.) Nees, Syll. Ratisb. /, 4, 1828;

Ridl. Mat. 3, 175, 1907; Flor. Mai. 5, 249, 1925;

Burkill. l.c. 2, 1333, 1935; Bor, l.c. 516, 1960.

Poa chinensis Linn. Sp. PI. 69, 1753.

A weed of cultivation throughout Malaya.

142. Leptochloa panicea (Retz.) Ohwi. Bot. Mag. Tokyo, 55, 311*

1941; Bor, l.c. 517, 1960.

Poa panicea Retz . Obs. Bot. 11, 1783.

Leptochloa polystachya Ridl. non Kunth, Flor. Mai. 5*

248, 1925; Burkill, l.c. 2, 1333, 1935.

A weed of rice fields in the north.

143. Lepturus repens (G. Forst.) R. Br. Prodr. FI. N. Holl. 207,

1810; Ridl. Mat. 3, 182, 1907; Flor. Mai. 5, 254, 1925;

Bor, l.c. 585, 1960.

Rottboellia repens G. Forst. Prodr. 9, 1786.

Recorded from the east coast.

144. Lophatherum gracile Brongn. in Duperr. Bot. Voy. Coq. 50,

t.8, 1831; Ridl. Mat. 3, 181, 1907; Flor. Mai. 5, 253,

1925; Burkill, l.c. 2, 1364, 1935; Bor, l.c. 460, 1960.

A forest grass common throughout.

145. Massia triseta (Nees) Balansa in Morot, J. Bot. Paris, 4, 165,.

1890, Bor, l.c. 479, 1960.

Eriachne triseta Nees ex Steud. Syn. PI. Gl. 237, 1854;

Ridl. Mat. 3, 172, 1907; Flor. Mai. 5, 240, 1925; Hen¬

derson, l.c. 316, 1954.

A wiry perennial from Malacca to Pahang and north.

146. Microstegium ciliatum (Trin.) A. Camus, Ann. Soc. Linn.

Lyon, 68, 201, 1921; Jansen, l.c. 305, 1953; Bor, l.c.

193, 1960.

Pollinia ciliata Trin. Mem. Acad. Petersb. ser. 6, 2, 306,

1833; Ridl. Mat. 3, 156, 1907.

P. ciliata var. glabrata Ridl. Flor. Mai. 5, 198, 1925.

Pollinia gracilis Ridl. Mat. 3, 156, 1907.

Forest margins from Selangor northwards.

147. Microstegium geniculatum (Hayata) Honda, Mon. Poac.

Japon. 410, 1930; Jansen, l.c. 306, 1953.

Pollinia geniculata Hayata, 1c. PI. Formos. 7, 73, 1918.

167

Gardens Bulletin, S.

Pollinia hendersonii C. E. Hubb., Kew Bull. 79, 1927.

Microstegium hendersonii C. E. Hubb. in Hutchins, Fam.

FI. PL 2, 227, 1934; Burkill, Lc . 2, 1786, 1935.

A rare mountain species.

148. Microstegium nudum (Trin.) A. Camus Ann. Soc. Linn.

Lyon, 68, 201, 1921; Bor, l.c. 194, 1960.

Once collected from Fraser’s Hill, Pahang.

149. Microstegium vagans (Nees ex Steud.) A. Camus l.c. 200,

1921; Bor, l.c. 195, 1960.

Pollinia vagans Nees ex Steud. Syn. PL Glum. 1, 410,

1855.

Pollinia grata (Nees et Meyen) Hack. Monog. Phan. 6,

175, 1889; Ridl. Mat. 3, 156, 1907; Flor. Mai. 5, 198,

1925; Burkill, Lc. 2, 1786, 1935.

Ephebopogon gratus Nees et Meyen ex Steud. Nom. ed.

2, 1, 556, 1840.

Recorded from Penang and to the north.

150. Miscanthus sinensis Anderss. Oefvers. Kon. Vet-Akad. Forh.

165, 1855.

Miscanthus japonicus Ridl. non Anderss., Flor. Mai. 5,

194, 1925; Burkill, Lc. 2, 1479, 1935.

Recorded only from Perak.

15L Mnesithea cancellata Ridl. Flor. Mai. 5, 206, 1925.

Rottboellia cancellata Ridl. J.S.B.R.A. Soc. 59, 288, 1920.

Rare in the far north of Malaya.

152. Mnesithea geminata (Hack.) Ridl. Mat. 3, 163, 1907; Flor.

Mai. 5, 206, 1925; Jansen, l.c. 309, 1953.

Rottboellia geminata Hack. Oesterr. Bot. Zeitschr. 41, 48,

1891.

Sandy places on the east coast.

153. Mnesithea mollicoma (Hance) A. Camus, Bull. Mus. Hist.

Nat. Paris, 25, 57, 1919; Jansen, l.c. 308, 1953.

Rottboellia mollicoma Hance, J. Bot. 9, 134, 1871.

Mnesithea pubescens Ridl. J.S.B.R.A. Soc. 44, 207, 1905;

Mat. 3, 163, 1907; Flor. Mai. 5, 206, 1925; Burkill, Lc.

2, 1484, 1935.

Recorded from Johore.

154. Myriostachya wightiana (Nees) Hk. f. Flor. Brit. Ind. 7, 327,

1897; Burkill, l.c. 2, 1521, 1935; Jansen, l.c. 310,

1953; Bor, l.c. 518, 1960.

Leptochloa wightiana Nees ex Steud. Syn. PL Glum. 1,

209, 1854.

168

Vol. XIX. (1962).

Myriostachya wightiana var. longispicula Hk. f. l.c. 328,

1897; Ridl. Mat. 3, 180, 1907; Flor. Mai. 5, 245, 1925.

A grass of estuarine muds from Penang northwards.

155. Neyraudia rcynaudiana (Kunth) Keng ex Hitchc. Am. J. Bot.

21, 131, 1934; Bor, l.c. 518, 1960.

Arundo reynaudiana Kunth. Rev. Gram. 2, 275, t49, 1830.

Neyraudia madagascariensis (Kunth.) Hk. /., Ridl. Mat.

3, 176, 1907.

Triraphis madagascariensis Stapf ex Ridl. Flor. Mai. 5,

251, 1925; Burkill, l.c. 2, 2186, 1935; Henderson, l.c.

310, 1954.

A reed recorded as far south as Seremban & Malacca.

156. Oplismenus burmannii (Retz.) Beauv. Ess. Agrost. 54, 168,

169, 1812; Ridl. Mat. 3, 145, 1907; Flor. Mai. 5, 221,

1925; Burkill, l.c. 2, 1585, 1935; Jansen, l.c. 312, 1953;

Henderson, l.c. 336, 1954; Bor, l.c. 317, 1960.

Recorded from Penang and Kedah.

157. Optismcnus compositus (Finn.) Beauv. Ess. Agrost. 54, 168,

169, 1812; Ridl. Mat. 3, 145, 1907; Flor. Mai. J, 221,

1925; Burkill, l.c. 2, 1586, 1935; Jansen, l.c. 311,

1953; Henderson, l.c. 335, 1954; Bor, l.c. 318, 1960.

Panicum compositum Linn. Sp. PI. 57, 1753.

A grass of dry rocky spots in shade, widespread in

Malaya.

158. Oryza granulata Nees et Arn. ex Hk. f. Flor. Brit. Ind. 7,

93, 1896; Bor, l.c. 604, 1960.

Recorded from a river bank in northern Perak.

159. Oryza minuta Presl. Rel. Haenk. /, 208, 1830; Burkill, l.c.

2, 1593, 1935; Henderson, l.c. 321, 1954; Bor, l.c.

605, 1960.

Oryza latifolia Ridl. non Desv. Mat. 3, 148, 1907; Flor.

Mai. 5, 252, 1925.

“Padi burong” or wild rice.

160. Oryza rufipogon Griff. Notul. 3, 5, 1851; Bor, l.c. 605, 1960.

Oryza fatua var. longe-aristata Ridl. Flor. Mai. 5, 1925.

Oryza fatua of Burkill, l.c. 2, 1593, 1935.

Also “Padi burong.”

161. Oryza ridleyi Hk. f. Flor. Brit. ind. 7, 93, 1897; Ridl. Mat.

3, 148, 1907; Flor. Mai. 5, 252, 1925; Burkill, l.c. 2,

1593, 1935; Jansen, l.c. 312, 1953; Henderson, l.c.

321, 1954; Bor, l.c. 605, 1960.

On riverbanks throughout Malaya.

169

Gardens Bulletin, S.

162. Ottochloa nodosa (Kunth) Dandy, J. Bot. 69, 55, 1931; Bor,

l.c. 318, 1960.

Panicum nodosum Kunth, Rev. Gram. 1, 97, 1833; Hen¬

derson, l.c. 332, 1954.

Hemigymnia multinodis Stapf, Ridl. Flor. Mai. 5, 228,

1925; Bur kill, 1, 1138, 1935.

Hemigymnia fusca Ridl. Flor. Mai. 5, 228, 1925; Burkill,

l.c.

A perennial grass of swamps.

163. Panicum austro-asiaticum Ohwi, Act. Phytotax. Geobot. 2,

45, 1942; Jansen, l.c. 314, 1953; Bor, l.c. 324, 1960.

Panicum humile Nees ex Steud. non Thunb., in Syn. PI.

Glum. 84, 1854; Ridl. Mat. 3, 135, 1907; Flor. Mai. 5,

224, 1925; Burkill, l.c. 2, 1655, 1935.

A grass of northern Malaya.

164. Panicum brevifolium Linn. Sp. PI. 59, 1753; Burkill, l.c. 2,

1655, 1935; Jansen, l.c. 315, 1953; Henderson, l.c. 329,

1954, Bor, l.c. 324, 1960.

Panicum ovalifolium Poir in Lamk. Encyl. Bot. Suppl. 4,

279, 1816; Ridl. Mat. 3, 141, 1907; Flor. Mai. 5, 227,

1925.

Common throughout Malaya.

var. hirtifolium (Ridl.) Jansen, l.c. 315, 1953.

Panicum hirtifolium Ridl. Mat. 3, 141, 1907; Flor. Mai. 5,

228, 1925.

Rare in Selangor.

165. Panicum cambogiense Balansa in Morot, J. de Bot. 4, 142,

1890; Bor, l.c. 325, 1960.

Panicum luzonense Ridl. non Presl. Mat. 3, 136, 1907.

Panicum caesium Ridl. non Nees, Flor. Mai. 5, 225, 1925.

Panicum tuberculatum Burkill non Presl. l.c. 2, 1658,

1935.

Widespread in Malaya.

166. Panicum elegantissimum Hk. f. Flor. Brit. Ind. 7, 52, 1897;

Ridl. Mat. 3, 139, 1907; Flor. Mai. 5, 226, 1925; Bor,’

l.c. 325, 1960.

A rare grass from Perak.

167. Panicum maximum Jacq. Coll. Bot. 1, 76, 1786; Ridl Mat 3

140, 1907; Flor. Mai. 5, 1925; Burkill, l.c. 2, 1656,

1935; Bor, l.c. 327, 1960.

Guinea grass is widely cultivated as fodder.

168. Panicum montanum Roxb. FI. Ind. 1, 313, 1820; Ridl. Mat.

3, 139, 1907; Flor. Mai. 5, 227, 1925; Burkill, l.c. 2,

1657, 1935; Jansen, l.c. 306, 1953; Bor, l.c. 329, 1960.

Recorded from Selangor and northwards.

170

Vol. XIX. (1962).

169. Panicum paludosum Roxb. Flor. Ind. /, 310, 1820; Bor, l.c.

329, 1960.

Panicum prolijerum Ridl. non Lam., Mat. 3, 139, 1907;

Flor. Mai. 5, 225, 1925; Burkill, l.c. 2, 1657, 1935.

A floating grass of pools & swamps, rare.

170. Panicum perakense (Hk. f.) Merr. Philip. J. Sci. 7/C, 52,

1916; Ridl. Flor. Mai. 5, 226, 1925; Jansen, l.c. 317,

1953.

Panicum humidorum var.? perakense Hk. f. FI. Br. Ind.

7, 54, 1897; Ridl. Mat. 3, 137, 1907.

A rare grass recorded from Johore northwards.

171. Panicum repens Linn. Sp. PI. ed. 2, 87, 1762; Ridl. Mat 3,

139, 1907; Flor. Mai. 5, 225, 1925; Burkill, l.c. 2,

1657, 1935; Henderson, l.c. 329, 1954; Bor. l.c. 330,

1960.

Widespread in Malaya.

172. Panicum sarmentosum Roxb. FI. Ind. 7, 308, 1820; Ridl.

Mat. 5, 140, 1907; Flor. Mai. 5, 227, 1925; Henderson,

l.c. 332, 1954; Bor, l.c. 330, 1960.

“Rumput jangut Ali” is widespread throughout Malaya.

173. Paspalidium punctatum (Burm.) A. Camus, FI. Gen. Indo-

chin. 7, 419, 1922; Ridl. Flor. Mai. 5, 218, 1925;

Burkill, l.c. 2,1672, 1935; Jansen, l.c. 318, 1953; Bor,

l.c. 333, 1960.

Panicum punctatum Burm. FI. Ind. 26, 1768; Ridl. Mat. 3,

132, 1907.

Rare, recorded from Johore.

174. Paspalum cartilagineum Presl. Rel. Haenk. 7, 216, 1830;

Jansen, l.c. 320, 1953; Bor, l.c. 335, 1960.

Recorded from Pahang.

175. Paspalum commersonii Lam. Ill. 7, 175, t43, 1791; Ridl.

Flor. Mai. 5, 218, 1925; Jansen, l.c. 319, 1953; Bor,

l.c. 335, 1960.

Paspalum scrobiculatum Ridl. non Linn. Mat. 3, 213, 1907;

Burkill, l.c. 2, 1674, 1935; Henderson, l.c. 340, 1954.

Paspalum orbiculare Ridl. non Forst., Flor. Mai. 5, 217,

1925.

P. commersonii var. hirsutum Jansen, l.c. 319, 1953.

A weed widespread in Malaya.

176. Paspalum conjugatum Berg. Act. Helvet. Phys. Math. 7, 129,

t8, 1762; Ridl. Mat. 5, 124, 1907; Flor. Mai. 5, 218,

1925; Burkill, l.c. 2, 1673, 1935; Henderson, l.c. 339,

1954; Bor, l.c. 336, 1960.

“Rumput kerbau” is widespread in Malaya.

171

Gardens Bulletin, S.

177. Paspalum longifolium Roxb. Flor. Ind. /, 280, 1820; Ridl.

Flor. Mai. 5, 217, 1925; Bor, l.c. 339, 1960.

Paspalum platycoleum Ridl. Flor. Mai. 5, 217, 1925.

Widespread in Malaya.

178. Paspalum vaginatum Sw. Prodr. FI. Ind. Occ. 21, 1788;

Burkill, l.c. 2, 1675, 1935; (ef. Jansen l.c. 321, 1953);

Henderson, l.c. 339, 1954; Bor, l.c. 341, 1960.

Paspalum distichum Ridl. non Linn. Mat. 3, 124, 1907;

Mai. 5, 218, 1925.

A grass of saline mud around the coasts.

179. Pcnnisetum purpureum Schum. Beskr. Guin. PI. 44, 1827;

Burkill, l.c. 2, 1688, 1935; Henderson, l.c. 325, 1954;

Bor, l.c. 348, 1960.

“Napier fodder” is well established in many places in

Malaya.

180. Pennisetum clandestinum Hochst. ex Chiov. Ann. 1st. Bot.

Roma 8, 41, t5 f2, 1903; Burkill, l.c. 2, 1687, 1935;

Henderson, l.c. 325, 1954; Bor, l.c. 344, 1960.

Kikuyu grass is now established on hill stations in Malaya.

181. Perotis indica (Linn.) O. Ktze, Rev. Gen. PI. 787, 1891;

Burkill, l.c. 2, 1696, 1935; Henderson, l.c. 319, 1954;

Bor, l.c. 611, 1960.

Anthoxanthum indicum Linn. Sp. Pi. 28, 1753.

Perotis latifolia Ait. Hort. Kew 1, 85, 1789; Ridl. Mat. 3,

149, 1907; Flor. Mai. 5, 243, 1925.

“Rumput ekor kuching” is a common grass of maritime

sandy areas.

182. Phragmites karka (Retz.) Trin. ex Steud. Nom. Bot. cd. 2,

2, 324, 1841; Ridl. Mat. 3, 175, 1907; Bor, l.c. 416,

1960.

Arundo karka Retz. Obs. Bot. 4, 21 , 1786.

Phragmites communis Ridl. non Trin. Flor. Mai 5, 240,

1925; Burkill, l.c. 2, 1715, 1935, Henderson, l.c. 309,

1954.

“Tebu salah” is the common reed of rivers & tin tailings.

183. Poa annua Linn. Sp. PI. 68, 1753, Ridl. Flor. Mai. 5, 254,

1925; Jansen, l.c. 322, 1953; Henderson, l.c. 311,

1954; Bor, l.c. 555, 1960.

A European weed now established at hill stations.

184. Pogonatherum paniceum (Lam.) Hack. Allg. Bot. Zeit. 12,

178, 1906; Jansen, l.c. 333, 1953; Bor, l.c. 202, 1960.

Saccharum paniceum Lamk. Encycl. Meth. Bot. 1, 595,.

1785.

Ill

Vol. XIX. (1962).

Pogonatherum saccharoideum var. monandra Hack.

Monog. Androp. 193, 1889; Ridl. Mat. 3, 154, 1907;

Flor. Mai 5, 195, 1925; Henderson, l.c. 348, 1954.

Pogonatherum crinitum Trin. oj Bur kill, l.c. 2, 1780, 1935.

From Johore northwards “Bamboo grass” occurs in

mountain streams.

185. Polytrias amaura (Buse) O. Ktze. Rev. Gen. PI. 788, 1891;

Henderson, l.c. 348, 1954; Bor, l.c. 202, 1960.

Andropogon amaurus Buse in Miq. PI. Junghn. 360, 1854 .

Pollinia praemorsa (Nees) Hack. Monog. Androp, 189,

1889; Ridl. Mat. 3, 164, 1907.

Eulalia praemorsa (Nees) Stapf ex Ridl. Flor. Mai. 5, 197,

1925; Burkill, l.c. 1, 975, 1935.

Recorded from Singapore, Malacca, Kuala Lumpur and

Penang.

186. Pseudechinolaena polystachya (H.B.K.) Stapf. Flor. Trop.

Agr. 9, 495, 1919; Ridl. Flor. Mai. 5, 221, 1925; Burkill,

l.c. 2, 1812, 1935; Bor, l.c. 352, 1960.

Echinolaena polystachya H.B.K. Nov. Gen. & Sp. 1, 119,.

1816.

Rare in the mountains in Northern Malaya.

187. Pseudoraphis spinescens (R. Br.) Vickery, Proc. R. S.

Queensl. 62, 7, 69, 1952; Bor; l.c. 353, 1960.

Panicum spinescens R.Br. Prodr. 193, 1810.

Andropogon squarrosus Linn. f. Sup pi. 433, 1781; Ridl.

Mat. 3, 168, 1907.

Recorded from the Expt. Station, Telok Chengai, Kedah.

188. Racemobambos setifera Holttum, Gard. Bull. Sing. 15, 27 U

1956; l.c. 16, 134, 1958.

A rare scrambling bamboo of Johore.

189. Rhynchelytrum repcns (Willd.) C. E. Hubb. Kew Bull. 110„

1934; Bor, l.c. 355, 1960.

Tricholaena rosea Nees lnd. sem. Hort. Vratisl. 1835; Ridl.

Flor. Mai. 5, 235, 1925; Burkill, l.c. 2, 2177, 1935; Hen¬

derson, l.c. 325, 1954.

“Red-top” is now well established in many parts of Malaya.

173

Gardens Bulletin, S.

190. Saccharum arundinaceum Retz. Obs. Bot. 4 , 14, 1786; Ridl.

Mat. 3 , 153, 1907; Flor. Mai. 5 , 193, 1925; Burkill,

l.c. 2, 1924, 1935; Henderson, l.c. 344, 1954; Bor, l.c.

211, 1960.

“Teberau” is a wild sugarcane of sandy river banks in

Malaya.

191. Saccharum spontaneum Linn. Mant. 2, 183, 1771; Hender¬

son, l.c. 344, 1954; Bor, l.c. 214, 1960.

The reputed wild parent of sugar cane.

Burkill, l.c. 2, 1926, 1935, discusses the hybridisation of

S. spontaneum with the sugarcane S. officinarum Linn.

192. Sacciolepis indica (Linn.) Chase, Proc. Biol. Soc. Wash. 21,

8, 1908; Ridl. Flor. Mai. 5, 232; 1925; Burkill, l.c. 2,

1940, 1935; Henderson, l.c. 334, 1954; Bor, l.c. 357,

1960.

Air a indica Linn. Sp. PI. 1753 in “errata”.

Panicum indicum Linn, non Mill., Mant. PI. 2, 184, 1771;

Ridl. Mat. 3, 134, 1907.

Sacciolepis angusta Ridl. non Stapf, Flor. Mai. 5, 232,

1925; Burkill, l.c. 2, 1940, 1935.

Sacciolepis turgida Ridl. Flor. Mai. 5, 231, 1925; Burkill,

l.c. 2, 1941, 1935.

Widespread in Malaya.

193. Sacciolepis interrupta (Willd.) Stapf, Flor. Trop. Afr. 9, 757,

1920; Bor, l.c. 358, 1960.

Panicum interruptum Willd. Sp. PI. 1, 341, 1797.

Lankawi islands in padi fields.

194. Sacciolepis myosuroides (R. Br.) A. Camus FI. Gen. Indo-

chin. 7, 460, 1922; Ridl. Flor. Mai. 5, 232, 1925; Hen¬

derson, l.c. 335, 1954; Bor, l.c. 358, 1960.

Panicum myosuroides R.Br. Prodr. FI. N. FI oil. 189, 1810;

Ridl. Mat. 3, 134, 1907.

Throughout Malaya in swampy sites.

195. Schizachyrium brevifolium (Sw.) Nees. Agrost. Bras. 332,

1829; Ridl. Flor. Mai. 5, 210, 1925; Burkill, l.c. 2,

1975, 1935; Jansen, l.c. 336, 1953; Bor, l.c. 215, 1960.

Andropogon brevifolius Sw. Prodr. Veg. Ind. Occ. 26 ,

1788 ; Ridl. Mat. 3 , 165, 1907 ; Henderson, l.c. 346. 1954.

Sporadic north of Johore.

174

Vol. XIX . (1962).

196. Schizachyrium sanguineum (Retz.) Alst. Suppl. FI. Ceyl. 334,

1931; Bor, l.c. 216, 1960.

Rottboellia sanguined Retz . Obs. Bot. 3, 25, 1783.

Schizachyrium semiberbe Nees Agrost. Bras. 336, 1829;

Ridl. Flor. Mai. 5, 210, 1925; Burkill, l.c. 2, 1975, 1935.

Rare in the south.

197. Schizostachyum aciculare Gamble, Ann. Bot. Gard. Calc.

7, 117, pi. 104, 1896; Ridl. Mat. 3, 196, 1907; Flor.

Mai. 5 , 270, 1925; Burkill, l.c. 2, 1976, 1935; Holttum

Gard. Bull. Sing. 16, 39, 1958.

“Buioh padi” is known from Malacca, Negri Sembilan &

Selangor.

198. Schizostachyum brachycladum Kurz. J. As. Soc. Bengal. 39,

89 pi. 6, f.2, 1870; Burkill, l.c. 2, 1977, 1935; Holttum,

l.c. 45, 1958; Gilliland, Comm. Mai. Pits. 194, 1958.

Schizostachyum chilianthum Ridl. non Kurz . p.p., Mat. 3,

194, 1907; Flor. Mai. 5, 269, 1925; Burkill, l.c. 2, 1977,

1935.

“Buioh nipis” is widely cultivated in Malaya.

var. auriculatum Holttum, l.c. 47, 1958.

Cultivated in Singapore.

199. Schizostachyum gracile (Munro) Holttum, Kew Bull. 206,

1956; l.c. 37, 1958.

Melocanna gracilis Munro, Trans. Linn. Soc. 26, 133,

1868.

Schizostachyum tenue Gamble, Ann. Bot. Gard. Calc. 7,

114, pi. 100, 1896; Ridl. Mat. 3, 194, 1907; Flor. Mai. 5,

268, 1925; Burkill, l.c. 2, 1977, 1935.

Schizostachyum chilianthum Ridl. non Kurz. p.p.

“Buioh rappen” is wild & cultivated in Malaya.

var. erectum Holttum, l.c. 38, 1958.

Malacca and Negri Sembilan.

200. Schizostachyum grande Ridl. J.S.B.R.A. Soc. 82, 204, 1920;

Flor. Mai. 5, 271, 1925; Burkill, l.c. 2, 1977, 1935;

Holttum, l.c. 49, 1958.

“Buioh semeliang” is common in the foothills of the Main

Range.

175

Gardens Bulletin, S.

201. Schizostachyum jaculans Holttum Kew Bull. 494, 1953;

Gard. Bull. Sing. 16, 40, 1958.

Schizostachyum blumei Gamble non Nees, Ann. Bot. Gard.

Calc. 7, 116, pi 112, 1896; Ridl. Mat . 3, 195, 1907;

Flor. Mai 5, 272, 1925; Burkill, l.c. 2, 1976, 1935.

“Buloh sumpitan” is both wild and cultivated in Malaya.

202. Schizostachyum longispiculatum Kurz, J. As. Soc. Bengal,

39, 89, pi. 6, f.l, 1870; Ridl. Flor. Mai. 5, 270, 1925;

Burkill, l.c. 2, 1977, 1935; Holttum; l.c. 48, 1958.

Schizostachyum latifolium Gamble, Ann. R. Bot. Gard.

Calc. 7, 117, 1896; Ridl. Mat. 3, 196, 1907; Flor. Mai.

5, 270, 1925; Burkill, l.c. 2, 1977, 1935.

Ochlandra ridleyi Gamble, l.c. 127, 1896; Ridl. Mat. 3,

197, 1907; Flor. Mai. 5, 197, 1925; Burkill, l.c. 2, 1569,

1935.

Schizostachyum ridleyi (Gamble) Holttum, Gard. Bull.

Sing. 11, 296, 1947.

This bamboo is widespread in Malaya at the edge of forest

on river banks.

203. Schizostachyum terminale Holttum Gard. Bull. Sing. 15, 274,

1956; 16, 51, 1958.

Rare known only from one locality in Kedah.

204. Schizostachyum zollingeri Steud. Syn. PI. Glum. 332, 1854;

Ridl. Mat. 3, 195, 1907; Flor. Mai. 5, 269, 1925;

Burkill, l.c. 2, 1978, 1935; Holttum, l.c. 42, 1958.

Schizostachyum chilianthum Gamble non Buse p.p. Ann.

Bot. Gard. Calc. 7, 115, pi. 101, 1896; Ridl. Mat. 3,

194, 1907; Flor. Mai. 5, 269, 1925; Burkill, l.c. 2, 1977,

1935.

Melocanna zollingeri (Steud.) Munro, Trans. Linn. Soc.

26, 134, 1868.

“Buloh telor” is common north from Selangor.

205. Sphaerocarvum malaccense (Trin.) Pilg., Fedde Rep. Sp.

Nov. 45, 2, 1938; Henderson, l.c. 317, 1956; Bor, l.c.

583, 1960.

Panicum malaccense Trin. Gram. Panic. 204, 1826.

Sphaerocaryum elegans Nees ex Steud.; Ridl. Mat. 3, 170,

1907; Flor. Mai. 5, 241, 1925.

A forest path grass from the lowlands of Malaya.

176

Voi XIX. (1962).

206. Sderostachya ridleyi (Hack.) A. Camus, Ann. Soc. Linn.

Lyon. N.S. 71, 104, 1915; Burkill, l.c. 2, 1983, 1935.

Saccharwn ridleyi Hack. Oestr. Bot. Zeitschr., 41, 6, 1891;

Ridl. Mat. 3, 153, 1907.

Sclerostachyum ridleyi Stapf mss. of Ridl. Flor. Mai. 5,

194, 1925.

A local river reed in Pahang.

207. Setaria italica (Linn.) Beauv. Ess. Agrost. 51, 170, 178,

1812; Ridl. Mat. 3, 144, 1907; Flor. Mai. 5, 234, 1925;

Burkill, l.c. 2, 1999, 1935; Henderson, l.c. 323, 1954;

Bor, l.c. 362, 1960.

Panicurn italicum Linn. Sp. Pi 56, 1753.

“Sekoi” is cultivated by nomadic aboriginal tribes.

208. Setaria pallide-fusca (Schum.) Stapf & Hubb. Kew Bull. 259,

1930; Jansen, l.c. 339, 1953; Bor, l.c. 363, 1960.

Panicum pallide-fuscum Schum. Beskr. Guin. Pi 58, 1827.

Setaria glauca Hack., Ridl. J.S.B.R.A. Soc. 23, 23, 1891;

Mat. 3, 144, 1907.

Setaria rubiginosa (Stead.) Miq., Ridl. Flor. Mai 5, 234,

1925.

Setaria geniculata Burkill, non Beauv. ic. 2, 1999, 1935;

Henderson, ic. 323, 1956.

Common weed of waste ground.

209. Setaria palmifolia (Willd.) Stapf, J. Linn. Soc. Bot. 42, 186,

1914; Burkill, l.c. 2, 1999, 1935; Henderson, l.c. 324,

1954; Bor, l.c. 363, 1960.

Panicurn palmifolium Willd. ex Poir. in Lam. Encycl.

Suppi 4, 282, 1816.

Setaria plicata Cooke, Ridl. Flor. Mai 5, 235, 1925.

Widespread from N. Johore and northwards.

210. Sorghum propinquum (Kunth.) Hitchc. Lingn. Sci. J. 7, 249,

1931; Bor, l.c. 223, 1960.

Sorghum affine (Presl.) A. Camus. “Stapf mss. ,f of Ridl.

Flor. Mai 5, 195, 1925; Burkill, ic. 2, 2055,1935.

Riverbanks in the north, a reedlike grass.

211. Spinifex littoreus (Burm.) Merr. Philip. J. Sci. 7C, 229, 1912;

Burkill, l.c. 2, 2066, 1935; Henderson, l.c. 323, 1954;

Bor, l.c. 366, 1960.

Stipa littorea Burm . f. Fi Ind. 29, 1768.

Ill

Gardens Bulletin, S.

Spinifex squarrosus Linn. Mant. 2, 163, 1771; Ridl. Mat.

3, 147, 1907; Flor. Mai 5, 236, 1925.

Sandy seashores of the east coast of Malaya.

212. Spodiopogon velutinus (Jansen) Holttum Gard. Bull. Sing.

11, 297, 1947.

Recorded only from Cameron Highlands.

213. Sporobolus diander (Retz.) Beauv. Ess. Agrost. 26, 147,

1812; Ridl. Mat. 3, 170, 1907; Flor. Mai. 5, 244, 1925;

Burkill, l.c. 2, 2069, 1935; Henderson, l.c. 318, 1954;

Bor, l.c. 629, I960.

Agrostis diandra Retz. Obs. Bot. 5, 19, 1789.

A weed of open sunny dry places in Malaya.

214. Sporobolus indicus auctt. non (Linn.) R.Br. ef Bor, l.c. 630,

1960.

Sporobolus indicus Ridl. Mat. 3, 171, 1907; Flor. Mai. 5,

244, 1925; Henderson, l.c. 318, 1954.

Sporobolus berteroanus Hitchc. & Chase; Burkill, l.c. 2,

2069, 1935.

A widespread weed of drier sunnier places in Malaya.

215. Sporobolus piliferus (Trin.) Kunth, Enum. PI. /, 211, 1833;

Ridl. Mat. 3 , 171, 1907; Flor. Mai. 5, 244, 1925; Bor,

l.c. 632, 1960.

Once recorded from Malacca.

216. Sporobolus virginicus (Linn.) Kunth. Rev. Gram. /, 67,

1829; Bor, l.c. 634, 1960.

Agrostis virginica Linn. Sp. PI. 63, 1753.

Recorded from Singapore and Lankawi islands.

217. Stenotaphrum helfcri Munro ex Hk. f. Flor. Brit. Ind. 7, 91,

1897; Ridl. Mat. 3 , 146, 1907; Flor. Mai. 5, 220, 1925;

Burkill, l.c. 2, 2075, 1935; Henderson, l.c. 337, 1954;

Bor, l.c. 368, 1960.

Usually associated with limestone hills in the northern

States.

218. Themeda arguens (Linn.) Hack, in D.C. Monog. Phan. 6 ,

657, 1889; Ridl. Flor. Mai. 5, 212, 1925; Burkill, l.c. 2,

2146, 1935; Henderson, l.c. 351, 1954; Bor, l.c. 250,

1960.

Stipa arguens Linn. Sp. PI. ed. 2, 117, 1762.

Anthistiria arguens (Linn.) Willd. Sp. PI. 4, 901, 1806 ;

Ridl. Mat. 3, 168, 1907.

“Rumput misai adam” is recorded throughout Malaya.

178

Vol. XIX. (1962).

219. Themeda arundinacea (Roxb.) Ridl. Trans. Linn. Soc. J,

401, 1893; Bor, l.c. 250, 1960.

Anthistiria arundinacea Roxb. FI. Ind. I, 256, 1820.

Anthistiria gigantea var. arundinacea Hack, in DC Monog.

Phan. 6, 674, 1889; Ridl. Mat. 3, 169, 1907.

(Themeda subsericans Ridl. Flor. Mai. 5, 212, 1925 is as¬

sumed to be a hybrid between Th. arundinacea X Th ,

villosa cf. Bor. Ic. 252.

Rare recorded from Pahang.

220. Themeda villosa (Poir.) A. Camus in Lecomte FI. Gen. Indo-

Chine 9, 364, 1922; Ridl. Flor. Mai. 5, 212, 1925;

Burkill, l.c. 2, 2147, 1935; Henderson, l.c. 352, 1954;

Bor, l.c. 252, 1960.

“Rumput riong” is widespread in Malaya.

221. Thuarea involuta (Forst.) R. Br. ex R.&.S. Syst. Veg. 2, 808,

1817; Bor, l.c. 368, I960.

Ischaemum involutum G.Forst. FI. Ins. Austr. Prodr. 73,

1786.

Thuarea sarmentosa Pers. Syn. PI. 1, 110, 1805; Ridl. Mat .

3, 146, 1907; Flor. Mai. 5, 236, 1925; Burkill, l.c. 2, 2157,

1935; Henderson, l.c. 335, 1954.

On sandy coasts throughout just above high tide level.

222. Thyrsostachys siamensis Gamble, Ann. R. Bot. Gard. Calc.

7, 59, pi. 51, 1896; Holttum, l.c. 80, 1958.

Cultivated bamboo of Singapore & Penang.

223. Thysanolaena maxima (Roxb.) O. Ktze. Rev. Gen. PI. 794,

1891; Burkill, l.c. 2. 2160, 1935; Henderson, l.c. 321,

1954; Bor, l.c. 650, 1960.

Agrostis maxima Roxb. FI. Ind. 1, 319, 1820.

Thysanolaena agrostis (Roxb.) Nees, Edin. New Phil. J.

18, 180, 1835; Ridl. Mat. 3, 143, 1907; Flor. Mai. 5,

241, 1925.

“Buloh teberau” is also the name for this hillside reed from

Negri Sembilan northwards.

224. Vetiveria zizanioides (Linn.) Nash in Small, FI. Southeast

U.S. 67, 1903; Stapf, Kew Bull. 346, 1906; Burkill, l.c.

2, 2228, 1935; Henderson, l.c. 446, 1954; Bor, l.c. 258,

1960.

Phalaris zizanioides Linn. Mant. 2, 183, 1771.

Vetiveria odorata Virey, J. Pharm. 13, 501, 1827; Ridl.

Flor. Mai. 5, 208, 1925.

The “Kuskus” or “Vetiver” cultivated in gardens.

179

Gardens Bulletin, S

225. Zoysia matrclla (Linn.) Merr., Philip. J. Sci. 7C, 230, 1912,

Burkill, l.c. 2, 2308, 1935; Henderson, l.c. 318, 1954;

Gilliland, Comm. Mai. Pits. 222, 1958; Bor, l.c. 684,

1960.

Agrostis rnatrella Linn. Mant. 2, 185, 1771.

Zoysia pungens Willd. Ges. Nat. Fr. Berlin. N.S. 3, 441,

1801; Ridl. Mat. 3, 149, 1907; Flor. Mai. 5, 243, 1925.

Common as a maritime grass on sandy places.

The lawn Zoysia is a different species imported from the

Philippines.

180

The Species of Brackenridgea in the

Singapore Herbarium

By C. X. Furtado

Singapore

While comparing some recent collections, I noticed a great deal

of confusion in the Singapore herbarium in the determinations of

the specimens belonging to Brackenridgea. In Ridley’s Flora of

the Malay Peninsula 1 (1922), this material is referred to Gomphia

hookeri and G. corymbosa, but since the main distinction is made

on the colour of the flowers, whether they are red or white, many

errors have crept in the naming of the species. Further G. hookeri

was established on a Penang specimen which had a red fruiting

calyx. Some authors mistook this to mean that the species pro¬

duced red flowers. Actually the flowers are white, the red being

developed in the pedicel, calyx and disc after the flowers have

been fertilised and the petals shed. As a result all our recent mate¬

rial from Penang has been referred to G. corymbosa —a white

flowered shrub which does not seem to develop the red pigment

in the flowers after their being fertilised. Ridley overlooked the

important paper by van Tieghem (Ann. Sci. Nat. VIII, 1902 pp.

393-399) where three new species were added to the genus

Brackenridgea, besides the two mentioned by Ridley under Gom¬

phia. Two of these, B. rubescens and B. kingii are reduced to B.

palustris , described originally from Borneo; the third B. perak-

ensis has been reduced here as a synonym of B. hookeri. In addi¬

tion Singapore has been credited with a new species, B. denticulata.

I mention here two extra Malayan species, one, B. serrulata, be¬

cause it has leaves with spiny margins as in B. denticulata, and

second, B. foxworthyi (Elm.) comb. nov. because it is an addition

to the genus; it has also spinulose leaf margins.

Sections

Van Tieghem (l.c.) divided the genus into two sections: Capi -

tatae and Spiciformes. In the former the inflorescence is a com¬

pound globose head, consisting of an abbreviated axis bearing

short branches that produce flower-bearing stalks. The scales are

arranged spirally on the main axis and distichously on the lateral

branches. It is very difficult to detect the branches when the flowers

are young, though they become obvious in the fruiting stage and

later. The axis in the second section may be short or long, but is

always simple and bears its scales usually distichously. Sometimes

the older parts of the axis tend to produce branches bearing a

separate set of flowers, so that this branching cannot be confused

with the branching in the first section where all the flowers appear

to form one head, the main axis itself rarely producing flowers.

181

Gardens Bulletin , 5.

The Species

A. Sectio Capitatae van Tiegh. op. cit. (1902) 397.

1. Brackenridgea hookeri (PI.) A. Gray, New Gen. PI. (1853) 6

(fide cit. seq.); C. Muell, in Walp., Ann. Bot. Syst. IV (1857)

421; Gray in Proc. Ann. Acad. Ill (1857) 51; Gilg in Engl. u.

Prantl., Pflanzenf. Ill, 6 (1893) 143; v. Tiegh. in Ann. Sci.

Nat. VIII (1902) 395.

G. perakensis v. Tiegh. in op. cit. VIII (1902) 396 Syn. nov.

Gomphia hookeri Planch, in Hook, Lond. Joum. Bot. VI

(1847) 3; Walp., Ann. Bot. Syst. I (1848) 182; Bennett in Hook,

f., PI. Brit. Ind. I (1875) 525; King, Mat. FI. Mai. Pen. II (1893)

474; Ridl., FI. Mai. Pen. I (1922) 366 p.p.: basionym.

G. corymbosa Ridl. op. cit. I (1922) 367 p.p.

Ouratea hookeri (PI.) Burkill in Kew Bull. (1935) 318:

isonym.

Corner notes in his field label: Tree 60 ft. tall. Petals white.

Sepals filaments, ovary and style greenish white. The petals drop

off and the sepals, peduncle, ovary, carpel and disc become deep

rose-red. Kunstler’s field notes are as follows: Tree 40-60 ft.

high. Flower white with an yellow centre. Flower with young

fruits deep red, yellow inside. Md. Shah states that the tree is 25

ft. tall and that the inflorescence is cream coloured when young,

turning red when fruiting.

Malaya: Penang, Government Hill, alt. 2,500 ft. (Curtis 2,154,

= 2,154, and 1,147); Moniot’s Road, alt. 1,500 ft. (Haniff

2,409). Perak, Larut, alt. 1,500-2,000 ft. (Kunstler 7,310, iso-

holotype of B. perakensis). Malacca, Sungei Udang (Derry 935).

Negri Senibilan, Gunong Tampin, alt. 1,800 ft. (Burkill 3,167).

Johore, Mawai-Jemaluang Road (Corner 28,993); Gunong Pulai

(Henderson s.n.).

Borneo: Sarawak, Semengoh (Mead SAR 10); Bako National

Park at Bukit Gondol, alt. 700 ft. (Md. Shah P. 5,649).

The type of the species was a specimen collected by Phillips in

Penang, and characterised, as pointed out by van Tieghem, by its

peculiarly compound and globose inflorescence and entire leaves.

In the type of B. perakensis the flowers are young and so the com¬

pound character of its inflorescence is not easily seen; but in the

isotype specimens where the flowers are older, its true character

is recognized. Planchon had a specimen which had shed all its

flowers and had one fruiting pedicel. Hence he described the calyx

as reddish. Apparently this character has led Ridley to confuse

this species with the other which produces red flowers and to place

all the specimens with white flowers under G. corymbosa Ridl.,

leaving under G. hookeri none of the recent collections from

Penang. The collector’s field notes are given above to show that

the flowers are white at first and only later, when the petals have

dropped off, the sepals and other parts become red.

182

Vol. XIX. (1962).

2. Brackenridgea denticulata Furtado spec. nov.

Gomphia corymbosa Ridl. in FI. Mai. Pen. I (1922) 367 p.p.

A B. hookerii, cui affinissima, haec species foliis denticulato-

spinosis recedit. A B. serrulata conjus folia serrulata spinulosa

sunt, haec species inflorescentis capitato-compositis ut in B.

Jiookeri, haud simplicibus, facile distinguitur.

Malaya: Singapore, Bukit Timah (Ridley 10,738, holotypus

in SING.; Hullett 436 & 910; Sinclair 39,573, sterilis); loc. in-

cert. (Cantley’s collection).

The flowers are crowded on a compound axis as in B. hookeri,

but the leaves are spinulose in the margins. Without these spines,

the specimens would be easily confused with B. hookeri. Ridley

notes that it is a small tree and that its flowers are white. Hullett

(No. 910) notes that the fruit bearing calyx is red.

B. Sectio Spiciformes v. Tiegh. op. cit. VIII (1902) 397.

3. Brackenridgea corymbosa (King) v. Tiegh. in Ann. Sci. Nat.

VIII (1902) 395 & 397.

Gomphia hookeri PI. var. corymbosa King, Mater. FI. Mai. Pen.

II (1893) 475: basionym.

G. corymbosa (King) Ridl. FI. Mai. Pen. I (1922) 367 pro

parte basionymica.

A shrub 6-9 ft. tall, producing bifarous white flowers in

corymbs having a simple axis. The flowers are congested at the

apex of a longish inflorescence axis, but below them the axis show

the stalks of fallen flowers and the scales. Sometimes branches are

produced from this lower part of the axis, each bearing bifarous

flowers at its apex. The young fruit is described as pale green.

Malaya: Perak, Gopeng, alt. 500-1,000 ft. (Kunstler 4,673,

isoholotype in SING.).

4. Brackenridgea palustris Bartell, in Malpighia XV (1901) 165.

t.10; v. Tiegh. in Ann. Sci. Nat. VIII (1902) 395 & 397.

B. kingii v. Tiegh. op. cit. VIII (1902) 395: Syn. nov.

B. rubescens v. Tiegh. op. cit. VIII (1902) 395: Syn. nov.

Gomphia hookeri PI. sensu Ridl., FI. Mai. Pen. I (1902) 366

p.p.

Tree 15-60 ft. tall. Corymbs terminal, very short, with leaves

immediately below the rachis and without any long scaly rachis

below the flowers. Flowers red, bifarously arranged. Leaves vari¬

able in size, broadly or narrowly ovate or ovate-lanceolate,

usually brownish, 5-10 cm. long, 2-5 cm. wide. Fruit red.

Malaya: Kedah, Kedah Peak, alt. 3,000 ft. (Robinson & Kloss

5899); alt. 3,600 ft. (Cheang & Chang 656 & 663, red fruit and

flowers without petals red); Gunong Jerai (Mustafa CF. 20,734).

183

Gardens Bulletin, S,

Perak, Larut (Kunstler 6,396, holotype of B. kingii in K, flowers

(=fruiting calyx) dark red). Pahang, Praman (Ridley 1192, disc

red, berris black); Kuantan at Baloh (Yeop CF. 835). Malacca,

(Alviqs 876 as Pokoh Pcndarak & Pokoh Chenarahan). Singapore,

Tanjong Gol (Goodenough 1,957); Changi (Goodenough 2,072,

isotype of B. rubescens ); Tampenis Road (Ridley 4,807);

Tampenis river Ridley s.n., flowers deep red); Chan Chu Kang

(Ridley 5,896).

Sumatra: Siberut (Boden-Kloss 14,530, fruit red).

Borneo: Sarawak, Kuching Haviland 2,224 sepals and pedicels

pink).

This species was described from a specimen collected by Beccari

in Borneo (P.B. No. 3,472) by the lake of Kapuas.

The species show a good deal of variation in the size and shape

of the leaves, especially in the mountain collections. Cheang &

Chang 663 has rather longish leaves with a long tapering apex.

6. Brackenridgea serrulata Bartell. in Malpighia XV (1901) 163

t.9; v. Tiegh. in Ann. Sci. Nat. VIII (1902) 395 & 397.

Tree or shrub. Inflorescence simple with bifarous flowers which

are white at first, but become red later. Fruit bright red when ripe.

Leaves serrulate.

Borneo: Sarawak, Matang (Brooke 9,715, small tree, flowers

pale pink or white; fruit black; calyx crimson; Haviland 1,021,

timber tree); Bako National Park at Telok Asam, 400 ft. (Purse-

glove 4,919, shrub 8 ft. tall, fruits bright red).

This has serrulate leaves and white flowers as in B. denticulata,

but the inflorescence is simple, with bifarously arranged flowers.

7. Brackenridgea foxworthyi (Elm.) Furtado comb. nov.

Ochna foxworthyi Elm., Philipp. Leafl. Bot. V (1913) 1823;

Merr. Enum. Philipp. PI. Ill (1923) 68. basionym.

Philippines: Palawan, Victoria Mts. at Panacan (Sulit 12,425);

Bacungan, at Puerto Princesa (Edano 177).

The leaves are dentate or sometimes very obscurely so.

184

Name Changes in Bornean Araliads

By C. X. Furtado

Botanic Gardens, Singapore

1. Schefflera furfturacea Merr. MS.

Sch. borneensis Merr. in Joum. Str. Br. Roy. Asiat. Soc. 79

(1918) 27 basionym, non S. borneensis Merr. in Philipp. Joum.

Sci: Bot. 13 (1918) 100.

Owing to an error this species was published under a name that

had already been given by Merrill himself a few months earlier to

another plant having 5-foliolate leaves, much larger leaflets,

shorter and lightly pubescent inflorescence and shorter-stalked

umbels.

He referred to this earlier species in the introduction of a new

set of Shefflera spp. published in his second paper, and identified

with it a “specimen probably collected by Haviland” (p. 27) from

the Singapore herbarium. Yet in this second paper he used S. bor¬

neensis a second time for a different species which he obviously

intended to call Schefflera furfuracea Merr., a name written by

Merrill himself on the holotype in the Singapore herbarium. Its

leaves are described as 8-10 foliolate and is stated to have

“greatly elongated primary branches of the inflorescence which are

conspicuously brown-furfuraceous throughout and about 40 cm.

in length.”

Though these two species are manifestly distinct, unfortunately

Merrill failed to detect his error in preparing “A Bibliographic

Enumeration of Bornean Plants” (1921), so that here he united

the two species as one.

2. Schefflera petiolosa (Miq.) Harms.

Sch . racemosa Merr. in Journ. Str. Br. Roy. Asiat. Soc. 79

(1918) 30, non S. racemosa (Wight) Harms (1898). syn. nov.

Merrill’s binomial is a later homonym and so cannot be used.

Fortunately the species does not differ materially from S. petiolosa

(Miq.) Harms.

3. Arthrophyllum merrillianum Furtado nom. nov.

A. borneense Merr. in Univ. Calif. Publ. Bot. XV (1929) 231,

basionym, non A. borneense Baker (1896).

185

THE

GARDENS’ BULLETIN

SINGAPORE

Vol. XIX, Part II ^ 31st October, 1962

CONTENTS

Page

Corner, E. ). H.: The Classification of Moraceae . . .187

Ashton, P. S.: Some New Dipterocarpaceae from Borneo . .253

Whitmore, T. C.: Studies in Systematic Bark Morphology, III. Bark

Taxonomy in Dipterocarpaceae . . . . .321

To be purchased at the Botanic Gardens, Singapore

Price: $12

Published by Authority

PRINTED BY LIM BIAN HAN, ACTINO GOVERNMENT PRINTER, SINOAPOAE

The Classification of Moraceae

E. J. H. Corner

Botany School, University of Cambridge

New subgenera :—Trophis P. Br. subgen. Prototrophis Corner.

New sections :—Broussonetia L'Herit. sect. Allaeanthus (Thw.) Cor¬

ner; Madura Nutt. sect. Cardiogyne (Bur.) Corner, sect. Cudrania

(Tree.) Corner; Streblus Lour. sect. Bleekrodea (Bl.) Corner, sect.

Paratrophis (Bl.) Corner, sect. Phyllochlamys (Bur.) Corner, sect.

Pseudostreblus (Bur.) Corner, sect. Pseudotrophis (Warb.) Corner,

sect. Sloetia (Teysm. ct Binn.) Corner, sect. Taxotrophis (Bl.) Cor¬

ner; Trophis P. Br. sect. sect. Calpidochlamys (Diels) Corner, sect.

Maillardia (Frapp, et Duch.) Corner.

New series :—Maclura Nutt. sect. Cudrania (Tree.) Corner ser.

Connatae Corner, ser. Liberae Corner.

New species :—Streblus perakensis Corner. 5. solomonensis Corner.

New varieties :—Antiaris toxicaria Lesch. var. macrophylla (R. Br.)

Corner, v. welwitschii (Engl.) Corner; Broussonetia luzonica (Blanco)

Bur. v. glabra (Warb.) Corner; Madura amboinensis Bl. v. pauci-

nervia Corner; Streblus glaber (Merr.) Corner v. australianus Corner,

S. urophvllus Diels v. salicifolius Corner, S. asper Lour. v. monoica

(Gagnep.) Corner.

New specific combinations :—Broussonetia kurzii (Hook, f.) Cor¬

ner, B. zeylanica (Thw.) Corner: Madura africana (Bur.) Corner,

M. cochinchinensis (Lour.) Corner and var. pubescens (Tree.) Cor¬

ner, M. fruticosa (Roxb.) Corner, M. greveana (Baill.) Corner, M.

humberti (Leandri) Corner, M. regia (A. Chev.) Corner, M. thorelii

Gagnep.) Corner; Streblus anthropophagorum (Seem.) Corner, S.

crenatus (Gagnep.) Corner, S. elongatus (Miq.) Corner, S. glaber

(Merr.) Corner. S. ilicifolius (Vid.) Corner, S. indicus (Bur.) Cor¬

ner, S. insignis (Bl.) Corner, S. laxiflorus (Hutch.) Corner. S. tnada-

gascariensis (Bl.) Corner, S. smithii (Cheeseman) Corner. S. spinosus

(Bl.) Corner, S. tahitensis (Nadeaud) Corner, S. tonkinensis (Stapf)

Corner; Trophis branderhorstii (Diels) Corner, T. drupacea (Diels)

Corner, T. philippinensis (Bur.) Corner.

Genera fully reviewed :—Antiaris Lesch., Broussonetia L’Herit.,

Madura Nutt., Plecospermum Tree., Streblus Lour.

Introduction—Distinction from Urticaceae—Subdivision of

Moraceae (filaments; extrorse, introrse anthers; plicate leaves:

perianth; pistillode; ovary; fruit; seed; embryo; number of floral

parts; microscopic characters; inflorescence; spines)—Proto-

Moraceae—Affinity of Urticales—Geographical table of Morace-

ous genera and species—Definition of Moraceae—Key to the

tribes of Moraceae —Moreae—Artocarpeae—Olmedieae—Bro-

simeae—Dorstenieae—Ficeae—Genera excludenda—References.

187

Gardens Bulletin , S..

Introduction

No family has such small standardised flowers, yet such an

astonishing array of infructescences. Acquaintance with Morns

leads to the supposition that these tropical accomplishments are

recent developments of little significance to scientific theory. Ac¬

quaintance with the tropical plants shows, in contrast, that Morus

is one of the more derived and reduced genera conforming with

temperate simplicity. By denying itself so much in this way that is

tropical, botany loses its grip. Ficus and Artocarpus are two

vivid lights, but no student of the phylogeny of flowering plants

has seen them. As morphology shifts to the tropics, the family will

be appreciated. It holds many fascinating problems of vestigal

features, transference of function, and parallel evolution. My con¬

tribution now is merely an interim development of the static classi¬

fication into which the family has subsided. Much has still to be

learnt of the American genera, not only from the herbarium, but

from the forests in which they survive. There is, for instance, the

unique Palmolmedia described by Ducke in 1939.

Distinction from Urticaceae

This is a problem. In the diagnosis of Moraceae on p. 209 the

main differences are italicised. Chief are the orthotropous ovule

and the lack of latex in the Urticaceae. On the ovular test Cono-

cephaloideae must be transferred to Urticaceae and, this done,

the genera in each family hold together. Latex-tubes are said to

occur in the bark, at least the primary bark, of all Conocephaloi-

deae ( Cecropia , Coussapoa, Musanga, Myrianthus, Poikilos -

permum, and Pourouma ), but not in the leaves or inflorescences

and infructescences. This tribe may seem, therefore, intermediate

between the two families, but other features as the shape of the

stigma, the small seed, and the small embryo ally them with

Urticaceous, not Moraceous, genera. Trifling as these two distinc¬

tions may seem, they delimit two large families, and the seeming

triviality arises from lack of appreciation of the two characters.

Fatoua is not an Urticaceous herb, but a Moraceous parallel.

Cecropia is not a Moraceous tree, but an Urticaceous parallel.

The function of latex is unknown, yet it permeates as a capillary

system giant Moraceous trees from root-tip to stem-tip, and it is

initiated in the embryo. The function of ovular shape is also

unknown, but it relates to angiospermous ancestry. The evidence,

anatomical and taxonomical, suggests not that the direct ortho¬

tropous ovule is primitive, but that it is derived from the ana-

tropous ovule transferred to the base of the ovary and straightened

in conformity with its improved position for food-supply. In ana¬

tomical evidence, Bechter (1921) described the flexure of the

ovular vascular bundle in Boehmeria and Laportea, which first

ascends in the ovary-wall as if to an apical position, and then des¬

cends to the basal attachment. Certain tropical genera as

188

Vol. XIX. (1962).

Stenochasma and, indeed, Pourouma have sub-basal and even

laterally placed ovules which turn abruptly into the long ortho-

tiopous body. In this respect they appear intermediate with

Moraceae, but details of flower and fruit again refer them unques¬

tionably among Urlicaceae.

Urticaceae are derivatives of Proto-Moraceae which lost the

latex and in which the ovule was transferred from the lateral

(appendicular) position to the axile. Conocephaloideae are the

Urticaceous group with traces of Proto-Moraceous ancestry. Ac¬

tually there is no difficulty whatever in separating most Moraceae,

as Artocarpus or Ficus, from Urticaceae; the trouble arises in the

very derived and simplified genera as Dorstenia, Morus, and Stre -

blus, convergent with Urticaceous derivatives.

Lack of latex distinguishes also the Ulmaceae, the flowers of

which are generally functionally bisexual. The combination of

Ulmeae with Celtideae in one family introduces several of the

difficulties met in the subdivision of the Moraceae, especially the

two subfamilies Moroideae and Artocarpoideae. As I will show,

these two subfamilies cannot be maintained, at least in their pre¬

sent light, because the nature of the stamen, inflexed or straight

in bud, though useful enough in the material on which taxonomists

have to rely, has not the phyletic significance which practice would

attribute. The same solutions will probably fit Ulmaceae and, then,

the position of the Cannabiaceae will be ascertained.

Subdivision of Moraceae

To the student of Ficus the classification and, therefore, the

comprehension of Moraceae is confused with excessive genera.

The fig (syconium) is so characteristic that no one is willing to

make Ficus a tribe or family and raise its subdivisions to generic

or tribal rank. Further, there is a continuum through Ficus link¬

ing the first with the last. It is a genus which preserves a great

deal of its evolution, and its products may still be strung to¬

gether. But, suppose no other Moraceae existed and that most

modern Ficus had disappeared, then Ficus would be a family

with several distinct genera. In this light other Moraceae must be

considered; they are fragments only from a greater past.

An exception was Miquel. Where others have seen in Ficus

merely specific variety, and have failed, only too often, even to

dissect the flowers, Miquel studied deeply and was so impressed

with the great range of floral variation in the genus that he pro¬

posed many genera in place of the one. His prowess defeated him;

he was overwhelmed with sterile material for identification and

had to forgo his principle. Yet, if the flowers were on the outside

of the head, instead of inside the fig, many would have seen without

difficulty and have followed him and argued that a colossal mo¬

noecious banyan with unistaminate flowers and simple stigma

could not be congeneric with a slender dioecious creeper with

189

Gardens Bulletin , 5 .

bistaminate flowers and bifid stigma. The enormous bracteate fig

of Dammaropsis differs as much from that of F. carica as the

Olmedieae do from the Artocarpeae, yet Dammaropsis is also

sunk in Ficus. The point is not that there is so much variation in

this genus but that the extremes are linked by intermediates. For

instance, the group of F. hirta (2 stamens, peculiar seed, no cys-

toliths, palmate-leafed trees) is very distinct from that of F. subu-

lata (1 stamen, gall-ovary in the male flower, simple seed, cysto-

liths, simple-leafed epiphyte), but the remainder of their sections

Ficus and Sycidium link these extremes together.

f take Ficus as demonstrative, and where monoecism or dioec-

ism, staminal number, perianth-construction, stigmatic and seed-

details, ovary-position, microscopic differences in cystoliths,

gland-hairs, hypodermis, etc., and differences in habit and leaf are

the chief distinctions between genera, I treat them with suspicion.

Caution is needed, nevertheless, in recombining the fragments of

other Moraceae. There has been much parallel evolution, even

Sparattosyce simulating Ficus, so that similar effects need meticul¬

ous care in separation, and endless tables of comparison must be

prepared to sort out the details. Endless because it is always

found that much critical information is lacking and cannot be

obtained from the bits of the plants in herbaria; too often the

American and African plants are in default. Yet, the pieces of the

puzzle are so strange, so splendidly different from the banalities of

many other families, that the monography of the Moraceae is one

of the most exciting chapters in angiosperm taxonomy. To relate

a mulberry and a bread-fruit, one superficially the aggrandisement

of the other, and add the little Fatoua calls for a working hypo¬

thesis of flowering-tree evolution. A concept of Proto-Moraceae is

essential.

Filaments. The form of the filament in the flower-bud has been

taken as a primary means of distinguishing subfamilies or tribes.

The filament may be short and lengthen directly to project the

anther or it may be inflexed under tension until the flower opens

and the filament straightens and flicks the pollen from the anther.

Such is the Urticaceous stamen, and the difference between sub-

fam. Moroideae (inflexed filament) and subfam. Artocarpoideae

(straight filament). Certain facts discredit this criterion.

Because of their orthotropous ovule and general affinity in

other respects, the subfam. Conocephaloideae must be transferred

to the Urticaceae. This at once introduces the straight stamen

( Musanga, Myrianthus , Cecropia, Coussapoa, Pourouma) into the

Urticaceae. Poikilospermum has always been placed in the Urti¬

caceae because of its inflexed filament. This character holds only

for one part of the genus; the other part (Conocephalus Bl.) has

the straight stamen and has always been referred to the Moraceae.

It is acknowledged now that these two subgenera are congeneric;

some species, indeed, have such slightly inflexed filaments that it

is not clear on this point alone to which subgenus they should be

190

Vol. XIX. (1962).

referred. Then, Cudrania, placed in Artocarpoideae, has straight

filaments but, as 1 will show, is merely a section of Maclura

(Moroideae) with inflexed filaments; there is the reputed hybrid

Madudrania (M. pomifera X C. tricuspidata), but its filament

has not been described. Again, for the same reason Sorocea and

Clarisia are placed in Artocarpoideae where their racemose or

spicate inflorescences and non-syncarpous fruits are entirely ano¬

malous; in fact, they are nearer to Trophis (Moroideae) with in-

flexed filaments. Craterogyne (inflexed filaments) has been placed in

Dorstenieae and Trymatococcus (straight filaments) in Brosimeae,

yet other differences are so slight that I would refer them to one

genus of Brosimeae. Lastly, so far as I have been able to find out,

subinflexed filaments are described by Woodson (1960) in Baio-

carpus (Artocarpeae).

The inflexed filament has a small anther, often reniform or bi-

lobed. The flower has at least two stamens ( Craterogyne, Dorste-

nia), generally 3-5, and it has a pistillode against which the anthers

are pressed immovably in the bud. The filaments are at first short

and straight; in lengthening they are forced to curve outwards

against the bulging concave tepals and so they become, as it were*

inflexed; they never inflex. The inflexed filament, therefore, in this

as in other cases, such as the Caesalpinioideae, is related with the

immovability of the anther in bud. In some genera, nevertheless,

the pistillode is minute or even absent, and one must suppose that

the connivent anthers caused the so-called inflection.

The straight filament commonly bears a rather large anther,

often mucronate, typical of the normal angiosperm stamen. Its

flower is often unistaminate, but may be 2—5 merous, and there

may be 7-8 stamens ( Antiaris , Ficus). It rarely has a pistillode

(present in Ficus pr. p., Antiaropsis, Sparattosyce). In develop¬

ment the anther fills most of the bud-cavity, and not until the flower

begins to open does the filament elongate. Extended anther-growth

and retarded filament-growth distinguish this flower from that

with inflexed filament. Now which may be ancestral?

The straight filament is generally associated with the advanced,

capitate or condensed, inflorescence, as in Artocarpeae, Brosimeae,

Olmedieae, and Ficeae. The inflexed filament is associated with

the less advanced spicate, racemose, cymose, and paniculate in¬

florescences, an exception being the Dorstenieae. The straight

filament is more suited to the compacting of flowers because it

does not need the elbow-room for the explosive stamen. For this

reason the inflexed filament seems the more primitive. It is the

straight filament, however, which has the less specialised anther.

The minute anther of Artocarpus is clearly as derived as it is

diminished from the large anther shown by Parartocarpus. The

best developed anthers, strongly mucronate, occur in Ficus sect.

Rhizocladus (climbers). Likewise the small anther of the inflexed

filament must be derived. Possibly therefore the inflexed filament

191

Gardens Bulletin, S.

is derived from the stamen with small anther, in turn derived

from the stamen with large anther and straight filament, and one

suits one kind of inflorescence, the other another. It may be re¬

called that in all flowering plants the anther develops before the

filament which is an intercalary growth. The ability of the anther

to fill the bud-cavity determines the form of the filament. So the

large anther and necessarily short filament seems the primitive con¬

dition, and it brings the Moraceous stamen into line with the

theory of Parkin (1951).

It follows that the inflexed filament has been evolved in parallel

in different Moraceous and, even Urticaceous and Ulmaceous,

groups or genera. That such parallel evolution may well have

taken place is shown by the parallel occurrence of the extra¬

ordinary turbinate anther with transverse equatorial dehiscence

found in Ficus sterrocarpa Diels, Brosimum alicastrum Sw., and

Craterogyne kameruniana (Engl.) Lanjouw (Fig. 1, L, N).

I conclude that the form of the filament is not useful for pri¬

mary distinctions and that it leads to artifical grading. Further,

I conclude that the presence of the straight stamen in Moraceous

groups with condensed inflorescences shows that they are variously

derived from states with expanded inflorescences, and that any

attempt to relate them directly is short-sighted, because they are

the survivors of as varied Proto-Moraceae which have given rise

at any stage of inflorescence-evolution to forms with inflexed

filaments.

Extrorse, introrse anthers. Ficus, Antiaropsis, and Antiaris have

introrse anthers. Other genera, so far as I can determine, have

extrorse anthers. In those with inflexed filaments, the anthers are

often described as introrse, but in bud they are extrorse and only

after they have been pulled out and rotated on the filaments do

they become introrse. The distinction is important because the in¬

trorse anther would not be so effective in the wide-spread explo¬

sive stamen-mechanism.

Plicate leaves. Engler used this character to define subfam.

Moroideae in contrast to the inrolled or convolute leaf in sub¬

lam. Artocarpoideae. However, all the smaller elliptic leaves of

Moroideae are convolute as in Maclura and Streblus, and all the

larger palmate, pinnate, or cordate leaves of Artocarpoideae are

plicate, as F. carica, and Artocarpus incisa. Large leaves, plicate

in bud, seem to be the pachycaulous character, small leaves con¬

volute in bud the leptocaulous as in other families and, indeed,

genera.

192

Vol. XIX. (1962).

(Roxb.) Corner, with flanged and ridged filament curved at the

tip. K, Stamen with incurved filament, as in Moreae. L, Diagram

of the stamen of Ficus sterrocarpa Diels. M, Diagram of the

stamen of F. hesperidiiformis King. N, F, sterrocarpa. O, F.

h esperidiifor mis.

Perianth. The more or less free tepals of many genera contrast

with the gamophyllous or utriculate perianth of others, and both

with the state without perianth. All states occur in Ficus. To at¬

tempt to use them as primary distinctions leads to confusion.

Thus Artocarpus subgen. Pseudojaca, Hullettia , and Trymatococ -

cus are not to be classed together because they have practically

no perianth, any more than Strehlus sect. Bleekrodea , Malaisia ,

and Ficus sect. Sycocarpus are to be classed together because of

their utriculate perianth. These peculiarities have also been

evolved in parallel.

193

Gardens Bulletin , S.

Fig. 2. Male flowers of Moraceae to show the pistillodcs. x 10. A, Ficus

obscura Bl. B, F. henryi Diels, two flowers. C, F. wassa Roxb.

D, Plecospermum spinosum Tree. E, Streblus elongatus (Miq.)

Corner. F, Morns macroura Miq. G, Maclura thorelii (Gagnep.)

Corner.

Pistillode. This diminutive sterile structure of the male flower,

apparently indicative of the bisexual origin of the Urticalean

flower, poses several problems. It is best developed as a short

quadrate-columnar structure in Moreae and Urticaceae: against it

press the four anthers with inflexed filaments. In contrast it is a

minute, but very persistent, spike in Fatoua, Streblus (sect. Blee-

krodea , Pseudostreblus, Sloetia ), Sloetiopsis, and Neosloetiopsis .

It is absent from Clarisia and Sorocea. To separate these sections

of Streblus and classify them with Fatoua on this ground is un¬

reasonable in the face of their overwhelming resemblance in other

respects with the rest of Streblus. A similar pistillode occurs in

Antiaropsis and Sparattosyce in the Olmedieae and, sometimes,

as the merest vestige, in Parartocarpus and Treculia in the Arto-

carpeae, and Trymatococcus in the Brosimeae. In other Artocar-

peae, as Broussonetia, Maclura, and Plecospermum, the pistillode

is better developed as a small ligulate ovary with a minute style

and, even, stigmatic arms. Jn Helianthostylis, generally placed in

Artocarpeae but problematic, it is long, filiform, and exsert from

the flower. Finally, in Ficus , it varies from a vestigial, but clearly

194

Vol. XIX. (1962).

recognisable, ovary to a minute spike, or it is absent. In two cases,

however, namely Ficus henryi Diels and sect. Sycidium subsect.

Palaeomorphe , the pistillode functions as a gall-ovary with insect.

These are the nearest approach to the functionally bisexual flower

in the family. Of all the taxonomic characters in the family, the

pistillode is the most demonstrative of the value of vestigal

inheritance.

Fig. 3. Ovaries of Moraceae, X 7. A, Maclura cochinchinensis (Lour.)

Corner. B, M. thorelii (Gagnep.) Corner. C, M. fruticosa (Roxb.)

Corner. D, Streblus glaber (Merr.) Corner. E, S. asper Lour.

F, S. macrophyllus Bl. G, S. solomonensis Corner. H, Malaisia.

I, Ficus macrostyla Corner. J, F. deltoidea Jack. K, F. auran-

tiaca Griff. L, F. fistulosa Reinw. M, Broussonetia papyrifera

(L.) Vent. N, O, Plecospermunx andamanicum King. P, Arto-

carpus hirsutus Lam. Q, A. lanceifolius Roxb. R, A. vrieseanus

Miq. S, Parartocarpus venenosus Zoll. et Mor. T, Fatoua. U.

Prainea frutescens Becc. V, Antiaropsis. W, Antiaris toxic-aria

Lesch., the 1-flowered female inflorescence with inferior ovary.

195

Gardens Bulletin, S.

Ovary. There are three states. Firstly, the ovary is superficial.

■Secondly, it is sunk in loculi or sockets of the receptacle with the

perianth free from the walls of the loculus ( Artocarpus subgen.

Artocarpus, Craterogyne, Plecospermum ), or there is no perianth

(Artocarpus subgen. Pseudojaca, Hullettia, Parartocarpus).

Thirdly, the ovary is truly inferior and only the ovule is free in

the loculus of the receptacle ( Antiaris, Castilloa, Trophis, and

many Brosimeae); in several cases the ovary is partly inferior,

<( Sorocea, Maclura sect. Cudrania). The semi-inferior ovary oc¬

curs also in Ficus subgen. Urostigma sect. Malvanthera and this

shows that ovary-displacement has also suffered parallel evolution

and is not a primary criterion for classification.

Fruit. The ovary develops into a drupe. There are traces in

Morus of two loculi, each with an ovule, in the drupe, but in all

other genera, whether there are one or two styles, or one or two

stigmatic arms to the single style, ovary and drupe are unilocular

and uni-ovulate. The largest drupes with best developed pulpy

exocarp and woody endocarp occur, so far as known, in the ad¬

vanced genera Parartocarpus and Treculia. Modifications are

these:—

1. The drupe diminishes to a drupelet (Fatoua, Ficus ,

Dorstenia, Broussonetia).

2. The drupe develops a strongly thickened parenchymat¬

ous base and dehisces over the thin apex to eject the seed

(in endocarp) forcibly ( Streblus sect. Phyllochlamys, Sloetia,

Taxotrophis, and possibly Bleekrodea; Ficus spp., Dorstenia

spp.).

