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“THE GARDENS’ BULLETIN

SINGAPORE

BEE

. XXXII 1980

GONTENTS

Oe Ka

PAGES

ZEDGE

Zeow Ke SS

. E. Hortrum:

The Fern-Family of Thelypteridaceae in Malaya ... Fig “a - 1

F.M LL:

ae ‘Revision of Memecylon L. (Melastomataceae) from the Malay Peninsula 31

ef a

M. YEN & G. Lim:

_ Cercospora and Alhed Genera of Singapore and the Malay Peninsula ... 151

%3 ..

EPR DR ODE

> Wind,

oe OE

KE DZE\ DAKE DE

Published by Botanic Gardens

Parks & Recreation Department

Singapore

Printed by Amsterdam Type Printers, Singapore.

eA AO es FDR ORR OOD OOO

GARDENS’ BUI

EDITORIAL BOARD.

Chairman: Y.K. Wong, B.Sc. (Hons.) (Singapore), M. AL

M.S.L. Biol.

Editor: Y.S. Choo, B.Sc. (Malaya), Ph.D. (Monash), M.S.I. |

Members: S.E. Chua, B.S.A., M.S.A. (Toronto), Ph.D. (Singapore),

M.I. Biol., M.S.I. Biol.

CH. Kee, B.Sc. (Hons.), Dip. Bus. Admin.,

Cert. Edu. (Singapore), M.S.I. Biol.

H. Keng, B.A., M.Sc. (China), Ph.D. (California),

MS.L. Biol. |

S.Y. Geh, B.Sc. (Hons.) M.Sc. (Singapore),

Dip. Hort. Sc. (New Zealand).

Secretary and G.T. Choo, B. Hort. Sc. (Hons.) New Zealand.

Business Manager:

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All correspondence concerning advertisements and exchange shoul

addressed to: — ne

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Republic of Singapore.

THE GARDENS’ BULLETIN

SINGAPORE

VOL. XXXII 1980

PAGES

R. E. HOLTTUM:

The Fern-Family of Thelypteridaceae in Malaya 1

J. F. MAXWELL:

Revision of Memecylon L. (Melastomataceae) from the Malay Peninsula

J. M. YEN! & G. Lim?:

Cercospora and Allied Genera of Singapore and the Malay Peninsula 151

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Thelypteridaceae in Malaya 1

THE FERN-FAMILY THELYPTERIDACEAE IN MALAYA

R. E. HoLtrumM

Royal Botanic Gardens

Kew

SUMMARY

The genera and species of Thelypteridaceae in Malaya are here arranged as

in a monograph of the family prepared for Flora Malesiana, Series II (Pteridophyta)

Vol. 1, part 5, which is in process of publication simultaneously with the present

paper. New names and new combinations will date from Flora Malesiana and

not from the present paper, the object of which is to indicate the necessary

corrections in Holttum, A Revised Flora of Malaya Vol. 2 (dated 1954 but

published early in 1955, second edition 1968) to which reference is made under

every species. Apart from changes in generic concepts, the principal new in-

formation concerns the species named Thelypteris vicosa, Cyclosorus stipellatus

and Cyclosorus ferox in 1955. New descriptions are only provided where those

in the book are defective.

INTRODUCTION

The family Thelypteridaceae comprises about 1000 species, about 8% of

all known ferns. About 430 species are now known in Malesia. Early descrip-

tions of species were rarely adequate to identify specimens, and several names

were used confusedly in the 19th century. As a basis for the present work, types

of almost all species were examined and re-described; all specimens in some

major herbaria have also been studied, and many recent unnamed collections.

This resulted in the discovery that some species had been wrongly named in

the book of 1955; also more critical study and new collections in Malaya indi-

cated that a few species should be subdivided. In 1971 I published a new scheme

of genera in the family, to which the Malayan species are here allocated. Sub-

sequently I published a series of monographs of most of the genera, except the

largest (Sphaerostephanos). Some taxonomists prefer to regard all species in

the family as members of the genus Thelypteris; the correct name for each species

in that genus, if published, is here indicated.

A CONSPECTUS OF THE GENERA

It was evident to me that the system of genera adopted in the book of 1955

was not a natural one; for example, the species there named Thelypteris immersa

and Cyclosorus extensus are closely allied and should not be in different genera;

similarly, the species named Abacopteris lineata is closely related to Cyclosorus

glandulosus. Within both Thelypteris and Cyclosorus is a mixture of species of

diverse relationships. Ching established some new genera in 1963, but some of

them are wholly or mainly in mainland Asia, and in Malesia there are very

numerous additional species which need to be fitted into the picture. My treat-

ment was thus more elaborate than Ching’s. The following conspectus shows

how I think the genera are related. It is not easy to show this in a way that

will also serve as a key by which specimens may be allocated to their correct

genera; an artificial key for this purpose (covering Malayan genera only) follows.

The inter-relations of the groups are still very uncertain, and need to be con-

sidered in the context of the whole family which is world-wide in distribution

with the greatest abundance of species in the wet tropics.

2 Gardens’ Bulletin, Singapore XX XIII (1980)

1. Group of Phegopteris (Holttum, Gard. Bull. Singap. 29: 145, with key to

species. > 1977).

Fronds bipinnate or pinnate with deeply lobed pinnae; veins often branching,

not reaching margins of leaflets; upper surface of costae prominent, not grooved;

indusium present or absent; chromosome numbers 30, 31, 35.

Genera: (Phegotperis, not in Malaya), Pseudophegopteris, Macrothelypteris,

Metathelypteris.

2. Group of Coryphopteris.

Fronds pinnate with deeply lobed pinnae and simple free veins which all

reach the margin; sporangia short-stalked, lacking hairs or glands distally; indusia

present; small spherical resinous glands usually present on lower surface of pinnae;

some acicular hairs in some species consisting of several cells; caudex erect

(Coryphopteris), or slender and long-creeping (Parathelypteris). Chromosome

number 31 or 32.

3. Group of Trigonospora.

- Caudex erect; fronds pinnate with deeply lobed pinnae and free veins (except

one species in Burma); indusia present; sporangia lacking glands or hairs distally;

spores trilete with discontinuous perispore of minute papillae; plants of rocky

stream-beds; chromosome number 36.

Genera: Trigonospora (Asia and W. Malesia), Menisorus (Africa).

4. Group of Thelypteris.

Caudex prostrate, short or long; fronds simply pinnate with lobed pinnae;

no reduced basal pinnae; veins free or anastomosing, simple except in Thelypteris,

all reaching the margin; persistent Seales, often broad, preserit on lower surface

of costae; sori indusiate or not; sporangia lacking glands or hairs distally but

bearing a large spherical red gland at the end of a hair on sporangium-stalk,

similar glands sometimes present on lower surface of pinnae; chromosome number

36 (35 in Thelypteris).

Genera: Thelypteris (not Malayan), Cyclosorus, Ampelopteris, Mesophlebion.

5. Group of Chingia.

Caudex massive, erect (sometimes short-creeping in Plesioneuron); long narrow

scales at base of stipe, often throughout stipe and rachis; fronds pinnate, pinnae

variously lobed; no reduced basal pinnae; veins in Chingia always oblique, several

pairs passing to the sides of a long sinus-membrane which is always prominent

on the lower surface; sori close to costules in Chingia, often exindusiate; sporangia

often bearing small capitate hairs or small spherical glands; chromosome number

36.

Genera: Chingia Plesioneuron (in Eastern Malesia and the Pacific).

6. Group of Sphaerostephanos.

Caudex erect or short-creeping (long-creeping in S. unitus only); fronds simply

pinnate, pinnae deeply lobed to entire; reduced pinnae present, sometimes very

Thelypteridaceae in Malaya 3

small with long aerophores, except Pronephrium; veins anastomosing in the less

deeply lobed species, sinus membranes usually distinct, short or long; sessile

spherical glands or small capitate hairs present on sporangia of many species

and often also on surface of pinnae; spores with many separate small thin wings

or with a median translucent wing and cross-wings; chromosome number 36.

Genera: Pneumatopteris, Sphaerostephanos, Pronephrium.

7. Group of Christella.

Caudex erect or short- to long-creeping; fronds simply pinnate; pinnae deeply

to shallowly lobed; lower pinnae in Christella gradually reduced but the basal

ones never very small; swollen aerophores lacking or rare; glandular hairs on

pinnae, if present, + elongate; sporangia lacking glands or hairs distally, in

Christella always with an elongate unicellular gland on the stalk; perispore con-

sisting of rather thick tubercles or ridges; chromosome number 36.

Genera: Christella, Amphineuron.

KEY TO THE GENERA

1. Upper surface of costae not grooved; veins not reaching margin

2. Fronds bipinnate

3. Slender septate acicular hairs present on lower surface of axes of frond

| SRS os Se ae ee eke ee ae 1. Macrothelypteris

3. Acicular hairs on axes of frond all uicellular ... 2. Pseudophegopteris

2. Fronds simply pinnate

a peeaieia bearing, -sctac s.34i..2tte. JAS. 2. Pseudophegopteris

Seporaneia, lacking- Sctae ..2... fics. ccs ced eee ste ee 3. Metathelypteris

1. Upper surface of costae grooved; veins all reaching margin

5. Much-reduced basal pinnae lacking, or 1-2 pairs inconstantly present or

absent

6. Veins all free

i Caudex tstender,.long-creeping | ii.%.....4.....6.0.5- 4. Parathelypteris

7. Caudex erect or short-creeping, not slender

8. Caudex quite erect; basal basiscopic vein of each group arising

from costule, not from costa

9. Fronds commonly more than 100 cm long; copious minute

yellow glands on lower surface of veins ... 15. Amphineuron

9. Fronds rarely over 50 cm long; glands on lower surface, if

present, otherwise

10. Plants of mountain ridges; resinous glands usually pre-

sent on lower surface of pinnae ...... 5. Coryphopteris

Gardens’ Bulletin, Singapore XX XIII (1980)

10. Plants of rocky stream-beds; no glands present ............

alee. LEM. AOSRG TR, See 6. Trigonospora

8. Caudex short-creeping; basal basiscopic vein of each group arising

from: costa, not: from:.costulesiis-.iae eee 7. Mesophlebion

6. Veins anastomosing

11. Plants of wet places; caudex long-creeping; flat scales present on

lower surface of costae, at least on young fronds

12. Plants of open swamps; fronds not proliferous; indusia present

« SMBS AGS oc Nee Qf Ru SOL a 8. Cyclosorus

12. Plants of banks of streams or ditches; fronds freely proliferous;

Indusia Tacks... x. sasmeines ces anol mee en: 9. Ampelopteris

11. Plants not growing in wet places; caudex erect or short-creeping;

no flat scales on lower surface of costae

13. Caudex erect; stiff narrow scales at base of stipe; sori close

to costules; a long prominent sinus-membrane present .........

be cdasodvesedeudens ot oy uGelMlOig of Ut aW Rte Raven ae 10. Chingia

13. Caudex short-creeping; scales otherwise; sori rarely close to

costules; no long sinus-membrane prominent on lower surface

14, Pinnagss cremate. Werk ae eee 13. Pronephrium

14. Pinnae distinctly lobed

15. In larger pinnae, at least 3 pairs of veins anastomosing

or passing to sides of sinus-membrane; a few spheri-

cal yellow glands present on lower surface of costules

and Veines, 4) seh gen le 12. Sphaerostephanos

15. In larger pinnae, at most 2 pairs of veins thus; glands,

if present, otherwise

16. Copious hairs 1 mm long on lower surface;

elongate red glands present on lower surface of

W BLINS. 1 .5'an oc Saks deca aroha ort no etaale' onraeie 14. Christella

16. Most hairs much shorter; glands on veins usually

abundant, small and colourless or minute and

yellow 2.02, 7o By eae ene 15. Amphineuron

5. Much-reduced basal pinnae present; transition from normal to reduced

pinnae abrupt or gradual

17. Spherical yellow glands present, at least on indusia or sporangia, in

most species throughout lower surface of pinnae .......................405

Livnevel oh lanes qltabit. 3s Pe eRs eee 12. Sphaerostephanos

17. Spherical yellow glands lacking r |

18. A few pairs of lower pinnae gradually reduced ... 14. Christella

18. An abrupt transition from normal to reduced pinnae at base of

frond ....2230.5. 2h Shee ee 11. Pneumatopteris

Thelypteridaceae in Malaya 5

1. MACROTHELYPTERIS CHING

1. Macrothelypteris torresiana (Gaud.) Ching; Holttum, Gard. Bull. Singap. 29

(1977) 148.

Basionym: Polystichum torresianum Gaud. in Freycinet Voy. Bot. (1824) 333.

Name in Holttum 1955: Thelypteris uliginosa (Kunze) Ching, p. 241.

Correct name in Thelypteris: T. torresiana (Gaud.) Alston, Lilloa 30 (1960) 111.

Status in Malaya: as Holttum 1955.

2. PSEUDOPHEGOPTERIS CHING

1. Pseudophegopteris rectangularis (Zoll.) Holttum, Blumea 17 (1969) 19; Gard.

Bull. Singap. 29 (1977) 149.

Basionym: Polypodium rectangulare Zoll., Syst. Verz. (1854) 37, 48.

Name in Holttum 1955: Thelypteris oppositipinna (v.A.v.R.) Ching, p. 239.

Correct name in Thelypteris: T. rectangularis (Zoll.) Nayar & Kaur, Comp. to

Beddome (1974) 72.

Status in Malaya: as Holttum 1955.

2. Pseudophegopteris paludosa (BI.) Ching; Holttum, Gard. Bull. Singap. 29

(1977) 149.

Basionym: Polypodium paludosum Bl., Fl. Jav. Fil. (1851) 192.

Name in Holitum 1955: Thelypteris brunnea (Wall.) Ching; p. 240.

Correct name in Thelypteris: T. paludosum (Bl.) K. Iwats., Acta Phytotax.

Geobot. 19 (1961) 11.

Status in Malaya, with note on synonym: as Holttum 1977, l.c.

3. METATHELYPTERIS CHING

1. Metathelypteris flaccida (Bl.) Ching; Holttum, Gard. Bull. Singap. 29 (1977)

147.

Basionym: Aspidium flaccidum Bl., Enum., Pl. Jav. (1828) 161.

Name in Holttum 1955: _ lacking.

Correct name in Thelypteris: T. flaccida (Bl.) Ching, Bull. Fan Mem. Inst. Biol.

Bot. 6 (1936) 336.

Status in Malaya: increasing. on bare earth banks by roadsides in sheltered

places at Cameron Highlands, c. 1500 m. See notes on the two forms of

this species in Holttum 1977.

Distribution: W. Java, Sumatra (?), Ceylon & S. India, N.E. India to Yunnan.

6 Gardens’ Bulletin, Singapore XX XIII (1980)

2. Metathelypteris gracilescens (Bl.) Ching, Acta Phytotax. Sinica 8 (1963) 305.

Basionym: Aspidium gracilescens Bl., Enum. Pl. Jav. (1828) 155.

Name in Holttum 1955: _ lacking.

Correct name in Thelypteris: T. gracilescens (Bl.) Ching, Bull. Fan. Mem. Inst.

Biol. Bot.°6 (1936) 327.

Misidentified as M. decipiens (Clarke) Ching in Holttum 1977: 147.

Status in Malaya: known only from a single collection from G. Batu Brinchang.

This species occurs in Mainland Asia from the Darjeeeling district in N.

Bengal to southern Japan, and has been found in Malesia on mountains at about

1800 m from Sumatra to New Guinea, usually (as on G. Batu Brinchang) on

wet rocks near a waterfall. M. decipiens (at Darjeeling) differs from Malesian

specimens of M. gracilescens in having shorter fronds (those of the latter in

Malesia commonly 25-30 cm long) with fewer pinnae, the basal pinnae largest,

and veins mostly forked. The Malayan specimen (Molesworth Allen 5005) has

more forked veins than is usual in Malesian specimens but otherwise does not

differ. There are not many places in Malaya where wet rocks by waterfalls

occur at 1800 m.

3. Metathelypteris dayi (Bedd.) Holttum in Nayar & Kaur, Comp. to Beddome

(1974) 205; Holttum, Gard. Bull. Singap. 29 (1977) 148.

Basionym: Nephrodium dayi Bedd., Journ. Bot. 25 (1877) 323.

Name in Holttum 1955: Thelypteris singalanensis (misidentification), p. 243.

Correct name in Thelypteris: T. dayi (Bedd.) Nayar & Kaur, Comp. to Beddome

(1974) 59.

Status in Malaya: as Holttum 1955.

Distribution: throughout Malesia except E. Java and Lesser Sunda Islands.

4. PARATHELYPTERIS CHING, ACTA PHYTOTAX. SINICA 8 (1963) 300.

The type of this genus is P. glanduligera (Kunze) Ching, a species based

on a specimen collected near Canton. The lower pinnae of its fronds are not

reduced. It appears to be related to Coryphopteris but has along slender creep-

ing rhizome. Ching has also included in this genus two species with lower pinnae

gradually much reduced: the Malayan P. beddomei and P. nipponia (Fr. & Sav.)

Ching from China and Japan. There are two species in North America, one

like P. beddomei in habit, the other like P. glanduligera. I think that two genera

are here confused; comparative studies of plants from Asia and N. America

are needed to resolve this situation.

1. Parathelypteris beddomei (Bak.) Ching, Acta Phytotax. Sinica 8 (1963) 302.

Basionym: Nephrodium beddomei Bak., Syn. Fil. (1867) 267.

Correct name in Thelypteris and name in Holttum 1955, p. 240: T. beddomei

(Bak.) Ching. |

Status in Malaya: now frequent at Cameron Highlands by roadsides which are

sheltered by forest, in places where there is a seepage of water.

Thelypteridaceae in Malaya 7

5. CORYPHOPTERIS HOLTTUM, BLUMEA 19 (1971) 33; 23 (1976) 18-47.

The type of this genus is C. viscosa (Bak.) Holttum, of which the type speci-

men was collected near the top of Mt Ophir by Hugh Cuming in 1839. The

genus comprises about 47 species almost all of which occur in Malesia, on

mountain ridges where the soil is peaty and very acid. All species have an

erect caudex and free veins, the basal pinnae not or little reduced: almost all

have red resinous glands on one or both surfaces of pinnae (not easy to see on

dried specimens). In 1955 I treated Thelypteris viscosa very broadly, mentioning

variation among the specimens so named; here they are divided into four species.

KEY TO THE SPECIES IN MALAYA

1. Base of stipe bearing pale, firmly cylindrical septate hairs

2. Stipe and abaxial surface of rachis densely covered with spreading hairs

many of which are septate

Saeeitinac 701 Gk a. 8 CIN st oo hota cee 1. C. unidentata

Perm 925 60 17 (eM ik feed. ci Ro eS 2. C. tahanensis

2. Stipe above base and abaxial surface of rachis less densely hairy with

ROM het ENE IM Soc 0 glee l'on, 5 «Sw OSA gametes Gop iwmuee staat 3. C. hirtipes

1. Base of stipe not bearing such hairs

4. Sessile glands present between veins on upper surface of pinnae

5. Stipe-scales thin, less than 1 mm wide above base; lower surface

of costae bearing many acicular hairs .................. 4. C. viscosa

5. Stipe-scales firm, 1 mm or more wide; lower surface of costae

Dearne short capitate “hairs: ...2..i0ss02.)..54.-.0090s 5. C. gymnopoda

6. Upper surface of pinnae lacking hairs between veins ............

OS at LS EE ORES Na an es PES ROE PT PELE var. gymnopoda

6. Upper surface of pinnae bearing acicular hairs between veins

7. Lower surface between veins, and indusia, lacking acicular

ee Saye cote 28 Canes ns Bag ane vnsg ts var. bintangensis

7. Lower surface between veins bearing copious short erect

TT kT Un let eel A Pea Te mS A var. humilis

4. Sessile glands lacking between veins on upper surface of pinnae

8. Acicular hairs present on lower surface of costae; glands present,

at least on costae

9. Septate hairs present on upper surface of rachis and costae

10. Septate hairs on upper surface of rachis and costae less

than 0.5 mm long; stipe-scales 3 mm long, thin ............

a Oia ee PCE Cath ak bt ore rs nat cele rawness 6. C. arthrotricha

8 Gardens’ Bulletin, Singapore XX XIII (1980)

10. Septate hairs on upper surface of rachis and costae 1 mm

or more long; stipe-scales to 7 x 1 mm, rigid, hair-pointed

OD. ATE EIT, LAS Ee 2. C. tahanensis

9. Septate hairs absent on upper surface of rachis and costae ......

hs. cee. aes. ea hee ee 7. C. pectiniformis

11. Hairs on lower surface of costae unicellular, less than

L mata TOMS yids neg bn teh vse Poke ee var. pectiniformis

11. Hairs on lower surface of costae septate, to 1.5 mm long

ncenelrve nods dude dene) sedge) th cilia ete ie tne an var. hirsuta

8. Acicular hairs lacking or rare on lower surface of costae; no glands

present -on lower. surfaces. .iges52305, ue. dee 8. C. badia

1. Coryphopteris unidentata (Bedd.) Holttum, Blumea 23 (1976) 26.

Basionym: Lastrea unidentata Bedd., Handb. Suppl. (1892) 53.

Name in Holttum 1955, p. 251, correct in Thelypteris: T. unidentata (Bedd.)

Holttum.

Status in Malaya: known only from G. Bubu, G. Bintang and G. Inas in Perak

(not found at Cameron Highlands, wrongly so reported in Holttum 1955).

2. Coryphopteris tahanensis Holttum, Blumea 23 (1976) 33.

Name in Holttum 1955: Thelypteris viscosa, in patt.

Correct name in Thelypteris: none published.

Differs from C. viscosa: scales at base of stipe 7-8 x 1 mm, rigid, acuminate;

glands lacking on upper surface of pinnae; upper surface of rachis and costae,

at least near apex of frond, bearing septate hairs 1 mm long, multicellular hairs

1.5 mm long sometimes also present on lower surface. One specimen from G.

Tahan has the lower surface of rachis and costae covered with hairs almost

2 mm long.

Three collections from c. 1800 m on G. Tahan, one from G. Batu Brinchang

and one from G. Lari Tembakau.

3. Coryphopteris hirsutipes (Clarke) Holttum in Nayar & Kaur, Comp. to

Beddome (1974) 203.

Basionym: Nephrodium gracilescens var. hirsutipes Clarke, Trans. Linn. Soc.

II Bot. 1 (1880) 514, t. 67.

Name in Holttum 1955: Thelypteris herbacea Holttum, p. 254.

Correct name in Thelypteris: T. hirsutipes (Clarke) Ching, Bull. Fan Mem.

Inst. Biol. Bot. 6 (1936) 314.

Base of stipe bearing spreading septate hairs 1-3 mm long; some septate

hairs also present on upper surface of rachis and costae; hairs on lower surface

very variable, sometimes all unicellular; glands present on lower surface of

Sumatran specimens, not on Malayan ones.

Thelypteridaceae in Malaya 9

In Malaya, known from three collections: G. Tahan, near Wray’s Camp

(915 m); G. Padang, Trengganu: G. Hijau at 1370 m, “locally abundant”

(Molesworth Allen).

4. Coryphopteris viscosa (Bak.) Holttum, Blumea 19 (1971) 33.

Basionym: Nephrodium viscosum Bak. Syn. Fil. (1867) 264.

Name in Holttum 1955: Thelypteris viscosa (Bak.) Ching, p. 252, in part.

Status in Malaya: only known with certainty from Mt Ophir.

Stipe-scales c. 4 x 0.5 mm, thin; lamina to 30 cm long, tapering gradually

distally; pinnae 25 pairs or more, closely placed: basal pinnae to 1.4 cm wide,

narrowed towards their bases; largest pinnae 5.5 x 1.2 cm, lobes nearly entire;

glands present on both surfaces between veins; acicular hairs present on lower

surface of rachis and costae, also very narrow scales; distal sori often +

asymmetric.

There are apparently distinct varieties of this species in Western Sarawak.

Some specimens from the Main Range in Malaya which I have named C. gymno-

poda are perhaphs intermediate between that species and typicai C. viscosa.

5. Coryphopteris gymnopoda (Bak.) Holttum, Blumea 23 (1976) 29.

Basionym: Nephrodium gymnopodum Bak., Trans. Linn. Soc. II Bot. 4 (1894)

ponte

Name in Holttum 1955: Thelypteris viscosa, in part.

Correct name in Thelypteris:none published.

Synonyms: JLastrea ridleyi Bedd. and L. robinsonii Ridl.

Var. gymnopoda differs from C. viscosa as follows: stipe-scales firm, dark,

1-1.5 mm wide; pinnae 15-18 (rarely to 25) pairs, well spaced: pinna-lobes mostly

crenate: lower surface of costae bearing capitate hairs and glands, acicular hairs

usually lacking; distal sori rarely asymmetric.

Status in Malaya: found on G. Tahan and at scattered localities on the

Main Range, especially in open places. The type came from Mt Kinabalu, where

this species is abundant at 1800 m and higher. The types of Lastrea ridleyi and

L. robinsonii are small plants.

Var. bintangensis Holttum, Blumea 23 (1976) 30. Upper surface between

veins covered with very short acicular hairs in addition to glands.

Status in Malaya: only known from the type collection on G. Bintang.

Var. humilis Holttum, Blumea 23 (1976) 30. Fronds small; both upper and

lower surfaces between veins covered with short acicular hairs.

Status in Malaya: several collections from exposed places at Cameron High-

lands at about 1500 m; their condition is possibly due to exposure; more study

is needed.

10 | Gardens’ Bulletin, Singapore XXXIII (1980)

6. Coryphopteris arthrotricha Holttum Blumea 23 (1976) 33.

Name in Holttum 1955: Thelypteris viscosa, in part.

Correct name in Thelypteris: none published.