3. The perianth (always persistent) becomes fleshy, often

at the expense of the exocarp (Morus, Ampalis, Pachytrophe ,

Maclura, Artocarpus subgen. Artocarpus).

4. The receptacle becomes pulpy to form the fleshy syn-

carp (most Artocarpeae, Brosimeae, Olmedieae, and Ficus).

These modifications have occurred in parallel evolution. The

second is highly peculiar because it re-instates dehiscence by a

transverse, not longitudinal, rupture in the indehiscent drupe. It

is a problem whether this is the original fruit-form in Streblus or

a subgeneric product. In some species of Ficus, without syste¬

matic significance, and in Dorstenia the crustaceous endocarp is

squeezed cut of the pulpy drupelet.

196

Vol. XIX. (1962).

Seed. By seed I imply not only the correct botanical usage but

the practical. 1 intend the endocarp and its enclosed true seed:

strictly it is a pyrene, not an achene, but everyone calls these

bodies the seeds. There are small seeds 0.5-3 mm. long, and

large seeds 4-20 mm. long. The one genus with both kinds is

Ficus; its seeds are mostly 1-2 mm. long, but seeds 3-5 mm. long

occur in F. deltoidea and its immediate allies. According to my

durian-theory (Corner, 1954) the large seed is primitive. This

conclusion agrees in general with the primitive state of the straight

filament, for most genera with this character are tropical forest-

plants with large seeds. Small seeds preponderate in genera, in¬

cluding Urticaceae, with inflexed filaments, both tropical and

extra-tropical ( Broussonetia, Monts), and also herbaceous ( Dors -

tenia, Fatoua). The exception is Ficus which is the small-seeded

derivative with straight stamens, adaptable in consequence to

secondary vegetation and the epiphytic habit. The conclusion em¬

phasizes again the retention of primitive features in Artocarpeae,

Brosimeae, and Olmedieae with advanced inflorescence.

Embryo. There is such diversity in embryo-characters that it

would seem that they should be of prime importance in classifica¬

tion. From this standpoint I have retired after much trial. There

can be no doubt from the impossibility of fitting embryo-charac¬

ters into a satisfactory scheme of classification that they represent

also parallel evolution. Small seeds have small simple embryos

with relatively long radicle and flat, thin cotyledons ( Brousso¬

netia, Dorstenia, Fatoua, Ficus, Morus)\ they are not primitive

but neotenic, derived from the complicated embryos of the large

seeds. The variety is shown in Fig. 4, 5. There is every gradation

from curved embryos with plicate-conduplicate cotyledons to

straight embryos with two thick, flat cotyledons and those with

such very unequal cotyledons as to be functionally monocoty-

ledonous. Streblus and Maclura, even Ampalis and Pachytrophe,

reveal what has happened, for they show the range from rather

thin, plicate-conduplicate cotyledons, which most resemble the

foliage leaf with plicate vernation (otherwise lost in these genera),

to the simple state with incumbent radicle ( Streblus) or accumbent

( Ampalis, Maclura). The trend is roughly parallel with the dimi¬

nution of seed-size. Then comes thickening of the cotyledons,

leading to the hypogeal, non-photosynthetic state, accompanied by

loss of folding and by unequal development of the two. There are

considerable differences depending on the exact shape of the

197

Gardens Bulletin , S.

ovule, whether anatropus or campylotropous, the unequal deve¬

lopment of the ovary or ovule after fertilisation, and the orienta¬

tion of the embryo about the sagittal plane of the ovule. These

are intricate points which require copious material for elucida¬

tion, together with seedling-studies. They have been worked out

in some species of Artocarpus, Parartocarpus , and Prainea by

Jarrett (1959, 1960). The conclusion must be that the small and

simple embryos of Broussonetia, Ficus , and Monts are parallel

with those of Urticaceae; that the fleshy cotyledons specialise as

hypogeal food-stores for the plumule; and that the primitive state

with the cotyledons resembling the foliage leaf is as rare as in

most other families. Similar argument will apply to the Ulmaceae.

Fig. 4. Moraceous embryos, X 5. A, Madura africana (Bur.) Corner.

B, M. pomifera (Raf.) Schneid. C, M. cochinchinensis (Lour.)

Corner. D. M. thorelii (Gagnep.) Corner. E, Broussonetia papy-

rifera (L.) Vent. F, Morus macroura Miq. G, Antiaropsis deci -

piens K. Schum. H, Trophis branderhorstii (Diels) Corner. I,

Ficus deltoidea Jack. J, F. hesperidiiformis King. K, F. rhizo -

phoriphylla King. L, F. grossularioides Burm. f.

198

Vol. XIX. (1962).

Fig. 5. Moraceous embryos, x 5: those of Artocorpus and Parartocarpus,

X 1. A, Plecospermum spinosum Tree. B, Streblus insignis (Bl.)

Corner. C, S. asper Lour. D, S. indicus (Bur.) Corner. E, S.

elongatus (Miq.) Corner. F, Artocorpus integer Linn. f. G, A.

heterophyllus Lam. H, A. elastic us Reinw. I, A. glaucus Bl. J,

A, anisophyllus Miq. K, Parartocarpus venenosus Zoll. et Mor.

The embryos may be classified as follows:—

A. Cotyledons rather thin.

1. Cotyledons conduplicate, plicate; radicle long.

Broussonetia kurzii; Madura sect. Cardiogyne, sect. Cudrania pr.p.;

Pachytrophe, Sparattosyce, Streblus sect. Paratrophis pr.p.

2. Cotyledons conduplicate, not plicate.

Ficus pr.p.

3. Cotyledons flat or curved, not folded,

a. Radicle incumbent.

i. Embryo longitudinal in fruit.

Broussonetia, Streblus sect. Paratrophis pr.p.

ii. Embryo transverse in fruit.

Dorstenia, Fatoua, Ficus pr.p., Morus.

199

Gardens Bulletin, S..

b. Radicle obliquely incumbent to accumbent.

Ampalis, Maclura sect. Madura and sect. Cudrania pr.p.

c. Embryo straight.

Ficus pr.p.

B. Cotyledons thick, fleshy.

1. Radicle long, often incumbent.

a. Cotyledons conduplicate.

i. Cotyledons subequal.

Antiaropsis, Streblus sect. Paratrophis pr.p.

ii. Cotyledons very unequal.

Treculia.

b. Cotyledons not folded, equal.

Parartocarpus, Trophis.

2. Radicle short, curved: cotyledons not folded.

a. Cotyledons equal.

Artocarpus subgen. Artocarpus pr.p. (radicle incumbent or ac¬

cumbent.).

b. Cotyledons unequal.

Artocarpus subgcn. Artocarpus pr.p., Malaisia, Plecospermum,

Sorocea; Streblus sect. Pseudostreblus, Streblus and Taxotrophis.

3. Radicle very short, almost none; embryo straight or slightly curved.

a. Cotyledons equal.

A nonocarpus, Antiaris, Artocarpus subgen. Pseudojaca, Castilloa>.

Hulletia, Noyera, Prainea.

b. Cotyledons unequal.

Streblus sect. Bleekrodea and sect. Sloetia.

Number of Floral Parts. In Ficus there are many differences in

number of stamens (1-8) and tepals (0-8); the style may have one

or two stigmatic arms, and the gall-stigma may differ. These nu¬

merical points have been exaggerated into generic characters in

other tribes, chiefly because their species are more relictual and

the trivialities appear profound. In the light of Ficus, the pen-

tamery of Pseudostreblus and the trimery of Sloetia are not gene¬

ric differences from the tetramerous Streblus.

Microscopic characters. The striking taxonomic differences in

the occurrence of cystoliths, hypodermis, crystals, resin-cells,

gland-hairs, papillate hairs, and so on, in the leaf were first ex¬

plained by Renner (1907). Their value is shown also by Ficus,

where they have variously subgeneric, sectional, and specific rank.

The temptation is to believe that they may be generic characters

in the fragmentary tribes. A detail worth pursuing is the nature

of the resin-cells which occur in the mesophyll of Artocarpus,

Maclura sens, lat., Malaisia, and Prainea. These cells are known

to be absent from Broussonetia, Plecospermum, and Parartocar¬

pus, all of which I classify in the Artocarpeae, but there are still

many genera in which they have not yet been studied. In Prainea

and Artocarpus subgen. Artocarpus , yellow resin-cells are scat¬

tered in the mesophyll. In Artocarpus altilis (Miq.) J.J.Sm. they

200

voi. xix. {m2).

are aggregated into compact glands (masses of yellow resin-cells)

in the leaf-teeth. In Maclura they are aggregated into 2-7 yellow

glands immersed in the mesophyll of the bracts and/or the tepals.

In Malaisia they form a glandular cylinder, or ring in transverse

section, in the mesophyll of the pericarp. In Prainea, further,

they form small masses in the mesophyll of the perianth. Yet,

from Artocarpus subgen. Pseudojaca, with the exception of A.

•altilis, the resin-cells are absent.

Fig. 6. Diagrams of the inflorescences of Moreae (A-G) and Artocarpeae

(H-R). A, Fatoua. B. Streblus ( Bleekrodea ) insignis (Bl.) Cor¬

ner. C, S. madagascariensis (BI.) Corner. D, S. ( Pseudostreblus)

indicus (Bur.) Corner. E, S. ( Sloeiia) elongatus (Miq.) Corner.

F, S. ( Streblus) asper Lour. G, Streblus sect. Pseudotrophis and

Taxotrophis (pedicellate flowers) and sect. Paratrophis (sessile

flowers). H, Maclura seel. Maclura. I. Broussonetia. J, Malaisia.

K, Maclura sect. Chloroxylon . L. Maclura sect. Cudrania. M,

Plecospermum. N, Parartocarpus. O, Artocarpus , male. P. sub¬

gen. Artocarpus. Q, Artocarpus subgen. Pseudojaca. R, Treculia.

201

Gardens Bulletin, S.

Fig. 7. Diagrams of inflorescences of Dorstenicae (A), Brosimeae (B, C)„

Olmedieae (D-l), and Ficus (J-P). G, H, 1-flowered female in¬

florescences of Olmedia and Antiaris respectively. I, Male and

female inflorescences of Sparattosyce. J, Ficus subgen. Urostigma

sect. Malvanthera, the ovaries immersed in the fig-wall. K, gall-

fig of Ficus subgen. Ficus sect. Kalosyce, with disperse pedi¬

cellate male flowers. L, multibracteate seed-fig of Ficus subgen.

Ficus sect. Sycocurpus. M, Ficus subgen. Urostigma, male, gall-,

and seed-flowers disperse in the same fig. N, Ficus subgen. Syco-

morus, with male flowers ostiolar and sessile. O. P, Ficus sub¬

gen. Ficus, gall-fig with ostiolar male flowers and seed-fig,

dioecious.

Inflorescence. The shape and structure of the inflorescence

seems to be the only satisfactory basis for primary classification.

Ficus is one example, Brosimeae another, because they are cons¬

picuous. Far less obvious is the unity of the relatively unspecia¬

lised panicles, racemes, spikes, and heads of Moreae. Because the

advanced inflorescences resulting in syncarps must have been

derived from less advanced inflorescences, as shown by the

Moreae, the tendency is to regard this tribe as primitive. Its

genera, however, are all advanced in many respects, and much

more can be learnt about Proto-Moraceae from the other tribes.

202

Vol. XIX. (1962).

When defined broadly, these tribes stand by themselves and point

as variously back to Proto-Moraceae. The splitting of genera and

tribes in a confusion of floral detail has obstructed a clear view

of the family. Indeed, so far as clarification is now needed, the

question must be considered whether there is not just one genus

for each tribe, comparable with Ficus. The point, as already ex¬

plained, is that Ficus, as the most recent ebullition of the family,

still shows a fairly continuous spectrum of evolution. The other

tribes have merely fragments of their spectra. It is the lack of

detailed knowledge to bridge these gaps which makes the putting

together of these genera at present highly dubious. I have joined

several genera into Streblus and have felt inclined to join all into

one genus Morus for the Moreae. 1 have joined three or four

genera in Madura, and united Allaeanthus with Broussonetia.

Compare, then, the small genera Malaisia, Prainea, and Plecos-

permum, and it will be seen how difficult it is to place them with

satisfaction. Artocarpus, Parartocarpus, and Treculia introduce

another set of differential details.

A peculiar point about the spicate inflorescences of Moreae

and some Artocarpeae is the presence of a sterile longitudinal

strip on one side, to which Baillon first drew attention. It needs

investigation. It does not occur in the racemose or paniculate

inflorescences with pedicellate flowers, and it is absent from the

capitate inflorescences, though vestigial in the subcapitate female

inflorescence of Malaisia .

I have united Broussonetieae with Artocarpeae because I can

find no sharp separation. Perhaps this sterile groove is significant.

It occurs in the spicate male inflorescences of Broussonetia, Mac -

lura, and Malaisia, but not in the spicate inflorescences of

Artocarpus.

Spines. Short shoots ending in a spine (stem-tip) distinguish

Streblus sect. Phyllochlamys, sect. Pseudotrophis (excluding S.

macrophyllus B1.), and sect. Taxotrophis, and also Madura (ex¬

cept for the African trees of sect. Chlorophora). In M. africana

(Cardiogyne ) the spines end short leafy shoots: in the others the

spines are leafless, or nearly so. It seems that the feature has de¬

veloped in parallel in these two genera, just as the spinous leaf-

edge of Streblus ilicifolius parallels that of Sorocea. It is remark¬

able that the habit has not developed in any other genus, though

in Ficus there is the solitary example of a spinous leaf-edge in F.

macilenta King var. ilicifolia Comer.

Proto-Moraceae

To present an archetype is often considered mere speculation.

Then, what is it that systematists imply when they put into genera

and tribes the remnants of ancient groups? Must they be blind

to the pictures this sort of pattern will show? Every taxon im¬

plies, unless one disbelieves evolutionary theory, an antecedent.

203

Gardens Bulletin , S.

The more numerous the taxa, the clearer should become the

antecedent. Taxonomy which denies itself this duty to speculate

grows more and more minute and is well-labelled microtaxonomy.

Without a hypothesis of origins, the taxonomist cannot appreciate

what he is doing. I often think that taxonomy is a larder of no¬

tions awaiting thought.

The following is my view on the ancestors of Moraceae, sup¬

ported by reasons.

Fig. 8 . Proto-Moraceae in reconstruction. Pinnate-leafed pachycaul stem,

X 1/6. Male and female flowers, x 3. Fruits, one with echi-

nulate drupe, the other with smooth drupe and spinous peri¬

anth-segments, x 1. Seed (endocarp) in section. X 1.

Characters of Proto-Moraceae. —Monoecious, pachycaul, pin¬

nate-leafed; stipules foliaceous, amplexicaul; latex present. Inflo¬

rescences cymose-paniculate; flowers unisexual, pedicellate, ape-

talous, male and female in the same inflorescence; perianth with

204

Vol. XIX. (1962).

5-8 imbricate tepals. Male flowers with the stamens opposite the*

tepals; anthers large, mucronate; filaments straight in bud; pistil-

lode as a sterile ovary. Female flowers (?with abortive stamens);

ovary superior, with 2 loculi or one; styles 2, long, or one with

two long stigmatic arms; ovule one in each loculus, subapical,

anatropous. Drupe large, with woody endocarp, surrounded at the

base by the persistent spicate tepals. Seed with membranous testa,

enclosed in the endocarp, with little endosperm. Embryo curved;

radicle long, incumbent; cotyledons foliaceous, plicate-condupli-

cate; germination epigeal. Leaf with cystoliths on both sides; hypo-

dermis ? none; stomata superficial on the lower side; microscopic

gland-hairs multicellular, ? subglobose or peltate; hairs papillate.

Monoecious. This state is indicated as the primitive in Ficus

subgen. Urostigma, Pharmacosycea, and Sycomorus, and there is

clear evidence of the derivation of the male and female inflores¬

cences, leading ultimately to dioecism, in Streblus sect. Bleekrodea

and sect. Streblus , in Treculia and in Parartocarpus.

Pachycaul, megaphyllous, with foliose stipules. This ancestral

state is shown by Ficus sect. Sycidium, Sycocarpus, and Adenos -

perma, perhaps also by sect. Galoglychia (subgen. Urostigma).

It is shown for Olmedieae in Palmolmedia. Except for the foliac-

ous stipules it is the state of Artocarpus anisophyllus and A .

incisus. It is indicated in Bagassa, allied with Maclura, in Brous-

sonetia, and in Dorstenia. Specialisation in all these genera dis¬

playing pachycaul origin leads to the leptocaul and the willow-

leafed habit in their more derived species. The point is important

because it means that the family Moraceae and its tribes arose as

short pachycaul trees and that such genera as Ficus, Artocarpus,

Maclura, Broussonetia, and Antiaris have been independantly

evolved into large trees, as some of them have been evolved into

climbers. Comparative anatomy must learn from the anatomy of

pachycaul plants, knowledge and, indeed, appreciation of which is

still so deficient. The communities of banyans, which assemble on

the ruins of civilisations, are not primitive fig-forests, the nature

of which is to be seen rather in the secondary vegetation of pachy¬

caul figs in Borneo, New Britain, and Bougainville Island.

Inflorescence. When traced from their inception, the flowers

even of Ficus and Dorstenia are found to arise in a cymose pattern

(Bernbeck, 1932). Intercalation of intemodes into capitate inflo¬

rescences leads back to the racemose-cymose state, and this leads,

by branching, to the paniculate. The cymose monoecious remnants

as Streblus sect. Bleekrodea, Brosimeae, and Dorstenieae indicate

that there was a central female flower, terminal and surrounded

by male flowers as in Euphorbiaceous inflorescences. Conversely,

condensation of such an inflorescence with or without dioecism

leads to the variety of the Moraceae. The strong protogyny in bi¬

sexual Moraceous inflorescences also indicates the cymose cons¬

truction with terminal female flower.

205

Gardens Bulletin , S.

Flower-structure. The conclusions, argued on preceding pages,

agree with those of Bechtel, based on vascular anatomy. Proble¬

matic is the antitepalous position of the stamens. It suggests miss¬

ing petals of which there is no external sign in the Urticales. In

Ulmus americana , Bechtel found abortive vascular bundles be¬

tween and alternating with those of the tepals and stamens, as

internal vestiges of petals; also, in this flower of floral formula

P(8)/A8, there were six abortive vascular bundles internal to and

alternating with the staminal; the ovary has four vascular bundles.

Thus, the Ulmaceous flower gives evidence of 5 phyliotaxis in

perianth and androecium, descending to i in the gynoecium. In

most Moraceae this has become ± phyliotaxis throughout the

flower, as P 2+2 A2 + 2 Gl, with two styles or stigmatic bran¬

ches. Such falling phyliotaxis is proof that the Proto-Moraceous

flower is descended from one of massive construction with many

parts.

Fruit. Of an antecedent drupe there can be no doubt, but the

Proto-Moraceae must be considered along with other flowering

plants. The syncarps of Artocarpus subgen. Artocarpus , of Parar-

tocarpus, and of Treculia strongly resemble the fruit of the durian

iDurio zibethinus). The spines of the durian are outgrowths

of the ovary located beneath peltate scales; the pulp of the

durian-seed is a true aril, its testa the outer integument. The

spines of the Moraceous syncarps are thickened perianths, tepals

or bracts; the peltate scales are bracts with peltate tops; the seed

is a pvrene, the endocarp of which forms a false testa; the pulp

is the outer part of the ovary-wall or the perianth. Transfer, then,

the testa-lignification of Durio from the outer integument to the

endocarp, the aril-pulpiness to the exocarp, the spinous character

to the tepals and bracts, and there is the transformation into

Parartocarpus or Treculia. In Artocarpus subgen. Artocarpus the

perianth is divided into two parts, a proximal and a distal, both

free from their neighbours but fused together with them laterally

by a middle layer; the proximal part becomes the pulpy false aril

and the distal part becomes the spine. A simple transfer of func¬

tion, or gene-action, by one or two tissue-layers away from the

ovule occurs and the durian-fruit is converted into the Moraceous

syncarp. This false fruit is a syncarp not of carpels but of syn-

carpous ovaries; it is a second-order fruit, and thus every durian-

feature is displaced outwards from its primary position to a

secondary. That is, the stem-apex forms not floral primordia to

become large flowers, but floral primordia so reduced and neo-

tenic that they become, as it were, single uni-ovulate carpels which

function as arillate seeds, and the reduced tepals arc converted into

spines. It requires the crowding of primordia on an embryonic

condensed inflorescence for such displacement of differentiation.

Thus it is the elaborate syncarp of Artocarpeae, not the simple

drupe of Moreae, which has the primitive characters.

206

voi. xix. (mi).

Fig. 9. Fruit of Durio (upper left) and syncarps of Artocarpus subgen.

Artocorpus (upper right), subgen. Pseudojaca (lower right), and

Parartocarpus (lower left), diagrammatic, X i. The pulpy part

of each fruit is dotted, namely aril (Durio), pericarp ( Pararto -

corpus), perianth (subgen. Artocarpus), and the whole syncarp-

tissuc (subgen. Pseudojaca). Note, in subgen. Artocarpus. the

utricular perianths (perforated by the style) are fused in their

middle part, the strands between the fertile flowers being sterile

perianths; in Parartocarpus and subgen. Pseudojaca, the ovaries

are inferior in sockets of the receptacle and the strands betwe-n

the fruiting ovaries are receptacular tissue, in Artocarpus the

peltate interfloral bracts are shown.

207 ^

Gardens Bulletin, S~

It is necessary to argue this way because the durian-theory

(Corner, 1954) shows that, for the modern flowering plant (not

for a hypothetical pteridosperm ancestor), the arillate seed and

spinous pericarp are primitive features. To suppose that the

simple drupe of Moraceae is primitive and the syncarp a novel

durian-anologue is to beg the question; this reasoning implies that

Moreae never had the durian-features while Artocarpus dis¬

covered them anew only to lose them again in its subgen. Pseudo -

jaca which in all respects is advanced upon subgen. Artocarpus.

Sapindaceae show the derivation of spineless drupes from spiny

pericarps ( Nephelium ), and what are the sublamellate excres¬

cences on the indehiscent fruit of Planera (Ulmaceae) if not

durian-vestiges? That spines are a morphological entity, as vas-

culated epidermal outgrowths, the durian shows. That spines have

biochemical precursors in the genotype is shown by the action of

gall-insects which provoke durian-like galls in plants which seem

to have lost the ability to produce spines, e.g. Artocarpus cana -

rana Miq. which is galled Hopea wightiana (Dipterocarpaceae).

Recently in North Borneo I found a small shrub (sterile) which

bore a terminal pedunculate gall resembling a small durian-fruit

4-5 cm. long, but filled with flies. The day will come when the

biochemistry of durianology is revealed and genetics will

apprehend.

The lost history of Moraceae now begins to appear. It is not in

the advanced leptocaul states of Moreae but in the pachycaul

states of Artocarpeae, Olmedieae, Dorstenieae, and Ficeae. These

tribes are not derived from Moreae, but all from Proto-Moraceae.

Morus can be likened to sect. Ficus; behind Morus lies as much

lost history as there is present evidence in Ficus.

Affinity of Urticales

Several botanists as Weddell, Hallier (1905), Bessey, and Bechtel

have favoured affinity with Malvales. The Proto-Moraceous re¬

construction, Fig. 8, is a remarkable parallel with Euphorbiaceae

of the same alliance. Critical would be the microscopic structure

of the seed-coat, but all Moraceous and Urticaceous seeds seem

to be enclosed in endocarp (? transferred testa), and do not develop

typical testa-structure. Then Hallier (1912), Tippo (1939), and

Hutchinson (1959) refer Urticales to the affinity of Hamameli-

dales, which may lead back to a similar pachycaulous ancestor.

It is necessary to consider pachycaulous ancestry because, as

Moraceae show, family-characters, even tribal and generic, of

flower and fruit must have been evolved in this state of flowering

plant evolution. The pachycaulous is, in fact, the chief phase of

flowering plant evolution which, as would be expected from its

primitiveness, is widely missing.

208

Vol. XIX. (1962).

Geographical Table of Moraceous Genera and Species

Gen./Spp.

America

Africa

Asia and

Australasia

Tribes

10/68

4/19

6/13

4/36

Moreae

15/95

8/20

2/7

7/68

Artocarpeae

18/69

13/58

2/5

4/6

Olmedieae

8/44

«/6

7/38

. •

Brosimeae

1/120

1/80?

1/40?

i/i

Dorstenieae

1/1000?

1/150?

1/250?

1/600?

Ficeae

53/1397

27/333

18/353

17/711

Total Moraceae

Australasia has been included with Asia in this Table because

it has only one peculiar genus, namely Sparattosyce with two spe¬

cies in New Caledonia; it has also six species of Streblus along

with a few wide-spread species of Artocarpus, Antiaris, Fatoua,

and Malaisia.

Excluding Ficus and Dorstenia, as satisfactory genera, the ratio

of genera to species in America is 1/4, in Africa 1/3.7, and in

Asia 1/7.3. The number of genera in America and Africa may

well be halved, but these low ratios indicate that, excepting

Dorstenia and Ficus, the genera are relics.

Moraceae Lindl.

Veg. Kingd. (1847) 266; Bureau, DC. Prodr. 17 ( 1873) 211; Engl.

Nat. Pflanzenf. 3, 1 (1894) 66; Veg. Erde IX, vol. 3, 1 (1915) 17;

Dalle Torre et Harms Gen. Siph. (1900) 120; Lemee, Diet. Phan. 7

(1939) 184; 8b (1943) 8, 14, 17 .—Artocarpoceae Bur. DC. Prodr. 17

(1873) 280.—subfarn. Artocarpoideae A.Br. ex Aschers. FI. Prov.

Brandenb. 1 (1864) 57; Engl., l.c. 80; D. Torre et Harms, l.c. 121.—

subfarn. Moroideae Engl. l.c. 70; D. Torre et Harms, l.c. 120.

Trees, shrubs, climbers, or herbaceous ( Dorstenia, Fatoua).

Latex present, generally milky. Leaves pinnate, palmate, incised,

or simple, stipulate. Inflorescences axillary, typically paired, uni¬

sexual or bisexual, monoecious or dioecious, paniculate, racemose,

spicate, capitate, or urceolate. Flowers small, unisexual, apetal-

ous; tepals 8-0, generally 4, free or gamophyllous, imbricate or

valvate, persistent; stamens isomerous and antitepalous, or re¬

duced to 3, 2, or 1; filaments straight, free or connate, or inflexed;

anthers large, mucronate, to small, bilobate and non-mucronate,

varying crescentic to turbinate (with transverse, equatorial dehis¬

cence); pistillode present in male flower or not; ovary 1-, rarely

2-, locular, superior to inferior, or immersed in sockets in the

209

Gardens Bulletin , S ..

inflorescence-axis; styles 2 or 1 with two stigmatic arms, or one

stigmatic arm; stigma not capitate or peltate; ovule 1, anatropous

or campylotropous, apical or subapical. Fruits drupaceous, dis¬

crete or more or less connate in large fleshy syncarps. Seeds large

or small, invested in endocarp; testa membranous or disintegrated.

Embryos various , curved or straight; radicle long or short, in¬

cumbent or accumbent; cotyledons plicate, conduplicate, or plane,

foliaceous or thickened, equal to very unequal. Lamina with or

without cystoliths. Chromosome haploid numbers 12, 13, 14, with

intrageneric polyploidy. Gen. 53, pantropic, few temperate; spe¬

cies c. 1400 (Ficus c. 1000 spp.).

Key to the Tribes of Moraceae

1. Flowers inside urceolate receptacles (syconia), numerous; styles not

extruded from the receptacles. Blastophagous with sterile insect-in-

habited female flowers. Stamens, when 2 or more, with introrse

anthers; filaments straight in bud. Female flowers stalked or sessile

Ficeae.

1. Flowers not so enclosed or, if so, the styles exserted from the receptacle,

not blastophagous. Stamens mostly extrorse. Female flowers mostly

sessile.

2. Inflorescences bisexual, discoid or compressed-elongate, globose or

urceolate. Not spinous.

3. Female flowers numerous in each receptacle, each surrounded by

many sessile male flowers; stamens 1-3, mostly 2, the filaments

inflexed in bud; ovaries immersed in sockets. Receptacles flat¬

tened, discoid to variously elongate, bracteate round the margin,

forming syncarps. Seeds small. Herbs or suffrutescent

Dorstenieac..

3. Female flower solitary in the centre of each receptacle, surrounded

by many sessile male flowers; stamens 1-4, the filaments straight

in the bud or, rarely, inflexed; ovaries free or immersed. Recep¬

tacles discoid, globose, or urceolate, not elongate, forming syn¬

carps. Seeds rather large. Trees, shrubs . Brosimeae .

2. Inflorescences unisexual or, if bisexual, not discoid. Trees, shrubs,

climbers, rarely herbs.

4. Female inflorescence racemose or spicate with slender axis or 1-

flowered, or the inflorescence bisexual (cymose or spicate),

neither capitate nor discoid; ovaries mostly free, if inferior then

not in sockets. Male inflorescence paniculate, racemose, spicate

or capitate, often with a sterile groove; stamens 5-1, filaments

mostly inflexed in bud, in some cases straight; pistillode gene¬

rally present. Spinous or not . Moreae.

4. Female inflorescence capitate or thickly spicate, mostly syncarpous:

ovaries often immersed in sockets of the receptacle or connate

with it. Male flowers mostly without pistillode.

5. Male and female inflorescences discoid, obconic, or urceolate,

with an involucre of 1-several bracts; female inflorescences

varying 1-flowered. Syncarpous or not. Stamens 8-1, the fila¬

ments straight in bud. Not spinous. Olmediene.

5. Male and female inflorescences of different shape, involucrate or

not. Female inflorescence thickly spicate to capitate-globose,

never 1-flowered, syncarpous. Male inflorescence paniculate,

racemose, spicate. or capitate; stamens 4-1, the filaments

straight or inflexed in bud. Spinous or not .... Artocarpeae.

210

Vol. XIX. (1962).

Tribe Moreae Gaud.

Voy. Freyc. (1826); Endl. Prodr. FI. Norf. (1833) 40; Gen. PI.

(1841) 277; Suppl. 1, 1375; Suppl. 4 (1842) 33; Miq. Mart. FI. Bras.

4, 1 (1852) 154; Bur. DC Prodr. 17 (1873) 234; Bail!. Hist. PI. 6

(1875) 167; Benth et Hook. Gen. PI. 3 (1880) 343 (sub Urticaceae);

Engl. Nat. Pflanzenf. 3, 1 (1894) 72; Veg. Erde IX, 3, pt. 1 (1915);

Dalle Torre et Harms, Gen. Siph. (1900) 120.— Soroceac Miq.

Mart. FI. Bras. 4, I (1852) 111 (subtrib.); Bur. DC Prodr. 17

(1873) 288 .—Strebleae Bur. id. 215; Benth. et Hook. I.c. 344 (ut

subtribus); Dalle Torre et Harms l.c. 120.— Fatoueae Benth. et Hook.

I.c. 344 (ut subtribus); Engl. Nat. Pflanzenf. 3, 1 (1894) 71; Dalle

Torre et Harms, I.c. 120.— Eumoreae Benth. et Hook. I.c. 345.

Inflorescence paniculate, cymose. racemose, spicate, or (male)

capitate, (the axis not expanded into a discoid, urceolate, globose,

or stoutly spicate receptacle), bisexual or mostly unisexual. Fe¬

male inflorescence cymose, racemose, spicate, or 1-flowered, never

capitate. Filaments incurved or (Sorocea, Clarisia) straight in bud;

anthers extrorse; pistillode commonly present. Ovary generally

superior, inferior in a few genera. Fruit drupaceous, never in syn-

carps. Trees, shrubs, or ( Fatoua) herbs, spinous or not, monoeci¬

ous or dioecious. Gen. c. 10, species c. 68, pantropical or (Moms)

temperate.

Key to the Genera of Moreae

1. Seeds small, 1-2 mm. wide, somewhat compressed; endocarp crustace-

ous. Embryo with long, incumbent, transverse radicle; cotyledons

flat. Male flowers with pistillode; filaments 4, inflexed in bud.

2. Herb. Inflorescence cymose, bisexual or unisexual. Madagascar, Asia,

Australasia ..... Fatoua.

2. Trees. Inflorescence racemose or spicate, unbranched, unisexual. Dioe¬

cious or monoecious. Asia, North America. Moras.

1. Seeds 4-12 mm. wide, rounded. Radical not transversely elongate; coty¬

ledons often folded or much thickened, often unequal. Leaves dis¬

tichous. Spinous or not.

3. Male flower without pistillode. Filaments straight in bud. Female

perianth utricular, 4-dentate. Ovary (drupe) inferior. Unarmed or

with spinous-dentate leaf. America.

4. .Stamens 4, Male inflorescence racemose, spicate or ( Paraclarisia )

capitate .... Sorocea.

4. Stamen 1. Male inflorescence spicate . Clarisia.

3. Male flower with pistillode. Filaments (3-5, mostly 4) inflexed in bud.

5. Fruiting perianth fleshy as in Moras. Madagascar

Ampalis, Pachytrophe.

5. Fruiting perianth not fleshy, but the drupe thinly fleshy or with

thickly fleshy base.

6. Ovary (drupe) inferior. Unarmed. America, Malaysia, Mada¬

gascar .. Trophis.

6. Ovary (drupe) superior. .Spinous or not. Asia, Australasia,

(Africa ?) . Streblus.

(Africa, Neosloetiopsis, Sloetiopsis).

Too many genera on insufficient and invalid grounds trouble

this small tribe. I have reduced eleven to sections, or synonyms,

of Streblus, and I have considered reducing Streblus to Trophis ,

and all genera to Morus. The variety in the tribe is certainly much

less than that of Ficus, but insufficient knowledge and the difficulty

in assessing relic plants such as Fatoua , Morus, Ampalis, and

Pachytrophe render wholesale reduction at this stage unwise.

Gardens Bulletin,

The tribe represents the relics of that part of the family in which

the female inflorescence, though sometimes reduced to the one-

flowered state, is not specialised in form through thickening of

the axis.

Sexuality. The bisexual inflorescence of Fatoua and Streblus

sect. Bleekrodea has been used to separate these two genera into’

a tribe Fatoueae. The Madagascan species of Fatoua, however,

has unisexual inflorescences and sect. Bleekrodea merges through

Streblus tonkinensis (Teonongia Stapf) with S. asper. The inflo¬

rescences of S. tonkinensis vary from the condition of sect. Blee¬

krodea with the female flower surrounded by males to the condi¬

tion in S. asper , though monoecious, but Gagnepain has described

var. monoica of S. asper as having a female flower in the male

cluster. Then, because of its bisexual spike, Sloetia has been

placed in Dorstenieae, where it is entirely out of place. In flower,

fruit, seed, and leaf-structure Sloetia and Bleekrodea are closely

allied and both, by analogy with Ficus, are merely sections of

Streblus.

Sterile groove in male spike. As already noted, this feature oc¬

curs in all spicate inflorescences of Moreae and in Broussonetia,

Maclura, and Malaisia of Artocarpeae. It has yet to be explained

morphologically. It would seem to be a sign of affinity, yet the

racemose and paniculate inflorescence of Streblus sens, lat., Tro-

phis, and Maclura lack the sterile groove. In spite of its peculia¬

rity it is difficult to give it a primary value in classification.

Capitate male inflorescence. The inflorescence of shortly pedi¬

cellate male flowers in S. asper seems a good generic character

until it is realised that the insertion of a female flower into the

centre of it, as may happen in S. asper var. monoica, turns it into

the bisexual cyme of Streblus sect. Bleekrodea, with 5. tonkinensis

as the bridge. Streblus sect. Phyllochlamys has also a capitate

male inflorescence, but the fruit-characters ally the single species

with sect. Taxotrophis and indicate that the capitate male inflo¬

rescence is here a condensed raceme. S. crenatus (sect. Pseudotro -

phis) and 5. perakensis (sect. Paratrophis) have almost capitate

male inflorescences, but they have the sterile groove which indi¬

cates that they are derived from condensed spikes. It must be

concluded that the condensation of the male inflorescence has

occured independently in several lines of the genus.

Female inflorescence. Unlike the male, the female inflorescence

never condenses into a head; that is the character of Artocarpeae.

Instead, reduction leads to fewer flowers and ends with the solitary

pedunculate female flower, arrived in parallel in Streblus sect.

Streblus, Phyllochlamys, Pseudostreblus, Taxotrophis, Pseudotro-

phis, and Paratrophis, in Trophis sect. Maillardia, and, perhaps,,

in the African Neosloetiopsis.

212

Vol. XIX. (1962).

Fig. 10. Fruits in section, x 5. Streblus ( Bleekrodea) insignis (BI.) Cor¬

ner, left, with embryo. Streblus ( Sloetia ) elongatus (Miq.)

Corner, right, with thickened fruit-base; the lignified endocarp

hatched.

Pulpy fruit-base. The drupes of Streblus sect. Phyllochlamys,

Sloetia, and Taxotrophis have a strongly swollen base and they

actually dehisce crescentically over the apex in a plane at right

angles to that of the style. When this happens in S. elongatus, and

presumably in the other species, the seed is forcibly ejected by the

snapping together of the two sides of the swollen, white, edible

and sweet-tasting base. It seems a good character for a genus, but

there are transitions with slightly fleshy base as in sect. Bleekrodea

(drupe dehiscent at the apex), and in sect. Pseudotrophis and

Paratrophis in which the drupe is mostly, if not always, indehis-

cent. The drupelets of Ficus are often fleshy at the base and, in a

few species, dehisce at the apex to liberate the seed into the fig-

cavity, but the character has no classificatory value. Nevertheless,

it may be a primary distinction between Streblus and Morus,

though most species of Streblus seem to have lost it.

Pulpy tepals. Together with the small seed, the pulpy fruiting

tepals are the chief distinction of Morus. The Madagascan Am-

palis and Pachytrophe have also pulpy tepals, but this character

occurs in Maclura with which they seem more nearly allied. Pulpy

tepals occur, too, in many groups of Ficus, particularly sect. Ficus.

The character seems to contrast with that of Streblus, and this is

the main reason why I refrain from uniting Streblus with Morus.

213

Gardens Bulletin, S~

Pistillode. This feature offers three states which may be a basis

of classification:—

1. Pistillode quadrate-columnar; Morns , Ampalis, Pachy-

trophe; Streblus sect. Streblus, Taxotraphis, Pseudotrophis,

and Paratraphis; Traphis.

2. Pistillode minute, spicate, subulate, or conic; Fatoua ,

Neosloetiopsis, Slaetiopsis; Streblus sect. Bleekrodea, Pseu-

dastreblus, and Sloetia.

3. Pistillode none; Clarisia , Sorocea.

In Artocarpeae with pistillode it is the second kind, which

appears as the degeneration of the sterile ovary towards the final

state of complete absence. The quadrate-columnar state is that

typical of Urticaceae with inflexed stamens. Streblus, therefore,

seems to show stages in the loss of the pistillode.

Ampalis Bojer and Pachytrophe Bur.

Bojer, Hort. Maurit. (1837) 291; Bur. DC Prodr. 17 (1873) 234;

Leandri, FI. Madag. fam. 55 (1952) 6, 9.— Streblus Lour, subgen..

Parastreblus Bl. Mus. Bot. Lugd. Bat. 2 (1856) 80 (= Ampalis).

These two small genera of Madagascar resemble Streblus in

many ways but the fruiting tepals become fleshy as in Morus and,

indeed, as in Madura. Ampalis has the embryo of Madura f con¬

sisting of two small flat cotyledons and a rather long accumbent

radicle. Pachytrophe has an embryo like that of several species

of Streblus sect. Paratrophis, with foliaceous conduplicate coty¬

ledons and incumbent radicle, but this embryo also occurs in

Madura sect. Cardiagyne . The ovule of Pachytrophe is attached

to a much thickened placenta which becomes a woody plug at the

side of the seed, and this feature occurs in various Moraceae,

without mark of affinity, such as Broussonetia kurzii, Pararto-

carpus, Streblus salomanensis, Treculia , and Trophis brander-

horstii. It seems as if both genera are related with Madura , as

relics of its less specialised ancestry, and, if so, this will necessitate

a re-definition of Artocarpeae, or the fusion of it with Moreae.

Compare, however, Neaslaetiopsis.

Morus L.

The ovate-cordate lamina of Morus , with plicate vernation,

corresponds with the state of tree-evolution seen in Ficus subgen.

Sycomorus and in various groups of subgen. Ficus. From it the

simpler, inrolled, leaf of Streblus has been derived.

214

Vol. XIX. (1962).

M. macroura Miq. PI. Jungh. 1 (1851) 42.—M. laevigata Wall,

ex Hook. FI. Br. Ind. 5 (1888) 492.— M. alba L. var. laevigata

Bur. DC Prodr. 17 (1873) 245.

I can see no distinction whatever between these two species

and, therefore, reduce M. laevigata. The species becomes, then,

a wide-spread tree from Sikkim to Hainan with a relict status in

Sumatra, as a mountain tree, and, perhaps, in Java though it may

have been introduced to this country from Sumatra. Comparable

is the group of Ficus hirta in Ficus sect. Ficus.

Streblus Lour.

FI. Cochinch. (1790) 615; Blume. Mus. Bot. Lugd. Bat. 2, (1856)

79, f. 30; Miq. FI. In. Bat. 1, 2 (1859) 277; Bur. DC Prodr. 17

(1873) 218; Baill. Hist. PI. 6 (1875) 146, 195; Benth. et Hook. Gen.

PI. 3 (1880) 359; Engl. Nat. Pflanzenf. 3, 1 (1894) 78, f. 57 A-C.—

Achymus Vahl ex Juss. Diet. Sc. Nat. 1, suppl. (1816) 31.— Achimus

Poir. id. 5 (1827) 51.— Epicarpurus Bl. Bijdr. (1824) 488.— Al-

brandia Gaud. Voy. Freyc. Bot. (1826) 509.— Albradia D. Dietr.

Syn. PI. 5 (1852) 280.— Calius Blanco, FI. Filip. (1837) 698; Bur.

DC Prodr. 17 (1873) 21%—Bleekrodea Bl. Mus. Bot. Lugd. Bat. 2

(1856) 87; Bur. DC Prodr. 17 (1873) 254; Baill. Hist. PI. 6 (1875)

147, 198; Benth. et Hook. Gen. PI. 3 (1880) 358; Engl. Nat. Pflan¬

zenf. 3,1 (1894) 71, f. 50 C,D; Leandri, FI. Madagasc. fam. 55

(1952) 5.— Paratrophis Bl. Mus. Bot. Lugd. Bat. 2 (1856) 81; Bur.

DC Prodr. 17 (1873) 235; Baill. Hist. PI. 6 (1875) 143, 191; Benth.

et Hook. Gen. PI. 3 (1880) 364; Engl. Nat. Pflanzenf. 3, 1 (1894)

72; Cheeseman, Man. N. Zeal. FI. (1906) 631.— Taxotrophis Bl.

Mus. Bot. Lugd. Bat. 2 (1856) 77; Miq. FI. Ind. Bat. 1,2 (1859)

278; Bur. DC Prodr. 17 (1873) 216; Baill. Hist. PI. 6 (1875) 146,

195; Benth. et Hook. Gen. PI. 3 (1880) 358; Engl. Nat. Pflanzenf. 3, 1

(1894) 77; Hutchinson. Kew Bull. (1918) 147.— Sloetia Teysm. et

Binn. ex Kurz, J. Linn. Soc. Bot. 8 (1864) 168; Bur. DC Prodr. 17

(1873) 257; Baill. Hist. PI. 6 (1875) 147, 198; Benth. et Hook. Gen.

PI. 3 (1880) 365; Engl. Nat. Pflanzenf. 3, 1 (1894) 79 .—Diploeos

Bur. DC Prodr. 17 (1873) 215; Baill. Hist. PI. 6 (1875) 148. 199.

Phyllochlamys Bur. DC Prodr. 17 (1873) 217; Baill. Hist. PI. 6

(1875) 146, 196; Benth et Hook Gen. PI. 3 (1880) 359; Engl. Nat.

Pflanzenf. 3, 1 (1894) 77.— Pseudomorus (End!.) Bur. Ann. Sc.

Nat. ser. 5, 11 (1869) 372; DC Prodr. 17 (1873) 249; Baill. Hist.

PI. 6 (1875) 143, 191; Benth et Hook. Gen. PI. 3 (1880) 364; Engl.

Nat. Pflanzenf. 3, 1 (1894) 72; Skottsberg, Act. Hort. Gotob. 15

(1944) 347; Stearn, .1. Arn. Arb. 28 (1947) 426.— Pseudostreblus

Bur. DC Prodr. 17 (1873) 219; Bail!. Hist. PI. 6 (1875) 146, 195;

Benth. et Hook. Gen. PI. 3 (1880) 357; Hook. FI. Br. Ind. 5 (1888)

487; Engl. Nat. Pflanzenf. 3, I (1894) 71.— Uromorus Bur. DC Prodr.

17 (1873) 236.— Pseudotrophis Warb. Bot. Jahrb. 13 (1891) 294;

Engl. Nat. Pflanzenf. Nachtr. 1 (1897) 119. —Teonongia Stapf.

Hook. Ic. PI. 30 (1911) t. 2947; Gagnep. FI. Gen. I.-C. 5 (1928)

710, f. 86.— Dimerocarpus Gagnep. Bull. Mus. Hist. Nat. Paris 27

(1921) 441; Bull. Soc. Bot. Fr. 70 (1923) 204; FI. Gen. I.-C. 5

(1928) 704. —Diplothorax Gagnep. Bull. Soc. Bot. Fr. 75 (1928) 98;

FI. Gen. I.-C. 5 (1928) 705. —Chevalierodendron Leroy, Compt. Rend.

Ac. Sci. Paris 227 (1948) 146.— [B<dunostreblus Kurz = Sorocea, vid.

Jarrett, J. Arn. Arb. 39 (1958) 107.J.

215

Gardens Bulletin, S.

Trees, unarmed or spinous, monoecious or dioecious. Leaves

distichous, mostly oblong-elliptic to lanceolate, denticulate or en¬

tire, mostly short-petiolate, without glands. Inflorescences axillary,

bisexual or unisexual, pedunculate, varying cymose. racemose,

spicate, and capitate, or the female flower solitary. Male flower

with a pistillode, 3-5 merous, mostly 4-merous; tepals more or

less valvate, free or shortly joined; filaments inflexed in bud;

anthers small, reniform, extrorse. Female flowers 4-merous; tepals

decussate, imbricate, free or more or less joined; ovary superior.

Drupe rather large, 5-14 mm. wide, thinly fleshy, often with a

thickened fleshy base, dehiscent or not, the thin tepals investing

the drupe or not. Seed large, round, invested by the thin mem¬

branous endocarp; embryo various. Cystoliths amphigenous,

hypergenous, or none, not papillate except sect. Streblus; micros¬

copic gland-hairs various; hairs simple, aseptate, smooth except

sect. Streblus; stomata superficial. Species 22, Asia, Australasia,

(Africa ?); 13 in Malaysia.

As reconstituted, this genus represents the existing remains of

the Proto-Moraceae with most generalised inflorescence. Vegeta-

tively it has advanced to the applanate foliage with distichous

leaves. The inflorescences show every reduction to the spicate

condition, and the female to the one-flowered state. The fruit

seems to retain traces of dehiscence. Sterile the genus may be

confused with some groups of Ficus , but the leaves invariably

lack glands. Trophis, in the Old World, differs not only in flower

but in the hypogenous cystoliths. See, also, sect. Sloetia.

sect. Bleekrodea (Bl.) Corner stat. nov.— Bleekrodea Bl.

Unarmed, monoecious. Inflorescence bisexual, composed of 1-2

female flowers surrounded by more or less pedicellate male flowers,

solitary. Male flower with 4-5 tepals, gamophyllous in the lower

half; pistillode minute, conic. Female perianth utricular, minutely

4-dentate, investing the ovary; style and stigmatic arms long.

Drupe with a thinly fleshy base, dehiscent, enclosed in the en¬

larged utricular perianth; endocarp not or scarcely differentiated.

Cotyledons thick, fleshy, unequal, not folded; radicle short. Cys¬

toliths amphigenous, not papillate; microscopic gland-hairs 1-4-

celled, cruciate; hairs smooth, often hooked at the tip; epidermal

cells undulate-stellate. Species 2, Borneo, Madagascar. Type, S.

insignis (Bl.) Corner.—The inflorescence recalls Fatoua, but in

all other respects the section is near sect. Streblus.

sect. Paratrophis (Bl.) Corner stat. nov.— Paratrophis Bl.;

Pseudomorus (Endl.) Bur.; Uromorus Bur.; Chevalierodendron

Leroy.

216

VoL XIX. (1962).

Unarmed, dioecious. Male spike unbranched, with a sterile

groove; flowers sessile, 4-merous, 4-partite, valvate; pistillode

quadrate-columnar. Female spike many to 1-flowered; flowers in

2 rows, sessile. Bracts ovate-lanceolate or peltate-reniform. Drupe

without fleshy base, the tepals little enlarged (except S. peraken-

sis), indehiscent. Embryo curved; cotyledons equal, foliaceous and

conduplicate to small and flat, not much thickened; radicle rather

long, incumbent. Cystoliths hypergenous or none, not papillate;

microscopic gland-hairs various; hairs smooth; epidermal cells

generally polygonal (subundulate in S. solomonensis ); hypodermic

none or 1-celled ( S. glaber, S. perakensis, S. urophyllus). Species

9, Asia, Australasia. Type, S. heterophyllus (Bl.) Corner.

sect. Phyllochlamys (Bur.) Corner stat. nov.— Phyllochlamys Bur.

Twigs spinous. Dioecious. Male inflorescence capitate or very

shortly racemose, without interfloral bracts; flowers pedicellate,

4-merous, tepals gamophyllous in the lower half; pistillode qua¬

drate-columnar. Female flower solitary. Drupe with thick fleshy

base, ? dehiscent, much exceeded by the enlarged foliaceous tepals.

Cotyledons thick, fleshy, subequal; radicle short. Cystoliths hyper¬

genous, not papillate, or also scattered on the lower side of the

lamina along the veins; gland-hairs ?; prismatic crystals abundant

in the sheath of the veins; epidermal cells mainly polygonal;

hypodermis 1-celled. Species 1, Asia, S. taxoides (Heyne) Kurz.—

This section is very close to sect. Pseudotrophis and Taxotrophis.

sect. Pseudostreblus (Bur.) Corner stat. nov.— Pseudostreblus Bur.

Unarmed. Monoecious. Male inflorescence subspicate, branched,

with a sterile groove; flowers sessile to shortly pedicellate, 5—

merous; tepals free, imbricate; pistillode minute, conic. Female

flower solitary, pedunculate; tepals 4. Drupe not thickened at the

base, ? indehiscent, invested by the much enlarged tepals. Embryo

curved; cotyledons fleshy, unequal; radicle short. Cystoliths none;

microscopic gland-hairs clavate, multicellular; hairs smooth,

microscopic; epidermal cells undulate; hypodermis none. Species

1, Asiatic mainland, Hainan; S. indicus (Bur.) Corner.

sect. Pseudotrophis (Warb.) Corner stat. nov.— Pseudotrophis

Warb.; Dimerocarpus Gagnep.

Unarmed or with spinous twigs. Dioecious. Male inflorescence

spicate, unbranched, with a sterile groove; bracts ovate-lanceolate

to peltate-reniform; flowers sessile, 4-merous; pistillode quadrate-

columnar. Female inflorescences racemose, 8-1-flowered; flowers

more or less pedicellate. Drupe with thick fleshy base, dehiscent*

more or less covered by the enlarged tepals. Embryo curved;

217

Gardens Bulletin, S.

-cotyledons thick, fleshy, unequal, the smaller spathulate or cylin-

dric and surrounded by the large; radicle short. Cystoliths amphi¬

genous, not papillate; microscopic gland-hairs I-many celled, capi¬

tate; hairs smooth; epidermal cells polygonal to undulate; hypo-

dermis 1-celled. Species 3, Asia. Type, Pseudotrophis laxiflora

Warb. — S. ilicifolius (Vidal) Corner.

sect. Sloetia (Teysm. et Binn.) Corner stat. nov.— Sloetia Teysm.

et Binn.

Unarmed. Monoecious. Inflorescences spicate, bisexual or male,

with a sterile groove. Male flowers sessile, many, trimerous; pis-

tillode minute, spicate. Female flowers 0-4 per spike, 4-merous.

Drupe with thick fleshy base, dehiscent, more or less covered by

the enlarged tepals. Cotyledons fleshy, generally very unequal, but

varying equal; radicle very short. Cystoliths amphigenous, small,

not papillate; microscopic gland-hairs 4-8-celled, cruciate to capi¬

tate-peltate; hairs smooth; upper epidermal cells polygonal; hypo-

dermis none. Species 1, Malaya, Sumatra; S. elongatus (Miq.)

Corner.

sect. Streblus.— Achymus Vahl (Achimus Poir.); Albrandia Gaud.

(Albradia D. Dietr.); Calius Blanco; Epicarpurus Bl.; Teonon-

gia Stapf; Diplothorax Gagnep.

Unarmed. Monoecious or dioecious. Inflorescence bisexual and

capitate with a central sessile female flower surrounded by males,

or unisexual with the male capitate and the female as a solitary

pedunculate flower. Male inflorescence with few small bracts sur¬

rounding the capitulum. not between the flowers; tepals 4-5, free

or shortly joined; stamens 4 (-2); pistillode small, subconic,

columnar to quadrate, with 2 minute stigmata. Female flower

with 1-2 minute bracts at the base of the peduncle, sometimes

one on the peduncle, and 2 larger bracteoles at the base of the

perianth; tepals 4, free; style short, with 2 long stigmatic arms.

Drupe dehiscent or not, enclosed at first in the enlarged tepals,

without fleshy base. Embryo curved; cotyledons thick, fleshy, equal

to very unequal; radicle short, incumbent. Cystoliths amphigenous,

papillate as the hairs; microscopic gland-hairs 1-4-celled, capitate;

no crystal-cells in the epidermis; epidermal cells undulate-stellate.

Species 2, South-east Asia; type, S. asper Lour.

sect. Taxotrophis (Bl.) Corner stat. nov.— Taxotrophis Bl.;

Diplocos Bur.; Pseudotrophis Warb.

Twigs spinous. Dioecious. Male inflorescence racemose, un¬

branched, without a sterile groove; bracts ovate-lanceolate;

flowers 4-merous, pedicellate, often unevenly spaced; pistillode

quadrate-columnar. Female inflorescence shortly racemose, few-

flowered to I-flowered. Drupe without a fleshy base. Cotyledons

subequal, folded, radicle long; or cotyledons fleshy, unequal,

radicle short. Cystoliths ? Species 3, Asia. Type, S. spinosus (Bl.)

Corner = Taxotrophis javanica Bl.

218

Vol. XIX. (1962).

Key to the sections of Streblus

1. Inflorescence cymose, often condensed, bisexual, with 1-2 protogynous

female flowers in the centre; or the male inflorescence capitate (with¬

out a sterile groove) and the female flower solitary. Cotyledons very

unequal, one strongly thickened or both fleshy and equal; radicle

short.

2. Spinous tree. Fruiting tepals much exceeding the drupe with fleshy

base. Male inflorescence capitate; pistillode quadrate-cylindric.

Female flower solitary. Dioecious. Ceylon to Timor

Phyllochlamys.

2. Unarmed. Fruiting tepals equal to or shorter than the drupe.

3. Female perianth with free tepals. Inflorescence bisexual or the male

capitate and the female flower solitary; pistillode conic to colum¬

nar. Monoecius or dioecious. Ceylon to Celebes .... Streblus.

3. Female perianth utricular, covering the drupe but free. Inflores¬

cence bisexual. Pistillode minute conic. Madagascar, Borneo

Bleekrodea.

1. Male inflorescence elongate-racemose, simple or branched, or spicate.

Female flowers spicate or solitary.

4. Male inflorescence branched racemose-spicate, with a sterile groove;

flowers 5-merous; pistillode minute conic. Female flower solitary,

4-merous; tepals covering the large drupe. Cotyledons unequal,

fleshy. Monoecious, unarmed. India, China, Thailand

Pseudostreblus.

4. Male inflorescence unbranched; flowers 4-merous or 3-merous.

5. Spike bisexual with a sterile groove; male flower 3-merous; pistil¬

lode minute conic; female flower 4-merous. Drupe dehiscent

with fleshy base, covered by the tepals. Cotyledons equal, fleshy,

or one much reduced. Unarmed. Malaya, Riouw, Sumatra

Sloetia.

(Male flower 4-merous. East Africa. Sloetiopsis).

5. Spikes unisexual, dioecious. Flowers 4-merous. Pistillode quadrate-

columnar.

6. Male and female flowers pedicellate, the racemes without a ste¬

rile groove. Spinous. Ceylon to Flores . Taxotrophis.

6. Male flowers sessile, the male spike with a sterile groove.

7. Female flowers more or less pedicellate. Drupe fleshy at the

base, dehiscent, more or less covered by the enlarged tepals.

Cotyledons thick, unequal, one large and fleshy. Spinous or

not. India to Molucca Isl. Pseudotrophis.

7. Female flowers sessile, more or less distichous, or solitary and

pedunculate. Drupe not fleshy at the base, ? indehiscent,

mostly not covered by the tepals. Cotyledons various. Un¬

armed. Malaysia, Australasia . Paratrophis.

(West Africa; pistillode minute, spicate .. Neosloetiopsis).

Streblus sect. Bleekrodea

Key to the Species

Male flowers pedicellate, rather laxly clustered, equal to or exceeding the

female flower and fruit. Radicle very short. Stipules indurate, persis¬

tent. Lamina 8-21 X 3-8 cm. Borneo . S. insignis.

Male flowers subsessile, in clusters shorter than the female flower and

fruit. Radicle shortly elongate, incumbent on the smaller cotyledon.

Lamina generally small. Varying deciduous. Madagascar

S. rnadagascariensis.

S. insignis (Bl.) Corner comb. nov .—Bleekrodea insignis Bl.

Mus. Bot. Lugd. Bat. 2 (1856) 88, t.28. Type, Korthals, G.

Sakoembang, Borneo, at Leiden.

219

Gardens Bulletin , S.

S. madagascariensis (Bl.) Corner comb. nov.— Bleekrodea

madagascariensis BL id. 88; Leandri, FI. Madagasc. fam 55

(1952) 5.

Streblus sect. Paratrophis

Key to the Species

1. Bracts reniform or peltate. Male inflorescences often long, female often

many-flowered, never 1-flowered. Lamina more or less membranous,

usually without hypodermis. Micronesia, Melanesia, Polynesia.

2. Cotyledons flat, rather small; radicle incumbent. Lamina often sca-

brid, the basal nerves often as long as the lateral; cystoliths none.

Male spike 2-18 cm. long; female 5-22 mm. long, 2-9-flowered.

Monoecious or dioecious . S. pendulinus.

2. Cotyledons foliaceous, conduplicate, the radicle incumbent in the

sinus of the cotylcdpns. Lamina smooth, basal nerves shorter than

the lateral.

3. Male spikes 1-5 cm. long; female 6-25 mm. long, 3-8-flowered.

Drupe 4 mm. wide, red. Leaves often small; cystoliths abundant

on the upper side. New Zealand . S. heterophyllus.

3. Male and female spikes longer, many-flowered. Lamina often with

1-3 intercostals; cystoliths none.

4. Male and female spikes 10-20 cm. long. Lamina 9-30 X 3-15

cm.; lateral nerves 7-14 pairs; petiole 15-65 mm.

S. anthropophagorum.

4. Spikes -10 cm. long. Lamina 8-20 X 3-10 cm.; petiole 10-20

mm.

5. Stigmatic arms very short. Lateral nerves 8-11 pairs

S. solomonensis.

5. Stigmatic arms -1.5 mm. long, distinct.

6. Lateral nerves 10-15 pairs. Drupe black .... S. tahitensis.

6. Lateral nerves 6-8 pairs. Drupe red . S. smith'd.

1. Bracts ovate-lanceolate, sessile. Male and female spikes 10-25 mm.

long, often few-flowered, or the female flower solitary. Lamina nar¬

rowly elliptic to lanceolate, mostly thinly coriaceous and with a

hypodermis (1-cell thick); petiole 2-12 mm.; intercostals none.

7. Fruiting tepals much enlarged, covering the drupe; stigmata very short.

Male spike -10 mm. long. Female flower solitary. Cotyledons

somewhat thickened, conduplicate; radicle rather short. Cystoliths

hypergenous, sparse, or none. Prismatic crystals in the lower spider-

mis along the veins. Malaya . S. perakensis.

7. Fruiting tepals scarcely enlarged, not concealing the drupe. Male

spike 10-25 mm. long. Female spike 1-9-flowered. Cotyledons

flat, not conduplicate; radicle rather long, incumbent. Cystoliths

none. Small sphaerocrystals in the lower epidermal cells.

8. Lamina dentate, shortly acuminate, often stiffly coriaceous; lateral

nerves 6-13 pairs. Male flowers 3-7 per spike, 2.5-3.5 mm.

wide in bud, subglomerate. New Guinea . S. urophyllus.

8. Lamina entire or crenulate, rather long attenuate or acuminate or

caudate; lateral nerves 9-18 pairs. Male flowers often numerous,

smaller, not glomerate. Malaya to New Guinea and Queensland

S. glaher.

S, anthropophagorum (Seem.) Corner comb. nov.— Trophis

anthropophagorum Seem. FI. Vit. (1868) 258, t. 68.— Uro-

morus anthropophagorum (Seem.) Bur. DC Prodr. 17 (1873)

220

Vol. XIX . (1962).

236.— Paratrophis anthropophagorum (Seem.) Benth. et Hook,

ex Drake Ill. Ins. Mar. Pacif. f. 7 (1892) 296.— P. ostermeyri

Rechinger, P. viridissima Rechinger, P. zahlbruckneri Rechin¬

ger, Fedd. Rep. 5 (1908) 130.

Distr. Fiji, Samoa, Cook Isl., Niue.

The very long, many-flowered, male and female inflorescences

and the large leaves, often with rather long petioles, distinguish

this species. In Fiji and Samoa it varies into forms with shorter

inflorescences, which seem exactly intermediate with S. tahitensis;

such are the three species of Rechinger, described from Upolu

(Samoa), and not specifically distinct. To this complex belong S.

smithii and, possibly, S. solomonensis. They need field-study. For

instance, it is not clear if the long inflorescences and large leaves

are borne on saplings, and that they become smaller as the tree

grows larger. Does S. anthropophagorum become as large a tree,

—20 m. high, as S. solomonensis ?

This group of species suggests a Pacific centre for the genus

^treblus, whence the species have migrated and, in the Morace-

ous manner, become smaller in leaf and inflorescence, as they

have diverged. The genus has reached Hawaii but not, apparently,

the American tropics.

S. glaber (Merr.) Corner comb. nov.— Gironniera glabra Merr.

Philip. J. Sci. 1 (1906) suppl. 42.— Paratrophis glabra (Merr.)

v. Steen. J. Bot. 72 (1934) 8.— Chevalierodendron glabrum

(Merr). Leroy, Compt. Rend. Ac. Sci. Paris 227 (1948) 146.—

Aphananthes negrosensis Elmer, Leafl. Philip. Bot. 2 (1909)

575 -— Pseudostreblus caudatus Ridley, J. Fed. Mai. St. Mus. 6

(1915) 54.— Streblus laevifolius Diels, Bot. Jahrb. 67 (1935)

171.—( Excoecaria baccifera Elm. ined., Elmer 22014).

var. glaber

Distr. Malaya, Borneo, Philippine Isl., Celebes, New Guinea;

mountain forest, 700-2,500 m. alt.

var. australianus Corner var. nov.— Paratrophis australiana

C. T. White, Contr. Arn. Arb. 4 (1933) 15.

Alabastra mascula minora -1 mm. lata, plura. Lamina crenulata.

Distr. Queensland; Kajewski 1378, 1383, Herberton Range.

In leaf, inflorescence, and fruit, S. glaber is the most attenuate

species of the section and it has the widest range.

S. heterophyllus (Bl.) Corner comb. nov.— Epicarpurus microphyl-

lus Raoul, Ann. Sci. Nat. ser. 3, 2 (1844) 117.— Taxotrophis

microphylla (Raoul) F.v.M. Fragm. Phyt. Austral. 6 (1868)

193.— Paratrophis microphylla (Raoul) Cockayne, Bot. Notes

Kennedy’s Bush and Sci. Res. (1915) 3.— Paratrophis hetero-

phylla Bl. Mus. Bot. Lugd. Bat. 2 (1856) 81.

221

Gardens Bulletin, S.

var. heterophylla

Male spikes -25 mm. long; female spikes 6-12 mm. long, 3-8-

flowered. Drupes 1 (-3) per spike, 4 mm. wide. Lamina small,

-35 mm. long, the juvenile lamina pinnatifid to serrate.

Distr. New Zealand.

var. elliptica Kirk.— Paratrophis heterophylla Bl. var. elliptica

Kirk, Tr. N.Z. Inst. 29 (1897) 500, t. 46.— Trophis opaca Banks

et Sol. ex Hook. f. FI. Nov. Zel. 1 (1853) 224.—P. hanksii

Cheeseman, Man. N. Zeal. FI. (1906) 633.— P. opaca Druce,

Rep. Bot. Exch. Cl. Br. Isles (1916-17) 639.

Male spikes -5 cm. long; female spikes -25 mm. long, 8-25-

flowered. Drupes several per spike, -6 mm. wide. Lamina -8.5

cm. long, crenate-dentate.

Distr. New Zealand.

That this is only a variety of S. heterophyllus is shown by the

hybrids recorded by Allen as X Paratrophis micropaca (Gene-

tica 9, 1927, 145; id. 7, 1925, 290, f.3).

The combination S. microphyllus is antedated by Kurz’s syno¬

nym of S. taxoides.

S. pendulums (Endl.) F.v.M. Fragm. Phyt. Austral. 6 (1868) 192.

—Morus pendulina Endl. Prodr. FI. Norf. (1833) 40.— M.

brunoniana Endl. Atakta Bot. (1835) t. 32.— Streblus bruno-

nianus (Endl.) F. v. M. l.c. (1868) 192— Pseudomorus bruno¬

niana (Endl.) Bur. Ann. Sci. Nat. ser. 5, 11 (1869) 373.—

Pseudomorus sandwicensis Degener, FI. Hawaii. (1938) 21/22.

— Ps. brunoniana (Endl.) Bur. var. australiana Bur., v. obtu-

sata Bur., v. pendulina (Endl.) Bur., v. scabra Bur. l.c. (1869)

373.— Ps. brunoniana v. sandwicensis (Degener) Skottsberg,

Act. Hort. Gotob. 15 (1944) 347.— Ps. pendulina (Endl.)

Steam, J. Arn. Arb. 28 (1947) 427.

This is a variable plant. When young, it has large leaves with

10-16 pairs of lateral nerves. Larger trees have smaller leaves

with 7-12 pairs of lateral nerves. The larger leaves are ovate-

elliptic to ovate-oblong; the smaller are elliptic to elliptic-lanceo¬

late and less toothed. I doubt if the three or four varieties which

have been made are more than such differences in growth, though

some collections have scabrid leaves. Almost identical differences

occur in Malaisia scandens and are not to be distinguished in the

field. Of S. pendulinus I have seen 33 collections from the whole

range of the species and have been unable to draw any sharp line.

Leaves typical of P. sandwicensis occur on collections from Mic¬

ronesia and the side-twigs of these have the leaves of var. austra¬

liana; collections from Queensland may have the subacute leaves

of v. obtusata (New Caledonia). There is no sectional distinction

between Pseudomorus and Paratrophis.

222

Vol XIX . (1962).

Fig. 11. Streblus perakensis Corner. Leaves, X L A, coll. Forest Ranger

s.n., Chikus, 1933; B, Ridley 16171. Fruit and embryo in sec¬

tion, X 6; coll. For. Dept. 27338.

S. perakensis Corner sp. nov. Fig. 11.

Arbor parva glabra inermis, foliis distichis. Ramuli 1-1.5 mm.

crassi, fusco-brunnei. Stipulae 1.3-3.5 mm. longae, caducae.

Lamina 2.5-12 X 0.7-4 cm., anguste elliptica v. lanceolata v.

suboblongata, ad apicem mucronatum attenuata v. acuminata, ad

basim anguste cuneata v. anguste rotundata, integra v. subdenti-

culata, coriacea, laevis, sicco brunnea; costis lateralibus utrinsecus

8-15, patentibus, subtus elevatis, inarcuatis, intercostis nullis;

costis basalibus brevibus; petiolo 2-7 mm. longo. Inflorescentia

mascula axillaris 1-3, immatura -5 mm. longa, breviter pedun-

culata, uno latere sterilis, flores c. 6 sessiles secundos gerens;

bracteis 1 mm. longis, ovatis, obtusis; tepalis 4, extus puberulis;

staminibus 4; pistillodio minuto, 4-angulato. Flos femineus soli-

tarius, breviter pedunculalus. Drupa c. 7 mm. lata, tepalis de-

cussatis ovato-subacutis puberulis duobus eternis brevioribus

obtecta; pedunculo 4-9 mm. longo, bracteis 1-2 parvis basim

versus pracdito; tepalis externis 5-7.5 mm. longis, internis 8-10

mm.; stylo nullo v. brevissimo; stigmatibus 2 subulatis 0.7 mm.

longis; endocarpio tenuissimo subsclerotico. Embryon curvatum;

radicula brevi recta; cotyledonibus inaequalibus, minori a majori

implicato. Cystolitha nulla v. pauca hypergena.

Distr. Malaya (Perak, Pahang); For. Dept. 27338 (Chikus

Forest Res., leg C. L. Carrier, fructu; typus, herb. Singapore);

Forest Ranger s.n., Chikus Forest Res. March 1933, lelimau ,

floribus masculis; Ridley 16171, Kuala Teku.

223

Gardens Bulletin, S.

S . crenato armato comparanda, differt costis lateralibus pluri-

bus, stylo subnullo, drupa basim versus haud incrassata, coty-

ledonibus subaequalibus, cystolithis nullis v. sparsis.

S. smithii (Cheeseman) Corner comb. nov.— Paratrophis smithii

Cheeseman Trans. N.Z. Inst. 20 (1888) 148; Man. New Zeal.

FI. (1906) 631.

Distr. New Zealand.

This is near to S. anthropophagorum and S. tahitensis .

Fig. 12. Streblus solomonensis Corner. Leaf, X i. Flower, X 6. Fruit and

embryo, X 4. Coll. Kajewski 2396.

S. solomonensis Corner sp. nov. Fig. 44.

Arbor 20 m. alta, glabra, foliis distichis. Ramuli 1.5-2 mm.

crassi, fusco-olivacei dein pallide ochracei. Stipulae -6 mm. longae,

caducae. Lamina 8-12 X 3-9 cm., elliptica, saepe subasym-

metrica, apice -14 mm. longo acuminata, basi cuneata, integra,

membranacea, laevis, sicco griseo-viridis; costis lateralibus utrin-

secus 8-11, subtus vix elevatis; intercostis 1-3 laxis; costis basa-

libus brevibus; petiolo 10-20 mm. longo. Inflorescentia mascula ?