Stipe dark at base, paler distally; scales at base of stipe c. 3 mm long, thin;

lamina 25-45 cm long; pinnae c. 20 pairs, well spaced, commonly 8 x 1.6 cm;

pinnalobes entire to dentate; lower surface of rachis and costae bearing minute

acicular and capitate hairs, on costae also red glands and linear scales; few glands

between veins; hairs on upper surface of rachis and costae hardly 0.5 mm long

but many of them septate (2-4 cells), upper surface between veins glabrous or

with a few short hairs, not glands.

Status in Malaya: this is the common species on the Main Range, in ridge

forest at 1200-1500 m; known also from Sumatra.

7. Coryphopteris pectiniformis (C. Chr.) Holttum Blumea 23 (1976) 34.

Basionym: Dryopteris pectiniformis C. Chr. Gard. Bull. Str. Settl. 4 (1929) 379.

Name in Holttum 1955, p. 253, and correct name in Thelypteris: T. pectiniformis

(C. Chr.) Ching.

Status in Malaya: in forest on Taiping Hills at 1250-1400 m; a few records

from the Main Range and G. Padang in Trengganu.

Var. hirsuta Holttum, Blumea 23 (1976) 34. Differs from the typical form

of the species as follows: fronds smaller with pinnae to 6.5 cm long; lower

surface of costules and veins bearing acicular hairs 1.5 mm long which are

septate; hairs on indusia 0.5 mm long.

Status in Malaya: only known from the ridge connecting Fraser’s Hill to

Pine... Trees Hill,

8. Coryphopteris badia (v.A.v.R.) Holttum, Blumea 23 (1976) 44.

Basionym: Dryopteris badia v.A.v.R., Bull. Jard. Bot. Btzg IH, 16 (1914) 9.

Correct name in Thelypteris: 7. badia (v.A.v.R.) Ching, Bull. Fan Mem. Inst.

Biol. Bot. 10 (1941) 250.

Stipe dark, glossy, at base often bearing a tangled mass of slender hairs

which are golden brown when dry; lamina of frond variable according to habitat,

10-65 cm long; pinnae thick and rigid when dry, nearly all distinctly stalked,

basal acroscopic lobe free or nearly so, lower surface quite glabrous apart from

a few hair-like scales on costae, no glands; upper surface of costae bearing rigid

dark brown hairs, no other hairs on upper surface and no glands; indusia glabrous.

Status in Malaya: known only from one small plant found on G. Ulu Kali

at 1800 m by Mrs A.G. Piggott; other collections from N. Sumatra, Sarawak,

Sabah. Sulawesi and New Guinea, at 1400-2500 m, usually growing in moss-

cushions, sometimes on branches of trees in moss-forest. The largest known

fronds are from an epiphytic plant collected in N. Sumatra.

mn #F

Thelypteridaceae in Malaya 11

6. TRIGONOSPORA HOLTTUM, BLUMEA 19 (1971) 29.

Plants of rocky stream-beds; caudex short, erect; basal pinnae not reduced;

veins free; sori indusiate; sporangia lacking glands or hairs; spores trilete.

This is a genus of about 10 species, mostly in mainland S.E. Asia and Ceylon.

1. Trigonospora ciliata (Benth.) Holttum, Blumea 19 (1971) 29.

Basionym: Aspidium ciliatum Benth., Fl. Hongkong. (1861) 455.

Name in Holttum 1955, p. 250, correct in Thelypteris: T. ciliata (Benth.) Ching.

Correction to Holttum 1955. The type of this species is a specimen from

Hong Kong described by Bentham, who adopted a name given by Wallich to

a plant from Nepal.

Status in Malaya: on rocks in stream-beds in the flood-zone, at many places

on the Main Range, also Taiping Hills and G. Tahan, from low altitudes to

1500 m; formerly abundant above the Parit Falls at Cameron Highlands.

7. MESOPHLEBION HOLTTUM, BLUMEA 19 (1971) 29,

EXCL. SUBG. PLESIONEURON.

This group of species is described in Holttum 1955 as the group of Thelypteris

crassifolia. Apart from changes in nomenclature, the only difference in the pre-

sent treatment is recognition of M. beccarianum as a species distinct from M.

chlamydophorum. This is an interesting genus which would be worth a detailed

cytotaxonomic study. As reported in 1955, two Malayan species are tetraploid;

a diploid has recently been found in Sarawak (a new species, not known in

Malaya).

KEY TO THE SPECIES IN MALAYA

1. Stipe and lower part of rachis bearing many stiff spreading scales ............

oe eae cada Sp Se panwcdinnn coeaen's 1. M. trichopodum

1. Upper part of stipe and rachis lacking large scales

2. Sterile pinnae lobed not more than half-way to costa .....................2..

WE i Saat 2 ae Re ee, Pape eee eee ree ore 2. -M. motleyanum

2. Sterile pinnae lobed more than half-way to costa

3. Indusia firm, glabrous, covering the sorus almost to maturity; fertile

pinnae lobed to 1-1.5 mm from costa ............ 3. M. beccarianum

3. Indusia + hairy, in most cases not covering the sori to maturity;

pinnae lobed less deeply

4. Fronds rarely dimorphic; pinnae thin, rarely more than 2 cm

wide; indusia always conspicuous ...... 4. M. chlamydophorum

4. Fronds usually dimorphic; pinnae rigid, fertile to 3 cm, sterile

to 4 cm wide; indusia varied, thin and shrivelled when dried

RE eC, eA vive una ee sabisten desea’ 5. M. crassifolium

12 Gardens’ Bulletin, Singapore XX XIII (1980)

1. Mesophlebion trichopodum (C. Chr.) Holttum, Blumea 72 (1975) 226.

Basionym: Dryopteris trichopoda C. Chr., Ind. Fil. (1905) 298, new name for

Nephrodium polytrichum Bak., Journ. Bot. 29 (1891) 107, non Schrad. 1824.

Name in Holttum 1955: Thelypteris paleata (Copel.) Holttum, p. 249.

Correct name in Thelypteris: none published.

The description of 1955 needes no revision for Malayan plants. These are

smaller than the type specimen of N. trichopodum from Sarawak but otherwise

not different. More collections are needed from Sarawak and Sumatra; possibly

distinct varieties exist in Sarawak.

In Malaya known from few places, mostly beside streams in forest: Penang

Hill at 650 m (Richmond Pool); Taiping Hills; by the Sungei Tahan at K. Teku,

50 m; by the Sungei Jeriau below Fraser’s Hill at 1070 m; on G. Angsi at 600 m.

2. Mesophlebion motleyanum (Hook.) Holttum in Nayar & Kaur, Comp. to

Beddome (1974) 209.

Basionym: Nephrodium motleyanum Hook., Syn. Fil. (1867) 266.

Name in Holttum 1955, correct in Thelypteris: T. motleyanum (Hook.) Holttum,

p. 247.

Status in Malaya: as in Holttum 1955 for lowland plants, but I now think that

mountain plants, smaller and firmer in texture than lowland ones (mentioned

under T. crassifolia in 1955) probably belong to this species; more study is

needed.

3. Mesophlebion beccarianum (Cesati) Holttum, Blumea 22 (1975) 230.

Basionym: Nephrodium beccarianum Cesati, Atti Acad. Napoli 7, no 8 (1876) 23.

Name in Holttum 1955: Thelypteris chlamydophora, in part, p. 246.

Correct name in Thelypteris: T. beccariana (Cesati) Reed, Phytologia 17 (1968)

263.

Fronds firm; lower pinnae with stalks 6-10 mm long; pinnae 12-18 cm long,

1.3-2.5 cm wide (sterile ones largest), lobed to 1-1.5 mm from costa; basal

basiscopic vein arising from costa near its costule; lower surface of costae usually

hairless but with some scales which are early caduous; sori inframedial; indusia

large, firm, glabrous, covering the sori almost to maturity; spores papillose.

Status in Malaya: specimens from Ulu Langat, in hill forest at 750 m, and

from G. Angsi, are closely similar to the type and other Bornean specimens;

some specimens from Perak and Penang differing in having some short acicular

hairs on the lower surface of costae and on indusia may be hybrids. In Sarawak,

the species seems to be very uniform.

4. Mesophlebion chlamydophorum (C. Chr.) Holttum, Blumea 22 (1975) 321.

Basionvm: Dryopteris chlamydophora C. Chr., Gard. Bull. Str. Settl. 4 (1929)

to) BB4.

Thelypteridaceae in Malaya 13

Name in Holttum 1955, correct in Thelypteris: T. chlamydophora (C. Chr.)

Ching, p. 246.

Status in Malaya: as in Holttum 1955.

The spores are papillose, as in M. beccarianum.

5. Mesophlebion crassifolium (BI.) Holttum, Blumea 22 (1975) 232.

Basionym: Aspidium crassifolium Bl., Enum. Pl. Jav. (1828) 158.

Name in Holttum 1955, correct in Thelypteris: T. crassifolia (Bl.) Ching, p. 246.

In mountain forest in Malaya at 700-1800 m, variable; pinnae of sterile

fronds are usually much wider than those of fertile fronds; stalks of lower pinnae

very variable in length; indusia thin, with short hairs, sometimes rather large,

sometimes small; spores with small wings, as in M. motleyanum. There may

be hybrids between this and M. motleyanum and M. beccarianum; one specimen

from Fraser’s Hill has spores intermediate between Nari and papillose, and

rather large, thin, short-hairy indusia.

8. CYCLOSORUS LINK, HORT. REG. BOT. BEROL. 2 (1833) 128.

A genus of few species, pantropic, always in open places in freshwater

swamps. A characteristic feature is the presence of a large red glandular cell at

the end of a hair on the stalks of sporangia; similar glandular cells also occur

on the lower surface of pinnae.

1. Cyclosorus interruptus (Willd.) H. Ito, Bot. Mag. Tokyo 51 (1937) 714,

nomen tantum.

Basionym: Prteris interrupta Willd., Phytographia (1794) 13, t. 10, fig. 1.

Name in Holttum 1955: Cyclosorus gongylodes (Schkuhr) Link, p. 261.

Correct name in Thelypteris: T. interrupta (Willd.) K. Iwats., J. Jap. Bot. 38

(1963) 314 nomen tantum.

Status in Malaya: as Holttum 1955.

The type of this species, in the Willdenow Herbarium at Berlin, was collected

in Southern India. The species has had many names; see Holttum, Amer. Fern

Journ. 63 (1973) 81. Unfortunately Willdenow’s type has in recent years been

wrongly identified with the species here named Amphineuron terminans.

9. AMPELOPTERIS KUNZE, BOT. ZEIT. 6 (1848) 114.

This is a monotypic genus, occurring throughout the wetter parts of the

tropics from West Africa to N.E. Australia and New Caledonia.

1. Ampelopteris prolifera (Retz.) Copel., Gen. Fil. (1947) 144.

Basionym: Hemionitis prolifera Retz., Obs. Bot. 6 (1791) 38.

Name in Holttum 1955: Ampelopteris prolifera (Retz.) Copel., p. 298.

Correct name in Thelypteris: T. prolifera (Retz.) Reed, Phytologia 17 (1968)

306.

14 Gardens’ Bulletin, Singapore XX XIII (1980)

Sori exindusiate; between the sporangia are conspicuous red glands which

are borne at the ends of hairs on sporangium-stalks, as in Cyclosorus and Mesop-

hlebion; broad scales present on the lower surface of costae of young fronds,

soon lost; forked or branched hairs, often sparse, present on lower surface of

costae and rachis; see also description in Holttum 1955.

Status in Malaya: as in Holttum 1955; see also Molesworth Allen, Gard.

Bull. Singapore 17 (1959) 261 for reports of this species by the Muda and

Kinta rivers. There have been few collections in other parts of Malaya; condi-

tions necessary for establishment of growth of prothalli and sporophytes need

to be studied.

10. CHINGIA HOLTTUM, BLUMEA 19 (1971) 31

Caudex erect, usually massive on well-grown plants; stipes covered, at least

near base, with long narrow scales; basal pinnae not reduced; pinnae lobed up

to half-way to costa; veins all oblique, 1-2 pairs anastomosing and usually several

pairs passing to the sinus-membranes which are prominent on the lower surface;

sori near costules; indusia small or lacking; sporangia sometimes bearing capitate

hairs or small glands; spores black, minutely tuberculate.

A genus confined to Malesia and the Pacific (to Tahiti); 18 species known.

Corrections are here made to the description of the previously-known Malayan

species, and an additional one is described.

KEY TO THE MALAYAN SPECIES

1. Stipe and at least the lower part of abaxial surface of rachis bearing copious

stiff scales or their persistent bases; pinnae thin, bearing variously abundant

small sessile glands on their lower surface ..................... 1. C. sakayensis

1. Stipe scaly near base only; pinnae rigid; more or less abundant short capitate

hairs present on lower gurlace ..d.7i.1. iced ee aes 2. C. perrigida

1. Chingia sakayensis (Zeiller) Holttum, Flora Malesiana.

Basionym: Nephrodium sakayense Zeiller.

Name in Holttum 1955: Cyclosorus ferox, p. 265.

Correct name in Thelypteris: T. sakayensis (Zeiller) Reed, Phytologia 17 (1968)

314.

The type of this species, at Paris, is the upper part of a frond of a young

plant, partly fertile; it was collected near Ipoh. In 1955 I had not seen it,

and made no reference to the species. A complete young plant collected by

Mrs A. G. Piggott on G. Telapak Burok, Negri Sembilan, along with mature

plants, closely matches Zeiller’s type. As noted in 1955, Malayan plants named

Cyclosorus ferox differ in several ways from the type of Aspidium ferox Bl. from

Java. Important differences are the distinctly flat scales on the rachis, the pre-

sence of small indusia and of copious small sessile glands on the lower surface

of pinnae between veins.

Thelypteridaceae in Malaya 15

2. Chingia perrigida (v.A.v.R.) Holttum, Kalikasan 3 (1974) 27.

Basionym: Dryopteris perrigida v.A.v.R., Bull. Jard. Bot. Btzg IH, 10 (1914) 27.

Correct name in Thelypteris: none published.

Plants smaller than those of C. sakayensis, with rigid fronds having veins

prominent on the lower surface; stipe-scales abundant near base only, no scales

on rachis; pinnae to 30 x 2.3 cm, lobed 1/3 towards costa or a little more

deeply; lower surface of pinnae bearing short capitate hairs on all parts and

a variable number of acicular hairs on costules and veins; sori with small indusia.

Status in Malaya: only known from open places by roadsides on G. Ulu

Kali (Genting Highlands) at 1500-1700 m, first collected by Mrs A. G. Piggott

in 1974. The type of the species was collected on G. Merapi, near Bukit Tinggi,

in Sumatra.

11. PNEUMATOPTERIS NAKAI, BOT. MAG. TOKYO 47 (1933) 179.

A genus of about 80 species, confined to the tropics of the Old World apart

from one aberrant species in New Zealand; mainly forest plants, a few confined

to limestone; differing from Sphaerostephanos in lacking spherical yellow glands,

in lacking also abundant acicular hairs on lower surfaces; small capitate hairs

are present on lower surfaces and/or sporangia of some species; the lower surface

of dried specimens, between veins, is + pustular; aerophores are always some-

what swollen, much elongate in a few species, including the type, P. callosa.

KEY TO THE MALAYAN SPECIES

1. Reduced pinnae at base of frond all distinctly leafy

2. Reduced pinnae 6-10 pairs, the upper ones not auricled ..................

eet E Das gr nce coho e lig, STILL seks “RVs sk SEK L So os 1. P. truncata

2. Reduced pinnae 2-3 (-4) pairs, strongly auricled on their acroscopic

Se Sela ea 3 NE a Ra ihe onde ahem he 2. P. ecallosa

1. Reduced pinnae represented by prominent aerophores, each with a very

Ram MRE att NR STIG as 7 acs ae ot gis aug oness ors ance ns 3. P. callosa

1. Pneumatopteris truncata (Poir.) Holttum, Blumea 21 (1973) 314.

Basionym: Polypodium truncatum Poir., Encycl. Meth. 5 (1804) 534.

Name in Holttum 1955: Cyclosorus truncatus (Poir.) Farwell, p. 266.

Correct name in Thelypteris: T. truncata (Poir.) K. Iwats., Mem. Coll. Sci.

Univ. Kyoto, B, 31 (1964) 33.

Status in Malaya: as Holttum 1955.

The type specimen of this species is labelled Brazil, and I accepted this

statement in 1955, but no species of Pneumatopteris is known from the New

World. ‘The distribution of P. truncata is: S. India and Ceylon (where it is

tetraploid), Western Malesia and the Philippines. Plants in Northern India are

diploid, with smaller fronds, more strongly toothed pinna-lobes and rather large

capitate hairs on sporangia.

16 Gardens’ Bulletin, Singapore XX XIII (1980)

2. Pneumatopteris ecallosa (Holttum) Holttum, Blumea 21 (1973) 310.

Basionym and name in Holttum 1955: Cyclosorus ecallosus Holttum, Gard.

Bull. Singap. 11 (1947) 269.

Correct name in Thelypteris: T. ecallosa (Holttum) Reed, Phytologia 17 (1968)

274.

Status in Malaya: now abundant in thickets by small streams at Cameron

Highlands; collected also at Genting Highlands at 1500 m.

3. Pneumatopteris callosa (Bl.) Nakai, Bot. Mag. Tokyo 47 (1933) 179.

Basionym: Aspidium callosum Bl., Enum. Pl. Jav. (1828) 152.

Name in Holttum 1955: Cyclosorus dicranogramma (misidentified, p. 267).

Correct name in Thelypteris: T. callosa (Bl.) K. Iwats., Mem. Coll. Sci. Univ.

Kyoto, B, 31 (1964) 34.

Status in Malaya: collected only three times, at Cameron Highlands and at

Genting Highlands, at 1500 m, near stream in forest.

Very young fronds are covered with a layer of mucilage through which the

aerophores project.

12. SPHAEROSTEPHANOS J. SM. IN HOOK. GEN. FIL. (1839) T. 24.

The type species of this genus is S. asplenioides J. Sm. (= S. polycarpus),

of which John Smith had a defective specimen; he thus made a wrong statement

as to the structure of the indusium. Later he corrected this statement and

remarked that the species did not differ significantly from others, which he named

Nephrodium, apart from its elongate sorus. For the same reason I transferred

S. polycarpus to Cyclosorus in 1955. In 1971, when breaking up Cyclosorus as

defined by Ching and Copeland, I limited Sphaerostephanos to a group of species

characterized by the presence of spherical yellow sessile glands, which occur

usually on the lower surface of pinnae, sometimes confined to costules and veins,

or to indusia or sporangia. Similar glands occur in some species of Pronephrium,

but the latter differ in having almost entire normal pinnae and no reduced pinnae

at the base of fronds. One Malayan species, S. norrisii, has only 1-2 pairs of

very small basal pinnae on its larger fronds, none on smaller ones. S. peltochlamys

has almost entire pinnae like those of Pronephrium but several pairs of reduced

basal pinnae; it may be of hybrid origin.

KEY TO THE MALAYAN SPECIES

1. Sori distinctly elongate

2. Reduced pinnae 6-12 pairs, normal pinnae lobed 1/4 — 1/3 towards

ot! ee ro Se ee 1. S. larutensis

2. Reduced pinnae.many pairs; normal pinnae lobed more than half-way

QO OOO oes aks cant cdtesccice sedan gedibsiiend een anennennnnnn 2. §S. polycarpus

Thelypteridaceae in Malaya 17

1. Sori not or little elongate

3. Spherical glands abundant on lower surface of pinnae

4. Hairs on lower surface of costae and costules appressed or distinctly

antrorse

5. Caudex slender, long-creeping; hairs on lower surface of costae

SUPER APIOUCNSAL 18 EO ee. tac cucetea 3. §. unitus

5. Caudex erect; hairs on lower surface of costae closely appressed

Beer PREORS 8 oe ised GIR Saisie d eck oeince 4. §. porphyricola

4. Hairs on lower surface of costae and costules erect or nearly so

6. Pinnae lobed less than half-way to costa; 2-3 pairs of veins

NS is POPE GE iy Brg ee ee 5. S$. penniger

6. Pinnae lobed at least half-way; only one pair of veins truly

anastomosing

7. Lower surface of costae and rachis glabrous; lowest normal

pinnae not or little narrowed at their bases ..................

6. S latebrosus

7. Lower surface of rachis and costae bearing short erect

hairs; lower normal pinnae much narrowed towards their

Dee ate eas cate Cee eee RE 7. S. heterocarpus

3. Spherical yellow glands lacking or rare on lower surface of pinnae

8. Lower surface of costae and costules covered with slender appressed

hairs

9. Rachis of sterile fronds bearing thick curved brown hairs on

both surfaces; basal veins not always anastomosing

10. Pinnae lobed less than half-way to costa; basal pair of

veins often anatomosing ................... 8. S$. pterosporus

10. Pinnae lobed 3/5 — 2/3 to costa; basal veins rarely

Pir ieecrenamrnte tats =< Jat 7 P22 77.2 es. 2igkd.22. cgsten~ Ee var. altilobus

9. Hairs on rachis pale; basal veins always anastomosing .........

S. hendersonii

Peeeeceescecseceesssescseveesesceesesesnaseeeeseveeeseveseneseeseeone

8. Lower surface of costae and costules not covered with appressed

hairs

11. Normal pinnae entire or crenate ............ 10. S. peltochlamys

11. Normal pinnae lobed 2/5 towards costa ......... ll. S. norrisii

1. Sphaerostephanos larutensis (Bedd.) C. Chr., Ind. Fil. Suppl. Ill (1934) 172.

Basionym: Nephrodium larutense Bedd., Handb. Suppl. (1892) 73.

Name in Holttum 1955: Cyclosorus larutensis (Bedd.) Ching, p. 284.

18 Gardens’ Bulletin, Singapore XX XIII (1980)

Correct name in Thelypteris: T. larutensis (Bedd.) Reed, Phytologia 17 (1968)

286.

Status in Malaya: as Holttum 1955.

2. Sphaerostephanos polycarpus (Bl.) Copel., Univ. Cal. Publ. Bot. 16 (1929) 60.

Basionym: Aspidium polycarpum Bl., Enum. Pl. Jav. (1828) 156.

Name in Holttum 1955: Cyclosorus polycarpus (Bl.) Holttum, p. 283.

Correct name in Thelyptersis: T. polycarpa (Bl.) K. Iwats., Mem. Coll. Sci.

Univ. Kyoto B, 31 (1964) 32.

Status in Malaya: as Holttum 1955.

There is a variety with long hairs on the lower surface of costae, collected

on Taiping Hills. Plants at 1500 m on Genting Highlands appear little different

from lowland plants.

3. Sphaerostephanos unitus (L.) Holttum, Journ. S. Afr. Bot. 40 (1974) 165.

Basionym: Polypodium unitum L., Syst. Nat. ed. 10, 2 (1759) 1326 excl. syn.

Name in Holttum 1955: Cyclosorus unitus (L.) Ching, p. 260.

Correct name in Thelypteris: T. unita (L.) Morton, Amer. Fern Journ. 49 (1959)

113.

Status in Malaya: as Holttum 1955.

There are two varieties of this species, one distributed in East Africa,

Mascarene Islands, Ceylon and Western Malesia, the other from Borneo and

the Philippines eastwards to Fiji.

4. Sphaerostephanos porphyricola (Copel.) Holttum, Kalikasan 4 (1975) 59.

Basionym: Dryopteris porphyricola Copel., Philip. J. Sci. 7C (1912) 60.

Name in Holttum 1955: Cyclosorus porphyricola (Copel.) Ching, p. 271.

Correct name in Thelypteris: 7. porphyricola (Copel.) Ching, Bull. Fan Mem.

Inst. Biol. Bot. 6 (1936) 287.

Status in Malaya: as Holttum 1955, but exclude reference to plants from

Kinabalu.

The fertile pinnae of this species are up to 14 x 2.5 cm (sterile ones some-

times larger), with abruptly short-acuminate apex.

5. Sphaerostephanos penniger (Hook.) Holttum in Nayar & Kaur, Comp. to

Beddome (1974) 209.

Basionym: Nephrodium pennigerum Hook., Spec. Fil. 4 (1862) 82.

Name in Holttum 1955: Cyclosorus megaphyllus (Mett.) Ching, p. 268.

Thelypteridaceae in Malaya 19

Correct name in Thelypteris: T. megaphylla (Mett.) K. Iwats., Mem. Coll. Sci.

Univ. Kyoto B, 31 (1964) 34.

Status in Malaya: as Holttum 1955; but caudex is never truly erect.

Blume wrongly gave the name Aspidium pennigerum (based on Polypodium

pennigerum Forster; a New Zealand species) to plants of the present species in

Java. Hooker copied his description but recognized that Blume has misused

Forster's name. In so doing, Hooker in effect created a new name; he retained

Forster’s name for the New Zaland plant, still in Polypodium because it had no

indusia. The name Thelypteris pennigera (Forst.) H. H. Allan applies to the

New Zealand fern, so that the later epithet megaphylla (given by Mettenius to

correct Blume’s mistake) has to be used if this species is transferred to the genus

Thelypteris.

6. Sphaerostephanos latebrosus (Mett.) Holttum in Nayar & Kaur, Comp. to

Beddome (1974) 209.

Basionym: Aspidium latebrosum Mett., Farngatt. IV (1858) 104.

Synonym: WNephrodium glaucostipes Bedd., Handb. Suppl. (1892) 80.

Name in Holttum 1955: Cyclosorus heterocarpus var. glaucostipes, p. 271.

Correct name in Thelypteris: T. latebrosa (Mett.) Reed, Phytologia 17 (1968)

287.

When writing my book of 1955 I had not seen specimens with the base

of fronds intact. Such specimens from the type collection, at Calcutta, show

that the basal normal pinnae are hardly narrowed at their bases, and the reduced

pinnae are all much larger than in S. heterocarpus. This species is still known

in Malaya only from two collections in Perak: near Gopeng, and at the foot

of G. Idong, both in lowlands. The type of A. latebrosum Mett. was from Java.

7. Sphaerostephanos heterocarpus (Bl.) Holttum in Nayar & Kaur, Comp. to

Beddome (1974) 280.

Basionym: Aspidium heterocarpon Bl\., Enum. Pl. Jav. (1828) 155.

Name in Holttum 1955: Cyclosorus heterocarpus (Bl.) Ching, p. 269.

Correct name in Thelypteris: | T. heterocarpa (Bl.) Morton, Amer. Fern Journ.

49 (1959) 113.

Status in Malaya: as Holttum, except that I think the plants mentioned which

had young fronds covered with mucilage belonged to another species, perhaps

S. latebrosus.

Lowland plants are all small, with pinnae less than 1.5 cm wide, lobed about

half-way to the costa; I cannot see a sharp distinction between these and mountain

plants which have wider more deeply lobed pinnae; more field study is needed.

8. Sphaerostephanos pterosporus (v.A.v.R.) Holttum, Flora Malesiana.

Basionym: Dryopteris pterospora v.A.V.R.

20 Gardens’ Bulletin, Singapore XX XIII (1980)

Name in Holttum 1955: Cyclosorus stipellatus, in part, p. 272.

Correct name in Thelypteris: T. pterospora (v.A.v.R.) Reed, Phytologia 17

(1968) 307.

In Holttum 1955 this species and S. hendersonii were confused with S.

stipellatus (Bl.) Holttum; all three species lack glands on the lower surface of

pinnae, have appressed hairs on lower surface of costae and long thick aerophores

at the bases of pinnae. The true S. stipellatus, in Java and Sumatra, has less

deeply lobed pinnae. S. pterosporus is distinguished by thick hairs on both

surfaces of the rachis. There are two varieties, as distinguished by the above key.

Var. pterosporus: pinnae lobed rather less than half-way to costa; basal

veins sometimes meeting to form excurrent veins, sometimes just touching each

other at the base of a sinus-membrane. This variety has been found at low

altitudes in the southern parts of Malaya: near Malacca (an old collection by

Pinwill), near Kajang in Trengganu (Corner) and at low altitudes on G. Pulai

and G. Muntahak in Johore. The type came from Sumatra; there are specimens

also from G. Matang in Sarawak.

Var. altilobus Holttum, Flora Malesiana: pinnae deeply lobed; veins rarely

anastomosing. Known only from two collections from Taiping Hills at 750-

900 m (Kunstler and Molesworth Allen), the recent one found along the pipe-

line path near the Tea Gardens.

9. Sphaerostephanos hendersonii Holttum, Flora Malesiana.

Name in Holttum 1955: Cyclosorus stipellatus, in part, p. 272.

This species agrees with S. pterosporus in most characters but has pale hairs

on the rachis and more deeply lobed pinnae. The basal veins are always at a

wide angle to the costule and always anastomose to form an excurrent vein to

the sinus. Three collections are known from forest at Cameron Highlands at

1500 m; one from North Sumatra appears to be conspecific.

10. Sphaerostephanos peltochlamys (C. Chr.) Holttum, Flora Malesiana.

Basionym: Dryopteris peltochlamys C. Chr.

Name in Holttum 1955: Abacopteris peltochlamys (C. Chr.) Holttum, p. 295.

Correct name in Thelypteris: T. peltochlamys (C. Chr.) Reed, Phytologia 17

(1968) 303.

Status in Malaya: — still known only from one collection from low country in

Kelantan.

IT have now seen many collections from Sumatra. Glands are often present

on sporangia and sometimes on indusia; indusia are always large and usually

glabrous but sometimes have a few hairs; small fronds usually lack reduced

basal pinnae, but several of these are always present on large fronds. This

species is anomalous both in Sphaerostephanos and in Pronephrium (A bacopteris);

as above noted, it might be of hybrid origin.

Thelypteridaceae in Malaya 21

11. Sphaerostephanos norrisii (Rosenst.) Holttum, Flora Malesiana.

Basionym: Dryopteris norrisii Rosenst. (type from Malaya).

Name in Holttum 1955: Cyclosorus toppingii, p. 280 (misidentification).

Synonym: Nephrodium pennigerum var.malayense Bedd., Handb. Suppl. (1892)

74.

Correct name in Thelypteris: none published.

Status in Malaya: as Holttum 1955.

The locality at which the type was collected by Norris is not recorded, but

probably it came from the Taiping Hills like the specimens cited by Beddome

and two more recent ones, at about 1000 m. The only other collection is my

own from Cameron Highlands in forest near the path leading down from

Robinson’s Falls to the Ringlet valley.

Additional information: glands are usually present on the lower surface of

costules and veins in the pinna-lobes, sometimes also on indusia and sporangia;

reduced basal pinnae are present only on large fronds.

Distribution outside Malaya: the type of Dryopteris falcinella v.A.v.R. from

Sumatra is quite identical with that of D. norrisii; specimens from Mindanao

agree in essentials with those of Malaya and Sumatra but differ in producing

fertile fronds of a much smaller size and in the hairs on the upper surface

between veins being short and suberect; a few specimens from Borneo are like

those from Mindanao. Dryopteris toppingii Copel. from Mt Kinabalu differs in

a total lack of giands and of reduced pinnae, and in glabrous indusia. A few

collections of D. toppingii have been made in other parts of Borneo. Copeland

gave the name Sphaerostephanos toppingii to a quite different species; another

name has therefore to be found for the Kinabalu plant which in 1955 I confused

with S. norrisii.

13. PRONEPHRIUM PRESL, EPIM. BOT. (1851) 258.

Caudex short-creeping; fronds lacking reduced basal pinnae; pinnae sub-

entire or crenate, usually with several pairs of anastomosing veins; lower surface

between veins often pustular when dry; spherical glands like those of Sphaero-

stephanos sometimes present on lower surfaces of pinnae or on sporangia; in

section Grypothrix hairs on the lower surface and on sporangia are hooked and

the sori are exindusiate; the species P. repandum is intermediate, with exindusiate

sori and sporangia bearing many long stiff straight hairs.

This genus, as at present arranged, may be composite; more study of species

over a wider area is needed; 57 species are known in Malesia and several more

in mainland Asia.

KEY TO THE MALAYAN SPECIES

1. No hooked hairs on any part of the plant

2. Sori indusiate (indusia in some cases small)

22 Gardens’ Bulletin, Singapore XX XIII (1980)

3. Pinnae to 30 cm long, not or little dimorphous; veins 12-15 pairs

os iednrie aan ave wiehewapanenascwesks bekey ORE ae ee 1. P. asperum

3. Pinnae much shorter; fronds dimorphous; veins not over 10 pairs

4. Lower surface of pinnae between veins bearing glands .........

Haid... bbdH..bi idee. Se 2. P. glandulosum

4. Lower surface of pinnae between veins lacking glands

5. Pinnae 4-5 pairs, their bases not or little auricled .........

2 acudeéeelanedeicns dca UuGe os aceeeen ec 3. P. menisclicarpum

5. Pinnae to 10 pairs, distinctly auricled ...... 4. P. pletatum

2. ‘Sor exindusiate 00.04...) ate ee 5. P. repandum

1. Hooked hairs present, at least on caudex

6. Caudex slender, long-creeping

7. Fronds” trifQuare 0.55 om anise nindsde eae 6. P. triphyllum

7. Pinnae more than one pair, irregular ............... 7. P. X parishii

6. Caudex thicker, short-creeping with fronds close together

8. Pinnae to 2 cm wide, their bases narrowly cuneate ................4

5 wine ad ships owes sisdalsbindas Aikguh apne cenEentie idvine meni 8. P. salicifolium

8. Pinnae much wider, their bases broadly cuneate ....................00:

s ncastiib-edmave igre hod « vincaitis ne Goh ligiaas abel Gia aan nnn nn 9. P. rubicundum

1. Pronephrium asperum (Presl) Holttum, Blumea 20 (1972) 112.

Basionym: Goniopteris aspera Presl, Tent. Pterid. (1836) 183.

Name in Holttum 1955: Abacopteris multilineata var. malayensis, p. 297.

Correct name in Thelypteris: T. aspera (Presl) Reed, Phytologia 17 (1968) 261.

Status in Malaya: as Holttum 1955.

2. Pronephrium glandulosum (BI.) Holttum, Blumea 20 (1972) 118.

Basionym: Aspidium glandulosum Bl., Enum. Pl. Jav. (1828) 144.

Name in Holttum 1955: Cyclosorus glandulosus (Bl\.) Ching, p. 278.

Correct name in Thelypteris: T. malayensis (C. Chr.) Reed, Phytologia 17 (1968)

291.

Status in Malaya: as Holttum 1955.

The name Thelypteris glandulosa belongs to a tropical American species, listed

by Christensen as Dryopteris glandulosa in his Index Filicum, 1905; he there

proposed a new epithet for the Malayan species.

Thelypteridaceae in Malaya 23

3. Pronephrium menisciicarpon (Bl.) Holttum, Blumea 20 (1972) 111.

Basionym: Aspidium menisciicarpon Bl., Enum. Pl. Jav. (1828) 142.

Name in Holttum 1955: Abacopteris menisciicarpa (Bl.) Holttum, p. 290.

Correct name in Thelypteris: T. menisciicarpa (Bl.) K. Iwats., Acta Phytotax.

Geobot. 21 (1965) 171.

Status in Malaya: as Holttum 1955.

4. Pronephrium peltatum (v.A.v.R.) Holttum, Flora Malesiana.

Basionym: Dryopteris peltata v.A.v.R.

Name in Holttum 1955: Abacopteris lineata, p. 292 (misidentification)

Correct name in Thelypteris: T. peltata (v.A.v.R.) Reed, Phytologia 17 (1968)

303.

Status in Malaya: as Holttum 1955 but excluding the specimens from Patani,

which are Pronephrium affine (Bl.) Holttum (a species not found in Malaya).

P. affine has thinner fronds, crenate pinnae and much larger indusia; it

occurs in the lowlands of Java and Sumatra, apparently on rocks by streams.

The type of P. peltatum was collected in Sumatra. It differs from Malayan

specimens in having distinctly crenate pinnae and small glabrous indusia. In

Flora Malesiana the Malayan plants are named var. peninsulare Holttum; they

are tetraploid. The true Pronephrium lineatum (Bl.) Pres! does not occur in

Malaya.

5. Pronephrium repandum (Fée) Holttum, Blumea 20 (1972) 109.

Basionym: Goniopteris repanda Fée, Gen. Fil. (1852) 251.

Name in Holitum 1955: Abacopteris urophylla, p. 296.

Correct name in Thelypteris: none published.

Status in Malaya: as Holttum 1955.

The name urophylla, adopted in 1955, is later than repanda so a change to

the latter is made; the types of both names came from Penang and undoubtedly

represent the same species.

6. Pronephrium triphyllum (Sw.) Holttum, Blumea 20 (1972) 122.

Basionym: Meniscium triphyllum Sw. in Schrad. Journ. Bot. 1800 2 (1801) 16.

Name in Holttum 1955: Abacopteris triphylla (Sw.) Ching, p. 287.

Correct name in Thelynteris: T. triphylla (Sw.) K. Iwats., Mem. Coll. Sci. Univ.

Kyoto B, 31 (1965) 190.

Status in Malaya: as Holttum 1955.

24 Gardens’ Bulletin, Singapore XX XIII (1980)

7. Pronephrium X parishii (Bedd.) Holttum, Blumea 20 (1972) 123.

Basionym: Meniscium triphyllum var. parishii Bedd., Handb. (1883) 399.

Name in Holttum 1955: Abacopteris triphylla var. parishii, p. 287.

Name in Thelypteris: T. triphylla var. parishii K. Iwats. l.c., p. 191.

Status in Malaya: on river banks; S. Kerling in Perak, S. Cheka in Pahang.

The specimens are variable in the number, size and spacing of pinnae, and

have the appearance of hybrids, one parent being P. triphyllum, the second un-

known. Similar plants have been found in Peninsular Thailand, southern Burma

and southern Vietnam; I have seen none with ripe sporangia. Plants from

Pahang cultivated in Singapore retained their variable character.

8. Pronephrium salicifolium (Hook.) Holttum, Blumea 20 (1972) 123.

Basionym: Meniscium salicifolium Hook., Ic. Pl. 10 (1854) t. 990.

Name in Holttum 1955: Abacopteris salicifolia (Hook.) Holttum, p. 288.

Correct name in Thelypteris: T. salicifolia (Hook.) Reed, Phytologia 17 (1968)

Status in Malaya: as Holttum 1955.

This is certainly related to P. triphyllum and T. rubicundum, but has hooked

hairs only on the caudex and bases of stipes; the fronds are almost glabrous.

9. Pronephrium rubicundum (v.R.v.R.) Holttum, Blumea 20 (1972) 123.

Basionym: Phegopteris rubicunda v.A.v.R., Bull. Jard. Bot. Btzg III, 2 (1920)

162.

Name in Holttum 1955: Abacopteris rubicunda (v.A.v.R.) Holttum, p. 292.

Correct name in Thelypteris: T. rubicunda (v.A.v.R.) K. Iwats., Mem. Coll.

Sci. Univ. Kyoto B, 31 (1965) 195.

Status in Malaya: as Holttum 1955.

14. CHRISTELLA LEV., FL. DE KOUY-TCHEOU (1915) 472;

HOLTTUM, TAXON 20 (1971) 533.

Caudex erect or short-creeping; fronds in all species except C. parasitica

with a few pairs of lower pinnae gradually decrescent, lowest not very small; aero-

phores not or little evident; pinnae always bearing many acicular hairs (very

short in two species) and in some species elongate glandular or short capitate

hairs; veins anastomosing in all Malayan species; sori indusiate; sporangia lacking

glands or hairs distally but bearing on their stalks elongate yellow or reddish

unicellular glandular hairs; perispore consisting of thick tubercles or ridges.

A genus of about 50 species in the warmer parts of the Old World, with

a group in the New World and Africa which still needs correlating with the

former. A full account of species in the Old World is given in Holttum, Kew

Bull. 31 (1976) 293 ff.

—

Thelypteridaceae in Malaya 25

KEY TO THE MALAYAN SPECIES

1. Caudex long-creeping; reduced lower pinnae not auricled ...... L., G. qrda

1. Caudex not long-creeping; lower pinnae, if reduced, distinctly auricled

2. Hairs on lower surface of costa and costules all minute, hardly distin-

guishable with a X 10 lens

3. Caudex erect; reduced pinnae c. 8 pairs, broad and deeply iobed

ALOR RU ARS SS TSS A A ed ee LE 2. C. papilio

3. Caudex short, suberect; reduced pinnae fewer, not deeply lobed

EN aR PRON: LU Lr a eA SOE, «soe nes ld SEALY. 3. C. subpubescens

2. Hairs on lower surface of costae and costules distinct, usually of varied

length

4. Pinnae lobed much more than half-way to costa; hairs on lower

surface of costae commonly | mm or more long

5. Basal pinnae not or little reduced; red glands present on lower

sarace Or COSLUIES ANG. VeMms te. ee 4. C. parasitica

5. Basal pinnae gradually decrescent and more widely spaced; no

red -dlands’ on lower Suiface®). :.c.28h kt 5. C. hispidula

4. Pinnae lobed half-way to costa or a littke more deeply; hairs on

lower surface of costae rarely 0.5 mm long ............ 6. C. dentata

1. Christella arida (D. Don) Holttum in Nayar & Kaur. Comp. to Beddome

(1974) 206.

Basionym: Aspidium aridum D. Don, Prodr. Fl. Nepal (1825) 4.

Name in Holttum 1955: Cyclosorus aridus (D. Don) Ching, p. 259.

Correct name in Thelypteris: T. arida (D. Don) Morton, Amer. Fern Journ.

a9 41959)" 113;

Status in Malaya: as Holttum 1955.

The most distinctive features of this species are the elongate glands on veins

in the pinna-lobes, and the lack of acroscopic basal auricles on the reduced

pinnae. This species is reported to be diploid in northern India.

2. Christella papilio (Hope) Holttum in Nayar & Kaur, Comp. to Beddome

(1974) 208.

Basionym: Nephrodium papilio Hope, Journ. Bombay Nat. Hist. Soc. 12 (1899)

Gra.) def 1,

Name in Holttum 1968: Cyclosorus papilio (Hope) Ching, p. 633.

Correct name in Thelypteris: T. papilio (Hope) K. Iwats., Mem. Coll. Sci. Univ.

Kyoto B, 31 (1965) 175.

Status in Malaya: known from Cameron Highlands (below Ringlet), Bukit

Hitam and Ginting Simpah, in forest at 300-900 m.

26 Gardens’ Bulletin, Singapore X X XIII (1980)

The massive, quite erect, caudex is distinctive. Specimens from Malaya and

Thailand all have large ellipsoid red glands on the lower surface of costules and

veins; in other respects they agree with the type from northern India.

3. Christella subpubescens (BI.) Holttum, Webbia 30 (1976) 193: Kew Bull. 31

(1976) 323.

Basionym: Aspidium subpubescens Bl., Enum. Pl. Jav. (1828) 149.

Names in Holttum 1955: Cyclosorus sumatranus (v.A.v.R.) Ching, p. 275; C.

latipinna (Benth.) Tard., p. 276.

Correct name in Thelypteris: T. subpubescens (Bl.) K. Iwats., Mem. Coll. Sci.

Univ. Kyoto B, 31 (1965) 173.

Status in Malaya: as Holttum 1955, but see note below on union of the two

species.

The common Singapore fern described as Cyclosorus sumatranus in 1955

agrees with the types of both Dryopteris sumatrana y.A.v.R. (1908) and of

Aspidium subpubescens Bl. (1828), also with specimens named D. sumatrana by

van Alderwerelt at Bogor. Plants named Cyclosorus latipinna in 1955 were from

stream-banks in low country in Perak and Pahang; in this situation they are

washed away by floods before they attain their full growth. Similar plants away

from streams (and cuitivated plants taken from stream-banks) are larger, and

most of them have fronds stiffer in growth than the Singapore plants; some of

them also have elongate red glands on the lower surface of veins (a gland-bearing

plant has also been found in Singapore). Bentham’s type and other specimens

of latipinna from Hong Kong lack glands and are closely similar to some stream-

bank plants from Perak. A cytotaxonomic study of cultivated plants taken from

various sources is needed to clarify the situation. It should be noted that a

Singapore plant cultivated at Leeds was found to be tetraploid; it probably had

one parent in common with C. dentata which is also a tetraploid.

4. Christella parasitica (L.) Lev., Fl. de Kouy-tchéou (1915) 475.

Basionym: Polypodium parasiticum L., Spec. Plant. (1753) 1090.

Name in Holttum 1955: Cyclosorus parasiticus (L.) Farw., p. 281.

Correct name in Thelypteris: 7. parasitica (L.) K. Iwats., Mem. Coll. Sci. Univ.

Kyoto B, 31 (1965) 172.

Status in Malaya: as Holttum 1955.

Plants in Ceylon similar to those in Malaya proved to be tetraploid. It is

probable that hybrids between this species and C. dentata occur in Malaya.

5. Christella hispidula (Decne) Holttum, Kew Bull. 31 (1976) 312.

Basionym: Aspidium hispidulum Decne, Nouv. Ann. Mus. Hist. Nat. Paris 3

(1834) 346.

Name in Holttum 1955: Cyclosorus contiguus (Rosenst.) Ching, p. 282.

Thelypteridaceae in Malaya 27

Correct name in Thelypteris: T. hispidula (Decne) Reed, Phytologia 17 (1968)

283. .

Status in Malaya: as Holttum 1955, and 1968 p. 633.

Aspidium hispidulum, based on a specimen from Timor at Paris, is the oldest

name for this pantropic diploid species, which is somewhat variable in different

countries. In Malaya the lower pinnae are distinctly decrescent and more widely

spaced. Plants from Ceylon have been hybridized with the tetraploids C. dentata

and C. parasitica.

6. Christella dentata (Forsk.) Brownsey & Jermy, Brit. Fern Gaz. 10 (1973) 338.

Basionym: Polypodium dentatum Forsk., Fl. Aegypt. — Arab. (1773) 1835.

Name in Holttum 1955: Cyclosorus subpubescens, p. 273 (misidentification).

Correct name in Thelypteris: T. dentata (Forsk.) E. St John, Amer. Fern Journ.

26 (1936) 44.

Status in Malaya: rather common everywhere in somewhat exposed places, not

in forest.

This is a very variable and widely distributed tetraploid; in Malaya it pro-

bably hybridizes with both C. parasiticus and C. subpubescens.

15. AMPHINEURON HOLTTUM, BLUMEA 19 (1971) 45; 23 (1977) 205.

Lower pinnae always much narrowed towards their bases; reduced pinnae,

one or two pairs, inconstantly present at the base of fronds in some species;

pinnae lobed; veins in several species (notably in A. opulentum) not constantly

anastomosing; glandular hairs on lower surfaces pear-shaped, very small and

often pale yellow, or larger and colourless; indusia present; sporangia lacking

glands or hairs distally, glandular hairs on stalk apparently variable; spores as

in Christella.

This is a small: genus extending from East Africa to the Pacific.

KEY TO THE MALAYAN SPECIES

1. Basal veins often or always anastomosing: caudex not erect

2. Pinnae lobed less than half-way to costa; basal veins always anastomosing

SE Retr is A es Ke ee a nic wi Sine we bad Sack - weal od 1. A. terminans

2. Pinnae lobed more than half-way; basal veins sometimes free ............

6 DOS DE Ee RG Pepe EN a POS Beall a Te Oe PE 2. A. opulentum

Eoasat evetmsentwiays irec, caudex erect: 2... 20.....-2se08.....0... 3. A. immersum

1. Amphineuron terminans (Hook.) Holttum, Amer. Fern Journ. 63 (1973) 82.

Basionym: Nephrodium terminans Hook., Spec. Fil. 4 (1862) 73.

Name in Holttum 1955: Cyclosorus interruptus, p. 262 (misidentification).

Correct name in Thelypteris: T. terminans (Hook.) Tagawa & K. Iwats., Acta

Phytotax. Geobot. 26 (1975) 169.

Status in Malaya: as Holttum 1955.

28 Gardens’ Bulletin, Singapore X X XIII (1980)

Colourless glandular hairs are usually present on the lower surface of veins,

the distal ones largest; their presence is sometimes the best way of distinguishing

old fronds of this species from those of the true Cyclosorus interruptus.

2. Amphineuron opulentum (Kaulf.) Holttum, Blumea 19 (1971) 45; 23 (1977)

2142.

Basionym: Aspidium opulentum Kaulf., Enum. Fil. Chamisso (1824) 238.

Name in Holttum 1955: Cyclosorus extensus (Bl.) Ching, p. 264.

Correct name in Thelypteris: T. opulenta (Kaulf.) Fosberg, Smiths. Contr. Bot.

8 (1972) 3.

Status in Malaya: as Holttum 1955.

3. Amphineuron immersum (BI.) Holttum in Nayar & Kaur, Comp. to Beddome

(1974) 203.

Basionym: Aspidium immersum Bl., Enum. Pl. Jav. (1828) 156.

Name in Holttum 1955 and correct name in Thelypteris: T. immersa (Bl.) Ching,

p. 243.

Status in Malaya: as Holttum 1955S.

LIST OF NAMES IN HOLTTUM 1955 AND 1968

WITH EQUIVALENTS IN THE PRESENT PAPER

Abacopteris lineata (Bl.) Ching = Pronephrium peltatum (v.A.v.R.)

Holttum

Abacopteris menisciicarpa (BI.) = Pronephrium menisciicarpum (BI.)

Holttum Holttum

Abacopteris multilineata (Hook.) = Pronephrium asperum (Presl)

Ching var. malayensis Holttum Holttum

Abacopteris peltochlamys (C. Chr.) = Sphaerostephanos_ peltochlamys

Holttum (C. Chr.) Holttum

Abacopteris rubicunda (v.A.v.R.) = Pronephrium rubicundum (v.A.v.R.)

Holttum Holttum

Abacopteris salicifolia (Hook.) = Pronephrium salicifolium (Hook.)

Holttum Holttum

Abacopteris triphylla (Sw.) Ching = Pronephrium triphyllum (Sw.)

Holttum

Abacopteris urophylla (Hook.) Ching = Pronephrium repandum (Fée)

Holttum

Ampelopteris prolifera (Retz.) Copel. unchanged

Cyclosorus aridus (D. Don) Ching = Christella arida (D. Don) Holttum

Thelypteridaceae in Malaya

Cyclosorus contiguus (Rosenst.)

Copel.

Cyclosorus dicranogramma (v.A.v.R.)

Holttum

Cyclosorus ecallusus Holttum

Cyclosorus extensus (Bl.) Ching

Cyclosorus ferox (Bl.) Ching

Cyclosorus glandulosus (BI.) Ching

Cyclosorus gongylodes (Schkuhr)

Link

Cyclosorus heterocarpus (Bl.) Ching

var. glaucostipes (Bedd.)

Cyclosorus interruptus (Willd.) Ching

Cyclosorus larutensis (Bedd.) Ching

Cyclosorus latipinna (Hook.) Tard.

Cyclosorus megaphyllus (Mett.)

Ching

Cyclosorus papilio (Hope) Ching

Cyclosorus parasiticus (L.) Farw.

Cyclosorus polycarpus (BI.) Holttum

Cyclosorus porphyricola (Copel.)

Ching

Cyclosorus stipellatus (BI.)

Ching p.p.

stipellatus p.p.

Cyclosorus subpubescens (BI.) Ching

Cyclosorus sumatranus (v.A.v.R.)

Ching

Christella hispidula (Decne) Holttum

Pneumatopteris callosa (Bl.) Nakai

Pneumatopteris ecallosa (Holttum)

Holttum

Amphineuron opulentum (Kaulf.)

Holttum

Chingia sakayensis (Zeiller) Holttum

Pronephrium glandulosum (BI.)

Holttum

Cyclosorus interruptus (Willd.) H. Ito

Sphaerostephanos heterocarpus (BI.)

Holttum

Sphaerostephanos latebrosus (Mett.)

Holttum

Amphineuron terminans (Hook.)