Inflorescentia feminea -15 mm. longa, in fructu -5 cm. longa,

axillaris 1-2, floribus sessilibus distichis; pedunculo 2-6 mm.

longo; bracteis 0.5-0.7 mm. longis, reniformibus subpeltatis,

minute fimbriatis, inferioribus multis sterilibus; tepalis 4 decus-

satis, 1 mm. longis, ovatis subacutis v. obtusis; stylo brevissimo;

224

Vol XIX. (1962).

stigmatibus 2, 0.5 mm. longis. Drupae 3-5 per infructescentiam,

7-8 mm. latae, tepalis vix amplificatis 1.5 mm. longis; endocarpio

ligneo tenui. Embryon curvatum; radicula recta longa; cotyledoni-

bus subaequalibus, plicatis. Cystolitha nulla.

Distr. Bougainville Isl., Guadalcanal Isl.; Kajewski 2006, Bou¬

gainville Isl., Koniguru, Buin, 800 m. alt., keru; Kajewski 2396,

Guadalcanal Isl., Berande River, torgapbagi (typus, herb. Kew).

This is said to be a common tree in rain-forest, the young

leaves being cooked and eaten, the timber used for houses. The

extremely short style and stigmatic arms distinguish it.

S. tahitensis (Nadeaud) Corner comb. nov.— Uromorus tahitensis

(Nadeaud) Bur. DC Prodr. 17 (1873) 237.— Pseudomorus

brunoniana var. tahitensis Nadeaud, En. PI. Tahiti, p. 43.—

Paratrophis tahitensis (Bur.) Benth. et Hook, ex Drake, Ill.

Ins. Mar. Pacif. f. 7 (1892) 296.

Possibly a variety of S. anthropophagorum. I have seen no

material.

S. urophyllus Diels, Bot. Jahrb. 67 (1935) 172.

var. urophyllus

Distr. New Guinea, 2,400-3,000 m. alt.

Closely allied with S. glaber and perhaps a high mountain state

of it.

var. salicifolius Corner v. nov.

Frutex 3 m. altus. Lamina 5-14 X 0.6-1.5 cm., oblongo-

lanceolata, acuta v. subacuminata, basi cuneata, denticulata v.

integra, coriacea; costis lateralibus utrinsecus 18-25 rectangulatis;

petiolo 3-7 mm. longo.

Distr. New Guinea; L. J. Brass 30220, Terr. New Guinea, East¬

ern Highlands, Mt. Wilhelm, east slopes, 3,000 m. alt., typus herb.

Leiden.

Streblus sect. Phyllochlamys

S. taxoides (Heyne) Kurz, For. FI. Burm 2 (1877) 465.— Trophis

taxoides Heyne ap. Roth, Nov. PI. Ind. Or. (1821) 368.— T.

taxijormis Spreng. Syst. Veg. 3 (1826) 902.— T. spinosa Roxb.

FI. Ind. 3 (1832) 762.— Epicarpurus timorensis Decne. Nouw.

Ann. Mus. Hist. Nat. 3 (1834) 499, t. 21.— E. spinosus (Roxb.)

Wight, lc. Ind. 6 (1835) 7, t. 1962 (partim).— Taxotrophis

roxburghii Bl. Mus. Bot. Lugd. Bat. 2 (1856) 78.— Phylloch¬

lamys spinosa (Roxb.) Bur. DC Prodr. 17 (1873) 218.— Stre¬

blus microphyllus Kurz, Prel. Rep. For. Pegu (1875) App. A,

p. cxviii; App. B, p. 84 (in clavi); For. FI. Burm 2 (1877) 464

(in clavi).—5. taxoides (Heyne) Kurz var. microphylla Kurz,

For. FI. Burm. 2 (1877) 465.— Phyllochlamys wallichii King,

Hook. FI. Br. Ind. 5 (1888) 489.— P. taxoides (Heyne) Kds.

225

Gardens Bulletin, S.

Exkursionfl. Java 2 (1912) 89.— P. taxoides var. parvifolia

Merr. Philip. J. Sci. 17 (1920) 247.— P. tridentata Gagnep. FI.

Gen. I.-C. 5 (1928) 714.— Taxotrophis poilanei Gagnep. id.

701.

Distr. Ceylon, India, Indochina, Thailand, Hainan, Malaya,

Java, Lesser Sunda Isl. to Timor, Celebes (S.E.), Mindoro,

Palawan.

I am indebted to Dr. Chew Wee Lek for the synonymy of this

species.

Streblus sect. Pseudostreblus

S. indicus (Bur.) Corner comb. nov.— Pseudostreblus indicus Bur.

DC Prodr. 17 (1873) 219.

Distr. Khasia Mts., North Thailand, Yunnan, Kwangsi-Kwang-

tung border, Hainan.

This seems a local or rare plant. I have seen merely six col¬

lections:—Hook. f. et Thompson (Epicarpurus 8), Kerr 6453,

Sino-Soviet Exp. (1957) 8137 (Yunnan), F. C. How 43790 and

73223 (Hainan), and W. T. Tsang 22641 (Kwangsi). The last

collection gives “fairly common in dry soil; fruit yellow”.

Streblus sect. Pseudotrophis

Key to the Species

1. Unarmed. Lamina membranous-chartaceous, acuminate, with 1-6 inter-

costals. Male spike 40-140 mm. long. Cotyledons somewhat un¬

equal. Twigs and lamina often drying yellowish. Cystoliths amphi-

genous; no crystal cells in epidermis. S. macrophyllus.

1. Spinous. Lamina coriaceous, acute or attenuate. Male spike shorter.

Cotyledons very unequal, the inner not folded.

2. Male spike 10-50 mm. long; female 2-6-flowered; bracts reniform.

Lamina 6-20 X 3.5-10 cm., often spinous-dentate. Cystoliths

hypergenous; single prismatic crystals abundant in the upper epi¬

dermal cells and along the veins beneath . S. ilicifolius.

2. Male spike 5-10 mm. long; female flower solitary; bracts ovate. La¬

mina 3—9 x 1—4 cm., acute, crenate-denticulate, base rounded to

cordate. Cystoliths amphigenous; crystal cells none in epidermis

S. crenatus .

(Unarmed. Leaf-base cuneate. Drupe-base not thickened. Cys¬

toliths none or few and hypergenous; crystal-cells along the veins

on the underside of the leaf. S. perakensis ).

S. crenatus (Gagnep.) Corner comb. nov.— Taxotrophis crenata

Gagnep. FI. Gen. I.-C. 5 (1928) 702, f. 82.

Distr. Indochina (Cambodia, Laos, Cochinchina).

This comes very near to S. taxoides, but the male spike, though

very short, has a sterile groove and the fruiting tepals are not

larger than the drupe. Nevertheless it seems impossible to distin¬

guish sterile plants and those with young female flowers.

226

Vol. XIX. (1962).

S. ilicifolius (Vidal) Corner comb. nov.— Taxotrophis ilicifolius

Vidal, Rev. PI. Vase. Filip. (1886) 249.— Pseudotrophis laxi -

flora Warb. Bot. Jahrb. 13 (1891) 294.— Taxotrophis obtusa

Elmer, Leafl. Philip. Bot. 5 (1913) 1813.— T. triapiculata Gam¬

ble, Kew Bull. (1913) 188.

Distr. Chittagong, Burma, Indochina, Thailand, Hainan,

Malaya, Philippine Isl., Celebes, Molucca Isl., Key Isl., Timor.

I am indebted to Dr. Chew Wee Lek for this synonymy. I have

also seen the type of P. laxiflora Warb., loaned from Berlin; it

is the more easterly state of S. ilicifolius with few or no spinous

leaf-teeth.

S. macrophyllus Bl. Mus. Bot. Lugd. Bat. 2 (1856) 80.— Diplocos

? macrophyllus (Bl.) Bur. DC Prodr. 17 (1873) 216 .-Para-

trophis mindanaensis Warb. Perk. Fr. FI. Philip. 1 (1904) 165.

— P. caudata Merr. Philip. J. Sci. 1 (1906) Suppl. 183.— Taxo¬

trophis mindanaensis Elm. Leafl. Philip. Bot. 5 (1913) 1815.—

T. balansae Hutch. Kew Bull. (1918) 151.— Dimerocarpus

brenieri Gagnep. Bull. Mus. Hist. Nat. Paris 27 (1921) 441.

Distr. Indochina, Malaya, Borneo, Celebes, Philippine Isl.

The types have been studied by Dr. Chew Wee Lek and myself.

The species has been reduced erroneously to S. ilicifolius.

Streblus sect. Sloetia

S. elongatus (Miq.) Corner comb. nov. Artocarpus elongatus Miq.

FI. Ind. Bat. Suppl. (1861) 172, 419.— Sloetia sideroxylon

Teysm. et Binn. ex Kurz, J. Linn. Soc. Bot. 8 (1864) 168, t. 13.

— SI. pinangeana D. Oliver, Hook. Ic. PI. (1886) t. 1531.— SL

wallichii King, Hook. FI. Br. Ind. 5 (1888) 493.

Distr. Malaya (Penang to Singapore), Riouw Archipelago,

Sumatra.

This well-known timber-tree of Malaya is strangely limited in

distribution and seems not to occur in Borneo. Its affinities are

also strange. The bisexual spicate inflorescence with few female

flowers is repeated only in the insufficiently known Sloetiopsis of

East Africa (4-merous male flowers; fruit unknown). If the male

and female flowers were on different inflorescences S. elongatus

would agree with sect. Pseudotrophis, as S. macrophyllus, except

for the minute conic pistillode and the 3-merous male flowers.

Neosloetiopsis (West Africa), also insufficiently known, may re¬

present this dioecious state, but it may be allied rather with

Ampalis. On the other hand, if the bisexual spike were contracted

into a head, the affinity would be with sect. Bleekrodea , except

for the perianth. Then, again, if the bisexual spike were extended

into a branched raceme, sect. Sloetia would be the antecedent con¬

dition to sect. Pseudostreblus. Thus, this timber-tree is not a

unique monotypic genus but a central species in the complex of

Streblus.

227

Gardens Bulletin, S.

Streblus sect. Streblus

Key to the Species

1. Lamina smooth, membranous, lanceolate-acuminate, base attenuate.

Monoecious, the male inflorescence with or without a female flower.

Male flowers with 4-5 tepals and stamens; pistillode quadrate. Fe¬

male tepals 3-4 mm. long (anthesis), ? the fruiting tepals not re¬

flexed; stigmatic arms 3-4 mm. long. Drupe dehiscent

S. tonkinensis.

1. Lamina typically rough, chartaceous-coriaceous, attenuate at the apex,

often narrowly rounded-cuneate at the base. Tepals 4. Pistillode

columnar, not quadrate, with 2 minute stigmata. Female tepals 2 mm. >

long (anthesis), reflexed in fruit; stigmatic arms lengthening 6-12

mm. in fruit. Drupe indehiscent Cotyledons very unequal .. S. asper.

2. Dioecious, rarely monoecious. Male head without a female flower;

stamens 4 . v. asper.

2. Monoecious. Male head often with a female flower in the centre.

Stamens 2 . v. monoica.

S. asper Lour. var. asper .—Diplothorax tonkinensis Gagnep. Bull.

Soc. Bot. Fr. 75 (1928) 98.

Distr. Ceylon, India, South China, Hainan, Indochina, Thai¬

land, Malaya (north), Sumatra (north), Java, Bali, Lombok,

Sumbawa, Celebes, Philippine Isl.

This species is absent from the Riouw-pocket of south Malaya,

south Sumatra, and, apparently, all of Borneo. I do not give the

full synonymy which is well-known. The type of D. tonkinensis

consists of a twig with male inflorescences and another with fe¬

male. The female perianth consists of four tepals, and is not utri-

culate as Gagnepain described.

var. monoica (Gagnep.) Comer stat. nov.— Streblus monoicus

Gagnep. Lee. Not. Syst. 14 (1950) 36.

This is intermediate with S. tonkinensis and comes from the

same region (Laos, Xieng-Kouang; Spire 176 and 200).

S. tonkinensis (Dub. et Eberh.) Corner comb. nov.— Bleekrodea

tonkinensis Dub. et Eberh. Compt. Rend. Ac. Sci. Paris 114

(1907) 631; Bull. Econ. Indoch. 10, 868-870 (ut Streblus sp.);

id, 13, 175 (ut Bleekrodea).—Teonongia tonkinensis (Dub. et

Eberh.) Stapf Hook. Ic. PI. 30 (1911) t. 2947; Gagnep. FI.

Gen. I.-C. 5 (1928) 710, f. 86.

Distr. Tonkin, along the Chinese frontier and in Ninh-Binh and

Hoa-Binh south of the Red River.

This is very like S. asper, with which Gagnepain partly confused

it. Indeed, in view of S. asper v. monoica, it is not clear how dis¬

tinct 5. tonkinensis may be. The joining of the inner tepals of

the female flower is more marked than in S. asper, but not a

specific difference.

228

Vol. XIX. (1962).

Streblus sect. Taxotrophis

Key to the Species

1. Female spike 2-6-flowered; style 0.5-1 mm. long, the stigmatic arms

2 mm. long. Bracts ovate. Fruiting tepals generally enlarged. Embryo

with long radicle and subequal, folded cotyledons. Lateral nerves

6-9 pairs.. S. zeylanicus.

1. Female flower solitary. Radicle short, the cotyledons very unequal.

2. Bracts ovate. Style 3-4 mm. long, the stigmatic arms 1-2 mm. Fruit¬

ing tepals not or little enlarged. Lamina drying yellow-brown;

lateral nerves 6-10 (-12) pairs . S. spinosus.

2. Bracts peltate. Style very short, the stigmatic arms 1 mm. long.

Fruiting tepals enlarged. Lamina often with spinous teeth; lateral

nerves 10-14 pairs . S. laxiflorus.

S. laxiflorus (Hutch.) Corner comb. nov.— Taxotrophis laxiflora

Hutch. Kew Bull (1918) 151.— T. eherhardtii Gagnep. FI. Gen.

I.-C. 5 (1928) 700.

Distr. Indochina, Malaya (Perlis, Kedah, Penang, Perak).

Examination of the type of T. eherhardtii by Dr. Chew Wee

Lek showed that it differed in no way from T. laxiflora. Whether

the pistillode is 3-lobed, 4-lobed, or practically entire, though

emphasized by Gagnepain, is immaterial.

S. spinosus (Bl.) Corner comb. nov.— Urtica spinosa Bl. Bijdr.

(1825) 507.— Taxotrophis javanica Bl. Mus. Bot. Lugd. Bat. 2

(1856) 77, t. 26.—T. spinosa (Bl.) v. Steen. Backer Bekn. FI.

Java 6 (1948) 9.

Distr. Sumatra, Java, Lesser Sunda Isl. to Timor.

S. zeylanicus (Thw.) Kurz, For. FI. Burm. 2 (1877) 464.—

Epicarpurus zeylanicus Thw. Kew J. 4 (1852) 1; id. 3 (1851)

t. 11. Taxotrophis zeylanica Thw. En. PI. Ceyl. (1864) 264.—

Diplocos zeylanica (Thw.) Bur. DC Prodr. 17 (1873) 215.—

Taxotrophis caudata Hutch. Kew Bull. (1918) 149.

Distr. Ceylon, India, Burma, China, Indochina.

I am indebted to Dr. Chew Wee Lek for the synonymy of this

species, which he has checked from the types.

Species Incertae Sedis

S. mitis Kurz, For. FI. Burm. 2 (1877) 464; Hook. FI. Br. Ind.

5 (1888) 489.

Unknown to Hooker, who wrote “probably not a Streblus”.

I have seen no specimen.

229

Gardens Bulletin, S.

Trophis P. Br.

Nat. Hist. Jamaica (1756) 357; Linn. Syst. Nat. ed. 10 (1759)

1289; Sp. PI. ed. 3, 2 (1764) 1451; Trecul, Ann. Sci. Nat. ser. 3, 8

(1847) 146; Bur. DC Prodr. 17 (1873) 251; Baill. Hist. PI. 6 (1875)

143, 192; Benth. ct Hook. Gen PI. 3 (1880) 365; Engl. Nat. Pflan-

zenf. 3, 1 (1894) 73; Engler, Bot. Jahrb. 40 (1908) 543, f.l; Woodson

and Schcry, Ann. Mo. Bot. Gdn. 47 (1960) 119, f. 37; ( nomen con-

servandum).—Bucephalon L. Sp. PI. (1753) 1190; ed. 3, 2 (1764)

1661; ( nomen rejiciendum).—Maillardia Frapp, et Duch. in Maillard,

Notes sur Tile Reunion (1863) annexe P, 3; Bur. DC Prodr. 17

(1873) 220; Leandri, FI. Madagasc. fam. 55 (1952) 15.— Calpido-

chlamys Diels, Bot. Jahrb. 67 (1935) 173.—? Skutchia Pax et Hoffm.

ex Morton, J. Wash. Ac. Sci. 27 (1937) 306.

Trees or shrubs, unarmed, dioecious. Leaves distichous. Stipules

paired, small, caducous. Inflorescences axillary, solitary or paired,

pedunculate, paniculate, racemose, or spicate and then with a

sterile groove; bracts very small. Flowers 4-merous. Male tepals

valvate, joined below; filaments inflexed in bud; pistillode quad¬

rate-columnar. Female tepals small, free; ovary inferior, the ovule

immersed in the receptacle; style short, with 2 long stigmatic arms.

Drupe inferior, the persistent tepals little enlarged, fleshy with

chartaceous to ligneous endocarp, indehiscent. Seed rounded to

oblong, 5-12 mm. wide, or the endocarp angled. Embryo straight

or curved; cotyledons fleshy, not folded, equal or subequal; ra¬

dicle very short to rather long, incumbent. Species 11, Trop.

America (3), Madagascar (5), Malaysia (3). Type; T. americana

Linn.

subgen. Prototrophis Corner subgen. nov.

Inflorescentiae paniculatae v. racemosae, floribus pedicellatis,

bracteis minutis. Embryo ?. Typus, T. mexicana Bur.

subgen. Trophis

Inflorescentiae spicatae, floribus sessilibus, masculae sulco ste-

rili praeditae, bracteis peltatis.

sect. Trophis. —Endocarpium chartaceum. Embryon rectum;

cotyledonibus crassis, subglobosis, aequalibus; radicula brevissima.

Spica feminea floribunda. Stigmata longiuscula, ex integro puber-

rula. Cystolitha amphigena. Typus, T. americana L.

sect. Calpidochlainys (Diels) Corner stat. nov.;—Endocarpium

ligneum, 0.2-0.3 mm. crassum, toro ligneo in loco hili obturatum.

Embryon curvatum; cotyledonibus crassiusculis, curvatis, haud

plicatis, subaequalibus; radicula longiuscula, incumbenti. Spica

feminea pauciflora. Stigmata brevia, unilateraliter papillosa.

Cystolitha hypogena. Typus, T. drupacea (Diels) Corner. Asia,

spp. 3.

sect. Maillardia (Frapp, et Duch.) Corner stat. nov.;—Endocar¬

pium ? tenue. Embryon ? curvatum.; cotyledonibus valde inae-

qualibus, majore plicato, alterum amplectenti; radicula brevissima.

Flores feminei solitarii v. binati. Cystolitha hypergena. Madagas¬

car, spp. 4; Reunion, sp. 1.

230

Vol. XIX . (1962).

Key to the species of sect. Calpidochlamys

1. Female inflorescence 15-40 mm. long, many-flowered. Endocarp not

angled. Lamina acuminate, base cuneate; lateral nerves 9-16 (20)

pairs . T. philippinensis.

L Female inflorescence 30-70 mm. long, with few fertile flowers. Endo¬

carp 3-angled, pyramidal.

2. Lamina acuminate, base rounded; lateral nerves 15 pairs

T. drupacea.

2. Lamina attenuate or subacuminate, base cuneate; lateral nerves 8-10

pairs . T. branderhorstii .

T. branderhorstii (Diels) Corner comb. nov.— Calpidochlamys

branderhorstii Diels, Bot. Jahrb. 67 (1935) 173.

Distr. Neth. New Guinea.

Possibly not separable from T. drupacea. I have examined the

type, Branderhorst 273.

T. drupacea (Diels) Corner comb. nov.— Calpidochlamys

drupacea Diels, Bot. Jahrb. 67 (1935) 173.

Distr. North-east New Guinea (Sepik-district).

I have found no specimen of this.

T. philippinensis (Bur.) Corner comb. nov.— Uromorus philip-

pinensis Bur. DC Prodr. 17 (1873) 237; Vidal Cat. PI. Prov.

Manila (1880) 43.— Paratrophis philippinensis (Bur.) F. Vill.

Noviss. App. Blanco FI. Filip, ed. 3 (1880) 198; Vidal, Phan.

Cuming. Philip. (1885) 145; Rev. PI. Vase. Filip. (1886) 250;

Merr. En. Philip. 2 (1923) 36.— Sloetia minahassae Kds. N.O.

Cel. Versl. (1898) 612, 645; Suppl. 2, t. 5; Suppl. 3, p. 3.—

Paratrophis grandifolia Elm. Leaf!. Philip. Bot. 5 (1913) 1814;

Merr. En. Philip. 2 (1923) 36.

Distr. Philippine Isl. (widespread), Celebes, Flores, Soemba,

Halmahera, New Guinea, New Britain.

Artocarpeae R. Br.

Bot. Congo (1818) 454; Gaud. Voy. Freyc. Bot. (1826) 511;

Bartl. Ord. Nat. (1830) 104; Endl. Gen. PI (1841) 278; Suppl. 1,

1375; Suppl. 4 (1842) 36; Miq. Mart. FI. Bras. 4, 1 (1852) 79, 81;

Bur. DC Prodr. 17 (1873) 281, 284; Baill. Hist. PI. 6 (1875) 167;

Benth. et Hook. 3 (1880) 343, 346 .—Broussonetieae Gaud. Voy.

Freyc. Bot. (1826) 508; Bur. DC Prodr. 17 (1873) 221; Benth. et

Hook. Gen. PI. 3 (1880) 345 (subtribus); Engl. Nat. Pflanzenf. 3,

1 (1894) 73; Dalla Torre et Harms, Gen. Siph. (1900) 120.—

Euartocarpeae Tree. Ann. Sci. Nat. ser. 3, 8 (1847) 77, 108; Benth.

et Hook. Gen. PI. 3 (1880) 347; Engl. Nat. Pflanzenf. 3, 1 (1894)

80; Dalla Torre et Harms, Gen. Siph. (1900) 121.

231

Gardens Bulletin, S.

Inflorescences unisexual (in a few cases bisexual), monoecious

or dioecious. Female inflorescence globose-capitate or thickly

spicate, many-flowered, syncarpous or not; involucre of one row

of bracts or none; ovaries free or immersed in sockets, or connate

with the receptacle. Male inflorescences paniculate, racemose,

spicate, or capitate, with or without a sterile groove; stamens

4—1; filaments straight or inflexed in bud; anthers extrorse; pistil-

lode present or not. Seed small or large. Embryos various. Trees,

shrubs, climbers, spinous or not. Gen. 15 (Asia 7, Africa 2,

America 8); spp, 95 (Asia 68, Africa 7, America 20).

Key to the Asiatic Genera of Artocarpeae

1. Female tepals 4, free (at least in the distal half), well-developed, de¬

cussate, imbricating, fleshy in fruit. Bracts and/or tepals with 2-7

immersed yellow glands. Seed compressed, with ligneous endocarp.

Dioecious, often spiny . Maclura.

1. Female perianth differently constructed, without immersed yellow

glands.

2. Male inflorescence raccmose-spicate, unbranchcd, with a sterile

groove; filaments inflexed in bud; pistillodc present. Female peri¬

anth utricular; ovaries not in receptacular loculi; styles long. Em¬

bryo curved. Dioecious, unarmed.

3. Syncarp globose, thickly set with slender stalked bracts of various

shapes more or less covering the drupes. Seeds 2-3 mm. long;

endocarp crustaceous to ligneous. Cotyledons equal, condupli-

cate to plane; radicle long, accumbent. Stipules membranous

Broussonetia.

3. Syncarp with few strongly projecting drupes each invested by the

utricular perianth, the bracts short. Seed 6-7 mm. long; endo¬

carp membranous. Cotyledons very unequal, the large thickly

fleshy and folded. Climber . Malaisia.

2. Male inflorescence clavate or capitate; if spicate, then without a

sterile groove and with the filaments straight in bud.

4. Male flowers 4-merous; filaments inflexed in bud; pistillode pre¬

sent. Female perianth reduced to 4 minute teeth; ovaries em¬

bedded in sockets of the receptacle; styles long. Embryo curved,

transversely elongate; cotyledons unequal, conduplicatc, fleshy;

radicle short. Dioecious spiny climbers . Plecospermum.

4. Male flowers with 4-1 stamens, the filaments straight in bud; pis¬

tillode none. Styles often short. Embryo elongate longitudinally.

Unarmed trees (? Prainea scandens),

5. Dioecious. Male and female heads similar, globose. Perianths

utricular; ovaries superficial. Stamen 1. Drupes projecting

strongly from the syncarp. Embryo straight; cotyledons thick

equal, plane; radicle very short . Prainea

5. Monoecious. Drupes embedded in the more or less fleshy syn¬

carp.

6. Male heads usually of different shape from the female, vary¬

ing spicate to clavate or capitate; flowers superficial; sta¬

men 1. Female perianth utricular, much reduced, or none;

ovaries sunk in receptacular sockets or concealed beneath

the layer of connate utricular perianths. Embryo straight

or slightly curved; cotyledons thick, equal or unequal; ra¬

dicle short ...... A rtocarpus.

6. Male and female heads similar, globose, distinctly involucrate

with bracts. Stamens 2-3, sunk like the ovaries in recepta¬

cular sockets, the filaments and style projecting between

the free conic or truncate tepals. Embryo curved; radicle

rather long, ascending. Parartocarpus.

232

Vo\. XIX. (1962).

Artocarpus J. R. and G. Forster

Char. Gen (1776) 101, t. 51; Jarrett, J. Arn. Arb. 40 (1959) 113-

368; id. 41 (1960) 73-140.

Tiiis genus presents the extreme reduction of the male inflore¬

scence as a spike or head of minute unistaminate flowers with

tubular perianth and minute anther. The female inflorescence re¬

tains a well-developed, if unusual, utricular perianth in subgen.

Artocarpus, but the perianth disappears more or less entirely in

subgen. Pseudojaca. The ovaries are superior in subgen. Arto¬

carpus , though hidden from the outside by the connate middle

parts of the perianths, but they become inferior in sockets of the

receptacle in subgen. Pseudojaca. The fruiting perianths of sub¬

gen. Artocarpus become fleshy, at least in their proximal parts

surrounding the ovaries, thus making the false aril around the

false seed. In subgen. Pseudojaca the pulpy character of the fruit

is transferred to the receptacular tissue which forms, thus, the

whole pulpy body of the false fruit. Ripe syncarps of subgen.

Artocarpus must be tom open to eat the false aril; those of sub¬

gen. Pseudojaca can be bitten directly, as an apple. This kind of

syncarp resembles that of Plecospermum, but there are so many

differences between it and Pseudojaca in other respects that the

two must be regarded as parallel conclusions in Moraceous fruits.

The state of Pseudojaca is presumably derived by simplification

from that of Artocarpus by loss of the free internal parts of the

perianths followed by loss of the free external parts. Detailed

developmental studies of these female inflorescences are needed

in a variety of species to clarify the exact construction of the

syncarps.

Broussonetia L’Herit. ex Vent.

Tabl. Regn. Veg. 3 (1799) 547 (gen. conserv.); Bl. Mus. Bot.

Lugd. Bat. 2 (1856) 85; Miq. FI. Ind. Bat. Suppl. (1861) 415; Bur.

DC Prodr. 17 (1873) 223; Baill. Hist. PI. 6 (1875) 143, 192, f.

102-107; Benth. et Hook. Gen. PI. 3 (1880) 361; Engl. Nat. Pflan-

zenf. 3, I ( 1894) 76, f. 54, 55 .—Broussonetia Ortega, Nov. PI. Descr.

(1798) 61, t. 7.— Papyri us Lam. Ill. Gen. PI. (1798) t. 762.— Al-

laeanthus Thw. Hook. J. Bot. Kew Gard. Misc. 6 (1854) 302, t. 9b;

Bur. DC Prodr. 17 (1873) 222; Benth. et Hook. Gen. PI. 3 (1880)

361; Hook. FI. Br. Ind. 5 (1888) 490; Engl. Nat. Pflanzenf. 3, 1

(1894) 74 (ut Allacanthus ).—Smithiodendron Hu, Sunyatsenia 3

(1936) 106.

Dioecious unarmed trees, shrubs, or climbers, often (? always)

deciduous. Leaves spirally arranged, opposite, or distichous,

palmately lobed to undivided, generally cordate to rounded or

widely cuneate at the base, dentate to entire, membranous; inter-

costals numerous; stipules membranous. Bracts and perianth with¬

out immersed yellow glands. Male inflorescence racemose to spi-

cate, with a sterile groove; bracts ovate to clavate, truncate, or

umbonate, the interfloral bracts generally stalked, those bordering

the sterile groove sessile; flowers shortly pedicellate to sessile;

233

Gardens Bulletin, S.

tepals 4, joined in the lower half, valvate; filaments 4, inflexed

in bud; pistillode spicate, minute. Female inflorescence capitate,

globose, without a sterile groove; bracts as in the male, becoming

stipitate in fruit; perianth utricular with 2-4 small lobes or teeth;

ovary sessile or becoming stipitate in fruit (sect. Broussonetia );

style 1 (-2), long. Syncarp globose, 10-25 mm. wide, the flowers

not connate, ripening orange-red, more or less pulpy. Drupes

thinly pulpy, invested by the thinly pulpy, yellow to red, perianth.

Seeds 1.7-2.5 mm. long, ovoid-compressed, faintly keeled; endo-

carp crustaceous to ligneous, smooth or asperate. Embryo curved;

cotyledons equal, thinly fleshy, conduplicate to flat; radicle long,

incumbent. Cystoliths hypergenous, hypogenous, or none, often

as cystolith-hairs; microscopic gland-hairs 1-3-celled, rather

large, capitate; hairs smooth or papillate, often hooked (especially

microscopic hairs); sphaerocrystal-cells in the lower epidermis

along the veins, or none. Species ? 7.

sect Broussonetia

Drupe stipitate within the sessile perianth. Seed slightly com¬

pressed, papillate-asperate, crustaceous, the keel double at the

base. Cotyledons flat. Cystoliths hypergenous or none. Leaves

spirally arranged to distichous. Spp. 4; type, B. papyrifera (L.)

Vent.

sect. AUaeanthus (Thw.) Corner stat. nov.— Allaeanthus Thw.

Drupe sessile. Seed compressed, smooth, ligneous, the keel not

double. Cotyledons conduplicate or flat. Cystoliths hypogenous.

Leaves distichous. Male bracts often ovate, sessile. Spp. 3; type,

B. zeylanica (Thw.) Corner.

There are no major differences between these sections, which

are not generically distinct. There appears to be an undescribed

species from West Borneo, represented by the sterile collections

NIFS bb 8015 and 8016; possibly it belongs to sect. Allaeanthus ,

but its leaves have no cystoliths.

Key to the Species of sect. Allaeanthus

1. Climber. Syncarp -10 mm. wide. Seeds with the endocarp much thick¬

ened into a lip at the hilar end. Cotyledons conduplicate corrugated.

Lamina crenate-dentate; lateral nerves 3-6 (-8) pairs; stipules

entire. Male spike -5 cm. long . B. kurzii.

1. Trees. Syncarps -20 mm. wide. Seeds without the thickened lip to the

endocarp. Lateral nerves 7-15 pairs.

2. Lamina serrate; stipules denticulate. Male spike -6 cm. long. Coty¬

ledons ? corrugate . B. zeylanica.

2. Lamina and stipules entire. Male spike 10-26 cm. long. Cotyledons

flat . B. luzonica.

B. kurzii (Hook, f.) Corner comb. nov.— Allaeanthus kurzii

Hook. f. FI. Br. Ind. 5 (1888) 490; Kurz, For. FI. Burm. 2

(1877) 466 (ut Malaisia tortuosa); Gagnep. FI. Gen. I.-C. 5

(1928) 721.

234

Vol. XIX . (1962).

Distr. Assam, Upper Burma, Yunnan, Laos, Cochinchina,

Thailand.

This large deciduous climber is at once distinguishable from

Malaisia by the numerous close intercostals.

B. luzonica (Blanco) Bur. DC Prodr. 17 (1873) 224,— Moms

luzonica Blanco, FI. Filip. (1837) 703.— Broussonetia luzoni-

ensis Blanco, FI. Filip, ed. 2 (1845) 488.— Allaeanthus luzoni-

cus (Blanco) Vidal, Noviss. App. (1880) 198.

var. luzonica

Pubescent.—Philippine Isl. (Luzon).

var. glabra (Warb.) Corner comb, nov.— A. glaber Warb. Perk.

Fragm. FI. Philip. 1 (1904) 166.— A. luzonicus (Blanco) Vidal

var. glaber (Warb.) Merr. En. Philip. 2 (1923) 37.

Puberulous, soon glabrous.

Distr. Luzon, Basilan, Mindoro, Celebes (Gorontalo, Moena).

B. zeylanica (Thw.) Corner comb. nov.— Allaeanthus zeylanicus

Thw. Hook. J. Bot. Kew Gard. Misc. 6 (1854) 302, f. 9b;

Worthington, Ceylon Trees (1959) pi. 425.

Bureau described the cotyledons of this species, which is known

only from Ceylon, as strongly corrugated-foliaceous. It seems that

they must resemble those of B. kurzii, but all the seeds of the

collections which I could examine, including the type, were empty.

Madura Nutt.

Gen. Amer. PI. 2 (1818) 233 (gen. conserv.); Bur. DC Prodr. 17

(1873) 227; Benth. ct Hook. Gen. PI. 3 (1880) 363; Engl. Nat.

Pflanzenf. 3, 1 (1894) 74; Lemee, Diet. Phan. 4 (1932) 230.—

Vanieria Lour. FI. Cochinchin. (1790) 564.— Toxylon, Joxylon Raf.

Am. Month. Mag. (1817) 118 and (1818) 188, 19 5.—Chlorophora

Gaud. Voy. Freyc. Bot. (1826) 509.— Fusticus Raf. New FI. Am. 3

(1836) 43.— Sukaminea Raf. id. 44.— Cudrania Tree. Ann. Sci. Nat.

ser. 3, 8 (1847) 122 (gen. conserv.); Bur. Ann. Sci. Nat. ser. 5, 11

(1869) 377; DC Prodr. 17 (1873) 285; Bail!. Hist. PI. 6 (1875) 145.

194; Benth. et Hook. Gen. PI. 3 (1880) 374; Hook. FI. Br. Ind. 5

(1888) 538; Engl. Nat. Pflanzen. 3, 1 (1894) 82, f. 60B, C; Renner,

Bot. Jahrb. 39 (1907) 361.— Cudranus Rumph. ex Miq. FI. Ind.

Bat. I, 2 (1859) 290; Rumph. Herb. Amb. V, 22, t. 15, 16.— Cardio-

gyne Bur. DC Prodr. 17 (1873) 232.— Milicia Sim, For. FI. Port. E.

Africa (1909) 97, t. 72.

Dioecious trees, shrubs, climbers, generally with axillary spines.

Leaves spirally arranged to distichous, simple, without glands.

Male inflorescences paniculate, spicate with a sterile groove, or

capitate. Female inflorescences capitate or shortly spicate. Bracts

and/or tepals with 2-7 immersed yellow glands. Bracts spathulate

to clavate or none. Male flowers 4-merous; tepals free; anthers

small, extrorse to laterally dehiscent; filaments inflexed or straight

in bud; pislillode subulate or as a slender sterile ovary or none.

Female flowers sessile, free or connate proximally; tepals 4, de¬

cussate, obtuse, free or gamophyllous to the middle, fleshy in fruit;

235

Gardens Bulletin, S.

ovary free or sunk in sockets in the receptacle; stigmatic arms 1-2,

short or long. Drupes very thinly pulpy, enclosed in the fleshy

perianth, generally not projecting, often concrescent below into a

fleshy syncarp. Seed compressed, with more or less ligneous en-

docarp 2.5-10 mm. long. Embryo curved; radicle long, asym¬

metrically incumbent to accumbent; cotyledons equal, folded or

not, rather thin. Cystoliths amphigenous or none; microscopic

gland-hairs 4-celled, capitate; generally with sphaerocrystal cells

in the lower epidermis, at least along the veins; hypodermis none;

hairs smooth. Species 12, America, Africa, Asia.

Key to the Sections of Maclura

1. Filaments straight in bud. Style or stigmatic arms short, 1-2.5 mm.

long. Male heads capitate. Spines axillary, not on short leafy shoots.

Cotyledons folded or flat. Cystoliths none. Trees, shrubs, climbers.

Asia, Australasia . sect. Cudrania.

1. Filaments inflexed in bud. Style or stigmatic arms 8-20 mm. long.

2. Male heads capitate. Ovaries immersed in sockets of the receptacle.

Style undivided. Syncarp fleshy. Cotyledons much folded. Cysto¬

liths none. Spines ending short leafy shoots, or decurved with 1-2

scale-leaves. Climber. Africa . sect. Cardiogyne.

2. Male heads not capitate. Ovaries not in sockets. Cotyledons flat, small.

Trees.

3. Male inflorescence paniculate, ebracteate; flowers pedicellate. Stig¬

matic arms 1-2. Syncarp depressed subglobose, 8-14 cm. wide.

Cystoliths none. Generally spiny. America .... sect. Maclura.

3. Male inflorescence spicate with a sterile groove; flowers sessile.

Style undivided. Syncarp globose to shortly oblong, smaller.

Cystoliths amphigenous. Spinous or not. America, Africa

sect. Chlorophora.

sect. Maclura.— Maclura subgen. Eumaclura Bl. Mus. Bot. Lugd.

Bat. 2 (1856) 82.

Tree, generally spinous. Inflorescences apparently ebracteate.

Male inflorescence paniculate; flowers pedicellate; tepals free; fila¬

ments inflexed in bud; pistillode ? none. Female tepals with 2

immersed yellow glands; ovaries not in sockets of the receptacle;

stigmatic arms 1-2, 9-20 mm. long. Syncarp massive, fleshy, the

drupes immersed. Seeds 9-10 mm. long. Cotyledons flat, thin;

radicle obliquely incumbent. Cystoliths none. Species 1, M. ponii-

fera (Raf.) Schneid. ( M. aurantiaca Nutt.).

sect. Chlorophora (Gaud.) Bur. DC Prodr. 17 (1873) 228.—

Chlorophora Gaud., Milicia Sim.

Trees spinous or not. Inflorescences bracteate; bracts and tepals

with immersed yellow glands. Male inflorescence spicate with a

sterile groove; tepals free; filaments inflexed in bud; pistillode as

a small flattened ovary. Ovaries not embedded in sockets of the

receptacle; style undivided, 8-10 mm. long. Syncarp -20 mm. wide.

Seeds 2.5-4 mm. long. Cotyledons flat, thin; radicle accumbent.

Cystoliths amphigenous, often sparse below. Species 5, America,

Africa. Type, M. tinctoria (L.) Don.

America:— M. tinctoria (L.) Don.

236

Vol. XIX . (1962).

Africa:— M. excelsa (Welw.) Bur.

M. regia (A. Chev.) Corner comb. nov.— Chlorophora

regia A. Chev. Bull. Soc. Bot. Fr. 58 (1912) 209.

Madagascar:— M. greveana (Baill.) Corner comb. nov.—

Ampalis greveana Baill. Grandid. Hist. Madag. PI. (1891) t.

293A.— Chlorophora greveana (Baill.) Leandri, Mem. Inst. Sci.

Madag. ser. B, 1 (1948) 18; FI. Madag. fam. 55 (1952) 12, f. ITT

1 - 8 .

M. humberti (Leandri) Corner comb nov.— Chloro¬

phora humberti Leandri, l.c. (1948) 20; l.c. (1952)

14, f. Ill 9.

This section needs further resolution. The American species is

generally spiny and has globose female head and syncarp; there

may be two species. The African and Madagascan species are

unarmed. M. greveana has a globose female head and syncarp,

but the other three have distinctly oblong female heads and syn-

carps. In M. excelsa, too, the lamina is less reduced than in other

species of the genus and has 14-18 pairs of lateral nerves. Chlo¬

rophora was first sunk in Maclura by Endlicher (Gen. PI. 1841,

277).

sect. Cardiogyne (Bur.) Corner stat. nov.— Cardiogyne Bur.

Climber with short leafy shoots ending in a spine, or the de-

curved axillary spine with 1-2 scale-leaves. Inflorescences brac-

teate; tepals with 2 immersed yellow glands. Male inflorescence

capitate; flowers sessile; filaments inflexed in bud; pistillode ligu-

late. Female perianths 4-partite, gamophyllous below and connate

laterally at the base; ovaries sunk in sockets of the receptacle;

style undivided, -18 mm. long. Syncarp -20 mm. wide, the drupes

concealed. Seed 5-6 mm. long. Cotyledons much folded; radicle

obliquely incumbent. Cystoliths none. Species 1, Africa.

M. africana (Bur.) Corner comb. nov.— Cardiogyne ajricana

Bur. DC Prodr. 17 (1873) 232.

sect. Cudrania (Tree.) Corner comb. nov.— Cudrania Tree.,

Vanieria Lour., Cudranus Rumph. ex Miq.— Maclura Nutt.

subgen. Leptosura Bl. Mus. Bot. Lugd. Bat. 2 (1856) 83.

Trees, shrubs, climbers, with leafless axillary thorns in place of

short shoots. Male and female inflorescences capitate, both brac-

teate; bracts subulate, lanceolate, spathulate, to subpeltate, with

2 immersed yellow glands, often 2-4 bracts per flowers, often

adnate to the tepals, usually several sterile bracts at the base of

the capitulum. Tepals free or gamophyllous in the lower half,

with 2-7 immersed yellow glands in each. Male flowers with the

filaments straight in bud; pistillode subulate. Female perianths

free or connate in the lower part; ovaries free or embedded in

sockets of the receptacle; style short, undivided or with two short

237

Gardens Bulletin, S.

arms. Syncarp fleshy, -40 mm. wide, the drupes generally con¬

cealed. Seeds 4-9 mm. long. Cotyledons broad with infolded marg¬

ins and obliquely incumbent radicle, or small and flat with ac-

cumbent radicle. Cystoliths none. Species 5, Asia, Australasia.

Type, Maclura cochinchinensis (Lour.) Comer.

Ser. Liberae Corner ser. nov.

Flores feminei liberi, ovario superficiali, stylo saepissime bi-

lobato. Flores masculi filamentis quam antheris exsertis bis usque

quater longioribus. Semina 5-9 mm. longa. Radicula accumbens.

Species 2. Typus, M. fruticosa (Roxb.) Corner.

Ser. Connatae Corner ser. nov.

Flores feminei partibus inferioribus connati, tepalis basim versus

gamophyllis, ovario in cavo parvo receptaculi immerso, stylo in-

diviso raro ramulo dentiformi praedito. Flores masculi filamentis

quam antheris brevioribus v. vix longioribus, antheris vix exsertis.

Semina 3-5 mm. longa. Cotyledones marginibus inflexis, radicula

eorumdem uno lateri incumbenti. Species 3. Typus, M. cochin¬

chinensis (Lour.) Comer.

Key to the species of sect. Cudrania

1. Female flowers free; ovary superficial; style mostly with 2 stigmatic

arms. Filaments 2-4 times as long as the exsert anthers

ser. Liberae.

2. Tepals free; style less than 0.5 mm. long; stigmatic arms thick, blunt;

filaments flattened, anther minute. Syncarp -15 mm. wide, irre¬

gularly lobate from the separately projecting fruits. Seed 5-9 mm.

long; endocarp 0.3 mm. thick. Cotyledons subequal, rather thick.

Twigs, petioles, puberulous to glabrous. Lamina long acuminate,

base cuneate; lateral nerves 4-5 pairs, oblique, scarcely inarching;

basal nerves 1/3-2/3 lamina; intcrcostals 3-8, regular

M. jruticosa.

2. Perianth tubular in the lower half; style 1.5 mm. long; stigmatic

arms slender, as long; filaments cylindric; anthers 1.5 mm. long.

Syncarp -4 cm. wide, fleshy, evenly subglobose. Seed 7-8 mm.

long; endocarp 0.6-0.7 mm. thick, very hard. Cotyledons rather

thin. Twigs, petioles, underside of veins fulvous brown tomentose.

Lamina obtuse to acute, base rounded or subcordate; lateral nerves

4-6 pairs, inarching; basal nerves not elongate; intercostals 3-5,

faint ..«. M. thorelii.

1. Female flowers connate at the base, the perianth tubular in the lower

part; ovaries in sockets of the receptacle; style simple, rarely with a

minute branch. Filaments shorter or slightly longer than the not or

scarcely exsert anther. Syncarp 15-25 mm. wide, fleshy, evenly sub-

globose. Seed 3-5 mm. long; endocarp 0.2 mm. thick. Cotyledons

thin, with infolded edges; radicle obliquely incumbent. Basal nerves

of lamina short ..ser. Connatae.

3. Lateral nerves inarching strongly, 6-9 (-14) pairs; intercostals none

or indistinct; petiole 3-15 mm. Tepals closely villosulous, all

with 2 immersed glands. Filaments very short. Climber

M. cochinchinensis.

3. Lateral nerves ascending, slightly inarching; intercostals 2-5, distinct.

Filaments 1-11 times as long as the anthers. Petiole often longer.

238

Vol XIX. (1962).

4. Tree or shrub. Lamina-base rounded to widely cuneate; lateral

nerves 3-5 pairs, often very oblique; petiole -35 mm. long.

Male heads 7-10 mm. wide; tepals with 2 immersed glands

M. tricuspid at a.

4. Climber. Lamina-base cuneate; petiole -20 mm. long. Male heads

5-7 mm. wide; tepals (male) thinly puberulous, generally with¬

out immersed glands . M. amboinensis.

5. Lateral nerves 6-9 pairs . v. amboinensis.

5. Lateral nerves 4-6 pairs . v. paucinervia.

M. amboinensis Bl. Mus. Bot. Lugd. Bat. 2 (1856) 84.—

Cudrania amboinensis (BL) Miq. FI. Ind. Bat. 1, 2 (1859) 290

(ut Cudranus). — C. grandifolia Merr. Philip. J. Sci. 18 (1921)

52.

var. amboinensis

Lateral nerves 6-9 pairs.

Distr. Thailand, Malaya, Sumatra, Java, Borneo, Celebes, Am-

boina (type, leg. Zippel), New Guinea.

var. paucinervia Corner var. nov.—C. grandifolia Merr.

Costae laterales utrinsecus 4-6.

Distr. Philippine Islands, New Guinea. Typus, Elmer 15530

(Luzon, Mt. Bulusan, prov. Sorsogon, det. Cudrania javanensis

Tree.; herb. Kew).

M. cochinchinensis (Lour.) Corner comb. nov.— Vanieria cochin -

chinensis Lour. FI. Cochinch. (1790) 564.— Cudrania javanen¬

sis Tree. Ann. Sci. Nat. ser. 3, 8 (1847) 123, pi. 3, f. 76-85.

—C. obovata Tree. id. 126.— Maclura javanica BL, M. timorensis

Bl. Mus. Bot. Lugd. Bat. 2 (1856) 83.—C. cochinchinensis

(Lour.) Kudo et Masumune, Ann. Rept. Taihoku Bot. Gard. 2

(1932) 272.—C. cambodiana Gagnep. Bull. Soc. Bot. Fr. 72

(1925) 808; FI. Gen. I.-C. 5 (1928) 729, f. 89, 90.

Distr. Ceylon, India, China, Japan to New Britain, Australia,

and New Caledonia.

There are several other well-established synonyms of this com¬

mon species.

var. pubescens (Tree.) Corner comb. nov.— Cudrania pubescens

Tree. Ann. Sci. Nat. ser. 3, 8 (1847) 125.

Twigs and underside of leaves brown hairy or velvety, but with

all transitions to the glabrous state of v. cochinchinensis.

var. bancrofti Bailey, Compr. Cat. Queensl. PI. (1910) 504.

Foliage variegated.

M. fruticosa (Roxb.) Corner comb. nov.— Batis fruticosa Roxb.

FI. Ind. 3 (1832) 763.— Cudrania fruticosa Wight ex Kurz,

For. FI. Burm. 2 (1877) 434; Hook. FI. Br. Ind. 5 (1888)

539.— Vanieria fruticosa (Wight) Chun, J. Arn. Arb. 8 (1927)

21 .

239

Gardens Bulletin, S.

This species is remarkable in two ways. The very tips of the

flattened filaments are contracted and slightly inflexed, as a ves¬

tige of the state with normally inflexed filaments. Then, as many

of the female flowers seem sterile or do not fruit, the drupes pro¬

ject individually from the infructescence, each in its own decussate

fleshy perianth, simulating the syncarp of Malaisia.

M. thorelii (Gagnep.) Corner comb, nov.— Cudrania thorelii

Gagnep. Bull. Soc. Bot. Fr. 72 (1925) 810; FI. Gen. I.-C. 5

(1928) 725.

This species approaches ser. Connatae in the tubular lower half

of the perianth. It is remarkable for the large, strongly exsert an¬

thers, 1.5 mm. long, with filaments 3-4 mm. long, and for the

large syncarp, —4 cm. wide. If there were 2—3 stamens, instead of

4, the species would be truly artocarpoid, but the flattened seed

with thick endocarp and the embryo place it in Maclura.

M. tricuspidata Carr. Rev. Hort. (1864) 390, f. 37; id. (1872)

56, f. 7.— Cudrania triloba Hance, J. Bot. 6 (1868) 49; Hook.

Ic. PI. (1888) t. 1792.—C. tricuspidata (Carr.) Bur. Lavallee

Arb. Segrcz. (1877) 243.— Morus integrifolia Lev. et Van. Bull.

Ac. Interm. Geogr. Bot. 17 (1907) 210; Rehder, J. Arn. Arb.

17 (1936) 73.— Vanieria tricuspidata (Carr.) Hu, J. Arn. Arb. 5

(1924) 228.— V. triloba (Hance) Satake, J. Fac. Sci. Tokyo

sect. 3, Bot. 3 (1931) 497.

The inflorescences and syncarp of this species seem identical

with these of M. cochinchinensis. It seems that many twigs, parti¬

cularly the upper ones, are unarmed. As a silk-worm tree it is

said to be as good as Morus but too thorny for convenient

picking.

Macludrania hybrida Andre (Rev. Hort. 1905, 362) is said

to be a hybrid between M. pomifera and this species. The few

sterile specimens that I have seen in herbaria could not be dis¬

tinguished from M. pomifera.

Malaisia Blanco

FI. Filip. (1837) 789 .—Dumartroya Gaud. Voy. Bonite (1844)

t. 97 .—Cephalotrophis Bl. Mus. Bot. Lugd. Bat. 2 (1856) 75, t. 27.

The position of this monotypic genus is problematic. The male

flower and embryo resemble those of Streblus sect. Sloetia. The

female flower resembles that of Streblus sect. Bleekrodea. The

male inflorescence fits Streblus but the contracted female inflores¬

cence, which is not exactly capitate, and the fleshy syncarp, parti¬

cularly, agree rather with Prainea. Furthermore, the yellow resin-

cells of the drupe-wall in Malaisia agree with the yellow glands

of Maclura and the resin-cells of Artocarpus. The genus seems

a relic of the ancestors of Prainea and Artocarpus.

240

Vol XIX. (1962).

The sole species is M. scandens (Lour.) Planch. The second

species, which is generally dragged in without enquiry, is M. pube-

rula (Miq.) Bur., but after examining the type I can find no other

difference than the 4-merous male flower, which seems at most

varietal. Similar variation occurs in Streblus sect. Sloetia.

Par art near pus Baill.

Adansonia 11 (1875) 294; Jarrett, J. Arn. Arb. 41 (1960) 320.

This genus is well-named because it shows the durian-factors

developed in parallel with those of Artocarpus. In Parartocarpus

the free bracts and tepals become the spines of the syncarp, not

the tips of the utricular perianths; it is the ovary-wall or exocarp

of the drupe which becomes the false aril of the false seed, not.

the proximal parts of the perianth. In fact, the drupe of Pararto¬

carpus in size and succulence, with well-developed endocarp, is

more primitive than any drupe or drupelet of Moraceae. The

straps” between the drupes of Artocarpus subgen. Artocarpus

are the undeveloped perianths of unfertilised or sterile flowers; in

Parartocarpus they are straps of receptacular tissue between the

flowers as in subgen. Pseudojaca. The large anthers and 2-3

stamens indicate, also, a less advanced state than Artocarpus.

Thus Parartocarpus has advanced to the sunken ovaries of sub¬

gen. Pseudojaca while remaining less advanced in other respects.

It is an off-shoot of pre-Artocarpus which has attained the syn¬

carp of subgen. Pseudojaca by parallel evolution.

Plecospermum Tree.

Ann. Sci. Nat. ser. 3, 8 (1847) 124, pi. 4, f. 121-126.

Spiny dioecious climbers, some of the twigs modified into leaf¬

less spines. Leaves distichous. Inflorescences capitate, pedunculate;

bracts small, lanceolate-obtuse, both bracts and tepals without

immersed glands; flowers sessile, 4-merous. Male perianth shortly

gamophyllous; filaments inflexed in bud; pistillode filiform or as

p minute ovary. Female heads with the ovaries immersed singly

in sockets of the receptacle; tepals minute, round the mouth of the

socket; ovary free from the receptacular wall; styles 1-2, long,

undivided. Syncarp fleshy, bulging from the included seeds, with

many aborted flowers. Seeds 7-9 mm. long, plump; endocarp

membranous. Embryo large, curved; radicle very short; cotyledons

very unequal, both folded and fleshy, the larger surround¬

ing the smaller, both with long stalks. Cystoliths none. Species 2

f-3). India, Ceylon, Burma, Indochina, Thailand, Andaman Isl.

This is another problematic relic, related perhaps with Malaisia.

The two species resemble Maclura sect. Cudrania in climbing

habit, leaf, and spines, but the seed and embryo are those of

Malaisia , and the inflorescences are an advance on this genus

towards Artocarpus subgen. Pseudojaca. Experimental hybridisa¬

tion may elucidate the affinity.

241

Gardens Bulletin, S.

Key to the species of Plecospermum

Glabrous or finely puberulous. Lamina 5-8 cm. long. Style 1, 3-5 mm.

long. Syncarp 10-15 mm. wide, finely brown velvety . . P. spinosum.

1 wigs, stipules, young leaves, inflorescences, and syncarp brown tomen-

tose. Lamina 6-14 x 3—8 cm. Styles 1 or 2, 10-13 mm. long. Syn-

carps 15-30 mm. wide . P. andamunicum.

P. andainanicum King, Hook FI. Br. Ind. 5 (1888) 491—

Cudrania poilanei Gagnep. Bull. Soc. Bot. Fr. 72 (1925) 809;

FI. Gen. I.-C. 5 (1928) 724 (excl. Poilane 10038, floribus mas-

culinis = Maclura cochinchinensis).

Distr. Laos, Annam, Cochinchina, Burma, Thailand, Andaman

Isl.

To judge from herbarium-material, this seems to be deciduous.

P. spinosum Tree, l.c.; Hook. FI. Br. Ind. 5 (1888) 491; Alston,

Kandy FI. (1938) 34, f. 183.— Trophis spinosa Heyne, Willd.

Sp. PI. 4 (1806) 734.— T. aculeata Roth, Nov. Sp. PI. (1821)

368.— Batis spinosa Roxb. FI. Ind. 3 (1832) 762.

Distr. Ceylon, India.

Evergreen ?

P. bureaui Richter, Termeszetrarzi Futezek 15 (1895) 296.

This is unknown to me.

Species Excludendae

P. cuneifolium Thw. is Madura cochinchinensis (Lour.) Comer.

P. laurifolium Bail!, is Pachytrophe ohovata Bur. var. laurifolia

Leandri, Mem. Inst. Sci. Madag. ser. B, 1 (1948) 16; FI.

Madag. fam. 55 (1952) 11, f.2.

P • obovatum Bur. ex Richter is Pachytrophe obovatum Bur.

Prainea King

Hook. FI. Br. Ind. 5 (1888) 546; Jarrett, J. Arn. Arb. 40 (1959) 30.

This genus agrees with Artocarpus in the capitate male and

female inflorescences, the tubular perianths, the single stamen

with straight filament and small anther, and the short stigmatic

arms. Yet it is dioecious and the syncarp, so unlike that of Arto¬

carpus, resembles closely that of Malaisia. Further, both Prainea

and Malaisia have membranous endocarp. The perianth of Prainea

has many clusters of yellow resin-cells in its mesophyll, such

as occur in the ovary (pericarp) wall of Malaisia, but not,

so it seems, in the floral parts of Artocarpus. On the other hand,

Prainea has the peltate (4-8-celled) microscopic gland-hairs and

mesophyll resin-cells of subgen. Artocarpus and it lacks the cyslo-

liths of Malaisia; this last point, however, is of very doubtful value

and there are very rudimentary cystoliths along the underside of

the veins in some species of Prainea. A microscopic character of

Prainea is the strongly stellate form of the epidermal cells of the

leaf.

242

Vol. XIX. (1962).

A peculiarity of Prainea, described by Jarrett, lies in the man¬

ner in which the ovule is moved from a subapical position in the

ovary at anthesis to a sub-basal position in fruit. It is possible,

therefore, that the embryo is developed transversely inside the

ovule though longitudinally in relation to the ovary-cavity, such

as happens in Plecospermutn. Parartocarpus has the same pecu¬

liarity, but it differs in so many ways from Prainea that the

similarity seems to be parallel.

According as one or other character is emphasized, so Prainea

inclines to Malaisia or to Artocarpus, and retains its own character.

Treculia Decne ex Tree.

Ann. Sci. Nat. ser. 3, 8 (1847) 108.

This genus, even its best known species T. africana, requires

much more investigation. It is placed near Artocarpus , but it

lacks the characteristic female perianths of subgen. Artocarpus

and it does not have the sunken ovaries of subgen. Pseudojaca.

The slender uinbonate-peltate bracts of the syncarp much resem¬

ble those of Broussonetia. T. africana is said to be dioecious but

solitary specimens in Malaya fruit and seed; their fruit-heads are

strongly protogynous and then develop copious male flowers with

dehiscent stamens and, apparently, viable pollen, while the seeds

are forming; some small heads are wholly male. As a monoecious

and partly monoclinous plant it has the character antecedent to

Brosimeae. The male flower with 2-4 stamens and large anthers

resembles that of Parartocarpus but retains a utricular perianth.

The drupe resembles that of Parartocarpus, but the embryo is

orientated the other way round, having the radicle away from the

raphe-side of the seed. The genus may be a parallel with Arto¬

carpus, but, to show how inadequate published descriptions may

be, there seems from the literature little to choose between the

African Acanthotreculia, generally sunk in Treculia, and subgen.

Pseudojaca.

Olmedieae Tree.

Ann. Sci. Nat. ser. 3, 8 (1847) 126; Miq. Mart. FI. Bras. 4, 1

(1852) 79; Bur. DC Prodr. 17 (1873) 281; Benth. et Hook. Gen. PI.

3 (1880) 346; Engl. Nat. Pflanzenf. 3, 1 (1894) 83; Dalla Torre et

Harms. Gen. Siph. (1900) 121.

Inflorescences unisexual, discoid, obconic, or urceolate, with an

involucre of 1-several rows of bracts, the male many-flowered,

the female many to 1-flowered; flowers sessile. Stamens 8-1;

filaments straight in bud; anthers introrse or extrorse; pistillode

present or not. Ovaries superficial or immersed, lnfructescence

syncarpous or not. Seeds large. Embryo various. Monoecious or

dioecious. Trees, shrubs, not spinous. Gen. 18 (America 13, Africa

2, Asia 4); species 69 (America 58, Africa 5, Asia 6).

243

Gardens Bulletin , S.

Antiaris Lesch.

Ann. Mus. Hist. Nat. Paris 16 (1810) 478, t.22 (gen. conserv).

Seventeen species have been described. After studying the

abundant collections in London, Paris, Leiden, Singapore, and

Bogor, I have reduced the number to four, namely A. africana,

A. toxicaria, and the two Madagascan species A. humbertii and

A. madagascariensis. Several of the proposed species are clearly

synonyms, but the variability of the wide-spread A. toxicaria is

the real cause of the superfluity and, indeed, of the uncertainty.

On broad principles only one species may be recognised because

there are no essential differences, merely variations of size and

shape, in leaf and fruit. Thus, the small leaf distinguishes A . hum¬

bertii; the ovate-elliptic, tapered leaf may distinguish A. toxicaria

var. macrophylla; the prominent venation distinguishes A. afri¬

cana; but, without fruit, A. madagascariensis cannot be distin¬

guished from A. toxicaria . These distinctions are supported

geographically and ecologically. A. africana belongs to the drier

forests of west and central Africa, where its timber is recognised

as different from that of A. toxicaria. The Madagascan species

belong to the xerophytic scrub on calcareous and sandstone rock.

In contrast, A. toxicaria is a primary rain-forest tree of the low¬

land hylea and, in this capacity, has the widest and most continu¬

ous range of any Moraceous plant, from West Africa to South

China and Fiji. It is absent from Madagascar and other Mascarene

islands, from the Australian mainland, and from New Caledonia.

It reaches 1,500 m. alt. in North Borneo and New Guinea and

its occurrence in South China suggests a certain adaptability to a

seasonal climate. On the periphery of its range occur the other

species.

The nearest allies of the genus seems to be the American

Olmedia and Pseudolmedia, both with a single superficial ovary,

more or less gamophyllous perianth, and distinctly exsert anthers.

Ogcodeia differs more in having several flowers in the female in¬

florescence. The little known Mesogyne of Africa may be another

species of Antiaris, but its stamens are extrorse and the ovary is

partly superior. Though placed by Engler in Dorstenieae, the fila¬

ments of Mesogyne are straight in the bud and the inflorescences

accord well with Antiaris.

Habit. A. toxicaria is a large tree. The form cultivated in the

Solomon Islands, Fiji, and Tonga is a shrub or small tree. Its

leaves and large fruits place it in var. macrophylla, collections of

which from New Guinea and New Hebrides seem to show all

gradations to large trees. Perhaps, nevertheless, there is a dwarf

form. A. africana is also a large tree. The Madagascan species

are small trees or shrubs. The evidence of field-notes shows that

all the species have intermittent growth and that, as adult plants,

they become deciduous.

244

Vol. XIX. (1962).

Saplings. The saplings of A. toxicaria have brown-hispid or

setose-scabrid leaves and twigs. The leaves are larger, thinner,

more obovate, more attenuate to the base, and more toothed than

the adult leaves which, with the twigs, seem to become more

shortly hairy and, finally, practically glabrous. Nevertheless, there

may be persistently hairy and persistently less hairy varieties. A.

rufipila Miq. was based on a fertile, brown-hispid specimen. The

sapling form seems identical in A . africana. In both the adult

foliage becomes coriaceous, but the feature is much more pro¬

nounced in A. africana. Since specimens of saplings are com¬

monly collected for identification, they make the wide use of A.

toxicaria desirable.

Fruit. The size and shape of the fruit, or drupe (omitting the

stalk), is important. A. africana and A. toxicaria have ellipsoid

or pyriform drupes, the size of which increases from Africa to

Polynesia. There is some overlap but all African collections have

small drupes, 10-15 X 7-12 mm. when dried, compared with

the Far Eastern drupes, 20-45 X 17-26 mm., when dried. In

western Malaysia there is considerable variation but no fruits so

large as the eastern. Celebes seems the boundary. Here are trees

with small fruits indistinguishable from the African var. welwits-

chii, trees with large fruits and leaf-shape approaching var. macro-

phylla, and others exactly resembling the intermediate var.

toxicaria. Celebes is also a meeting ground for east-west variation

in figs, e.g. F. tinctoria. The Madagascan species have nearly

spherical drupes almost as large as those of A. toxicaria var.

macrophylla; and this shows that they are not the same species

as on the African mainland. Now, in Ficus as a rule, the further

a species is from its geographical centre of alliance the smaller

are its fig-fruits. Applying this rule to Antiaris, A. toxicaria must

have spread westwards from eastern Malaysia and Australasia

and to this far eastern centre of alliance the Madagascan species

relate. The progressive xerophily of the leaf in A. toxicaria con¬

forms with this westward trend. Yet, Antiaris has no close ally

in eastern Malaysia, and it is absent from New Caledonia, the

figs of which are predominantly allied in subgen. Pharmacosycea

with those of tropical America. Nevertheless, as Trophis shows

the alliance between America and Polynesia or Eastern Malaysia,

so may Antiaris at the end of its evolutionary course.

The fruit-variation is summarised in the accompanying table.

Two collections from Soemba are without fruit and cannot be

placed exactly. One collection from east Flores (Solor Isl.) is sterile

but appears to be var. macrophylla. Of four collections from

Timor, two belong to var. macrophylla (N1FS bb. 20,062, and

27,073), the others being without fruit.

245

Gardens Bulletin , S.

Table of Fruit-sizes in Antiaris

(Measurements refer to the dried drupe, excluding the stalk)

—

Drupe (mm.)

Country

Leaf-shape

A. africana

10-15 x 9-10

Africa

as in var.

toxicaria

A. toxicaria

v. welwitschii

10-15 X 7-12

Africa

do.

v. toxicaria

15-25 x 11-20

Continental

Asia to

Celebes

obovate,

acuminate

12-20mm. long

Celebes

do.

v. macrophylla

20-25 x 17-20

Moluccas

Ovate-

elliptic.

attenuate

25-38 x 20-26

New Guinea

do.

30-45 x 22-25

Solomon TsI.

Fiji

do.

20-24 x 17-19

Philippines

do.

A. humbertii

20-30 x 20-30

Madagascar

as in var.

toxicaria

A. madagascariensis

do.

Toxicity . There is much conflicting evidence on the toxicity of

the latex and seeds of A. toxicaria. That many trees have poison¬

ous latex, used for arrow-poison in West and Central Malaysia,

is well known. The poison is caused by cardiac glycosides, parti¬

cularly antiarin. Yet, there are reports of harmless trees (A.

innoxia Bl.) and there seem to be no reports of such arrow-poisons

from the trees of Africa, New Guinea, and Melanesia. Bisset in¬

forms me in lift, that in these cases the glycoside-content is prob¬

ably very low. The fruits of A. toxicaria are eaten with impunity

though those of the Madagascan species are said to be poisonous.

Bisset (1957) found essentially the same cardiac glycosides in

Antiaris , Antiaropsis, and the American Ogcodeia. This chemical

evidence suggest the derivation of the American allies of Antiaris

from the New Guinea centre of Antiaropsis.

246

Vol. XIX. (1962).

Key to the species and varieties of Antiaris

1. Drupes subglobose 20-30 mm. wide. Madagascar.

2. Lamina -4x3 cm., harshly scabrid; lateral nerves 5-6 pairs

A . humbertii.

2. Lamina larger, becoming more or less smooth; lateral nerves 8-14

pairs . A. madagascariensis.

1. Drupes ellipsoid or pyriform.

3. Costal and intercostal venation strongly raised on the underside of

the scabrid, coriaceous lamina. Drupe 10-15 mm. long

A. africanct.

3. Intercostal venation not or scarcely raised. Adult lamina subscabrid

or smooth . A. toxicaria.

4. Lamina ovate-elliptic, attenuate. Drupe 20-45 X 17-26 mm. Mo¬

luccas, Philippines, eastwards . v. macrophylla.

4. Lamina elliptic-obovate, acuminate.

5. Drupe 12-25 mm. long. Asia to Celebes. v. toxicaria.

5. Drupe 10-15 mm. long. Africa.v. welwitschii.

A. africana Engl. Bot. Jahrb. 33 (1902) 119; Keay, FI. W. Trop.

Afr. ed. 2, 1, pt. 2 (1958) 612.

Lamina as in A. toxicaria v. toxicaria but strongly coriaceous,

strongly marked beneath by the raised costas and intercostals.

Male inflorescence -15 mm. wide. Drupe 10-15 X 9-10 mm.,

ellipsoid.

West and central Africa, in drier forest.

An earlier name may well be A. challa (Schw.) Engl., though

this is generally reduced to A. toxicaria v. welwitschii. Saplings

of A. africana seem indistinguishable from those of A. toxicaria.

A. humbertii Leandri, Not. Syst. 13 (1948) 175; FI. Madag. fam.

55 (1952) 32, f. vi.

Lamina -4X3 cm., small, cordate at the base, subacuminate,

harshly scabrid-hairy; lateral nerves 5-6 pairs, not prominent

beneath. Drupe 20-30 mm. wide, subglobose, poisonous.

Madagascar; shrub or small tree in xerophytic bush.

A. madagascariensis H. Perrier, Arch. Bot., Bull 1-4 (1927) 70;

Leandri, FI. Madag. fam. 55 (1952) 32, f. vi.

Lamina as in A. toxicaria v. toxicaria. Drupe 20-30 mm. wide,

subglobose, poisonous.

Madagascar; on sandstone and calcareous rocks, more or less

xerophytic.

247

Gardens Bulletin, S.

A. toxicaria Lesch. l.c.

v. toxicaria

Lamina mostly elliptic obovate, acuminate, base cordate to

cuneate and often somewhat asymmetric, thinly to rather strongly

coriaceous, subscabrid to smooth when adult. Male inflorescence

—20 mm. wide. Drupe 12-25 X 11-20 mm., ellipsoid to pyriform.

Distr. Ceylon, India, Burma, South China, Hainan, Indochina,

Thailand, Andaman Isl., Malaya, Sumatra, Java, Borneo, Celebes;

rainforest to 1,500 m. alt.

The leaves vary greatly in size and shape even on the same twig.

Possibly each new shoot bears a regular succession of differing

leaves.

var. macrophylla (R.Br.) Corner stat. nov.— Antiaris macro -

phylla R.Br. Flind. Voy. 2 (1814) 602, t. 5.— A. bennettii Seem.

Bonplandia 9 (1861) 259; id. 10 (1862) 3, t. 7. Arbor toxicaria

mas, Rumph. Herb. Amb. 2, 263, t. 87.

Lamina ovate-elliptic or elliptic, attenuate to the acute apex,

base cordate to widely cuneate, scabridulous to smooth. Drupe

20-45 X 17-26 mm. Tree or shrub.

Distr. Philippine Isl. Molucca Isl., Timor, (? Flores, Solor Isl.),

(? Soemba), New Guinea, Solomon Isl., New Hebrides, Fiji, Tonga,

North Australia (Company’s Isl. off Arnhem Land); lowland rain¬

forest.

The shrubby plant of Melanesia may be a cultivar spread by

sea-farers.

var. welwitschii (Engl.) Comer stat. nov.— Antiaris welwitschii

Engl. Bot. Jahrb. 33 (1902) 118; Keay, FI. W. Trop. Afr. ed. 2,

1, pt. 2 (1958) 613.— A. usambarensis Engl. id. 119.— A. kers-

tingii Engl. Veg. Erde 9, 3 (1), (1915) 33.—? Ficus challa Schw.

Bull. Herb. Boiss. 4, app. 2 (1896) 144.—? A. challa (Schw.)

Engl. Veg. Erde 9, 3 (1), (1915) 33.— A. challa Blatt. Rec. Bot.

Surv. Ind. 8 (1923) 446.

Leaf as in v. toxicaria . Male inflorescence -15 mm. wide.

Drupe 10-15 X 7-12 mm.

Distr. Tropical Africa; lowland forest tree.

I maintain this variety in deference to the geographical homo¬

geneity, the larger fruited specimens which occur in Asia not

having been collected from Africa, but specimens from Celebes

match those from Africa almost exactly in leaf and fruit;

in no way can I distinguish the leaf of this variety from that of

var. toxicaria. Possibly the male inflorescence is slightly smaller.

Several herbarium sheets show that the variety is monoecious, as

in the whole genus.

A. challa may be A. ajricana. I have been unable to see the

type.

248

Vol. XIX. (1962).

List of species and synonyms in Antiaris

A. africana Engl.

A. bennettii Seem, is A. toxicaria Lesch. var. macrophylla (R.Br.)

Corner.

A. challa (Schw.) Engl, is A. africana Engl, or A. toxicaria Lesch.

var. welwitschii (Engl.) Corner.

A. challa Blatt. is A. toxicaria Lesch. var. welwitschii (Engl.)

Corner.

A. dubia Span, is A. toxicaria Lesch. var. toxicaria.

A. hum berth Leandri.

A. innoxia Bl. is A. toxicaria Lesch. var. toxicaria.

A. kerstingii Engl, is A. toxicaria Lesch. var. welwitschii (Engl.)

Corner.

A. macro phylla R.Br. is A. toxicaria Lesch. var. macro phylla

(R.Br.) Corner.

A. madagascariensis H. Perrier.

A. palembanica Miq. is Artocarpus dad ah Miq.

A. rufa Miq. is A. toxicaria Lesch. var. toxicaria.

A. saccidora (Nimmo) Dalz. is A. toxicaria Lesch. var. toxicaria.

A. toxicaria Lesch.

A. turbinifera Hemsl. is not Antiaris. The type at Kew from the

Solomon Is!, seems to be a mixture of fallen leaves, flowers, and

fruits from as many different trees.

A. usambarensis Engl, is A. toxicaria Lesch. var. welwitschii (Engl.)

Corner.

A. welwitschii. Engl, is A toxicaria Lesch. var. welwitschii (Engl.)

Corner.

A. zeylanica Seem, is A. toxicaria Lesch. var. toxicaria.

Antiaropsi K. Schum.

FI. Kaiser-WilhelmsI. (1889) 40; Laut. et K. Schum. FI. Deutsch.

Schutzgeb. Suds. (1901) 267; Diels, Bol. Jahrb. 67 (1935) 174.

So far as concerns the female inflorescence this genus is central

in the Olmedieae. The superior ovaries, the free tepals, and the

interfloral bracts render it the least specialised of the tribe. If this

inflorescence were bisexual, containing male and female pedicel¬

late flowers, and became urceolate, it would represent a big step

towards the receptacle of Ficus. Sparattosyce Bur., of New Cale¬

donia, represents dioecious Antiaropsis with such an urceolate

receptacle splitting open at anthesis of the male flowers and at

maturity of the syncarp; at anthesis of the female receptacle the

249

Gardens Bulletin, S.

styles project through its ostiole in a manner not known in Ficus,

though conceivably its ancestral state. Sparattosyce has been

placed in the Ficeae, but is not blastophagous and its anthers are

extrorse; in Antiaropsis and Ficus they are introrse, which is un¬

usual in the family.

Antiaropsis is a leptocaul tree. If converted into an ancestral

pachycaul, it would have the possibilities of the non-blastophag-

ous Ficitf-ancestor. I have defined Ficus not only on its inflores¬

cence but on its blastophagy, and have, therefore, transferred

Sparattosyce to the Olmedieae. The work of Bisset (1957) on the

cardiac glycosides of Moraceae has shown that Antiaris , Antia¬

ropsis, Castilloa, Ogcodeia, and Pseudolmedia have much the

same glycosidal content, particularly in possessing antiarin and

antioside, but that Sparattosyce, as he informs me in litt., does

not have them.

There may be a second species of the genus, which passes as A.

decipiens var. parvifolia Diels. One collection of this (T. McAdam

s.n., Wau, Terr. New Guinea, 1939) has a small female inflores¬

cence, 4-5 mm. wide, with a single female flower. Whether it is

merely depauperate or a new species, the collections are insuffi¬

cient to decide, but it shows how the one-flowered state of Antiaris

may have arisen

Brosimeae Tree.

Ann. Sci. Nat. ser. 3, 8 ( 1847) 138; Miq. Mart. FI. Bras. 4, I

( 1852) 79; Bur. DC Prodr. 17 (1873) 282; Benth et Hook. Gen. PI.

3 (1880) 346; Engl. Nat. Pflanzenf. 3, 1 (1894) 87; Dalla Torre et

Harms, Gen. Siph (1900) 122.

Inflorescences bisexual, discoid, capitate-globose or urceolate,

more or less involucrate. Female flower central, solitary, surround¬

ed by numerous sessile, superficial male flowers. Stamens 4-1;

filaments indexed or straight in bud; anthers (? extrorse), varying

subglobose with transverse equatorial dehiscence; pistillode none.

Ovary superficial or more or less immersed in the receptacle. Seed

medium-size to large. Embryo (so far as known) straight; radicle

short; cotyledons not folded, equal or unequal. Trees, unarmed.

Genera 8 (Africa 7, America 1); species 44 (Africa 38, America

6 ).

1 include in this tribe Craterogyne Lanjouw, which was placed

in Dorstenieae.

Dorstenieae Gaud.

Voy. Freyc. (1826) 510.

I consider that this tribe contains only the genus Dorstenia. 1

refer Craterogyne to Brosimeae and Mesogyne to Olmedieae.

250

Vol. XIX. (1962).

Ficeae Gaud.

Voy. Freyc. (1826) 510.

I consider that this tribe contains only the genus Ficus. I refer

Sparattosyce to Olmedieae.

Genera Excludenda

Conocephaloideae: Cecropia L., Coussapoa Aubl., Musanga

R.Br., Myrianthus Beauv., Poikilospermum Zipp., and Pourouma

Aubl. to be transferred to Urticaceae.

Stenochasma Miq. belongs with Urticaceae.

Botrymorus Miq. is Pipturus Wedd. (see Koord. et Val., Bijdr.

12, 1910, 724).

Metatrophis F. Brown, Bull. Bish. Mus. Honolulu n. 130 (1935)

34.—This may be Urticaceae or Euphorbiaceae.

In conclusion, I express my gratitude to Dr. Chew Wee Lek y

of the Singapore Botanic Gardens, for his critical and detailed

help in the treatment of Conocephaloideae, Streblus sens, lat., and

Urticaceae in general.

References

Bechtel, A. R. (1921). The Floral Anatomy of the Urticales. Am. J. Bot. y

386-410.

Bernbeck, F. (1932). Vergleichende Morphologie der Urticaceen und

Moraceen Infloreszenzen. Goebel, Bot. Abhandl. 19, 1-100.

Bisset, N. G. (1957). Cardiac Glycosides, Part III. Ann. Bogor. 2, 219-

223.

Corner, E. J. H. (1954). The Durian Theory Extended. PhytomorphoJogy

4, 263.

Corner, E. J. H. (1960). Taxonomic Notes on Ficus, 1-5. Gard. Bull. Sing.

17, 368-485; id. 18, 1-69.

Hallier, H. (1905). Provisional scheme of the natural (phylogenetic)

system of Flowering Plants. New Phytol. 4, 151-162.

Hallier, H. (1912). L’origine et le systeme phyletique des Angiospermes.

Arch. N6erl. Sci. Exact. Nat. ser. 3, 1, 146-234.

Hutchinson, J. (1959). Families of Flowering Plants.

Jarrett, F. M. (1959). Studies in Artocarpus and Allied Genera. I, Gene¬

ral Considerations; Journ. Arn. Arb. 40, 1-29. II, A Revi¬

sion of Prainea; id., 30-37. Ill, A Revision of Artocarpus

subgen. Artocarpus; id., 113-368.

Gardens Bulletin , S

Jarrett, F. M. (1960). Studies in Artocarpus and Allied Genera. IV, A

Revision of Artocarpus subgen. Pseudojaca; id, 41, 73-140.

V, A Revision of Parartocarpus and Hullettia; id., 320-340.

Parkin, J. (1951). The Protrusion of the Connective beyond the Anther.

Phytomorph. 1, 1-8.

Renner, O. (1907). Beitrage zur Anatomic und Systematik der Artocar-

peen und Conocephaleen, inbesondere der Gattung Ficus. Bot.

Jahrb. 39, 319-448.

Tippo, O, (1938). Comparative Anatomy of the Moraceae and their pre¬

sumed Allies. Bot. Gaz. 100, 1-99.

Trecul, A. (1847). Memoire sur la Famille des Artocarpees. Ann. Sci.

Nat. ser. 3, 8, 38-157.

Weddell, H. A. (1857). Considerations Generates sur la Famille des

Urticacees. Ann. Sci. Nat. ser. 4, 8, 307-398.

Woodson, R. E. (1960). Flora of Panama (Moraceae). Ann. Mo. Bot.

Gard. 47, 135.

252

Some New Dipterocarpaceae from Borneo

By P. S. Ashton

Botany School, Cambridge University

A Forester’s Manual of Brunei’s Dipterocarps is in preparation.

The present paper provides descriptions for species which will be

included in the Manual but which have not as yet been published.

Of these several have been recognised for some time past by forest¬

ers; Hopea vaccinifolia, Shorea geniculata, and S. myrionerva were

recognised but not described by Symington over twenty years ago,

and these names were used by him and subsequent collectors on

herbarium sheets. Wyatt-Smith prepared, but did not complete,

a paper on the three Parashorea species, and I am grateful to him

for handing this over to me for completion. S. rubra, S. agami, and

S. flaviflora were names suggested by G. H. S. Wood, and prepara¬

tory notes were made by him for description of the latter. S. sloo-

teni was described by him in ms. as S. vanslooteni, and S. pilosa

as S. tomentosa. The very distinct S. isoptera was named by Wood

in ms. as Neohopea isoptera, but I am of the opinion that its affi¬

nities are too close to Shorea to merit generic status.

Kepong code letters were assigned to several of the species here

described:

P. macrophylla

—

P. sp. P.

P. parvifolia

—

P. sp. D.

P. smythiesii

—

P. sp. C.

S. acuta

—

S. sp. V.

S. agami

S. sp. L. and S. sp. G*

S. amplexicaulis

S. sp. P.

S. faguetioides

—

S. sp. T.

5. minor

*=-

S. sp. X 3

S. revoluta

S. sp. V

S. rubella

S. sp. A 3 and S. sp. H s

S . rubra

-=»

S. sp. J 3

S. slooteni

S. sp. N x

I am particularly grateful for the continued encouragement and

advice of Mr. B. E. Smythies, Conservator of Forests at Kuching,

and one time State Forest Officer, Brunei, who has put his great

knowledge of Borneo’s Dipterocarps at my disposal; to Mr. E. J.

H. Comer, f.r.s., for advice on numerous points and for correcting

the proofs; to Inche Yakin bin Long, State Forest Officer, Brunei,

253

Gardens Bulletin , S.

for his enthusiastic support and for his aid in obtaining for me the

type material of Hopea garangbuaya , Parashorea macrophylla, and

Shorea bullata; and to Dr. W. Meijer, Forest Botanist, Sandakan,

for his close co-operation and for supplying notes on several spe¬

cies. Lastly, but by no means least, I wish to record the friendly and

ever helpful partnership of Ladi anak Bikas, Asah anak Unyong,

Mujah anak Guang, and Sengelang anak Nantah, who climbed the

trees and obtained the collections on which these descriptions are

largely based, and with whom I spent three very happy years

exploring the forests of Brunei.

Thanks are also due to Mr. Brian Golding of the Botany School,

Cambridge University, for his painstaking preparation of the plates.

This paper is the first of several which will be jointly published

with Dr. Meijer, in order to bring our greatly increasing knowledge

of Borneo Dipterocarpaceae up to date.

HOPEA FLUVIAL1S Ashton s.n.

H. pedicellatae (Brandis) Sym. et H. griffithii Kurz affinis sed

lobis minoribus calycis in fructu et lamina differt.

Ramuli, perulae, stipulae, petiolique griseo-brunneo-tomentosi;

lamina primo griseo-brunneo-tomentosa, dein glabrescens. Ramuli

apicem versus -1.5 mm. diam., glabrescentes, leves, lenticellis

parvis inconspicuis punctati; internodis 8-20 mm. longis. Gemmae

-1 mm. longae, ovoideae. Stipulae -2 mm. longae, lineares,

fugaces. Lamina 7-12 X 2.8-4.8 cm., lanceolato-ovata, tenuis,

basi anguste vel late cuneata, subaequali, apice in acumen caudato

-1.5 cm. longum attenuato; costis lateralibus primariis utrinsecus

c. 10 confertis angustis, angulo 60°-80° exorientibus, ad marginem

curvatis, secundariis vix minus evolutis, basalibus marginem

secundum ad 1/3 laminae elongatis; intercostis obscuris reticulatis;

costa media subtus applanata vel paullum elevata, supra pro¬

minent. Petiolus 7-10 mm. longus, -1.5 mm. diam., sicco niger.

Lamina delapsa fusco-brunnea, costa intensius vel nigrescenti.

Racemi -6 cm. longi, semel vel bis ramosi, stipitis basi in fructu

-1.25 mm. diam., 1-3 axillares, raro terminales, ex integro teretes,

semper breviter griseo-pubescentes; ramulis -1.5 cm. longis, -7

flores gerentibus; bracteolis -2 mm. longis, ovato-acuminatis,

glabrescentibus. Alabastrum -4 mm. longum, ellipsoideum, bre¬

viter pedicellatum. Calyx glabrescens, ad marginem setulosus; lobis

extemis 2 ovato-acuminatis, intemis 3 tenuioribus, elliptico-

mucronatis. Corolla in vivo viridi-flava, petlis lineari-lanceolatis,

externe conferte pubescentibus, intus glabris. Stamina 15, verticil-

lis 3, bina cum solis alternata; filaments basim versus applanato-

dilatatis, supeme gradatim filiformibus; antheris subglobosis, locu-

lis interioribus quam exterioribus paullum minoribus; aristis

quam antheras 2-plo longioribus. Ovarium ovoideum glabrum;

stylo filiformi, tereti, ovarium aequanti; sine stylopodio distincto.

254

Vol. XIX. (1962).

Plate 1. Hopea fluvialis.

a, Flowering twig, b, Flower bud. c, Sepals (outer below, inner above), d, Petal, e, Ovary,

f, Stamens (lateral view above, frontal below), g, Fruit, h, Nut.

(a-f from Kep 34492; g, h, from Brun 3394).

Gardens Bulletin , 5.

Pedicellus in fructu -4 ram. longus, basi -1 mm. diam., superne

incrassatus. Fructus in racemo erectus. Calyx glaber; lobis longi-

oribus 2-5.5 X 1 cm., spatulatis, subacutis, basim versus angus-

tatis -4 mm. latis, partibus basalibus -7X5 mm., dilatatis, non

incrassatis, vix saccatis; lobis brevioribus 3, inaequalibus, 1-2.5

cm. longis, acutis, hastatis, basibus saccatis nucem obtegentibus,

adpressis. Nux -15X6 mm., glabra, anguste ovoidea, ad stylo-

podium breve acutum attenuata.

Collections. Indonesian Borneo: bb 20448, Puruktjau. Brunei:

Brun 3394, 5668, 794, 412, Kuala Belalong; Brun 5219, S. Bela-

long above Kuala Empan; Brun 125, 127, Kuala lngei; Brun 722,

Kuala Temburong Machang-Kuala Sekurop, Temburong; Kep

34492, Pajarai, S. Belait; Kep 34523, S. Melayan, Belait (Holo-

typus in Herb. Kepong).

HOPEA GARANGBUAYA Ashton s.n.

H. cernuae T. et B., M. mengerawan Miq. affinis, sed lamina

maiore latiore multo crassiore, coriacissima, floribus maioribus,

staminum aristis minute papillosis differt.

Ramuli, perulae, stipulae, petioli, laminaeque ex integro glabri.

Ramuli apicem versus -2 mm. diam., robusti, teretes, leves, fusco-

rubro-brunnei, lenticellis minutis orbicularibus punctati. Gemma

-2 X 1 mm. ovoidea. Stipulae -4 mm. longae, lineares, fugaces.

Lamina 11-16 X 6-10 cm., late ovata, crasse coriacea, basi

obtuse, apice in acumen -1.2 cm. longum angustum subabrupte

attentuato, margine paullum revoluto, costis lateralibus primariis

utrinsecus 8-11, curvatis, dryobalanoidiis, sed prominentibus.

angulo 60°-70° exorientibus; costis lateralibus secundariis bre-

vibus; intercostis distinctis, conferte scalariformibus, a costa media

angulo 90° exorientibus; costa media lata, subtus prominenti

tereti, supra paullum elevata. Petiolus 2-2.5 cm. longus, -2.5 mm.

diam., robustus, sicco niger. Lamina delapsa ex integro flavo-

olivacea. Racemi -9 cm. longi, ordine semel ramosi, stipitis basi

in fructu -1.5 mm. diam., terminales vel 1-2 axillares, ex integro

teretes, glabri; ramulis -3 cm. longis, -6 flores secundos gerentibus;

bracteis bracteolisque ignotis. Alabastrum —6 X 2.5 mm. magnum,

ellipsoideum, distincte pedicellatum. Calyx extus glabrescens, intus

glaber, margine breviter setoso; lobis externis 2 ovato-acuminatis;

lobis internis 3 tenuibus, suborbicularibus obtusis. Petala -1.2

cm. longa, lineari-lanceolata, acuta, partibus in alabastro expositis

glabrescentibus, aliter glabra. Stamina 15, verticillis 3, bina cum

solis altemata; filamentis gracilibus, attenuatis; antheris oblongis,

loculis exterioribus quam interioribus paullum maioribus; aristis

quam antheras 2-3-plo longioribus, basi tuberculis glandulosis

prominentibus punctatis. Ovarium ovoideum basi glabra, dimidio

apicali conferte setoso, in stylum attenuato; stylopodio indistincto.

256

VoL XIX. (1962).

/Mm

Plate 2. Hopea garangbuaya.

a, Flowering twig, b, Flower bud. c, Sepals (outer above, inner below), d, Petal, e, Ovary

and stamens, f, Stamens (left, lateral; right posterior, view), g, Left, fruit; nut.

(a-e from Brun 2006a; f, g, from 2006c).

257

Gardens Bulletin, S .

stylo ovarium aequanti, filiformi, attenuate, dimidio basali con-

ferte setoso, aliter glabro. Pedicellus in fructu c. 3 mm. longus,

basim fructui versus incrassatus. Calyx glaber, conferte imbricatus;

lobis longioribus 2, 7 X 1.5 cm., plus minus loratis, obtusis, parti-

bus basalibus 1 cm. latis, non incrassatis, tenue saccatis, subauri-

culatis; lobis minoribus 3, -2 X 1.2 cm., ovatis, acutis basibus ut

in lobis longioribus. Nux -20 X 9 mm., ovoidea, attenuata, glabra,

stylopodio -4 mm. longo apiculato.

Collections. Brunei: S 1658, 2055, 5768, Batu Apoi; Brun 3347,

Brun 2006a (Holotypus in Herb. Kew), Ulu Senuko, Labu; Brun

713, Ulu S. Labu.

r / !

HOPEA VACCINIFOLIA Ridl. ex Ashton :—Hopea vaccinifolia

Ridl. nomen nudum.

H. bracteatae Burck affinis, sed racemis brevissimis, laminis mi¬

nimis, corolla revoluta, staminis 10 differt.

Ramuli, perulae, stipulae, petiolique conferte pallide griseo-

brunneo-pubescentes. Ramuli apicem versus -0.5 mm. diam.,

graciles, plurimum horizontale alterne ramosi, dein glabrescentes,

leves, fusco-rubro-brunnescentes; internodis -8 mm. longis. Gem¬

mae minutae. Stipulae -1.25 mm. longae. breviter hastatae,

acutae, fugaces. Lamina 1-2.5 (3) X 0.4-1.2 cm., chartacea, apice

breve obtuso, vel in arboribus maturis -5 mm. longis late acumi-

natis; costis lateralibus primariis utrinsecus c. 6, subdryobalanoidi-

is, obscurissimis; costis lateralibus secundariis angustissimis; inter-

costis reticulatis; costa media angusta, subtus applanata vel

paullum elevata, supra depressa. Petiolus -2 mm. longus, -0.5 mm.

diam., gracilis, sicco fusco-brunneo-puberulus vel nigrescens.

Lamina delapsa ex integro pallide griseo-brunnescens, sicca invo-

luta. Racemi -1.2 cm. longo, inconstanter semel ramosi, stipitis

basi in fructu 0.5 mm. diam.; -2 axillares, breves, teretes, glabri;

ramulis brevibus, -3 flores gerentibus; bracteolis minutis, deltoi-

diis, subpersistentibus, glabris. Alabastrum parvum, ovoideum,

subsessile. Calyx ad apicem marginibus setosis, aliter glaber; lobis

extends 2 ovatis, acutis, lobis internis 3 late suborbicularibus, ob¬

tusis, quam illis tenuioribus. Corolla in vivo fusco-vinosa, rotata

vel reflexa; petalis oblongis, obtusis, glabris, multo-contortis,

aetate expandi basim versus tubiforme imbricatis. Stamina 10,

subaequalia; filamentis basim versus dilatatis, supeme abrupte atte-

nuatis filiformibus; antheris subglobosis, loculis exterioribus quam

interioribus paullum longioribus; aristis gracilibus, quam antheras

2-plo longioribus, apicem stylopodii attingentibus. Ovarium stylo-

podiumque cylindricum, truncatum, podio apicali puberulo, aliter

glabrum; stylo brevi, abrupto. Pedicellus in fructu c. 1 mm. longus.

Calyx glaber; lobis subaequalibus, -4 X 3.5 mm., ovatis acutis

vel obtusis incrassatis, tenue saccatis. Nux -8X6 mm., ovoidea,

glabra, ad apicem mucronata.

258

Vol. XIX. (1962).

Plate 3. Hopea vaccinifolia.

a, Flowering twig, b, Leaf, c. Flower bud. d. Sepals (left, outer; right, inner), e, Corolla.

f ’ Petal, g, Ovary, h, Stamens (1, frontal; 2, lateral; 3, from inside), i. Fruit.

(a-h from Brim 3035; i from Kep 35567).

259

Gardens Bulletin , S.

Collections. Brunei: Kep 30614, 30630, Andulau F.R.; Kep

35567. Pengkalan Ran; SFN 10494. Brun 3035. Labi Rd. M.5; Brun

140, R. Ingei. Sarawak: Brun 5564, Hose 583 (Holotypus in herb.

Kew) Marudi.

HOPEA WYATTSMITHII Wood ex Ashton:—//, wyattsmithii

Wood, msc.

H. glaucescentis Sym. affinis, sed lobis calycis in fructu quam

nucem brevioribus subaequalibus differt.

Ramuli, perulae, stipulae, petioli laminaeque ex integro glabri.

Ramuli apicem versus -1 mm. diam., fusco-rubro-brunnei, leves;

internodis 1.5-3 cm. longis. Gemma -1.5 mm. diam., ovoidea.

Stipulae -2 mm. longae, lineares, fugaces. Lamina 9-14 X 5.5-9

cm., late ovata vel elliptica, basi late cuneata, raro obtusa, apice

in acumen caudatum -2 cm. longum abrupte attenuato; costis

lateralibus primariis utrinsecus 4-8, remotis, propter costas la¬

terals secundarias breves prominentes diffusis et irregulariter dis¬

poses, basi 2-3 utrinsecus primo ad costam mediam decurrenti-

bus, dein rectis, aliter curvatis, angulo 40°-50° exorientibus; in-

tercostis remotis, scalariformibus, a costa media > 90° exorienti¬

bus; costa media angusta, subtus applanata, supra acuta. Petiolus

1.2-1.7 cm. longus, -2 mm. diam., sicco niger. Lamina delapsa

ex integro pallide griseo-brunnea. Racemi -6 cm. longi, semel

ramosi, stipitis basi in fructu -1 mm. diam., terminales vel -2

axillares, ex integro glabri, teretes vel paullum compressi; ramulis

-1.5 cm. longis, -6 flores secundos gerentibus; bracteolis parvis,

deltoideis, glabris. Alabastrum -3 mm. longum, subglobosum.

Calyx glabrescens, margine breviter setoso, expansus; lobis sub¬

aequalibus, late ovatis vel suborbicularibus, obtusis. Petala

oblongo-lanceolata, acuta, partibus in alabastro expositis conferte

tomentosis, aliter glabris. Stamina 15; verticillis 3, bina cum solis

alternata; filamentis basim versus dilatatis, sub antheris abrupte

attenuatis; antheris late oblongis, loculis subaequalibus; aristis

angustis, quam antheras 2-plo longioribus. Ovarium stylopodium-

que pyriforme, ovario glabro, stylopodio quam ovarium paullum

longiori, breve puberulo, style brevi glabro. Pedicellus in fructu

-2.5 mm. longus, -2 mm. diam. Calyx nuxque glaber, calyce quam

nucem breviore; lobis 5 subaequalibus, -10 X 8 mm., ovatis, sub-

acutis, paullum saccatis, incrassatis, conferte inbricatis, ad nucem

adpressis, foramine apicali -3 mm. diam. apice nucis exposito. Nux

-1 X 1 cm., ovoidea, apice acuto, stylopodio vestigiali.

Collections. North Borneo: San 15061, Beaufort (Holotypus in

Herb. Kew); San 16467, Tawau. Brunei: Brun 885, Ulu Tutong

at the second stone rapid; S 5589, Brun 833, Andulau F.R.; Kep

35475, S. Biang: S 5809, Bt. Patoi.

260

Vol. XIX. (1962).

Plate 4. Hopea wyattsmithii.

a, Fruiting twig, b, Flower bud. c, Sepals (left, inner; right, outer), d, Petal, e, Stamens

(1, frontal; 2, from inside; 3, lateral), f, Ovary, g, Fruit.

(a, g, from San 16467; b-f from San 15061).

261

Gardens Bulletin, S.

PARASHOREA MACROPHYLLA Wyatt Smith ex Ashton:—

P. macrophylla Wyatt Smith, nomen nudum.

Lamina maxima subtus glauca, costis lateralibus multis, ramulis

compressis, cicatricibus stipularum amplexicaulibus, lobis calicis

in fructu longissimis distinguitur.

Ramuli, perulae, externi stipularum, petiolaeque valde breviter

conferte plane pallide ochraceo-tomentosi, intemi stipularum lami-

naeque glabri. Ramuli apicem versus -12 X 5 mm., compressi,

dein terescentes, pallide rubro-brunnescentes; cicatricibus stipu¬

larum angustis amplexicaulibus; cicatricibus foliorum ellipticis,

magnis, prominentibus; foliis altemis; internodis 1-4 cm. longis.

Gemma -9X8 mm., linearis, acuta, glaucescens. Stipulae -15

X 2.5 cm., lineares, subacutae, caducae. Lamina 30-50 X 16-24

cm., tenuis, oblongo-elliptica, subtus argenteo-glauca, basi sub-

cordata, apice obtuso vel breve acuminato, abrupte attenuate;

costis lateralibus utrinsecus 28-36, confertis, rectis, sed ad mar-

ginem curvatis, angulo 25°-50° exorientibus basi excepta, subtus

prominentibus, glabris, non glaucis, plicis obscuris inter costas sub-

persistentibus; intercostis angustis, conferte scalariformibus, a

costis lateralibus diagonale exorientibus; costa media crassa, sub¬

tus tereti, supra vix prominent! Petiolus 3-5 cm. longus, -5 mm.

diam., teres, sicco pallide ochraceo-tomentosus. Lamina delapsa

supra ochraceo-brunnescens, subtus pallide purpureo-brunnescens,

glaucescens, costis pallide fuscis. Inflorescentiae paniculatae, -6

cm. longae, bis opposito-ramosae, stipitis basi in fructu -4 mm.

diam., terminales vel axillares, rigidae, rectae, breviter pallide gri-

seo-brunneo-pubescentes; bracteis -4 X 2.5 cm., magnis, ovatis,

acutis, saccatis, externe conferte breviter pubescentibus, intus

glabris; bracteolis -12X8 mm., ut in bracteis. Flores maturi ig-

noti. Alabastra ellipsoidea, calyce externe conferte, intus sparsim

tomentoso, lobis subaequalibus, anguste ovatis, acutis; petalis

oblongis, obtusis; staminis 15 subaequalibus; filamentis brevibus

deltoidiis; antheris lineare-oblongis, loculis internis 2 quam exter-

nis 2 paullum brevioribus; aristis prominentibus, angustis, atte-

nautis, quam antheris 1-plo longioribus; ovario conico, breviter

pubescenti, ad stylopodium prominens anguste cylindricum atte¬

nuato. Pedicellus in fructu brevis. Calyx in fructu sparsim pube-

rulus, dein glabrescens, basim versus confertius; lobis longioribus

3, -22 X !8 cm., anguste spatulatis, crassis, fragilibus, ad api¬

cem subacutis, basim versus -5 mm. latis gradatim attenuatis,

partibus basalibus -16 X 8 mm., ellipticis, saccatis, ad nucem

adpressis; lobis brevioribus 2,-12 X 0.7 cm., linearibus, inae-

qualibus, basim versus -5 mm. latis gradatim attenuatis, partibus

basalibus -5X5 mm., quam illos plure saccatis. Nux -2.5 X 1.2

cm., ellipsoidea, conferte griseo-brunneo-tomentosa, stylopodium

versus attenuata, stylopodio -6 mm. longo, lineari.

262

Vol. XIX. (1962).

Plate 5. Parashorea macrophylla.

a, Flowering twig, b, Fruit.

(a from Brun 2002; b from s.n., S. Tubai.

263

Gardens Bulletin. S .

Collections. Sarawak: No. 81848, S. Naut, Bakong; Brun 5423,

S. Sut, Rejang; s.n., S. Tubai, Limbang; Pursglove 5207, Tau

Range; S 1281, Baram. Brunei: Brun 901, S. Kebubok, Tutong;

Brun 3136, Ulu Biang; San 17377, Kep 48483, S. Belalong; Kep

30388: S. Motong; Brun 2002: Kuala Belalong (Holotypus in

Herb. Kew).

PARASHOREA PARVIFOLIA Wyatt Smith ex Ashton:—

Parashorea parvifolia Wyatt Smith, nomen nudum.

Lamina parva applanata non glauca, costis lateralibus utrinsecus

c. 9 angustis, curvatis, vix elevatis, intercostis remotis scalariformi-

bus, petiolo longo gracili glabro distinguitur.

Perulae stipulaeque sparsim breviter pallide flavo-brunneo-

tomentosae, ramuli stipulaeque primo breviter pallide flavo-brun-

neo-tomentosi dein glabrescentes. Ramuli apicem versus -1 mm.

diam., teretes, apicem aversus glabrescentes leves fusco-brunnei,

lenticellis minutis orbicularibus sparsi; cicatricibus stipularum

brevibus, angustis, sursum vel deorsum directis; intemodis 1-2.5

cm. longis. Gemma -3X1 mm., anguste lanceolata, acuta. Sti-

pulae -3.5 mm. longae, angustae, acutae, fugaces. Lamina 6-9

X 3-4.5 cm., elliptica vel ovata, non glaucescens, basi late

cuneata, apice in acumen anguste vel caudate —1.5 cm. longum

attenuato; costis lateralibus utrinsecus 8-10, angustis curvatis, sub-

tus prominentibus, laxis, angulo 50°-60° exorientibus; intercostis

scalariformibus vel paullum reticulatis, laxis, a costis lateralibus

c. 90° exorientibus; costa media angusta, subtus prominenti, supra

depressa. Petiolus 1-1.8 cm. longus, -1 mm. diam., dimidio distali

incrassato, geniculatus, sicco nigrescens. Lamina delapsa supra

flavo-brunnea, subtus pallide fusca. Racemi -4 cm. longi, semel

ramosi, stipitis basi in fructu -1.5 mm. diam., terminales vel axil-

lares, teretes, recti, penduli, sparsim breviter pallide flavo-brun-

neo-tomentosi; bracteis bracteolisque ignotis. Alabastrum -4.5 X

3 mm., ellipsoideum, obtusum. Calyx exteme conferte griseo-

brunneo-tomentosus, intus glaber; lobis lineari-deltoideis, subae-

qualibus, subacutis, ad corollam in alabastro non adpressis. Corolla

in vivo alba, petalis ellipticis obtusis, partibus in alabastro expositis

breviter tomentosis. Stamina 15, interiora 5 quam exteriora 10

paullum longiora stvli parte quartaria apicali superata; filamentis

basim versus applanato-dilatata, sub antheris subabrupte attenua¬

te; antheris lineare oblongis; aristis brevibus, supra antheras

paullum extrusis. Ovarium subglobosum, conferte tomentosum;

stylo quam ovarium 3-plo longiore, filiformi, tertia parte basali

breviter pubescenti, aliter glabro. Pediccllus in fructu -2 mm.

longus, brevis. Calyx in fructu apicem versus glabrescens, basim

versus puberulus; lobis longioribus 3-8.5 X 1.7 cm., late spatu-

latis, obtusis, basim versus -3 mm. latis attenuatis; partibus basali-

bus —7 mm. lato, non dilatatis, paullum incrassatis, valvatis, a nuce

264

Vol. XIX. (1962).

Plate 6. Parashorea parvifolia.

a, Fruiting twig, b, Sepals (right, outer; left, inner), c, Petal, d, Stamen (from inside)

e, Stamen (frontal view), f. Ovary, g, Fruit.

(a from Brim 3074; b-f from Brun 3013; g from Brun 3381).

265

Gardens Bulletin, S.

maturitanti repulsis; lobis brevioribus 2, —7.5 cm. longis, subae-

qualibus, saepe illos subaequalibus. Nux-1.5 X 1.3 cm., ellipsoi-

dea, obtusa, pallide verrucoso-lenticellata; stylopodio —1 mm.

longo, brevi, abrupto.

Collections. Sarawak: Haviland 2810/2331, Belaga, Rejang

(Holotypus, in Herb. Kew); Pickles 3431, R Kenaban, Plieran;

S A 0308, Ulu Baram; Brun 3159, Ulu Limbang. Brunei: Kep

34567, Pak Andaki; Kep 37057, Kiudang, Tutong; S 1693 S.

Kiaput, Tutong; Brun 2604, Kuala Temburong Machang; Brun

2528, Ulu Temburong-Medamit watershed; Brun 3013, 3074,

Bangar; Brun 3381, 455, S 5731, 5732, Kuala Belalong. North

Borneo: San A 4331, Kota Belud; Kep 80463, Pangi. Indonesian

Borneo: bb 10165, 17811, 18165, 18314, 17954, Tidung; bb 11772,

Bulungan.

PARASHOREA SMYTHIESII Wyatt Smith ex Ashton:— P.

smythiesii Wyatt Smith, nomen nudum.

P. lucidae Kurz affinis, sed lobis calycis in fructu longioribus,

lamina maiore, petiole breviore, conferte breviter tomentoso, costis

lateralibus utrinsecus 14-18 difTert

Ramuli, perulae, petiolique semper pallide flavo-ochraceo-tomen-

tosi; costae intercostae laminaque subtus sparsim scabrido-tomen-

tosi. Ramuli apicem versus -3 mm. diam., teretes vel striati; dein

teres glabrescentes, leves, fusco-brunnescentes; cicatricibus stipu¬

late -1.5 mm. longis, angustis, plus minus horizontalibus; inter-

nodis 1-2 cm. longis. Gemma -5X3 mm., ovoidea obtusa.

Stipulae -10x4 mm., late hastatae, subacutae. Lamina 10-15

X 4-7 cm., tenuis, subtus argenteo-glauca, elliptico-oblonga, basi

obtusa, apice in acumen -5 mm. longum plus minus abrupte

attenuato; costis lateralibus utrinsecus 14—18, rectis, angulo 30°-

45° exorientibus; inter costas obscuris plicatis persistentibus; inter-

costis scalariformibus distinctis; a costis lateralibus c. 90° exorienti¬

bus; costa media subtus prominenti, supra angusta paullum dep-

ressa Petiolus 1.5-2.2 cm. longus, -2.5 mm. diam., brevis, cras-

sus, sicco pallide brunneo-tomentosus. Lamina delapsa supra

flavo-brunnea, subtus pallide hebes purpuro-brunnea. Racemi -10

cm. longi, bis ramosi, stipitis basi in fructu -1.5 mm. diam.; termi-

nales vel axillares, angulati vel compressi, semper breviter pallide

griseo-brunneo-tomentosi; ramulis brevibus, -5 flores gerentibus;

bracteis -8X3 mm., oblongis, obtusis, externe breviter pubescenti-

bus, intus glabrescentibus; bracteolis -4X3 mm., late ellipticis,

alabastrum obtegentibus adpressis, externe conferte tomentosis.

Alabastrum -7X4 mm., ovoideum, obtusum. Calyx externe bre¬

viter conferte pallide griseo-brunneus; lobis anguste ovatis, lobis

externis 3 obtusis, quam lobis internis paullum longioribus et

angustioribus; lobis internis 2 acutis, subacuminatis. Corolla in

vivo alba; petalis ellipticis obtusis, partibus in alabastro expositis

266

Vol. XIX. (1962)

Plate 7. Parashorea smythiesii.

a, Flowering twig, b, Fruit, c, Sepals (outer, left; inner, right), d, Petal, e, Ovary,

f. Stamen (frontal view), g, Stamen (inside view), h. Stamen (fronto-lateral view),

(a, c-h, from Brun 2000; b from Kep 35456).

267

Gardens Bulletin, S.

breviter tomentosis. Stamina 15, interiora 5 quam exteriora 10

paullum longiora; filamentis basim versus applanato-dilatatis, sub

antheris abrupta attenuatis; antheris lineare oblongis, prope apicem

styli attingentibus; aristis brevibus, apices antherorum non super¬

antibus. Ovarium ovoideum, conferte tomentosum; stylo filiformi,

quam ovarium 3-plo longiore, tertia parte basali breviter tomen-

tosa, aliter glabra. Pedicellus in fructu -2 mm. longus, brevis.

Calyx in fructu apicem versus glabrescens, basim versus puberulus;

lobis longioribus 3, -10 X 1.7 cm., anguste obtusis, basim versus

-2.5 mm. latis gradatim attenuatis; partibus basalibus -6 X 3.5

mm., paullum saccata, incrassata; lobis brevioribus 2, -70 X 7

mm., spathulatis, ad basim angustis. Nux -13 X 9 mm., ovoidea,

conferte fulvo-tomentosa, ad stylopodium lineare -4 mm. longum

subabrupte attenuata.

Collections. Indonesian Borneo: bb 18320, 17890, Tidung.

North Borneo: San 18604, 18720, Tawau; San A 1732, A 1703,

San A 4091, Beaufort; San A 3284, Sipitang, Kep 80261,

S. Lukutan; San A 3967, Kalabakan, Tawau. Sarawak: S 123, S.

Miah, Balleh, Rejang; S A 424, Balleh. Brunei: Kep 35488,

Brun 3016, 3169, Bt. Biang; S 1667, Biang; Kep 35456, 35445,

35448, S 2102, 2104, Kep 35458, Bt. Patoi; San 17071, S 5738,

Brun 472, 2000a (Holotypus in Herb. Kew), 3129, Kuala Belalong;

S 1657, Bt. Apoi; Kep 80138, Peradayan F.R.; Brun 308, Batu

Apoi-Sebatu watershed; Brun 5217, S. Belalong above Kuala Em-

pan; S 5590, 5593, Ulu Senuko, Labu.

SHOREA ACUTA Ashton s.n.

S. macropterae Dyer affinis, sed lamina maiore coriaceore, costis

prominentioribus, tomento differt.

Ramuli, perulae, stipulae, petiolique semper conferte, breviter,

plane, pallide brunneo-tomentosi; costae subtus costaque media

supra primo conferte breviter plane pallide brunneo-tomcntosae,

dein glabrescentes. Ramuli apicem versus -4 mm. diam., dein

teretes glabrescentes, leves, pallide roseo-brunnescentes; internodis

1-4 cm. longis; cicatricibus stipularum primo -3 X 1 nun.,

cuneatis, transversis. Gemma 6-10 X 4-6 mm., ovoidea, sub-

acuta. Stipulae -12 X5 mm., oblongae, obtusae. Lamina 14-26

X 6.5-12 cm., ex integro glaber, basi obtusa vel late cuneata,

apice in acumen -2 cm. longum anguste attenuato; intercostis

lateralibus utrinsecus 10-13, subtus prominentibus, basi angulo

90° aliter 45°-55° exorientibus; intercostis angustis, sinuatis,

conferte scalariformibus, costis lateralibus diagonale exorientibus;

costa media subtus prominenti, tereti, supra angusta, paullum

depressa. Petiolus 1.5-2 cm. longus, 3-4 mm. diam., crassus, sicco

pallide brunneo-pubescens vel nigrescens. Lamina delapsa supra

pallide roseo-brunnea, subtus luteo-ferruginea. Racemi -18 cm.

268

Vol XIX . (1962)

O !Mm

Plate 8. Shorea acuta.

a, Flowering twig and leaf, b, Sepals (outer, above; inner, below), c, Ovary and androe-

cium. d, Ovary, e, Fruit.

(a-d from Brun 3076; e from s.n., Bt. Teraja).

269

Gardens Bulletin, 5.

longi, semel vel bis ramosi, stipitis basi in fructu -3 ram. diam.,

terminates vel axillares, ex integro compressi, semper breviter

pallide brunneo-pubescentes; ramulis -6 cm. longis, -9 flores dis-

tichos gerentibus; bracteolis -3.5 X 2.5 mm., ellipticis, obtusis,

externe breviter flavido-brunneo-tomentosis, intus glabris,

caducis. Alabastrum -9X4 mm., anguste ellipsoideum, obtusum.

Calyx externe breviter conferte pallide flavido-alutaceo-tomentosus,

intus glaber; lobis subaequalibus, ovatis, subacutis. Corolla in

vivo fusco-rubra, partibus in alabastro expositis pallide brunneo-

tomentosis; petalis linearibus, subacutis. Stamina 15, verticillis in-

aequalibus 3, interna 5 longissima ovarium paullum superantia;

filamentis ad antheras gradatim attenuatis, aristis brevibus, dein

reflexis. Ovarium stylopodiumque quam stylum c. 3-plo longiore

anguste conicum vel ovoideum, apicem versus asperum tomento-

sum, basim versus breviter tomentosum; stylo filiformi, glabro.

Pedicellus in fructu -1 mm. longus, -2.5 mm. diam., fructu sub-

sessili. Calyx in fructu basim versus breviter pubescens, aliter

glabrescens; lobis longioribus 3,-15 X 2.8 cm., spatulatis,

obtusis, basim versus —1.5 cm. latis attenuatis; partibus basalibus

-1.8 cm. lato, ad pedicellum abrupte terminals, discis centralibus

-8X7 mm. ovatis saccatis incrassatis, auriculis duobus laterali-

bus; lobis brevioribus 2, -8 X 0.7 cm., linearibus, inaequalibus,.

partibus basalibus ut in lobis longioribus. Nux -3X2 cm.,

ovoidea, conferte pallide badio vel pallide alutaceo-tomentosa,

ad stylopodium brevem acutum attenuata.

Collections. Sarawak: S 1922, S. Bakong, Baram; Brunei: Kep

30505, Brun 500, Bt. Biang; Kep 47042, ‘Brunei’; Kep 35545,

Pengkalan Ran; S 1924 (Holotypus in Herb. Kew), S. Liang; Kep

30579, Ukong; S 1906, 2171, Kep 80088, 30596, 30607, 30628,.

80096, S 4950, Brun 3291, 3076, 3164, San 17474, Andulau

F.R.; s.n., Bt. Teraja.

SHOREA AGAMI Ashton s.n.

S. virescentis Parijs affinis, sed floribus minoribus, basi fructus

impressa, laminis maioribus coriaceoribus differt.

Ramuli, perulae, stipulae, petioli, costaque media subtus primo

pallide brunnescentes, dein glabrescentes. Ramuli apicem versus

2-3 mm. diam., leves, fusco-brunnescentes vel nigrescentes, teretes,

saepe lenticellis minutis pallidis punctati; cicatricibus stipularum

brevibus inconspicuis; internodis 1-3 cm. longis. Gemma c. 4 X

2.5 mm., ovoidea vel falcata, obtusa, paullum pubescens vel gla¬

brescens, stipulae c. 10 X 3 mm., oblongae, obtusae, externe

breviter pubescentes, intus glabrae, caducae. Lamina 10-15 X

6-10 cm., late ovoidea vel oblonga, coriacea, basi obtusa vel

cuneata, apice in acumen late deltoideum 0.5-1 cm. longum sub-

abrupte attenuato; costis lateralibus utrinsecus 9-13, remotis.

270

Vol XIX. (1962)

a, Fruiting twig, b, Sepals (left, inner; right, outer), c, Petal, d, Stamen (frontal view)

e, Ovary.

(a from San 15402, b-e from Kep 30525).

271

Gardens Bulletin, S~

curvatis, ad basim laminae angulo c. 90°, apicem versus eiusdem

angulo 40° exorienlibus. Petiolus 1—1.5 cm. longus, 1.5-2 mm.

diam., sicco niger. Lamina delapsa supra fusco-brunnescens, subtus

ex integro viride-brunnescens. Racemi 6-10 cm. longi, semel vel

bis irregulariter ramosi, stipitis basi in fructu -1.5 mm. diam.,

terminales vel axil lares, basim versus teretes, apicem versus angu-

lati, primo breviter pallide fulvo-tomentosi, dein glabrescentes;

ramulis brevibus, -5 flores gerentibus; bracteis fugacibus; brac-

teolis -3 mm. longis, anguste ellipticis, subacutis, externe breviter

pallide alutaceo-fulvo-tomentosis, intus puberulentibus, caducis

Alabastrum -5 X 2.5 mm., parvum, anguste ellipsoideum, sub*

acutum. Pedicellus c. 0.5 mm. longum. Calyx conferte pallide

flavido-alutaceo-tomentosus; lobis externis 3 anguste ovatis, sub-

acuminatis; lobis internis 2 prominente caudato-acuminatis, quam

lobis externis 2/3-plo longioribus angustioribus. Petala parva,

ovato-elliptica, acuta, partibus in alabastro expositis conferte tom-

entosis, aliter externe puberulentia. Stamina 15, verticillis subae-

qualibus 3, filamentis ut antheras longis, paullum angustis, at-

tenuatis, antheris oblongis, apicem versus paullum attenuatis;

aristis quam antheras c. 3-plo longioribus, gracilibus, apicem styli

attingentibus. Ovarium ovoideum, conferte tomentosum, stylo

quam ovarium paullum longiori, dimidio basali sparsim tomen-

toso, dimidio distali glabro, crasse filiformi, apicem versus ob¬

scure trifurco. Pedicellus in fructu -3 mm. longus, brevis, crassus.

Calyx primo breviter pubescens, dein glabrescens; lobis longiori¬

bus, 3, 10-12 X L5-2.2 cm., spatulatis, obtusis, basim versus

-5 mm. latis gradatim attenuatis, partibus basalibus 1-1.5 cnn

latis, multum saccatis, incrassatis; lobis brevioribus 2, -6 X 0.4

cm., inaequalibus, linearibus, anguste acutis, partibus basalibus

ut in lobis longioribus. Nux -2 X 1.5 cm., laber, ovoidea, ad

stylopodium 3-4 mm. longum attenuata, vasi ad pedicellum im-

presso.

Collections. Brunei: Kep 35449, Bt. Patoi; Kep 30483, S. Ke-

duan; Kep 34401, S. Lumut; Kep 37249, Bangar; Kep 48108,

Labi Hills; Kep 48153, B. Rampayoh; Kep 48160, S. Lumut; Kep

48206, Bt. Gatas; Kep 48209, 48211, Balai Tarunan; Kep 48491,

Brun 3092, S 1939, Andulau F.R.; Brun 3135, Kuala Temburong

Machang; Brun 211, S. Topi, Ulu Belait; Kep 30525 (Holotypus

in Herb. Kepong), Labi Hills; Brun 739, Kuala Sekurop; S 1632,

Kenapol, Labi Hills F.R.; S. 1636, Labi Hills F.R. North Borneo:

San 16947, 16987, Beaufort; San 17797, Sibubu R., Sipitang; S

A379, S. Rayoh, Sabah; San A 3129, A 3477, A 3131, 15385,

15402, Sandakan; San A 3160, Kudat. Indonesian Borneo: Meijer

2464, Tarakan; bb 13617, Karangan R., NW of Sangkulirang; bb

17963, 17870, 33002, Tidung. Sarawak: S 1537, Bt. Meringgit,

Merapok, Lawas

272

Vol. XIX. (1962)

SHOREA AMPLEXICAULIS Ashton s.n.

S. beccarianae Burck affinis, sed ramulis angustioribus vix com-

pressis, cicatricibus stipularum amplexicaulibus, petiolo breviore

differt.

Novellae primo pallide alutaceo-pubescentes vel brunneo-tomen-

tosae, dein glabrescentes vel sparsim tomentosae. Ramuli apicem

versus -2x3 mm. diam., paullum compressi, dein teretes, graci-

lcs, pallide griseo-alutacei, glabri, leves; intemodis 1—3 cm. longis;

cicatricibus stipularum amplexicaulibus, c. 0.5 mm. latis, horizon-

talibus. Gemma 10-15 X 3-5 mm., hastata, acuta. Stipulae -25

X 8 mm., hastatae, subacutae, subpersistentes. Lamina 11-21 X

5—8 cm., elliptica, coriacea, in vivo novella magenta, basi cuneata,

apice in acumen —1 cm. longum attenuato; costis lateralibus utrin-

secus 9-12, angulo 50°-70° exorientibus, subtus prominenti;

intercostis distinctis, scalariformibus, remotis; costa media subtus

prominenti, tereti, supra plus minus applanata. Petiolus 1.5—2 cm.

longus, -2 mm. diam., brevis, sicco pallide griseo-brunnea. Lamina

delapsa supra pallide purpureo-brunnescens, subtus rubro-brun-

nea. Racemi —24 cm. longi, semel ordine ramosi, stipitis basi in

fructu -3 mm. diam., terminales vel axillares, laxi, ex integro

graciles, compressi, primo saepe aureo-tomentosi, dein glabres¬

centes; bracteis -13 X 8 mm., lanceolatis, acutis, glabris, fugaci¬

ous; bracteolis -11X5 mm., oblongis, obtusis, externe glabres-

centibus vel puberulentibus, intus glabris, subpersistentibus.

Alabastrum -10 X 3.5 mm., lanceolatum, obtusum. Calyx

externe conferte breviter pallide griseo-tomentosus, intus glaber;

lobis subaequalibus, anguste deltoideis, obtusis, basi paullum ex¬

panses. Corolla in vivo pallide flava, petalis lineare-lanceolatis, sub-

acutis, partibus in alabastro expositis breviter pubescentibus.

Stamina 15, verticillis 3, interna 5 quam alia paullum longiora;

filamentis latis, verticillis conjunctis, sub antheris abrupte atte-

nuatis; antheris anguste oblongis; aristis quam antheras 2-3-plo

longioribus, non reflexis. Ovarium ovoideum, dimidio distali

puberulenti vel glabrescenti, dimidio basali glabro; stylopodio in-

distincto; stylo quam ovarium paullum longiore, filiformi, glabro.

Pedicellus in fructu -2 mm. longus, -4 mm. diam. Calyx in fructu

breviter puberulens vel glabrescens, ad basim impressus; lobis

longioribus 3,-18 X 3 cm., oblongo-spatulatis, obtusis, basim

versus -2 cm. latis attenuatis, partibus basalibus -1.6 X 2.2 cm.,

late ovatis, saccatis, incrassatis; lobis brevioribus 2, -13 X 0.8

cm., basi ut in lobis longioribus. Nux -3.7 X 2.5 cm., late ovoidea,

plane conferte pallide aureo-alutaceo-tomentosa, ad apicem coni-

cum abrupte attenuata, stylopodio brevi acuto.

Collections. North Borneo: San 15071, 15122, 15072. Beaufort:

San 15106, Pangi. Indonesian Borneo: bb 29636, 29666, 29663,

29702, 31420, 31419, Muara Tewe: bb 32780, 32781, 32774,

32779, Tarakan; Meijer 2218, Kostermans 8711, Nunukan, Tara-

kan; bb 29306, 29309, Bulungan; bb 31260, 31200, 31201,

273

Gardens Bulletin , 5.

2 Cm

2Cm

>Plate 10. S. amplexicaulis.

a, Flowering twig, b, Leaf, c, Sepals (left, inner; right, outer), d, Ovary and androecium.

e. Ovary, f, Fruit, g, Nut.

(a-e from Jacobs 5323; f, g, from Brun 3261).

274

Vol. XIX. (1962)

31202, 31203, 29636, 27023, 28121, 27006, 27024, 30208,

30209, 30207, 26429, Melawi. Brunei: Kep 30374, Temburong;

Brun 751, Base of Bt. Tanggoi, Temburong; Brun 3151, 3006,

Bangar; S 5719, Kuala Belalong; Kep 30572, 30574, 35550,

35551, Tutong R.; Kep 28653, Bt. Kerita, Ladan Hills; S 1675,

Mile 3, Ladan Hills; Kep 30633, Bt. Ladan; Kep 37086, 37087,

Bt. Arong F.R.; Kep 32590, Balai Rotan; Kep 34559, Kuala Sela-

dan, Tepiku; Kep 34566, Pak Andaki; Kep 34584, Pak Gabal;

Kep 34596, Bt. Menggelang; Kep 48201, Balai Halim; Kep 80094,

S 1860, 1927, 1937, 5580, Brun 555, 3261 (Holotypus in Herb,

Kew), Andulau F.R.; Kep 30386, Kep 30387, S 1633, 1937, Kep

30389, 30460, 39638, Labi Hills F.R.; Kep 48184, Bt. Kenapol,

Labi; S 2131, Bt. Teraja. Sarawak: S 247, Semengoh F.R.; S 6502,

Simanggang; Clemens 21965, Sarawak; S 1683, Lawas; Jacobs

5323 Belaga.

SHOREA ANDULENSIS Ashton s.n.

S. pauciflorae King affinis, sed corolla ianthina, ovario glabro*

lamina minore subtus glauca differt.

Ramuli, perulae, costa media supra, stipulae, petiolique, breviter

conferte pallide flavido-alutaceo-tomentosi, costaeque subtus

sparsius eodem tomentosi. Ramuli apicem versus 1.5-2 mm. diam.,

primo compressi, dein teretes, sicco nigrescentes, lenticellis minutis

orbicularibus pallidioribus punctati, leves, glabri, angusti; inter-

nodis c. 1-1.5 cm. longis, brevibus; cicatricibus stipularum ineon-

spicuis. Gemma 3-6 X 1.5—3 mm., ovoidea, compressa, subacuta.

Stipulae -14 X 5 mm., hastatae, subacutae. Lamina 6-9 X 3-5

cm., elliptico-ovata. basi obtusa, apice in acumen -8 mm. longum

attenuato; in arboribus maturis subtus glauca; costis lateralibus

utrinsecus 10-13, rectis, marginem versus abrupte curvatis, subtus

angustate prominentibus, angulo 40°-50° exorientibus basi ex-

cepta; intercostis angustis, scalariformibus, sinuatis, a costis

lateralibus diagonale exorientibus. Petiolus 1-1.2 cm. longus,

1-1.5 cm. latus, sicco cremeo-glauco-tomentosus, rugosus. Lamina

delapsa supra pallide purpureo-brunnescens, subtus plus minus

glauca, dein saepe cere glauca attrita fusco-rufo-brunneam,

niargine saepe paullum revoluto. Racemi -12 cm. longi, semel vel

bis ramosi, stipitis basi in fructu -2, 5 mm. diam., terminales vel

axillares, ex integro teretes vel paullum compressi, semper breviter

conferte griseo vel pallide ferrugineo-tomentosi; ramuli —10 flores

distichos gerentibus; bracteis bracteolisque -3 X 1-25 mm., ovatis,

subacutis, externe breviter griseo-pubescentibus, intus glabres-

centibus. Alabastrum —8 X 2.5 mm., anguste ovoideum, sub-

acutum, calyce expanso. Calyx externe conferte breviter tomen-

tosus, intus puberulens; lobis subaequalibus, ovatis, acutis vel

paullum acuminatis, internis 2 quam externis 3 paullum acumina-

tioribus. Corolla in vivo pallide ianthina, ad marginem pallidior;

275

Gardens Bulletin, 5.

Plate 11. S. andulensis.

a, Flowering twig, b. Fruit, c, Part of lamina undersurface, d, Sepals (1, outer; 2, inner)*

e, Ovary and androecium. f, Ovary.

(a-f, from Brun 3263).

276

Vol. XIX. (1962).

petalis lineari-lanceolatis, anguste acutis, nectis, partibus in ala-

bastro expositis breviter tomentosis, basim versus saccatis. Stamina

15, verticillis 3, bina cum solis alternata; interna 5 quam alia vix

longiora; filamentis basim versus applanato-dilatatis, ad medium

abrupte attenuatis, antheris late oblongis; aristis quam antheras

3-plo longiori, rectis, non reflexis, internis 5 prope apicem styli

attingentibus. Ovarium parvum, globosum, glabrum, stylopodio

indistincto; stylo quam ovarium 2-plo longiori, filiformi, dimidio

basali sparsim tomentoso, aliter glabro. Pedicellus in tructu c. 3

mm. longus, c. 1 mm. diam., gracilis. Calyx in fructu breviter

pubescens, basim versus confertius pubescens, in aetate maturandi

sicco aureo-luteus; lobis longioribus 3, -5 X 1*2 cm., late

spatulatis, obtusa, basim versus -2.5 mm. latis attenuatis, partibus

basalibus c. 6 X 4 mm., anguste ovatis, incrassatis, saccatis; lobis

brevioribus 2, -2.5 X 0.2 cm., linearibus, subaequalibus, basim

versus ut in lobis longioribus. Nux —1.4 X 0.7 cm., ovoidea,

breviter pallide griseo-alutaceo-pubescens, ad stylopodium —1 mm.

acutum abrupte attenuata.

Collections. Brunei: Brun 5435, 3263 (Holotypus in Herb. Kew),

3030, 3275, San 17523, Andulu F.R.; Brun 3332, Bt. Patoi; Brun

878, Bt. Bedawan; S 2122, Bt. Teraja; Brun 173, R. Ingei. Sara¬

wak: S 5637. Bt. Bubong Rumah, Lawas; Brun 809, Sourabaya-

Merapok track, Kayangeran East F.R., Lawas.

SHOREA ANGUSTIFOLIA Ashton s.n.

S. xanthophylla Sym. affinis, sed petiolo pubescenti, lamina

minore, coriaceore, sine costis lateralibus differt.

Perulae petiolique semper pilis brevissimis pallide brunneis

punctati; ramuli stipulaeque pilis minutis pallidis brunneis primo

sparsim punctati, dein glabrescentes. Ramuli apicem versus 1.5—2

mm. diam., recti, teretes, pallide flavido-brunnescentes, rugosi, fissi,

facie floccis parvulis chartaceis exfolia. Gemma 0.5—1.5 X 1

mm., parva, ovoidea, obtusa. Stipulae —3 mm. longae, lineares,

fugaces. Lamina novella in vivo fusco-violescens; lamina 8—14 X

2.5-4 cm., ovato-lanceolata, coreacea, basi cuneata, apice in

acumen 1—1.5 cm. longum gradatim attenuato; costis lateralibus

utrinsecus 8-9 (10), vix prominentibus, curvatis, remotis, angulo

50°—60° exorientibus; costa media subtus vix elevata, intercostis

conferte scalarifocmibus. Petiolus 6-8 mm. longus, c. 1.5 mm.

latus, brevis fissis minutis, sicco distincte pallide brunneus vel

pallide cremeo-griseus. Lamina delapsa griseo-viridis, non crispata

sed margine saepe undulato. Racemi —10 cm. longi, ordine semel

vel bis ramosi, stipitis basi in fructu —1.5 mm. diam., terminates

vel axillares, graciles, laxi, ex integro teretes, breviter pallide

griseo-alutaceo-pubescentes; ramulis —2 cm. longis, —8 flores dis-

tichos gerentibus; bracteolis parvis, fugacibus. Gemma -2.5 X 1-5

mm., lanceolata, acuta. Calyx exteme breviter tomentosus, intus

277

Gardens Bulletin, S.

Q.25Mm

Plate 12. S. angustifolia.

05Mm

a, Flowering twig, b, Sepals (above, outer; below, inner), c, Corolla, d, Stamens (left,

frontal view; right, lateral view), e, Ovary.

(a-e from Brun 778).

278

Vol. XIX. (1962).

glaber, lobis ovatis, acutis, lobis internis 2 quam externis 3

tenuoribus, ad basim costrictioribus. Corolla in vivo pallide flava,

aetate expandi basim versus imbricata, scyphiformis, apicem

versus expansa; petalis linearibus, partibus in alabastro expositis

breviter tomentosis, apicem versus nectis. Stamina 15, in verticillis

3 inaequalibus, bina cum solis alternata; filamentis basim versus

dilatatis, apicem versus filiformibus; antheris oblongis; aristis quam

antheras 1—2-plo longioribus, apicem styli attingentibus, apicem

versus ciliatis. Ovarium ovoideum, basi glabra, aliter breviter

tomentosum, stylopodio conico, tomentoso, stylo brevi glabro;

stylo styloporioque ovarium aequanti. Fructus ignotis.

Collections. North Borneo: Kep 80302, Gaya F.R. Indonesian

Borneo: bb 17915, 17919, 17760, Tidung; bb 12583, E. Kutai;

bb 31774, Melawi; bb 11050, 21201, 21199, 21184, 21158,

21187, 21155, 22754, Puruktjau; bb 18988, 19555, Berau; bb

11200, Bulungan; bb 20143, Sampit; bb 20627, 20629, Boven

Mahakam; bb 19987, Boven Dayak. Brunei: Brun 3393, 5682,

Kuala Belalong; Brun 520, Bt. Patoi; Brun 748, Kuala Sekurop,

Temburong; Brun 778, Kuala Temburong Machang, Temburong

(Holotypus in Herb. Kew).

SHOREA ASAHI Ashton s.n.

S. maxwellianae King, S. biawak Ashton aflinis, sed lamina coria-

cea, costa subtus supraque applanata differt.

Novellae primo brevissime pubescentes, dein glabrescentes,

stipulae persistenter. Ramuli apicem versus c. 0.75 mm. diam.,

graciles, teretes, fusco brunnei vel nigri, lenticellis minutis palli-

dioribus conferte punctati; cicatricibus stipularum parvis, horizon-

talibus, inconspicuis. Gemma c. 1.5 X 1 nun., parva, falcata, sicco

nigrescens. Stipulae -5 mm. longae, anguste deltoideae, fugacae.

Lamina 6-10 X 3-5 cm., ovata, coriacea, non glauca, basi late

cuneata, subaequali, apice in acumen c. 1 cm. longum anguste

attenuate; costis lateralibus utrinsecus 6-7, angustis, curvatis, ap-

planatis, angula c. 60° exorientibus; costa media indistincta, subtus

supraque applanata; intercostis scalariformibus, confertis, angus-

tissimis, indistinctis, a costa media angulo 90° exorientibus vel

sursum directis, marginem versus a costis lateralibus angulo c.

90° exorientibus. Petiolus c. 1 cm. longus, c. 1 mm. diam., sicco

nigrescens. Lamina delapsa supra obscure purpureo-brunnescens,

subtus pallide flavido-brunnescens, ex integro nitens. Racemi -12

crn * longi, semel ramosi, stipitis basi in fructu -1 mm. diam.,

terminates vel axi Hares, ex integro graciles, teretes, recti, conferte

cremeo-tomcntosi; ramulis -7 mm. longis, brevibus, ‘zigzag’, -4

Hores secundos conferte gerentibus; bracteolis —2 mm. longis,

e Hipticis, acutis, puberulentibus, fugacis. Alabastrum —2 mm.

■°ngum, globosum. Calyx externe breviter pubescens, intus

glaber; lobis externis 3 ovato-deltoideis, subacutis; lobis internis

279

Gardens Bulletin, S.

a, Flowering twig, b, Flower bud. c, Sepals (inner above; outer below), d, Petal*

e, Stamen (from inner side), f, Ovary.

(a-f from Kep 37183).

280

Vol XIX. (1962).

2 quam illis minoribus, latius ovatis, minoribus acutis. Corolla in 1

vivo alba; petalis elliptico-oblongis, acutis, externe breviter pub-

escentibus, intus puberulentibus. Stamina c. 30, subaequalibus;

filamentis basim versus applanato-dilatatis ad medium abrupta

attenuatis, apicem versus filiformibus, breviter barbatis; antheris

anguste oblongis, loculis interioribus quam exterioribus majoribus,

apice breviter barbato; aristis quam antheras 2-plo longioribus,

crassis, apicem versus barbatis. Ovarium stylopodiumque ovoi-

deum, conferte tomentosum, in stylum breve gracile glabrum at-

tenuatum. Pedicellus in fructu -2 mm. longus. Calyx sparsim alu-

taceo-pubescens; lobis -4X4 mm. brevibus subaequalibus nucem

adpressis. Nux -11X6 mm. oblongo-ovoidea acuta conferte alu-

taceo-pubescens.

Collections. Brunei: Kep 37183, Bt. Bedawan (Holotypus in

Herb. Kepong); Brun 772, Kuala Temburong Machang, Ulu Tem-

burong; Brun 5632, Bt. Ulu Tutong; S 1183, Bt. Teraja. North

Borneo: San 24279, M. 5, Telupid road, Beluran.

SHOREA BIAWAK Ashton s.n.

S. maxwellianae King affinis, sed lobis calycis in fructu quam

nucem brevioribus differt.

Ramuli petiolique primo breviter pallide alutaceo-pubescentes,.

dein glaber, perulae semper iidem pubescentes, laminae glabrae.

Ramuli apicem versus -1 mm. diam., graciles, teretes, multum

ramosi, dein pallide vel fusco-brunnescentes, leves vel lineare

striati; internodis 1.2-3 cm. longis; cicatricibus stipularum

minutis, brevibus, inconspicuis. Gemma -1 mm. longa, parva.

Stipulae ignotae. Lamina 6-10 X 2.5-4.5 cm., tenuis, ovata,

basi obtusa vel late cuneata, saepe subaequali, apice in acumen

-1.5 cm. longum angustum vel caudatum attenuato; costis late-

ralibus utrinsecus 5-6, angustis, subtus paullum prominentibus,

supra paullum depressis, curvatis, angulo 50°-60 exorientibus;

domatiis parvis puberulentibus axillaribus; intercostis indistinctis,

scalariformibus, a costa media c. 90° exorientibus; costa media

subtus paullum prominenti, supra paullum prominenti vel appla-

nata. Petiolus 7-10 mm. longus, -1 mm. diam., gracilis, sicco

nigrescens. Lamina delapsa sicco ex integro pallide virido-brunnea.

Racemi -14 cm. longi, semel ramosi, stipitis basi in fructu -1

mm. diam., terminales vel 1-axillares, ex integro teretes, semper

breviter sparsim pallide cremeo-brunneo-pubescentes; ramulis -6

mm. longis, brevioribus, paullum secundis, -6 flores secundis con¬

ferte gerentibus; bracteolis -2 mm. longis, ovatis, acutis, bre¬

viter puberulentibus, fugacis. Alabastrum -1.5 mm. longum, glo-

bosum. Calyx externe breviter tomentosus, intus glaber; lobis

late ovatis, acutis, subaequalibus, internis 2 quam externis 3 paul¬

lum brevioribus, tenuioribus, latioribus. Corolla in vivo cremea,

petalis ellipticis, obtusis, externe breviter puberulentibus, intus

281

Gardens Bulletin, S.

Plate 14. S. biawak.

a, Flowering twig, b, Flower bud. c, Sepals (inner above, outer below), d, Petal, e,

f, Stamen (inner side), g, Fruit.

(a-f from Brun 3005, g from Brun 3369).

Vol. XIX. (1962).

glabris. Stamina c. 35; filamentis gracilibus, glabris, attenuatis;

antheris anguste oblongis, apicem versus attenuatis; loculis in-

terioribus quam exterioribus paullum maioribus, apice breviter

barbato; aristis quam antheras paullum brevioribus, conferte barb-

btis. Ovarium stylopodiumque ovoideum vel pyriforme, conferte

tomentosum, stylo brevi glabro. Pedicellus in fructu -2 mm. longus,

brevis. Calyx quam nucem brevior; lobis subaequalibus, -7 X ^

mm., suborbicularibus, obtusis, tenuibus, ad nucem conferte ad-

pressis, semper breviter pallide alutaceo-puberulentibus. Nux -13

X 10 mm., subglobosa, conferte pallide alutaceo-pubescens, apice

in stylopodio -1.5 mm. lineari abrupte attenuato.

Collections. Brunei: Brun 3356, 3351, Ulu Senuko, Labu; Brun

3385, Kuala Belalong; Brun 3005, 3369 (Holotypus in Herb. Kew)

Bangar.

SHOREA BULLATA Ashton s.n.

5. flemniichii Sym. atlinis, sed lamina maiore, sparsius tomen-

tosa, paullum bullata, staminum aristis longissimis undulatis, stylo

longiore differt.