Holttum

Sphaerostephanos larutensis (Bedd.)

Car.

Christella subpubescens (BIl.) Holttum

Sphaerostephanos penniger (Hook.)

Holttum

Christella papilio (Hope) Holttum

Chrisiella parasitica (L.) Lév.

Sphaerostephanos polycarpus (BI.)

Copel.

Sphaerostephanos porphyricola

(Copel.) Holttum

Sphaerostephanos pterosporus

(v.A.v.R.) Holttum

Sphaerostephanos hendersonii

Holttum

Christella dentata (Forsk.) Brownsey

& Jermy

Christella subpubescens (BI.) Holttum

30 Gardens’ Bulletin, Singapore XX XIII (1980)

Cyclosorus toppingii (Copel.) Ching

Cyclosorus truncatus (Poir.) Farw.

Cyclosorus unitus (L.) Ching

Thelypteris beddomei (Bak.) Ching

Thelypteris brunnea (Wall.) Ching

Thelypteris chlamydophora (Rosenst.)

Ching

Thelypteris ciliata (Benth.) Ching

Thelypteris crassifolia (Bl.) Ching

Thelypteris herbacea Holttum

Thelypteris immersa (Bl.) Ching

Thelypteris

Holttum

motleyana (Hook.)

Thelypteris oppositipinna (v.A.v.R.)

Ching

Thelypteris paleata (Copel.) Holttum

Thelypteris

Ching

var. eglandulosa Holttum

pectiniformis (C. Chr.)

var. hirsuta Holttum

Thelypteris singalanensis (Bak.)

Ching

Thelypteris uliginosa (Kunze) Ching

Thelytperis unidentata (Bedd.)

Holttum

Thelypteris viscosa (J. Sm.) Ching

p.p.

viscosa p.p.

Sphaerostephanos norrisii (Rosenst.)

Holttum

Pneumatopteris truncata (Poir.)

Holttum

Sphaerostephanos unitus (L.) Holttum

Parathelypteris beddomei (Bak.)

Ching

Pseudophegopteris paludosa (BI.)

Ching

Mesophlebion chlamydophorum

(Rosenst.) Holttum

Trigonospora ciliata (Benth.) Holttum

Mesophlebion crassifolium (BI.)

Holttum

Coryphopteris hirsutipes (Clarke

Holttum

Amphineuron immersum (BL.)

Holttum

Mesophlebion motleyanum (Hook.)

Holttum

Pseudophegopteris rectangularis (Zoll.)

Holttum

Mesophlebion trichopodum (C. Chr.)

Holttum

Coryphopteris pectiniformis (C. Chr.)

Holttum

Coryphopteris pectiniformis

Coryphopteris pectiniformis var.

hirsuta Holttum

Metathelypteris day (Bedd.) Holttum

Macrothelypteris torresiana (Gaud.)

Ching

Coryphopteris unidentata (Bedd.)

Holttum

Coryphopteris viscosa (J. Sm.)

Holttum

Coryphopteris arthrotricha Holttum

Coryphopteris gymnopoda (Bak.)

Holttum

Coryphopteris tahanensis Holttum

REVISION OF MEMECYLON L.

(MELASTOMATACEAE)

FROM THE MALAY PENINSULA

J. F. MAXWELL

Botanic Gardens, Singapore

32 Gardens’ Bulletin, Singapore XX XIII (1980)

CONTENTS

Page

Abstract me oe ee dick ded a tnt a 33

Introduction ae Eee ee ae of Lid re ia 34

Literary review of Memecylon from the Malay Peninsula si ee. 33

Salient Morphological Characters of Meimecylon

Habit ane ae ne a6 i fe ae oe 36

Bark and wood ee bad a ae 7a Le A 36

Branchlets me cr As a a: ae oud ie 29)

Interpetiolar ridge or line... ay ia Si aU er 37

Leaves ae as ee bse en a. a ee a

Inflorescence ~~... oe has ae: ee ae a a 39

Calyx ve si ah x ies we _ ag 40

Petals se me e +8 a sat rk Re 4]

Stamens a ro ee es A nate a a 4]

Gynoecium kde AS: ce "y 33 Ents ie % 42

Fruit Ds i ee Ep i ite . i 43

Distribution and abundance We Da ree iz ae 44

Taxonomic Section

Generic description... hye he ee mt , yok 44

Key to flowering material... mY oe Me oa a 45

Key to fruiting material ee wks nh an sah i Sl

Taxa (descriptions and critical notes) ep ie ee a 57

Insufficiently known taxa _... a aa i le Pe 107

General Conclusions... Sate ih a She — +e ca 108

Table 1. Review of salient morphological characters... veh ei 110

Index to collections... ay. oo oa ad io va Ae 113

Index to taxa ie oy Ses ry me ay se =. lee

Reduced taxa from the Malay Peninsula a” bi eas a 126

Illustrations :

Line drawings ... oT she iv a . si so 1

Photographs _... re ne is ak va fe ain 142

Revision of Memecylon L. 33

ABSTRACT

This revision of Memecylon of the Malay Peninsula includes twenty-eight

species, three varieties, and three imperfectly known taxa. No new taxa are

proposed here. However, twenty-two taxa have been reduced to new synonyms,

along with one new combination. Separate keys to flowering and fruiting material,

critical taxonomic notes, detailed analyses of various organs of taxonomic import-

ance, and an index to collections are included. Simple line drawings of the floral

parts and other salient features of each taxon have been prepared to supplement

the descriptions.

ACKNOWLEDGEMENT

In November 1976, when Dr. Hsuan Keng (University of Singapore) suggested

that I study Memecylon, I eagerly accepted the project since I knew that it

would be a challenge. Indeed the work has proved to be very interesting and

rewarding.*

I am most grateful to Prof. A.N. Rao and Dr. Hsuan Keng fore their assist-

ance, helpful suggestions, and encouragement during my research and preparation

of the revision.

Drs. Robert Geesink at Leiden, Netherlands invited me to study from May

to July 1977 at the Rijksherbarium, Leiden where a major portion of the research

for this paper was completed. Many other botanists there provided inspiration

and encouragement for me to continue my work. I would also like to thank

the Keeper and staff of the Kew herbarium for their hospitality during my six

day visit and forthe loan of many specimens. I also borrowed several collections

from the British Museum (Natural History) and I would like to express my

thanks to the Keeper of that herbarium.

Dr. Chang Kiaw Lan, Curator of the Singapore herbarium, has assisted in

numerous ways and I am deeply grateful for her kindness and friendship. I

would also like to thank Dr. B. C. Stone, University of Malaya, for editing this

paper. I acknowledge gratefully the work of Dr. C. X. Furtado in the Singapore

herbarium for he organized and sorted out the specimens of Memecylon there

saving me weeks of effort and considerable frustration.

Finally, I must express my thanks to Mr. Vijay Kumar Natarajan for draw-

ing, and to Mr. Jumali Kafrawi and Mrs. Lucia Sunderaj Krueger for preparing

and arranging the illustrations.

* M.Sc. thesis, University of Singapore (Feb. 1978).

34 Gardens’ Bulletin, Singapore XX XIII (1980)

INTRODUCTION

King, Gamble, and Ridley’s “‘Materials for a Flora of the Malay Peninsula”

(1889-1907), followed by Ridley’s “Flora of the Malay Peninsula” (1922-1925)

are two monumental works which have served several generations of botanists

and foresters well, but are now in need of a thorough revision. With the

accumulation of numerous collections from throughout the Malay Peninsula since

the end of World War II and better knowledge of the floras of neighbouring

areas — especially those of Sumatra and Borneo, a revised flora of the Malay

Peninsula is urgently needed. Although excellent revisions of many families

have appeared in “‘Flora Malesiana”’ (edited by Prof. Dr. C.G.G.J. van Steenis)

since 1948, the gigantic scope and high standards of the project will take years

(generations ?) to complete. The Melastomataceae, a very large and complex

family, was inadequately treated in the earlier floras and now needs to be re-

studied. Memecylon, one of the larger genera of Melastomataceae from the

Malay Peninsula, has perplexed botanists for many years and even with several

excellent papers to work with, it is still very difficult to be certain about the

identities of many taxa. An effort has been made, therefore, to be as careful

and critical as possible in this revision of the genus for the Malay Peninsula.

It is hoped that this presentation will clear some of the confusion and mis-

understanding that has built up through the years concerning Memecylon in the

region.

Revision of Memecylon L. 35

REVIEW OF EARLIER WORK ON MEMECY LON

FROM THE MALAY PENINSULA

The first species of Memecylon described from the Malay Peninsula was

M. acuminatum Sm. in 1813, followed by M. caeruleum Jack in 1820. With

the publication of the Wallich Catalogue in 1831 and distribution of specimens

collected at Penang and Singapore a few years earlier (about 1822), five more

distinct taxa, including what are now known as M. cinereum King (M. umbellatum,

4109 in part), M. wallichii Ridl. (M. depressum Benth., 4101 in part) M. mega-

carpum Furtado (4102), and M. edule Roxb. var. ovatum (Sm.) Cl. (M. grande

Retz., 4103 in part; M. umbellatum, 4109B; M. laxiflorum Wall. ex Ridl., 4472)

were recorded from the Peninsula. Roxburgh added M. amplexicaule Roxb.

from Penang, in his ‘Flora Indica” (II, 260) of 1832. Clarke, in Hooker’s

“Flora of British India’? (II, published in 1879), added M. elegans Kurz var.

dichotoma Cl. (=M. dichotomum (Cl.) King var. dichotomum), M. maingayi Cl.

(=M. excelsum Bl.), M microstomum Cl. (=M. amplexicaule Roxb.), M. cam-

panulatum Cl., M. pubescens (Cl.) King, M. acuminatum Sm. var. flavescens Cl.,

M. grande Retz. var. horsfieldii Cl. (=M. oleifolium Bl.), and M. malaccense

(Cl.) Ridl.

The first systematic treatment of Memecylon for the Malay Peninsula was

published by King in 1900 with 28 species and 4 varieties included. Four of

these taxa (3 species and | variety) were described by King from specimens from

the Andaman and Nicobar Islands and only one of them (M. kurzii King =M.

excelsum Bl.) is found on the Peninsula.

Ridley (1910-1920) published 9 new species of Memecylon from the Malay

Peninsula, 7 of which were included in his “‘Flora of the Malay Peninsula” I

(1922). Ridley, however, did not include M. corticosum Ridl.; and he also

reduced M. eugeniflora Ridl. to M. dichotomum (Cl.) King var. eugeniflorum

(Ridl.) Ridl., and M. pulchellum Ridl. to M. pauciflorum Bl. In his treatment,

31 species and 3 varieties of Memecylon are included.

Craib described 3 species and 2 varieties of Memecylon from lower Thailand

in 1930-31 viz. M. brandisianum Craib (=M. oleifolium Bl.), M. constrictum

Craib (q.v. M. minutiflorum Mig.), M. dissitum Craib (= M. cantleyi Ridl.),

M. gracilipes Ridl. var. rotundatum Craib. (=M. dichotomum (Cl.) King var.

rotundatum (Craib) Maxw.), and M. pauciflorum BI. var. brevifolium Craib. The

last taxon is insufficiently known and is probably synonymous with the typical

variety.

Bakhuizen van den Brink’s exhaustive study of Melastomataceae (1943-45)

included a detailed analysis of Memecylon from the Malay Archipelago. He

included 36 species (14 of which were described as new) and 3 varieties. A

total of 25 of these taxa are found in the Malay Peninsula.

The second edition of Corner’s ‘““Wayside Trees of Malaya” (1952) includes

brief descriptions of 9 taxa with useful field notes, mostly complied from his

own observations.

36 Gardens’ Bulletin, Singapore XX XIII (1980)

The latest study of Memecylon for the Malay Peninsula was by Furtado

in 1963. While not an exhaustive or complete study, eight species of Memecylon

are discussed critically. M. acuminatissimum Bl. (=M. oleifolium BIl.), M.

hepaticum Bl., and M. multiflorum Bakh. f. (=M. campanulatum Ci.) are in-

cluded as new records from the Malay Peninsula; and M. megacarpum Furtado

which is a new name for M. pulchrum Cogn.

Salient Morphological Characters of Memecylon.

HABIT

All taxa of Memecylon from the Malay Peninsula are woody. From the

field notes for many collections examined it is apparent that there is considerable

variation in the habit of many taxa. M. malaccense Ridl. and M. pauciflorum

Bl. var. brevifolium Craib, both rather poorly known taxa, have only been re-

corded as being shrubs; while 18 other taxa have been collected in flowering

or fruiting stages as shrubs and treelets up to 4 m tall, or trees up to about

1S m. M. caeruleum Jack, M. cantleyi Ridl., M. cinereum King, M. fruticosum

King, M. garcinioides Bl., M. globosum Bakh. f., M. hullettii King, and M.

wallichii Ridl. have bee noted most commonly as shrubs. Some taxa which

have often been collected as treeleits are M. dichotomum (Cl.) King and M.

corticosum Ridl. Other taxa which have been recorded as small trees more

often than as shrubs or treelets include: M. megacarpum Furtado, M. lilacinum

Z. & M., and M. oligoneurum Bl. M. beccarianum Cogn. has been recorded

as a shrub, but most specimens were collected from trees.

Seven species have been recorded only as small to medium sized trees

(10-20 m tall) which, as far as I can determine, have never been collected in

flower or fruit as shrubs or treelets. These include: M. acuminatum Sm., M.

amplexicaule Roxb., M. campanulatum Cl., M. excelsum BI., M. floridum Ridl.,

M. kunstleri King, and M. oleifolium Bl. The largest species collected (20-30 m

tall with a diameter up to 2 m and often buttressed) are: M. intermedium BL.,

M. minutiflorum Mig., M. paniculatum Jack, and M. pubescens (Cl.) King.

More field studies are required in order to understand the habit of many

taxa of Memecylon from the Malay Peninsula. Knowledge of the habits, growth

forms, and habitats of many taxa of Memecylon is inadequate.

BARK AND WOOD

Unfortunately, only a few specimens of Memecylon examined have bark or

wood samples attached. During the present work eight taxa of Memecylon

have been seen in the field and with the notes on several collections it seems

that most, if not all, taxa of Memecylon have similar bark — at least for those

collected as trees. The outer bark is typically thin (1-2 mm thick); with shallow,

vertical, closely spaced fissures and cracks, sometimes flaking or peeling off, and

of a greyish-brown to blackish colour. The slash inner batk is also thin, and

is orange to brown. The wood in larger trees is usually white to brown, very

dense, and sinks in water. In many taxa the wood is very durable and is used

for house posts, lumber, furniture, and fuel.

Figure 31 of the trunk of a mature specimen of Memecylon lilacinum Z.

& M. illustrates the typical nature of the bark in this genus.

Revision of Memecylon L. 37

BRANCHLETS

The branchlets of Memecylon are often very useful in the identification

of many species. Basically there are three different types of branchlets: cylindric,

grooved, and angled or winged. These branchlets are smooth and glabrous in

all taxa, and ultimately become cylindric.

There are 15 taxa with cylindric branchlets, e.g. M. amplexicaule Roxb.,

M. megacarpum Furtado, M. oleifolium Bl., etc. Many taxa have cylindric

branchlets which are often slightly flattened and shallowly grooved on the two

flattened faces immediately below the upper node; the branchlets become cylindric

below this short flattened region. Some examples include: M. acuminatum Sm..,

M. caeruleum Jack, and M. garcinioidzs Bl. In 10 taxa the branchlets are

flattened and grooved on the two compressed faces for the entire length of the

upper internode. Older branchlets become cylindric, or nearly so, but there is

often a trace of the grooves. Several excellent examples are: M. campanulatum

Cl., M. cinereum King, and M. minutiflorum Mig. Vegetative specimens of

M. campanulatum Cl. and M. caeruleum Jack can be distinguished on the basis

of a groove in the former and cylindric branchlets in the latter. Each groove

is bordered by 2 sharp ridges and in many instances the groove is widened (that

is the branchlet is less compressed), thus giving the branchlet a 4-angled appear-

ance. M. edule Roxb., M. pauciflorum Bl., and M. lilacinum Z. & M. frequently

show this feature. The upper node in M. campanulatum Cl. and M. edule Roxb.

is often angled and tapers to 2 grooves on the upper internode. The grooved

branchlets of M. floridum Ridl. is a very good distinguishing feature separating it

from M. acuminatum Sm. which has cylindric branchlets. Also M. megacarpum

Furtado, with grooved branchlets, can immediately be distinguished from M.

excelsum Bl. and M. beccarianum Cogn. which have cylindric branchlets. The

remaining 6 taxa of Memecylon have sharply 4-angled to 4-winged branchlets

and upper nodes. M. dichotomum (Cl.) King var. dichotomum and M. dicho-

tomum (Cl.) King var. rotundatum (Craib) Maxw. have 4 wings below the upper

node which tapers to 4 angles. M. corticosum Ridl., M. wallichii Ridl., and

M. fruticosum King have distinctly 4-winged internodes. M. paniculatum Jack,

because of its 4- angled to 4-winged branchlets, is readily distinguished from

M. kunstleri King which has cylindric internodes.

INTERPETIOLAR RIDGE OR LINE

At first glance it appears that Memecylon has stipular scars, however these

interpetiolar ridges or lines, best seen on upper nodes, are merely raised zones

of articulation at the base of each petiole which appear separated (by a groove)

on the upper node and coalesce on the lower nodes to form a connecting ridge

or line which becomes indistinct on older branchlets.

The growing tip of each branchlet is distinct and is flanked on each side

at the base by a pair of leaves which are never connected by stipules, lines, or

ridges.

LEAVES

The leaves of all taxa of Memecylon are opposite, simple, entire, and glabrous.

The texture may be chartaceous to thick coriaceous and seems to be constant,

and often a diagnositc feature, of each taxon. Memecylon dichotomum (C1.)

King var. dichotomum and M. dichotomum (Cl.) King var. rotundatum (Craib)

38 Gardens’ Bulletin, Singapore XX XIII (1980)

Maxw. have very thin (chartaceous) blades while all other taxa have thicker

blades. Among the 12 taxa with sub-coriaceous blades are: M. corticosum Ridl.,

M. megacarpum Furtado, M. paniculatum Jack, and M. pubescens (Cl.) King.

Three species have blades which range in texture from sub-coriaceous to coria-

ceous viz. M. cantleyi Ridl., M. intermedium Bl., and M. oleifolium Bl. The

remaining 14 taxa have coriaceous blades and include, for exampie: M. edule

Roxb. var. edule, M. edule Roxb. var. ovatum (Sm.) Cl., M. globosum Bakh. f.,

M. lilacinum Z. & M., M. pauciflorum Bl., and M. wallichii Ridl. Very often

the blades of M. amplexicaule Roxb., M. caeruleum Jack, and M. campanulatum

Cl. are thick coriaceous.

The shape of the blades varies from lanceolate (e.g. M. corticosum Ridl.)

to sub-orbicular (e.g. the limestone form of M. dichotomum (Cl.) King). Blade

tips vary from broadly rounded (e.g. M. amplexicaule Roxb. and M. caeruleum

Jack) to acuminate-caudate (M. acuminatum Sm.). Cordate blade bases are

found in several taxa, e.g. M. amplexicaule Roxb., M. caeruleum Jack, M. wallichii

Ridl., etc.; while in others it is often broadly rounded, e.g. M. cinereum King

and M. fruticosum King; or decurrent, e.g. M. floridum Ridl. and M. oleifolium

BI. The blades of all taxa of Memecylon from the Malay Peninsula (except

M. oligoneurum Bl. which is trinerved) have a single main nerve which, in all

taxa (including the three main nerves of M. oligoneurum Bl.), is sunken on the

dorsal surface and raised below where it is thickest near the petiole and tapers

to the apex. The secondary veins range in number from about 6 to about 25

pairs which frequently parallel the mid-nerve for a few mm _ before bending

sharply at 45° towards the intramarginal nerve. In most taxa where the venation

is visible the distal end of each secondary nerve tends to curve towards the

apex of the blade before merging with the intramarginal nerve. The intramarginal

nerve parallels the margin (generally at a distance from 1-5 mm) in a nearly

straight (e.g. M. garcinioides Bl. and M. pubescens (Cl.) King) or breadly looping

line (e.g. M. dichotomum (Cl.) King, M. excelsum Bl., and M. wallichii Ridl.).

The intramarginal nerve is usually of the same thickness and degree of pro-

minence as the secondary veins.

For covenience the venation types are grouped into five general categories

in this paper: invisible, invisible to obscure, obscure, distict, and prominent. The

veins are typically invisible in 5 taxa — all of which have thick blades viz.

M. amplexicaule Roxb., M. caeruleum Jack, M. campanulatum Cl., M. pauciflorum

Bl. var. pauciflorum, and M. pauciflorum Bl. var. brevifolium Craib. There are

9 taxa which have invisible to obscure venation, e.g. M. acuminatum Sm., M.

edule Roxb. var. edule, M. edule Roxb. var. ovatum (Sm.) Cl., M. globosum

Bakh. f., M. malaccense (Cl.) Ridl., M. lilacinum Z. & M., etc. M. cantleyi Ridl.,

M. floridum Ridl., M. garcinioides Bl., and M. intermedium Bl. have obscure

venation; while 5 taxa have distinct (that is, slightly more prominent) venation,

e.g. M. dichotomum (Cl.) King and M. hullettii King. Prominet venation is

common in 8 taxa where the secondary veins are often raised below, e.g. M.

corticosum Ridl., M. excelsum Bl..M. paniculatum Jack, and M. wallichii Ridl.

The leaves of Memecylon have been a source of confusion in many taxa

and in numerous instances new species have been distinguished on this basis

alone. With detailed analyses of other characteristics of various species it is

obvious that many taxa have similar inflorescences, flowers, and fruits; there-

fore with voriable leaf morphologies it has often been difficult to determine some

species solely on this characteristic. Memecylon caeruleum Jack, M. dichotomum

(Cl.) King, M. lilacinum Z. & M., and M. oleifolium Bl. are examples where

Revision of Memecylon L. 39

the leaves are variable and in many instances intermediate specimens can be

shown to link some of the new synonyms proposed here to the accepted species.

Among the most striking examples include the reduction of M. floribundum BI.

to a new synonym of M. caeruleum Jack, and the numerous new synonyms

under M. oleifolium BI.

The branches and leaves of many species of Eugenia L. (Myrtaceae) (e.g.

E. polyantha Wight and E. scortechinii King) often resemble those of several

species of Memecylon. Aside from obvious differences in the inflorescences,

flowers, and fruit, Eugenia blades often have closer secondary venation, and dark

glandular dots the latter being absent in Memecylon and affording the most

reliable distinction in vegetative specimens.

Memecylon oligoneurum BI., the only species of Memecylon from the Malay

Peninsula with trinerved blades, is easily confused with Pternandra Jack. In

addition to well defined floral differences, the secondary veins in Pternandra are

generally more distinct and reticulate, and the branchlets usually have a darker

epidermis which tends to peel off. Strychnos L. (Loganiacaae) is in several ways

similar vegetatively to M. oligoneurum Bl. and Pternandra; however its climbing

habit immediately distinguishes it from these two genera.

The petioles of Memecylon generally range from 2-6 mm in length, 1-3 mm

in width, and are usually grooved on the upper surface. M. edule Roxb. var.

edule, with petioles typically shorter than M. edule Roxb. var. ovatum (Sm.)

Cl. is perhaps the only example seen presently where petioles are used to dis-

tinguish taxa. Aside from their relative lengths, the petioles have few charac-

teristics which can be used to distinguish Malayan taxa.

INFLORESCENCE

The basic structure of the inflorescence in Memecylon is cymose. From this

arrangement there are many variations ranging from a panicle of cymes to sessile

glomerules. The most complex inflorescences in some species have distinct 3rd

and 4th order axes, in addition to the pedicels (e.g. M. oleifolium Bl. and M.

paniculatum Jack). Contraction of the ultimate axes has led to more compact

inflorescences where the 4th or 3rd order axes are reduced yielding glomerules

on the tips of lower axes (e.g. M. edule Roxb. and M. pubescens (Cl.) King);

or umbels (M. hulletti King) due to the flattening of the nodes. Further reduc-

tion has resulted in the shortening of the secondary axes so that they appear

glomerulate or up to c. 1 mm long on the tips of the primary axes (e.g. M.

intermedium Bl. and M. minutiflorum Miq.). With the shortening of the primary

axes the inflorescence becomes very compact (e.g. M. cinereum King. M. oligo-

neurum Bl., and M. lilacinum Z. & M.). Finally, in several species (e.g. M.

amplexicaule Roxb. and M. campanulatum Cl.) the primary axes are indistinct

and the pedicels are clustered on a tubercle.

A pair of bracts, is present at the base of each axis and these are situated

at 90° to the axis below them. They are usuaily of an ovate shape (lanceolate

in M. garcinioides Bl.), acute at the tip, usually less than | mm long, and are

either persistent (e.g. M. caeruleum Jack and M. dichotomum (Cl.) King) or,

as in most taxa, fall off before the petals mature. Fallen bracts and bracteoles

leave distinct scars at the base of each axis. The bracts and bracteoles have

not been very useful taxonomic aids except in those few taxa in which they

are persistent.

40 Gardens’ Bulletin, Singapore XX XIII (1980)

The nodes of the inflorescence are typically flattened, laterally expanded,

and often reflexed at the sides. This feature is most readily seen in taxa with

large inflorescences (e.g. M. oleifolium Bl. and M. hullettii King). Subsequent

axes are arranged at right angles to lower axes with the central axis being longer

than the lateral ones.