Stipulae externe, ramuli, petioli, costae subtusque semper

aspere scabrido-fulvo-cristato-tomentosi, stipulae intus breviter

pubescentes. Ramuli apicem versus c. 2 mm. diam., primo cristato-

tomentosi, dein glabri, tenue chartaceo exfoliati, pallide brunnei,

teretes, multum ramosi; internodis 1-2 cm. longis, brevibus; cica-

tricibus stipularum brevibus, transversis, tomento obscuris. Gemma

3-4 X 2-3 mm., late ovoidea, subacuta. .Stipulae -7X2 mm.,

anguste hastatae, acutae. Lamina 6.5-10 X 3-4.5 cm., elliptico-

vel oblongo-obovata, bullata; saepe tenue concava, basi anguste

obtusa vel late cuneata attenuata, apice obtuso vel in acumen -6

mm. longum breviter attenuato; margine revoluto; costis laterali-

bus utrinsecus 10-12, angulo variabile exorientibus; intercostis laxe

scalariformibus, costatis, a costis lateralibus 90° exorientibus;

costis supra depressis. Petiolus 0.8-1 cm. longus, c. 1.5 mm. diam.,

brevis, sicco fulvo-tomentosus. Lamina delapsa supra pallide

griseo purpurescens, subtus fusco-rubro-purpurescens; tomento

cristato distincte aureo-fulvo. Racemi -11 cm. longi, semel vel bis

(terminales) ordine ramosi; stipitis basi in fructu -1.5 mm. diam.,

terminates vel axillares, ex integro teretes, laxi, ad basim conferte

rubro-brunneo-tomentosi, ad apicem brevius pallidius tomentosi;

ramulis -3.5 cm. longis, -8 flores plus minus secundis gerentibus;

bracteis ignotis; bracteolis -6 X 3.5 mm., late ellipticis, obtusis,

breviter pallide griseo-puberulentibus. Alabastrum -15X3 mm.,

lineare obtusum. Calyx parvus, externe conferte pallide griseo-

tomentosus, intus glaber; lobis subaequalibus, late ovatis vel

suborbicularibus. Corolla in vivo pallide cremea, basim versus

rosea; petalis linearibus, obtusis, partibus in alabastro expositis

conferte tomentosis, corolla delapsa paullum connata. Stamina 15,

283

Gardens Bulletin , S.

Plate 15. S. bullata.

a, Flowering twig, b, Part of lamina undersurface, c, Flower bud. d, Outer sepal (from

outside, left; from inside, right), e, Petal, f, Androecium and gynoecium. g, Gynoecium.

h, Fruit.

(a-g from Brun 2003a; h from Brun 2003c).

284

Vol XIX . (1962).

verticillis confertissimis 3, bina cum solis alternata, antheris inter¬

ims 5 supra aliis extrusis, externis 10 subaequalibus; filamentis basi

connatis, applanato-dilatatis, ad medium abrupte attenuatis,

apicem versus filiformibus; antheris oblongis; aristis quam antheras

-4-plo longioribus, apicem styli superantibus, gracillimis, sinuato-

crispulatis. Ovarium stylopodiumque ovoideum, conferte breviter

tomentosum; stylo quam illis prope 2-plo longiori, dimidio basali

incrassato, setoso. Pedicellus in fructu -1 mm. longus. Calyx

basim versus sparsim pallide alutaceo-pubescens, apicem versus

glabrescens; lobis longioribus 3, -9 X 1.3 cm., spatulatis, obtusis,

basim versus angustatis -5 mm. latis; partibus basalibus -6 X 6

mm., prominente saccatis incrassatis; lobis brevioribus 2, 2—5 X

0.3 cm., linearibus, obtusis, partibus basalibus ut in lobis longi¬

oribus. Nux -10 X 7 mm., ovoidea, breviter conferte cremeo-

tomentosa, acuta.

Collections. Brunei: Brun 924, Andulau F.R.; S 5773, Bt.

Biang; Brun 2003 (Holotypus in Herb. Kew), S. 5798, Bangar.

SHOREA DOMATIOSA Ashton s.n.

S. atrinervosae Sym. affinis, sed fructu maiore, lamina tenuore,

petiolo angustiore, longiore, costis lateralibus angustioribus,

domatiis prominentibus differt.

Domatiae stipulae externeque pubescentes, aliter novelli glabri.

Ramuli apicem versus c. 1.5 mm. diam., graciles, multum ramosi,

sicco saepe compressi vel rugosi, fusco-brunnei vel nigrescentes,

leves, lenticellis minutis pallidis rotundatis punctati; internodis 1-2

cm. longis; cicatricibus stipularum brevibus, pallidis, horizontal-

ibus, inconspicuis. Gemma c. 4 X 2 mm., ovoidea, acuta. Stipulae

c. 6 X 2.5 mm., ovatae, anguste obtusae, basi paullum constricta,

caducae. Lamina 6.5-10 X 3-7 cm., late ovata vel obovata,

tenuis, subtus in arboribus maturis glaucescens, basi obtusa vel

subcordata, apice in acumen c. 8 mm. longum subabrupte at¬

tenuate; costis lateralibus utrinsecus (8)9-10(12), angustis, cur-

vatis, subtus paullum prominentibus, angulo 45°-65° exorienti-

bus, domatiis axillaribus prominentibus; intercostis angustis, con-

fertis, scalariformibus, sinuatis. Petiolus 1.5-2.5 cm. longus, c. 1

mm. diam., gracilis, geniculatus, dimidio apicali incrassato, sicco

nigrescens. Lamina delapsa supra purpureo-grisea, subtus in

arboribus novellis ex integro griseo-brunnea, in arboribus maturis

glauca costis nigris. Racemi, bractei, bracteolique ignoti. Ala-

bastrum -1.3 X 4 mm., fuseforme. Calyx externe breviter pube-

scens, intus glaber; lobis externis 3 late ovatis subacutis, internis

2 anguste ovatis, acutis. Petala linearia, in alabastro vix contorta,

obtusa, partibus in alabastro expositis sparsim pubescentibus, aliter

glabra. Stamina c. 25-30; filamentis basim versus applanatis. ad

antheris attenuatis; antheris late oblongis, loculis internis 2 quam

externis 2 brevioribus; ad apicem sparsim setosis; aristis quam

285

Gardens Bulletin, S.

O 2 Mm O 0.5Mm O

q O 2 Mm

05Mm

O 0.5 Mm

O /Cm

Plate 16. S. domatiosa.

a, Leafy twig, b, Fruit, c. Nut. d, Sepals (left, inner; right, outer), e, Petal, f, Stamen

(frontal view), g, Stamens (lateral view), h, Ovary.

(a from S 1782; b, c, from San 16601; d-h from San 21492).

286

Vol XIX. (1962).

antheris i-plo longioribus, crassis, ± conferte setosis. Ovarium

ovoideum, conferte tomentosum basi excepta; stylopodio quam

ovarium. 1^-plo longiori, longissimo, basim versus pubescenti,

aliter glabro. Pediceilis in fructu -1 mm. longus, crassus. Calyx

glaber; lobis longioribus 3,-13 X 3 cm., late spatulatis, obtusis,

basim versus angustatis -8 mm. latis; partibus basalibus -1.8 X

1.6 cm., ellipticis saccatis incrassatis; lobis minoribus 2, -7 X 1.2

cm., anguste spatulatis, partibus basalibus ut in lobis longioribus.

Nux -3.5 X 2.5 cm., ovoidea, conferte breviter cremeo-pubescens,

ad apicem acutum attenuata, stylopodio -1 cm. longo filiformi.

Collections. North Borneo: San A 308, Saliwagon M. 64, Beau¬

fort; San 15164, A 306, A 302, 16604, 16601 (Holotypus in

Herb. Kepong), Ulu Sipitang; San 21492, Tawau; Sarawak: S 1782,

Bt. Berayong, Lawas. Brunei: S 5726, Brun 3395, 5665, 5210,

Kuala Belalong; Brun 305, S. Batu Apoi-Sebatu watershed; Brun

888, Base of Bt. Bedawan; Kep 39645, Labi F.R.

SHOREA FAGUETIOIDES Ashton s.n.

Shoreae faguetianae Heim affinis, sed lamina tenuiore, latiore,

basi saepe subcordata, petiolo longiore, differt.

Ramuli novell interdum pallide alutaceo-tomentosi, interdum

glabri, perulae, stipulae laminaeque glabri. Ramuli apicem versus

-3 mm. diam., primo subcompressi, sicco nigri, dein teretes, pallide

brunnescentes, minute striati vel rugulosi. Gemma 3-4 X 2.5

mm., obtuso-ovoidea, glabra, sicco nigra. Stipulae -1.7 X 0.6

cm., anguste deltoideae, saccatae, ad apicem obtusae, tarde

caducae. Lamina 12-18 X 5-7 cm., anguste ovata, tenuis, basi

late cuneata vel subcordata, subaequali, apice in acumen -2 cm.

longum gradatim attenuato; costis lateralibus utrinsecus 10-15,

angustis sed conspicuis, 60°—70° exorientibus, intercostis scalari-

formibus, a costis lateralibus diagonale exorientibus. Petiolus 2.2

-3 cm. longus, -2 mm. diam., sicco, niger. Lamina delapsa fusco-

virido-brunnescens, costis subtus promincnte nigris, sicca involuta.

Racemi -15 cm. longi, ordine 2-3 ramosi, stipitis basi in fructu

-1.2 mm. diam., terminales vel axillares, ex integro teretes, laxi,

semel breviter pallide griseo-brunneo-pubescentes, sicco rugosi;

ramulis -2 cm. longis, -7 flores gerentibus; bracteolis parvis, fuga-

cibus. Alabastrum-2.5 X L5 mm., parvum, ellipticum, obtusum.

Calyx externe conferte breviter pubescens, intus glaber; lobis ova-

tis, acutis, internis 2 quam externis 3 paullum minoribus, tenuiori-

bus, ad basim constrictioribus. Corolla aetate florendi pallide

cremea, ad basim imbricata et gracile scyphiformis, apicem versus

expansa; petalis linearibus, apices versus spiraliter contortis, par¬

tibus in alabastro expositis pubescentibus. Stamina 15, verticillis

3 inaequalibus, bina cum solis alternata; filamentis angustis, at¬

tenuate, basi dilatata, paullum gibbosa, apice filiformi; antheris

subglobosis; aristis angustis, quam antheras 3-plo longioribus,

287

Gardens Bulletin, S.

O 0.5 Mm

Plate 17. S. jaguetioides.

a, Fruiting twig, b, Inflorescence, c, Flower, d, Sepals (left, inner; right, outer), e, Ovary,

f, Stamens (frontal view), g, Stamen (lateral view).

(a from S 1771; b-g from S 1804).

288

Vol. XIX. (1962).

apicem versus paullum ciliatis, internis 5 apice stylo superantibus.

Ovarium ovoideum, sparsim tomentosum, stylopodio cylindrico,

quam ovarium breviore confertius tomentoso, stylo glabro. Fructus

subsessilis. Calyx in fructu ad pedicellum attenuatus, basim versus

puberulens, apicem versus glabrescens; lobis longioribus 3, -7 X

-1.5 cm., late spatulatis, tenuibus, obtusis, basim versus -4 mm.

latis angustatis; partibus basalibus -5 mm. lata, prominente tuber-

culatis, incrassatis; lobis brevioribus 2, -5 X 0.7 cm., aliter ut in

lobis longioribus. Nux -20 X 6 mm., oblongo-obovoidea, pallide

alutaceo-tomentosa, ad stylopodium -3 mm. longum angustum

attenuata.

Collections. Indonesian Borneo: bb 22875, G. Pering, Sanggau:

bb 29609, Long Bleh, W. Kutei; bb 18294, Tidung. Sarawak:

S A 0301, Bt. Ngili, Sadong; S A 069, Nanga Mujan, Batang Ai;

S 1804, Ulu Berawan, Limbang; S 1771, Bt. Pengalih, Lawas

(Holotypus in Herb. Kew). Brunei: Kep 35674, Labi Hills F.R.;

Kep 35466, Bt. Patoi; Kep 80137, Peradayan F.R.; S 1664, Biang;

S 5805, Bangar; Brun 3340, 5645, S 5810, Bt. Patoi; S 5733, 5702,

Kuala Belalong.

SHOREA FLAVIFLORA Wood ex Ashton:— 5. flaviflora Wood

msc.

S. kunstleri King affinis, sed floribus maioribus, calyce glabro

fimbriato incrassato, antheris magnis reniformibus, lamina attenua-

tiore undulata differt.

Ramuli, petioli, laminaeque primo pallide griseo-pubescentes,

cristis minutis adpressis stellatis, dein glabrescentes, perulae sti-

pulae externeque semper pubescentes, stipulae intus semper puber-

ulentes. Ramuli apicem versus 2-3 mm. diam., ex integro teretes,

leves, dein glabri, pallide brunnei; internodis 1.5-4 cm. longis;

cicatricibus stipularum c. 1.5-2 mm. longis, angustis, sursum

directis, ramulos semicircumdatis. Gemma 6-12 X 1.5-3 mm.,

falcata, acuta. Stipulae -24 X 7 mm., anguste hastatae, fugacac.

Lamina 12-18 X 5.5-7 cm., tenuis, anguste ovata, basi obtusa

vel anguste cuneata, apice in acumen -1.5 cm. longum angustum

attenuato; costis lateralibus utrinsecus 6-7, curvatis, remotis,

angustis, subtus prominentibus, domatiis parvis axillaribus, angulo

40°-50° exorientibus; costa media supra applanata; intercostis

angustissimis, a costis lateralibus diagonale exorientibus, conferte

scalariformibus. Petiolus 2-3 cm. longus, c. 1.5 mm. diam., sicco

rugoso-nigrescens. Lamina delapsa supra griseo-brunnescens, sub¬

tus ferrugineo-vel rubro-nigrescens, margine undulato. Racemi -15

cm. longi, bis ramosi, stipitis basi in fructu -2 mm. diam., ter-

minales vel axillares, pendentes, ex integro teretes, sicco rugosi,

brcviter sparsim pallide griseo-pubescentes vel glabrescentes;

ramuli -12 flores distichos gerentibus; bracteolis -5 X 2.5 mm..

289

Gardens Bulletin , S.

/Cm

Plate 18. S. .flaviflora.

a, Flowering twig, b, Flower bud. c, Outer sepal, d, Corolla, e, Ovary and androecium.

f, Ovary, g. Fruit.

(a-f from Brun 5664; g from Brun 5211).

290

Vol. XIX. (1962).

ellipticis, obtusis, externe conferte breviter pallide griseo-tomen-

tosis, intus sparsim pubescentibus. Alabastrum -14 X 4.5 mm.,

fuseiforme, obtusum. Calyx glaber, margine fimbriato excepto;

lobis aequalibus, late ovatis, incrassatis, ad corollam conferte ad-

pressis. Corolla in vivo fusco-fiava, petalis lineare-lanceolatis,

acutis, basi firmiter connatis, multum contortis nectisque, partibus

in alabastro expositis breviter tomentosis, aetate florendi dimidio

basali imbricato late scyphiformi. Stamina 15, verticillis inaequa-

libus 3; filamentis basim versus dilatato-applanatis, apicem versus

abrupte attenuatis, filiformibus; antheris magnis, reniformibus,

loculis interioribus quam exterioribus paullum minoribus; aristis

quam antheris 3-plo longioribus. Ovarium pyriforme, breviter con¬

ferte tomentosum; stylo ovarii stylopodiique aequali, dimidio supra

antheris longissimis superanti, filiformi, glabro. Pedicellus in fructu

c. 3 mm. longus, c. 2.5 mm, latus. Calyx glaber; lobis longioribus

3,-13 X 3 cm., late spatulatis, obtuso, basim versus -11 mm.

latis gradatim attenuatis, partibus basalibus -1 X 12 cm.,

abrupte prominente saccatis, incrassatis; lobis brevioribus 2-9 X

1.3 cm., subaequalibus, basim versus ut in lobis longioribus. Nux

-2.5 X 1.3 cm. (stylopodio incluso), ovoideum, ad stylopodium

-10 X 2 mm. gradatim attenuatum, breviter conferte pallide

alutaceo-tomentosum.

Collections. North Borneo: San 15098, 15109 (Holotype in

Herb. Kepong), Pangi, Beaufort; San 16389, Ranau; San 16475,

Balong. Sarawak: S 3529, Upper Plieran. Brunei: Brun 5664,

5211, Bt. Belalong; Brun 5276, Ulu Ropan-Belalong watershed;

S 5739, Kuala Belalong; Brun 781, Kuala Temburong Machang;

Brun 736, Kuala Sekurop, Temburong.

SHOREA GENICULATA Sym. ex Ashton:— S. geniculata Sym.

nomen nudum.

S. inappendiculatae Burck affinis, sed nuce maxima globosa,

lobis calycis in fructu brevibus subaequalibus, lamina late ovata,

petiolo longissimo differt.

Novellae ex integro glabrae. Ramuli apicem versus c. 3 mm.

diam., crassi, multum ramosi, teretes, pallide brunnei vel nigre-

scentes, leves; internodis 2.5-4 cm. longis, nodis prominentibus,

cicatricibus petiolarum orbicularibus, elevatis, cicatricibus stipula-

rum 2-3 mm. longis, rectis, pallidis, sursum directis, vix distinctis.

Gemma 4-7 X 1-1.5 mm., linearis vel falcata, acuta, sicco nigre-

scens. Stipulae -10 X 3 mm., anguste oblongae, acutae, fugacae.

Lamina 11-17 X 7-13 cm., late ovata vel suborbicularis, cor-

iacea, subtus glauca, basi obtusa vel subcordata, apice in acumen

-8 mm. longum abrupte attenuato vel obtuso; costis lateralibus

utrinsecus 9-11, angulo 40°-50° exorientibus; intercostis sinuatis,

conferte scalariformibus angustis. Petiolus 4-6.5 cm. longus, c. 2

291

Gardens Bulletin, S.

Plate 19. S .. geniculata.

a. Flowering twig, b. Fruit, c. Petal, d, Sepals (left, outer; right, inner), e, Ovary, f, Sta¬

mens (frontal view), g. Stamens (lateral view).

(a, c-g from Brun 3264, b from Brun 1037).

292

Vol. XIX. (1962).

mm. diam., sicco nigrescens. Lamina delapsa supra pallide viride-

brunnescens, subtus glauca : costis nigrescentibus. Racemi -12 cm.

longi, semel ordine ramosi, stipitis basin in fructu -2 mm. diam.;

terminales vel axillares, ex integro plus minus teretes, primo con-

ferte breviter pallide alutaceo-pubescentes, dein glabrescentes;

ramulis -1.5 cm. longis, -4 flores secundos gerentibus; bracteolis

-4 mm. longis, oblongis, puberulentibus, fugacis. Alabastrum -20

X 4 mm., magnum, lineare-Ianceolatum. Calyx externe breviter

pubescens, intus glaber; lobis ovatis, acutis, externis 3 quam

internis 2 paullum longioribus, brevioribus. Corolla in vivo cremea,

petalis -2.5 cm. longis, linearibus, ex integro pubescentibus. Sta¬

mina c. 55, subaequalia; filamentis basi applanato-dilatatis, apicem

versus filiformis, attenuatis; antheris oblongis, glabris, loculis in-

terioribus quam exterioribus paullum minoribus; aristis quam an-

theras paullum brevioribus, sed prominentibus, crassis, setosis.

Ovarium styopodiumque late ovoideum, conferte tomentosum, basi

glabro, apice in stylum breve latum trifurcatum glabrum abrupte

attenuato; stigmate minuto. Pedicellus in fructu -3 mm. longus,

-4 mm. diam., brevis. Calyx quam ngcem brevior, sparsim pallide

griseo-alutaceo-pubescens; lobis c. 1.5 cm. longis latisque, aequa-

libus, incrassatis, deltoideis, subacutis, apicibus ad basim nucis

adpressis. Nux -5 cm. longa lataque, globosa, conferte breviter

pallide griseo-alutaceo-tomentosa, sicco longitudine striata, trans¬

verse rugosa, apice obtusa, stylopodio c. 1.5 mm. longo brevi.

Collections. Brunei: S 1852, Tapangan Sapi; Kep 37074, Bt.

Ligi; S 1902, 1929, Kep 80092, 48486, 48452, 47041, Brun 3264

(Holotypus in Herb. Kew), 1037, 919, 3060, 3188, 618, San

17470, Andulau F.R.; Kep 35553, 35549, Pengkalan Ran; Kep

12988, Tutong R., hills west of Ukong.

SHOREA ISOPTERA Ashton :—Neohopea isoptera Wood msc.

Shoreis aliis facile potest distinguere; alabastro globoso, stamini-

bus 15 glabris, filamentis latis brevibus, aristis latis brevibus, stylo

brevissimo, receptaculo lato tenui, lobis calycis expansis subaequa-

libus distinguitur.

Novellae glabrae, racemis exceptis. Ramuli apicem versus -1.5

mm. diam., ex integro graciles, fusco-rubro-brunnei vel nigrescentes,

leves, teretes; internodis 2-4 cm. longis; cicatricibus stipularum

brevibus, inconspicuis. Gemma -1 mm. longa, parva, globosa,

sicco nigrescens. Stipulae caducae, ignotae. Lamina 9-16 X 5-8

cm., ovata, basi late cuneata vel obtusa in petiolum -2 mm. de-

currenti; apice in acumen -1.5 cm. longum angustum subabrupte

attenuato; margine undulato; costis lateralibus utrinsecus 9-11,

curvatis, subtus paullum elevatis, supra depressis, angulo 45°-65°

exorientibus; intercostis scalariformibus, sinuatis, angustis; costa

media subtus tereti, paullum prominenti, supra plus minus ap-

planata. Petiolus 1.3-2 cm. longus, -1.5 mm. latus, sicco nigre¬

scens. Lamina delapsa ex integro hebete virido-brunnea, supra

293

Gardens Bulletin, S.

O 0.25Mm

Plate 20. S. isoptera.

0.2 Mm

0.5Mm

a, Flowering twig, b, Fruit, c, Flower bud. d, Sepals (left, outer; right, inner), e, Petal,

f, Androecium. g, Stamens (from within), h, Outer stamen (frontal view), i. Ovary,

(a, c-i from Brun 3018; b from Brun 3343).

294

Vol XIX. (1962).

paullum rubrior. Racemi -11 cm. longi, bis ordine ramosi, stipitis

basi in fructu -1.5 mm. diam., terminales vel 1-plures axillares,

ex integro teretes vel paullum commpresses, laxi, super breviter

griseo-puberulentes; ramosis -5 cm. longis; ramulis -1 cm. longis,

brevibus, -6 flores secundos conferte gerentibus; bracteis ignotis;

bracteolis -1 mm. longis, linearibus, pubescentibus, fugacis. Ala-

bastrum -1.5 mm. longum, globosum. Calyx externe breviter

pubescens, intus glaber; lobis subaequalibus, imbricatis, acutis;

externis 3 ovatis, intends 2 deltoideis. Corolla in vivo cremea,

petalis externe conferte tomentosis, intus breviter pubescentibus,

oblongis, obtusis, multum contortis. Stamina 15, verticillis 3, bina

cum solis alternata; filamentis crassis compressis, ad antheras vix

attenuatis; antheris oblongis glabris, 4-loculis, loculis subaequa¬

libus vel exterioribus quam interioribus paullum maioribus; aristis

brevibus, crassis, glabris, apices antherorum non superantibus.

Ovarium stylopodiumque ovoideo-conicum, basi glabro; apicem

versus breviter pubescenti, in stylo brevi glabro abrupte termina-

tum; stigma minuta. Pedicellus in fructu -1.5 mm. longus, brevis.

Calyx ad basim puberulens, apicem versus glabrescens; lobis 5

subaequalibus, -5.5 X L5 cm:, spatulatis, obtusis, in aetate

maturo rotatis, basim versus -5 mm. latis attenuatis; partibus

basalibus paullum expansis, imbricatis; receptaculo -9 mm. diam.,

tenui. Nux -9 X 10 mm., ovoidea, latior quam longa, breviter

pallide-griseo-brunneo-pubescens, apice in stylopodium -4 mm.

longum acutum subabrupte attenuato.

Collections. Sarawak: S 7978, Lundu. North Borneo: San 4394,

3955, Bettotan; San 15111, Beaufort; San A 4318, Tenom San A

307, Saliwagon, M. 64, Beaufort; San 16603, Sipitang; San 15202

(fr. excl.), 15222, Pangi. San A 4336: Ko'ta Belud. Brunei: Brun

5676, Kuala Belalong; S 5563, Bt. Peradayan; Kep 80132, 80133,

Peradayan F.R.; Brun 3343, 3018 (Holotypus in Herb. Kew),

Bt. Patoi.

N.B. The fruit of San 15202 represents Dryobalanops sp.

SHOREA LADIANA Ashton s.n.

5. maxwellianae King, 5. biawak Ashton affinis, sed lobis calycis

in fructu quam nucem brevioribus, subaequalibus, lamina maiore

coriacea, costis differt.

Ramuli petioli novellaeque breviter pubescentes, perulae semper

griseo-tomentosae, lamina glabra. Ramuli apicem versus 1.5-2

mm. diam., graciles, teretes, pallide brunnei, multum ramosi, leves

vel rugulosi, internodis 2-4 cm. longis; cicatricibus stipularum

parvis, paullum deorsum curvatis, inconspicuis. Gemma c. 1 X

0.5 mm., parva, conica. Stipulae ignotae. Lamina 10-14 v 4.5-

7.5 cm., ovata, utrinsecus aequalis vel subaequalis, coriacea, basi

295

Gardens Bulletin , S.

Plate 21. S. Indiana.

a, Flowering twig, b, Fruit, c, Flower bud. d, Sepals (left, outer; right, inner), e, Ovary,

f, Stamens (inner side).

(a, c-f, from S 11076; b from Brun 2622).

296

Vol. XIX. (1962).

obtusa vel late cuneata, apice in acumen 1-1.5 cm. longum

gradatim attenuate); costis lateralibus utrinsecus 5-6, subtus pro-

minentibus, remotis, curvatis, angulo 40°-50° exorientibus; inter-

costis angustis, indistinctis, conferte scalariformibus, a costa media

90° exorientibus; margine saepe paullum revoluto. Petiolus 1-2.2

cm. longus, c. 1.25 mm. diam., sicco nigrescens. Lamina delapsa

supra fusco-rubro-brunnescens, subtus pallidius melleo-brun-

nescens, ex integro nitens. Racemi -15 cm. longi, semel vel bis

ramosi, stipitis basi -2 mm. diam., terminales vel -2 axillares,

recti, ex integro paullum compressi, laxi, semper breviter pallide

cremeo-griseo-pubescentes; ramulis -8 mm. longis, brevibus,

‘zigzag', -9 flores secundos conferte gerentibus; bracteolis -1.5

mm. longis, ovato-deltoideis, acutis, breviter pubescentibus, fugacis.

Alabastrum -1.5 mm. longum, globosum (Alabastrum maturum

ignotum). Calyx externe conferte tomentosus, intus glaber; lobis

suborbicularibus, subaequalibus. Petala late elliptica. Stamina 30-

35; filamentis brevibus, attenuatis; antheris oblongis, loculis in-

terioribus quam exterioribus maioribus, apice breviter barbato;

aristis brevibus, barbatis. Ovarium ovoideum, conferte tomento-

sum, stylo brevi glabro. Pedicellus in fructu -3 mm. longus,

gracilis, tomentosis. Calyx quam nucem brevior, lobis 5 subaequa¬

libus, c. 8 X 8 mm., late ovatis, breviter puberulentibus, subacutis,

incrassatis, tenue saccatis, conferte ad nucem adpressis. Nux -1.7

X 1.4 cm., obovoideo-globosa, conferte breviter pallide alutaceo-

tomentosa, apice in stylopodium c. 1 mm. longum angustum

acutum abrupte attenuato.

Collections. Sarawak: S 11076 (Holotypus in Herb. Kew) 0188,

1625, Semengoh F.R. Brunei: Brun 5543, 2629, 2622, Andulau

F.R.

SHOREA MONTICOLA Ashton s.n.

S. pauciflorae King affinis, sed lamina maiore, incrassatissima,

subtus aureo-lepidota, costis lateralibus utrinsecus 13-16 differt.

Ramuli, petioli, et costae subtus primo breviter pallide flavido-

alutaceo-tomentosi, dein glabrescentes, perulae semper similiter

tomentosae. Ramuli apicem versus -3 mm. diam., terctes, crassi,

lenticellis minutis pallidis ellipticis verruculosis punctatis, pallide

brunnei; internodis 1.5—5 cm. longis; cicatricibus stipularum sub-

amplexicaulibus vel amplexicaulibus, 0.5 mm. latis, pallidis, pro-

minentibus. Gemma 5—10 X 2—4 mm., anguste ovata, acuta,

falcata. Stipulae ignotae. Lamina 8—13 X 5-8 cm., multum coria-

cea, elliptica, primo aureo-lepidota; basi obtusa vel late cuneata,

apice in acumen -5 mm. longum, brevem, late deltoideum,

abrupte attenuato; costis lateralibus utrinsecus 13—16, ad margi-

nem curvatis, angulo 45°-55° exorientibus; intercostis angustis,

conferte scalariformibus, a costis lateralibus diagonale exorienti¬

bus; costa media subtus prominenti, acuta, supra glabra depressa.

297

Gardens Bulletin , S.

Plate 22. S. monticola.

a, Twig with young fruit, b, Part of lamina undersurface, c, Sepals (below, outer; above,

inner), d, Ovary and androecium. e, Stamen (frontal view), f, Ovary.

(a and b from Clemens 31885, c-f from Hallier 678).

298

Vol. XIX. (1962)

Petiolus 2-2.5 cm. longus, c. 2 mm. diam., sicco niger. Lamina

delapsa supra nitens, pallide griseo- vel luteo-brunnea; subtus

griseo- vel aureo-lepidota, costis subtus griseis vel nigrescentibus.

Racemi -12 cm. longi, semel ramosi, stipitis basi in fructu -2 mm.

diam.; terminales vel axillares, teretes, conferte pallide flavido-

brunneo-pubescentes; ramulis -12 flores gerentibus; bracteis

bracteolisque ignotis. Alabastrum -7X3 mm., lanceolatum, sub-

acutum. Calyx externe breviter pubescens, intus glaber; lobis

externis 3 anguste ovatis, obtusis, quam internis 3 li-plo longiori-

bus; lobis internis 2 latius ovatis, acutis, basibus constrictis. Petala

oblonga, obtusa, partibus in alabastro expositis breviter tomentosis.

Stamina 15, verticillis 3, interna 5 quam alia paullum longiora;

filamentis basim versus applanato-dilatatis, ad medium abrupte

attenuatis, apicem versus filiformibus; antheris late oblongis; aristis

quam antheras 2—3-plo longioribus, gracilibus. Ovarium stylo-

podiumque pyriformibus, breviter tomentosum; stylo ovarium

aequanti, filiformi, glabro. Fructus maturus ignotus. Pedicellus in

iructu -1 mm. longus, -3 mm. diam., brevissimus. Calyx basim

versus puberulus, aliter glabrescens; lobis maioribus 3, c. 7 X 1.5

cm., spatulatis, obtusis, supra basi vix attenuatis; partibus basali-

bus -1.5 X 1.5 cm., incrassatis, tenue saccatis; lobis minoribus 2,

-3.5 X 0.6 cm., lineare oblongis, partibus basalibus ut in lobis

longioribus. Nux —1.6 X 1.6 cm., ovoidea, acuta, conferte breviter

tomentosa.

Collections. Indonesian Borneo: Hallier 678, Summit of G.

Semedoen. North Borneo: San 16704, Gunong Lumaku, 1650 m;

San 16914, Kundasan, 1200 m; Clemens 31885, Penibukan,

Kinabalu, 1650 m (Holotypus in Herb. Kew), s.n. (J. Wyatt

Smith), Kg. Kangeran, 800 m; San A 4353, Penibukan, Keningau,

1000 m; San A 4372, Penibukan, Keningau, Trus Madi, 1350 m;

Kep 80446, Tambunan, 1000 m. Sarawak: S 9135, G. Penrissen,

W Sarawak, 1000-14000 m. Brunei: Brun 2360, 2377, Gunong

Pagon Periok, 1600 m.

SHOREA MYRIONERVA Sym. ex Ashton:— S. myrionerva

Sym., nomen nudun.

S. macropterae Dyer, S. slooteni Ashton affinis, sed lamina lon-

gissima, tenui, costis lateralibus multis supra applanatis, subtus

sparsim tomentis, racemis brevibus axillaribus saepe ramifloris,

lobis calycis in fructu prominentius auriculatis distinguitar.

Ramuli petiolique semper aspere cristis pallide badiis punctati,

costae subtus, perulae, stipulaeque brevius cristatae; lamina dein

glabrescens. Ramuli apicem versus 2.5-3.5 mm. diam., crassi, recti,

vix ramosi, teretes, sicco rufo-tomentosi; internodis 2-5 cm. longis;

cicatricibus stipularum -2 X 1 mm., pallidis, connatis. Gemma

299

Gardens Bulletin , S.

Plate 23. S. myrionerva.

a, Flowering twig, b, Fruit,, c, Nut. d, Sepals (outer, left; inner right), e, Ovary, f, Sta¬

men (from inside).

(a, d-f, from Jacobs 5371; b and c from S 1519).

300

Vol. XIX. (1962)

“10 X 7 mm., compressa, perulis foliaceis numerosis laxe com-

posita. Stipulae -15 X 7 mm., subpersistentes, in arboribus novel-

lis persistentioribus, late hastatae, basi subauriculata, apice sub-

acuto; in vivo magentae. Lamina 12-22 X 4.5-9 cm., anguste

onlonga, tenuis, in vivo primo magenta, basi obtusa, apice in acu¬

men -8 mm. longum late deltoideum abrupte attenuato; costis

lateralibus utrinsecus 24-28, angustis, confertis, subtus prominen-

tibus, apicem versus curvatis; intercostis anguste scalariformibus;

a costis lateralibus angulo 90° exorientibus; costa media supra

depressa. Petiolus 1.25-2 cm. longus, 2-2.5 mm. diam., brevis,

crassus, sicco nigrescens. Lamina delapsa subtus fusco-rufo-brun-

nescens, supra fuscius-purpureo-brunnescens, semper plus minus

applanata. Racemi -8 cm. longi, semel ramosi, stipitis basi in

fructu -2 mm. diam., axillares, raro terminales, saepe ramiflori, ex

integro teretes, sicco striati, laxi, basim versus cristis badiis spar-

sim punctati; ramulis brevibus —5 flores distichos gerentibus;

bracteis -8 X 1.5 mm., lineare-lanceolatis, acutis; bracteolis -5

X 3 mm., ovatis, obtusis, externe puberulentibus, intus glabris.

Alabastrum —5X3 mm., ellipticum, rotundatum. Calyx externe

breviter conferte pubescens, intus glaber; lobis subaequalibus,

ovatis, obtusis, externis 3 quam intends 2 paullum angustioribus.

Corolla in vivo fusco-rubra, marginem versus alba (Jacobs), petalis

anguste ovatis, partibus in alabastro expositis conferte flavido-

aureo-tomentosis. Stamina 15, verticillibus inaequalibus 3, interna

5 quam alia multum longiora, prope apicem styli extensa; fila-

mentis crassis, ad antheras gradatim attenuatis; aristis brevibus,

dein reflexis. Ovarium stylopodiumque anguste conicum vel pyri-

forme, breviter pubescens; stylo quam ovarium stylopodiumque

paullum breviori, late filiformi, glabro. Pedicellus in fructu -4 mm.

longus, c. 3 mm. diam., glaber, ad basim fructus dilatatus. Calyx

glaber; lobis longioribus 3,-17 X 2.5 cm., oblongis, tenue char-

taceis, obtusis, basibus ad centra saccatis, incrassatis, auriculis

-1.5 cm. latis tenuibus lateralibus 2; lobis brevioribus 2, subae¬

qualibus, -8 X 07 cm., linearibus saccatis nec auriculatis. Nux

-2.5 X L2 cm., glabra, glaucescens, ellipsoidea, ad stylopodium

c. 1.5 mm. longum acutum celeriter attenuata.

Collections. North Borneo: Kep 80268, 80269, S. Lukutan Beau¬

fort. Sarawak: S 0920/1928, S. Segan, Bintulu; S A 0314, Long

Bunau, Pata; S A 0421, Balleh; S 1519, Merapok, Lawas (Holo-

typus Herb. Kepong); S 1805, Ulu Berawan, Limbang; S 2901, R.

Kenaban, Upper Plieran; Kep 79349, Lambir Hills; S 00279,

Beseri F.R., S. Babong, Suai; Jacobs 5371, Belaga. Brunei: Kep

30480, Ulu Belait; Kep 48268, Andulau F.R.; Kep 39630, Batu

Apoi; Brun 5218, S. Belalong above Kuala Empan, Ulu Tem-

burong; Brun 5200, San 17378, Kuala Belalong.

301

Gardens Bulletin, S.

SHOREA PATOIENSIS Ashton s.n.

S. gibbosae Brandis lamina similis, sed fructus glaber, calyx

quam nucem brevior, lonis subaequalibus differt.

Stipulae perulaeque breviter fimbriatae, gemmae novellae ramu-

lique in arboribus maturis saepe teniussime pulverulento albo-

cerati, foliae glabrae. Ramuli apicem versus c. 1 mm. diam.,

graciles, teretes, leves vel rugulosi, sicco fusco-rubro-nigrescentes

vel nigri. Gemma c. 3 X 1 mm. (in arboribus novellis -4X2

mm.), anguste ovoidea, obtusa, perulis distincte expansis, non

adpressis; sicco nigrescentibus. Stipulae c. 8 X 2 mm., oblongae,

breve acutae. Lamina 5-8 X 2-3.5 cm., tenuis, ovata, utrinsecus

subaequalis, saepe costa media oblique inflecta, basi plus minus

late cuneata, juxta petiolum -1 mm. decurrenti, apice in acumen

anguste c. 1 cm. longuin gradatim attenuato; costis lateralibus

utrinsecus 7-9, angustis quanquam prominentibus, curvatis, angulo

35°-45° exorientibus; costa media supra paullum elevata, subtus

prominente elevata; intercostis conferte scalariformibus. Petiolus

7-10 mm. longus, c. 1 mm. diam., gracilis, sicco nigrescens. Lamina

delapsa griseo-brunnescens, sursum quam deorsum pallior, lutior,

sicca involuta. Racemi -9 cm. longi, semel vel bis ramosi, stipitis

basi in fructu -2 mm. diam., terminales vel axillares, paullum

breves confertique, graciles, ex integro teretes, semper breviter

pallide griseo-brunneo-pubescentes; ramulis -1 cm. longis, -6

flores gerentibus; bracteolis minutis, fugacis. Alabastrum -3.5 X

1.5 mm., lanceolatum, acutum. Calyx conferte rubro-brunneo-

tomentosus; lobis minimis, suborbicularibus, obtusis; lobis internis

2 quam lobis extends 3 paullum tenuioribus, basi costrictis.

Corolla in vivo fusco-flava, aetate florendi basim versus imbricata

scyphiformis, apicem versus revoluta expansa necta; petalis lineari-

bus, partibus in alabastro expositis breviter pubescentibus. Stamina

15, verticillis 3, bina cum solis alternata, inaequalibus; filamentis

paullum brevibus, basim versus dilatatis, ad medium abrupte atte¬

nuate, apicem versus filiformibus; antheris subglobosis; aristis

antheras aequantibus, apicem versus breviter ciliatis, apicem styli

vix attingentibus. Ovarium ovoideum, breviter pubescente; stylo-

podio cylindrico pubescenti, stylo brevi glabro. Pedicellus in fructu

c. 1 mm. longus, brevis. Calyx breviter caduce pallide flavido-alu-

taceus, margine semper fimbrillato; lobis 3-5 mm. longis latisque,

late ovatis, ad basim nucis adpressis, basim versus conjunctis scy-

phum -5 mm. diam. factis. Nux -15 X 8 mm., parva, oblongo-

ellipsoidea, striata, aetate maturandi prope glabra, apice obtuso.

Collections. Brunei: Brun 3324 (Holotypus in Herb. Kew), 3017,

Kep 85465, S 5811, Bt. Patoi; Kep 80127, 80113, 80139, S 5562,

Peradayan F.R.; S 5757, 5710, 5746, Kuala Belalong. Sarawak:

S 1807, Bt. Tegaria, UIu Berawan, Limbang.

302

Vol XIX. (1962)

Plate 24. S. patoiensis.

a, Flowering twig, b, Fruit, c, Sepals (inner, above; outer below), d. Corolla, e. Ovary,

f. Stamens (frontal view), g, Stamens (lateral view).

(a, c-g, from Brun 3324; b from Brun 3324).

303

Gardens Bulletin, S.

SHORE A PILOSA Ashton s.n.:— S. tomentosa Wood msc; non

S. tomentosa Hort. Bog. ex Pierre; non S. tomentosa Miq. ex

Brandis.

S. pinangae Scheff., S. beccarianae Burck, affinis, sed lamina

minore, tomento conferto aureo, cicatricibus stipularum amplexi-

caulibus, domatiis prominentibus tomentosis differt.

Ramuli, perulae, petioli, costa media supra subtusque, et lamina

costaeque subtus semper conferte scabrido-aureo-tomentosi; lamina

supra stipulaeque puberulentes vel glabrescentes. Ramuli apicem

versus 2-2.5 mm. diam., paullum compressi, dein teretes, fusco-

brunnei, multum ramosi; intemodis 2-3 cm. longis; cicatricibus

stipularum angustis, prope transversis, amplexicaulibus. Gemma

3-7 X 1.5-3 mm., ovoidea, obtusa vel subacuta. Stipulae -3.5

X 1.5 cm., in vivo magenta, paullum persistentes, hastatae, acutae,

basi paullum constricta. Lamina 10-17 X 4—7.5 cm., ovata vel

elliptica, paullum coriacea, basi obtusa, paice in acumen -1.2

cm. longum anguste deltoideum subabrupe attenuato; margine

saepe paullum revoluto; costis lateralibus utrinsecus 12-15 con-

fertis, paullum curvatis, angulo 45°-50° exorientibus; doma¬

tiis pilosis axillaribus; intercostis conferte scalariformibus, a costis

lateralibus c. 90 exorientibus. Petiolus 1-1.2 cm. logus, c. 2 mm.

diam., brevis, sicco aureo-brunneo-tomentosus. Lamina delapsa

subtus aureo-brunneo-tomentosa, supra pallide rubro-brunnea.

Racemi -14 cm. longi, semel ramosi, stipitis basi in fructu -2 mm.

diam., terminales vel axillares, semper ex integro aureo-tomentosi,

teretes; bracteis, bracteolis, floribusque ignotis. Pedicellus in fructu

-3 mm. longus diametroque, crassus. Calyx glabrescens, lobis

longioribus 3, -17 X 2.5 cm., subaequalibus, oblongo-spatulatis,

obtusis, basim versus -1.2 cm. latis attenuatis; partibus basalibus

c. 1.2 X 15 cm., latis, incrassatis, multum saccatis, ad nucem

obtectum adpressis; lobis brevioribus 2, -12 X 1 cm., aliter ut in

lobis longioribus. Nux c. 2 X 1.7 cm., ovoidea, ad apicem anguste

conicum abrupte attenuata, conferta breviter pallide alutaceo-

tomentosa; stylopodio brevi acuto.

Collections. North Borneo: San A 1748, Beaufort; San 16721,

Ulu Mendalong, Sipitang (Holotypus in Herb. Kew). Brunei:

Brun 5091, Mile 7, Kuala Abang Rd., Brun 2636, Ulu Lumut,

Andulau F.R.; Kep 80108, Labu Estate; San 17528, S 1930,

1932, Andulau F.R.; S 5777, Bt. Biang.

SHOREA REVOLUTA Ashton s.n.

5. parvifoliae Dyer affinis, sed racemis longioribus rigidioribus

rectioribus, lamina maiore coriaceiore subtus nitenti, costis laterali¬

bus prominentioribus laxioribus, margine prominenti revoluto

differt.

304

Vol XIX. (1962)

5 Mm

^ a te 25. S. pilosa.

a, Leafy branch, b, Fruit, c, Nut.

(a from Brun 5091; b and c from San 16721)

305

Gardens Bulletin, S.

Plate 26. S. revoluta.

a, Twig with young fruit, b, Fruit.

(From S 5914).

306

Vol. XIX . (1962)

Ramuli, perulae, stipulae extemae, petiolique semper sparsim

pallide brunneo- vel fulvo-scabrido-cristato-tomentosi, stipulae

intus plane pubescentes, costae subtus, costa media supra brevius

tomcntosi vel glabrescentes. Ramuli apicem versus 2-3 mm. diam.,

primo teretes vel paullum striati compressi, dein teretes, crassi, glab¬

rescentes, fusco-brunnei, lenticellis minutis orbicularibus pallidis

punctati; intemodis 1.5-4 cm. longis; cicatricibus stipularum

primo -2 mm. longis, horizontalibus, angustis, inconspicuis.

Gemma 4—7 X 3-5 mm., ovoidea, compressa, obtusa vel sub-

acuta. Stipulae -15 X 7 mm., ellipticae vel oblongae, obtusae.

Lamina 10-15 X 5.5-10 cm., ovata, coriacea; basi obtusa vel

subcordata, apice in acumen -1.5 cm. longum angustum attenuato,

margine rcvoluto; costis lateralibus utrinsecus 9-12, curvatis, sub¬

tus prominentibus, remotis, basim versus angulo c. 90° exorienti-

bus, apicem versus angulo c. 45° exorientibus; domatiis axillaribus

parvis glabris; intercostis angustis, scalariformibus, a costis laterali¬

bus diagonale exorientibus; costa media angusta, supra paullum

depressa. Petiolus 1.2-1.5 cm. longus, c. 2 mm. diam., crassus,

sicco brunneo-tomentosus vel nigrescens. Lamina delapsa supra

flavido-brunnea, subtus fusco-ferruginea. Racemi -22 cm. longi,

semel vel bis ramosi, stipitis basi in fructu 2-3 mm. diam., termi-

nales vel axillares, rigidi, conferte pallide brunneo-vel fulvo-sca-

brido-cristato-tomentosi; bracteis, bracteolis, floribusque ignotis.

Pedicellus in fructu —1.5 mm. longus, gracilis. Calyx in fructu

puberulens; lobis longioribus 3, 7.5 X 1.3 cm., tenuibus, spatu-

latis, obtusis, basim versus -3 mm. latis attenuatis, partibus

basalibus c. 6 X 5 mm., ovatis, incrassatis, saccatis; lobis breviori-

bus 2, -35 X 2 mm., subaequalibus, linearibus, acutis basibus ut

in lobis longioribus. Nux —15X8 mm., ovoidea, pallide alutaceo-

pubescens; stylopodio c. 1.5 mm. longo, brevi, attenuato.

Collections. North Borneo: San 15135, Mengalong F.R.;

Sarawak: s.n., Meragang F.R., Lawas; S 5618, Bt. Bubong Rumah,

Lawas; S 5914 (Holotypus in Herb. Kew), Bt. Sagan, Limbang.

Brunei: S 5814, Brun 5646, Bt. Patoi; S 5782, Biang.

SHOREA RUBELLA Ashton s.n.

Antherae ovariumque Sect. Anthoshoreae Heim similes, sed

arista staminis brevissima, filamentis loratisque differt; in illis

notationibus sine propinquatibus.

Perulae stipulaeque externe semper plane minute pallide roseo-

alutaceo-pubescentes, stipulae intus puberulentes; ramuli, petioli,

laminaeque primo similiter tomentosi, dein glabrescentes. Ramuli

apicem versus 2-3 mm. diam., apicem versus sicco striati

compressi, dein leves, teretes, crassi, pallide brunnei; internodis

2-4 cm. longis, multum ramosis; cicatricibus stipularum primo

c. 1.5 mm. longis, plus minus transversis, anguste cuneatis. Gemma

6-9 X 2-3 mm., anguste ovoidea vel paullum falcata. Stipulae

307

Gardens Bulletin , S.

O 0.5Mm

Plate 27. S. rubella.

a, Flowering branch, b, Sepals (outer, below; inner, above), c, Petal, d, Ovary, e, Sta¬

men (frontal view), f, Stamen (lateral view).

(a-f from Brun 3078).

308

Vol. XIX. (1962)

-18 X 6 mm., hastatae vel falcatae, acutae. Lamina 9.5-14 X

6-8 cm., late ovata, crasse coriacea, subtus cremeo-glauca, basi

obtusa, apice in acumen c. 1 cm. longum plus minus abrupte

attenuato; margine saepe paullum revoluto; costis lateralibus

utrinsecus 5-7, prominentibus, curvatis, angulo c. 40°-50° ex-

orientibus, saepe domatiis axillaribus minutis; intercostis a costis

lateralibus c. 90° exorientibus, conferte scalariformibus; costa me¬

dia supra applanata vel paullum depressa. Petiolus 2.2-3.5 cm.

longus, c. 2.5 mm. latus, sicco glaucus vel nigrescens. Lamina de-

lapsa supra roseo-brunnea, nitens, subtus cremeo-glauca (in

arboribus novellis rubro-brunneus). Racemi -15 cm. longi, semel

vel bis ramosi, stipitis basi in fructu -2 mm. diam., terminales vel

axillares, sicco striati, semper ex integro breviter conferte roseo-

brunnescente-pubescentes; ramulis -5 cm. longis, -5 flores dis-

tichos gerentibus; bracteis bracteolisque ignotis. Alabastrum -7

X 3 mm., anguste ovoideum, acutum. Calyx externe breviter con¬

ferte pallide roseo-griseo-pubescens, intus glaber; lobis ovatis

subaequalibus, internis 2 quam externis 3 paullum latioribus,

paullum brevioribus. Corolla in vivo pallide rosea, basim versus

fuscior; petalis oblongis, obtusis, vix contortis, delapsis vix con-

natis, partibus in alabastro expositis breviter tomentosis. Stamina

15, subaequalia, verticillis 3, bina cum solis alternata, circum

ovarium conferta, filamentis loratis, sub antheris abrupte attenua¬

te, apicem ovarii attingentibus; antheris filamentos aequalibus,

lineare-oblongis, angustis; aristis brevibus crassis, paullum reflexis.

Ovarium parvum, globosum, glabrum; stylopodio indistincto,

stylo quam ovarium 3-plo longiore, glabro, filiformi. Fructus

maturus ignotus. Calyx in fructu novello pubescenti; lobis maiori-

bus 3 quam minoribus 2 multum longioribus. Nux ovoidea, con¬

ferte pubescens.

Collections. North Borneo: San A 1750, Beaufort. Sarawak:

S 9474, S. Iran, Pelagus. Brunei: S 1909, Belait; S 1907, Arbo¬

retum, Kampong Gana; S 1925, Kep 36027, 80084, 81925,

48487, 48460, San 17564, Brun 3078 (Holotypus in Herb. Kew),

616, Andulau F.R.; Kep 30574, S. Tutong; Kep 30585, Tutong;

Kep 48162, S. Lumut; Kep 34529, S. Tepiku; Kep 34578, Pak

Gabal; Kep 35569, Pengkalan Ran; Kep 35667, Labi Hills F.R.

SHOREA RUBRA Ashton s.n

5. parvifoliae Dyer, S. dasyphyllae Foxw. affinis, sed lamina,

coriaceore, tomento conferte fusco-ferrugineo differt.

Ramuli, perulae, stipulae, petioli, costa media supra, lamina

subtusque semper conferte fusco-rufo-brunneo-tomentosi, stipulis

intus quam externe brevius sparsius tomentosis. Ramuli apicem

versus -1.5 mm. diam., teretes, multum ramosi, dein rufo-brun-

nescentes, leves, lenticellis parvis pallidis orbicularibus punctati;

internodis 1-1.5 cm. longis; cicatricibus stipularum primo c. 1 mm.

longis, brevibus, tenuibus, deorsum curvatis. Gemma 4-7 X 3-4.5

309

Gardens Bulletin, S.

Plate 28. S. rubra.

a, Flowering twig, b, Part of lamina (undersurface), c, Sepals (outer, above; inner,

below), d, Ovary and stamens, e, Ovary, f, Fruit.

(a-e from S 9475; f from S 2112).

310

Vol. XIX. (1962)

mm., compressa, ovoidea, late acuta. Stipulae -16 X 5 mm.,

hastatae, basi attenuata, subacutae. Lamina 8-12 X 4-7.5 cm.,

late ovata; basi obtusa, apice in acumen -1.3 cm. longum (saepe

brevem) abrupte attenuato, margine paullum revoluto; costis

lateralibus utrinsecus 11-13, subtus prominentibus, ad apices cur-

vatis, angulo 40°-50° exorientibus; intercostis angustis, conferte

scalariformibus, a costis lateralibus diagonale exorientibus; costa

media subtus prominenti, tereti, supra angusta paullum depressa.

Petiolus 1-1.4 cm. longus, c. 1.5 mm. diam., sicco rufo vel pur-

pureo-griseo-tomentosus. Lamina delapsa supra purpurcscens,

subtus rufescens, dein indistincte purpurescens vel grisea. Racemi

-14 cm. longi, semel vel bis ordine ramosi, stipitis basi in fructu

-2 mm. diam., terminales vel axillares, ex integro teretes, recti,

striati, conferte fusco-rufo-brunneo-tomentosi; ramulis brevibus,

confertis, -5 flores distichos gerentibus; bracteolis -4 X 3.5 mm.,

late ovatis, subacutis, circum alabastro imbricatis, externe breviter

pallide alutaceo-pubescentibus, intus sparsium pubescentibus.

Alabastrum -7X4 mm., ellipsoideum vel subglobosum, obtusum.

Calyx externe conferte pallide griseo-tomentosus, intus glaber;

lobis externis 3 deltoideo-ovatis obtusis, lobis internis 2 late ovatis

acutis quam illis minoribus. Corolla in vivo pallide flavida, parti-

bus in alabastro expositis breviter tomentosis; petalis lineari-

oblongis, subacutis. Stamina 15, verticillis 3 inaequalibus, interna

5 quam alia prope 2-plo longiora; filamentis externis 10 ad an-

theras gradatim attenuatis, internis 5 late loratis, sub antheris

abrupte attenuatis; antheris subglobosis; aristis quam antheras bre-

vioribus, dein paullum reflexis. Ovarium stylopodiumque ovoideo-

conicum, conferte tomentosum basi excepto; stylo quam ovarium

breviore glabro. Pedicellus in fructu c. 1 mm. longus diametroque,

brevis. Calyx in fructu breviter tenue puberulus vel glabrescens;

lobis longioribus 3, -11 X 1.6 cm., anguste spatulatis, anguste

obtusis, basim versus -4 mm. attenuatis; partibus basalibus -8 X

6 mm, multum saccatis, incrassatis; lobis brevioribus 2, -4.5 X

0.4 cm., subaequalibus, linearibus, acutis, partibus basalibus ut in

lobis longioribus. Nux -10 X 9 mm., late ovoidea, alutaceo-

velutinata, in stylopodium -2 mm. longum acutum abrupte at¬

tenuata.

Collections. North Borneo: San 15124, 15085, Beaufort; San

17008, 17013, 17010, 17023, Tambunan; San 16638, 16556,

15187, 15190, 15193, Sipitang; San 15124, Pangi; San 16355,

Ranau; San 16577, Ulu Moyah. Brunei: S 1677, Kep 48492,

Andulau F.R.; S 1691, S. Kiaput, Tutong; S 2112, Bt. Teraja;

Brun 364, Ulu Katop-Ulu Senuko watershed, Ulu Batu Apoi; S

5754, Kuala Belalong; Brun 2615, Kuala Temburong Machang;

Brun 784, Kuala Sekurop. Sarawak: S 9475, S. Iran, Pelagus

(Holotypus in Herb. Kew); Jacobs 5464, Belaga; S 9413, Kuching;

S 117, S. Miah, Rejang; S 166, S. Merirai, Ulu Balleh; S 6525, G.

Menuku.

311

Gardens Bulletin, S.

SHOREA SLOOTENI Ashton:— S. vanslooteni Wood msc.

S. macropterae Dyer affinis, sed lamina maiori, costis lateralibus

multis supra depressis, ramulis crassis compressis, staminum aristis

externis 10 vix reflexis, ovario angusto, stylo longiore differt.

Ramuli, perulae, stipulae externe, petiolo laminaeque subtus

semper conferte pallide fulvo-scabrido-cristato-tomentosi, cristis

brevibus, duris, verrucosis; stipulae intus pubescentes, costa media

supra breviter pubescens. Ramuli apicem versus -5X3 mm.,

primo compressi, striati, dein teretes, crassi, basibus cristorum

persistentibus verrucosis; internodis 1-3 cm. longis; cicatricibus

stipularum primo - 3 mm. longis, basi -0.5 mm. lata, cuneatis,

paullum sursum directis. Gemma 6-10 X 5-8 mm., late ovoidea,

paullum compressa, obtusa. Stipulae -25 X 8 mm., oblongae,

obtusae. Lamina 11-22 X 4-7 cm., late oblonga, coriacea, basi

obtusa, apice in acumen 1.5 cm. longum abrupte attenuato;

costis lateralibus utrinsecus (25) 27 -34, confertis, subtus pro-

minentibus, supra depressis, ad marginem curvatis, angulo 50°-

60° exorientibus; intercostis conferte scalariformibus, prominen-

tibus, a costis lateralibus paullum diagonale vel ad 90° exorienti-

mus; costa media subtus prominenti, tereti, supra multum de-

pressa. Petiolus 1.7-2.5 cm. longus, c. 3 mm. diam., crassus,

sicco rufo-brunneo-tomentosus. Lamina delapsa supra pallide

ferruginea, subtus fusco-ferruginea, utrinsecus inflexa, scaphiformis,

Racemi -18 cm. longi, semel vel bis ordine ramosi, stipitis basi

in fructu -4 mm. diam., terminales vel axillares, ex integro crassi,

fragiles, plus minus compressi, conferte pallide fulvo-scabrido-

tomentosi, ramulis brevibus, -4 flores gerentibus; bracteolis -3 X

2 mm., ovatis vel deltoideis, obtusis, externe conferte flavido-

brunneo-pubescentibus, intus glabris, caducis. Alabastrum -12 X

7 mm., ovoideum vel ellipsoideum, obtusum. Calyx externe bre¬

viter conferte virido-brunneo-tomcntosus, intus glaber; lobis ex¬

ternis 3 ovatis, acuminatis, quam lobis internis 2 acutis longiori-

bus. Corolla in vivo pallide flava, petalis lanceolatis, subacutis,

partibus in alabastro expositis conferte breviter tomentosis. Sta¬

mina 15, verticillis 3, bina cum solis alternata; interna 5 basim

styli attingentia, quam aliis 2-plo longioribus, his inaequalibus

filamentis ad antheras gradatim attenuatis, in alabastrum paullum

nectis; antheris globosis; aristis internis 5 quam antheras breviori-

bus dein reflexis, externis 10 quam antheras 2-plo longioribus, vix

reflexis. Ovarium stylopodiumque anguste ovoideum vel lanceola-

tum, dimidio distali sparsim setoso, stylo ovarium prope aequali,

late filiformi, glabro. Pedicellus in fructu -3 mm. longus diame-

troque, brevis, crassus. Calyx in fructu nitens, breviter puberulens,

apicem versus sparsus; lobis longioribus 3, -17 X c. L75 (-4) cm.,

inaequalibus, lineare-oblongis, obtusis, basim versus vix attenuatis,

partibus basalibus ad pedicellum abrupte attenuatis, discis centris

-12 X 12 mm. saccatis incrassatis, nuce adpressis, auriculis

lateralibus 2, -1 cm. latis; lobis brevioribus 2, 5.5-12.5 X 0-7 cm.,

312

O kj

Vol. XIX. (1962)

^ ate 29. S. slootenii.

a, Twig with young fruit, b, Sepals (outer, left; inner, right), c, Stamens and Ovary,

d, Ovary, e, Fruit.

(a from S 11053; b-d from Brun 3070; e from S 1516).

313

Gardens Bulletin, S.

inaequalibus, linearibus, basibus ut in lobis longioribus. Nux -3.5

X 1.5 cm., ovoideum, stylopodio -4 mm. longo acuto gradatim

attenuato; conferte breviter pallide brunneo- vel autoceo-pubescens.

Collections. Sarawak: S 11053, 1616, 157, Semengoh F.R.; S

1516, Lawas. North Borneo: San 16922, Bt. Sipitang; San 15160,

Mengalong F.R. (Holotypus in Herb. Kew). Brunei: Kep 35560,

Tutong R.; Kep 35561, Pengkalan Ran; Kep 48470, S 5586, Brun

5436, 3270, 3070, Andulau F.R.; S 5807, Brun 3175, Bangar.

Indonesian Borneo: bb 7097, Sambas, W. Borneo; bb 14614,

14509, E. Kutei.

VATICA CORIACEA Ashton s.n.

Vatica teysmannianae Burck affinis, sed lamina coriaceore, apice

obtusa vel retusa, petiolo breviore crassiore, calyce ad basim dis-

secto, differt.

Novellae primo conferte breviter ochraceo-tomentosae, dein

glabrae; gemmae semper similiter tomentosae. Ramuli apicem ver¬

sus -5 mm. diam., crassi rugosi, dein fusci vel nigri glabri leves.

Gemma -7x5 mm., ovoidea subacuta. Stipulae -13X5 mm.

hastatae subacutae. Lamina 6.5-15 X 2.2-6 cm. coriacissima

obovata, basi anguste obtusa, apice obtuso vel retuso, margine

plus minus revoluto; costis lateralibus utrinsecus 10-11, supra

subtusque paullum elevatis indistinctis curvatis angulo 50°-60°

exorientibus; intercostis angustis; costa media crassa, subtus pro-

minenti, supra paullum elevata. Petiolus 1-1.5 cm. longus, -3 mm.

diam., sicco niger. Lamina delapsa ochraceo- vel fusco-brunnea, ex

integro leves. Racemi -20 cm. longi, semel ramosi, stipitis basi

in fructu -3 mm. diam., terminales vel axillares, primo pallide

ochraceo-tomentosi, dein glabrescentes, bracteis stipitatis saepe

foliatis, subpersistentibus, aliter fugacis. Alabastrum -1.8 mm.

longum; calyce conferte breviter pallide griseo-brunneo-pubescenti.

Flores aliter ut in Vaticis aliis. Pedicellus in fructu -7 mm. longus,

1 mm. diam., gracilis. Calyx externe breviter pubescens, intus glab-

rescens, ad basim dissectus; lobis longioribus 2, -7 X 2.3 cm.,

oblongis obtusis, apicem versus —3.5 mm. latis subabrupte atte-

nuatis; lobis brevioribus 3, -20 X -6 mm., hastatis, acutis. Nux

-8X8 mm., globosus, breviter puberulus vel glabrescens, rugu-

losus, saepe stylo —2 mm. longo lineari terminatus.

Collections. Sarawak: S 362, 372 (Holotypus in Herb. Kew), Bt.

Sebandar, Lundu; S 10296, 10296A, Bako N.P.; S 10390, 10391,

Bt. Serepah, Kuching; S 7295, Selang F.R.; S 6344, G. Pueh F.R.;

18/58, Bt. Kayangeran, Merapok, Lawas. Brunei: S 4134, Brun

5155, Berakas F.R.; Brun 5652, Bt. Patoi.

Vol. XIX. (1962)

/Mm

Plate 30. Vatica coriacea.

a, Fruiting twig, b. Leaf, c, Stamen, d, Ovary.

(a and b from S 10296a; c and d from S 10296).

315

Gardens Bulletin, S.

VATICA PARVIFOLIA Ashton s.n.

V. micranthae V.S1. affinis, sed lamina minore, costis lateralibus

vix elevatis, angustis, tomento longiore ferrugineo differt.

Ramuli, novellae, perulae, petiolique conferte ferrugineo-pulve-

rulento-tomentosi, stipulae breviter iidem. Ramuli apicem versus

-1 mm. diam., plurimum ramosi, dein glabrescentes, leves, fusco-

brunnescentes vel nigrescentes; internodis c. 1 cm. longis. Gemma

-3 X 1*5 mm., ovoidea, subacuta. Stipulae -5.5 X 1 mm.,

lineares, caducae. Lamina 2.8-6 X 1-2.3 cm., coriacea, anguste

ovata vel lanceolata, basi obtusa, apice in acumen caudatum -1

cm. longum attenuate; costis lateralibus utrinsecus c. 8, indistinctis,

applanatis, multum curvatis, angulo 60°-75° exorientibus; costa

media angusta, subtus prominenti, supra applanata. Petiolus 6-9

mm. longus, -1 mm. diam., sicco ferrugineo-tomentosus. Lamina

delapsa ex integro pallide flavido-brunnescens. Racemi -2 cm.

longi, semel ramosi, stipitis basi in fructu -1 mm. diam., terminales

vel axillares, ex integro teretes, conferte ferrugineo-pulverulento-

tomentosi. Alabastrum -6 mm. longum, parvum; calyce griseo-

brunneo-tomentoso; corolla in vivo alba; aristis brevibus, obtusis,

floribus aliter typicis. Pedicellus in fructu -3 mm. longus, -0.7

mm. diam., gracilis. Calyx in fructu glabrescens vel ad basim

puberulus; lobis ad pedicellum liberis; lobis longioribus 2, -6 X

1.7 cm., oblongis, anguste obtusis, supra basi revolutis, ad basim

abrupte constrictis, subaequalibus; lobis brevioribus 3, -10 X

7 mm., late ovalis, acutis, basi cordatis, multum revolutis. Nux

-5 X 3.5 mm., late ovoidea, obtusa, obtusa, breviter fulvo

pubescens.

Collections. Sarawak: Jacobs 5518, S 5127, Bako N.P.; S 371,

Bt. Sebandar, Lundu (Holotypus in Herb. Kew). Brunei: Brun

2007, Bl. Puan; Kep 34475, Badas.

VATICA VINOSA Ashton s.n.

V. dulitensis Sym. affinis, sed costis lateralibus pluribus promi-

nentibus, lobis calycis minoribus, incrassatioribus, revolutis, persis-

tente vinoso-tomentosis differt.

Ramuli, perulae, stipulae, petioli, costae subtusque plus minus

semper breviter plane fusco-purpureo-tomentosi, sub laminis

quadam tenus atterrentes. Ramuli apicem versus -1.5 mm. diam.,

graciles, plurimum ramosi, dein glabrescentes, pallide grisentes vel

brunnescentesque maculosi, saepe fissulis lamellulisque superfixi;

internodis 8-15 mm. longis. Gemma -1.5 mm. longa lataque,

ovoidea, obtusa. Lamina 6-15 X 1.5-4.5 cm., spatii varii, ellip-

tica vel lanceolata, basi anguste obtusa vel cuneata, apice in acu¬

men angustum -1.2 cm. longum attenuato; costis lateralibus pri-

mariis utrinsecus 12-20, angustis, subtus plus minus prominente

elevatis, curvatis, confertis angulo 50°-70° exorientibus, secun-

dariis brevibus angustis; costa media angusta, subtus prominenti.

316

Vol XIX. (1962)

Plate 31. Vatica parvifolia.

a. Fruiting twig, b, Leaf, c. Stamen (frontal view), d, Ovary, e, Fruit,

(a and e from S 371; b-d from Jacobs 5518).

317

Gardens Bulletin, S.

Plate 32. Vatica vinosa.

a, Fruiting twig, b, Part of lamina undersurface, c, Ovary, d, Stamen,

(a and b from Brun 764; c and d from Brun 3383).

318

Vol. XIX. (1962)

tereti, supra applanata. Petiolus 6-13 mm. longus, -1 mm. diam.,.

gracilis, sicco purpureo-tomentosus vel nigrescens. Lamina delapsa

ut integro pallide griseo-brunnescens. Racemi -7 cm. longi, semel

vel bis ramosi, stipitis basi in fructu -1.5 mm. diam., terminales vel

axillares, semel ramosi, conferte breviter plane purpureo-tomentosi.

Alabastrum -6 mm. longum, parvum. Calyce breviter cinereo-

purpurescens; corolla in vivo cremia, aristis brevibus obtusis, fiori-

bus aliter typicis. Calyx in fructu purpureo-tomentosus, lobis -9 X

8 mm., obovatis, plus minus revolutis, incrassatis, ad basim re-

flexis, apicem versus gradatim recurvis, rotatis. Nux -8 mm. longa

lataque, globosa, saepe stylopodio brevi apicali, conferte purpureo-

tomentosa.

Collections. Brunei: S 5753, Brun 3383, Kuala Belalong; Brim

357, Ulu Senuko-Ulu Katop watershed, Batu Apoi; Brun 764,

Kuala Sekurop, Temburong (Holotypus in Herb. Kew); Kep 48465,.

Andulau F.R.

319

Studies in Systematic Bark Morphology

III.* Bark Taxonomy in Dipterocarpaceae

By T. C. Whitmore

St. John’s College, Cambridge, England t

SUMMARY

Bark surface pattern and slash in Dipterocarpaceae were interpreted in

terms of structure and seven main Bark Types described (Whitmore 1962a,

b) which are general categories consisting of a number of Bark Manifesta¬

tions that differ slightly in structure, hence in surface pattern and slash,

from the general Type. The Bark Manifestations are practical categories

distinct in the forest and are described and keyed here. A key is given to the

Bark Types.

The kinds of variation in bark are described and their relative import¬

ance ascertained from detailed study of a few species. The main survey is

based on 103 species in 7 genera (appendix 1). Their bark is described

genus by genus, and the bark present throughout life shown diagrammetic-

ally (appendix 2 justifies the inductions on which the diagrams are based).

The use of bark for formal taxonomy and forest recognition is reviewed.

Bark provides valuable taxonomic information: Symington’s taxonomic

groups are in main confirmed, and in some cases extended, particularly to

include Bornean species. Comment is made on a number of outstanding

taxonomic problems of individual species and a few new problems indicated.

There is no evidence of adaptive bark differences between species of Rain

forest proper and drier forests.

Twenty-one species and many species groups can be recognised in the

forest; some previously confused Scaly barks are distinguished; little is

added to previous distinctions within Dipterocarpaceae. It is suggested that

the use of bark for forest recognition could be very considerably extended

in families less well known to foresters.

* For parts I and II see The New Phytologist, 1962.

t Present address: Department of Botany, The University, Southampton,

England.

321

Gardens Bulletin, S.

THE BARK MANIFESTATIONS

General

I have described (Whitmore 1962a) how the surface pattern and

slash appearance (oblique tangential section) of tree bark may be

interpreted in terms of internal structure divided for convenience

into four interacting bark components. 103 species in seven genera

of Malayan and Bornean Dipterocarpaceae have been studied

(listed in appendix 1); from these seven main Bark Types have

been described (Whitmore 1962a) and interpreted (Whitmore

1962b) in terms of the relative rates of formation of xylem (wood)

and phloem (bark) at the vascular cambium. These Bark Types

are general categories and consist of a number of Bark Manifesta¬

tions which have minor differences in the components from the

general Type and hence differ slightly in surface pattern and slash.

The Types are named after surface appearance {Smooth, Fig. 9a,

Dippled, Fig. 9c, d, Shallow Fissured, Fig. 10a, Deep Fissured,

Fig. I Ob, and Scaly, Fig. 9b) or slash ( Surface Rotten, Fig. 10c

and Laminate, Fig. lOd). In this paper the Bark Manifestations

and their taxonomic occurrence will be described.

Non floral characters are of established importance in the forest

identification of trees in the tropics and bark surface and slash

characters are important amongst others. Indeed in a family like

Dipterocarpaceae which seldom flowers floral characters are only

of secondary importance. Bark characters are at present used

empirically. It is possible that their use for forest recognition can

be extended now that the description of bark is rationalised and

that bark can be used in studying the taxonomy of the family

which is at present imperfectly understood.

The Bark Manifestations are practical categories. Deciding whe¬

ther one bark is sufficiently different from another to be described

as a separate Manifestation is partly subjective and the criterion

adopted is that the bark must be distinct in the forest from surface

or slash characters (a hand lens may be needed).

The Manifestations just like the Bark Types can be defined by

describing the bark components and the results of their inter¬

actions. Manifestations differ from each other in one or more of

the three grossly varying components, periderms, expansion tissue

and phloem proliferation tissue. Periderms play the most im¬

portant part in determining the surface appearance (Fig. 1 in

Whitmore 1962a) and Manifestations usually differ in this com¬

ponent, for instance Manifestations of Shallow Fissured bark. In

some barks however, for instance Manifestations of Surface Rotten

and Laminate barks, the nature and amount of phloem prolifera¬

tion tissue also exercises a very important effect on surface appear¬

ance. The fourth bark component, the secondary phloem, does

not vary.

322

Vol XIX. (1962)

Variation is continuous and within some Manifestations there

are bole barks, observed on a single species or a few trees, which

differ in a few features. These I have described as Variants of

the Manifestations, they are not always easily recognisable in the

forest. Some forest-distinctive barks have also been described as

Variants not Manifestations because they were suspected to be

atypical or because the material examined was poor.

A few Manifestations (Smooth—e, Smooth—f and Shallow Fis¬

sured—c, q.v.) have characters intermediate between two Bark

Types. I have placed these with the Bark Type they most nearly

resemble in bole appearance from a distance.

Some barks do not fit the main Bark Types but have bark com¬

ponents of unusual structure, hence they differ in surface pattern

and slash. These may be abnormal or may be additional Bark

Types not well represented in this incomplete survey of the Dip-

terocarpaceae. They are mentioned with the notes on the species

bearing them as they are of potential importance in understanding

the bark of the whole family, and reference is made to the full

diagnosis and discussion of status given in Whitmore (1960).

Names of Manifestations

The Manifestations are distinguished by a letter suffixed to the

Bark Type, thus Smooth—a, Scaly—g.

Scheme for description of Manifestations

The visual impression of the bark from a distance is similar for

all trees of a Manifestation but there are differences in some of

the details of surface and slash. The diagnoses reflect this vari¬

ability, some features are very precisely defined because they are

invariable, others more loosely.

The Bark Type descriptions (Table 1, Whitmore 1962a) are

indefinite because they cover the range of the Manifestations. In

the Manifestation diagnoses the features of the Type are not re¬

peated. The two descriptions together give a complete account of

any bark. In this paper the Key to the Bark Types gives the main

diagnostic features of each.

The description of each Manifestation follows the plan:

Diagnosis, with the most important features for identification

in italics, under the subheadings:

Periderm(s), Inner Bark Structure (tangential expan¬

sion and phloem proliferation tissues), Outer Bark Struc¬

ture (note that the rhytidome layers are usually developing

and sloughing all the time so the number present varies),

Sloughing and Surface (configurations, texture and

colour),

Variants, species by species.

323

Gardens Bulletin, S.

Occurrence, bole or crown and species group.

Photographs of the bole from a distance and close to and photo¬

graphs or diagrams of a transverse surface are given to the Mani¬

festations and Variants in Whitmore (I960), here Figs. 9 and 10

a selection of Manifestations of every Type.

Keys

There is a key to the Manifestations of every Bark Type making

use of the salient features, a guide to the diagnoses but not sup¬

planting them. The keys assist comprehension of the differences

between the Manifestations and make possible the identification of

the bark of any of the species examined or any others with one of

these Manifestations.

The terms used in the diagnoses of the Manifestations and in

the keys to the Bark Types and Manifestations are defined in

Whitmore (1960 and 1962a).

KINDS OF VARIATION IN BARK

The basic structures and processes, the same in all trees of a

species with bark at the same stage of development, must be dis¬

tinguished from variable features and there may be developmental

changes. Different features of the bark may vary at different taxo¬

nomic levels. The relative importance of these kinds of variation

was ascertained early in this survey from frequently collected

Shorea, Red Meranti group species, and from Shorea albida

(Meranti Pa’ang), Dipterocarpus costulatus, D. crinitus and D.

verrucosus. It is to some extent possible to distinguish inherited

and environmental variations, the former are of potential use for

forest recognition and taxonomy.

The variation expounded

1. The morphogenesis of bark throughout life was compared

on the stems of seedlings, saplings, poles and trees and crown

members of increasing girth. The Bark Type sequence is the same

but in the crown the changes take place at smaller girths. The

difference in phasing can be understood by postulating that the

development of a Bark Type depends on the age of a member not

its girth; poles grow at a more or less steady rate whereas there

is little increase in girth in the crown members of mature trees.

There is considerable variation between trees in the girths at

which the bark changes, this may be inherited or environmental.

Species differ in their Bark Type sequence and girths of change.

There are often considerable variations in details of the bark

between parts of an individual at the same stage, particularly be¬

tween bole and crown (see 5 below) but also between lower and

upper bole. These I ascribe to the difference in environment

between the lower levels and canopy level of rain forest.

324

Vol. XIX . (1962)

2. Inherited geographical variation. This would be expected to

be most conspicuous in polymorphic species, and individuals from

widely separated localities for instance Malaya and Borneo, per¬

haps as a sign of incipient speciation; however it proved to be un¬

common. The only species of many examined in both Malaya and

Borneo which shows it is Hopea beccariana and of Shorea lepro-

sula , S. macroptera, S. parvifolia, S. pauciflora and Dipterocarpus

costulatus examined all over Malaya only S. parvifolia shows

geographical variation. The only geographical variation due to dif¬

ferences in the environment noticed was the tendency for trees to

mature smaller and to become gnarled on ridges or in shallow or

sandy, infertile soil.

3. Variations between individuals due to the cyclical develop¬

ment and sloughing of rhytidome layers is common with Scaly

barks and may have inherited or environmental causes, in many

cases I have not examined sufficient material to say which.

4. Slight differences in environment might produce variations

in bark from tree to tree. Such variations are minimised in this

survey by using trees from within the more or less uniform environ¬

ment of Rain forest. There may be small inherited variations

which will not be distinguishable.

Within a locality in the Rain forest minor differences occur in

bark surface pattern, ridge width, fissure size, details of surface

texture and sculpturings, in scale shape and size and the number

of rhytidome layers present together. These limit the detail which

can be usefully recorded in forest descriptions. Bark structure also

shows variations, in periderm pattern and outer bark structure

(particularly the degree of rotting), perhaps dependent mainly on

micro-climate, in expansion tissue, dependent on the xylem/

phloem relative growth rate, and in phloem proliferation tissue, still

unexplained. Similar variations to these have been found between

trees from different localities.

5. Some exposed trees have been examined to discover the

potential variability of bark between open and Rain forest habi¬

tats but it would be difficult to study barks from such trees alone. 1

(i) The variations found in the open are that the outer bark

tends to be thicker with more rhytidome layers present

together. The thicker layer of dead tissues is subject to

stronger Assuring forces and desiccation is greater than

in the forest; thus barks become fissured or more

strongly fissured.

(ii) Environmental fluctuations are greater and the surface

tends to become rugose and grey in colour.

(iii) Bark Types tend to develop at smaller girths than in the

forest.

l- Hence difficult to study bark in England where most specimens of

many broad leaved species grow in hedgerows and park land.

325

Gardens Bulletin, S.

Similar differences were found between bark in the crown and

on the bole where there are additional differences due to the extra

strains set up by the bent shape and the exposure to wind of the

crown members.

THE PROCEDURE FOR EXAMINATION

The species were investigated in the laboratory in the groups

suggested by Symington (1943) which is the most natural subdivi¬

sion of the family available, and natural subgroups were investi¬

gated together.

Within each group the procedure adopted after some trial was

first to determine from the collected samples, the descriptions and

any photographs the Bark Types present on the boles of the sam¬

pled trees than to work out the Bark Type sequence with the help

of the published generalised descriptions (Foxworthy 1927, 1932,

Symington 1943, Wood 1957).

Tables showing Bark Type sequence through life, species by

species, are reproduced here, they also show at a glance the num¬

ber of individuals sampled and their girths. Details of their con¬

struction are given in appendix 2. In some species paucity of

material, particularly of small individuals, renders the Table

incomplete.

The second step in the analysis was carefully to examine the

samples of each Bark Type species by species and to record the

structure on a table by the presence or absence of separate fea¬

tures. This led to the concepts of bark structure described in

Whitmore (1962a). The diagnoses of the Manifestations and their

Variants were made from these analysis tables; several of them

are reproduced in Whitmore (1960) to show the magnitude of

variation within and between Manifestations and Bark Types.

Once the extent of variation between trees had been established

(see above) generalisations were made about the bark present at a

particular girth from the examination of only a few individuals.

The extent to which the data can be extrapolated to cover the

whole girth range is investigated in appendix 2.

For each species group a short note is given on the present

Taxonomic status, then follows the Argument with my findings

on Bark Type sequence, the Bark Manifestations and any un¬

usual features of bark structure, and finally the Conclusions which

can be drawn on the taxonomy of the group, its subdivisions and

particular critical species, and how the group can be distinguished

in the forest from others with which it has previously been

confused.

The Manifestations which are unique are noted and by referring

to the diagnoses of these Manifestations the reader can discover

the bark features whereby the trees can be recognised in the forest.

326

Vol. XIX. (1962)

KEY TO BARK TYPES

A Expansion tissue uniform, continuous at surface in wedges

and or fingers confluent externally into a pseudocortex,

B Some periderms forming scollop shaped rhytidomes

2-3 mm. thick of varying extent greater than 2-3

cm.; sloughing as scollops or large fractions thereof;

surface dippled, scroll marked

DIPPLED BARK Fig. 9c, d

(Shorea Meranti Damar Hitam, Vatica).

BB Periderms single or 2-3 very near surface and almost

touching; sloughing apparently absent; surface entire

SMOOTH BARK Fig. 9a

(young members of all groups; mature

members A nisoptera, Dipterocarpus,

Hopea Smooth and Mata Kuching,

Shorea).

AA Expansion tissue localised in wedges or in clusters of fin¬

gers, tangential extent at inner bark surface usually 30%

(rarely continuous, pseudocortex present); periderms not

parallel to cambium for more than 1-3 cm. on TS; rhy-

tidome of several thin layers completely and irregularly

superimposed; surface closely, more or less regularly, V

section fissured

SHALLOW FISSURED BARK Fig. 10a

(Shorea Red Meranti, Meranti Damar

Hitam, Meranti Pa’ang).

AAA Expansion tissue disperse, tangential extent at inner bark

surface about 10%,

C Periderms regular, always continuous, completely

separating living tissues from dead; rhytidome

layers extensive, sheet-like, outer bark clearly

demarcated,

D Periderms frequently penetrated by fibres;

2-10 rhytidome layers present together,

outer bark coherent, very dark; surface

coarsely, closely fissured; sloughing as

small, adherent, chunky scales

DEEP FISSURED BARK Fig. 1 Ob

(Hopea Fissued Barked).

DD Periderms not penetrated by fibres; 1-2 (3)

rhytidome layers present together; outer

bark otherwise; surface not coarsely fissur¬

ed, fissures scattered or close; sloughing

usually as large, loose or adherent, layer-

thick scales

327

Gardens Bulletin, S,

SCALY BARK Fig. 9b

(Balanocarpus, Dipterocarpus,

Dryobalanops, Hopea Scaly

Barked, Shorea Red Meranti,,

Balau and Meranti Damar

Hitam).

CC Periderms irregular, not always continuous or

completely separating living tissues from dead;

no regular rhytidome layers, outer bark appear¬

ance of progressive rotting from surface inwards,

E Compound laminae throughout bark;

periderms pale coloured; inner edge of

outer bark more or less parallel to

cambium

LAMINATE BARK Fig. lOd

(A nisoptera).

EE Compound laminae absent, simple lami¬

nae sometimes present; periderms dark

coloured; inner edge of outer bark fol¬

lowing surface configurations, not

parallel to cambium

SURFACE ROTTEN BARK

Fig. 10c

(»Shorea Meranti Pa’ang).

MANIFESTATIONS OF SMOOTH BARK

Key to the Manifestations of Smooth bark

A Compound laminae present in inner bark; stone cell clusters filling

pseudocortex; . g, A nisoptera.

AA Not as above,

B Expansion tissue in well defined wedges below regular superficial

fissures. 3 mm. wide X 2 mm. deep, 2 cm. apart.

C Periderm thick (2 mm.), reticulate, occupying all but a

narrow band of the pseudocortex

e, Shorea, Meranti Pa’ang.

CC Periderm not as above. f, Shorea, Meranti Pa’ang.

BB Expansion tissue in abundant fingers, surface not as above,

D Surface with finely striate, net-like pattern due to rup¬

turing of outer periderms; periderms paper thin;

h, all groups.

DD Surface not as above; periderms not paper thin,

E Surface quite smooth, without fine sculpturings,

F Restricted to twigs (and probably seedlings and

saplings too); surface purple brown; not

sloughing . a, all groups.

FF Mature boles; not purple brown; sometimes

sloughing

G Phloem proliferation tissue parenchyma

only

j, most Smooth and Mata Kuching

Hopeas.

328

Vol XIX. (1962)

GG Phloem proliferation tissue parenchyma

and conspicuous stone cells

some c, Shorea, Red Meranti.

EE Surface finely sculptured,

H Mature periderms radially striate,

hard, brittle, thick (c.0.4 mm.), can

be chipped away; surface some¬

times finely grid-cracked

d, Shorea , Meranti Damar

Hit am.

HH Mature periderms not as above, can¬

not be chipped away; surface never

grid-cracked,

I Surface rugulose or rugose due to

fine sculpturings

some c. Shorea,

Red Meranti.

II Surface fine sculpturings not as

above,

J Surface fine sculpturings in

form of fine, close, vertical

lines less than 1 mm. deep

and wide. k, Vatica.

JJ Surface fine sculpturings in

form of short, superficial

fissures to 1-2 mm. deep

and wide,

K Phloem proliferation tissue

usually of varying

amounts in different

tangential zones of fibr¬

ous part of bark, some¬

times white with silica

deposited,

h, Dipterocarpus.

KK Phloem proliferation tissue

uniform, never siliceous,

Ho pea vesquei (j).

MANIFESTATION Smooth—a

Diagnosis: Periderm: single superficial. Inner bark structure: mature bark

structure not yet developed; an earlier stage present. Sloughing: apparently

nil. Surface: quite smooth; purple brown.

Occurrence: Restricted to twigs up to 8 cm. girth, probably also on seed¬

lings and saplings though bark of these was not examined. Probably present

in all species groups; its apparent absence from some may be because no

small twigs were examined or because there had been no recent growth

flush and the twig bark had changed on ageing.

Discussion: Further examination may show that this is the bark present

before periderm has developed when the epidermis forms the surface. The

stages in the development of mature bark structure are described in Whit¬

more (1960 Chapter 8b).

MANIFESTATION Smooth—b

Diagnosis: Periderms: several, close, paper thin. Inner bark structure:

crown structure immature; bole as Type (expansion tissue of tangentially

elongate stone cells and or parenchyma; phloem proliferation tissue usually

extensive, large circular stone cells and parenchyma). Sloughing: as Type.

Surface: finely sculptured— outer periderms rupturing and forming an open

net pattern on the surface.

329

Gardens Bulletin, S.

Variants:

1. Shorea waltoni (Red Meranti): Inner bark: phloem proliferation

tissue: circular stone cells aligned in inconspicuous tangential sheets—c.f.

Smooth—c also a few conspicuous sheets of radially elongate stone cells.

2. Shorea, Meranti, Damar Hitam species: Periderm: thick on bole, to

0.3 mm.—c.f. Scaly—d.

Occurrence: Crowns and some young boles, perhaps all groups; seen in

Anisoptera; Balanocarpus; Dipterocarpus and Shorea all groups.

MANIFESTATION Smooth—c

Diagnosis: Periderms: 1-2 present together. Inner bark structure: as Type

(expansion tissue of tangentially elongate stone cells and parenchyma;

phloem proliferation tissue usually widespread of parenchyma and large

circular stone cells). Outer bark structure: thin rhytidome layers to 1-2

mm. thick. Sloughing: as Type. Surface: either finely sculptured rugose,

rugulose, or not sculptured, smooth with small lenticels.

Variant:

1. Shorea mecistopteryx (Red Meranti): Inner bark: phloem prolifera¬

tion tissue in the fibrous part as tangential sheets of radially elongate stone

cells, interrupted outwards by the broad expansion tissue fingers where these

merge with the pseudocortex, hence not forming long continuous sheets not

dominating appearance of cut transverse surface.

Occurrence: Restricted to boles; Shorea, Red Meranti.

MANIFESTATION Smooth—d

Diagnosis: Periderms: usually only one; phellem thick (to 0.4 mm.),

radially striate, soft and pale when first formed but soon becoming brittle

and dark; phelloderm absent, or thin (but occasionally to 0.2 mm.); phel¬

lem may be chipped away, sometimes to reveal the bright phelloderm—

the ‘scrape’. Inner bark structure: as Type (expansion tissue mainly of

parenchyma or small stone cells, phloem proliferation tissue absent or local¬

ised). Outer bark structure: mainly thin; periderm single, superficial; in

places, especially crown, a thin rhytidome (0.5 mm.) where parts of a

former periderm persist. Sloughing: as Type. Surface: periderm becoming

finely sculptured:—rugulose, rugose, finely grid-cracked (5 mm. X 3-4

mm.), or longitudinally striate (in crown). Lenticels along lines of peri¬

derm rupture, periderm tissue powdery here.

Occurrence: Bole and crown; Shorea, Meranti Damar Hitam.

MANIFESTATION Smooth—e

Diagnosis: Periderm: superficial, firm, dark sheet with inwards a zone,

2 mm. thick, of convoluted, anastomosing, dark, hard ‘phellem’ tissue in¬

terspersed with paler bands of ‘phelloderm,’ together making up a hard

dark reticulated layer. Inner bark structure: Pseudocortex only a few mm.

thick, the inner part of the very thick periderm probably formed from and

in pseudocortical tissue. Expansion tissue in well defined wedges, no

fingers. Phloem proliferation tissue: small, scattered, tangential laminae,

mentioned in all the published generalised descriptions but absent or only

slightly developed in the samples examined (then simple laminae, sheets of

stone cells); other phloem proliferation tissue restricted to outermost

phloem blocks. Outer bark structure: see periderm. Sloughing: apparently

absent. Surface: dark coloured, with regular, superficial fissures (3 mm.

wide X 2 mm. deep) above the wedges but not penetrating into inner bark;

ridges 2 cm. wide with small, scattered lenticels, otherwise smooth and

firm.

Occurrence: Bole, Shorea ochracea, and perhaps other species of Meranti

Pa’ang.

Discussion: The distinction of ‘phellem’ and ‘phelloderm’ is not certain as

no phellogens have been made out. This Manifestation is described from a

single tree. It is conceivable that the thick reticulated periderm, its main

distinctive feature, is a pathological condition, although the tree appeared

quite healthy, and that further research will show Smooth—e and —f to

be identical.

330

Vol. XIX. (1962)

MANIFESTATION Smooth— f

Diagnosis: Periderms: 1-2 present together. Inner bark structure: same

as Smooth—e with wider pseudocortex. Outer bark structure: thin rhyti-

dome layers, to 1-2 mm. Sloughing: apparently absent. Surface same as

Smooth — e.

Occurrence: Bole and crown; Shorea bracteolata, S. symingtoniana and

perhaps other species of Meranti Pa’ang.

Discussion: This Manifestation differs from Smooth—e, the other smooth

bark of the Merantis Pa’ang’s in the periderms and the outer bark which

resemble Smooth— c.

MANIFESTATION Smooth—g

Diagnosis: Periderm: single, superficial, 0.20 mm. thick, pale in colour.

Inner bark structure: tangential laminae of compound structure , throughout,

varying in stage of development from one phloem block thick and tangen¬

tially short to thicker and more or less continuous; laminae interrupted by

fingers and by pseudocortex where this penetrates inwards; pseudocortex

more or less filled with stone cell clusters , not laminate; expansion tissue

in broad fingers developing from pairs of rays. Sloughing: apparently

absent. Surface: superficial fissures 4-10 mm. wide X 2 mm. deep, wholly

periderm bounded, not in any way disrupting the overall smooth bole

appearance; pale, at periderm.

Occurrence: Crowns and boles; Anisoptera.

Discussion: Periderm same as Smooth—c except for the pale colour.

Laminae not developed in inner bark at 1 ft. girth on bole (A. scaphula),

developing at 3 ft. (A. laevis).

MANIFESTATION Smooth—h (Fig. 9a)

Diagnosis: Periderms: usually 2, sometimes 1 only; strongly small-undu¬

late on TS and LS. Phellem hard, sclerified, pale coloured; phelloderm

thick (to 0.5 mm.). Inner bark structure: as Type (expansion tissue:

parenchyma; phloem proliferation tissue: extensive, small stone cells not

in clusters greater than one matrix block, usually of variable amount in

different tangential zones of the bark, increasing outwards); silica some¬

times deposited in matrix. Sloughing: as Type. Surface: finely sculptured:

rough with short (to 5 cm.) close (few mm.) superficial fissures (1-2 mm.

wide and deep) developing in the thickness of the outer bark.

Occurrence: Boles of immature individuals and probably mature tree

crowns of Dipterocarpus.

Discussion: Inner bark: expansion tissue and phloem proliferation tissue

are the same as Scaly—e.

MANIFESTATION Smooth—j

Diagnosis: Periderms: as Smooth—c. Inner bark structure: as Type (ex¬

pansion tissues as Smooth—c; phloem proliferation tissue: parenchyma

only). Outer bark structure: as Smooth—c. Sloughing: as Type. Surface:

smooth, entire, no fine sculpturings.

Variant:

1. Hopea vesquei: Surface: finely sculptured—close regular superficial

fissures.

Occurrence: Boles, Hopea . Smooth and Mata Kuching.

MANIFESTATION Smooth—k

Diagnosis: Periderms: as Smooth—c. Inner bark structure: as Type (ex¬

pansion tissue as Smooth—c; phloem proliferation tissue: parenchyma only)

silica often deposited. Outer bark structure: as Smooth—c. Sloughing: as

Type. Surface: finely sculptured: close , fine, vertical lines less than I mm.,

deep and wide.

Occurrence: Boles, Vatica.

331

Gardens Bulletin, 5.

MANIFESTATION OF DIPPLED BARK

MANIFESTATION—a (Fig. 9c; 9d shows the much more strongly dip-

pled bark of Agathis alba )

Diagnosis: Periderms: pattern: as Type; structure: 0.2 mm. thick. Inner

bark structure: as Type. Outer bark structure: thin, or with 2 mm. thick

scollops, 2-6 cm. in diameter, varying between trees. Sloughing: as Type.

Surface: as Type.

Variants:

1. Shorea: Inner bark: phloem proliferation tissue absent.

2. Vatica: Inner bark: phloem proliferation tissue widespread, of paren¬

chyma, white and silicified.

Occurrence: Boles; Vatica and Shorea Meranti Damar Hitam (5. fague-

tiana and S. multi flora) where it is probably intermediate between Smooth

and Scaly bark.

MANIFESTATIONS OF SHALLOW FISSURED BARK

Key to the Manifestations of Shallow Fissured bark

A Expansion tissue localised in well defined wedges below fissures which

penetrate into inner bark (though usually lined by periderm(s)),

B Pseudocortex present in outer inner bark or outer bark;

outer bark essentially a superficial skin of close, anastomosing

or almost touching periderms; 1-3 mm. thick; sloughing ap¬

parently absent from boles; bole appears smooth-fissured

a, Shorea Red Meranti (bole & crown) and

Meranti Damar Hitam (crown only).

BB Pseudocortex absent; outer bark not a superficial skin; 3-11 mm.

thick; sloughing always present; bole appears coarsely fissured

b, Shorea, Red Meranti

e, Shorea talura, Meranti Pa'ang.

AA Expansion tissue not as above; fissures not penetrating into inner bark,

C Surface coarsely fissured; ridges rugose; outer bark to 17 mm.

thick, of up to 12 rhytidome layers; pseudocortex absent;

expansion tissue in clustered fingers

d, Shorea gibbosa, Meranti Damar Hitam.

CC Surface shallow fissured; ridges smooth; outer bark 1-3 mm.

thick, never more than 3-4 rhytidome layers; thin pseu¬

docortex present; expansion tissue in uniformly distributed

fingers . c, Shorea macroptera, Red Meranti.

MANIFESTATION Shallow Fissured—a

Diagnosis: Periderms: multiple structure, thickness variable (0.03 mm.-

0.20 mm.), phellems black and usually leathery, phelloderm either ap¬

parently absent, or 0.25 mm. thick. Several, close (0.5-1.0 mm. apart,

rarely more); pattern as Type. Inner bark structure: pseudocortex present;

expansion at first by fingers and wedges then by wedges only; phloem pro¬

liferation tissue: as Type. Outer bark structure: essentially a superficial

skin, 3 mm. thick or less, usually dipping in at, and bounding the fissures.

Sloughing: on boles apparently absent; in crowns absent or as layer-thick,

brittle, thin, scales, adherent before sloughing. Surface: as Type; fissures

penetrating inner bark; ridges smooth, sometimes microlichen flecked;

surface firm, dark coloured, at a periderm; bole smooth-fissured, becoming

coarsely fissured on old trees.

Variants:

1. Shorea parvifolia (Red Meranti): Periderms: phelloderm absent or

inconspicuous. Inner bark: no phloem proliferation tissue in fibrous part.

There are two phases:

(a) Trees to about 6 ft. girth.

Bole appearance: smooth, shallow fissured, blotched with mic¬

rolichen flecks. Fissures wide-V section, shallow (5-7 mm. wide

X 2 mm. deep), regular; ridges: flat, firm, with slightly raised

edges.

332

Vol. XIX. (1962).

( b ) Trees above about 6 ft. girth.

Bole appearance: variably, coarsely fissured. Fissures wider

and deeper, less regular than before (to 10 mm. wide by 5 mm.

deep); the ridges lose their continuous vertical and smooth ap¬

pearance and sometimes fissure further, the surface becomes

rugose but remains quite firm.

A conspicuous feature of this species is what Symington (1943) called

‘small warty portions of the bark’. These are often present, mainly on young

trees to about 6 ft. girth, damar often drips from them. They are formed

by the bole swelling over a zone (average size 30 cm. broad X 15 cm.

high but sometimes tangentially greater) and bursting the normal outer

bark which remains as a rough fringe to the swelling which becomes

surfaced by fresh periderms.

2. Shorea leprosula (Red Meranti): Periderms: distinct red brown phel-

loderms present. Inner bark: phloem proliferation tissue in scattered tan¬

gential sheets; a few dammar channels present in phloem rays stretching

from xylem to surface. Outer bark: periderms develop beneath fissures

first and spread outwards beneath ridges, hence there are more periderms at

fissure sides than under ridge centres; periderms, ruptured by subsequent

fissure increases and only 1-2 innermost are continuous beneath fissures;

periderms furthest apart and hence rhytidome layers thickest at fissure

edges, becoming closer towards ridge centres. Subsequent enlargement of

expansion tissue wedges below fissures forces fissure edge tissue outwards

so that ridges become slightly concave in section. There are two phases:

(a) Trees to about 5 ft. girth.

Bole appearance: smooth-fissured; fissures 5 mm. wide X 3 mm.

deep, ridges 2-3 cm. wide with small lenticels and raised marks

continuous across several ridges.

(b) Trees above about 5 ft. girth.

Bole appearance: coarsely fissured, surface rough. Fissures be¬

come compound, getting progressively wider, to as much as 20

mm.; smooth ridges progressively wider separated. Tissue inter¬

polated at fissures sometimes form a mass of cork tissue which

comes to fill the fissures except for a narrow central groove and

may protrude above the general level so emphasising the con¬

cavity of the ridees.

Occurrence: Crowns, Shorea Red Meranti and Meranti Damar Hitam

(S. gibbosa ); boles, Shorea Red Meranti, S. parvifolia and S. leprosula.

Discussion: Sloughing: outer bark almost free of fibres, no strong connec¬

tions between rhytidome layers although periderms of multiple structure,

thus sloughing easy, down to a periderm and outer bark fairly thin.

MANIFESTATION Shallow Fissured—b

Diagnosis: Periderms: multiple structure, thickness variable (0.03 mm-

0.40 mm.), phellems black and usually leathery, phelloderms absent or in

conspicuous isolated lenses, 0.4 (1.0) mm. thick, and also sometimes in

strips along fissure edges. Pattern: as Type. Inner bark structure: as Type

(expansion tissue in wedges, tangential extent 30 (25-30) %). Outer bark

structure: 3 or more rhytidome layers usually present together, thickness

variable, outer bark 3 mm. or more, rarely as much as 11 mm., thick.

Sloughing: ridge-wide, chunky or layer-thick elongate scales, adherent be¬

fore sloughing. Surface: as Type, ridges concave in section when phello-

derm strips have formed; fissures penetrating inner bark; ridge surfaces

firm or not, powdery. Bole appearance coarsely fissured.

Variant: Shorea rubella (Red Meranti): Periderms: much larger scollops

of tissue enclosed, 4 cm. wide X 5-7 mm. thick; brown not black coloured.

Outer bark: very thick, 10-14 mm. Surface: fissures coarser than above.

Occurrence: Restricted to boles; Shorea, Red Meranti.

Discussion: Sloughing: the scales are chunky, and adherent before slough¬

ing because there are no definite localised planes of sloughing at or near

the multiple structure, fibre-penetrated periderms.

333

Gardens Bulletin, S.

MANIFESTATION Shallow Fissured—c (Fig. 10a)

Diagnosis: Periderms: single sheets; several, undulate, anastomosing, vari¬

ously separated. Inner bark structure: narrow band of pseudocortex; ex¬

pansion tissue in uniformly distributed fingers —uniseriate files of tan¬

gentially elongate stone cells; phloem proliferation tissue extensive, small

circular stone cells formed in most phloem blocks. Outer bark structure:

1-3 mm. thick, only partially penetrated by fissures. Sloughing: small

chunky or layer-thick, thin, scales; sloughing sometimes apparently absent.

Surface: regular, superficial fissures with slightly raised edges disrupting the

overall smooth appearance of bole; ridges smooth, surface firm, powdery

or not.

Occurrence: Boles only, Shorea macroptera (Red Meranti).

Discussion: Periderm separation variable hence outer bark from ‘surface

skin’ of —a to thicker, like —b. Surface pattern is similar to S. parvifolia,

—a.

MANIFESTATION Shallow Fissured-—d

Diagnosis: Periderms: thick (0.4 mm.), dark brown, single sheets;

anastomosing on TS; undulate but not anastomosing on LS. Inner bark

structure: expansion tissue in fingers localised in clusters, of tangentially

elongate stone cells only, tangential extent about 40%; phloem prolifera¬

tion tissue localised, of parenchyma. Outer bark structure: many (to 12)

rhytidome layers; to 17 mm. thick; formed of unevenly overlapping scollop-

section strips of thickness varying from 1.5-4.0 mm. Sloughing: elongate,

chunky scales. Surface: fissures not penetrating inner bark; ridges rugose,

with loose powdery surfaces; bole appears coarsely fissured.

Occurrence: Boles only, Shorea gibbosa (Meranti Damar Hitam).

Discussion: Outer bark coherent and long persistent, no definite sloughing

planes develop hence scales are chunky, contrast with Scaly —d (same

species group) which has sloughing planes within the phellems.

MANIFESTATION Shallow Fissured—e

Diagnosis: The only sample available was too badly damaged to ascertain

characters of the outer bark: the inner bark structure is of this Bark Type.

There is a full discussion in Whitmore (1960, pp. 304-6). Periderms: pre¬

sent, more or less continuous. Inner bark structure: as Type (expansion

tissue in wedges of 30% tangential extent; phloem proliferation tissue

absent). Outer bark structure:—. Sloughing:—. Surface: as Type; bole

coarsely boat-shape fissured.

Occurrence: Boles, Shorea talura (Meranti Pa’ang); crown not examined.

MANIFESTATION OF DEEP FISSURED BARK

MANIFESTATION—a (Fig. 10b)

Diagnosis: See Table 1 (Whitmore 1962b) and key to the Bark Types

p. 327 above.

Variant:

1. Shorea beccariana (Malayan trees 180 & 197): Periderms: closely

anastomosing on LS and TS. Outer bark structure: rhytidome layers

narrow imbricating. This is so similar to the Bornean tree seen (488) and

to the other species in all respects except the closely anastomosing peri¬

derms (the anastomoses do not coincide with the fissure edges) that, on

the evidence available, it is included in the same Manifestation; further

material is needed to decide whether the periderm difference is consistent

and if there are other differences, hence whether this really is the same

Manifestation.

Occurrence: Fissured Barked Hopeas.

334

Vol. XIX. (1962).

MANIFESTATIONS OF SCALY BARK

Key to the Manifestations of Scaly bark

A Phellem 2 zoned, inner dark blackish brown and outer biscuit coloured

layers; surface at periderm, pale when new becoming very dark by

erosion. g, Dryobalanops.

AA Phellem 1 zoned, dark or pale; surface at periderm or not,

B Expansion tissue in radially short fingers, more numerous out¬

wards,

C Phloem proliferation tissue entirely parenchymatous; radial-

oblique fingers sometimes present; phelloderm pale, cons¬

picuous and relatively thick (0.3 mm.)

c. Shorea, Balau.

CC Phloem proliferation tissue with conspicuous, pale stone

cells; radial oblique fingers absent; phelloderm inconspi¬

cuous, thin, or absent,

D Surface grid-cracked . b, Shorea, Red Meranti.

DD Surface often with more or less regularly distributed

close fissures but no cross cracks

a, Shorea, Red Meranti.

BB Expansion tissue, if present, in fingers stretching from near

cambium to surface and becoming wider outwards,

E Phloem proliferation tissue always extensive

usually of varying amounts in different tangential

zones sometimes distorting secondary phloem

structure, sometimes white, silicified; surface with

powdery bloom . e, Dipterocarpus.

EE Phloem proliferation tissue absent or extensive,

constant in amount throughout thickness of bark,

never distorting secondary phloem structure; sur¬

face shiny without powdery bloom,

F Phellem 0.4 mm. thick; 0, 1, or, rarely, 2 rhy-

tidome layers present together

d, Shorea, Meranti Damar Hitam.

FF Phellem 0.1-0.2 mm. thick; 1-2 rhytidome

layers present together, rarely 0

f, Scaly Hope a, Balanocarpus.

MANIFESTATION Scaly—a

Diagnosis: Periderms: several; on TS sometimes anastomosing about

every 5-7 cm.; on LS anastomoses greater than 15 cm. apart; usually simple

structure, rarely multiple; phelloderm absent or present, continuous and

inconspicuous or in isolated, conspicuous lenses. Inner bark structure: ex¬

pansion tissue in radially short fingers, more abundant outwards, tangential

extent about 5% only; phloem proliferation tissue extensive, a few cons¬

picuous simple laminae usually present, of small radially elongate stone

cells, more or less uniform in amount throughout bark, not distorting

secondary phloem. Outer bark structure: up to 3, occasionally more, thick

(2-3 mm.), sheet-like rhytidome layers present together. Sloughing: as

several layers of thick, more or less elongate, layer-thick or chunky scales.

Surface: firm, powdery; usually with more or less regularly distributed and

close square section fissures; ridges variously rough; bole more or less re¬

gularly fissured.

Variants:

1. Shorea kunstleri: Periderms multiple sheets in structure.

335

Gardens Bulletin, S .

2. Shorea andulauensis: Periderms: structure: phellems locally undu¬

late, single at scollop bottoms, multiple at their sides. Inner bark structure:

expansion tissue apparently absent; phloem proliferation tissue localised

into tangential sheets of small, circular stone cells. Outer bark structure:

rhytidome layers scollop shaped, to 17 cm. long X 5 cm. wide X 2-4 mm.

thick in middle; overlapping unevenly on TS and LS, but more or less

forming continuous sheets; tissues becoming loosely powdery on ageing.

Sloughing: rhytidome layers sloughing entire as scollop shaped scales, or

Assuring and shedding in pieces. Surface: fissures narrow, localised in

zones.

Discussion: The surface appearance is within the range of Scaly—a the

differences in inner bark structure can only be made out with difficulty in

the forest so this is considered to be a Variant not a different Manifestation.

3. Shorea singkawang, S. leptoclados and butts and overmature boles

of other Red Meranti species: Periderms: one superficial, in places with

another scolloping one. Outer bark structure: heterogeneous, thin areas,

and areas with one rhytidome layer. Surface: heterogeneous, smooth with

scattered scales or with zones of irregularly fissured, elongate-scaly bark.

4. Shorea leptoclados: Outer bark tissues becoming pale, cream colour¬

ed, free from tannins. Sloughing: papery flakes, free or in bundles. Surface:

dull purple, either at periderm; or not, cream (note similarity with Variant

2 of Scaly—c).

5. Shorea pauciflora (563): Sloughing: as Variant 4.

Occurrence: Bole and crown; Shorea Red Meranti; crown S. alhida

(Meranti Pa’ang).

MANIFESTATION Scaly—b

Diagnosis: Periderms: several, scolloping regularly on TS and LS; phel-

loderm apparently absent. Inner bark structure: expansion tissue as Scaly—a;

phloem proliferation tissue: widespread, of circular stone cells conspicuous

like sand grains on cut TS; sometimes in tangential sheets. Outer bark

structure: 2-3 scollop shaped rhytidome layers, average size 3 cm. wide X

9 cm. long. Sloughing: of all rhytidome layers, variously as ridge-wide,

layer-thick or slightly chunky, more or less square scales. Surface: rough,

powdery; grid-cracked, cracks 2—3 cm. apart.

Occurrence: Boles: Shorea acuminata and S. quadrinervis (Red Meranti).

MANIFESTATION Scaly—c

Diagnosis: Periderms: pattern variable, TS and LS either distantly or not

scolloping, or, rarely, closely scolloping and forming small (c 4 cm.)

scollop shaped rhytidome layers. Structure: periderm thickness variable

(mean 0.40 mm. (0.01-1.20 mm.)); phellem dark brown or black, phel-

loderm continuous, pale , conspicuous and relatively thick (mean 0.3 mm.

(0.00-0.80 mm.)) sometimes of layers of alternating orange and cream

tissues. Inner bark structure: expansion tissue: tangential extent 8 (0-18)

%, in fingers of tangentially elongate stone cells becoming more abundant

outwards and often clustered; radially oblique stone cell fingers often pre¬

sent; extensive parenchymatous phloem proliferation tissue, sometimes

becoming more abundant outwards. Outer bark structure: rhytidome

layers sheets, or elongate strips, or rarely scollop shaped; 0,1,2 or occa¬

sionally more present together; thickness varying from 1.5-7.0 mm.; tis¬

sues usually same colour as inner bark, rarely becoming pale. Sloughing:

scales layer-thick, usually large, either cracking or Assuring before slough¬

ing, or not—hence either elongate or isodiametric. Surface: usually near

periderm, surface dull, powdery, with hue of dead phloem tissues, or

sometimes at or weathering down to a periderm, then very dark in colour;

fissures absent, or present and as variable as the Type (sometimes mere

cracks); sometimes surface scroll marked.

Variants: Some of the Variants differ considerably from the Manifesta¬

tion but were only seen on one or a few trees; if they prove typical of

their species their status need reconsidering.

336

Vol. XIX. (1962).

1. Shorea geniculata (one tree, perhaps a typical), S. lumutensis, S.

laevifolia and S. laevis (sometimes): Outer bark: of 2-4 rhytidome layers.

Sloughing: scales usually elongate, adherent. Surface: more or less regu¬

larly fissured, fissures of irregular section, penetrating several rhytidome

layers; hence bole impression ‘fissured*.

Discussion: The essential feature of this Variant is the thick outer bark;

this depends on the structure of the periderms. The property of the peri¬

derms which hinders sloughing of the first two species is not known. The

latter two have on all the trees examined thin (0.04 mm.) phelloderm or

none and fibres penetrating the mature periderms, which arc not continuous

but in 4 cm. lengths; hence the rhytidome layers persist for some time, not

sloughing until the fibres have rotted through.

2. S. guiso, S. ochrophloia: Outer bark tissue becoming pale, cream or

fawn coloured and loose, soft and powdery. Sloughing: not quite down to

the periderm; hence bole with a characteristic yellow-grey tinge.

Discussion: The essential feature of this Variant is the physiological

change which takes place in the rhytidome after periderm formation. Note

the similarity with Variant 4 of Scaly—a.

3. S. foxworthyi: A similar change to the above, but caused by weather¬

ing and only affecting the thin film of phloem tissue outside the outermost

periderm.

4. S. inappencliculata: Periderms: scolloping on TS and LS, cleanly rup¬

turing fibres. Outer bark: rhytidome layers 4-7 mm. thick X 3-4 cm.

wide X 7-12 cm. or more long. Sloughing: elongate, adherent, layer-thick

scales, whole scollops or large fractions of scollops. Hence: surface scroll

marked, fresh surfaces powdery, later eroding down to periderm, so colour

darkening. The two trees seen were both similar, with these unique features;

they were exposed at a tracksidc and are possibly atypical.

5. S. sp. now 269: Periderms with blackish phellem and cream

phelloderm; hence scrape strikingly coloured. Surface not powdery, very

dark coloured.

Occurrence: Boles, Shorea, Balau.

Discussion: This is a broadly phrased diagnosis which encompasses the

variable mature bark of the 17 species of the Balau group of Shorea ex¬

amined. The variations are mainly in periderm pattern and structure and

these cause variations in outer bark structure (thickness and shape of rhy¬

tidome layers), sloughing pattern (scale size, shape and persistence) and

surface (Assuring).

MANIFESTATION Scaly—d

Diagnosis: Periderms: seldom more than 2 present together, distantly

scolloping (33 cm.) on TS and LS; thick (to 0.4 mm.) mostly phellem,

phelloderm thin or apparently absent; becoming dark coloured, hard and

brittle on maturing. Inner bark structure: tissues yellowish becoming green

towards cambium; expansion tissue either apparently absent or in abundant

fingers, tangential extent 15 (4-30)%, each becoming wider outwards, of

parenchyma or small stone cells in which are sometimes set short radial

files of tangentially elongate stone cells; phloem proliferation tissue absent,

localised or widespread, of parenchyma or circular stone cells only. Some¬

times with damar channels exuding liquid damar when cut. Outer bark

structure: usually heterogeneous; partly with / (rarely 2) sheet like rhyti¬

dome layer(s), 3-4 mm. thick, partly with none, outer bark thin and super¬

ficially fissured; sometimes mainly thin. Sloughing: elongate, ridge-wide,

layer-thick scales; sloughing plane within periderm, scales loose hanging

before sloughing. Surface: at periderm, dark coloured, not powdery, some¬

times turning grey, usually heterogeneous, rhytidome a single layer or ab¬

sent; narrow square section fissures (less than 5 mm. wide X 5 mm. deep),

or cracks, irregularly distributed: marks of scollops remaining as scroll

marks (frequency variable) after sloughing: these fissured areas interpersed

with smooth areas, often finely lenticellate, which become larger on old

trees, to dominate appearance: sometimes with black or yellowish pendent,

fine damar stalactites. Visual impression: dark, heterogeneous, smooth and

scaly.

337

Gardens Bulletin, S .

Variants:

1. Shorea bala nocarpoides (355): Inner bark: expansion tissue occupy¬

ing 60% of surface, in abundant, wide fingers.

2. Shorea hopcifolia (306), S. multiflora (171): Inner bark: expansion

tissue apparently absent; phloem proliferation tissue extensive. Outer

bark: rhytidome layers 6-7 mm. thick. These are overmature trees (res¬

pective girths 12 ft. and 15 ft.). The cut surface of the inner bark is simi¬

lar to Scaly—a in appearance.

Occurrence: Boles, Shorea, Meranti Damar Hitam.

Discussion: Periderms: few fibres connect the inner and outer bark through

these thick periderms hence the rhytidome layers tend to hang loose before

sloughing. The sloughing plane is within but very near the outside of the

phellem.

Damar is often extruded through the inner bark and through cracks in

the innermost periderm and may spread out over the outer surface of this

periderm and so perhaps hasten the separation and sloughing of the rhyti¬

dome layer. Thus the surface may have a thin damar coating and this con¬

tributes to the dark, dull bole colour. In some species the damar is ex-

extruded as far as the outer surface of the bark where it hardens in con¬

tact with the air and forms conspicuous drips, diagnostic of their species

(see Symington, 1943).

MANIFESTATION Scaly—e (Fig. 9b)

Diagnosis: Periderms: 1 superficial periderm, in places with another

deeper 1 or rarely 2; not anastomosing or distantly, and forming very

large (50 cm. across) or, rarely, smaller (15 cm.) scollops; periderms pale;

average thickness 0.6 mm.; phellem usually hard and sclerified, continuous

phelloderm usually present, and fawn so not conspicuous or clearly de¬

marcated. Inner bark structure: expansion tissue variable in amount, appa¬

rently absent or in fingers of parenchyma, rarely with a few files of tangen¬

tially elongate stone cells too, usually widening outwards; occupying tan¬

gential extent 10 (0-30) %; some rays becoming wider than others; rarely

pseudocortex present; phloem proliferation tissue extensive, of small stone

cells amount usually variable in different tangential zones of the bark;

sometimes, especially in outer parts, distorting radial arrangement of

secondary phloem; silica sometimes deposited. Outer bark structure: hetero¬

geneous; thin areas and others with 1 (rarely 2) rhytidome layer(s).

Sloughing: scattered layer-thick scales, usually large and more or less iso-

diametric, loose hanging or adherent before sloughing. Surface: firm, pow¬

dery; fissures usually absent, except in species with several continuous rhy¬

tidome layers; usually with close set round lenticels: occasionally a few

scroll marks; surface sometimes gently undulate with depressions and bul¬

ges; visual impression: heterogeneous, smooth with variously scattered

scales.

Variants:

1. Dipterocarpus appendiculatus: Outer bark: several rhytidome layers

present together all over bole. Sloughing: of small loose hanging, elongate

scales all over bole. Surface: tendency to fissure over whole bole so visual

impression scaly-fissured.

2. D. costulatus, D. rotund ifolius: intermediate between the ‘typical*

species and D. appendiculatus.

On the available data the differences between D. appendiculatus and the

other species are not sufficiently clear cut to consider the bark as a different

Manifestation. Examination of other fissured and scaly-fissured species (for

instance D. chartaceous , D. ohtusifolius, D. rigidus, and D. tuberculatus —

the last three from Burma) might necessitate such a step.

3. D. crinitus: Periderm: 1.0 mm. thick with a very thick (0.8 mm.),

pale, easily visible phelloderm.

4. D. sublamellatus: Periderm: thin, 0.2 mm.

5. D. cornutus , D. verrucosus: Inner bark: phloem proliferation tissue

so distorting secondary phloem that a pseudocortex is formed in some

trees.

6. D. verrucosus: Periderm: thin, 0.1 mm.; rhytidome layers thin, only

2 mm.; surface tinged reddish.

Occurrence: Boles; Dipterocarpus.

338

Vol XIX. (1962).

Discussion: Inner bark; expansion tissue and phloem proliferation tissue

are the same as Smooth—h; further the expansion tissue is, like Scaly—d,

very variable in amount between individuals and species. Amount of phloem

proliferation variable between different trees and different species. Some¬

times tangentially adjacent, proliferated phloem blocks become aligned

into sheets just visible as fine lamellae to the unaided eye, these are much

finer than the ‘simple laminae’ of Scaly—a and —b. Tissues just below the

superficial periderm tinged green in a few samples presumably due to the

presence of chlorophyll. The differences between species in appearance is

largely due to differences in details of the periderm pattern and structure,

hence in the sloughing pattern (scale size, thickness, closeness, and whether

loose hanging or adherent), but there is considerable variation and over¬

lap between species. Surface: the size and closeness of the lenticels varies

between species.

MANIFESTATION Scaly—f

Diagnosis: Periderms: distantly scolloping, c.5 cm. on TS, greater than

17 cm. on LS; structure: 0.1-0.2 mm. thick, phellem hard, black, phel-

loderm absent or thin (0.04-0.02 mm.). Inner bark structure: expansion

tissue apparently absent or in fingers widening outwards or not, tangential

extent 10 (0—19) % also sometimes a few rays with short radial lengths

of tangentially elongate stone cells, increasing in number outwards;

phloem proliferation absent or present and extensive, parenchymatous.

Outer bark structure: rhytidome layers, broad sheets or elongate strips,

1 or 2 present together, or rarely none; outer bark tissues mid brown.

Sloughing: close or scattered, elongate or isodiametric, loose hanging or

adherent scales. Surface: dark coloured, commonly blackish brown, at a

periderm so not powdery, fissures present, as variable as the Type (q.v.);

scales not fissures dominating appearance.

Variants: The considerable variability between the species of Hopea

examined is discussed with the notes on the genus.

1. H. helferi: Surface powdery, not at a periderm.

2. Balanocarpus heimii: Inner bark structure: expansion tissue absent

except sometimes for a few clustered, short, parenchymatous fingers, more

abundant outwards; phloem proliferation tissue absent or present and more

abundant outwards. Outer bark structure: rhytidome layers elongate strips.

Sloughing: close, elongate, loose (rarely adherent) scales. Surface: close,

irregular section fissures.

Occurrence: Boles; Scaly Barked Hopea; Balanocarpus.

MANIFESTATION Scaly—g

Diagnosis: Periderms, pattern: TS and LS; either sheet like not scollop¬

ing or, occasionally, distantly (7 cm.) scolloping; structure: continuous not

penetrated by fibres at all or phellem (first formed) penetrated then phel-

loderm (later formed) rupturing fibres; thin (0.2 mm. (0.04-0.5 mm.)),

phellem two zoned with inner impervious dark blackish brown and outer

biscuit coloured layers; phelloderm absent or thin (0.02 (0.0.4) mm.),

inconspicuous, merging with phloem. Inner bark structure: expansion tissue

in fingers, tangential extent 9 (4-13) %, usually parenchyma, fingers either

becoming more numerous or widening outwards, sometimes a few connate;

widespread, scanty phloem proliferation tissue, forming small stone cells

and, or, parenchyma, and varying numbers of extensive, tangential sheets

of radially elongate stone cells, 0.2-0.4 mm. thick and visible as fine

lamellations to unaided eye. Outer bark structure: bole heterogeneous;

cither several, extensive, sheetlike (rarely scollop-form) rhytidome layers,

or 1, or 0, forming irregularly, never in zones, up bole; rhytidome layers

commonly continuous with inner bark at top end, otherwise completely

free. Sloughing: in one or more layers, irregularly over bole; scales elon¬

gate, commonly large (60 cm. X 10 cm.), curling up at their edges on

desiccating and loose hanging except at top ends, or smaller, or adherent;

sloughing plane within phellem , pustular lenticels often developing and

assisting rupture. Surface: fissures present or absent, square or irregular

section, irregularly scattered, small pustular lenticels often present, bright

reddish purple when new, hence fresh surfaces with purplish tinge; at a

periderm, always pale when new sometimes becoming very dark brown

or black by erosion of part of phellem.

339

Gardens Bulletin, S.

Variant:

1. Dryobalanops rappa: Periderms: like Scaly—c in structure.

Occurrence: Boles; Dryobalanops.

Discussion: The plane in which the periderm forms, and the pustular

lenticels actively assisting sloughing are unique to this bark. Scales are

able to curl up as the tissues dry out because no fibres connect one rhyti-

dome layer with another.

MANIFESTATIONS OF SURFACE ROTTEN BARK

Key to the Manifestations of Surface Rotten bark

A Laminae present in inner bark, usually long and occupying most of

bark, sometimes short and scattered; periderms almost continuous

though irregularly undulate.

B Fissures and cracks sparse or absent; outer bark with thick and

thin areas; laminae always throughout whole inner bark .... c.

BB Fissures present, though of irregular distribution and size; outer

bark thick everywhere; laminae as above or small and scat¬

tered . b .

AA Laminae absent; periderms absent or localised as narrow, elongate,

tangential strips at ridge sides . a, Shorea albidcu

MANIFESTATION Surface Rotten—a (Fig. 10c)

Diagnosis: Periderms: absent, or localised as scattered, elongate, tangen¬

tial strips penetrating into ridges from fissure sides. Inner bark structure:

expansion tissue apparently absent; phloem proliferation tissue, extensive

stone cells and parenchyma, but tangential laminae absent. Outer bark

structure: very variable in thickness, sometimes absent, to 9 mm. thick,

following surface contours; dark coloured, variously rotten tissue; not deli¬

mited inwards by periderm. Sloughing: either as elongate scales of vari¬

ous thicknesses and size, just bundles of fibres, seldom cut off by a peri¬

derm or, where outer bark soft and the fibres rotten, as more or less

isodiametric small, chunky scales. Surface: surfaces black and firm where

periderm bounded, elsewhere soft and powdery, fissures regularly distri¬

buted, in shape just irregular furrows in the phloem to 10 mm. deep X 5

(10) mm. wide X 20-30 cm. long, sides rough with torn fibre bundles;

ridges anastomosing; appearance fibrous-fissured.

Occurrence: Boles, Shorea albida only.

Discussion: The absence or only localised development of periderms

means that surface configurations are more or less completely controlled

by the inner bark structure, hence the obviously fibrous nature of the

ridges and scales. The absence of laminae is in marked contrast with

Manifestations—b and —c.

MANIFESTATION Surface Rotten—b

Diagnosis: Periderms: usually continuous, sometimes with scattered gaps,

pattern variable, together may form a closely anastomosing reticulum

throughout the outer bark; thickness variable, often strongly undulate.

Inner bark structure: phloem proliferation tissue: simple laminae present,

either scattered and short, or, more often, occupying most of the bark

and greater than 20 cm. in extent on TS and LS; interlaminar phloem

blocks often with parenchyma proliferation tissue, expansion tissue either

in fingers interrupting the laminae or apparently absent and the laminae

continuous sheets. Outer bark structure: no well defined rhytidome layers;

usually with continuous periderm at boundary with inner bark, this boun¬

dary irregular and not parallel to cambium. Sloughing: elongate, layer-

thick or chunky scales. Surface: surface firm and black and at a periderm

or not, and powdery; appearance often dominated by scales not fissures;

fissures irregular furrows of variable section and penetration; irregular dis¬

tribution; variable between individuals.

Occurrence: Boles; Shorea, Meranti Pa’ang, most species.

340

Vol. XIX. (1962).

Discussion: The anastomosing, reticulate periderm pattern is very similar

to Smooth—e (also Meranti Pa'ang); it differs in the thinner and more

widely separated periderms. Inner bark: identical with Surface Rotten—c

where strongly laminate. Surface: the ridges do not have the roughly fibr¬

ous structure of Surface Rotten—a.

Shorea bracteolata, the only species with this Manifestation collected

several times, had some trees with scattered and other with extensive lami¬

nae. The amount of lamination did not vary much between dilferent parts

of a tree, nor did it seem to be correlated with girth. This evidence sug¬

gests that the amount of lamination is variable between individuals of a

species.

MANIFESTATION Surface Rotten—c

Diagnosis: Periderms: continuous, extensive, every one lying along the

side of a stone cell sheet. Inner bark structure: expansion tissue appa¬

rently absent or as scattered fingers. Phloem proliferation tissue: strongly

developed simple laminae more or less filling inner bark , between them the

phloem blocks sometimes with parenchyma proliferation tissue. Outer bark

structure: heterogeneous; thin, or thick with sheet like rhytidome layer(s);

inner edge more or less parallel to cambium. Sloughing: isodiametric,

usually layer-thick scales. Surface: surface dark, firm at a periderm; loose

scales in one or occasionally two layers, close or scattered over bole sur¬

face; cracks and fissures sparse or absent.

Occurrence: Shorea , Meranti Pa’ang, on butts, and lower boles some¬

times extending over whole boles.

Discussion: This Manifestation is dominated by the inner bark structure;

the strongly developed laminae determine the arrangement of the periderms,

hence the structure of the outer bark and the sloughing pattern, as well as

the surface configurations. This Manifestation may perhaps be either an

overmature stage or a reaction of the tree to peculiar environmental

conditions.

MANIFESTATIONS OF LAMINATE BARK

Key to the Manifestations of Laminate bark

A Surface closely (1-3 cm.) fissured, ridges rough, with large, powdery,

brownish-yellow areas; sloughing of small scales; laminae coarse

(about 0.45 mm. thick) . a. section Pilosae.

A A Surface with widely spaced (5-8 cm.) fissures, or smooth with scat¬

tered fissures, ridges more or less flat, mostly firm, yellowish-cream

in colour; sloughing of mid-sized, flat scales; laminae fine (about

0.25 mm. thick) . b, section Glabrae.

The evidence on which these Manifestations are distinguished is

fully discussed in Whitmore (1960, p. 205).

MANIFESTATION Laminate—a (Fig. lOd)

Diagnosis: Periderms: usually continuous; one more or less delimiting

inner edge of outer bark, remains of another one eroding away nearer sur¬

face. Inner bark structure: as Type; laminae coarse, about 0.45 mm. thick,

about five fibre blocks included. Outer bark structure: as Type. Sloughing:

small, chunky or layer-thick scales, 1-3 (5) cm. wide. Surface: fissures

close, ridges, 1-3 (5) cm. wide, surfaces usually very rough; either at a

periderm and firm, yellowish-cream or not and powdery, brownish-yellow.

Occurrence: Anisoptera, Pilosae section (? all).

MANIFESTATION Laminate—b

Diagnosis: Periderms: continuous; one superficial, with or without an¬

other deeper one too. Inner bark structure: as Type; laminae fine, about

0.25 mm. thick, about 4 fibre blocks included. Outer bark structure: locally

thin, thick areas as Type. Sloughing: flat, ridge-wide (5-8 cm.) scales,

usually layer-thick, less often chunky; some areas not sloughing. Sur¬

face: wide spaced fissures; ridges 5-8 cm. wide usually more or less

flat; surface may become smooth with scattered fissures and scales; usually

at a periderm, firm, yellowish-cream.

Occurrence: Anisoptera, Glabrae section.

341

Gardens Bulletin, S.

ANISOPTERA

Taxonomy

Anisoptera Korth is a comparatively small clearly defined genus

of about 14 distinct species although the specific limits are some¬

times unclear (A. oblonga). Two sections of the genus are recog¬

nised with very distinct floral characters (Heim 1822, Symington

1943, Wyatt-Smith 1955), section Glabrae, A. laevis and A. sea -

phula, and section Pilosae, with all the other known species.

Argument

The Bark Type sequence is shown on Fig. 1. All the species are

characterised by Laminate bark on mature trees and have the

diagnostic appearance of the Type. There is no evidence that

Laminate bark is succeeded by another Type, the Manifestation

may change.

Conclusions

This examination confirms the findings of previous botanists

that Anisoptera is a homogeneous group of species, Laminate bark

is not found elsewhere. Fig. 1 shows a clear difference between

the two generic sections. A. laevis and A. scaphula differ from the

other species in:

1. The very large size reached (Wood, 1957 (under Shorea

superbd) records that the largest known dipterocarp is

an Anisoptera probably A. laevis),

2. Manifestation Laminate—b, not Laminate—a, is develop¬

ed on trees between 6 ft. and 11 ft. girth. Even if

Laminate—b changes later to —a the two sections

differ in the girth range where they both occur.

This clear cut difference has not previously been recognised,

neither Symington (1943) nor Wyatt-Smith (1955) distinguish be¬

tween species on bark characters; it coincides with the accepted

taxonomic division and should be easy to see in the forest (see

key to Manifestations); in particular the two commonest Malayan

species A. curtisii and A. laevis can be distinguished on bark

characters alone.

BALANOCARPUS

Taxonomy

Balanocarpus Bedd. had become a repository for dipterocarps

with wingless fruits, long recognised unsatisfactory, before Syming¬

ton (1933, 1938) removed most of the species to more natural

systematic positions in Shorea and Hopea. The only Malayan

species left is B. heimii King with no close relatives and as similar

342

Vol. XIX. (1962).

Girth in feet

Pilosae

Glabrea

curtisii

a»

oblonga

scaphula

\ x

\ \

•

\ l

\ \

s '

*1-

***

+ +

■f-

4 -

-r

;♦

:♦

KEY

to Flgs.1-8

girth of

change of

bark type

Smooth

bark

Dippled

bark

ditto where

there is

considerable

overlap

every

individual

and Its

manifestation

Where

manifestation^

is uncertain

or tree is

heterogeneous)

see legend ?

to figure r

•

Shallow

Fissured

bark

Deep

Fissured

bark

Scaly

bark

maximum

usual girth

Surface

Rotten

bark

Laminate

bark

Fig. 1. ( A ) Anisoptera and (B ) Balanocarpus, bark sequence through life

and Key to Figs. 1-8.

Construction explained in appendix 2.

343

Gardens Bulletin, S.

to the Euhopea and Pierrea groups of Hopea as to Balanocarpus

Bedd. Desch (1941) includes B. heimii with Hopea in his account

of dipterocarp timbers and a recent analysis of the anthocyanins

of the leaves (Bate-Smith and Whitmore 1959) shows the two

genera to be similar m these biochemical characters.

Argument

The Bark Type sequence is shown on Fig. 1. The sapling has

the common Manifestation Smooth—b. Scaly bark has developed

by 3 ft. girth and is very similar to Scaly Hopea bark, it is con¬

sidered a Variant of the same Manifestation, Scaly—f.

The inner bark structure is unique with fibre blocks of different

sectional size in different tangential sheets, so that the cut trans¬

verse surface sometimes appears streaked. Faint radial striations

are visible on radial longitudinal surfaces and ripple marks on

tangential longitudinal ones, a feature otherwise seen only in

Hopea ferrea.

Conclusions

The mature bark is the same Manifestation as the field group

Scaly Barked Hopea with fine structural differences, it can be

confused in the forest; it is distinct from Shorea Balau with which

it has sometimes been confused. Scaly Barked Hopeas have vari¬

able bark and from the material available I have been unable to

decide whether this is taxonomically significant. Without further

investigation I cannot equate B. heimii to one or several of these

species but they are no doubt, as Symington remarked, closely re¬

lated. Most, of the Scaly Barked Hopeas are Euhopea and 1 con¬

clude therefore that the true relationship of B. heimii is here

rather than with Pierrea (the Smooth Barked field group).

DIPTEROCARPUS

Taxonomy

Dipterocarpus Gaertn. f. is a well defined homogeneous genus

of about 80 species. There are five artificial sections differing in

fruit characters, these do not have the same significance as the

natural subgroups of Shorea , all the species have much in common.

Argument

Forest observations of 25 species show that the 11 described

here cover the range of the genus. The Bark Type sequence is

shown on Fig. 2. Manifestation Smooth—b is a widely occurring

bark, it is probably intermediate here to Smooth—h, unique, and

to Scaly—e, also unique.

344

Vol XIX. (1962).

Girth in feet

DIPTEROCARPUS

cornutus

costulatus

crinitus

dyeri

fagineus

kerrii

pseudofagineus

rotundifolius

sublamellatus

verrucosus

\ \

b s

N '

\ k

\ >

b N

\ N

-G_

\ '

—

s k

\ '

“WT

r<2"

G-

g g

—

g”

•

w~w

—

g”

—

G“

e‘

—

•

—

G-

—

__ _

•

<2-

—

L #-

—

•

—

1 w

DRYOBALANOPS

Fig. 2. Dipterocarpus and Dryobalanops, bark sequence through life.

Construction explained in appendix 2; key on Fig. 1.

* Dryo. rappa G. 15 ft.: See text.

345

r* 1 1 , • i i t 1 ?, i 1 V /\ r °pp a

Gardens Bulletin, S.

Conclusions

Previous findings are confirmed, all the species of Dipterocarpus

in Malaya have similar bark, the same Manifestations with some

minor Variants, in the same sequence. The slash of mature tree

bark. Scaly—e, is distinctive in the forest due to various unique

features of the inner bark structure. The surface pattern though

can be confused with Scaly—a, Variant 3. A number of species

are distinguished as Variants. From its bark Dipterocarpus is a

natural homogeneous genus which cannot be divided. I am unable

to draw any sharp and constant distinction between the bark of

the closely related species D. fagineus and D. pseudojagineus.

DRYOBALANOPS

Taxonomy

Dryobalanops Gaertn. f. is a homogeneous well defined genus

of nine known species.

Argument

The Bark Type sequence, Fig. 2, shows that an unique Mani¬

festation Scaly—g is present on mature trees of the four species

examined and the published descriptions show it on three more.

One tree ( D . rappa, girth 15 ft.) had a bark with different peri¬

derm pattern hence surface appearance; it is thought to be ab¬

normal and overmature (diagnosis, Whitmore 1960, pp. 373-4).

Conclusions

Dryobalanops has an unique Manifestation on mature trees,

individual trees differ from each other particularly in details of

the sloughing pattern as is common in Scaly barks, and this varia¬

tion overlaps any differences there may be between species. The

genus can be distinguished in the forest on bark characters from

Dipterocarpus, Shorea Balau, and the Scaly Barked Hopeas which

it resembles (Symington 1943). Previous conclusions are confirm¬

ed, on bark characters Dryobalanops is a natural homogeneous

group of species.

HOPEA

Taxonomy

Hopea Roxb. (about 100 species) is, like Shorea, a heterogene¬

ous genus and there is considerable variation between species in

forest, herbarium and wood anatomical characters. The last com¬

plete revision was by Brandis (1895), now manifestly inadequate;

Symington (1934, 1939) made what revisions he could but was

hampered by the poor knowledge of the Bornean flora; his tenta¬

tive subdivisions were not of equal status and some species can¬

not be referred to any of them. These subdivisions do not corre¬

late well with the timber or forest characters and so in 1943 he

346

Vol. XIX . (1962),

Smooth

M.K

Fissured

Scaly

Girth in feet

griff ithii

nervosa

latifolia

vesquei

beccariana

mengarawan

sulcata

*♦-

helferi

nutans

odorata

sangal

semicuneata

Is!

\ n

s N

^ N

\ s

N '

\ '

\ N

s N

\ \

\ N

N N

\ \

s \

■\

\ N

, \

s \

X-

✓

• k

/s'

•

\ >

/'/

—

"2r

"T”

\ j

—

•

✓ /

—

Fig. 3. Hopea, bark sequence through life.

Construction explained in appendix 2; key on Fig. 1. M.K.

Mata Kuching group.

347

Gardens Bulletin, S.

proposed ‘tentative arbitrary field groups’ cutting across the

generic subdivisions and using forest characters of buttress, stilt

roots, bark and damar exudate. The groups are:

1. Smooth Barked Hopeas

2. Mata Kuching Hopeas (a segregate of 1 with damar in

globular or stalactitic exudations)

3. Fissured Barked Hopeas

4. Scaly Barked Hopeas.

Argument

The study of the bark of natural groups of species in Diptero-

carpaceae shows that, with the exception of Shorea Meranti

Pa’ang, all or most of the species of the groups have many bark

characters in common. Hence the bark survey of Hopea is of

particular interest. If it is possible to show the species of Syming¬

ton’s field groups have similar bark this will suggest that they are

the natural groups and the information will assist in the pending

revision of the genus and the description of new Bornean species

(Wood for instance has described a new genus, Neohopea, in

manuscript). Unfortunately I have not much material, in some

cases only a single sample of a species, nevertheless if these sam¬

ples have a lot of bark characters in common the induction is

clear; I cannot though propose any new groupings.

The Bark Type sequence is shown on Fig. 3. I have no material

showing the scaly bark reported (Symington, 1943) for large trees

of the Smooth and Mata Kuching groups.

All the species have similar secondary phloem structure except

in the Scaly Barked group where there are differences between

species in the shape and size of fibre blocks in cross section and

whether they have a surrounding crystal sheath or not.

Conclusions

The species with similar bark coincide exactly with Symington’s

‘tentative arbitrary field groups’ and there are no bark features

common to the whole genus. The conclusion is that these, rather

than the existing taxonomic sections, are the natural subgroups of

Hopea.

1. The Smooth Barked and Mata Kuching Hopeas have

together a unique Manifestation, j, of Smooth bark and are

indistinguishable. The barks of two trees (girths 5 ft. and 6

ft.) of undescribed species (sp. A (the code name at Kepong)

and sp. 324 (not matched at Kepong, specimen deposited

there)) and of the Bornean H. vesquei with Smooth bark were

examined and undoubtedly belong to one or other of these

groups; the two former species are not shown on Fig. 3.

348

Vol. XIX. (1962).

2. Fissured Barked Hopeas all have the same Manifesta¬

tion, Deep Fissured—a. of a unique Bark Type. Further re¬

search might show that the Bornean tree of H. beccariana

(488) is not the same species as the Malayan ones (180, 197).

3. Scaly Barked Hopeas are on the evidence available one

Manifestation, f, of Scaly bark but there are differences in

surface pattern (Symington, 1943) suggesting there may be

other Manifestations; this is the only species group in which

differences in secondary phloem between species have been

seen. Further data is needed to see whether this group is

homogeneous and natural.

SHOREA

Shorea Roxb., the largest and economically most important

genus of Dipterocarpaceae, has about 170 species. It is not a

homogeneous genus but contains representatives of several more

or less closely related groups some of which might be considered

worthy of generic rank. The established taxonomic sections

(Brandis, 1895) are patently unsatisfactory today. Symington

(1943) using Malayan material divided Shorea into four natural

subdivisions differing considerably from Brandis 1 sections, but was

not prepared to give them definite botanical names or status until

the whole genus was revised taxonomically. This revision still has

not been made. Desch (1936, 1941) independently established four

groups based on timber characters which closely correspond to

Symington’s subdivisions, but Bate-Smith and Whitmore (1959)

found no significant differences between the groups in the antho-

cyanin pigments of mature leaves. I have studied the bark of

Symington’s natural subdivisions separately.

Shorea, Balau group

Taxonomy

This group approximates to section Eushorea, Brandis (1895),

but has never been satisfactorily defined botanically. Symington

(1943), mainly on Malayan species, describes three botanical sub¬

groups indistinguishable in the forest and from wood anatomy.

Argument

The Bark Type sequence is shown on Fig. 4. An unique Scaly

bark (Manifestation—c) develops from Smooth bark (Manifesta¬

tion—b) at small girth and persists. The bark of sp. nov. 555 is

quite distinct, and of uncertain affinity. 1 In the crown there is an

1- Inner bark as Scaly but periderm pattern unique; surface appearance

reminiscent of Tristania (Myrtaceae)—see Whitmore (1960, p. 32). A

herbarium specimen of this tree, collected at Lungmanis, North Borneo, is

at CGE.

349

Gardens Bulletin , 5.

Ciliata

Barbata

Girth in feet

collina

0»

foxworthyi

geniculata

guiso

lumutensis

Cs)

•*->

meadiana

ochrophloia

glauca

’>

inappendiculata

sp.nov. 555

sp.nov. 569

\ \

b s

\ s

\ \

\ x

s \

N s

•

\ N

-Jv

\ '

—

•

““

—

•

___

mmm

★

—

•

—

— —

—

•

—

i j

18'

—

cnI

Fig. 4. Shorea, Balau group, bark sequence through life.

Construction explained in appendix 2; key on Fig. 1.

* S. sp. nov. 555 G. 6 ft.: see text.