The pedicels are usually in clusters, but sometimes reduced to one on the

tip of each ultimate axis. M. caeruleum Jack and M. dichotomum (C1.) King

are good examples. Many authors have erroneously interpreted this as a pedicel

with a pair of bracts near the middle. The primary axis is typically flattened,

usually with 2 distinct grooves or 4-angled, and arranged so that the flattened

surfaces face the branch and petiole. The axes are glabrous except in M.

beccarianum Cogn., M. hullettii King, and M. pubescens (Cl.) King. The inflore-

scences arise from leafy nodes in most species, however in a few taxa (e.g. M.

edule Roxb. var. ovatum (Sm.) Cl., M. excelsum Bl., and M. corticosum Ridl.,

the inflorescence originates from leafless nodes or from older branches. The

position, size, and structure of the inflorescence is of paramount importance in

the classification and identification of all taxa Memecylon.

CALYX

In addition to the anthers, the morphology of the calyx in all taxa of

Memecylon offers one of the most useful aids in identification. Memecylon

flowers are all pedicelled and in many taxa there is a distinct constriction above

the ovary which flares out to form a funnelform or campanulate calyx tube.

M. amplexicaule Roxb., M. intermedium BI., and especially M. minutiflorum

Mig. are some species which have prominent ovaries. Entirely campanulate

calyces without a noticeable ovary externally are less frequent within the genus,

but are typical for M. excelsum Bl. and M. oleifolium BI.

All taxa of Memecylon from the Malay Peninsula have glabrous flowers,

however 3 taxa (viz. M. dichotomum (Cl.) King, M. dichotomum (Cl.) King var.

rotundatum (Craib) Maxw. and M. minutiflorum Mig.) have a papillose (or almost

muricate) calyx. Sometimes M. cinereum King and M. heccarianum Coga. (from

Borneo) also have a papillose calyx. The margin of the calyx also provides

another feature which serves as a useful aid in identification. It ranges from

entirely truncate, with 4 minute cusps or undulations, to distinctly 4-lobed. There

are 9 taxa with an entirely truncate calyx margin, e.g. M. caeruleum Jack, M.

campanulatum Cl., M. excelsum Bl., M. dichotomum (Cl.) King, ete. Generally

speaking, the taxa in this group have a thickened calyx tube, while those taxa

with cusps or lobes have thinner tubes. In 7 taxa the margin varies from trun-

cate to cuspidate or undulate. Very often the young calyx in several taxa has

4 cusps which become very obscure when the calyx tube matures. Some examples

are: M. edule Roxb. var. edule, M. edule Roxb. var. ovatum (Sm.) Cl.; and

M. pubescens (Cl.) King (truncate to cuspidate); M. globosum Bakh. f. and M.

megacarpum Furtado (truncate to undulate).

There are 8 taxa which have 4 distinct cusps on the margin of the calyx

tube, e.g. M. garcinioides, Bl., M. minutiflorum Miq., and M. paniculatum Jack.

Finally, 5 taxa have 4 often acuminate calyx lobes, e.g. M. cinereum King, and

M. lilacinum Z. & M. M. acuminatum Sm. var. acuminatum, with a 4-lobed

calyx margin, is easily separated from M. acuminatum Sm. var. flavescens Cl.

which has a truncate margin.

Revision of Memecylon L. 4]

The internal septa found in all taxa of Memecylon are a vestige of the

extraovarial chambers which are more highly developed in other genera of

Melastomataceae, e.g. the tribe Dissochaeteae (Naud.) Triana. Most taxa have

8 distinct septa and 8 grooves in addition to a thinner line dividing each groove

(initially between the anther locules). In several taxa (e.g. M. cantleyi Ridl.)

the septa dividing the anther locules are almost as prominent as the ones sepa-

rating the anthers, thus the calyx has 16 septa and 16 grooves. The septa and

grooves in all taxa do not extend below the ovary and the manner in which

the anthers fit into these structures is most readily seen in mature buds. Aside

from M. oligoneurum Bl., which has very faint ridges and shallow grooves, the

septa and grooves have not proven to be very useful for identification purposes.

PETALS

There are 4 equal, symmetric, and glabrous petals in all taxa of Memecylon

from the Maiay Peninsula. Their shapes range from oblong to sub-orbicular

and they are often reflexed at maturity. The tips are broadly rounded to obtuse,

or acute; the bases clawed or truncate. Texture ranges from thin to thick

coriaceous, and in thicker petals the midrib is often thickened or dorsally keeied

There is some variation in petal morphology and colour (white, pink, blue, purple)

but these are not always reliable for identification purposes since, in comparison

to numerous vegetative and other floral characteristics, they are not as easy to

distinguish taxonomically as these other structures. ‘therefore, the petals nave

not been referred to very often in the key to flowering material.

STAMENS

The stamens of Memecylon are relatively smali and simple, and along with

those of Astronia and Pternandra are the least specialised anthers within the

family. All species of Memecylon have 8 equal, glabrous stamens with flattened

filaments and variously shaped, 2-locular, medifixed anthers which open by a

vertical slit. The connective, in most species, is well developed and in most taxa

there is also a crateriform gland. The anthers are usually described as being

dolabriform (axe-shaped), but this term is inadequate for comparative studies.

In many species the anthers in bud are not sufficiently developed to provide

recognizable distinctions, however in those species which lack a connective gland,

plus M. garcinioides Bl. and M. wallichii Ridl. which have unique shapes, helpful

clues can be found. The mature anthers provide the most important structural

evidence for the identification of several species, e.g. M. lilacinum Z. & M., M.

floridum Ridl., and M. pauciflorum BI.

The mature anthers of Memecylon can be separated into 4 basic groups:

“J”? or “‘C” shaped, “‘U”’ shaped, uniquely shaped, and glandless anthers. Mature

anthers with a “J’’ or ‘‘C” shape are characteristic of 18 taxa. Memecylon

excelsum Bl. and M. megacarpum Furtado have the largest anthers with very

thick connectives, but for the other taxa the variation is difficult to describe.

Several species (e.g. M. acuminatum Sm., M. caeruleum Jack, M. edule Roxb.,

etc.) have obviously different anthers and form seems to vary with different

stages of maturity. The variability of the curved anthers is not completely

known because of the lack of sufficient flowering material and detailed field

observations. There is a tendency for the anthers to curve from a nearly linear

shape in bud to crescent shape and then to a “C’’-shape. In some taxa, (eg.

M. edule Roxb. var. ovatum (Sm.) Cl.) some anthers are actually ‘‘U”’-shaped

due to more extensive bending. All these anthers have a thickened connective;

42 Gardens’ Bulletin, Singapore XX XIII (1980)

a distinct, approximately central gland, and locules which are smaller than the

connective.

There are 6 species with *“‘U’’-shaped mature anthers viz. M. cantleyi Ridl.,

M. fruticosum King, M. kunstleri King, M. paniculatum Jack, M. pauciflorum

Bl., and sometimes M. oleifolium Bl. M. lancifolium Ridl., which is insufficiently

known. also has (bud) anthers with this shape. In M. cantleyi Ridl. and M.

oleifolium Bl. some anthers show an asymmetric “‘C” outline. The anthers in

bud of these 7 species are usually more curved than those of the “J” or “C”’-

shaped anthers described above, however the present understanding of this group

is also incomplete. Indeed, some anthers of M. paniculatum Jack, perhaps due

to immaturity, are “‘C’’-shaped and resemble mature anthers of M. acuminatum

Sm. More observations of anther forms in these two groups are necessary in

order to understand both the variation in and the relationships among the various

taxa.

Anthers in the following taxa show unique shapes: M. garcinioides Bl. and

M. hullettii King (orbicular) and M. wallichii Ridl. (linear). Orbicular anthers

have a very reduced connective about as large as the locules with a small, often

obscure gland situated on a membranous extension of the connective. M. hullettii

King is, in addition, distinct from all other species of Memecylon by virtue of

its broadly rounded, subsessile leaves and compound — umbellate inflorescence.

The anthers of M. garcinioides Bl. are the most reduced among those studied,

and along with M. hullettii King are also the smallest in all taxa of Memecylon

from the Malay Peninsula. M. intermedium Bl. often has similar anthers, espe-

cially in bud, but the connective in most specimens is less reduced, thus giving

the anther a “‘J” shape. The two species can be easily distinguished by their

leaves and calyx margins. The linear shape of the bud and mature anthers of

M. wallichii Ridl. immediately distinguishes this species from M. dichotomum

(Cl.) King and M. corticosum Ridl. which have ‘‘C’’-shaped anthers and are

similar vegetatively. The locules of M. wallichii Ridl. are about the same size

as the connective which has a distinct, centrally located gland.

There are 5 species of Memecylon from the Malay Peninsula that have

glandless anthers, triangular or cuneate in form, which in many respects, resemble

those of Astronia smilacifolia Triana. These are: M. cinereum King, M. floridum

Ridl., M. malaccense (Cl.) Ridl., M: lilacinum Z. & M., and M. oligoneurum BI.

Apart from M. oligoneurum Bl., which is readily distinguished from all

other species of Memecylon in the region by its 3-nerved blades, the other 4

species in this group are often confused with other species. Both young and

mature anthers can be used to distinguish (1) M. floridum Ridl. from M.

acuminatum Sm., (2) M. lilacinum Z. & M. from M. garcinioides Bl. and M.

pauciflorum Bl., and (3) M. malaccense (Cl.) Ridl. from M. dichotomum (C1.)

King and M. acumiantum Sm. These species are difficult (except M. garcinioides

BI.) to distinguish solely by anther morphology, therefore various other charac-

teristics must be used to correctly identify each.

GYNOECIUM

The stigma is typically minute and is often difficult to distinguish from the

slender, glabrous style which is usually not much longer than the stamens and

merges with the top of the ovary without any disc or basal swellings. The

ovary is unilocular with very short free central (perhaps indistinguishable from

basal) placentation with 2-20 ovules attached in a whorl on a flattened, peltate

Revision of Memecylon L. 43

placenta. Memecylon oligoneurum Bl., as far as can be determined, has the

lowest ovule number (2), while most other taxa have at least 6 ovules in each

gynoecium. The number of ovules might ultimately provide a useful means to

help distinguish various taxa.

FRUIT

There are two basic fruit shapes in Memecylon viz. globose, or nearly so

(22 taxa) and oblong to elliptic (7 taxa). The fruits of M. acuminatum Sm. var.

flavescens Cl. and M. pauciflorum Bl. var. brevifolium Craib have not been

studied since fruits have yet to be described or collected from these two varieties.

However, it is most likely that they are globose as in their typical varieties.

Memecylon lilacinum Z. & M. usually has depressed, oblate, globose fruit; but

sometimes it is elliptic.

Within each group there are usually sufficient vegetative differences and

structural features of the infructescence to distinguish the taxa. In some species,

however, (e.g. M. lilacinum Z. & M., M. globosum Bakh. f., and M. excelsum

BI.) the fruit provides the most reliable diagnostic feature. Because flowers and

fruits are rarely found together on the same specimen (except in M. caeruleum

Jack) matching flowering material with fruiting specimens has often been difficult.

A good example is with M. minutiflorum Mig. and M. intermedium Bl. which

are easily confused unless collections are available. A similar situation exists

between M. excelsum Bl. and M. megacarpum Furtado, which have oblong and

globose fruit, respectively; but frequently have similar vegetative features. There

seems to be no correlation between vegetative characteristics, nature of the in-

florescence, and morphology of the anthers between species with globose or

oblong to elliptic fruit; except that the five species with glandless anthers all

have globose fruits.

The exocarp in dry specimens varies from smooth (e.g. M. cinereum King

and M. intermedium Bl., both with globose fruit; or M. caeruleum Jack with

elliptic fruit) to moderately rough (e.g. M. dichotomum (Cl.) King, and M. edule

Roxb., both with globose fruit, and M. paniculatum Jack which often has ovate

fruit). Very rugose or pustulate surfaces are typical of M. lilacinum Z. & M.,

and M. amplexicaule Roxb., both having globose fruit; and M. excelsum BI.

with oblong to elliptic fruit.

The colour of immature fruit is generally green, changing to yellowish, pink

or reddish, and finally in many taxa dark purple to blackish when ripe. The

dry colour ranges from black (e.g. M. edule Roxb. var. ovatum (Sm.) Cl. and

M. cinereum King) to tan-greenish (e.g. M. lilacinum Z. & M.), while M. cantleyi

Ridl. has fruit which dry light brown to tan.

The pericarp varies from very thin (c. 0.25 mm thick, e.g. M. cinereum King

and M. intermedium Bl.), moderately thick (c. 0.5 mm, e.g. M. edule Roxb. and

M. oligoneurum Bl.), to very thick (c. | mm or more and gritty in M. amplexi-

caule Roxb., M. lilacinum Z. & M. and M. megacarpum Furtado). M. floridum

Ridl. has a fibrous pericarp which immediately distinguishes it from the other

species. Generally speaking. species with ovate to elliptic fruits have a thinner

pericarp, the thickest being in M. excelsum BI. (c. 0.5 mm_ thick).

The calyx remnant, which is always present on immature and mature fruit,

varies somewhat in size and shape. A distinctly raised, crown-like remnant is

44 Gardens’ Bulletin, Singapore XX XIII (1980)

found in all taxa except M. amplexicaule Roxb. where it is flattened in the

plane of the fruit. All fruits have an areolus or space on the top enclosed by

the calyx remnant. The internal septa, most obvious in flowers, are also usually

distinct in fruits. The style in all taxa falls off with the petals and stamens

soon after the fruit begins to develop.

DISTRIBUTION AND RELATIVE ABUNDANCE

Memecylon is found in all the Malaysian States and Singapore. Several

taxa are found throughout the region, e.g. M. amplexicaule Roxb. and M. lilacinum

Z. & M., while others, e.g. M. malaccense (Cl.) Ridl., M. kunstleri King, and

M. globosum Bakh. f. are comparatively rare and have only been found in a

few localities.

Based on the data gathered from herbarium specimens, I have grouped

the taxa from the Malay Peninsula in four categories regarding their relative

abundance:

A. Very Common: M. amplexicaule Roxb. M. caeruleum Jack, M. dicho-

tomum (Cl.) King var. dichotomum, M. edule Roxb., var. edule, M.

edule Roxb. var. ovatum (Sm.) Cl., M. excelsum Bl., M. megacarpum

Furtado, M. minutiflorum Mig., M. lilacinum Z. & M., and M. oleifolium

BL;

B. Common: M. cantleyi Ridl., M. garcinioides Bl., M. intermedium BL,

M. oligoneurum Bl., M. paniculatum Jack, M. pauciflorum Bl. var.

pauciflorum, M. pubescens (Cl.) King, and M. wallichii Rid.

C. Not Common: M. acuminatum Sm. var. acuminatum, M. campanulatum

Cl., M. cinereum King, M. floridum Ridl., M. fruticosum King, and

M. hullettii King; and

D. Rare: M. acuminatum Sm. var. flavescens Cl., M. corticosum Ridl.,

M. dichotomum (Cl.) King var. rotundatum (Craib) Maxw., M. globosum

Bakh. f., M. kunstleri King, and M. malaccense (Cl.) Ridl.

MEMECYLON L.

Linnaeus, Sp. Pl. I (1753) 349; Smith in Rees Cyclopedia 23:4 (1813); Jack,

Malay Misc. 1:5 (1820) 26, II (1822) 62; Wallich Catalogue (1831); De Candolle,

Prodr. III (1828) 5; Roxburgh, Fl. Ind. ed. 2, II] (1832) 260; Blume, Mus. Bot.

Lugd. Bat. I:23 (1851) 353; Naudin, Ann. Sc. Nat. ser. H1:18 (1852) 264;

Miquel, Fl. Ned. Ind. suppl. I, Sumatra (1860) 323; Triana, Trans. Linn. Soc.

28 (1871) 155; Kurz, For. Fl. Brit. Burma I (1877) 515; Clarke in Hk. f., FI.

Brit. Ind. Il (1879) 553; Cogniaux in DC., Monogr. Phanerog. 7 (1891) 1130;

King, J. As. Soc. Beng. 69, Il: 1 (1900) 71 (Mat. Fl. Mal. Pen., 479); Ridley,

Fl. Mal. Pen. I (1922) 810; Craib, Fl. Siam. Enum. 1:4 (1931) 702; Bakhuizen

van den Brink, f., Med. Mus. Bot. Utrecht 91 (1943) 333 and Rec. Trav. Bot.

Neerl. 40 (1943-45) 333; Corner, Wayside Tr. Mal. I (1952) 448; Furtado, Gard.

Bull. Sing. 20 (1963) 119, -Backer & Bakhuizen van den Brink f., Fl. Java I

(1963) 371; Maxwell, Mal. Nat. J. 34:1 (1980).

Revision of Memecylon L. 45

Trees or terrestrial shrubs; branchlets cylindric, flattened and 2-grooved, or

4-angled to 4-winged; smooth, glabrous. Leaves simple, opposite, glabrous; blades

chartaceous, subcoriaceous, or thick coriaceous; entire; lanceolate to suborbicular;

with 1 main nerve (except in one species 3-nerved), secondary venation pinnate,

distinct to invisible, intramarginal nerves similar; petioles c. 1 mm to 15 mm long.

Stipules absent, interpetiolar ridge or line distinct on upper nodes, obscure to

invisible on older nodes. Inflorescence cymose, arising from leafy or leafless

nodes, less frequently terminal; the axes with up to 4 ramifications, or reduced

to multiflowered glomerules or umbels; glabrous or less commonly puberulent.

Calyx tube campanulate to funnelform, constricted or not above the ovary; margin

truncate, with 4 minute cusps, or undulations, to 4-lobed; tube glabrous or papil-

lose outside, with 8 internal ridges or lines, extraovarial chambers absent. Petals

4, thin or thick coriaceous; oblong, obovate, to orbicular; with or without a

thickened mid-line, usually reflexed at maturity, frequently colourful. Stamens

8, equal, glabrous; filaments flattened; anthers globose, axe-shaped, or curved

ina “J”, “C”’, or “U” shape; connective often thickened, with or without a

gland on the connective; 2-locular, each opening by a vertical slit. Stigma

minute, style slender; ovary inferior with one locule, with basal to free central

placentation; ovules 2-20, whorled on a flattened placenta. Fruit a_ berry,

globose or elliptic to ovoid; 1, rarely 2, seeded; pericarp juicy, fibrous, or gritty;

areolus distinct, raised or not; exocarp ripening red to purplish.

The genus Memecylon was established by Linnaeus in 1753 with a brief

description of M. capitellatum L. from Ceylon. Since then over 300 species

have been described from tropical Africa, Asia, Australia, and the Pacific Islands.

Memecylon has been traditionally included in the Melastomataceae by most

botanists, however a separate family. Memecylaceae DC. has been accepted by

Airy Shaw (Willis Dict. Fl. Pl. & Ferns) which includes four genera: Memecylon,

Axinandra, Mouriri, and Votomia — the former two being found in the Malay

Peninsula and-neighbouring countries; and the latter two from Central and Scuth

America, and the West Indies. Airy Shaw notes that Memecylaceae is more

or less intermediate between Myrtaceae and Melastomataceae. In this work

accord is made with most other botanists in maintaining Memecylon in the

Melastomataceae since the internal septae in the calyx tube and the morphology

of the stamens (especially the anthers) indicate relationship to Astronia and

Pternandra very closely and not to any genus in the Myrtaceae.

Memecylon, however differs from most other genera of Melastomataceae in

having uninerved blades (except M. oligoneurum Bl. and a few species from

Africa and Ceylon which have 3-nerved Blades); 8 equal, unappendaged anthers

with locules opening by slit (instead of pores as in most other genera); a unilocular

ovary with free central (almost basal) placentation; fewer ovules (up to 20); and

a (usually) single seeded berry. Therefore, the separation of Memecylon as a

subfamily, Memecyloideae, appears justified.

Key to the species of Memecylon from the Malay Peninsula based on

flowering material

1. Inflorescence sessile or nearly so; glomerulate with primary axes indistinct

or up to 2 mm long; leaf blades with 1 or 3 main nerves

2. Leaf blades with 1 main nerve;*calyx with distinct internal ridges;

ovules 6-20; anthers with a gland’

46 Gardens’ Bulletin, Singapore XX XIII (1980)

3. Branchlets prominently angled or winged, especially below the upper

node

4. Blade venation prominently raised on the undersurface; in-

florescence from leafy or leafless nodes (often ramiflorous);

pedicels ¢,, 3, mj dong. .<J..2+,.phie: ealeeese M. corticosum Ridl.

4. Blade venation obscure, not prominently raised on the under-

surface; inflorescence from leaf axils; pedicels 1.5-5 mm long

py ee ene een ene eee er RE EE M. fruticosum King

3. Branchlets cylindric

5. Blades sessile or very shortly (1-3 mm) petioled

6. Inflorescence glomerulate, primary axes absent; blades

thick coriaceous, veins obscure to invisible ............

sce sian Le aes § 02k SE ea M. amplexicaule Roxb.

6. Inflorescence cymose, primary axes at least 0.5 mm

long; blades subcoriaceous to coriaceous, venation in-

visible or distinct

7. Flowers 3-5 per inflorescence, bracts persistent;

pedicels 1-1.5 mm long; blades cordate at the

base, venation invisible\).tiqws... Bie a

5 ae M. dichotomum (Cl.) King var. dichotomum

(limestone form)

7. Flowers usually more than 5 per inflorescence,

bracts caduccus, pedicels 2-3 mm long; blades

narrowed at the base, venation distinct ............

nalgdbian ha deb cua eeciavine «cen MARTE M. megacarpum Furtado

5. Petioles over 3 mm long

8. Calyx truncate; anthers with a gland; inflorescence

mostly from leafless nodes

9. Blades thick coriaceous, 7-10 cm long, venation

INVIGIDIS. .0..... speek cata M. campanulatum Cl.

9. Blades coriaceous, 12—26 cm long, venation distinct

us pics one coh head ka a Ri daca ae M. excelsum BI.

8. Calyx with 4 broadly triangular lobes; anthers without

a gland; inflorescence from leaf axils .....................

ovale p's on sudie d avaee sy SOOO is beeen nna M. cinereum King

2. Leaf blades with 3 main nerves; calyx with faint internal ridges; ovules

2; anthers glandiegs ¢...)..:....-s00: nag aepneye ete M. oligoneurum Bl.

1. Inflorescence peduncled, primary axes at least 2.5 mm long; leaf blades with

] main nerve

10. Inflorescence 4-8 cm long, secondary axes at least 2 mm long, or if not

developed then the inflorescence is umbellate

Revision of Memecylon L. 47

11. Inflorescence umbellate; calyx tube muricate-papillose externally

12. Inflorescence a compound umbel, lowest internode of the pri-

mary axes 4-5 cm long, secondary axes 2-15 mm long; venation

meee. Stele) sshelt ... M. dichotomum (Cl.) King var.

rotundatum (Craibd) Maxw.

11. Inflorescence an open panicle of cymes or cymose, lower internodes

of the inflorescence 1-3 cm long, secondary axes distinct; venation

obscure or prominent; pedicels and calyx smooth, not muricate-

papillose

13. Veins prominent, raised on the undersurface

14. Branchlets cylindric

15. Inflorescence axes glabrous ............ M. oleifolium BI.

15. Inflorescence axes pubescent

16. Leaf blades 8-15 by 4-6 cm; petiole 1.5 mm thick

PENN etre ae i M. pubescens (Cl.) King

16. Leaf blades 14-30 by 6-8 cm; petiole 3 mm thick

Pr Peet ete ses atts M. beccarianum Cogn.

14. Branchlets 4-angled to 4-winged ... .. M. paniculatum Jack

13. Veins obscure, not raised below

17. Blades 6-13 by 2.5—5 cm; petiole 2-3 mm long, 1-1.5 mm

thick; inflorescence 2-4.5 cm long with sharply 4-angled

ght een i at ttn aguas | sean ees M. kunstleri King

17. Blades 10-20 by 5-10 cm; petiole 7-10 mm long, 2-3 mm

thick; inflorescence 7-9 cm long with flattened or cylindric

CRS es ete eee, cases M. oleifolium BI.

10. Inflorescence a contracted cyme or umbel, 0.44 (5) cm long, secondary

and/or tertiary axes up to 2 mm long, often indistinct

18. Inflorescence axes slender, 1 mm or less thick, or flattened and

1-2 mm wide; blades acute, cuneate, or rounded and usually decu-

rent at the base

19. Inflorescence sub-sessile or up to 1.5 cm long

20. Leaf blades somewhat narrowed, rounded, or shallowly

cordate at the base, not er only slightly decurrent at the

base

21. Veins conspicuous and often slightly raised below;

upper branchlets sharply 4-angled to 4-winged; anthers

with a gland

22. Upper internodes 4-winged, oftering tapering to

4-angles; bracts thin, caducous; pedicels clustered

on each secondary axis; calyx smooth

PaE:

Gardens’ Bulletin, Singapore XX XIII (1980)

23. Anthers ‘‘U’’-shaped; blades lanceolate or

ovate, 5-13 cm long ...... M. fruticosum King

23. Anthers slightly curved to almost “‘C’’-shaped;

blades lanceolate, 10.5—-18 cm long ............

et eR M. corticosum Ridl.

Upper internodes 4-angled tapering to cylindric;

bracts thickened, persistent; pedicels solitary on

each secondary axis; calyx papillose ...............

Wve M. dichotomum (Cl.) King var. dichotomum

21. Veins obscure to invisible; upper branchlets cylindric,

often 2-grooved or somewhat 4-angled below the upper

node; anthers with or without a gland

24.

Calyx thin, 1.5-2 mm long; anthers without a

plagigl <7. hig cece meee M. malaccense (Cl.) Ridl.

Calyx thick, at least 2 mm long; anthers with a

distinct gland

25. Branchlets drying brown to blackish; calyx

truncate: bracts persistent) ...:..\....l: ese

SoheeeteCi. Pe nae oc eanaane M. caeruleum Jack

25. Branchlets drying tan; calyx with 4 broad

lobes; bracts caducous .,......:¢1.5 eee

20. Leaf biades narrowed and often decurrent at the base;

venation obscure to invisible, or prominently raised on the

undersurface; branchlets not 4-winged (often 4-angled in

M. lilacinum and M. excelsum)

26. Upper branchlets grooved on 2 sides, each flanked by

2 ridges, or sharply 4-angled (becoming obscure on

older branches and appearing cylindric)

27.