350

Vol XIX. (1962).

intermediate gridcracked stage between Smooth and Scaly which

is unique and diagnostic. In S. laevifolia, S. laevis and 5. glauca

the secondary phloem fibre blocks are of various sizes as seen on

transverse section. Intercalary proliferation tissue from rays and

phloem blocks has aligned many of them into slightly convex,

short, imbricating, tangential plates with smaller blocks at the

ends than centre, the included parts of the phloem rays are not

widened. S. geniculata approaches this condition. The pellucid

conducting phloem occupies about one-third of the inner bark,

more than in any other group examined.

Conclusions

Sixteen Balau species examined have the unique Manifestation

Scaly—c, one species seen once has a bark of uncertain affinity.

Although there is the considerable and overlapping environmental

variation between species and individuals which is common

amongst Scaly barks a number are quite distinct Variants and

more could probably be distinguished by further study.

There are a number of constant differences between this Scaly

bark and Scaly Hopea, Balanocarpus and Dryobalanops with

which it has sometimes been confused in the forest. Some species

and small groups of species are distinctive (Symington has already

described the Malayan ones).

The species examined are of the Barbata and Ciliata botanical

subgroups, these are not clearly distinct on bark characters, there

is overlapping variation. Thus three of the four Barbata species

(S. glauca, S. laevifolia and S. laevis) have a unique arrangement

of the fibre blocks but the Ciliata species S. geniculata approaches

this condition; two Barbata species (5. laevifolia, S. laevis) and

two Ciliata species (5. geniculata, S. lumutensis) have fissured,

thick outer bark. S. maxwelliana (Barbata) has neither of these

features.

The bark of the new undescribed species ( sp . nov. 555 and

sp. nov. 569 1 ) and S. inappendiculata (Borneo) does not have

either of the Barbata features and hence they are probably in the

Ciliata subgroup; the two undescribed species, if typical, have

specific bark.

Shorea, Meranti Pa’ang group

Taxonomy

A natural homogeneous group corresponding closely to section

Anthoshorea Brandis, though Brandis enumerated here many spe¬

cies now known not to belong.

1- Herbarium specimens at BO, CGE, L, SAN and SING.

351

Gardens Bulletin, S,

Fig. 5. Shorea, Meranti Pa’ang group, bark sequence through life.

Construction explained in appendix 2; key on Fig. 1.

352

Vol XIX. (1962).

Argument

The survey initially presented difficulties not encountered in

other groups as there is much greater variation between boles and

between the parts of each bole, so that samples from the butts of

standing trees are less often typical of the whole bole. The Bark

Type sequence is shown on Fig. 5. Six species change from Smooth

to Surface Rotten bark, unique to the group, before mature girth is

reached and remain at this; the surface appearance of this Bark

Type is highly variable. Three species retain Smooth bark through¬

out life; of these 1 examined S. symingtoniana and found the

Manifestation unique (Smooth— f); S. bentongensis and S. deal -

bata are shown on Fig. 6 on the basis of Symington’s (1943) des¬

cription. S. ochracea remains Smooth with a unique Manifestation

(Smooth—e) until at least 8 ft.; the bark at larger girths remains

undetermined. All these Manifestations have features of the peri¬

derms and of the secondary phloem structure in common (see the

diagnoses). S. talura has fissured bark, provisionally assigned to

the Shallow Fissured Type as a unique Manifestation, e.

Conclusions

No bark features are common to all the species. Four of the

eleven have different, unique Bark Manifestations on mature trees

(5. albida Surface Rotten—a, S. ochracea, Smooth—e, S. syming¬

toniana, Smooth—f, S. talura, Shallow Fissured—e), the other

seven have Surface Rotten, Manifestation—b or —c, and are indis¬

tinguishable in the forest. S. talura, incompletely studied, seems to

differ completely from the other species which do have a few

features in common. Surface Rotten bark is quite distinct from

Laminate ( Anisoptera ), see key to the Bark Types, though the two

have sometimes been confused.

There are no bark features by which the group as a whole is

recognisable in the forest, most species have variously developed

simple laminae, though not always as conspicuous as the published

descriptions suggest, throughout most of their life, hence the slash

is pale yellowish.

Even the six species with Surface Rotten—b bark are not easily

told by surface appearance because this Manifestation has ex¬

tremely variable surface appearance.

Meranti Pa’ang, then, is a heterogeneous group on bark. S.

albida, tentatively divided into two botanical forms with different

ecology and timber is not divisible and the species belongs to this

group on bark despite its timber classification as a Red Meranti

or Red Selangan (= Balau) (Browne, 1955, pp. 137, 151).

353

Gardens Bulletin, S.

Fig. 6. Shorea, Meranti Damar Hitam group, bark sequence through life.

Construction explained in appendix 2; key on Fig. 1.

* S. multiflora: Dippled -a at 3 ft., Smooth -d at 4 ft. ‘E4’ is the

code name at Kepong for an undescribed Malayan species, my

reference collection is Whitmore 655 (at CGE).

354

Vol. XIX . (1962).

Shorea, Meranti Daniar Hitam group

Taxonomy

This is a natural homogeneous group assembled by Symington

(1938) from various Shorea and Balanocarpus species. He named

it Richetia for convenience though it has not been validly pub¬

lished or given rank such as genus or section.

Argument

The Bark Type sequence is shown on Fig. 6. There is a general

similarity, bark changing from Smooth (unique Manifestation—d)

to Scaly (unique Manifestation—d) at about 3 ft. girth, with some

variations. Dippled bark was seen on S. faguetiana at 9 ft. and

S. multiflora at 3 ft. and 7 ft. girth. On the latter it was at girths

intermediate to Scaly on the former well within the girth range

of Scaly bark; nevertheless the structure indicates that it is prob¬

ably always intermediate to Scaly on a particular tree (Whitmore

1962a, b). This Manifestation is shared with Vatica, from which

there are only fine differences.

S. gibbosa alone possesses a Shallow Fissured bark usually re¬

placed by Scaly on very large trees, the Manifestation, d (Fig. 6),

is unique.

Conclusions

An unique bark, Scaly—d, is present on all these Meranti

Damar Hitam species at some stage of their mature life and holds

them together as a homogeneous botanical group. Smooth—d

and Shallow Fissured—d barks are also unique and some of their

features are shared with Scaly—d, this adds to the homogeneity.

Smooth—d and Scaly—d are easily recognised in the forest and

Dippled sometimes so. The Scaly species cannot be told from

each other. S. gibbosa with Shallow Fissured—d is the only species

recognisable from its bark.

Shorea, Red Meranti group

Taxonomy

This is the most heterogeneous of Symington’s groups and can¬

not be identified with any single defined botanical group, never¬

theless there is ample justification for recognising it as natural.

Symington recognised three subgroups in Malaya based on

floral characters, these he noted are of taxonomic rather than

practical value because the differences have not been correlated

with differences in wood anatomy. He noted some bark differences

and predicted that there might be other forest differences, a pre¬

diction which has been amply justified.

355

Gardens Bulletin , S.

Kawang

Tembaga

Girth in feet

pinanga

C»

beccariana

waltoni

macroptera

sp. nov. SAR5798

parvifolia

leprosula

dasyphylla

argentifolia

curtisii

rubella

rubra

-X

\ N

X \

•

X S

n n

x '

x N

\ N

X N

X \

\ N

s*'

|/‘

/l/l

• i

i t t i

i 11

1 b*

\ #

-v.

X N

11 1

;.!

11 1

a/b

l i

i 1 1

k 1

i i

1 l i

l'l

I 1 1

ill

'*!

i:*

. X

•a

rV~

' l

1 1

ZM>

i 1

'•!

'i i

■ i

• i

i i

a“

■ i .

!!■

‘ |

"m 4 "

l‘ 1

1 I 1

h '

★

i";

1 11

.a-

Fig. 7. Shorea, Red Meranti group, bark sequence through life.

Construction explained in appendix 2; key on Fig. 1.

*S. beccariana G. 9 ft., S. macroptera G. 12 ft., S. macrantha

G. 4 ft.: these differ in periderm pattern, hence in surface and

slash, and are thought to be abnormal because overmature or

exposed trees; for full diagnosis and discussion see Whitmore

(1960), pp. 246, 248-249, 266-7. *S. singkawang: at least some

356

Vol XIX. (1962)

Nemes u

\aa

^ \

\ \

V \

\ v

W V

I**

[i«i

l'l

kL>

"b-

•i

fo¬

trees change bark in the girth range 5-10 ft. to bark with a pale,

dippled, rugose surface, not mentioned in published descriptions

so perhaps atypical; see Whitmore (1960), pp. 264-5. *5. curtisii

and S. smithiana: see text.

357

Gardens Bulletin, S.

Argument

The Bark Type sequence is shown on Fig. 7. The species ex¬

amined have been arranged in a number of subgroups each with

a common sequence:

1. The Kawang subgroup (left of figure) retains Smooth

bark of the unique Manifestation Smooth—c throughout life

or at least well into mature life. In this subgroup are S. bee -

cariana, S. mecistopteryx, S. pinanga, S. waltoni, S. gysbert-

siana, S. macroptera , S. sp. nov. SAR 5798.

2. Tembaga subgroup (centre) has some species changing

from Smooth to Shallow Fissured bark of the unique Mani¬

festation—a at about 2 ft. girth (S. parvifolia, S. dasyphylla ,

S. rubella , S. leprosula, S. argentifolia ), and other species be¬

coming Scaly later (5. ovata, S. rubra). S. curtisii has a bark

intermediate in structure between Shallow Fissured and Scaly

at small girths, this is diagnosed in Whitmore (1960 p. 259).

3. Nemesu subgroup (right) with Smooth bark changing

to Scaly (unique Manifestation—a), without a fissured phase,

at less than 1 ft. girth and persisting through life, old trees

sometimes smooth with only scattered scales. The species are:

S . pauciflora, S. kunstleri, S. singkawang, S. macrantha, S.

leptoclados, S. platyclados, S. andulauensis.

4. S . acuminata and S. quadrinervis form a small group

remaining Smooth to about 3 ft. girth then passing through

a short-lived Shallow Fissured phase which by 4 ft. girth

develops into a unique Manifestation (Scaly—b) of Scaly

bark which persists for the rest of life.

5. S. smithiana has a bark of completely different cons¬

truction from any other dipterocarp examined 1 which cannot

be related to any of the seven Types.

The full arguments for including the species in these groups are

given in Whitmore (1960).

Conclusions

Red Meranti is a group heterogeneous in bark but the slash of

all species has a general similarity. There are three main bark sub¬

groups and a fourth small subgroup within which the Bark Type

sequence and Manifestations are the same and S. smithiana does

not fit.

*• Pseudocortex present; expansion tissue in wedges; sheet like rhytidome

layers; sloughing apparently absent. Full diagnosis and illustrations Whit¬

more (1960 pp. 271-2).

358

Vol. XIX. (1962)

Within each subgroup there are variations between individual

species; this is the most variable group studied, 11 out of the 26

examined can be distinguished from each other on gross bark mor¬

phology. In the forest the subgroups and distinctive species are re¬

cognised, the similar slash is insufficient for recognition of the

whole group.

The Malayan species examined show a close correspondence be¬

tween the bark subgroups and Symington’s floral subgroups.

All the species of his Pauciflora subgroup fit my Nemesu sub¬

group (mature trees Scaly) except S. acuminata which must be

considered as a separate subgroup with the closely related Bornean

S. quadrinervis.

Most of the species of his Parvifolia subgroup fit my Tembaga

subgroup (mature trees Shallow Fissured), the exceptions are S.

macrantha and S. singkawang which on the available evidence I

put in Nemesu and S. macroptera in Kawang.

No species occur in Malaya of my Kawang bark subgroup ex¬

cept S. macroptera which is not typical of it.

Thus although wood anatomy does not support Symington’s

floristic subgroups bark characters do. We may predict that when

the genus is revised the Red Merantis will be formally divided

along the lines suggested by Symington for the Malayan species

and confirmed and extended here.

I can comment as follows on a number of outstanding taxo¬

nomic problems of individual Red Meranti species.

1. I suspect (Browne 1955, personal observation (Whit¬

more 1960), Comer observations) two taxa are confused

under the name S. beccariana, one from Sarawak with Scaly

bark with laminae on mature trees and one from Brunei and

North Borneo Smooth on mature trees. The former, if it

develops Scaly bark at small girth, belongs to my Nemesu

bark subgroup, the latter is a Kawang species.

2. Shorea parvifolia is a polymorphic species which

Symington (1943) considered to have several, more or less

distinct forms through its range. He tentatively defined three

forms in Malaya which, however, I found difficult to distin¬

guish in the herbarium and 1 am unable to distinguish be¬

tween them on bark though confirm a known difference in

the bark of the Selangor form.

3. Symington (1943, footnote p. 85) notes that S. par¬

vifolia, S. dasyphylla and S. ovata are very closely related

and liable to be confused where they grow together. They are

however easily distinguished on bark.

359

Gardens Bulletin, S.

4. S. rubra Wood Msc. known from sterile collections

from Brunei has been suspected to be an extreme form of

S. parvifolia, but it has different bark. It is included in my

Tembaga bark subgroup because of this putative relationship

(Whitmore 1960 p. 261).

5. S. acuminata and S. quadrinervis have similar bark and

are distinct from other species.

The taxonomic relationships from their bark of the new Bor¬

nean species examined are:—Kawang subgroup: S. waltoni

Ashton (Msc.), S. sp. now SAR 5798; Tembaga subgroup: S. ru¬

bella Ashton (Msc.), S. rubra Wood (Msc.); Nemesu subgroup:

S. andulauensis Ashton (Msc.).

VATICA

Taxonomy

Vatica Linn, is a clearly defined natural group of some 90 spe¬

cies, closely related to but distinct from Cotylelobium. There are

striking differences between species in fruits and some taxonom¬

ists have been tempted to recognise several genera, but leaf, flower

and wood anatomy are all similar and subdivision is not justified.

Van Slooten (1927) in the most recent monograph recognises

several subgenera but Symington (1943) reduces even these to

artificial sections (cf Dipterocarpus ).

Argument

The Bark Type sequence is shown on Fig. 8. A unique Mani¬

festation, Smooth—k is present on most species. Scaly bark oc¬

curs sometimes on big trees (more commonly than Symington,

1943 suggests); my material was too poor to spot the Manifesta¬

tion (starred on Fig. 8). Dippled bark—a was seen once, differing

in fine structure from Shorea Meranti Damar Hitam also bearing

it. Scroll marks referred to in the published descriptions can deve¬

lop with either Scaly or Dippled bark. Symington describes the

phelloderm as reddish or purplish, this was not seen.

Conclusions

The species are very similar and cannot be distinguished from

each other on bark characters though in favourable conditions the

genus can be recognised. The two ‘subgenera’ covered by the

survey (Isauxis (Arn.) Brandis and Synaptea (Griff.) Brandis) are

indistinguishable, what differences there are in Bark Type se¬

quence between species run across this division, reinforcing

Symington’s suggestion that the ‘subgenera’ are no more than

artificial sections.

360

Vol. XIX. (1962)

Synaptea

Isauxis

Girth in feet

cinerea

cuspidata

maingayi

nitens

oblongifolia

wallichii

\ '

\ s

\ '

\ N

V, '

. V

Cv.,

•a'

■3T

V-

• *

Fig. 8. Vatica, bark sequence through life.

Construction explained in appendix 2; key on Fig. 1.

* V. cuspidata G. 5 ft., V. oblongifolia G. 4 ft. and 5 ft.: see text.

361

Gardens Bulletin, S.

TABLE 1. Taxonomic occurrence of Bark Types and Manifestations on

boles.

SMOOTH BARK

DIPPLED BARK

SHALLOW FIS¬

SURED BARK

DEEP FIS¬

SURED BARK

a *

a Shorea, Meranti

Damar Hitam

and Vatica

a Shorea, Red

Meranti

a Hopea, Fis¬

sured

b **

b Shorea, Red

Meranti

c Shorea, Red

Meranti

c Shorea, Red,

Meranti

d Shorea, Meianti

Damar Hitam

d Shorea, Meranti

Damar Hitam

e Shorea, Meranti

Pa’ang

e Shorea, Meranti

Pa’ang

f Shorea Meranti

Pa’ang

g Anisoptera

h Dipterocarpus

j Hopea, Smooth

and Mata

Kuching

k Vatica

SCALY BARK

SURFACE

ROTTEN BARK

LAMINATE

BARK

a Shorea, Red

Meranti

a Shorea, Meianti

Pa’ang

a Anisoptera

b Shorea, Red

Meranti

b Shorea, Meranti

Pa’ang

b Anisoptera

c Shorea, Balau

c Shorea, Meranti

Pa’ang

d Shorea, Meranti

Damar Hitam

e Dipterocarpus

f Hopea, Scaly and

Balanocarpus

g Dryobalanops

* seedlings and saplings, probably all species groups.

** probably all species groups, mainly yourg plants.

362

Vol. XIX. (1962).

CONCLUSIONS ON BARK TAXONOMY

General

Table 1 shows the occurrence on boles of the seven main Bark

Types; it is seen that three (Deep Fissured, Surface Rotten, Lami¬

nate) are restricted to single species groups, a fourth, Shallow Fis¬

sured, to Shorea, and the other three are found in more than one

species group but that with very few exceptions the Bark Mani¬

festations are restricted.

Hence, as already noted, it is at the level of the Manifestations

that most of the differences between species groups and species

occur and so at this level that there are differences in bark useful

for identifying trees.

Only Manifestation Smooth—a. Smooth—b, Dippled—a Scaly

—f occur in more than one species group.

1 have discovered that the fine details of secondary phloem struc¬

ture (except Scaly Hopea, q.v.), and often the structure of the

expansion tissue, phloem proliferation tissue and periderms also,

are constant within a species group so that all the Manifestations

of every species group are alike in these fine details. This applies

to Dippled—a and Scaly—f and to some extent to Smooth—b

barks too; these Manifestations occur in more than one species

group but with microscopic differences.

Smooth—a bark is found on all seedlings, saplings and small

twigs, and has not been fully studied, Smooth—b bark is mainly

restricted to small members. The tangential strains in these parts

are highest (see Whitmore 1962b) and the resulting extensive ex¬

pansion tissue obscures the differences between species groups.

Formal taxonomy

The species have been studied in the most natural groups yet

recognised, which are those proposed by Symington (1943) but

not all validly published. This survey has shown differences in

bark between the groups and similarities within them. The taxo¬

nomic groups and the bark groups are identical suggesting that

these are indeed the natural groups of Dipterocarpaceae. I do not

consider that the differences 1 have detected between groups are

the result of inadequate sampling (between 2/5 and 1/10 of the

species of the groups) but that 1 have sampled enough species for

my conclusions to have this taxonomic application. The danger

of arguing in a circle (from studying taxonomic groups, finding

Manifestations are restricted to them, then arguing that these must

therefore be the natural groups) is obviated by the caution exer¬

cised in creating Manifestations, any incompletely understood

bark has been described as a Variant of a described Manifestation .

363

Gardens Bulletin, S.

Within the species groups variation is at two levels, between

taxonomic subgroups and between individual species, and is in

the girth at which the bark changes and in the Manifestations or

Variants present. The conclusions 1 draw have already been de¬

tailed with the notes on each species group. In summary I find

that I agree on the whole with Symington’s conclusions on the

taxonomic nature of the groups. I find bark differences accen¬

tuate the taxonomic divisions of Anisoptera and Shorea, and with¬

in Shorea, where I agree on the whole with his subdivisions of Red

Meranti and extend them, and find the other divisions distinct

though of varying homogeneity. I agree with him that Diptero-

carpus and Vatica are unnaturally subdivided, that Balanocarpus

should not be kept separate from Hopea and within Hopea itself

I find his ‘tentative arbitrary field groups’ correlate better with

bark characters than the taxonomic subdivisions. Dryobalanops

I confirm is a homogeneous natural genus. The groups of Shorea

and some of those of Hopea are as distinct as the other genera on

their bark, but one needs to consider carefully the purpose of one’s

taxonomy before creating new genera to accommodate these

differences.

Concerning individual species:

1. Ten new undcscribed species have been examined and

their relationships discovered from their bark (see Hopea and

Shorea, Red Meranti, Meranti Damar Hitam and Balau).

2. I can state, about known problems of taxonomy, that:

(i) Dipterocarpus fagineus and D. pseudofagineus appear

neither more nor less similar than other Diptero¬

carpus species, although they are taxonomically

very close related.

(ii) The closely related Shorea parvifolia, S . ovata and S.

curtisii (Red Meranti) are clearly distinct.

(iii) Shorea rubra is distinct from the closely related S.

parvifolia.

(iv) Shorea albida is a White Meranti despite its timber

classification as Red Meranti or Balau; the two

tentative taxonomic forms are indistinguishable.

3. Geographical Variation.

(i) Infraspecific.

(a) Two taxa of different distribution seem to be

confused under the epithet Shorea becca -

riana (Red Meranti).

( b ) Of the three putative Malayan forms of S. par¬

vifolia only one has slightly different bark

from the others.

364

Vol . XIX. (1962)

and Bornean trees of Hopea beccaricma

(Fissured Hopeas).

(ii) Interspecific. It has been suggested (Richards, 1952,

p. 58) that species restricted to tropical Rain forest

often have thinner bark than species from drier

forests. In this survey, however, no systematic dif¬

ferences have been noticed between dipterocarp

species of Burma, Indochina, Thailand and extreme

north-west Malaya—the Burmese floristic element

—which occur in dry, deciduous, dipterocarp

forests and fire climax Schima -bamboo forests

(Symington 1943) 1 and species of more easterly dis¬

tribution restricted to the lowland, everwet, ever¬

green, Rain forests of Malaya, Sumatra and Bor¬

neo. A few species occur in both these floristic

divisions (e.g. Dipterocarpus kerrii). If there is a

difference in bark between members of these floris¬

tic elements it is infraspecific or phenotypic and is

not universal; there are bark differences between

exposed and forest living trees of the same species

already described above.

These conclusions on the formal taxonomy of the species and

species groups of Dipterocarpaceae will have to be weighed with

the other evidence when the family is revised, a task which, as

in Symington’s day, still awaits the full collection and description

of the Bornean dipterocarps.

Forest Recognition

Bark Types cannot be confused; they are quite different in

details of bark surface pattern and slash and usually in impres¬

sion from a distance. Bark Manifestations are by definition distinct

in the forest. Thus it follows from the occurrence of Bark Types

and Manifestations (Table 1) that nearly all the species groups

are distinct in the forest in the bark present on mature boles and

sometimes from pole bark too.

The exceptions are:

Scaly—f, found on Balanocarpus and the Scaly Barked

field group of Hopea , but these taxa should perhaps be

lumped.

1* These species are Sf\orea hypochra, S. sericeiflora, S. talura (Meranti

Pa’ang), Dipterocarpus dyeri, and Hopea ferrea, H. helferi, and H. odorata

(Scaly Barked Hopea).

365

Gardens Bulletin, S.

Dippled bark: Shorea , Meranti Damar Hitam and Scaly

Barked Hopea —but small trees only as this is a transient

Manifestation.

The homogeneous species groups have a distinctive bark ges¬

talt from which they can be recognised in the forest. In hetero¬

geneous groups the subgroups or the individual species are recog¬

nised separately. In this survey these gestalts have been analysed

more completely than previously, in all out of the 103 species

examined mature trees of 21 species can be recognised from bark

alone and many of the others can be identified down to a small

group of species (of recognised taxonomic status or not) within

their group. Not many species have been added to the list which

Symington (1943) and Wood (1957) distinguish but the descrip¬

tions are fuller; a few differences which do not seem to hold are

contradicted.

Clear differences are stated between the Scaly bark of Shorea

Balau, Dipterocarpus, Scaly Hopea plus Balanocarpus and Dryo-

balanops which have sometimes been confused in the forest

(Symington, 1943); the reasons for the similarity of these Manifes¬

tations were analysed in Whitmore (1962a).

New distinctions are made and old confusions are sorted out

because of the critical descriptive scheme and glossary adopted

(Whitmore 1960 part 3). The Dipterocarpaceae because of their

paramount importance are better known in the forests of Malaya

and Borneo than any other family, there are no works compar¬

able to Symington (1943) and Wood (1957) for any other family

(except perhaps Henderson and Wyatt-Smith, 1956, on Calo-

phyllum (Myrtaceae )) and it is therefore likely that the scheme

and glossary developed for this survey would extend the value of

bark for forest recognition very considerably were it applied to

other families of trees.

ACKNOWLEDGEMENTS

Primarily I should like to thank Mr. E. J. H. Corner who first

interested me in tropical botany and who suggested and supervised

this study. Special thanks are also due to Professor H. B. Gilliland

(Singapore) and Dr. K. R. Sporne (Cambridge) who supervised

me part of the time and to the Conservators of Forests and their

staffs in Malaya and British Borneo for stimulating discussion and

considerable help in the forests. The investigations were financed

by grants from St. John’s College, Cambridge, University of Cam¬

bridge (Frank Smart Studentship), Colonial Development and

Welfare funds (grant CDW(R)806) and the Ministry of Education

(State Studentship), and were conducted in the Botany Schools of

Cambridge and Singapore and the forests of Malaya and Borneo.

366

Vol. XIX. (1962)

REFERENCES

Bate-Smith, E. C. and Whitmore, T. C. (1959) Chemistry and taxonomy

in the Diptcrocarpaceae. Nature, Lond. 184: 795-796.

Brandis, D. (1895) An enumeration of the Dipterocarpaceae. J. Linn. Soc.

(Bot.) 31: 1-148.

Browne, F. G. (1955) Forest trees of Sarawak and Brunei. Kuching.

Desch, H. E. (1936) Commercial timbers of the Malay Peninsula: I The

genus Shorea. Malay For. Rec. 12.

Desch, H. E. (1941) Dipterocarp timbers of the Malay Peninsula. Malay.

For. Rec. 14.

Foxworthy, F. W. (1927) Commercial timber trees of the Malay Penin¬

sula. Malay For. Rec. 3.

Foxworthy, F. W. (1932) Dipterocarpaceae of the Malay Peninsula.

Malay. For. Rec. 10.

Heim, F. (1892) Recherches sur les Dipterocarpacees.

Henderson, M. R. & Wyatt-Smith, J. (1956) Calophyllum Linn. Gdns/

Bull. 15: 285-377.

Richards, P. W. (1952) The tropical rain forest. Cambridge.

Slooten, D. F. van (1927) The Dipterocarpaceae of the Dutch East

Indies: IV The genus Vatica. Bull. Jard. bot. Buitenz.

Seric 3: 9: 66-137.

Symington, C. F. (1933) Notes on Malayan Dipterocarpaceae: I. Gdns/

Bull. 7: 129-159.

Symington, C. F. (1934) Notes on Malayan Dipterocarpaceae: II. Gdns/

Bull. 8: 1-40.

Symington, C. F. (1938) Notes on Malayan Dipterocarpaceae: IV. Gdns/

Bull. 9: 319-354.

Symington, C. F. (1939) Notes on Malayan Dipterocarpaceae: V. Gdns/

Bull. 10: 336-386.

Symington, C. F. (1943) Foresters’ Manual of Dipterocarps. Malay. For.

Rec. 16 Singapore, (published by the Japanese as ‘Syonan

(2603)’).

Whitmore, T. C. (1960) The systematic morphology of bark in the Dip¬

terocarpaceae. Dissertation for degree of Doctor of Philo¬

sophy, Cambridge University. (Copies in University Library

and Botany School, Cambridge, England, and Forest Re¬

search Institute, Kepong, Malaya).

Whitmore, T. C. (1962a) Studies in systematic bark morphology: I Bark

Morphology in Dipterocarpaceae. New Phytol. 61: 191-207.

Whitmore, T. C. (1962b) Studies in systematic bark morphology: II

General features of bark construction in Dipterocarpaceae.

New Phytol. 61: 208-220.

Wood, G. H. S. (1957) Common dipterocarp trees of North Borneo. Un¬

published manuscript of North Borneo For. Rec. 5. Privately

circulated from Sandakan to far eastern Forest Departments.

Wyatt-Smith, J. (1955) The Dipterocarpaceae of Brunei, North Borneo

and Sarawak: I Anisoptera. Malay. Forester 18: 70-79.

367

APPENDIX 1

Alphabetical list of species studied

The groups of Shorea are indicated as R (Red Meranti), D (Meranti

Damar Hitam), P (Meranti Pa’ang) and B (Balau). Numbers refer to my

own series, the location of the herbarium specimens is given in the text.

Anisoptera Korth.

A. curtisii Dyer

A. laevis Ridl.

A. megistocarpa V.S1.

A. oblonga Dyer

A. scaphula (Roxb.) Pierre

Balanocarpus Bedd.

B. heimii King.

Dipterocarpus Gaertn.f.

D. appendiculatus Scheff.

D. cornutus Dyer

D. costulatus V.S1.

D. crinitus Dyer

D. dyerj Pierre

D. fagineus Vesq.

D. kerrii King

D. pseudofagineus Foxw.

D. rotundifolius Foxw.

D. sublamellatus Foxw.

D. verrucosus Foxw. ex V.S1.

Dryobalanops Gaertn.f.

D. aromatica Gaertn.f.

D. lanceolata Burck

D. oblongifolia Dyer

D. rappa Becc.

Hopea Roxb.

H. beccariana Burck

H. ferrea Lanessan

H. griffithii Kurz.

H. helferi (Dyer) Brandis.

H. latifolia Sym.

H. mengarawan Miq.

H. nervosa King

H. nutans Ridl.

H. odorata Roxb.

H. sangal Korth.

H. semicuneata Sym.

H. sulcata Sym.

H. vesquei Heim

H. sp. nov. ‘A’.

H. sp. nov. 324.

Shorea Roxb.

S. acuminata Dyer (R).

S. albida Sym. (P).

S. andulauensis Ashton (Msc.) (R.)

S. argentifolia Wood (Msc.) (R).

S. balanocarpoides Sym. (D).

S. beccariana Burck (R).

S. bentongensis Foxw. (P).

S. blumutensis Foxw. (D).

S. bracteolata Dyer (P).

S. ciliata King (B).

S. collina Ridl. (B).

S. curtisii Dyer ex King (R).

S. dasyphylla Foxw. (R).

S. dealbata Foxw. (P).

S. dolichocarpa V.S1. (D).

S. sp. nov. ‘E4* (D).

S.? elliptica Burck (B).

S. faguetiana Heim. (D).

S. foxworthyi Sym. (B).

S. geniculata Sym. (B).

S. gibbosa Brandis (D).

368

Vol XIX. (1962).

S. glauca King (B).

S. guiso (Blanco) Bl. (B).

S. gysbertsiana Burck (R).

S. hopeifolia (Heim) Sym. (D).

S. hypochra Hance (P).

S. inappendiculata Burck (B).

S. kunstleri King (R).

S. laevifolia (Parijs) Endert (B).

S. laevis Ridl (B).

S. lamellata Foxw. (P).

S. leprosula Miq. (R).

S. leptoclados Sym. (R).

S. lumutensis Sym. (B).

S. macrantha Brandis (R).

S. macroptera Dyer (R).

S. materialis Ridl. (B).

S. maxima (King) Sym. (D).

S. maxwelliana King (B).

S. meadiana Sym. (B).

S. mecistopteryx Ridl. (R).

S. multiflora (Burck) Sym. (D).

S. ochracea Sym. (P).

S. ochrophloia E. J. Strugnell (B).

ex Sym. (R).

S. ovalis (Korth) Bl. (R).

S. ovata Dyer (R).

S. parvifolia Dyer (R).

S. pauciflora King (R).

S. platycarpa Heim. (R).

S. platyclados V.S1. ex Foxw. (R).

S. pinanga Scheff. (R).

S. quadrinervis V.S1. (R).

S. resina-nigra Foxw. (D).

S. resinosa Foxw. (P).

S. rubella Ashton (Msc.) (R).

S. rubra Wood (Msc.) (R).

S. sericeiflora Fischer et Hutch. (P).

S. singkawang (Miq.) Burck. (R).

S. smithiana Sym. (R).

S. symingtoniana Wood (Msc.) (P).

S. talura Roxb. (P).

S. waltoni Ashton (Msc.) (R).

S. sp. nov. SAR 5798 (R).

S. sp. nov. 555 (B).

S. sp. nov. 569 (B).

Vatica Linn.

V. cinerea King.

V. cuspidata (Ridl.) Sym.

V. lowii King emend. Sym.

V. maingayi Dyer

V. nitens King

V. oblongifolia Hk.f.

V. wallichii Dyer

369

APPENDIX 2

Note on construction of figures 1-8 showing relation between

girth and Bark Type on the bole

These figures show at a glance the Bark Type sequence on the bole

throughout the life of the tree, the number of individuals sampled and

their girths. There is a column for every species and these are arranged

across the figure from left to right; increasing girth is down the page.

The Bark Types present over the whole girth range are shown by differ¬

ent shadings, as far as they can be reasonably induced from the data.

Every individual sampled is shown by a dot and its Bark Manifestation

indicated where determined. Starred individuals are explained in the legends.

No Manifestation is shown when an individual is heterogeneous, beyond

the normal range of its Bark Type, or undetermined; the reader must refer

to Whitmore (1960) for detailed discussion.

The Bark Type sequence of a species and the Manifestation of an indi¬

vidual are discussed more fully in Whitmore (1960) where necessary; it is

not possible to reproduce these details here.

The Bark Type sequence has been induced from the published general¬

ised descriptions and my own observations. These two sources are comple¬

mentary; in most cases they indicate identical bark at every girth. In some

cases accurate personal observations make it possible to define an otherwise

vague published description. In the absence of personal observations I can¬

not always interpret the published descriptions with certainty as the terms

used are not defined.

The following rules have been followed in inducing the Bark Type

sequence throughout life:—

1. There are often considerable differences from one individual to

another in the girth at which the Bark Type changes; this accounts for

the occasional discrepancies between the generalised description of the

species and the description of an individual tree. On the figures the

bark is shown to change Type at a median girth between the extremes

observed; where the range is considerable an intermediate zone is

shown.

2. It is unlikely that Scaly, Surface Rotten and Laminate barks are

succeeded on very big trees by any others and, although the published

generalised description may not describe the bark of very big trees and

no personal observations were made, they are extrapolated to the

maximum usual girth of the species. It is possible and likely, however,

that the Bark Manifestation changes, first on the butt, later spreading

up the bole as the tree ages. Other Bark Types are extrapolated up¬

wards an arbitrary 1 ft. above the largest individual observed (see 4.

below), unless the published generalised description is sufficiently pre¬

cise to warrant greater extrapolation.

3. The Bark Type of poles (individuals below about 3$ ft. girth)

can only be induced from my own observations as the published gene¬

ralised descriptions are restricted to mature trees. In many cases no

observations were made, but the bark is likely to remain Smooth to

at least 1 ft. girth and is so shown on the figures. The girth at which

Smooth bark is replaced is usually somewhere in the girth range of

large poles (girth range 1 ft.-3i ft. for emergent species). No Bark

Type is shown for large poles or small trees unless individuals were

seen.

370

Vol. XIX. (1962).

4. I have extrapolated the Bark Type of the smallest tree examined

downwards an arbitrary 1 ft. in girth, where the Bark Type on small

individuals is in doubt. This allows for an error in estimation of the

girth (not likely to be greater than + 9 ins.) and for the variation

expected between individuals (see above).

5. No Bark Type precedes Smooth so no downward limit is fixed

for the extrapolation from the smallest Smooth barked tree seen.

6. The maximum usual girth, calculated from the published data,

is shown on the figures where possible, by an horizontal line (this is

considerably less than the maximum recorded girth); the bark present

above this girth is only shown for species where individuals were seen.

371

C d

Fig. 9. Examples of Bark Manifestations.

(a) Smooth -h ( Dipterocarpus) , note surface fine sculptur-

ings; ( b ) Scaly -e ( Dryobalanops ); (c) Dippled -a

{Shorea, Meranti Damar Hitam); (d) Coarsely

Dippled hark of Agathis alba (Lam.) Foxw.

Fig. 10. Examples of Bark Manifestations.

(a) Shallow Fissured -c ( Shorea , Red Meranti); (/>) Deep

Fissured -a ( Hopea , Fissured Barked); (c) Surface

Rotten -a ( Shorea , Meranti Pa’ang), note fibrous

fissured surface; ( d ) Laminate -a ( Anisoptera,

Pilosae), note scaly-fissured surface and laminate

slash.

THE

GARDENS’ BULLETIN

SINGAPORE

Vol. XIX, Part III ^ IOth December, 1962

CONTENTS

Page

Anne Johnson: On a small collection of Bryophytes collected by

Professor H. B. Gilliland in Eastern New Guinea 373

Annf. Johnson: A Short Note on some Soil Algae from New Guinea 375

Anne Johnson: Precursory Studies on the Epiterranean Soil Algae of

Singapore and Malaya . . .379

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

Addendum II . . . , 385

To be purchased at the Botanic Gardens, Singapore

Price: $3

Published by Authority

PRINTED BV I.IM BIAN HAN, ACTING GOVERNMENT PRINTER, SINGAPORE

1962

On a small collection of Bryophytes collected by

Professor H. B. Gilliland in Eastern New Guinea

By Anne Johnson

Small collections of mosses from West New Guinea have been

recorded by Dixon, Brotherus, Fleischer and Reimers; while a

large collection from Mt. Wilhelmina was made by Brass and

Myer-Drees and recorded by Bartram (1942). A small collection

of liverworts was collected by Takari Tuyama in Vogelkop in 1943

and was recorded by Hattori (1951). Records of an extensive col¬

lection of mosses from Eastern Papua were made by Bartram

(1957). Records from the Territory of New Guinea of mosses

collected by the Count and Countess Nils and Greta Gyldenstolpe

in the Mt. Hagen district were given by Bartram (1953). Further

collections in the Highlands of Eastern New Guinea were made

by Hoogland in 1953 and Robbins in 1957. These collections

were recorded by Bartram (1959).

The bryophyte flora of New Guinea is immensely rich and very

imperfectly known (Bartram, 1959). Although it has a consider¬

able affinity with the flora of the rest of Malaysia, there are a

large number of endemic species; while alpine and sub-alpine

species show a close relationship with the flora of Australia, Tas¬

mania and New Zealand.

Professor Gilliland’s collection was made in September 1960 in

two places:— Chimbu and the Daulo Pass (6,000-8,170 ft.) Aus¬

tralian Territory of New Guinea. The numbers assigned to the

bryophytes are accession numbers of the Bryophyte Herbarium of

the University of Singapore.

Hepaticae

Marchantia polymorpha L.—Daulo Pass, no. 687.

Marchantia geminata Nees.—Daulo Pass, nos. 703, 710; “forma

typica”, with female heads bearing exceedingly long purple

hairs, ventral scales similar to “Tjibodas type” (Burgeff, 1943).

Riccardia pinguis (L.) Gray—Chimbu, no. 696.

Nardia hasskarliana (N. ab Es.) O. Lind.—Daulo Pass, no. 693.

Plagiochlia teysmanni Sande Lac.—Daulo Pass, nos. 714, 711.

Eulejeunea Spr. sp. —Daulo Pass, no. 702, a slender species,

hyaline papilla proximal to apical tooth, the lobes several

times larger than the lobules.

Musci

Fissidens zollingeri Montg.—Chimbu, no. 700.

Campylopodium euphorocladium (C.M.) Besch.—Daulo Pass

no. 706.

Campylopus richardii Brid.—Daulo Pass, nos. 704, 713.

373

Gardens Bulletin, S.

Thysanomitrium comosum Hsch. et Rw.—Daulo Pass, no. 691.

Leucobryum subsanctum Broth.—Daulo Pass, nos. 701, 712.

Barbula consanguinea (Thw. et Mitt.) Jb.—Chimbu, no. 697.

Mniomalia semilimbata (Mitt.) C. Mull.—Daulo Pass, no. 694.

Symphysodon splendens (Hsch. et Rw.) Broth.—Daulo Pass,

no. 709.

Neckeriopsis gracilenta (Lac.) Fleisch.—Chimbu, no. 698.

Ectropothecium aurifolium Bartr.—Daulo Pass, no. 707.

Ectropothecium falciforme (Dz. et Mb.) Jaeg.—Daulo Pass,

no. 690.

Ectropothecium sp.—Chimbu, no. 708.

Isopterygium albescens (Schwaeg.) Jaeg.—Daulo Pass, no. 705.

Pogonatum junghuhnianum (Dz. et Mb.) v.d.B.—Daulo Pass,

no. 688.

Pogonatum microphyllum (Dz. et Mb.) v.d.B. et Lac.—Chimbu,

no. 695; Daulo Pass, no. 692.

Of the species listed above, Marchantia polymorpha and Riccardia

pinguis are cosmopolitan, while Thysanomitrium comosum Hsch.

et Rw. has a general tropical distribution. Ectropothecium auri¬

folium is endemic being previously recorded from Lake Habbema

in West New Guinea (Bartram, 1942). The majority of the rest of

the species are Malaysian in distribution some extending as far as

the Pacific Islands ( Fissidens zollingeri, Campylopodium euphoro-

cladium , and Mniomalia semilimbata ), Japan ( Campylopus

blumii) or East Africa ( Campylopodium euphorocladium). The

alpine and sub-alpine element is not represented, since no collec¬

tion was made above 8,170 ft.

Relevant Papers

Arens, P. (1953) Bemerkungen zu einigen von C. E. Carr in Neuguinea

gesamelten Moosen. Blumea 7 (2) 339-341.

Bartram, E. B. (1942) Third Archbold Expedition Mosses from the Snow

Mountains, Netherlands New Guinea. Lloydia 5 245-292.

Bartram, E. B. (1953) Additional mosses from N.E. New Guinea.

Svensk. Bot. Tidskr. 47 (3) 397-401.

Bartram, E. B. (1957) Mosses of Eastern Papua, New Guinea. Brittonia

9 (1) 32-56.

Bartram, E. B. (1959) Contributions to the mosses of the highlands of

Eastern New Guinea. Brittonia 11 (2) 86-98.

Burgeff, H. (1943) Gcnctische Studien an Marchatia, p. 282, Jena.

Dixon, H. N. (1937) On a small collection of mosses from New Guinea.

Annales Bryologici 10 16-19.

Fleischer, M. (1900-1922) Die Musci der Flora von Buitenzorg, vols. 1-4,

Leiden.

Hattori, S. (1951) On a small collection of Hepaticae from Dutch New

Guinea. Bot. Mag. (Tokyo) 64 112-119.

Meijer, W. (1959) Notes on species of Riccardia from their type localities

in Western Java. Jour. Hattori Bot. Lab. 21 61-78.

Noguchi, A. (1953) Mosses of Mt. Sarawatet, New Guinea. Jour. Hattori

Bot. Lab. 10 1-23.

Schiffner, V. (1900) Die Hepaticae der Flora von Buitenzorg. vol. 1,

Leiden.

Van der Wijk, R. ( 1 958) Precursory Studies on Malaysian Mosses. II. A

Preliminary Key to the Moss Genera. Blumea 9 (1) 143-186.

374

A Short Note on some Soil Algae from New Guinea

By Anne Johnson

Small samples of soil from the Australian Territory of New

Guinea were received together with bryophytes collected by Pro¬

fessor H. B. Gilliland in 1960. These samples were kept in poly¬

thene bags and, owing to transport difficulties, did not reach this

laboratory until one year after their dispatch. The soil was care¬

fully separated from the bryophytes which have been described in

a previous paper. Two sets of soil cultures were set up:—

(i) moist cultures, i.e. samples of soil placed in sterile petri-

dishes and exposed to continuous light;

(ii) liquid cultures in nutritive medium, following the method

of John (1942).

In both types of culture there was vigorous growth of algae and

there was considerable correspondence between algae which ap¬

peared in moist and liquid culture of the same sample.

Four samples were obtained from localities on the Daulo Pass,

6,000-8,000 ft.; while a fifth was obtained from Chimbu. In all

cases the soil was collected from 1-2 cm. of the surface.

Chlorophyceae

CWorococcales

Chlorococcum htimicolum (Naeg.) Rabenh.

Liquid culture, soil from Chimbu.

Seenedesmus obliquus (Turp.) Kutz

Liquid culture, soil from Chimbu.

Dactylococcus stage of Seenedesmus obliquus (Turp.) Kutz. (see

Grintzesco (1902))

Liquid culture, soil from Daulo Pass, 6,000-8,000 ft.

Cyanophyceae

Chroococcales

Chroococcus varius A. Br.

Liquid culture, soil from Daulo Pass, 6,000-8,000 ft.

Chamaesiphonales

Dcrmocarpa hemisphaerica Setchell & Gardner

Moist culture, epiphytic on Lyngbya mesotrichia Skuja from

Daulo Pass, 6,000-8,000 ft. *

Nostocales

Lyngbya mesotrichia Skuja

Moist culture and liquid culture from Daulo Pass, 6,000-8,000

ft.

Phormidium jadinianum Gomont

Moist and liquid culture from Daulo Pass, 6,000-8,000 ft.

375

Gardens Bulletin, S.

Symploca elegans Kutzing ex Gomont

Moist culture, soil from Chimbu.

Nostoc linckia (Roth) Bornet ex Born, et Flah.

Moist and liquid culture, soil from Daulo Pass, 6,000-8,000 ft.

Nostoc microscopicum Carm. ex Born, et Flah.

Moist culture, soil from Daulo Pass, 6,000—8,000 ft.

Scytonema pseudopiinctatum Skuja

Moist culture, soil from Daulo Pass, 6,000-8,000 ft.

Discussion

The pH of the samples of soil from the Daulo Pass and Chimbu

was about 5.3 when they reached the laboratory. The pH of the

nutritive solution used in the liquid culture was 5.6. There was no

change in the pH of the moist culture over a two month growing

period. Most of the liquid cultures showed a gradual rise of pH

to 6.2 in some cases. In a few liquid cultures with sparse growth

there was little change in pH.

While cultures of soil algae are of little direct value from an

ecological point of view, they are almost essential in obtaining a

complete algal flora of a soil (Tiffany, 1951). In the soil, algae are

so very small and widely dispersed they are easily overlooked. In

the moist cultures, samples of soil were given additional light

over 24 hours/day which led to enhancement of algal growth;

while in liquid cultures additional nutrients were also supplied.

These cultures tell us little of the relative abundance of the differ¬

ent algae in the natural habitat, but they do indicate which species

of algae are present in these particular soils.

The results given above indicate the occurrence of eight species

of blue-green algae but only two species of Chlorophyceae. No

diatoms were recorded. This may be due to the scarcity of diatoms

in the habitat, or to the fact that the method of culture is unsuit¬

able for diatom growth. Their complete absent even from moist

cultures suggests the former. Lund (1945) has indicated that dia¬

toms are absent in acid soils.

The relative large numbers of species of Cyanophyceae in tro¬

pical soils was noted for Ceylon by Fritsch (1907, 1907*) and in

Lahore and Simla by Ghose (1923). This was considered to be

due to their preference for high temperatures, damp atmosphere.

Their blue-colour may act as a screen against intense light while

lowland forms are able to withstand desiccation. Highly mucilagin¬

ous blue-green algae (e.g. Nostoc spp.) are most suited to damper

upland conditions. Fritsch (1907) found slimy forms at Nuwara

Eliya and Hakgalla at a height of 6,000 ft. and more.

376

Vol. XIX. (1962).

References

Fritsch, F. E. (1907) The Role of Algal Growth in the Colonisation of

New Ground and in Determination of Scenery. Georgr. Jour.

30 531-548.

Fritsch, F. E. (1907*) A general consideration of the sub-aerial and

Freshwater Algal Flora of Ceylon. Proc. Roy. Soc. (Lond.)

B 79 197-254.

Ghose, S. L. (1923) A Systematic and Ecological Account of a Collection

of Blue-Green Algae from Lahore and .Simla. Jour. Linn.

.Soc. Bot. 46 333-346.

Grintzesco, J. (1902) Recherches experimentales sur la morphologie et

physiologic de Scenedesmus acutus Meyen, Bull. Herb. Bois-

sier 11, 2 217 et seq.

John, R. P. (1942) An Ecological and Taxonomic Study of the Algae of

British Soils. I The distribution of surface-growing algae.

Annals Bot., N.S. 6 323-349.

Lund, J. W. G. ( 1945) Observations on soil algae. New Phytol. 44 196-

219.

Tiffany, L. H. (1951) Ecology of Freshwater Algae. Chronica Botanica

27 303-305.

377

Precursory Studies on the Epiterranean Soil Algae of

Singapore and Malaya

By Anne Johnson

Introduction

Algae are present in Malayan soils to a depth of several centi¬

meters or sometimes to one meter. Deeply buried subterranean

forms probably have their origin from the soil surface having been

washed down by the rain, aided by the movement of soil animals

and by cultivation. These algae are undoubtedly in complete dark¬

ness and must survive as saprophytes (Tiffany, 1951). In general

they are of little importance in soil economy. On the other hand,

the truly epiterranean forms which remain at or near the soil sur¬

face are exceedingly important since they can increase the organic

content of the surface layers of the soil by photosynthesis, while

certain Nostocaceae can contribute to the fertility of the soil by

nitrogen fixation (De, 1939). Soil algae are exceedingly important

ecologically particularly in tropical regions as first colonisers of

bare ground. Many algae assist in the actual disintegration of a

rock surface to form soil, in addition to forming an organic matrix

suitable for higher plant growth. (Fritsch, 1907; Treub, 1888).

Investigation of surface algae in Europe include those of Petersen

(1935); Bristol (1920); James (1935) and John (1942). In Ame¬

rica Coyle (1935); Booth (1941); Lowe & Moyse (1934); Moore

& Carter (1926) and many others have made studies of the soil

flora of different districts. Valuable contributions to our know¬

ledge of the soil algae of Ceylon were made by Fritsch (1907,

1907 a, 1907 b). Ghose gives an account of the Cyanophyceae of

Lahore and Simla (1923). As far as I am aware the only paper on

Malayan soil algae is that of Bristol (1919).

II. The Soil Samples and their mode of collection and culture

The technique of collection and culture was very similar to that

used by John (1942), but various details were modified to suit

local conditions.

Soil samples were taken from the top, second, third and fourth

centimeter below the surface of the soil with the usual precautions

to prevent contamination or mixing of the layers. Each sample

was examined microscopically and any algae were recorded. For

each soil sample two series of cultures were set up. Moist cultures

consisted of soil placed to a depth of 1 cm. in sterile petri dishes,

moistened with sterile distilled water, and exposed to continuous

light. Liquid cultures consisted of small quantities of soil introduced

379

Gardens Bulletin, S.

into a nutritive solution in conical flasks under sterile conditions.

The nutritive solution used was Knop’s solution (Pringheim, 1951,

p. 349) at either full- or half-strength. Liquid cultures were also

exposed to continuous light supplied by flourescent strips of the

warm daylight type.

Sub-cultures were made (i) by re-inoculating fresh Knop’s solu¬

tion in flasks, (ii) by plating on to Knop’s agar or peptone agar,

(iii) by inoculation with a sterile needle on to agar slopes in tubes.

III. General description of soils examined

Soil samples were taken from the following six localities:—

Locality 1. Hard exposed ground, University of Singapore.

This soil consisted of dark brown excrusting surface layer

about 2 mm. thick, beneath which was a red clay soil about

3 cm. deep. At 3-4 cm. depth the soil was lighter in colour.

The pH of all soil samples was approximately 6.4.

Locality 2. Exposed silt, 300 yards from the sea, Dungun,

Trengganu.

This soil is a medium brown silt with a fair amount of

organic content. The first 4 cm. are more or less homogene¬

ous, apart from a very narrow surface layer. The pH of all

soil samples was approximately 6.1.

Locality 3. Exposed cleared building site, Cluny Road, Singa¬

pore.

The first three centimeters were of a medium brown colour

and consisted of about 34 per cent silt and 62 per cent clay.

Beneath this was a heavier red soil with a slight increase in the

silt content. The pH of all soil samples was approximately 6.3.

Locality 4. Botanic Gardens Jungle, in forest with heavy leaf

litter.

The first centimeter consisted of a dark brown layer with

abundant humus, tree roots and dead leaves. The second cen¬

timeter consisted of a lighter zone but still with many tree

roots. Beneath this was a pale beige crumbly clay soil with

fewer roots. The pH of all soil samples was approximately 6.3.

Locality 5. Sunlit Path, MacRitchie Reservoir.

The soil was a homogeneous sticky clay of a pale gamboge

colour. The pH of all soil samples was approximately 5.3.

Locality 6. MacRitchie Reservoir Forest, under heavy leaf

litter.

The soil was a dark clay rich in tree roots and organic

matter. The pH of all soil samples was approximately 5.7.

380

Vol. XIX. (1962).

IV. Examination of initial samples

In soil taken from the natural habitat, soil algae are often sparse,

frequently only spores or more or less dormant cells are available;

and they are easily overlooked when present in small numbers in

a large mass of soil. (Tiffany, 1951). In soil from five of the above

localities no algae were observed in any of the top four centi¬

meters. However locality 1 showed an abundant growth of surface

algae in the field consisting of Scytonema javanicum (Kurtz) Born.,

together with Gloeotheca sp., Oscillatoria sp. and a little Micro¬

coleus vaginatus (Vouch.) Gomont. The second centimeter showed

a few coccoids and minute flagellates while no algae could be

found in the lower soil layers.

V. Examination of moist cultures

Moist cultures are the best imitation of natural conditions (John,

1942). The growth of soil algae is encouraged by exposure to light

over a twenty-four hour instead of a twelve hour period. After a

month or two the growth of soil algae on the surface becomes ap¬

parent. Later in some cases fern prothalli or seedlings of higher

plants make their appearance. The development of a surface layer

of algae is slower in soil from deeper layers.

The following is a list of algae which grew in the moist cultures

over a five month period. The number in brackets indicates the

mean depth from which the soil sample was taken.

Chlorophyceae

Pleurococcus naegelii Chodat

Sunlit path, MacRitchie Reservoir (2.5 cm.).

Trentepohlia forulosa De YVildemann

Sunlit Path, MacRitchie Reservoir (3.5).

Vaucheria sessilis (Vauch.) D.C.

Botanic Gardens Jungle (0.5).

Xanthophyceae

Tribonema bombycinum (Ag.) Derb. et Sol.

Sunlit Path, MacRitchie Reservoir (0.5).

Cyanophyccae

Chroococcus varius A.Br.

Sunlit Path, MacRitchie Reservoir (3.5).

Chroococcus sp.

University of Singapore (0.5).

Lyngbya sp.

Sunlit Path, MacRitchie Reservoir (0.5) and (3.5).

Lyngbya mesotrichia Skuja

Expose silt, Dungun (2.5).

Symploca elegans Kiitz.

Exposed cleared building site, Singapore (1.5).

381

Gardens Bulletin, S.

Microcoleus vaginatus (Vauch.) Gomont

Exposed silt, Dungun (1.5); University of Singapore (0.5).

Aulosira pseudoramosa Bharadw.

University of Singapore (0.5).

Scytonema javanicum (Kiitz.) Born.

University of Singapore (0.5), (1.5), (2.5); Exposed cleared

building site (3.5).

Tolypothrix fragilis (Gardn.) Geitler

University of Singapore (3.5).

Tolypothrix arenophila West, W. & GJS.

Exposed silt, Dungun (0.5).

In these cultures ten different species of Cyanophyceae, three

species of Chlorophyceae and one of Xanthophyceae were ob¬

served. There were no diatoms. These results are similar to those

of Esmarch (1914) who obtained mostly blue-green algae from

moist cultures and no diatoms.

It is to be noted that soil from MacRitchie Reservoir Forest

(Locality 6) did not develop any identifiable algae in moist cul¬

ture, although some minute colourless flagellates were observed.

The soil in this locality is in heavy shade and is covered with a

dense layer of large leaves throughout the year. It is not surprising

that the pigmented algae are relatively sparse. Similarly no algae

were obtained from the lower levels of the soil from the Botanic

Gardens jungle where similar conditions are found.

VI. Examination of Liquid Cultures

The conditions in liquid cultures differ markedly from those of

the natural environment. An additional supply of mineral sub¬

stances is supplied to the soil, in addition to an abundant supply of

water. Determination of the abundance of soil algae on growth in

cultures is subject to error in that they favour the more hydrophytic

plants (Petersen, 1935) in addition to selecting out those algae

which are particularly adapted to the culture environment. Never¬

theless, since all operations are carried out under sterile conditions,

the medium is autoclaved, etc., all algae which appear in such

cultures must have been present in the original soil habitat. There¬

fore this method indicates at least some of the algae spores, cells

and other bodies must occur in the natural environment but does

not indicate the relative abundance of these organisms.

The following is a list of the algae which grew in the liquid cul¬

tures. In most of the cultures a green or blue-green colour was

observable from seven to fourteen days after inoculation, but this

colour appeared earliest in those cultures of soil from nearest the

surface. The composition of the cultures was recorded at intervals

of one month. After two to three months staling phenomena com¬

menced and sub-culturing was necessary.

382

Vol. XIX. (1962).

Chlorophyceae

Palmella miniala Naeg.

Exposed cleared building site, Singapore (1.5), (2.5), (3.5).

Ourococcus bicaudatus Grobety

MacRitchie Reservoir Forest (3.5).

Pleurococcus naegelli Chodat

University of Singapore (0.5), (1.5), (2.5), (3.5); Exposed silt,

Dungun (0.5); Exposed cleared building site (0.5); Botanic

Gardens Jungle (1.5); Sunlit Path, MacRitchie Reservoir (0.5),

(1.5), (2.5), (3.5).

Trochisia aspersa (Reinsch.) Hansg.

MacRitchie Reservoir Forest (0.5), (1.5).

Vaucheria sessilis (Vauch.) D.C.

Botanic Gardens Jungle (0.5).

Cyanophyceae

Chroococcus multicoloratus Wood

Botanic Gardens Jungle (1.5), (2.5), (3.5).

Chroococcus varius A.Br.

Sunlit Path, McRitchie Reservoir (1.5), (2.5), (3.5).

Gloeotheca sp.

Exposed silt, Dungun (1.5).

Synechococcus cedrorum Sauvageau

MacRitchie Reservoir Forest (2.5), (3.5).

Oscillatoria sp. 1 . (1.3 wide, slightly capitate and twisted)

University of Singapore (0.5), (1.5), (3.5).

Oscillaoria sp. 2. (very coiled, strongly capitate)

Exposed silt, Dungun (0.5), (1.5), (2.5), (3.5).

Phormidium angustissimum West, W. et G.S.

Botanic Gardens Jungle (0.5).

Lyngbya mucicola Lemm.

Exposed cleared building site, Singapore (0.5), (1.5), (2.5), (3.5).

Microcoleus vaginatus (Vauch.) Gomont

University of Singapore (0.5), (1.5).

Microchaete tenera Thuret

University of Singapore (1.5).

Scytonema javanicum (Kiitz.) Born.

University of Singapore (1.5).

Summary and Conclusion

Samples of soil from five localities on Singapore Island and one

in the Federation of Malaya were cultured under sterile conditions.

“Moist cultures” were the best imitation of natural conditions but

continuous light was supplied. In these cultures three species of

the Chlorophyceae, one of the Xanthophyceae and ten of the

383

Gardens Bulletin, S.

Cyanophyceae were observed. There were no diatoms. No algae

appeared in the cultures from a forest locality.

“Liquid cultures” differed widely from the conditions of the

natural environment since they favoured the growth of hydrophytic

organisms particularly adapted to the nutrient medium supplied.

In these cultures five members of the Chlorophyceae and eleven

members of the Cyanophyceae were recorded.

All the algae recorded must have been present at least as spores

in the natural environment but no conclusions of their relative

abundance can be made.

References

Booth, W. E. (1941) Algae as pioneers in plant succession and their im¬

portance in erosion control. Ecology 22 38-46.

Bristol, B. M. (1920) On the algal-flora of some desiccated English soils.

Ann. Bot. 34: 36.

Bristol, B. M. (1919) On a Malayan form of Chlorococcum humicola

(Nacg) Rabenh. Jour, Linn. Soc., Bot. 44 473-82.

Coyle, E. E. (1935) Algae of some Ohio soils. Abstr. Doctoral Diss.,

Ohio State Univ. 17 217-227 (fide Tiffany, 1951).

Esmaech, F. (1914) Untersuchungen liber die Verbreitung der Cyanophy-

ceen auf und in verscheidenen Boden. Hedwigia 55 224.

Fritsch, F. E. (1907) The role of algal growth in the colonisation of new

ground and in the determination of scenery. Gcogr. Jour.

30 531-548.

Fritsch, F. E. (1907a) A general consideration of the sub-aerial and

freshwater algal flora of Ceylon. Proc. Roy. Soc. London.

B 79 197-254.

Fritsch, F. E. (1907b) The subaerial and freshwater algae of the tropics.

Ann. Bot. 21 235-275.

Ghose, S. L. (1923) A systematic and ecological account of a collection of

blue-green algae from Lahore and Simla. Journ. Linn. Soc.

London, Bot. 46 333-346.

James, E. J. (1935) An investigation of the algal growth in some naturally

occurring soils. Beih. Bot. Centralbl. 53 A 419.

John, R. P. (1942) An ecological and taxonomic study of the algae of

British soils. I The distribution of surface growing algae.

Ann. Bot., N.S., 6 323-349.

Lowe, C. W. & Moyse, A. V. (1934) An investigation of some Manitoba

soils for the presence of soil algae. Trans. Roy. Soc. Canada

28 119-152.

Moore, G. T. & Carter, N. Further studies on the subterranean algal flora

of the Missouri Botanic Garden. Ann. Missouri Bot. Gdn. 13

101-140 (1936).

Peterson, J. B. (1935) Studies in the biology and taxonomy of soil algae.

Dansk. Bot. Archiv. 8 no. 9.

Tiffany, L. H. (1951) in G. M. Smith, Manual of Phycology, Ch. 15 303-

306.

Treub, M. (1888) Notice sur la nouvelle flore de Krakatau. Ann. Jard.

bot. Buitenzorg 7 213.

384

Taxonomic Notes on Ficus L., Asia and Australasia

Addendum II

By E. J. H. Corner

Botany School, University oj Cambridge

Summary

New species and varieties.—subgen. Urostigma, F. microcarpa Linn,

f. var. rigo (Bailey) Corner; subgen. Ficus sect. Ficus, F. aureocor-

data Corner; sect. Rhizocladus Endl., F. ampulliformis Corner, F. hypo-

brunnea Corner, F. convexa Comer, E. ovatacuta Comer; sect.

Kalosyce, F. diandra Corner; sect. Sycocarpus Miq., F. moderata

Comer, F. virescens Corner, F. megaleia Corner var. multinervia Cor¬

ner and var. subuncinata Corner, F. uncinata Becc. var. subbeccarii

Comer F. scortechinii King var. lanceata Corner.

New subsections.—subgen Ficus sect. Sycocarpus Miq. subsect.

Papuasyce Corner (type, F. itoana Diels) and subsect. Lepidotus Cor¬

ner (type, F. griffithii Miq.).

Sect. Sycocarpus subsect. Pomifera Corner is reduced to sect. Neo-

morphe King ser. Auriculatae Corner.

The recent Royal Society Expedition to North Borneo, 1961,

has produced three interesting new species and four new varieties,

illustrative of the richness of the Bornean flora in this genus. One

species, F. diandra, was found within a few miles of Kuching.

Exploration of New Guinea forests is producing now a crop of

climbing species of sect. Rhizocladus, and I have taken the oppor¬

tunity to illustrate the figs of some species of this intricate section.

subgen. Urosigma Miq.

F. microcarpa Linn. f. var. rigo (Bailey) Corner comb. nov.—

F. rigo Bailey, Queensl. Agric. J. 1 (1897) 235.— F. retusa L.

var. rigo (Bailey) Diels, Bot. Jahrb. 67 (1935) 183.—A few

trees of this variety are planted in the main streets of Port

Moresby. They differ from typical F. microcarpa in the absence

of copious aerial roots from the branches, by the rather broad

and obtuse leaves, and by the slightly larger figs which ripen

yellow to orange without becoming pink, red, or purple. In the

herbarium it would seem impossible, without field-notes, to dis¬

tinguish v. rigo from v. latifolia (Miq.) Comer.

subgen. Ficus sect. Ficus subsect. Eriosycea (Miq.) Corner

ser. Auratae Corner subser. Auratae Corner

Corner, Card. Bull. Sing. 17 (1960) 420.

F. aureocordata Corner sp. nov.—Frutex foliis spiraliter dispositis.

Ramuli et petioli pilis rigidis aureobrunneis 1-2 mm. longis ves-

titi; lamina supra appresse pallide scabrido-pilosa; nervi et

nervuli subtus pilis pallidis -0.5 mm. longis vestiti. Ramuli 5-6

mm. crassi. Lamina -32 X 21 cm., ovato-cordata, apice -15

385

Gardens Bulletin , S.

mm. longo subacuminato, basi late et alte cordato, ciliato-den-

tata, membranacea, supra scabrida, subtus hispidula, sicco fla-

vescens; costis lateralibus utrinsecus 5-6, subtus valde elevatis,

glandulis axillaribus praeditis, marginem versus furcatis et in-

arcuatis; intercostis subtus elevatis, breviter pilosis, haud

velutinatis; areolis planis glabris; costis basalibus utrinsecus

2-3, -i laminae elongatis, glandulis basalibus 2-4 praeditis;

petiolo 8-12 cm. longo. Receptacula 17 mm. lata, axillaria ses-

silia subconoidea, pilis flavidis 1-2 mm. longis dense hispida;

bracteis basalibus ?, eruptis; bracteis lateralibus paucis, -3 mm.

longis, appresse pilosis; bracteis apicalibus erectis, appresse

pilosis, ut in cono 3 mm. alto confertis; setis intemis -2.5 mm.

longis, aureis, copiosis; cellulis scleroticis in pariete tenui nullis.

Text-Fig. 1„ F. aureocordata Corner, leaf, X i; fig- X 2; male and gall-

flowers, x 10. (Hans Winkler 895).

386

Vol XIX. (1962).

Flores masculi in ordinibus duobus instructi, prope apicem ses-

siles bistaminati, alteri pedicellati unistaminati; tepalis 4 spathu-

latis albidis; staminibus haud v. vix mucronatis, liberis, pilis

confertis ad basim filamentorum. Flores cecidiophori sessiles v.

pedicellis setosis —4 mm. longis praediti; tepalis 3-4, albidis,

apicem versus dense setosis, lineari-spathulatis; ovario flavidulo

sessili. Flores feminei ? Cystolitha nulla. Hans Winkler PI. bom.

895, Bt. Raja (112° 30' E, 0° 30' S), Borneo occid., 1,250 m.

alt. in silva; typus, herb. Leiden.

Though known from the single collection, this is a distinct spe¬

cies, peculiar in retaining lateral bracts on the fig. It comes in

leaf-shape between F. bruneiensis Comer and F. eumorpha Cor¬

ner, but the male flowers ally it with F. aurata Miq. It increases

the number of species of ser. Auratae to ten, of which nine are

endemic in Borneo.

subgen. Ficus sect. Rhizocladus Endl. ser. Ramentaceae

Corner subser. Pantonianae Corner

Comer, Gard. Bull. Sing. 18 (1960) 4.

F. ampulliformis Corner sp. nov. (Text-Fig. 2, H)—Scandens, foliis

distichis. Ramuli, petioli, costae subtus pilis albidis v. brun-

neolis -1 mm. longis appressis tenuiter vestiti. Ramuli 2 mm.

crassi. Stipulae -6 mm. longae, caducae, appressc pilosae.

Lamina 2.5-5 X 1.5-3 cm., elliptica, subacuta, basi cuneato,

valde coriacea marginibus incurvatis, scabrida, sicco dura fus-

cobrunnea; costis lateralibus utrinsecus 4, obliquis, subtus ele-

vatis, inarcuatis; intercostis 0-3, laxis, subtus vix elevatis; costis

basalibus utrinsecus 1, ad ± laminae; petiolo 5-10 mm. longo.

Receptacula axillaria binata hispidula, subglabrescentia; pedun-

culo 3-5 mm. longo; bracteis basalibus mox caducis; pedicello

0.5-1.5 mm. longo; corpore receptaculi 8-10 mm. lato, dep-

resso-globoso, ostiolo ad basim papillae apicalis 1 mm. altae

immerso; setis intemis nullis, vel paucis minutis; cellulis sclero-

ticis in pariete copiosis. Flores masculi pedicellati, ordinibus 2

v. 1 instructi; antheris submucronatis. Flores cecidiophori sessi¬

les v. pedicellati; tepalis 3-4, lineari-spathulatis, rubris, liberis;

ovario sessili ochraceo subindurato. Cystolitha amphigena.

F. A. W. Schram, BW 7743, Kebar valley, West New Guinea,

in silva c. 700 m. alt.; typus herb. Leiden.

This species is very distinct in its subseries from the small, stiff,

subacute, and scabrid leaves with few veins and from the charac¬

teristic shape of the fig which recalls that of F. villosa Bl.

F. hypobrunnea Corner, sp. nov. (Text-Fig. 2, F)—Scandens, foliis

distichis. Ramuli, stipulae, petioli, costae subtus, et receptacula

pilis brunneis v. flavidis 0.5-1 mm. longis appressis vestiti.

Ramuli 1.5-2 mm. crassi. Stipulae -8 mm. longae, dense seri-

ceae. Lamina 2.5-5.5 X 1.2-2.5 cm., lanceolato-elliptica, sub¬

acuta, basi angustato rotundato v. subcuneato, coriacea laevis,

387

Gardens Bulletin, S.

marginibus vix incurvatis; costis lateralibus utrinsecus 5-6,

oblique inarcuatis, supra vix impressis; intercostis 2-5, subtus

leniter elevatis; areolis planis; costis basalibus utrinsecus 1 (-2),

ad 1/3-i laminae; petiolo 3-6 mm. longo. Receptacula axillaria

solitaria, dense appresse pilosa, maturitate rubra; pedunculo l

mm. longo; bracteis basalibus 1 mm. longis, ovatis, caducis;

pedicello 1.5 mm. longo; corpore receptaculi 7 mm. lato, sicco

Text-Fig. 2. Figs of sect. Rhizocladus, X 3. A, F. amblisyce Corner, three

figs (v. Roycn 3775). B, F. pantoniana King var. colobocarpa Diels,

two figs (Clemens 2124). C, F. ftoccifera Diels, (Carr 13493). D, F.

camptandra Diels, (Buwalda 4993). E, F. pantoniana King, (Cart-

15830). F, F. hypobrunnea Corner, (BW 9283). G, F. baeuerleni King

var. vulcaniformis Corner, (NGF 3458). H, F, ampulliformis Corner,

(BW 7743).

388

Vol. XIX. (1962).

subellipsoideo, ostiolo minuto immerso; setis intemis copiosis,

-0.5 mm. longis, albidis; cellulis sclerotics in pariete copiosis.

Flores feminei sessiles; tepalis 3-4 rubris, anguste spathulatis v.

ovatis, liberis. Semina 1.2-1.5 mm. longa, compressa, anguste

alata. Cystolitha amphigena. F. A. W. Schram, BW 9283,

Oereb, div. Hollandia, New Guinea, in silva; typus, herb.