Pedicels, ovary, and calyx minutely papillose, tex-

ture roughened; petals obtuse to acute at the tip

rae DC eA ee M. minutiflorum Miq.

Pedicels etc. smooth; petals caudate- acuminate at

the tip

28. Anthers without a gland; flowers usually more

than 5 per inflorescence

29. Petals lanceolate to oblong .................

Nylon sie M. lilacinum Zoll. & Mor.

29. Petals broadly ovate

30. Inflorescence 5-7 mm long, pedicels

c. 1 mm long, blades drying black

5 «GR 4S eae M. cinereum King

Revision of Memecylon L.

30.

Inflorescence 10-15 mm long, pedi-

cels c. 4 mm long, biades drying

STCORICNE oc wos M. floridum Ridl.

28. Anthers with a distinct gland: flowers c. 5

per inflorescence

oi Blages Sa DY 1-25 CM os i snccgcncceenies=>

er eee

M. pauciflorum Bl. var. pauciflorum

DE Teens ee Toy Cs TCR 2 oats 3

M. pauciflorum BI. var. brevifolium Craib

26. Upper branchlets cylindric, often slightly flattened,

shallowly grooved, or somewhat 4-angled below the

upper node

oan

Calyx lobes reduced to 4 extremely minute points

or completely truncate; petals broadly ovate to

sub-orbicular, acute or mucronate at the tip

33. Calyx c. 1-1.5 mm long, c. 1.5 mm wide

33.

34. Bud and mature anthers circular in out-

line, with the gland on a thin extension

of the connective; calyx with 4 minute

Ormelsic. ae.) .. M. garcinioides BI.

34. Bud and mature anthers not circular in

outline, gland not as above or absent

35.

ad.

Bud and mature anthers curved in

a “C” shape, gland on the thickened

COnneChive, “Calyx ‘truncate. !).J.22...:.

a2) eres aes M. acuminatum Sm. var.

flavescens Cl.

Bud and mature anthers straight,

without a gland; calyx with 4 short,

triangular cusps ... M. floridum Ridl.

Calyx 2-3.5 mm long, 3-5 mm wide; bud

anthers curved in a ““C” or hook shape, gland

on the thickened connective distinct

36. Calyx campanulate, not constricted above

the ovary

of.

Leaf blades thick coriaceous, vena-

tion obscure to invisible ...............

Leaf blades subcoriaceous, venation

visible and raised on the _ under-

Barta 4 oi, wane M. excelsum BI.

Gardens’ Bulletin, Singapore X X XIII (1980)

36. Calyx flattened campanulate to funnel-

shaped, sharply constricted above the

Ovary: 2.4 ee M. constrictum Craib

(q.v. M. minutiflorum Miq.)

32. Calyx with 4 distinct cusps, each c. 0.25 mm high,

or distinctly 4-lobed, the lobes undulate or 1/3

as long as the calyx tube; petals lanceolate-oblong,

or elliptic-oblong, the tips not mucronate

38.

Leaf veins invisible; petals lanceolate-oblong

and longer than wide, caudate-acuminate at

the tip; anthers without a gland ...............

pani atend, eee M. lilacinum Zoll. & Mor.

Leaf veins prominent; petals broadly elliptic

to oblong, wider than long; acute, broadly

rounded, or truncate at the tip; anthers with

a Gland Ny, Vor. Se een M. excelsum Bl.

19. Inflorescence 1.5-—5 cm long

39. Bud and mature anthers circular in outline with the gland

along the entire surface of a thin, flat connective appendage

Lee le an cae Sa aera M. garcinioides Bl.

39. Bud and mature anthers “J”, “C’, or ‘“‘U”’-shaped; gland

centrally located on the thickened connective

40. Calyx with 4 triangular cusps or lobes; venation of

blades visible on the undersurface ......................06

Rian hae M. acuminatum Sm. var. acuminatum

40.

Calyx truncate or with 4 minute cusps or 4 undulate

lobes; blade venation very obscure to invisible on the

undersurface

41.

Inflorescence with 3-10 flowers; petals thin with

visible. venation. ici. cake M. cantleyi Ridl.

Inflorescence usually with more than 10 flowers;

petals coriaceous, with obscure to invisible veins

42. Branchlets cylindric below the upper node

43. Blades 6-9 by 2-3 cm, petiole 3-4 mm

long; inflorescence 1.5—2 cm long, secon-

dary axes not developed (glomerulate on

the primary axis) or up to 2 mm long;

calyx with 4 minute cusps or lobes ...

+ edict Uasiiglin cosas M. intermedium BI.

43. Blades (6) 9-21 by (2.5) 3-10 cm, petiole

6-9 mm long; inflorescence 1.5—5 cm

long, secondary axes 2-12 mm _ long;

calyx truncate ......... M. oleifolium BI.

Revision of Memecylon L. 5]

42. Branchlets 2-grooved to 4-angled below the

upper node

44. Petiole 4.5-7.5 mm long, 2.5-4 cm wide;

inflorescence (0.5) 1.5=2 cm long .........

eae ghee 2 eat M. edule Roxb. var. edule

44. Petiole 6-15 mm long; blades 5-12 cm

long, 3—6.5 cm wide; inflorescence up to

KS ight ER nite oe re esa

M. edule Roxb. var. ovatum (Sm.) Cl.

18. Inflorescence axes robust, flattened or not, 1.5-2 mm thick: blades

broadly rounded to cordate at the base, less frequently with a

slightly cordate-auriculate base

45. Veins on leaf blades raised on the undersurface, intramarginal

nerve distinct, blades 14-27 cm long; midnerve of petals slightly

raised dorsally, margins with a narrow thin portion (less than

1/10 mm wide)

46. Anther locules extending all along one side of the con-

nective and upper part of the other, connective shorter

than the locules, gland minute, round; branchlets 4-angled;

shrub or tregeup tee-in tae 73 oo M. wallichii Ridl.

46. Anther locules on the enlarged tip of the connective, con-

nective longer than the locules, gland elongate, prominent;

branchlets cylindric; trees 10-15 m tall ... M. excelsum BI.

45. Veins on leaf blades obscure to invisible on the undersurface,

intramarginal nerve invisible; blades 8-14 cm long; mid-nerve

of petals prominently keeled dorsally, margins with a prominent

thin portion (at least 0.25 mm wide) ...... M. caeruleum Jack

Key to the species of Memecylon from the Malay Peninsula based on

fruiting material

1. Mature fruit subglobose to globose, i.e. about as long as wide

Mature fruit 1.2-2 cm diameter

Axes of infructescence indistinct, glomerulate; areolus flat, not raised

above the plane of the fruit; leaf blades thick coriaceous, veins

o_O tg oli tal ed och ee a Riles ae Ae M. amplexicaule Roxb.

Axes of infructescence at least 3 mn long, cymose; areolus distinctly

raised above the plane of the fruit, often flattened; leaf blades sub-

coriaceous, veins distinct

4. Upper branchlets 4-angled

5. Leaf blades 5-10 cm by 2-5 cm; secondary axes 3, distinct

he te ee M. dichotomum (Cl.) King var. dichotomum

5. Leaf blades 14-25 by 3.5—10 cm; secondary axes indistinct,

clustered at the tips of the primary axes .....................

A EERE EE S12 Ee M. wallichii Ridl.

Gardens’ Bulletin, Singapore XX XIII (1980)

Upper branchlets cylindric

Leaf blades 15-20 by 6-7.5 cm, the veins slightly raised

on the undersurface; areolus slightly (1 mm) raised ......

j120)}. sonmmepiotld... case M. megacarpum Furtado

Leaf blades 7-10 by 3.5—5 cm, the veins not raised on the

undersurface, often invisible; areolus flattened in the plane

of thé frait!.a2t not, raised 1. 5a, M. globosum Bakh. f.

Mature fruit 2-12 mm diameter

7. Upper branchlets cylindric, flattened and grooved on 2 sides, or

somewhat 4-angled near the nodes

Veins of blades sunken above, raised on the undersurface

Fruit smooth when dry; blades 14-30 cm by 6-8.5 cm;

petiole. c. 3 mm stick sc. wtleeeiee: M. beccarianum Cogn.

Fruit rough when dry; blades 8-12 by 4-5.5 cm; petiole

15 oman | Ae bas ea eieoak frat M. pubescens (Cl.) King

Veins of blades obscure to invisible on both surfaces

10. Upper branchlets somewhat flattened wiih a vertical groove

flanked by two ridges on opposite sides of the petioles,

or 4-angled throughout

11. Fruit flattened at the poles; pericarp 1-2 mm thick

and gritty, often grooved near the base

12. Blades drying black above, brown on the under-

surface; fruit drying black; infructescence glome-

rulate, up to 10 mm.long ...4..c.Jusisue see

i oie ele Ree ane anne M. lilacinum Zoll. & Mor.

12. Blades drying brown above; green, tan, to yello-

wish on the undersurface; fruit drying tan to

grey-greenish; infructescence cymose, the axes

10-15 mm long ............ M. minutiflorum Miq.

11. Fruit globose throughout, usually not flattened at the

poles; pericarp up to 0.5 mm thick, not gritty or

grooved

13. Primary axes of the infructescence less than

1.5 mm long

14. Exocarp rough when dry; blade undersurface

and fruit drying tan, yellowish, to grey-

greenish 43 .Jieskeudiwe M. minutiflorum Miq.

14. Exocarp smooth when dry; blade undersurface

and fruit drying brown

Revision of Memecylon L.

13.

15. Infructescence from the axils of older

leaves or from leafless nodes; primary

axes 6-19 mm long, secondary axes 4-5

mm long ...... M. edule Roxb. var. edule

15. Infructescence from the axils of upper

leaves; primary axes 1.5-2 mm _ long,

secondary axes indistinct and reduced to

a tubercle c. 0.5 mm long

16. Leaf blades 3-6 by 1-3 cm .........

M. pauciflorum Bl. var. pauciflorum

16. Leaf blades 1.2-1.7 by up to 1 cm

Be Pan M. pauciflorum Bl. var.

brevifolium Craib

Primary axes of the infructescence less than

1.5 mm long

17. Leaf blades and fruit drying black ............

Fo PRE ORE e e es TP Ae Tee M. cinereum King

17. Leaf blades drying browish above, greenish

on the undersurface; fruit drying greenish-

DEO WH dni Say M. edule Roxb. var. edule

10. Upper branchlets cylindric, somewhat 4-angled, or flattened

with a shallow groove on 2 sides, below the upper node

18. Blades with 3 main nerves from the base; exocarp

with a grey-mealy texture when dry .....................

SUN er a ag Bel Bt ihe aye M. oligoneurum BI.

18. Blades with 1 main nerve (midvein) from the base;

exocarp not with a grey-mealy texture when dry

19.

he!

Infructescence an open, compound umbel; primary

axes 4-5 cm long; blades broadly rounded to

cordate at the base, petiole 1-2 mm long .........

Peer a i gtents Vasashded ote «scarceck M. hullettii King

Infructescence cymose, often very compact to

glomerulate; primary axes shorter; blades acute

and decurrent at the base; petiole more than

2 mm long (except M. constrictum)

20. Infructescence axes up to 5 mm long

21. Blade venation invisible; infructescence

from behind the leaves; primary axes up

to 1.5 cm long

22. Infructescence glomerulate, of 10 or

more pedicels, from behind the

leaves; blades thick coriaceous, pe-

Coke: S12 Ms iene nse cavn x

TE Ae eda oe M. campanulatum C1.

54 Gardens’ Bulletin, Singapore XX XIII (1980)

22. Infructescence cymose, of 3-5 pedi-

cels, from leaf axils; blades coria-

ceous; petiole c. 2 mm long .........

. M. dichotomum (Cl.) King var.

dichotomum (limestone form)

Blade venation visible on the under-

surface (often obscure); infructescence in

leaf axils; primary axes at least 3 mm

long

23. Petiole 2-4 by; 1-15 mn fae

sia ale. ee pha ene M. garcinioides BI.

23. Petiole ¢:°2°by 2-255 ami eee

ot tales ara eae M. constrictum Craib

(q.v. M. minutiflorum Miq.)

20. Infructescence axes at least 7 mm long

24.

Petiole 7-15 mm long; blades (5) 8-12.5

by 3-6.5 cm

25. Fruit drying tan-brown, exocarp

smooth, pericarp c. 0.2 mm _ thick

lores. Uedidack M. cantleyi Ridl.

25. Fruit drying black or greenish, exo-

carp rugose, pericarp 0.5—1 mm thick,

EG) RRS Med oe a M. edule Roxb. var.

ovatum (Sm.) Cl.

Petiole less than 7 mm long; blades 4—9

by 2-4.5 cm

26. Blades coriaceous, veins invisible

27. Infructescence mostly from be-

hind the leaves; blades drying

greenish to brown, obtuse to

acute at the. tipis teen ee

Peay’ M. edule Roxb var. edule

27. Infructescence mostly from leaf

axils; blades drying dark brown

to black, acuminate at the tip

Meer i ee M. intermedium BI.

26. Blades chartaceous to sub-coriaceous,

veins visible

28. Blades acute at the base; petiole

3-6 mm long

Revision of Memecylon L. 55

29. Fruit 6-7 mm _ diameter,

pericarp c. 1/3 mm thick;

blades broadly ovate to

oo 1 CR ee On A Oe Be

M. acuminatum Sm. var.

acuminatum

29. Fruit 10-12 mm diameter,

pericarp fibrous, 1-2 mm

thick, blades lanceolate-

To eh eee ee eee

28. Blades broadly rounded to shal-

lowly cordate at the base, petiole

1-2 mm long

30. Primary axes of the infruc-

tescence up to 6 mm long,

Pete 0) ae ne ee

M. dichotomum (Cl.) King

var. dichotomum

30. Primary axes of the infruc-

tescence 15-20 mm _ long,

Deri) hack»... Pevsoactier.

M. dichotomum (Cl.) King

var. rotundatum (Craib)

Maxw.

7. Upper branchlets generally 4-winged or at least distinctly 4-angled

along the entire length of the internodes

31. Petiole 2 mm thick; blades 11-26 by 3.5-14 cm, broadly

rounded to cordate at the base

32. Infructescence axes (total length) 3-4 cm, from behind the

leaves; trees 10-20 m tall ............ M. paniculatum Jack

32. Infructescence axes 1-1.5 cm long, generally from leaf

axils; shrubs or trees up to 6 m tall ...... M. wallichii Ridl.

31. Petiole 1-1.5 mm thick; blades 5-13 by 2-6 cm, rounded to

narrowed at the base

33. Blades thin coriaceous, veins usually visible (often obscure):

branchlets 4-winged to 4-angled

34. Primary axes up to 6 mm long, c. 1 mm thick

35. Upper branchlets 4-angled, tapering to cylindric;

bracts persistent; secondary axes not glomerulate,

Spree Aiea TUN MONEE No Ai Goss Sox Psa Shoe Cad bn bs <cau od

rhe M. dichotomum (Cl.) King var. dichotomum

56 Gardens’ Bulletin, Singapore XX XIII (1980)

35. Upper branchlets 4-winged, tapering to 4-angled;

bracts not present in fruit; secondary axes glome-

rulate or up to 1:mm Jong”... eee

34. Primary axes 15-20 mm long, c. 1/3 mm thick ......

en 4: M. dichotomum (Cl.) King var. rotundatum

(Craib) Maxw.

33. Blades coriaceous, veins invisible; branchlets 2-grooved to

AMMO sks in pe M. malaccense (Cl.) Ridl.

1. Mature fruit ovate to elliptic, ic. longer than wide

36. Infructescence axes up to 2 cm long, 3d and 4th axes usually not

developed

37. Mature fruit grooved near the base, often flattened at both ends,

areoluscockS mim. wide |. cc: teanc eee M. lilacinum Zoll. & Mor.

37. Mature fruit not grooved or flattened; areolus at least 2 mm wide

38. Fruit 8-10 mm long, areolus c. 2 mm wide

39. Blades coriaceous; ovate, acute, and often decurrent at

the base

40. Infructescence axes 6-10 mm !ong ... M. excelsum BI.

40. Infructescence axes at least 15 mm long ...............

wk a SOR is See a a M. oleifolium BI.

39. Blades subcoriaceous, lanceolate to narrowly ovate, rounded

to shallowly cordate at the base

41. Branchlets * cylingric). <e.ccemeaeoa M. kunstleri King

41. Branchlets 4-winged

42. Blades lanceolate, 10.5-18 cm long ..................

eee er M. corticosum Ridl.

38. Fruit 10-20 mm long, areolus 3-5 mm wide

43. Fruiting bracts persistent; exocarp smooth; blades coria-

ceous, veins mostly invisible ............ M. caeruleum Jack

43. Fruiting bracts caducous; exocarp rugose-pustulate; blades

subcoriaceous, veins distinct .................. M. excelsum BI.

36. Infructescence axes 2-8 cm long, 3d and 4th axes usually developed

Revision of Memecylon L. 57

44. Secondary veins sunken above, prominently raised on the under-

surface; branchlets drying brown, upper internodes usually 4-angled

ee emrte toe ie. fend os va i. M. paniculatum Jack

44. Secondary veins faint or invisible, not sunken above, invisible to

slightly raised on the undersurface; branchlets drying greyish to

khaki, upper internodes cylindric ..................... M. oleifolium BI.

fruit unknown: M. acuminatum Sm. var. flavescens Cl.

M. pauciflorum Bl. var. brevifolium Craib

1. Memecylon acuminatum Sm., Rees Cyclop. 23 (1813) var. acuminatum

Tree up to 15 m tall with a diameter of 15 cm; branchlets slender and often

hanging; upper internodes cylindric, often flattened and grooved below the nodes,

smooth, drying brown; blades subcoriaceous, broadly elliptic to ovate, acuminate

to caudate (up to 12 mm) at the tip, narrowed and decurrent at the base, mid-

nerve sunken above, raised below, secondary venation very obscure or invisible

on both surfaces; 5-7 cm long, 1.5-3.5 cm wide, dark glossy green drying

brownish to olive-green above, dull green below when dry, entirely glabrous;

petiole 5-7 mm long, c. | mm wide, glabrous; inflorescence from leaf axils,

loosely cymose with many branches, (1) 2-3 cm long, with (10) 20-30 flowers,

glabrous, axes flattened, bracts and bracteoles not seen, falling off before the

petals mature; primary axes solitary, 10-20 mm long with 1 or 2 nodes, c. 1 mm

wide at the base, secondary axes c. 5 from each primary node, 2-3 mm long

with 2-3 nodes, tertiary axes c. 2 mm long, 4th axes not developed or up to

0.5 mm long, pedicels 1-2 mm long; calyx campanulate, wider and becoming

flattened above the ovary, glabrous, c. 1 mm long, 1-1.5 mm wide, margin with

4 well developed triangular lobes, each c. 0.25 mm long; petals broadly ovate

to sub-orbicular, acute at the tip, truncate at the base, thickened with thinner

margins, c. 2-2.5 mm long and of similar width; anthers c. 1 mm long, crescent

shaped, with a prominent gland centrally located on the connective; stigma minute,

style c. 3.5 mm long, glabrous; fruit globose, often depressed, 6-7 mm diameter,

calyx remnant distinct, flat, areolus c. 4 mm wide; exocarp pale green when

immature, drying straw yellow with a slightly roughened texture, c. 0.3 mm thick

Figure 1: a. calyx and corolla bud, b. petals, c. bud stamen, d. mature

stamen

This species is sometimes confused with M. floridum Ridl. (q.v.) which has

glandless anthers, a shorter inflorescence, larger fruit, and angled branchlets.

M. intermedium Bl. (q.v.) has a shorter inflorescence with reduced, often

glomerulate, secondary axes. Even in specimens of M. acuminatum with very

reduced axes, e.g. Ridley 3297, 6413, 7309; and Foxworthy 1194, the nodes of

the secondary and tertiary axes are visible. The blades in M. intermedium tend

to dry black above, and brown below.

Kedah — Pulau Dayang Bunting: Corner sn, 13 Nov. 1941

Perak — Larut: King’s collector 3458, 6754

Kelantan — G. Stong: Symington 37731

Selangor — Bukit Kutu: Ridley 7309; Sempang Mines: Ridley 15591, 15618

58 Gardens’ Bulletin, Singapore XX XIII (1980)

Pahang — Ulu Rembau: Nur 11778

Malacca — Batu Tiga: Derry 1041; Gunong Ledang: Ridley 3297; Pokoh Magas:

Alvins 765; sine loc.: Griffith 2325/1

Johore — G. Belumut: Holttum 10779; Kluai Yong Estate: Corner 36295;

Gunong Pulai: Mat 3741; Peuyabong: Foxworthy 1194; Tana Runto: Ridley

4656; Tanjong Bunga: Ridley 6412, 6413, Sungai Bau: Ridley s.n. in 1894

Singapore — Ridley 4574, 6411

la. Memecylon acuminatum Sm. var. flavescens Cl. in Hk. f., Fl. Brit. Ind. II

(1879) 562.

This variety differs from the typical variety by the completely truncate calyx.

The differences, according to Clarke, are that var. flavescens has more rigid blades,

which dry yellowish; and shorter (c. 1 cm long) cymes. The anthers appear

to be quite similar to those of typical M. acuminatum. The blades, especially

with their rostrate tips, also compare well.

King (J. As. Soc. Beng. 69, If : I (1900) 81 and Mat. Fl. Mal. Pen. III,

489) and Craib (Fl. Siam. Enum. 1:4 (1931) 709) both considered var. flavescens

as a synonym of M. minutiflorum Mig. Their opinions are not accepted here

since the structure of the inflorescence, smooth calyx, and anthers all differ signifi-

cantly. The upper internodes of var. flavescens are cylindric and not 4-angled

or 2-grooved as with M. minutiflorum. The affinities of var. flavescens lie with

M. acuminatum and not M. minutiflorum; however since only 4 collections of

var. flavescens were studied during this research I cannot be certain of any varia-

tions or to which other species of Memecylon it may be related. Fruiting

material of var. flavescens has not been seen or described.

Figure la: a. calyx, b. petal, c. mature stamen

Penang

I.H. Burkill 2655: Curtis 815, 816

Malacca

Griffith 2325/2 (type K, L)

2. Memecylon amplexicaule Roxb., Fl. Ind. I (1832) 260 (M. amplexicaulis);

M. microstomum Cl. in Hk. f., Fl. Brit. Ind. If (1879) 557; King, J. As.

Soc. Beng. 69, If : I (1900) 79 (Mat. FI. Mal. Pen. HI, 487), syn. by Ridley,

Fl. Mal. Pen. I (1922) 815; Craib, Fl. Siam. Enum. I : 4. (1931) 703..

Tree 6-18 m tall, diameter 15-75 cm; bark smooth, grey-brown, thin,

shallowly fissured and cracked; slash inner bark thin, brown or orange; slash

wood brown or orange; bole straight, crown spreading; branches cylindric, smooth,

c. 2 mm thick, drying light brown, nodes thickened; blades coriaceous, ovate

or elliptic, obtuse or with an obtuse acumen c. 5 mm long, base rounded or

more often (especially in more mature or larger blades) cordate and clasping

the branch; 7-15 cm long, 3-8 cm wide; veins generally invisible above, invisible

or very obscure below, intramarginal nerve invisible; midnerve distinct and

sunken above, prominently elevated below, thickened near the petiole and taper-

: ee @¢ ew = oe | ee ee ee

Revision of Memecylon L. 59

ing towards the leaf tip; dark glossy green above, light yellowish-green below,

drying brown to greenish (often with a mottled pattern), greenish to lighter brown

below; petiole 0.5—2, less frequently up to 3 mm long, c. 2 mm-wide; leaf scars

prominent, becoming elevated on older branches; inflorescence sessile, glomeru-

late, in leaf axils, glabrous, composed of every reduced cymes united in compact

and globose fascicles; bracts and bracteoles lanceolate-ovate, acute, c. 0.5 mm long,

persistent; primary and secondary axes not apparent, united axes 5-8 (rarely 10)

mm diameter, entire inflorescence 1.5-2.5 cm wide; pedicles numerous, 15-25

per axil, 2-3 mm long, white to pinkish; calyx campanulate, c. 1.5-2 mm long,

1.5-2 mm wide, margin shallowly undulate or with 4 distinct, broadly triangular,

acute lobes; glabrous and smooth outside, prominently ridged inside, white to

pinkish; petals broadly ovate to suborbicular, acute at the tip, truncate at the

base, 1.5-2 mm long, 2 mm wide, thin with the venation visible and a prominent

and thickened dorsal midnerve, margins thinner, ventrally cocave at maturity,

white to cream coloured; filaments 2.5-3 mm long, glabrous, white; anthers

curved in a “‘)” shape, connective white, gland tan, distinct; style slender c. 4 mm

long, white; ovules 5—12, erect, reniform, and flattened; ovary slightly wider than

the pedicel, much narrower than the calyx, c. 1 mm wide; fruit globose, slightly

flattened at the apex, glabrous, 1.2-2 cm diameter, pedicels 2-3 mm, calyx

remnant sunken at the top without any tissue prcjecting above the plane of the

fruit, c. 2 mm wide, internal ridges generally obscure, style scar distinct; pericarp

c. 2 mm thick, pulpy, exocarp smooth, greyish-green to dark green and hard,

turning red, purplish and soft when ripe, drying tan to light brown with a

roughened texture, seed globose, 6-8 mm diameter, testa blackish, smooth.

Figure 2: a. calyx, b. petal, c. stamen

vernacular: nipis kulit, pantai ulat, kelat, daun pekan, dalak

properties: leaves boiled together with onions form a liquid which, when used

as a poultice and applied to the head, cures headaches

uses: charcoal

habitat: primary forest, swamp forest, 600-1200 m elevation

The glomerulate inflorescence and the thick blades are similar to M. multi-

florum which differs in having longer petioles, blades with acute bases, and a

truncate calyx. The anthers appear to be the same. M. caeruleum Jack (q.Vv.)

differs, among other traits, in having distinct primary axes, larger flowers, and

elliptic fruit.