Leiden.

This is near to F. amblisyce Corner, but differs in the densely

hairy, pedunculate and pedicellate fig, the smaller basal bracts, and

the copious internal bristles.

subgen. Ficus sect. Rhizocladus Endi. ser. Ramentaceae

Corner subser. Irritantes Corner

Comer, Gard. Bull. Sing. 18 (1960) 4.

F. convexa Corner sp. nov. (Text-Fig. 3, D)—Scandens, foliis dis-

tichis. Ramuli, petioli, et costae subtus primo pilis rigidis irri-

tantibus brunneis 1.5-2.5 mm. longis strigosi; nervulis pilis al¬

bidis puberulis. Ramuli 2 mm. crassi. Stipulae —10 mm. longae,

appresse puberulae, ad carinam pilis rigidis paucis affixis.

Lamina -8 X 3.8 cm., ovato-elliptica v. ovato-lanceolata, sub-

acuta v. obtuse subacuminata, basi cordato v. rotundato, rigide

coriacea, convexa, inter costas bullata, marginibus decurvatis,

subscabrida; costis lateralibus utrinsecus 5—7, supra impressis,

subtus valde elevatis; intercostis -7, supra valde impressis, sub¬

tus vix elevatis; areolis glabris v. puberulis; costis basalibus

- utrinsecus 1, ad *—1/3 laminae; petiolo 5-10 mm. longo;

bathyphyllis 1.5-3.5 cm. longis, ovato-cordatis, tenue coriaceis,

bullatis, aequalibus, costis lateralibus utrinsecus 3—4 praeditis.

Receptacula axillaria; pedunculo 0-1 mm. longo; bracteis

basalibus 1.5-2.5 X 2-3 mm., ovatis acutis caducis, sparsim

puberulis; pedicello 1—5 mm. longo; corpore receptaculi 7-9

mm. lato, subgloboso v. subellipsoideo, tenue appresse pilosa,

ostiolo alte immerso; setis intemis nullis v. paucis minutis; cellu¬

lis scleroticis in pariete crasso nullis. Flores feminei sessiles v.

pedicellati; tepalis 3-4 rubris liberis. Semina 1.5-2 mm. longa,

compressa, anguste alata. Cystolitha amphigena. Brass 31121,

in silvis montanis 2,400 m. alt., Mt. Otto, Terr. New Guinea;

typus, herb. Leiden. NGF 6350 (leg. H. S. McKee 1352),

Goroka, 2,500 m. alt., Terr. New Guinea. Comer et Womersley

s.n., Goroka, Daulo Pass, 2,400 m. alt., bathyphylla.

This may be a high mountain state of F. insculpta Summerh.,

but in addition to the smaller, more convex, and bullate leaf the

smaller fig is pedicellate, lacks the flattened apex of F. insculpta ,

and also the sclerotic cells in the fig-wall.

389

Gardens Bulletin, S.

oides (Wall.) King, (Sing. F.N. 28605). G, F. recurva Bl. var.

lasiocarpa Corner, (Sing. F.N. 27816). H, F. recurva Bl. var. elegan-

tior Corner, (Sing. F.N. 30126). I, F. recurva Bl. var. bridelioidex

Corner, (Sing. F.N. 25941). J, F. urnigera Miq., (Sing. F.N. 25948).

390

Vol. XIX. (1962).

Text-Fig. 4. Figs of sect. Rhizocladus, X 3. A, F. balanota Diels, (Schlech-

ter 17601). B, F. sageretina Diels, (Schlechter 18238). C, F. excavate

Wall., three figs, (Sing. F.N. 29990). D, F. araneosa King, (King’s

Coll. 3565). E, F. callicarpides Corner, (Anderson 9890). F, F. lanata

Bl., (Koord. 40149).

subgen. Ficus sect. Rhizocladus Endl. ser. Distichoideae Corner

Gard. Bull. Sing. 18 (1960) 5.

F. ovatacuta Corner sp. nov. (Text-Fig. 5, A) — Scandens, foliis laxe

spiraliter instructs v. subdistichis. Ramuli et petioli pilis appres-

sis brunneis -1 mm. longis vestiti, costa subtus sparsim appresse

pilosa v. glabra. Ramuli 1.5-2 mm. crassi. Stipulae 6-10 mm.

longae, glabrae, ad novellas persistentes. Lamina 3-6 X 1.5-

3.2 mm., ovata, acuminato-acuta, basi rotundato subcordato v.

late cuneato, coriacea, sicco brunnea; costis lateralibus utrinse-

cus 5-7, subtus vix elevatis; intercostis 0-2, laxis, areolis planis;

costis basalibus utrinsecus 1, brevibus; petiolo 5-11 mm. longo.

Receptacula axillaria glabra subsessilia rubescentia; bracteis

basalibus 1.5 mm. longis, ovatis subacutis glabris; pedicello vix

evoluto; corpore receptaculi 10-13 mm. lato (15—17 mm., vivo),

subgloboso, ostiolo piano; setis intends et cellulis scleroticis

nullis. Flores feminei sessiles v. pedicellati; tepalis 3-4, spathu-

latis, liberis, rubris. Flores neutri sessiles ostiolares. Semina ?

Cystolitha amphigena sed supra sparsa. Brass 31624, in silva

montana 1,950 m. alt. pr. Parosa, Okapa, Eastern Highlands,

Terr. New Guinea; typus, herb. Leiden.

The large subsessile figs, the ovate-acute leaves, and the ap-

pressedly hairy twigs readily distinguish this species which seems

to be nearer to F. distichoidea than to F. phatnophylla.

391

Gardens Bulletin, S .

Text-Fig. 5. Figs of sect. Rhizocladus ser. Distichoideac, X 3. A, F. ovata-

cuta Corner, a fig and two leaves (Brass 31624). B, F. phatnophylla

Diels, one leaf, (Ledermann 9406). C, F. agapetoides Diels, a fig and

three leaves, (Carr 15799). D, F. phatnophylla Diels var. glochidioides

Corner, (v. Leeuwen 10166). E, F. calodictya Summerh., one leaf,

(Hoogland 4801). F, F. distichoidea Diels var. megacarpa Corner, a fig

showing the large male flowers, and one leaf, (Carr 16031). G, F.

distichoidea Diels, (Carr 12112).

392

Vol. XIX. (1962).

Ficus L. subgen. Ficus sect. Kalosyce Miq.

ser. Apiocarpeae Corner

Corner, Gard. Bull. Sing. 18 (1960) 22.

F. diandra Corner sp. nov. (Text-Fig. 6)—Scandens, radicibus

affixa, glabra v. novellis puberulis brunneo-pruinosis. Ramuli

1-1.5 mm. crassi. Stipulae 4-7 mm. longae, persistentes.

Lamina 4-9 X 2-4.2 cm., elliptica v. ovato- v. obovato-elliptica,

acute, basi anguste cordata, saepe subinaequilateralis, coriacea

integra, haud scabrida; costis lateralibus utrinsecus 3-5, angulo

lato orientibus, inarcuatis, supra obscuris, haud impressis, sub-

tus vix elevatis, intercostis nullis: costis basalibus utrinsecus 1-2

brevibus: petiolo 4-20 mm. longo. Receptacula axillaria soli-

taria v. ad ramulos efoliatos -15 mm. longos, pallide marmo-

rata, maturitate rubro-aurantiaca; pedunculo 1.5-4 mm. longo;

bracteis basalibus 3, 1-1.5 mm. longis, ovatis acutis subcaducis;

pedicello nullo; corpore receptaculi 12-14 X 11 mm. (vivo)

ellipsoideo, minute brunneo-furfuraceo, ostiolo 2 mm. lato

leniter immerso, bracteis 3 apicalibus planis occluso; setis inter-

nis et cellulis scleroticis parietalibus nullis. Tepala 3-4 (masc.),

4-5 (cecid.), membranacea, pallide rosea. Flores masculi dis-

persi copiosi, fl. cecid. superantes; pedicellis 0.5-2 mm. longis;

tepalis lanceolatis, breviter gamophyllis. Flores cecidiophori

sessiles; tepalis lanceolato-filiformibus brevissime connatis;

ovario brunneo indurato sessili v. breviter stipitato; stylo breviter

bilobato. Flores fern. ? Cystolitha hypogena. Sarawak, ad basim

trunci Durionis zibethini , ad viam Kuching-Matang. S. 13740

(typus, herb. Cantab.), leg. E. J. H. Comer 26 Sept. 1961.

To place this species in sect. Kalosyce (one stamen) seems

wrong. Sect. Rhizocladus Endl. ser. Distichae Corner seems the

right place for the bistaminate flower. In other points, however,

as the marbled fig, the pedicellate male flowers, the membranous

tepals, and the bifid stigma, the species agrees with sect. Kalosyce,

and it is very close to F. warburgii Elm. which has usually one,

rarely two, stamens. F. warburgii differs specifically in the obtuse,

obovate lamina with cuneate base, the more numerous lateral

nerves (5-8 pairs), and the larger fig with long pedicel. Both spe¬

cies may, nevertheless, be relics, much diminished in stature, of the

bistaminate ancestors of sect. Kalosyce as would link it with ser.

Distichae and ser. Trichocarpeae. The indurate gall-ovary of both

species recalls that of F. hederacea Roxb. in ser. Distichae. Female

flowers and seeds are still unknown in both F. diandra and F.

warburgii.

393

Gardens Bulletin, S .

snbgen. Ficus sect. Sycocarpus Miq.

Corner, Gard. Bull. Sing. 18 (1960) 37.

The numerous species of this section are the most difficult to

classify. The discovery of two new and demonstrative species,

F. moderata and F. virescens, on Kinabalu raises the hope that

others will be found equally elucidating. F. virescens emphasizes

the importance of geographical distribution in a section of recent

evolution, as seems to be the state in Sycocarpus; its allies F.

treubii King, F. francisci Winkler, and F. cereicctrpa Corner are

394

Vol. XIX. (1962).

also endemic in Borneo. On this note I take the opportunity to

revise the classification of F. oligodon Miq. (F. pomifera Wall,

ex King).

subsect. Papuasyce Corner subsect. nov.—Semina 1.2—1.5 mm.

longa, lentiformia, subcompressa, vix carinata, hilo non promi-

nenti. Perianthium rubrum saccatum, ovaria feminea et cecidio-

phora albida obtegens. Stamina 2 (-1), libera. Receptacula

pedunculata, basi tribracteata, bracteis lateralibus nullis; setis

internis nullis; cellulis scleroticis in pariete receptaculi nullis v.

paucis. Folia plus minus coriacea integra spiraliter disposita.

Cystolitha hypogena. New Guinea, F. itoana Diels (typus), F.

microdictya Diels.

In my earlier paper, cited above, I instituted subsect. Pomifera

Comer, with F. oligogon as type, to include also F. itoana, F.

microdictya, and F. griffithii Miq. In the meantime I have learnt

from J. T. Wiebes, at the Rijksmuseum v. Natuurlijke Historic in

Leiden, who is studying the fig-insects that I have managed to

collect, that the fig-insects of F. oligodon agree with those of F.

auriculata Lour, in sect. Neomorphe King, whereas those of F.

itoana and F. microdictya agree with the fig-insects of sect. Syco-

carpus subsect. Auriculisperma Comer (Solomon Isl.) and subsect.

Dammaropsis (New Guinea). This confirms the botanical evidence.

F. oligodon has very much the same distribution on the Asiatic

mainland as F. auriculata, but F. oligodon extends down into the

Malay peninsula as far as the south of Selangor and Pahang. Here

it has the typical form which, in floral detail, agrees exactly with

sect. Sycocarpus, particularly with F. itoana. In Burma, Indochina,

and Thailand, however, it intergrades with F. auriculata in leaf-

shape, hairiness, and the critical construction of the perianth,

which is saccate and entirely covering the ovary in F. oligodon,

and composed of three short separate tepals in F. auriculata. The

fact is that, in this prime respect in the classification of the genus,

the one species belongs in sect. Sycocarpus and the other in sect.

Neomorphe; the allies of the first are with it on the Asiatic main¬

land, while its false allies are in New Guinea with their relatives.

The first reaction might be to join sect. Neomorphe with sect.

Sycocarpus, but this would be neither useful nor beneficial. Syco¬

carpus hangs together in enomious complexity in eastern Malaysia

and its origin is not to be sought in modem Neomorphe. The alter¬

native, which I assume, is that F. oligodon represents a parallel

in floral evolution to Sycocarpus. Confirmation lies in F. variegata

Bl. var. chlorocarpa King (sect. Neomorphe) which in south China

has also developed the saccate perianth covering the ovary and is

in fact so similar to F. oligodon that it is generally necessary to

examine the flowers in order to distinguish herbarium-material.

F. rivularis Merr. (Philippines) of sect. Ficus is another peculiar

species convergent in this way on Sycocarpus.

Accordingly, I reduce subsect. Pomiferae Corner and ser. Pami -

ferae Comer to sect. Neomorphe ser. Auriculatae Comer. I erect

395

Gardens Bulletin , S.

subsect. Papuasyce for the two New Guinea species, and another

subsect. Leptotus (below) for the aberrant F. griffithii. Neverthe¬

less, F. oligodon must still be keyed out in sect. Sycocarpus; its

exclusion from subsect. Papuasyce scarcely alters the definition.

Parallel evolution is rife in Ficus , but this is the most perplexing

case. Entomology has come to the rescue.

subsect. Lepidotus Corner subsect. nov.—ser. Lepidotae Corner

Gard. Bull. 18 (1960) 39.—Semina 1 mm. longa, lentiformia,

subcarinata, hilo non prominenti. Perianthium rubrum sacca-

tum, ovaria feminea et cecidiophora obtegens. Stamen 1. Recep-

tacula bracteis lateralibus praedita; setis internis et cellulis

scleroticis nuliis. Folia mcmbranacea spiraliter disposita. Cysto-

litha hypogena. Burma, Thailand, F. griffithii Miq.

subgen. Ficus sect. Sycocarpus Miq. ser. Tubercuiifasciculatae

Sata subser. Hispidae Corner

F. moderata Corner sp. nov. (Text-Fig. 7)—Arbor -9 m. alta, spar-

sim ramosa, cauliflora; foliis spiraliter instructis, interdum oppo¬

site. Ramuli et petioli pilis patentibus pallidis v. brunneolis

-1.5 mm. longis tenue vestiti; nervi subtus puberuli, glabrescen-

tes. Ramuli 3-4 mm. crassi, fistulosi. Stipulae -15 mm. longae,

caducae, lanceolatae, glabrae v. basim versus pilosae. Lamina

8-21 X 4.5-11 cm., elliptica v. obovata, subacuminata, apice

obtuso subcaudato -15 mm. longo, basi late cuneato v. anguste

subcordato, saepe leniter asymmetrico, integra v. apicem versus

dentata, laevis, tenue subcoriacea, sicco brunnea; costis laterali¬

bus utrinsecus 6-8, obliquis; intercostis 4-11, regulariter instruc¬

tis, subtus elevatis; costis basalibus utrinsecus 2, brevibus, raro

ad 1/3 laminae extensis; petiolo 9-25 mm. longo, glandula sub-

nodali praedito. Receptacula ad ramulos ramosos cauliflores

-5 cm. longos dense aggregata, maturitate flava, primo brun-

neolo-pilosa; pedunculo 4—14 mm. longo; bracteis basalibus 2-3

mm. longis, ovatis subacutis; pedicello nullo; corpore receptaculi

15-23 mm. lato (20-30 mm., vivo), subturbinato, bracteis

lateralibus ovatis apicibus incurvatis 1-3, 2-3 mm. longis, ad

parietem affixis, circum ostiolum 5-8 mm. latum bracteis

numerosis occlusum lateralibus pluribus insertis; setis internis

nuliis; cellulis scleroticis ut lamina angusta in pariete recep¬

taculi. Flores masculi ordinibus duobus instructi; stamen 1.

Flores cecidiophori sessiles v. pedicellati; perianthio saccato ova¬

rium primo includenti dein disrupto. Flores feminei sessiles v.

breviter pedicellati; perianthio ut collare brevissimo; stylo glabro

v. setis 1-2 praedito. Semina 1 mm. longa, carinata, hilo pro¬

minenti, obscure subtuberculata v. gibbosa. Cystolitha hypogena.

RSNB 2592, in silvis secundariis pr. Kundasan 1,200 m. alt.,

North Borneo, (typus, herb. Kew); RSNB 2594 (recept.

cecidioph.).

This has the leaves of F. congesta Roxb. and the yellow figs

with lateral bracts of F. hispida Linn. f. In bridging, therefore,

396

Vol. XIX. (1962).

the two subseries Congestae and Hispidae it is another example

of the importance of the region of Kinabalu to the study of Ficus.

F. hispida is known merely from two collections in Borneo, one in

West Koetai, the other without local detail, and I have not seen

it myself in Sarawak, Brunei, or North Borneo. F. congesta has

no certain record from Borneo.

Text-Fig. 7. F. moderata Corner; leaf, X i; figs, X 1; flowers and seeds,

X 10. (RSNB 2592, 2594).

397

Gardens Bulletin , S.

subgen. Ficus sect. Sycocarpus Miq. ser. Tuberculifasciculatae

Sata subser. Fulvidulae Corner

F. virescens Corner sp. nov. (Text-Fig. 8)—Arbor -13 m. alta cau-

liflora. Folia spiraliter instructa v. opposita. Ramuli, petioli,

stipulae, costae subtus, et receptacula juvenilia pilis fulvidulis

appressis 0.5 mm. longis vestiti; lamina supeme glabra. Ramuli

2-4 mm. crassi, lente glabrescentes, fistulosi. Stipulae -15 mm.

longae, caducae. Lamina 12-30 X 4-12 cm., anguste elliptica,

ad apicem acuminatum abrupte caudatum 12-30 mm. longum

angustata, basi cuneato symmetrico, integra v. subdenticulata,

tenue subcoriacea, supeme laevis, subtus subscabrida, in sicco

supeme grisea, subtus cinereo-viridis et valde punticulata;

costis lateralibus utrinsecus 5-7, subtus acute elevatis; inter-

costis -11, vix elevatis, confertis; costis basalibus utrinsecus 1

brevibus; petiolo 15-70 mm. longo, glandula subnodali praedito.

Receptacula ad ramulos efoliatos caulifloros sine internodis

elongatis -30 cm. longos, basi 1-3 cm. crassos, aggregata,

maturitate flavo-brunnea, glabrescentia; pedunculo 5-22 mm.

longo, glabro; bracteis basalibus 3, ternata, 2.5-4 mm. longis,

glabris; corpore receptaculi 14-15 mm. lato (20-22 mm., vivo),

subellipsoidea, bracteis lateralibus nullis, apicem versus 5-6-

costato, ostiolo inter 5-6 bracteis apicalibus umbonatis immerso,

ostiolo in statu juvenili bracteis numerosis parvis in disco -10

mm. lata praedito et bracteis 5-6 erectis marginalibus circum-

dato; setis intemis copiosis 0.5 mm. longis; cellulis scleroticis

in pariete receptaculi nullis. Flores masculi ordine uno instructi,

stamine uno. Flores cecidiophori juveniles perianthio saccato

ovarium tegenti. Flores feminei sessiles v. breviter pedicellati;

perianthio ut collare angusto stipitem ovarii complectanti; stylo

puberulo. Semina 1 mm. longa, hilo prominenti, reticulato-sub-

tuberculata. Cystolitha hypogena. RSNB 965, in silva prope

rivulum 900 m. alt., Kinabalu (east ridge), 25.vii.61, (typus,

herb. Kew.); RSNB 2543, pr. Hum. Liwagu 1,200 m. alt.,

29.viii.61.

This species resembles F. treubii King in so many respects that,

confronted with it in the herbarium, one might regard it as a

sapling state with radial arrangement of the leaves on stouter twigs

and incipient geocarpy. Having studied two living and full-grown

trees, I realise that it is distinct in both the radial construction and

in the cauliflory, and this conclusion is confirmed by such details

as the longer peduncle, larger basal and apical bracts, larger fig-

body, shorter female perianth, and less hairy style. It elucidates, in

fact, the peculiarity of F. treubii among the geocarpic figs of

Borneo in subser. Geocarpicae because it represents the ancestral

state with the more massive spiral construction and relates thereby

F. treubii with subser. Fulvidulae, of which F. cereicarpa Corner

and F. jrancisci Winkl. are also endemic in Borneo.

398

Vol. XIX. (1962).

The peculiarities of F. treubii are the absence of conspicuous

lateral bracts, the distinct peduncle and collar of basal bracts, the

abundant internal bristles, the oblique lateral nerves, and the

greenness of the dried leaf; these are the characters which ally it

with F. virescens. In both the sides of the seed are conspicuously

roughened whereas in the three other species of subser. Fulvidulae

they are nearly or quite smooth. This is a detail the importance of

which is not clear. In view of the close correspondence in other

respects and of the striking series of cauliflorous forms from large

multibracteate figs of F. cereicarpa to the small figs of F. virescens

399

Gardens Bulletin , S.

and the geocarpy of F. truebii with distichous leaves, I remove F.

treubii from subser. Geocarpicae, where I had tentatively placed it,

to subser. Fulvidulae.

F. virescens also resembles F. schwarzii Koord. (F. miquelii

auct.) and F. scortechinii King. They are both advanced species

with slender distichous construction of the twigs, as in F. treubii.

The mature fig of F. virescens is very similar to that of F. scor¬

techinii, while the young fig resembles that of F. schwarzii . The

ribbing of the fig-body towards the prominent peripheral apical

bracts in F. virescens and F. scortechinii indicates the incorpora¬

tion of the lateral bracts in the apical rosette; thus, this feature,

the significance of which had not been clear to me, marks the

change-over from the fig with lateral bracts to that without, and

places F. virescens in this respect between F. cereicarpa and F.

treubii. The same feature appears in the geocarpic F. ternatana

Miq. and F. pleyteana Comer, also of distichous habit with short,

pale brown, appressed hairs as in F. treubii. They both differ,

however, in the abundant sclerotic cells in the fig-wall, as well as

in the brown colour of dried leaves, and they are species of eastern

Malaysia from the Moluccas to New Guinea; they are probably

convergent with F. treubii, as the eastern F. arfakensis King is

convergent in the subseries of F. congesta. F. schwarzii may

belong to this subseries, where I have classified it, but the position

of F. scortechinii needs further investigation, particularly in

Borneo where the new variety v. lanceata here described suggests

that its more generalised form may yet exist.

subgen. Ficus sect. Sycocarpus Miq. ser. Tuberculifasciculatae

Sata subser. Geocarpicae Corner

F. megaleia Corner Gard. Bull. Sing. 18 (1960) 57. var. multi-

ncrvia Comer v. nov.—Ad omnes partes pili -1 mm. longi fulvi

dense appressi. Costae laterales utrinsecus 14-24; basales 4-6

plus 9-11. RSNB 970, in locis terrae delapsae, 900 m. alt., Kina¬

balu, 25.vii.61 (typus herb. Kew.).

This is probably a wide-spread variety on Kinabalu; it was col¬

lected several times without number by the Clemens, but the

lamina is too large for the making of convenient specimens. The

hanging pink young leaves are conspicuous.

var. subuncinata Comer var. nov.—Ramuli et petioli pilis fulvis

1-2 mm. longis curvato-ascendentibus vestiti. Costae laterales

utrinsecus 7-9; basales 3-4 plus 6-7. Receptacula sicca 10-12

mm. lata, semper glabra. RSNB 1576, in locis terrae delapsae,

1,500 m. alt., Kinabalu (east ridge), 31.vii.61, (typus herb. Kew.).

Six trees, to 3 m. high, were found. They had the large leaves

with strongly, and asymmetrically, cordate base as in F. megaleia

but the glabrous figs of F. uncinata, of which they would pass in

the herbarium as v. pilosior.

400

Vol. XIX. (1962).

F. uncinata Becc. var. subbeccarii Corner v. nov.—Ramuli,

petioli et costa (subtus) pilis appressis fulvis v. fusco-brunneis

0.5-1 mm. longis vestiti; subtus nervulis pallidis minoribus. Sti-

pulae 10-20 mm. longae, persistentes, lanceolatae. Lamina 10-30

X 4.5-10.5 cm., obovata, apice -15 mm. longa acuminata, basi

anguste asymmetrico, uno latere anguste cuneato, altero cuneato-

subcordato, integra, supeme pilis sparsis appressis scabrida; costis

lateralibus utrinsecus 4-7; intercostis -8, subtus elevatis, regula-

riter instructs; basalibus 2 plus 4. Receptacula sicca 14-20 mm.

lata, hispido-strigosa. Semina 0.6 mm. lata. RSNB 1619, ad ripem

flum. Mamut, 1,200 m. alt., 4.viii.61, (typus, herb. Kew.); RSNB

2541, ad ripem flum. Liwagu, 1,200 m. alt., 29.viii.61.

F. beccarii differs from the varieties of F. uncinata with hairy

figs, such as var. strigosa, only in the characters of the leaf, parti¬

cularly its smooth upperside and symmetric base. I have studied a

great many plants of both in the forest and seen no intermediate

until I met with these two plants on Kinabalu; they have the almost

symmetric leaf of F. beccarii but the scabrid and appressedly hairy

upperside of F. uncinata. Hasty collecting of these startling, but

awkward, geocarpic species gives the impression of a few distinct

species; actually they are all extremely variable and in need of

much further field-study.

subgen. Ficus sect. Sycocarpus Miq. ser. Tuberculifasciculatae

Sata subser. Tuberculifasciculatae

F. scortechinii King var. lanceata Corner var. nov.—Arbor —3

m. alta. Internodi et nervi in juventute pilis pallidis, plus minus

appressis, -0.5 mm. longis, sparsim vestiti, glabrescentes aliter

glabra. Folia opposita vel spiraliter instructa, dein subdisticha.

Lamina 6—20 X 1.6-4 cm., lanceolata, attenuato-acuta, basi

cuneato, apicem versus serrulata v. integra; costis lateralibus utrin¬

secus 6—10; petiolo 5—10 mm. longo. Receptacula ad ramulos

paniculato-ramosos -8 cm. longos, 4-6 mm. latos, basi 10 mm.,

in cumulis caulifloris -15 cm. latis affixa, maturitate rubrocarnea;

pedunculo 10-20 mm. longo; bracteis basalibus 0.5-1 mm. longo;

corpore receptaculi 7-9 mm. lato (11-13 X 10-12 mm., vivo),

saepe irregulariter tuberculato, ad apicem subdepressum valde cos-

tato; setis intemis et cellulis scleroticis nullis. Perianthium femi-

neum ut collare brevissimum; stylo puberulo v. subglabro. RSNB

2815, in silva, 1,500 m. alt., Kinabalu pr. Tenompok, 6.ix.61,

(typus herb. Kew.).

If I had not carefully collected this, 1 should have taken it to be a

mixture of the leaves of F. tarennifolia and the figs of F. scorte¬

chinii. It is less hairy than F. scortechinii, the figs being glabrous

from the first. Because of the characteristic ribbing of the figs and

of the manner in which they are borne, I refer the collection to

F. scortechinii , even though this species of Burma, Thailand, and

Malaya has not been found in Borneo. The warts, commonly

present on the figs, do not seem to be caused by insects.

401

INDEX

New Taxa and Names in Bold-faced Type.

Achimus, 215, 218.

Achnanthes, 24.

brevipes var. intermedia, 25.

crenuata, 25.

exigua, 25.

hauckiana var. rostrata, 25.

lanceolata, 26.

var. rostrata, 26.

lanceolatum, 26.

stauroneiformis. 26.

subhudsonis, 27.

tenuissima, 27.

tropica, 27.

weltereckii, 27.

Achymus, 215, 218.

Acroceras crassi-apiculatum, 148.

munroanum, 148.

ridleyi, 148.

sparsum, 148.

zizanioides, 148.

Agrostis diandra, 178.

matrella, 180.

maxima, 179.

procera, 160.

virginica, 178.

Aira indica, 174.

Albradia, 215, 218.

Albrandia, 215, 218.

Allaeanthus, 233.

kurzii, 234.

luzonicus, 235.

zeylanicus, 235.

Alloteropsis cimicina, 148.

Ampalis, 214.

Amphilophis glabra var. paurpera

151.

pertusus, 151.

Amphiprora lepidoptera, 63.

Amphora, 54.

aculiuscula, 55.

angusta, 55.

bitumida. 55.

coffaeiformis, 56.

var. aculiuscula, 55

var. salinarum, 55.

exigua, 56.

holsatica, 56.

var. malayana, 57.

libica, 57.

lineata, 55.

normani, 57.

ovalis var. libica, 57.

perpusilla, 57.

proteus, 58.

Andropogon aciculatus, 153.

amaurus. 173.

annulatus, 156.

brevifolius. 174.

caricosus, 156.

contortus, 163.

flexuosus, 154.

intermedius, 151.

lanceolatus, 149.

leschenaultianus, 161.

nardus, 154.

squarrosus, 173.

wightianus, 153.

Anomoconeis, 38.

brachysira, 38.

Anomoeoneis serians var. acuta, 39.

Anthers (Moraceae). 192.

Anthistiria arguens. 178.

arundinacea, 179.

gigantea var. arundinacea. 179.

Anthophysa, 135.

vegetans, 136.

Anthoxanthum indicum, 172.

Antiaris, 244.

Antiaris (List of Names), 249.

africana, 247.

bennettii, 248.

challa, 248.

humbertii, 247.

kerstingii, 248.

macrophylla. 248.

madagascariensis, 247.

toxicaria var. macrophylla. 248.

var. toxicaria, 248.

var. welwitschii, 248.

usambarensis, 248.

welwitschii, 248.

Antiaropsis, 249.

Aphananthes negrosensis, 221.

Apluda mutica, 148.

varia var. intermedia, 148.

Aristida adscens.onis, 149.

culionensis. 149.

setacea, 149.

Arthraxon lanceolatus, 149.

nudus, 149.

Arthrophyllum borneense, 185.

merrillianum. 185.

Artocarpaceae, 209.

Artocarpeae, 231.

(Inflorescences of), 201.

Ariocarpoideae, 209.

Artocarpus, 233.

elongatus. 227.

Arundinella setosa, 149.

Arundo donax. 149.

karka, 172.

reynaudiana, 169.

Ashton. P.S.: Some New Diptero-

carpaceae from Borneo, 253.

Aulosira pseudoramosa, 382.

Axonopus afifinis, 149.

compressus, 149.

var. aflinis, 149.

403

Gardens Bulletin, S.

Bacillaria paradoxa, 68.

ulna, 16.

Bacillariophyceae, 1.

Bam bo stricta, 156.

Bam bos arundinacea, 149.

Bambusa apus, 161.

arundinacea, 149.

aspcra, 155.

blumeana, 150.

burmanica, 150.

elegans, 150.

glaucescens, 150.

heterostachya, 150.

latispiculata, 150.

levis, 162.

magica, 150.

montana, 150.

multiplex. 150.

nana, 150.

pauciflora, 151.

ridleyi, 151.

Bambusa scandens, 158.

spinosa, 150.

vulgaris, 151.

wrayi, 151.

Barbula consanguinea. 374.

Bark Morphology (Studies in), 321.

Anisoptcra, 342.

Balanocarpus, 342.

Dipterocarpus, 344.

Dryobalanops, 346.

Hopea, 346.

Shorea, 349.

Vatica, 360.

Bark Taxonomy (Conclusions on

Dipterocarpaceae), 363.

Bark Types (Key to), 327.

Deep Fissured Bark, 334.

Dippled Bark, 332.

Laminate Bark, 341.

Scaly Bark. 335.

Shallow Fissured Bark, 332.

Smooth Bark. 328.

Surface Rotten Bark, 340.

Batis fruticosa. 239.

spinosa, 242.

Bathratherum nudum, 149.

Bibliographies, 78, 138, 251, 367, 374,

377, 384.

Bleekrodia. 215, 216.

insignis, 219.

madagascaricnsis, 220.

tonkinens’s. 228.

Bornean Araliads (Name Changes in),

185.

Bothriochloa intermedia, 151.

pertusa, 151.

Botrymorus. 251.

Brachiaria distachya, 151.

Brachiaria holotricha, 151. ■

mutica. 152.

paspaloides. 152.

reptans, 152.

Brackenridgca (The Species in the

Singapore Herbarium), 181.

sect. Capitatae, 182.

sect. Spiciformes, 183.

corymbosa, 183.

denticulata, 183.

foxworthyi, 184.

hookeri, 182.

kingii, 183.

palustris, 183.

rubescens, 183.

serrulata, 184.

Briza minor, 152.

Brosimeae, 250.

(Inflorescences of), 202.

Broussonetia, 233.

sect. Allaeanthus, 234.

sect. Broussonetia, 234.

kurzii, 234.

luzonensis, 235.

luzonica, 235.

var. glabra, 235.

var. luzonica. 235.

zeylanica, 235.

Broussonctieae, 231.

Bryophytes Collected by Prof. Gilli¬

land in Eastern New Guinea, 373.

Bucephalon, 230.

Calius, 215, 218.

Caloneis, 31.

ladogensis var. cuneata, 32.

Caloneis silicula var. gibba, 32.

var. minuta, 32.

Calpidochlamys. 230.

branderhorstii, 231.

drupacea. 231.

Campylodiscus, 75.

clypeus, 75.

Campylopodium euphorocladium,

373.

Campylopus richardii, 373.

Capiilipedium parviflorum, 152.

Cardiogyne, 235, 237.

africana, 237.

Cecropia, 251.

Cenchrus grandularis, 163.

lappaceus. 152.

Centotheca lappacea, 152.

var. longilamina, 152.

latifolia, 152.

longilamina, 152.

Centrales, 4.

Cephalostachyum malayanum, 156.

Cephalotrophis, 240.

Ceratoneis closterium, 65.

Chaetoceros, 11.

amanita, 11.

Chamaesiphonales, 375.

Chevalicrodendron, 215, 216.

glabrum, 221.

Chilomonas, 119.

paramecium, 119.

Choris barbata, 152.

dolichostachya, 152.

ridleyi, 161.

Chlorococcales, 375.

Chlorococcum humicolum, 375.

Chloromonadineae, 115.

404

Vol. XIX. (1962).

Chlorophora, 235, 236.

humbertii, 237.

regia, 237.

Chlorophyceae, 375, 381, 383.

Chromulina, 122.

sphaerica, 122.

Chroococcales, 375.

Chroococcus multicoloratus, 383.

sp., 381.

varius, 375, 381, 383.

Chrysamoeba, 122.

radians, 122.

Chrysodidynms, 128.

gracilis, 128.

synuroideus, 128.

Chrysophyceae, 120.

Chrysopogon aciculatus, 153.

collinus, 153.

orientalis, 153.

wightianus, 153.

Chrysopyxis, 126.

spp., 127.

Cocconeis, 23.

feuerbornii, 24.

thumensis, 24.

Cocconema lanceolata, 59.

stomatophorum, 60.

tumidum, 60.

Coelachne simpliciuscula, 153.

Coelorachis foveolata, 153.

glandulosa, 153.

helferi, 153.

Coix gigantea, 154.

lachryma-jobi, 154.

var. gigantea, 154.

Collelonema eximium, 30.

Conocephaloideae, 251.

Corner, E. J. H.: The Classification of

Moraceae, 187.

Taxonomic Notes on Ficus L.,

Asia and Australasia, Addendum

II, 385.

Coscinodiscus, 8.

antiquus, 8.

argus, 8.

decipiens, 8.

divisus, 9.

excentricus, 9.

griseus var. gallopagensis, 9.

lineatus, 9.

var. van-heurckii, 9.

striatus, 8.

symmetricus, 10.

Coussapoa, 251.

Craterogyne, 250.

Cryptomonas, 118.

erosa var. reflexa, 118.

ovata, 118.

phaseolus, 118.

Cryptophyceae, 117.

Cudrania, 235, 237.

amboinensis, 239.

cambodiana, 239.

cochinchinensis, 239.

fruticosa, 239.

grandifolia, 239.

javaensis, 239.

obovata, 239.

Cudrania poilanei, 242.

pubescens, 239.

thorelii, 240.

tricuspidata, 240.

triloba, 240.

Cudranus, 235, 237.

Cyanophyceae, 375, 381, 383.

Cyclotella, 7.

kuetzingiana, 7.

meneghiniana, 7.

striata. 8.

Cymbella, 58.

cuspidata, 58.

javanica, 59.

kolbei, 59.

lanceolata, 59.

sumatrcnsis, 59.

tumida, 60.

turgida, 60.

ventricosa, 60.

Cymbopogon calcicola, 154.

fiexuosus, 154.

nardus, 154.

Cynodon arcuatus, 154.

dactylon, 154.

tcner, 161.

Cynosurus aegypticus. 155.

corocanus, 159.

indicus, 159.

retroflexus, 158.

Cyrtococcum accrescens, 154.

carinatum, 155.

oxyphyllum, 154.

patens, 155.

trigonum, 155.

Dactylococcus, 375.

Dactyloctenium aegyptiacum, 155.

Dendrocalamus asper, 155.

dinosus, 155.

elegans, 155.

flagellifer, 155.

giganteus, 156.

hirtellus. 156.

pendulus, 156.

sinuatus. 156.

strictus. 156.

Dermocarpa hemisphaerica, 375.

Desmogonium, 23.

rabenhorstianum, 23.

Diandrochloa japonica, 159.

Diatoma tabulatum, 16.

Diatoms 1.

Dichanthium annulatum, 156.

caricosum, 156.

Digitaria adscendens, 156.

barbata, 157.

bicornis, 157.

chinensis, 157.

didactyla. 157.

longiflora, 157.

marginata. 157.

pertenuis, 157.

pusilla, 157.

405

Gardens Bulletin, S.

Dimeria ciliata, 157.

fuscens var. barbata. 157.

glabra, 157.

kurzii, 157.

leplorachis, 158.

ornithopoda, 158.

var. glabra. 157.

Dimerocarpus, 215, 217.

brenieri, 227.

Dinebra arabica. 158.

retroflexa, 158.

Dinobryon, 130.

bavaricum, 131.

var. affine, 131.

cylindricum var. ceylonicum, 131.

elongatum, 131.

var. affine, 131.

var. undulatum, 131.

inflatum, 132.

sertularia, 131.

var. thyrsoideum. 131.

sociale var. bavaricum. 131.

stipitatum, 131.

subsp. bavaricum, 131.

subsp. eustipitatum, 131.

var. affine, 131.

var. bavaricum, 131.

var. undulatum. 131.

thyrsoideum, 131.

Dinobryopsis, 130.

Dinochloa, 158.

montana, 150.

scandens, 158.

tjankorreh, 158.

Diplachne fusca, 158.

malayana. 158.

Diplocos, 215, 218.

macrophyllus, 227.

zeylanica. 229.

Diploneis, 34.

bombus var. minor. 34.

decipiens var. parallela, 34.

Diploneis elliptica var. linearis, 34.

interrupta. 34.

oblongella var. baltica. 35.

var. fossilis, 35.

ovalis, 35.

var. oblongella, 35.

parma, 35.

Diplosiga frequcntissima, 135.

Diplosigopsis frequentissima. 135.

Diplothorax, 215, 218.

tonkinensis, 228.

Dipterocarpaceae (Bark Taxonomy

in). 321.

(New Species from Borneo), 253.

Dorstenia, 250.

Dorstenieae (Inflorescences of), 202.

Dorstenieae, 250.

Dryobalanops, 346.

Dumartroya, 240.

Echinochloa colonum, 158.

crus-galli, 158.

var. frumentacea, 159.

frumentacea, 159.

stagnina, 159.

Echinolaena polystachya, 173.

Ectropothecium aurifolium, 374.

falciforme, 374.

sp., 374.

Eleusine aegyptiaca. 155.

corocana, 159.

indica, 159.

Embryos (of Moraceae), 197. 199.

Encyoncma turgidum, 60.

Endosigma exinium, 30.

Ephebopogon gratus, 168.

Epicarpurus, 215, 218.

microphyllus, 221.

spinosus, 225.

timorensis, 225.

zeylanicus, 229.

Epiterranean Soil Algae, 379.

Epithemia gibba var. parall.la, 63.

gibberula, 62.

musculus, 63.

Epipyxis, 130.

Eragrostis amabilis, 160.

atrovircns, 159.

chariis, 159.

elegantula, 159.

interrupta. 159.

japonica, 159.

malayana. 160.

pilosa, 160.

tcnella, 160.

unioloides, 160.

Eremochloa ciliaris, 160.

malayana. 160.

Eriachne pallescens, 160.

triseta, 167.

Erianthus speciosus, 161.

Eriochloa annulata, 160.

polystachya, 160.

procera, 160.

ramosa, 160.

Euartocarpeae, 231.

Euglena, 108.

acus var. mutabilis, 111.

agitis, 108.

Euglena cxilis, 108.

flava, 109

fusiform is. 111.

gracilis, 109.

intermedia var. klebsii, 110.

mutabilis, 111.

pisciformis, 108.

Euglenaceae, 108.

Euglenineae, 107.

Eulalia lan-pes, 161.

leschenaultiana, 161.

milsumi, 161.

praemorsa. 173.

ridleyi, 161.

speciosa, 161.

Eulejeuna sp.. 373.

Eumorcae, 211.

Eunotia. 17.

amphioxys, 64.

camelus. 17.

var. karveerensis, 17.

cancellata var. esseda, 18.

406

Voi XIX. (1962).

diodon var. minor, 18.

exigua, 18.

faba var. densestriata, 18.

flexuosa, 18.

formica, 19.

gracilis, 19.

grunowi var. uplandica

f. subundulata, 19.

indica, 20.

lunaris, 19.

var. capitata, 19.

major var. bidens, 20.

Eunotia major var. emarginata, 20.

var. hybrida

f. bidens, 20.

var. ind'ca

var. linearis, 20.

var. undulata, 21.

monodon var. alpina. 20.

var. constricta, 20.

var. major, 20.

var. undulata, 21.

papilio f. minor, 22.

pectinalis, 21.

var. incisa, 23.

var. minor, 21.

var. stricta, 19.

polydentula var. pcrminuta, 21.

praemonos var. inflata, 21.

var. monodon. 22.

var. tibetica, 22.

f. tibetica, 22.

praerupta var. bidens, 22.

var. curta f. monodon, 22.

var. inflata. 21.

var. monodon, 22.

var. tibet’ca, 22.

praerupta-monos f. undata, 22.

robusta, 22.

sarekensis var. minor, 22.

sudetica var. inc’sa, 23.

tridentula var. pcrminuta, 21.

Euphorbiaceae. 251.

Eustachys tenera, 161.

Excoecaria baccifera, 221.

Fatoucac, 211.

Festuca fusca, 158.

Ficeae, 251.

Ficus L., Asia & Australasia, 385.

Ficus (Inflorescences of), 202.

Ficus sect. Ficus, 385.

sect. Kalosyce, 393.

sect. Rhizocladus, 387, 389, 391.

sect. Svcocarpus, 394. 396, 398, 400

401.

ser. Apiocarpeae. 393.

ser. Auratae, 385.

ser. Distichoideae. 391.

ser. Lepidotae, 396.

ser. Ramentaceac, 387, 389.

ser. Tuberculifasciculatae, 396, 398

400, 401.

subgen. Ficus, 385, 387, 389, 391

393, 394. 396, 398. 400, 401.

subgen. Urostigma. 385.

subsect. Eriosycea, 385.

subsect. Lepidotus, 396.

subsect. Papuasyce, 395.

subser. Auratae, 385.

subser. Fulvidulae, 398.

subser. Geocarpicae, 400.

subser. Hispidae, 396.

subser. Irritantes, 389.

subser. Pantonianae, 387.

subser. Tuberculifasciculatae, 401.

aiiipiillifarmis, 387.

aureocordata, 385.

challa, 248.

convexa, 389.

diandra. 393.

hypobrunea. 387.

megaleia var. nniltinervia. 400.

var. subuncinata, 400.

Ficus microcarpa var. rigo. 385.

nioderata. 396.

ovataeuta. 391.

retusa var. rigo, 385.

rigo, 385.

scortechinii var. lanceata. 401.

uncinata var. subbeccarli, 401.

virescens. 398.

Filaments (of Urticaceae & Moraceae),

190.

Fissidens zollingeri, 373.

Flagellata (of Malayan Freshwaters).

105.

Floral Parts (of Moraceae), 200.

Fragilaria, 15.

intermedia, 15.

lapponica var. tetragonalis 15.

vaucheriae, 15.

virescens var. elliptica, 15.

Fruits (of Moraceae), 196.

Frustulia, 27.

appendrculata, 49.

javanica, 28.

rhomboides, 28.

var. saxonica, 28.

saxonica, 28.

serians, 39.

Furtado, C. X.: Name Changes in

Bornean Araliads, 185.

:The Species of Brackenridgea in

the Singapore Herbarium, 181.

Fusticus, 235.

Gaillonella granulata, 6.

italica, 6.

marchica, 6.

procera, 6.

Garnotia stricta, 161.

Gigantochloa apus, 161.

hasskarliana, 161.

heterostachya, 150.

kurzii, 162.

latifolia, 161.

latispiculata, 150.

levis, 162.

ligulata, 162.

maxima, 162.

ridleyi, 162.

407

Gardens Bulletin, S.

scortechinii, 162.

var. albovestita, 162.

verticillata, 162.

var. minor, 162.

var. viridis, 162.

wrayi, 162.

Gilliland, H. B.: A Checklist of

Malayan Grasses, 147.

Gironniera glabra, 221.

Gloeotheca sp., 383.

Gomphia corymbosa, 182, 183.

hookeri, 182, 183.

var. corymbosa. 183.

perakensis, 182.

Gomphonema, 60.

gracile, 61.

longiceps var. subclavata

f. gracilis, 61.

parvulu, 61.

subtile, 61.

subventricosum, 62.

Gonyostomum, 116.

depressum, 116.

semen. 116.

Gyrosigma, 29.

attenuatum. 29.

distortum, 29.

var. parkeri. 29.

excentricum. 30.

fasciolum var. arcuatum, 30.

scalproides var. eximium, 30.

spenccrii var. smithii, 31.

Hantzschia, 64.

amphioxys var. capitata, 64.

var. vivax, 64.

Haekelochloa granularis, 163.

Hemarthria protensa, 163.

vaginata, 163.

Hemigymnia fusca, 170.

multinodis, 170.

Hepaticae, 373.

Heteropogon contortus, 163.

Himantidium exiguum, 18.

faba, 18.

gracilis, 19.

majus, 20.

pectinalis, 21.

var. minor, 21.

Holcus pertusus, 151.

Hopea fltivialis. 254.

garangbuaya. 256.

vaccinifolia. 258.

wyattsmithii. 260.

Hyalobryon. 132.

Iauterbornii, 132.

Hymenachne aurita. 163.

pseudointerrupta, 163.

Ichnanthus pallens, 163.

vicinus, 163.

Imperata arundinacea, 164.

conferta, 164.

contracta, 164.

cylindrica, 163.

exaltata, 164.

Inflorescences (of Moraceae), 202.

Isachne albens, 164.

var. sylvcstirs, 165.

australis, 164.

confusa, 164.

globosa, 164.

javana, 164.

var. denticulata, 165.

var. saxicola, 165.

kinabaluensis, 164.

kunthiana. 164.

lankawiensis, 165.

miliacea, 165.

rigida, 164.

saxicola, 165.

semitalis, 164.

sylvestris, 165.

Ischaemum apricum, 165.

aristatum, 166.

barbatum, 166.

var. imbricatum, 165.

ciliare, 165.

commelynifolium, 165.

fieldingianum, 165.

fluviatile, 165.

goebelii, 165.

imbricatum, 165.

Ischaemum indicum, 165.

involutum, 179.

laeve, 166.

macrurum, 166.

maculatum, 165.

magnum, 166.

muticum, 166.

plumosulum, 166.

polystachym, 166.

rugosum, 166.

trimorense, 166.

Isochrysideae, 127.

Isopterygium albescens, 374.

Johnson, Anne: A Short Note on

Some Soil Algae from New

Guinea, 375.

:On a small collection of

Bryophytes collected by Pro¬

fessor Gilliland in Eastern

New Guinea. 373.

: Precursory Studies on the

Epiterranean Soil Algae of

Singapore and Malaya, 379.

Joxylon, 235.

Lagerheimia splendens, 124.

Lagurus cylindricus, 163.

Leersia hexandra, 166.

Lepocinclis, 111.

fusiformis. 111.

sphagnophila. III.

Leptaspis urccolata, 167.

Leptochloa chinensis, 167.

panicea, 167.

408

Vol. XIX. (1962).

polystachya, 167.

wightiana, 168.

Lepturus repens, 167.

Leucobryum subsanctum, 374.

Lophatherum gracile, 167.

Ludolfia glaucescens, 150.

Lyngbya mesotrichia, 375, 381.

mucicola, 383.

sp., 381.

Maclura, 235.

sect. Cardiogyne, 237.

sect. Chlorophora, 236.

sect. Cudrania, 237.

sect. Madura. 236.

ser. Connatae. 238.

ser. Liberae, 238.

subgen. Eumaclura, 236.

subgen. Leptosura, 237.

africana. 237.

amboinensis, 239.

var. amboinensis. 239.

var. paucinervia, 239.

cochinensis. 238, 239.

var. bancrofti, 239.

var. pubescens. 239.

excelsa, 237.

fruticosa, 238, 239.

greveana, 237.

humbertii. 237.

javanica, 239.

regia. 237.

thorelii. 240.

timorensis, 239.

tinctoria, 236.

tricuspidata, 240.

Maillardia, 230.

Malaisia, 240.

Malayan Grasses (A Checklist of),

147.

Mallomonas, 123.

acaroides. 123.

alpina, 125.

curta, 123.

litomesa var. curta, 123.

playfairii var. opisthiodonta, 124.

spherica, 124.

splendens, 124.

teilingioides. 125.

tonsurata var. alpina, 125.

var. dorsidentata, 126.

var. megalepis, 125.

Manisuris granularis, 163.

Marchantia geminata, 373.

polymorpha, 373.

Massia triseta, 167.

Mastogloia, 76.

exigua, 76.

Melocanna gracilis, 175.

zollingeri, 176.

Melosira, 5.

crenulata, 6.

var. italica, 6.

var. tenuis, 6.

granulata, 6.

itaiica, 6.

ordinata, 6.

roeseana, 6.

ruettneri, 7.

spinosa, 6.

Mesogyne, 250.

Mctatrophis, 251.

Metrotrichia, 116.

capitata, 117.

Microchaete tenera, 383.

Microcoleus vaginatus, 382, 383.

Microscopic characters (of leaves of

Moraceae), 200.

Microstegium ciliatum, 167.

geniculatum, 167.

hcndersonii, 168.

nudum, 168.

vagans, 168.

Milicia, 235, 236.

Milium cimicinum, 148.

compressus, 149.

globosum, 164.

Miscanthus japonicus, 168.

sinensis, 168.

Mnesithea cancel lata, 168.

geminata, 168.

mollicoma, 168.

pubescens, 168.

rupincola, 153.

Mniomalia semilimbata, 374.

Moraceae (The Classification of), 187.

(Subdivisions of), 189.

Moreae, 211.

(Inflorescences of), 201.

Moroideac, 209.

Morus, 214.

alba var. laevigata, 215.

brunoniana, 222.

integrifolia, 240.

laevigata, 215.

luzonica. 235.

macroura, 215. *

pendulina, 222.

Musanga, 251.

Musci, 373.

Myrianthus, 251.

Myriostachya wightiana, 168.

var. longispicula, 169.

Nardia hasskarliana, 373.

Nardus ciliaris, 160.

Navicula, 39.

aflinis, 33.

attenuata var. attenuata, 29.

ambigua, 42.

amphibola, 39.

ampliata. 33.

anglica, 39.

antiqua, 40.

apiculata, 41.

var. maculifera, 41.

arenaria, 40.

bicapitata var. hybrida, 51.

bogotensis, 50.

brachysira, 38.

brasiliensis var. minor, 40.

409

Gardens Bulletin , S.

braunii, 50.

brebisonii, 51, 53.

cancellata var. apiculata, 41.

confervacea, 41.

crucifera, 41.

cryptocephala var. lancettula, 42.

var. pumila, 42.

var. veneta, 42.

cuspidata var. ambigua, 42.

dicephala var. subcapitata, 40.

disparata, 42.

elegantoides, 42.

Navicula feuerborni, 43.

gastrum, 43.

var. anglica, 39.

gemina, 34.

glacialis var. septentrionalis, 43.

grevillei, 43.

halophila var. subcapitata, 44.

hemiptera, 51.

hitchcockii, 32.

hungarica var. luneburgensis, 44.

hustedtii, 44.

iridis, 33.

interrupta, 34.

lagerheimi var. intermedia, 44.

lanceolata var. arenaria, 40.

lancettula, 42.

lyra var. dilatata, 44.

var. ehrenbergii, 45.

lyra-typica, 45.

macilenta, 52.

mesolepta, 52, 53.

microcephala, 45.

minima, 45.

minuscula, 45.

oblongella, 35.

ostenlieldii, 45.

ovalis, 35.

var. fossilis, 35.

parvula, 53.

pinnularioides var. continua, 40.

placenta var. obtusa, 45.

platystoma. 46.

polyonca, 53.

producta, 33.

punctulata, 46.

Navicula pupula var. capitata, 46.

pusilla, 46.

radiosa var. minutissima, 47.

rhombica, 43.

rhomboides, 28.

rhyncocephala, 47.

rotaeana, 47.

schroeteri. 47.

serians. 39.

splendida, 74.

stauroptera, 53.

subtilissima, 48.

tenella var. minutissima, 47.

tenera, 48.

toornensis, 48.

tridentula, 48.

trituberculata, 48.

tumida var. gemina, 39.

uniseriata, 48.

veneta, 42.

ventricosa f. minuta, 32.

Neckeriopsis gracilenta. 374.

Neidium, 32.

eapitatum. 33.

var. prolongata, 33.

hitchcockii, 32.

iridis var. ampliatum, 33.

var. intercedens, 33.

productum, 33.

var. minor, 33.

Neohopea isoptera, 293.

Neyraudia madagascariensis, 169.

reynaudiana, 169.

Nitzschia, 64.

Nitzschia acicularis, 65.

amphibia var. acutiuscula, 65.

clausii, 70.

closterium, 65.

communis var. abbreviata, 65.

constricta, 65.

diducta, 65.

elongata var. minor, 64.

fonticola, 66.

gandersheimiensis, 66.

granulata, 66.

heufleuriana, 66.

ignorata, 66.

littoralis var. tergestina, 67.

lorenziana var. subtilis, 67.

marginulata, 67.

microcephala, 67.

navicularis, 67.

obtusa, 68.

var. brevissimo, 68.

var. scalpelliformis, 68.

palea, 68.

paradoxa, 68.

parvula, 68.

plana, 69.

punctata var. apiculata, 69.

var. malayana, 69.

romana, 70.

scalaris, 70.

sigma, 70

var. clausii, 70.

stagnorum. 71.

subtilis, 71.

surirelloidea, 71.

Nitzschia towutensis. 71.

tryblionella var. victoriae, 72.

vivax, 64.

Nostoc linckia, 376.

microscopicum, 376.

Nostocales, 375.

Ochlandra ridleyi, 176.

Ochna foxworthyi, 184.

Ochromonadeae, 129.

Ochromonas, 130.

hinterzartensis, 130.

Olmedieae, 243.

(Inflorescences of), 202.

Oplismenus burmanii, 169.

compositus, 169.

410

voi xix. (m2).

Orlhosira punctata, 6.

spinosa, 6.

Oryza fatua, 169.

var. longe-aristata, 169.

granulata, 169.

lati folia, 169.

minuta, 169.

ridleyi, 169.

rufipogon, 169.

Oscillatoria spp., 383.

Ottochloa nodosa, 170.

Ouralea hookeri, 182.

Ourococcus bicaudatus. 383.

Ovaries (of Moraceae), 195, 196.

Oxytenanthera auriculata. 162.

var. alba, 162.

var. efimbriata, 162.

nigrociliala. 161.

sinuala, 156.

Pachytrophe, 214.

Palmella miniata, 383.

Panicum accrescens, 154.

adscendens. 156.

amplexicaule. 163.

auritum, 163.

austro-asiaticum, 170.

brevifolium, 170.

caesium, 170.

cambogiense, 170.

cplonum, 158.

compositum, 169.

crus-galli, 158.

var. frumentaceum. 159.

var. stagninum, 159.

distachym, 151.

clegantissimum, 170.

hirtifolium, 170.

humidorum var. pcrakcnse, 171.

humile, 170.

indicum, 174.

interruptum, 174.

italicum, 177.

kunlhianum, 164.

luzonense, 170.

malaccense, 176.

maximum, 170.

montanum. 170.

munroanum. 148.

muticum, 152.

myosuroides, 174.

myuros, 163.

nodosum, 170.

ovalifolium, 170.

var. hirtifolium, 170.

Panicum oxyphyllum, 154.

pallide-fuscum, 177.

palmifolium, 177.

paludosum, 171.

patens, 154. 155.

perakense. 171.

pilipes, 155.

protiferum, 171.

punctatum. 171.

repens, 171.

reptans, 152.

ridleyi, 148.

sarmentosum, 171.

simpliciusculum, 153.

spinescens, 173.

trigonum, 155.

tuberculatum, 170.

vicinum, 163.

zizanioidcs, 148.

Papyrius, 233.

Parartocarpus, 241.

Parashorea macrophylla, 262.

parvifolia. 264.

smythiesii. 266.

Paratrophis, 215, 216.

anthropophagorum. 221.

australina, 221.

caudata, 227.'

glabra, 221.

grandifolia. 231.

heterophylla, 221.

microphylla, 221.

niindanaensis. 227.

ostermeyri, 221.

Paratrophis philippinensis, 231

smithii. 224.

viridissima, 221.

zahlbruckneri, 221.

Paspalidium punctatum, 171.

Paspalum bicorne, 157.

cartilagineum, 171.

commersonii, 171.

var. hirsutum, 171.

conjugatum, 171.

distichum, 172.

heteranthum, 157.

longiflorum, 157.

longi folium, 172.

orbiculare. 171.

platicaulon, 149.

platycaule, 149.

platycoleum, 172.

scrobiculatum, 171.

vaginatum, 172.

Pennales, 12.

Pennisetum clandestinum. 172.

purpureum, 172.

Pcranema, 113.

inflexum, 113.

Perancmaceae, 113.

Perianth (of Moraceae). 193.

Perotis indica, 172.

latifolia, 172,

Phacus, 111.

P- ^ * ks(,'

longicauda var. brevicaudata, 111

var. indica. 111.

meson. 111.

pleuronectes, 111.

Phalansterium, 134.

digitatum, 134.

Phalaris zizanioides. 179.

Pharus urceolatus. 167.

Phleum indicum. 165.

Phormidium angustissimum, 383.

jadinianum, 375.

Phragmites communis, 172.

karka, 172.

411

Gardens Bulletin, S.

Phyllochlamys, 215. 217.

spinosa, 225.

taxoidcs, 225.

var. parvifolia, 226.

tridentata, 226.

wallichii, 225.

Pinnularia, 49.

amphicephala, 50.

appendiculata var. budensis, 49.

biceps var. minor, 49.

bogotensis var. continua, 50.

borealis, 50.

braunii, 50.

var. amphicephala. 50.

brebissonii var. hybrida. 51.

brevicostata, 51.

gibba var. interrupta, 51.

hemiptera, 51.

hybrida, 51.

interrupta var. stauroneiformis, 49.

f. minor. 49.

legumen, 51.

macilenta, 52.

mesolepta, 52.

microstauron, 52.

Pinnularia microstauron var. amb-

igua. 52.

var. brebissonii, 53.

polyonca, 53.

parva, 53.

var. parvula, 53.

parvula, 53.

stauroptera var. subparallela. 53.

stomatophoroides var. nuda. 53.

stricta, 54.

tabellaria. 54.

trigonocephala, 54.

Pipturus, 251.

Pistil lodes (of Moraceae). 194.

Plagiochlia teysmanni, 373.

Plecospermum, 241.

andamanicum, 242.

bureani. 242.

cuneifolium, 242.

laurifolium, 242.

obovatum, 242.

spinosum, 242.

Pleurococcus nacgelii, 381, 383.

Pleurosigma, 31.

angulatum var. elongatum, 31.

arcuatum, 30.

attenuatum, 29.

var. caspia, 29.

delicatulum var. salinarum, 31.

distortum, 29.

elongatum, 31.

eximium, 30.

fasciola, 30.

parkeri, 29.

salinarum, 31.

spencerii var. smithii, 31.

Poa annua, 172.

atrovirens, 159.

chinensis. 167.

japonica, 159.

panicea, 167.

pilosa, 160.

tenella. 160.

Pogonatherum crinitum, 173.

paniccum. 172.

saccharoideum var. monandra. 173.

Pogonatum junghuhnianum, 374.

microphyllum, 374.

Poikilospermum, 251.

Pollinia ciliata, 167.

var. glabrata, 167.

geniculata, 167.

gracilis, 167.

grata, 168.

hendersonii, 168.

praemorsa. 173.

ridleyi, 161.

rupestris, 166.

vagans, 168.

Polytrias amaura. 173.

Poteriodendron, 133.

petiolatum, 133.

Pourouma, 251.

Prainea, 242.

Proto-Moraceae, 203.

Prowse. G. A.: Diatoms of Malayan

Fresh waters, 1.

: Further Malayan Freshwater

Flagcllata, 105.

Pseudechinolaenia polystachya, 173.

Pseudomorus. 215. 216.

brunoniana, 222.

var. australiana. 222.

var. obtusa, 222.

var. pendulina. 222.

var. sandwicensis, 222.

var. sea bra, 222.

pendulina, 222.

sandwicensis, 222.

Pseudoraphis spinescens, 173.

Pscudostreblus, 215, 217.

caudatus, 221.

indicus, 226.

Pseudotrophis, 215, 217.

laxifiora, 227.

Racemobambos setifera, 173.

Raphis oriental's, 153.

Rhaph domonas semen, 116.

Rhizaspidaceae. 114.

Rhizaspis, 114.

granulata. 114.

simplex, 114.

Rhizosolenia, 12.

eriensis, 12.

Rhopalodia, 62.

gibberula, 62.

musculus, 63.

parallela, 63.

Rhynchclytrum repens, 173.

Riccardia pinguis, 373.

Rottboellia cancellata. 168.

foveolata, 153.

Rottboellia geminata, 168.

glandulosa, 153.

helferi, 153.

mollicoma, 168.

412

Vol. XIX. (1962).

protensa, 163.

repens, 167.

sanguinea, 175.

Saccharum arundinaceum, 174.

confertum, 164.

paniceum, 172.

ridleyi, 177.

spontaneum, 174.

Sacciolepis angusla, 174.

indica, 174.

interrupta. 174.

myosuroides, 174.

turgida, 174.

Salpingoeca, 134.

frequentissima, 135.

Scenedesmus obliquus. 375.

SchefFlera borneensis, 185.

furfuracea, 185.

petiolosa, 185.

racemosa, 185.

Schizachyrium brevifolium, 174.

sanguineum, 175.

semiberbe, 175.

Schizonema comoides, 43.

eximium, 30.

grevillei, 43.

Schizostachyum aciculare, 175.

blumei, 176.

brachycladum, 175.

Schizostachvum chilianthum 175

176.

var. auriculatum, 175.

var. erectum, 175.

dumosum, 155.

clegans, 155.

gracile, 175.

grande, 175.

hasskarlianum, 161.

jaculans, 176.

latifolium, 176.

longispiculatum, 176.

ridleyi, 176.

subcordalum. 156.

tenue, 175.

term inale, 176.

zollingeri, 176.

Sclerostachya ridleyi, 177.

Sclerostachyum ridleyi, 177.

Scytonema javanicum, 382, 383.

pseudopunctatum, 376.

Seeds (of Moraceae), 197.

Sennia, 119,

parvula, 120.

Setaria geniculata, 177.

glauca, 177.

italica, 177.

pall'de-fusca, 177.

palmifolia, 177.

plicata, 177.

rubiginosa, 177.

Shorea acuta. 268.

agami, 270.

amplcxicaulis. 273.

andulensis. 275.

Shorea angustifolia. 277.

asahi, 279.

biavak. 281.

bullata, 283.

domatiosa. 285.

fagiietioides. 287.

flaviflora. 289.

geniculata, 291.

Indiana. 295.

monticola. 297.

myrionerva, 299.

patoiensis. 302.

pilosa. ?04.

revoluta, 304.

rubella. 307.

rubra. 309.

slootenii. 312.

tomentosa, 304.

vanslooteni, 312.

Skutchia, 230.

Sloetia. 215, 218.

minahassae. 231.

pinangeana. 227.

sideroxylon, 227.

wallichii, 227.

Smithiodendron, 233.

Soil algae from New Guinea (A Short

Note on), 375.

Soil Algae of Singapore & Malava.

379.

Sorghum affine. 177.

propinquum. 177.

Soroceae, 211.

Sparattosyce. 251.

Sphaerocaryum elcgans. 176.

malaccense. 176.

Sphcnella parvulu, 61.

Spines (of Moraceae), 203.

Spinifex littorcus. 177.

squarrosus, 178.

Spodiopogon velutinus, 178.

Sporobolus berieroanus, 178.

diander, 178.

indicus, 178.

piliferus, 178.

virginicus, 178.

Stamens (of Moraceae), 193.

Stauroneis, 36.

acuta. 36.

anceps, 36.

var. goth'ca, 36.

var. hyalina. 36.

var. pusilla, 38.

franconica, 38.

microstauron. 52.

obtusa, 37.

ova I is, 47.

phoenicenteron var. brevis. 37.

var. elegans. 37.

var. halmei

f. marginestriata, 37.

var. intermedia, 37.

var. vulgaris

f. intermedia. 37.

platystoma, 46.

pusilla var. franconica, 38.

rotaeana. 47.

413

Gardens Bulletin , S.

Stenochasma, 251.

Stenopterobia, 72.

intermedia, 72.

Stenotaphrum helferi, 178.

Stephanodiscus, 10.

biserial is. 10.

fenestralis. 11.

Stipa arguens, 178.

littorea, 177.

Stomatochone, 135.

infundibuliformis, 135.

Strebleae, 211.

Streblus, 215.

sect. Rleekrodea. 216, 219.

sect. Paratrophis. 216, 220.

sect. Phyllochlaniys. 217. 225.

sect. Pseudostreblus. 217, 226.

sect. P&eudotrophis, 217, 226.

sect. SJoetia, 218, 227.

sect. Streblus. 218, 228.

sect. Taxotrophis, 218, 229.

subgen. Parastreblus, 214.

anthropophagorum. 220.

asper. var. asper, 228.

var. monoica. 228.

brunnonianus, 222.

crenatus. 226.

elongatus. 227.

glaber. 221.

var. australianus, 221.

var. glaber, 221.

heterophylliis, 221.

var. elliptica, 222.

var. heterophyl'a. 222.

ilieifolius, 227.

indicus. 226.

insignis, 219.

laevifolius, 221.

Streblus macrophyllus. 225, 227.

niadagascariensis. 220.

mitis, 229.

pendulinus, 222.

perakensis, 223.

smithii. 224.

solomonensis. 224.

spinosus, 229.

lahitensis. 225.

taxoides, 225.

var. microphylla, 225.

tonkinensis. 228.

urophyllus, 225.

var. salieifolius. 225.

var. urophyllus. 225.

zeylanicus. 229.

Strombomonas, 113.

verrucosa var. chinensis. 113.

Sukaminea, 235.

Surirella, 72.

angusta, 72.

angusticostata, 73.

biseriata, 73.

constricta, 74.

capronii, 73.

intermedia. 72.

lemmcrmanni, 73.

linearis, 73.

muelleri, 74.

navicularis, 67.

robusta, 74.

var. splend.da. 74.

spiralis, 75.

splcndida. 74.

Surirella tenera, 75.

tenuissima, 75.

Symphysodon splendens. 374.

Symploca elegans, 376, 381.

Svnechococcus cedrorum, 383.

Synedra, 16

acicularis. 65.

aflinis var. acuminata, 16.

amphirhynchus. 16.

danica, 16.

longissima var. acicularis. 16.

lunaris, 19.

pa lea, 68.

scalaris, 70.

sigma, 70.

subtilis, 71.

tabulata var. acuminata, 16.

ulna var. amphirhynchus, 16.

var. chaseana, 16.

var. danica, 16.

vaucheriae, 15.

Synura, 129.

uvella, 129.

Taxotrophis. 215, 218.

balansae, 227.

caudatae. 229.

crenata, 226.

ilieifolius, 227.

javanica, 229.

microphylla, 221.

mindanaensis, 227.

obtusa, 227.

Taxotrophis poilanei. 226.

roxburghii, 225.

sp.nosa, 229.

triapiculata. 227.

zeyianica. 229.

Teonongia, 215, 218.

tonkinensis 228.

Thalassiosira, 10.

dccipiens, 8.

fluviatilis, 10.

Themeda arguens, 178.

arundinacea. 179.

villosa, 179.

Thuarea involuta. 179.

sarmentosa, 179.

Thyrsostachys siamensis, 179.

Thysanolaena agrostis, 179.

maxima. 179.

Thysanomitrium comosum. 374.

Tolypothrix arenophila. 382.

fragilis. 382.

414

Vol. XIX. (1962).

Toxylon, 235.

Trachelomonas, 112.

allia, 112.

chinensis, 113.

var. assuriensis. 113.

var. ovata, 113.

conspersa, 113.

indica, 112.

intermedia, 112.

volvocinopsis, 112.

Treculia, 243.

Trentepohlia torulosa, 381.

Trepomonas, 137.

rotans, 137.

Tribonema bombycinum, 381.

Tricholacna rosea, 173.

Triraphis madagascariensis, 169.

Trochisia aspersa, 383.

Trophis, 230.

sect. Calpidochlamys, 230.

sect. Maillardia. 230.

sect. Trophis, 230.

subgen. Prototrophis. 230.

Trophis. 230.

aculeata, 242.

anthropophagorum, 220.

branderhorstii. 231.

drupacea. 231.

philipplnensis. 231.

spinosa, 225, 242.

taxiformis, 225.

taxoides, 225.

Tropidoneis, 63.

lepidoptera, 63.

Turbonionas. 136.

gyrans, 136.

Tryblionella constricta, 65.

punctata, 69.

Urochloa paspaloides, 152.

reptans. 152.

Uromorus, 215, 216.

anthropophagorum, 220.

philippinensis, 231.

Urtica spinosa, 229.

Urticaceae (Distinction from Mora-

ceae), 188.

Urticales (Affinity of), 208.

Vacuolaria depressa, 116.

Vanieria, 235, 237.

cochinchinensis, 239.

fruticosa, 239.

tricuspidata, 240.

triloba, 240.

Vatica coriacea. 314.

parvifolia. 316.

vinosa, 316.

Vaucheria sessilis, 381, 383.

Vetiveria odorata, 179.

zizanioides, 179.

Volvox vegetans, 136.

Whitmore, T. C.: Studies in Syste¬

matic Bark Morphology, III.

Bark Taxonomy in Diptero-

carpaceae, 321.

Xanthophyceae, 381.

Zoysia malrella, 180.

pungens, 180.

415