Kedah — Bukit: Enggang: Everett 13747; Bukit Perak For. Res: Chan 13128,

13133, Everett 13684: Koh Mai For. Res.: Kiah 35182; Ulu Muda For. Res.:

Bray 11511; Kedah, Haniff & Sallah 10475

Penang — Curtis 766, 3592, s.n.

Perak — Kampong Tera: Ismail 95021; Padang Batang: Shing 17388; Ulu

Bubong: King’s coll. 10588; Ulu Kenderong: Hamid 11615

Kelantan — G. Rabong: Shah 2484; Kampong Gobek: Shah & Kadim 484:

Sungei Galas: Cockburn 7477; Sungei Merkeh; Shah & Shukor 3178

Trengganu — Jambu Bongkok For. Res.: T. & P. 12, 439 (3030); Sekayu For.

Res.: Shing 13538; Ulu Bendong: Corner 30003, 30188

60 Gardens’ Bulletin, Singapore XX XIII (1980)

Pahang — Aur For. Res.: Whitmore 3683; Bukit Blakang: Haniff 21047; Bukit

Tersit; Shah & Shukor 2628, Soepadmo 837; Fraser’s Hill: Henderson 11515,

Henderson & Nur 11212; Gunong Benom: Ismail 97802; Gunong Berembun:

Ismail 104897; Kuala Teku: Kiah 31740; Baloh For. Res.: Mahamud 0820;

Labis For. Res.: Ahmad & Shukor 522

Negri Sembilan — Gunong Angsi: Loh 17310; Jelebu: Everett 104909A

Malacca — Kedah Peak: Maingay 821 (1565) (type M. microstomum Cl.);

Nyalas: coll. ? 797; Sungei Udang: Alvins 24

Johore — Gunong Besar, Labis For. Res.: Everett 13994, 14098; Gunong Blumut:

Whitmore 8829; Kota Tinggi: Corner 30781; Kluai: Corner 29953; Labis For.

Res.: Suppiah 104957; Ma’okil For. Res.: Shah 3666, Shing 6861; Panti For.

Res): Everett) 13812

Singapore — Cantley sn; Henderson 36357; Ngadiman 37009; Ridley 2023, 9182,

15622

Further Distribution — peninsular Thailand: Surat (fide Craib, 703).

3. Memecylon beccarianum Cogn. in DC., Monogr. Phanero. 7 (1891) 1143.

Tree, 10-18 m; bark smooth, thin, reddish, brittle; branches cylindric, smooth,

c. 2 mm thick; blades sub-coriaceous, broadly lanceolate to elliptic, acuminate

at the tip, narrowed and slightly decurrent to somewhat rounded and shallowly

cordate at the base; venation pinnate, 15—20 pairs of nerves, sunken above, pro-

minently raised below; intramarginal nerve similar, 1.5—3 mm from the margin,

looping; midnerve sunken above, raised and tapering to the tip below. 14-30 cm

long, 6-8.5 cm wide; drying olive-green to brown above, greenish to light brown

below; entirely glabrous; petiole 6-9 mm long, 2-3 mm thick, glabrous; infore-

scence a panicle of cymes, many flowered, from leafy or upper leafless nodes;

3-10 cm long, axes 4-angled, often flattened and grooved, minutely muricate-

furfuraceus, becoming glabrous; bracts-and bracteoles lanceolate, acute 0.25—0.5 mm

long, caducous; primary axes usually solitary, 6.5-9 cm long, with up to 4 nodes,

secondary axes up to 2.5 cm long with 1 node, terminated by 15-20 pedicles,

1.5-2 mm long, clustered at the tip; calyx campanulate, often constricted above

the ovary, truncate with 4 minute cusps, at first minutely muricate-papillose,

becoming smooth, ridged internally, 1.5 mm high, 2 mm wide; petals thickened,

broadly ovate to suborbicular, acute at the tip, truncate at the base, margins

thinner, midvein slightly raised dorsally; 2 mm long, 2.5 mm wide, light blue;

filaments c. 2 mm long, anthers “‘C”’-shaped, gland distinct; style c. 4 mm long;

ovules c. 16; fruit globose, 7-8 mm wide, capped by the raised calyx remnant,

areolus c. 2 mm wide; exocarp thin, smooth, drying tan to black; pericarp c.

0.2 mm thick.

Figure 3: a. calyx and style, b. petal, c. stamen. Figure 32

Memecylon beccarianum Cogn. is very close to M. paniculatum Jack (q.v.)

and can be distinguished by the cylindric internodes in the former and 4-angled

to 4-winged branches in the latter. The structure of the inflorescence, calyx,

petals, and anthers are virtually the same. M. paniculatum is a rather variable

species and several other species have been reduced to synonyms of it. It is

quite possible that M. beccarianum Cogn. is also the same since the relative

Revision of Memecylon L. 61

nature of the branches is not a good feature meriting maintenance of a separate

taxon. As sufficient authentic material of M. beccarianum has not been seen

(types Beccari 1311, 1518, from Matang, Borneo) it is difficult to determine its

exact affinities with M. paniculatum merely from the orginal description and a

few specimens. Therefore, pending a detailed study of the two species, M.

beccarianum is maintained as a distinct species in this paper. As far as can

be determined M. beccarianum has only been collected in Borneo. The infore-

scence, especially the pubsecence, of M. pubescens (Cl.) King (q.v.) is also similar,

however the leaves of that species are smaller, more acute at the base, and there

are differences with the calyx and petals.

Memecylon oleifolium Bl. (g.v.) can be separated from M. beccarianum by

the cylindric inflorescence axes which are thicker, less branched, and the elliptic-

ovate fruit.

4. Memecylon caeruleum Jack, Mal. Misc. 1:5 (1820) 26: M. floribundum

Bl, Mus. Bot. Lugd.-Bat. 1:23 (1851) 36!; M. caeruleum Jack var.

floribundum (Bl.) Kurz, For. Fl. Brit. Burma I (1877) 511: M. cordatum

Wall. Cat. 4100E.

Shrub up to 4 m or less commonly a tree up to 12 m tall; bark thin, finely

fissued, grey-brown to blackish; branchlets cylindric, often flattened and shallowly

grooved on 2 faces below the upper node, smooth, 1.5-2.5 mm thick, drying

brown to blackish; blades coriaceous, ovate to elliptic, acute to obtuse at the

tip, often with a very minute cusp; broadly rounded, cordate, or acute at the

base; 8-16 cm long, 3.5-7 cm wide: venation very obscure to invisible on both

surfaces, midnerve sunken above, prominently raised and tapering below; dark

green above, green below, drying dark brown or blackish above, lighter brown,

less commonly greenish below; margin often inrolled below; glabrous; petiole

1-3 mm long, c. 2 mm thick in broadly rounded to cordate blades; 5-7 mm

long in acute blades and usually not as thick; glabrous; inflorescence cymose,

appearing glomerulate, 1-1.5 cm long, glabrous, many flowered, primary axes

2-3 from foliate nodes, 2-4 mm long, slightly flattened, c. 2 mm thick; secondary

axes 2-3 mm long, generally with one pedicel at the tip of each; pedicels

1-2 mm long; bracts ovate, acute, keeled on the back, up to 0.5 mm long,

persistent; calyx campanulate, often somewhat funnelform, smooth, glabrous;

margin truncate, thick, often cracking in 4 places (opposite the petals) after

flowering; 2-4 mm long, 2.5-3.5 mm wide; pink to purple; petals coriaceous,

broadly ovate to orbicular, acute or tipped with a minute mucro, truncate to

emarginate at the base, c. 2.5-3 mm long, 3-3.5 mm wide at the middle; mid-

nerve thickened and often keeled, margins thinner, reflexed at maturity, dark

purple, or deep blue, margins often whitish; filaments 2.5 mm long, deep blue;

anther curved in a “‘)’’ or “C” shape, c. 1 mm long, deep blue, locules white,

gland distinct, reddish or orange-brown; style 4-6 mm long, blue; ovules c. 10;

infructescence axes thicker, secondary axes often elongating 1-2 mm, total length

up to 2.5 cm; fruit ellipsoid, smooth, glabrous, 10-15 mm long, 6-9 mm wide,

crowned by the persistent calyx remnant, areolus 3-4 mm wide; exocarp thin,

pink to red, turning light blue, deep purple to black with deep purple juice

when ripe; pericarp pulpy and gritty, c. 2 mm thick, white.

Figure 4: a. inflorescence, b. petal, c. stamen, d. fruit and seed

62 Gardens’ Bulletin, Singapore XX XIII (1980)

There is considerable variation in the leaves of this species. Specimens

with broadly rounded to cordate blades with petioles 1-3 mm long and 2 mm

thick confer with Jack’s description of M. caeruleum. In M. floribundum BI.

the blades are generally narrowed at the base, however broadly rounded, but

not cordate, blades are often found associated with these narrowed blades (Noor

& Samsuri 10, is a good example). The petioles in M. floribundum, regardless

of the base of the blades, are always 5-7 mm long. The blades and leaves

of M. caeruleum Jack, specifically those described by Blume as M. floribundum,

are similar to those of M. campanulatum Cl. (q.v.) and M. edule Roxb. var.

ovatum (Sm.) Cl. (q.v.), however the former has a glomerulate inflorescence

(axes not distinct) and the latter has an elongate (1-4 cm) one; both differ in

having globose fruit. M. amplexcaule is easily distinguished from M. caeruleum

s.s., by its clasping leaf bases, smaller flowers, and globose fruit.

M. caeruleum s.s. is apparently restricted in distribution to southern Thailand,

the Malay Peninsula, and paris of Indo-China. M. floribundum s.s. ranges from

central Thailand to the Malay Archipelago. While the blades differ significantly

and there seem to be no intermediate shapes linking the two types; the inflore-

scences, flowers, and fruits are identical. There is, therefore, no reason in keeping

the species separate, therefore M. florubundum BI. has been reduced in this paper

to a synonym of M. caeruleum Jack on the basis of publication priority.

Kadim and Noor 660, from Pulau Tulai, Pahang; is an exceptional specimen

in that the infructescence is up to 2.2 cm long. The primary axes are solitary

and are up to 17 mm long with 1 or 2 nodes, the secondary axes are about

2 mm long and the pedicels are slightly shorter.

Thailand

Singora: Annandale 1662; Teratau Is.: Haniff & Nur 7479; Mohea, Trang:

Hamid 2959

Malay Peninsula

Perlis — Besih Hangat: Henderson 22881; Kaki Bukit: Kiah 35280

Kedah — Alor Star: Ridley 15057; Kedah Peak: Rao, Keng, Wee 77; Pulau

Langkawi: Bolinan 6787, Chew 138, Kerr 21762, Robinson 6328, Turnau 741,

Whitmore 12984; Pulau Rawei: Ridley 15777; Pulau Selang: Corner sn, 22 Nov.

1941; Pulau Timun: Henderson 29125; Tau: Ridley sn, June 1893; Yan: Ridley

sn, June 1893.

Wellesley — Bukit Jura: Ridley sn. Dec. 1895

Penang — Ahmad 1015: Curtis 54, Oct. 1884; Curtis 54, Aug. 1889; Curtis

54 (= 1507), 1507; King’s coll. 1687; Selvaraj 99660; Sinclair 39045; Wallich

4100E (type M. cordatum Wall.)

Perak — Pulau Jarajah: King’s Coll. 4972; Pulau Lanang: Seimund sn; Pulau

Pangkor: H.M. Burkill & Shah 253; Corner 31654; Foxworthy 1736; Ridley

sn; Yeob 1097; sine loc.: Scortechini 641, 1027

Trengganu — Pulau Kapas: Holttum 15216

Selangor — Kuala Selangor: Watson 16410

Revision of Memecylon L. 63

Pahang — Pulau Sepoi: Corner 25764, 29846; Pulau Tioman: Corner sn, 17

Aug. 1935: I.H. Burkill sn, June 1915; Henderson 18538; Pulau Tulai: Kadim

& Noor 660; S. Kampong Aur: Ahmad & Shukor 484

Negri Sembilan — Port Dickson: Strugnell 10988

Malacca — Bukit China: Goodenough 1522; Bukit Panan: Alvins 280; Gambaga

Batu: Hervey 2020; G. Kalan: Alvins sn, 19 Dec. 1885; Lanjong Kling: Ridley

218; Pang Kalan Miniak: Alvins sn, 19 Dec. 1885; sine loc.: Cuming 2322,

Griffith 2322, Maingay 809 (1423)

Johore — Hulu Sedil: Lake & Kelsall 4074, Pulau Pemanggil: Noor & Samsuri

Singapore — Chew 1463; Furtado 34854; Kassim 1489; Pangi sn; Walker sn.

Further Distribution — Indo-China, Burma, Thailand, Sumatra, Java (and

islands to north),, Anambas Islands, New Guinea (fide Craib and Bakh. f.).

5. Memecylon campanulatum Cl. in Hk. f., Fl. Brit. Ind. II (1879) 563; M.

multiflorum Bakh. f., Med. Mus. Bot. Utrecht 91 (1943) 343 and Rec. Trav.

Bot. Neerl. 40 (1943-45) 343; Furtado, Gard. Bull. Sing. 20 (1963) 121

syn. nov. |

Tree up to 20 m tall with a diameter of up to 60 cm, shortly buttressed;

bark closely fissured, soft, c. 2.5 mm thick, brown to grey; inner bark pink;

slash wood yellow; branches 2-grooved to slightly 4-angled below the upper

node, becoming cylindric, 2—2.5 mm thick, drying grey to light brown; blades

coriaceous, glabrous, elliptic to ovate, less frequently lanceolate, obtuse at the

tip, acute and decurrent at the base; venation invisible on both surfaces, intra-

marginal nerve very obscure to invisible, c. 2 mm from the margin, mid-vein

sunken above, raised and tapering below; drying tan to dark olive-brown above,

lighter brown below; 7—12.5 (15.5) cm long, 4-7 cm wide; petiole 5-8 mm long,

2 mm thick, glabrous; inflorescence glomerulate, up to 10 mm long, 5-10 mm

wide, glabrous, many flowered, on tubercles from older branches behind the

leaves; bracts and bracteoles lanceolate-ovate, acute, up to 0.5 mm long; primary

axes 0.5-2 mm long, secondary axes glomerulate or up to 1 mm long; pedicels

10 or more, c. 3 mm long, calyx narrowly campanulate, thickened, 1.5-2 mm

long, 1.5 mm wide, constricted above the ovary, margin truncate, flattening after

flowering, glabrous, smooth, yellow, drying dark brown to black; petals ovate,

acute at the tip, narrowed to the truncate base, c. 3 mm long, 1.5 mm wide,

thickened with thinner margins, midnerve dorsally. raised; filaments 4-4.5 mm

long, anthers ‘‘C’’-shaped, c. 1.5 mm long, gland prominent; style 2.5-3 mm

long; ovules 10-12; fruit subglobose, 7-8 mm diameter, on pedicles c. 2.5 mm

long, calyx remnant small, areolus c. 2 mm wide; exocarp green turning pink,

roughened and blackish when dry; pericarp c. 1 mm thick and gritty.

Figure 5: a. inflorescence axes, calyx and style; b. petal, c. stamen

In general appearance M. campanulatum very closely resembles M. edule

Roxb. var. ovatum (Sm.) Cl. (qg.v.) which has longer inflorescences (primary axes

at least 5 mm long), shallowly lobed to apiculate calyx, and petals that are more

acute at the tip. The anthers and fruit appear to be the same. There seems

to be a clear distinction between the lengths of the primary axes in these two

species, those of M. campanulatum are not more than 2.5 mm long and in var.

64 Gardens’ Bulletin, Singapore XX XIII (1980)

ovatum they are at least 5 mm (generally 1-3 cm) long. It is tempting to consider

M. campanulatum as a synonym of var. ovatum, but there are differences which

do not, as far as can be determined, merge to form a clear gradation of variation

between the two species. M. campanulatum, therefore, is maintained as a distinct

species in this paper with a strong suggestion that it may be the same as var.

ovatum. Kloss s.n., from Rawong, Selangor; is identical to the holotypes of

M. campanulatum and M. multiflorum except that the calyx has a 4-apiculate

margin. The petals and anthers of this specimen appear to be the same as the

two type collections. The type collection of M. multiflorum Bakh. f. (Achmad

1191, from Sumatra) is identical to the holotype of M. campanulatum at Kew.

Both type collections are in flower and fruits from other specimens collected

in Sumatra and the Malay Peninsula match. M. campanulatum is apparently

uncommon in both areas since there are only a few collections of this species

available.

The congested, glomerulate inflorescence and the thick leaves of M. cam-

panulatum are similar to those of M. amplexicaule Roxb. {(q.v.) which differs

in having a lobed calyx, nearly sessile and clasping leaves, and larger fruit.

Trengganu — Bukit Bauk: Hou 765; Jerangau For. Res.: Kochummen 2114

Selangor — Rawong For. Res.: Kloss sn

Pahang — Ulu Singei Anak, Endau: Cockburn 8105

Malacca — Ayer Panas: Derry 1184, Ridley 1184; sine loc.: Griffith 2325 (type)

Johore — Kluang For. Res.: Ng 98023; Ma’okil For. Res.: Sinclair 38992

Singapore — Ngadiman 36139

Further Distribution — Sumatra, Simeuloee Is. (fide Bakh. f.).

6. Memecylon cantleyi Ridl., J. Str. Br. Roy, As. Soc. 79 (1918) 72; M. dissitum

Craib, Kew Bull. 1930, 325 syn. nov.; M. steenisii Bakh. f., Med. Mus. Bot.

Utrecht 91 (1943) 355 and Rec, Trav. Bot. Neerl. 40 (1943-45) 355 syn. nov.

Shrub up to 3 m or a tree up to 15 m with a diameter up to 25 cm; bark

thin, smooth, finely fissured and flaking, brown to greyish, inner bark yellow

to brown; branchlets cylindric, often with two shallow grooves, sometimes with

four faint lines or four distinct angles, very smooth, c. 1.5 mm thick, light tan

to greyish when dry; blades coriaceous, less often subcoriaceous, lanceolate to

ovate, acuminate at the tp (acumen up to 1 cm long), acute at the base and

decurrent on the petiole for 2-3 mm, venation pinnate, 8-10 pair of nerves, very

obscure to invisible above, very obscure below, more distinct in thinner blades,

intramarginal nerve c. 2 mm from the margin, obscure to invisible, mid-vein

sunken above, raised and tapering below, texture smooth in thin blades, rough

in thicker ones when dry, 8-19.5 cm long, 3—7.5 cm wide; dark green above,

slightly yellowish-green below; drying brown-greenish above, greenish below;

petiole (5) 7-9 mm long, 1..25—1.5 mm wide; inflorescence 1.5—2.5 cm long, cymose

from leafy or leafless nodes, each with 3-10 flowers; primary axes | or 2 from

each axil, S-11 mm long, flattened, secondary axes not developed or up to 6 mm

long, pedicels clustered at the tips of the main axes, 3-5 mm long; calyx cam-

panulate, c. 2.5-3 mm high 3-4 mm wide, flattening after flowering, smooth and

glabrous outside, faintly ridged inside, margin truncate to somewhat undulate

Revision of Memecylon L. 65

with 4 minute (often indistinguishable) cusps, purplish-pink; corolla buds apiculate,

tips twisted; mature petals broadly ovate, 3-4 mm long, 3-4 mm wide, acute

at the tip and often provided with a mucro, shortly clawed at the base, thickened

with thinner margins, mid-nerve visible, lateral venation frequently visible, anthers

bent in an overturned “‘U”’ shape, gland distinct in immature anthers, rather

obscure and elongating in mature ones, anthers and filaments deep red-violet:

ovary distinctly narrower than the base of the calyx, ovules 6-8, infructescence

axes thickening and elongating 1-2 mm; mature fruit subglobose, c. 10 mm long,

8 mm wide, calyx remnant very prominent in immature fruits, less so when

mature; areolus c. 4 mm wide, internal ridges very faint; pinkish-white or sulphur

yellow, turning dark purple when mature, drying blackish when immature, light

brown to tan when mature, smooth with a thin (c. 0.25 mm) exocarp.

Figure 6: a. inflorescence, b. flower, c. petal, d. stamen

The relatively large flowers that are on short inflorescences plus the large,

subglobose fruit; and the flattened calyx in old flowers tend to characterise this

species.

Memecylon dissitum Craib is, in this paper, considered as being the same

as M. cantleyi Ridl. The types cf M. dissitum (Kerr 15408, from peninsular

Thailand) are in poor condition, nevertheless the holotype from Kew does have

a few small buds and the isotypes from BM and BK both have fruit. The

immature flowers, especially the anthers. match the buds of Sidek 380 — which

is M. cantleyi. The fruit on both isotypes are the same as on Chelliah 104387

and Sinclair & Kiah 39964 — both clearly being M. cantleyi. The leaves on

the type specimens of M. dissitum match those on the specimens of M. cantleyi

(including the holotype) examined. Craib (l.c. 325) notes that M. dissitum is

close to M. garcinioides Bl. (q.v.), however the structure of the inflorescence.

size of the flowers, fruit, and especially the anthers differ. The anthers of M.

garcinioides are quite unusual and those of M. dissitum are not at all similar.

Memecylon cantleyi Ridl. is most closely related to M. oleifolium B. (q.v.)

since the branches, especially the grey-tan colour when dry; leaves, and inflore-

scence structure are very similar. The flowers are also similar, however those

of M. oleifolium are much smaller and the fruit of M. cantleyi is smooth and

not roughened as with M. oleifolium.

Memecylon steenisii Bakh. f., from Sumatra, has also been reduced to a

new synonym of M. cantleyi Ridl. and is more remotely related to M. oleifolium

Bl. than Bakh. f. suggests. The holotype of M. steenisii (van Steenis 3373) and

the paratype (Achmad 1387) at Leiden both have inflorescences c. 1 cm long

each with 3-7 flowers. The size of the calyx (all buds) and the shape of the

petals, especially the texture, strongly resemble M. cantleyi. The bud anthers

resemble both M. cantleyi and M. oleifolium. These two specimens of M. steenisii

also have blades which have dried greenish to yellow-greenish as in M. cantleyi

— in contrast to M. oleifolium which dries brown.

The holotype of M. steenisii matches Yusoff 99141 and Suppiah 104811

(from Pahang and Kelantan, respectively) and is essentially identical to Chelliah

104387 (from Pahang) which has definite M@. cantleyi fruit. The flowering and

fruiting axes of all of these specimens are up to | cm long and are structurally

the same. M. oleifolium has longer axes which are typically over | cm long.

The holotype of M. dissitum Craib compares very closely in all vegetative and

flowering features to the two types of M. steenisii. All these specimens are in

bud and the size and shape of the inflorescences and buds are essentially identical.

66 Gardens’ Bulletin, Singapore XX XIII (1980)

Since the fruit of M. steenisii Bakh. f. has not been seen it is impossible

to be absolutely certain that it is a synonym of M. cantleyi. However, from the

evidence described above it is reasonably certain that this reduction is justified.

Thailand

Kanchanaburi, Kwae Noi Basin: Kostermans 940; Nakorn Sri Tammarat, Kao

Luang: Kerr 15408 (type M. dissitum Craib)

Malay Peninsula

Kedah — Bukit Enggang For. Res.: Sidek 380; sine loc.: Saaid 20661

Penang — Nauen sn, May 1940

Perak — Larut: King’s coll. 7635

Kelantan — Ulu Sat For. Res.: Suppiah 104811

Trengganu — Ulu Brang: Moysey & Kiah 33641; Sinclair & Kiah 39964

Selangor — Bukit Bauk For. Res.: Kochummen 2384

Pahang — Belingoo, Temerloh: Awang-Lela 4578; Bentong: Ahmad 5099;

Gunong Benom, Ulu Krau: Chelliah 104387, Yusoff 9914; Jambu Bongkok:

T. & P. 8 (2608); Jenderak Halt: Kadim & Mahmood 50; Jerantut: Holttum

24751; Raub: I.H. Burkill & Haniff 16219; Sungei Krau, Temerloh: Strugnell

22472, 22475; Tembeling: Henderson 24515; Titi Bungkor, Temerloh: Henderson

10671; Ulu Checka, Benom Forest: J.C. (Carrick) 1639 (2569); Ulu Chineas,

Kuala Lipis: I.H. Burkill & Haniff 17091

Negri Sembilan — Seremban: Alvins sn, 21 April 1886

Malacca — Ayer Panas: Derry 1240; sine loc.: Maingay 817 (1688)

Johore — Sungai Kayu: Kiah 32170; Sungai Sedili: Corner 36980

Singapore — Ahmad 1221, Cantley 148, Ridley 13012 (type), Shah 1733.

Further Distribution — Sumatra, Simeuloee Is. (fide Bakh. f.)

7. Memecylon cinereum King, J. As. Soc. Bengal 69, II: 1 (1900) 82 (Mat. FI.

Mal. Pen. III, 490); M. umbellatum Wall. Cat. 4109, in part.

Shrub 1-2 m or a tree up to 4 m tall; upper branches grooved and ridged

on 2 sides, or somewhat 4-angled with wider grooves, c. 1 mm thick; older

branches cylindric, smooth, thicker, younger branches drying dark brown to

blackish, older ones tan; blades subcoriaceous, ovate to lanceolate, long acumi-

nate to almost caudate at the tip (acumen up to | cm long), acute to cuneate

and often decurrent on the petiole; midnerve sunken above, raised and tapering

below, other venation generally invisible on both surfaces; 5.5-12 cm _ long,

2.5-5 cm wide, drying dark brown to blackish above, dark brown below, very

smooth on both sides, entirely glabrous; petiole c. 5 mm long, | mm thick;

inflorescence cymose, appearing glomerulate, 5-7 mm long, from leafy nodes,

glabrous; primary axes flattened, up to 1.5 mm long with 1 node, secondary axes

fused and indistinct or c..1 mm long, pedicels c. 1 mm long, flowers numerous

in each inflorescence; calyx campanulate and widened above the ovary, 1.5-3 mm

Revision of Memecylon L. 67

long, 2.5-3.5 mm wide, smooth or minutely papillose outside, ridged internally;

lobes distinct, broadly triangular, 0.25-1 mm long; pinkish or blue; petals broadly

ovate, acuminate to aristate at the tip, narrowed and truncate at the base, mid-

vein thickened but not keeled, 1.5-3.5 mm long, 1-1.5 mm wide, whitish or

purple; filaments c. 3 mm long; anthers triangular-cuneate, without a gland or

in bud anthers sometimes visible as a very minute bump, c. 1.5 mm long; ovules

6-10; fruit globose, 7-8 mm diameter, calyx remnant slightly raised, with a square

limb which is often flattened in the plane of the fruit, easily falling off; areolus

1.5-2 mm wide, internal ridges distinct; exocarp generally smooth or with a few

minute warts, c. 0.2 mm thick, black when dry; testa glossy, brown-tan when dry.

Figure 7: a. calyx and style, b. calyx, c. bud petal, d. mature petal,

e. stamen

M. cinereum King is very closely related to M. lilacinum with the inflore-

scence, calyx, and especially the glandless anthers. The blades and fruit differ.

M. malaccense (Cl.) Ridl. is probably even more closely related, and the only

major difference is with its rounded and shallowly cordate (c. 0.5 mm) blades

and shorter calyx lobes. Those of M. cinereum are acute and decurrent at the

base. The inflorescence, in basic structure, is similar, although less glomerulate

with M. malaccense. The petals, anthers, and fruit are the same.

Memecylon floridum Ridl. (q.v.) has slightly larger axes, but otherwise the

flowers are the same. The blades and especially the larger fruit with a fibrous

pericarp are major differences.

Two specimens of Wallich Catalogue 4109 at Kew, inserted in a type folder

of M. edule Roxb. var. ovatum (Sm.) Cl., are clearly M. cinereum. According

to Cogniaux (l.c. 1155, 1156) other parts of the collection belong to M. edule

var. ramiflora Triana, and M. edule var. ovatum.

Perak — Salama: King’s coll. 3143 (lectotype); Silma: Scortechini 2035 (syntype):

Ulu Kal: King’s coll. 10758 (syntype):; sine loc.: Scortechini 394 (syntype)

Trengganu — Gunong Padang: Moysey & Kiah 3390

Selangor — Ginting Simpah: Hume 9011; Pahang Track: Ridley 8615: Semangkok

Pass: I.H. Burkill 8888, Ridley 15570; Sungei Bulok: Goodenough 10602; Trig.

Station: Suillicee 8888; Weld’s Hill: Cubitt 5109, Hamid 9908

Pahang — Bentong: Best 13852; Temerloh: Henderson 10514, 10704

Singapore — Wallich Cat. 4109 (type M. umbellatum Wall., in part)

8. Memecylon corticosum Ridl., J. Fed. Mal. States Mus. 10 (1920) 92.

Shrub 2-3 m tall or treelet up to 3 m tall; branches distinctly 4-winged

throughout the upper internodes, nodes with wider wings (appearing auricled):

branches becoming cylindric with age, 2-3 mm thick, drying grey to tan; blades

sub-coriaceous, lanceolate to elliptic, acuminate at the tip (acumen 1-2 cm long);

rounded, and in larger blades slightly (c. 1 mm) cordate, less frequently narrowed,

at the base, venation pinnate, c. 15-20 pairs of nerves, sunken and obscure to

distinct above, raised, often prominently, below, intramarginal nerve similar, 4-6

mm from the margin, broadly looping, midnerve sunken above, prominently

raised and tapering below; 10-23 cm long, 3-9 cm wide; medium glossy green

68 Gardens’ Bulletin, Singapore XX XIII (1980)

above, dull green below, drying light greenish-brown above, light brown or

greenish below; petiole 2-3 mm long, 2-3 mm thick; inflorescence cymose, often

appearing glomerulate, from behind the leaves, often ramiflorus on thickened

(c. 1 cm) branches with tubercles 2-4 mm thick; c. 5 mm long; bracts and

bracteoles ovate, acute, thickened, c. 0-5 mm long, persisting; primary axes several

per axil, with 1 or 2 nodes, flattened, 1-4 mm long; secondary axes almost

indistinguishable and appearing fused or up to 0.5 mm long; pedicels 5-10,

2.5—3 mm long; calyx campanulate, often appearing funnelform with a constriction

above the ovary, margin truncate with 4 broad and short lobes; thickened, smooth,

glabrous, ridged internally, sometimes flattening after flowering, 1-1.5 mm _ long,

2-2.5 mm wide; pink, drying black; bud petals thick, broadly ovate, obtuse at

the tip, truncate at the base, margins thinner, 3-4 mm long, 4 mm wide, dark

crimson, white; anthers slightly curved to almost ‘“C’-shaped, purple with a

prominent gland; ovules 4-6; fruiting pedicels c. 5 mm lIcng; fruit pyriform to

obovate, capped by the slightly raised calyx remnant, areolus c. 3 mm wide;

9-11 mm long, 8-9 mm wide (at the widest point), exocarp pale greenish-white,

flushed purplish-violet when nearly ripe, drying smooth or with a somewhat

roughened texture; pericarp c. 0.5 mm thick.

Figure 8: a. upper node and internode, b. leaf, c. flower bud, d. calyx,

e. petal, f. stamen

The holotype of M. corticosum Ridl. (Kloss 7027, from peninsular Thailand)

has bud anthers which differ completely from M. wallichii and M. longifclium,

thus it is certain, even though the blades and branches of M. corticosum often

appear similar, that M. corticosum is distinct from these other two. The blades

on the holotype of M. corticosum have distinct venation, however in most other

specimens the venation is more prominently raised below. One specimen (Shah

& Noor 1838) has a few very immature buds on a small tubercle in a leaf axil,

however all the other specimens studied have the inflorescence behind the leaves

or on thickend branches.

The winged branches and general shape of the leaves suggest close affinities

with M. fruticosum King (q.v.) (including synonyms M. epiphyticum King and

M. tenuifolium Ridl.). All of these taxa are poorly known, thus their exact

relationships are difficult to determine. M. kunstleri King (q.v.), which is also

incompletely known, has elliptic fruit and similar bud stamens, but cylindric

branches.

Vegetative specimens are easily confused with M. wallichiit Ridl. (q.v.) and

in several collections, e.g. Henderson 19581, Moysey & Kiah 33738, 33856; the

distinction is not clear due to lack of flowers or fruit. The inflorescences in these

three collections are ramiflorus, thus they have been inserted under M. corticosum.

Henderson s.n., from Batu Papan, Kelantan is vegetative and could also be refer-

rable to M. wallichii.

Thailand

Bangtapan, Ratchaburi: Keith 492; Tasau, Chumpawn: Kloss 7027 (type)

Malay Peninsula

Kelantan — Bukit Batu Papan: Henderson sn, 8 July 1935; Gua Minik: Hender-

son 19581; Gua Panjang: UNESCO 609

Revision of Memecylon L. 69

Trengganu — Kuala Trengganu-Besut Road: Sinclair & Kiah 4058; Ulu Brang:

Moysey & Kiah 33738, 33856; Ulu Telemong For. Res.: Suppiah 11408

Pahang — Ulu Sungei Sat: Shah & Noor 1838

Johore — Sungei Juasseh, Labis: Ahmad 283

9. Memecylon dichotomum (Ci.) King in J. As. Soc. Beng. 69, II:I (1900) 75

(Mat. Fl. Mal. Pen. HI, 483). M. elegans Kurz var. dichotoma Cl. in Hk.

f., Fl. Brit. Ind. Hf (1879) 554; Cogniaux in DC., Monogr. Phan. 7 (1891)

1138; M. ridleyi Cogn. ex Ridl., J. Str. Br. Roy. As. Soc. 30 (1897) 85

nomen nudum; M. eugeniiflora Ridl. (‘‘eugeniflora’’), J. Str. Br. Roy. As.

Soc. 57 (1910) 48 syn. nov.; M. dichotomum (Cl.) King var. eugeniiflorum

(Ridl.) Ridl., Fl. Mal. Pen. I (1922) 812 syn. nov.

var. dichotomum

Tree 6-12 m tall with a diameter up to 30 cm, less commonly a treelet or

a shrub up to 3 m tall; bole irregular with many knots; bark thin, slightly fissured

and scaly, light brown; upper nodes slightly flattened, often somewhat 4-winged

or 4-auricled; internodes usually sharply 4-angled and usually tapering to cylindric,

older branches cylindric; 1-2 mm thick; blades chartaceous to subcoriaceous,

lanceolate or ovate, acuminate at the tip, rounded or shallowly cordate at the

base; venation pinnate, c. 8-15 (20) pairs of nerves, obscure above, distinct and

often slightly raised below; intramarginal nerve obscure to distinct, 3-4 mm from

the margin, broadly looping: mid-vein sunken above, raised and tapering below;

5-10 cm long, 2-5 cm wide; glossy greeen above, duller green below; drying

dark brown to greenish above, greenish to olive-green or brown below; entirely

glabrous; petiole 1-2 mm long, 1-1.5 mm thick (cordate based blades appearing

sessile), glabrous; inflorescence cymose, 5-7 mm long, glabrous, few (often 3)

flowered; primary axes 1 — several from leaf axils, 0.5—3 (exceptionally 5) mm

long, with 1 or 2 nodes, flattened and 4-angled; secondary axes 3-5, up to 3 mm

long, less commonly not developed; pedicels papillose, solitary on the tip of each

secondary axis, 0.5-1 mm long; bracts thick, lanceolate to ovate, acute, c. 1 mm

long, persistent; bracteoles similar, both usually minutely papillose; calyx cam-

panulate, often constricted above the ovary, becoming funnelform after flowering,

slightly thickened, truncate and often splitting irregularly at the margin, usually

minutely papillose outside, 3 mm long, 4 mm wide, pink to red; petals thick

coriaceous with thinner margins, broadly ovate to suborbicular, broadly rounded

to almost truncate at the tip, often with a minute mucro; narrowed and some-

what clawed or entirely truncate at the base; midnerve not apparent, 3-3.5 mm

long, 3.5-5 mm wide, whitish to pink, reflexed at maturity; filaments white,

3-4 mm long; anthers lilac, crescent shapped to ‘“‘C’’-shaped, I-1.5 mm _ long;

gland centrally located, prominent, red-brown; stigma minute, style 3-4 mm long;

ovules c. 8; fruit globose, 12-16 mm diameter, calyx remnant distinctly raised

(c. 1 mm) and thickened, areolus 4-5 mm wide; exocarp green when immature,

turning yellow flushed with rose-red, becoming blackish when ripe, edible, sweet,

smooth and thin in smaller and immature fruits, becoming minutely muricate-

papillose and black when dry; pericarp c. 1 mm thick, gritty.

Figure 9: a. branch, leaf, and internode:

b. bracts, calyx, and style; c. petal, d. stamen

A variable species with several forms which are not well understood. The

papillose pedicels and calyx recall M. minutiflorum Migq. (q. v.), but that species

has a very sharp constriction above the ovary, different branches, leaves, and

fruit. |

70 Gardens’ Bulletin, Singapore X XXIII (1980)

The two syntypes of M. eugeniiflora Ridl. have narrower and thicker blades

than the type collection of M. dichotomum, however both taxa have identical

branches, inflorescences, and flowers. Several collections clearly show that both

narrow and wide blades occur on the same plant, e.g. Ridley 13423 and others

with both distinct (dichotomum) and indistinct to invisible (eugeniiflora) venation,

e.g. Selvaraj 11189. There is no reason, therefore, to maintain Ridley’s taxon

as distinct from M. dichotomum. The papillose calyx and distinct bracts of

Ridley 14695 (syntype of M. eugeniiflora) match those of the type collection of

M. dichotomum.

Another species, M. ridleyi Cogn. ex Ridl., only differs from M. dichotomum

in the former having more rounded leaf bases and perhaps fainter venation on

the leaf undersurfaces. Two specimens (Ridley 2241, 2609) from Tahan River

and Kuala Tahan (respectively), Pahang) were collected within days of each other

and have “M. ridleyi Cogn.” written in Ridley’s writing on their original labels.

These are the only two specimens of M. ridleyi seen in the Singapore collection.

King (lc. 75) lists Ridley 2609 under M. dichotomum, and Furtado annotated

this specimen as M. dichotomum also. Ridley 2241 was annotated as M.

perakense by Furtado. There is no doubt that both of these specimens are M.

dichotomum. M. ridleyi is listed by Ridley in his compilation of Malay plant

names as “‘delima burong’’. Apparently Ridley never published a description of

this species, therefore it is a nomen nudum. Thus, not only are the differences

of this species artificial, but the name itself cannot be used due to nomenclatural

rules.

A form of M. dichotomum with cylindric or obscurely 4-angled branches,

coriaceous blades with shallowly (1-2 mm) cordate bases with invisible venation,

and a smooth calyx with a broadly 4-lobed margin each tipped with a minute

cusp is known from two collections on limestone from Kelantan. More material

of this form is desired for a better understanding of its exact taxonomic status

(UNESCO 520, Whitmore 4039).

uses: the liquid from boiled roots is drunk after childbirth, boiled roots used

for rheumatism, used for house posts which last for 5 or 6 years when exposed;

the fruit is stupefying, and the very heavy (sinks in water), durable wood is

used for making spring bows (belantek).

Kedah — Bukit Enggang For. Res.: Kochummen 2011; Gunong Baling: Corner

sn, 25 Nov. 1941; Gunong Jerai: Ridley 5752

Perak — Bujong Malacca: Ridley 9638; Gunong Tempurong:~ Allen & Kadim

494; Larut: King’s coll. 3239, 5036, 5297; Maxwell’s Hill: Ridley 2938, sn in

June 189?; Wray 2989; Temengo: Ridley 14695 and 14696 (types M. eugeniiflora

Ridl.); Ulu Kal: King’s coll. 10783

Kelantan — Gua Batu Goh: Whitmore 4039; Gua Musang: UNESCO 241;

Gua Panjang: UNESCO 520, Henderson 19595; Gunong Rabong: Shah 2486;

Kampong Gobeh For. Res.: Shah & Kadim 489; Sungai Ketchil, Batu Papan:

Nur & Foxworthy 12073; Sungai Lebir: Cockburn 7103; Temanga For. Res.:

Brown 52920

Selangor — Bukit Enggang: Symington 24199; Chadangan For. Res.: Chelliah

98204; Dusun Tua: Ridley 7331; Gading For. Res: Loh 13369; Klang Gates:

Ridley 13423; Kuala Lumpur: Curtis 2338; Sempang Mines: Ridley 15693;

Sungei Bulok: Jaamat 13908; Sungei Lalang Kajang: Symington 24075; Ulu

Klang Ampang: Gadoh 1661; Ulu Langat: Gadoh 959, Suppiah 11262

Revision of Memecylon L. 71

Trengganu — Bukit Kajang: Corner 30312, 30327; Gunong Tebu, Besut: Selvaraj

11189

Pahang — Bentong: Shah 194; Bukit Cheras: Henderson 25064; Cameron High-

lands: Nur 32862: Fraser’s Hill: I.H. Burkill & Holttum 8697, Cubitt 6522,

Henderson & Nur 11142, Kalong 22411, Kochummen 98156; Gali near Raub:

I.H. Burkill & Haniff 16249; Genting Highlands: Kochummen 16678; Whitmore

0925, 4629; Gunong Serudom, Kuantan: Shah, Sidek, Samsuri 3777: Jenderah

Halt: Kadim & Mahmood 27, Kadim & Nur 27; Kota Glanggi: Henderson

22403; Krau Game Res.: Whitmore 3231; Kuala Tahan: Ridley 2241 (syntype

M. ridleyi Cogn. ex Ridl.); Kwala Tembeling: Ridley sn (= 2241), Aug. 1891;

sn in 1891; Holttum 20535, Henderson 21772; Lesong For. Res.: Ahmad &

Shukor 421; Raub: Strugnell 22259; Sungai Nerling. Temerloh: Henderson 10565:

Tahan River: Ridley 2609 (syntype M. ridleyi Cogn. ex Ridl.); Sungai Tahan:

Kiah 31902, Mat sn in 1893; Taman Negara: van Balgooy 2479; Telom Valley:

Kiah & Strugnell 24023; Temerloh: Burn-Murdoch 164; Ulu Keniayam: Shah

1535; Ulu Sungei Sat: Shah & Noor 1777

Negri Sembilan — Gemas: I.H. Burkill 6387; Gunong Angsi: Nur 11572, Osman

23669; Bukit Tangga: Ridley sn on 17, 22, 27 Dec. 1920; Gunong Tampin:

Holttum 9574; Sungei Ujong: Alvins 1925, 2217, sn

Malacca — Mt. Ophir: Maingay 820 (2598); sine loc.: Alvins 119, Griffith 2324

(type), Maingay 818 (1424)

Johore — Gunong Belumut: Whitmore 8748: Khiang: Holttum 9279; Labis For.

Res.: Jumali 6711 (4257), Maxwell 77-372; Sungei Endau: Shah, Shukor, Ahmad

2624; Sungei Juasseh, Labis: Ahmad 273

Further Distribution — peninsular Thailand: Pattani (fide Craib).

9a. Memecylon dichotomum (Cl.) King var. rotundatum (Craib) Maxw. comb.

nov.; M. gracilipes Ridl. var. rotundatum Craib, Fl. Siam. Enum. 1:4 (1931)

708: M. gracilipes Ridl., J. Str. Br. Roy. As. Soc. 79 (1918) 72 not C. B.

Robinson, Phil. J. Sci. Bot. 6 (1911) 353: M. curtisii Burk. & Hend., Gard.

Bull. Str. Settl. III (1925) 377 syn. nov.; M. perakense Merr., Gard. Bull.

Str. Settl 8 (1935) 132 nom. illeg.

Tree up to 6 m tall or a shrub 1-3 m high; branchlets 4-angled and almost

winged below the upper nodes, often becoming cylindric in the lower parts of

the internode, cylindric in older branches, c. | mm thick; blades subcoriaceous,

ovate, acuminate at the tip (acumen 1-2 cm long); shallowly (1-2 mm) cordate

or less commonly broadly rounded at the base: venation pinnate, 6—12 pairs of

nerves, sunken above, faintly to distinctly raised below, intramarginal nerve 2-3

mm from the margin, looping, midnerve sunken above, raised and _ tapering

below; 5.5-10 cm long, 2.5-4.5 cm wide; drying dark brown to blackish above,

brown below; petiole c. 1 mm long, shorter than the cordate sinus at the base

of the blade: inflorescence a simple umbel composed of 3 flowers, solitary from

leaf axils, 20-33 mm long; primary axes 15-20 mm long, c. 0.3 mm thick,

glabrous; secondary axes subtended by a pair of lanceolate-ovate, acute bracts, c.

0.75 mm long, 1 or 3 per inflorescence, 14 mm long; each, when developed,

with 1 pedicel; pedicels 2-4 mm long, subtended by 2 lanceolate, acute bracteoles,

0.5-0.75 mm long; calyx campanulate, lobes broadly triangular, acute, sinuses

wide, lobes less distinct as the flowers mature and often appear almost truncate,

minutely papillose-muricate outside, ridged internally, c. 2 mm long, 2.5 mm

12 Gardens’ Bulletin, Singapore XX XIII (1980)

wide; petals ovate, acute, 1-1.5 mm long (ex description); anthers (only seen in

bud) crescent shaped, c. 2 mm long, gland obscure, not cupular, in many instances

it is merely a thickened area on the connective, probably even more obscure in

mature anthers; fruit globose, 15-20 mm diameter at maturity (ex description),

calyx remnant distinctly raised, areolus 1.5—2 mm wide; exocarp smooth, becoming

minutely muricate with maturity drying black; pericarp thin when immature,

thickening with maturity; testa glossy brown.

Figure 9a: a. inflorescence and calyx, b. bud stamens.

This variety is very closely related to M. fruticosum King (q.v.) and even

closer to the typical variety. A reduction of var. rotundatum to the typical

variety has not been made since there is variation in the inflorescences of the two

taxa, thus the two can always be easily distinguished on this basis. This is not

true with the leaves of both since they are variable.

There are two specimens in the Singapore collection (Haniff 14969, and

Henderson 23798) which fit Ridley’s description of M. gracilipes Ridl. viz. the

nearly sessile blades with round, often cordate bases; and the three flowered,

slender, long-peduncled umbel. Furtado annotated both of these as M. perakense

Merr. M. gracilipes Ridl. is a later homonym of M. gracilipes C. B. Rob. — an

entirely different species from the Philippines. M. gracilipes Ridl. was incorrectly

renamed as M. curtisii by Burkill and Henderson. At varietal status M. gracilipes

Ridl. var. rotundatum Craib is the oldest and only existing epithet for this taxon

and must be used as the correct name here. Merrill’s epithet cannot be used

since it was not only published after Craib’s var. rotundatum, but also has no

priority outside its species rank.

Curtis 1295, the type collection of M. gracilipes, matches Kerr 15766 which

is the type collection of var. rotundatum.

Thailand

Trang: Kerr 15766 (type M. gracilipes Ridl. var. rotundatum Craib), 19172

Malay Peninsula

Perak — Gunong Pondoh: Henderson 23798; Padang Rengas Reservoir: Haniff

14969; Waterloo Estate: Curtis 1295 (type M. gracilipes Ridl.)

10. Memecylon edule Roxb., Corom. Pl. 1:4 (1798) tab. 82, and FI. Ind. ed. 2,

(1832) 260; M. globiferum Wall. Cat. 4108.

var. edule

Tree up to 12 in tall with a diameter up to 15 cm, less frequently a shrub

up to 3 m tall; bark finely fissured, not flakikng, grey-brown; inner bark thin,

ochre; branchlets somewhat flattened and grooved on 2 sides, grooves often

widened, thus appearing 4-ridged or 4-angled, becoming cylindric with age, 1—1.5

mm thick; blades coriaceous. ovate, obtuse to acute at the tip, acute and decurrent

at the bise; venation pinnate, invisible or infrequently extremely obscure on

both surfaces; mid-nerve sunken above, raised and tapering below, 4.5-75 cm

long, 2.5-4 cm wide; dark green above, light green below; drying dark brown

above, brown below; petiole 4-6 mm long, 1—1.5 mm thick; inflorescence from

the axils of older leaves, sometimes from defoliate nodes, cymose, often con-

tracted and appearing glomerulate, glabrous, (0.5) 1.5-2 cm long, many flowered;

Revision of Memecylon L. 73

primary axis 1 or 2 from each axil, flattened and grooved or angled (1) 6-10 mm

long; often with 2 or 3 nodes; secondary axes usually 5, less frequently glome-

rulate, 4-5 mm long; tertiary axes 0-2 mm long; pedicels 2-3 mm long; calyx

campanulate, truncate with or without 4 minute cusps or 4 undulate lobes smooth

outside, ridged internally, 1-1.5 mm long, 2—2.5 mm wide, pinkish; corolla buds

conical, tips acute to shortly acuminate, twisted; petals coriaceous, broadly ovate

to oblong, acuminate at the tip, truncate at the base with a very short, broad

claw, dorsally keeled, margins with a wide thin zone, 1.5-2.5 mm long, c. 1.5 mm

wide, white, often with a pink hue; filaments c. 2 mm long, blue; anthers crescent

shaped, whitish to lilac, gland centrally located; stigma minute, style c. 3 mm

long, blue; ovules 8-14: fruit globose, often somewhat flattened, thus slightly

wider than long 5-9 mm wide, calyx remnant raised, areolus c. 1 mm wide,

exocarp green to yellow/green, turning pinkish, finally dark purple to blackish

when ripe, drying greenish-brown, often somewhat mottled, with a slightly rugose

texture; pericarp 0.5-1 mm thick, not gritty.

Figure 10: a. calyx and corolla bud, b. calyx and style, c. petal,

d. & e. stamens

habitat: open places, frequently found near the seashore

In many respects M. edule resembles M. lilacinum Z & M. (q.v.) but that

species has 4 distinctly apiculate calyx points, narrower and long acuminate petals,

no anther gland, and a grooved, flattened fruit with a very thick, gritty pericarp.

M. pauciflorum Bl}. (g.v.) is similar in flower structure to M. lilacinum, but has

a distinct anther gland similar to that of M. edule.

Thailand

La Tang Si: Annandale 1696

Malay Peninsula

Kedah — Pulau Langgun: van Balgooy 2349; Pulau Langkawai: Curtis 2627,

3691, sn in Feb. 1899; Haniff & Nur 7072: Keng et al. 126; Ridley 15804;

Robinson 6321; Telok Udang: Haniff 1027

Perak — Pangkor: Corner 31653

Kelantan — Gunong Brong: Shah & Ali 2891; Kota Baru: Ridley sn; Tumpat:

Corner sn on 22 April 1936

Trengganu — Kuala Dugun: T. & P. 349 (2949); Kuala Trengganu: Holttum

15158, Brelkerta sn on 1 Nov. 1939, Sinclair 40726; Mesang: Poore 6133

Selangor — Merchang: Poore & Merton 1133

Pahang — Badok For. Res.: Hou 763: Fraser’s Hill: Nur 11318, LH. Burkill

& Holttum 8937; Gunong Tahan: Wray & Robinson 5352; Pulau Duchong:

Corner sn on 22 Aug. 1935; Praman Pekan: Ridley = 1027, sn on 26 Aug.

1889; Rompin: Bidin 15442; sine loc.: Ridley 1027

Negri Sembilan — Cape Rachado: Mahmud bin Sider 13212, Kochummen 2484

Malacca — Gadek: I.H. Burkill 4477; Pulau Rumbia: Seimund sn; Sungai

Hudang: Derry 1028; Tanjong Bidara: Wong 2566; sine loc.: Alvins sn: Griffith

2326, 2327; Maingay 812

74 Gardens’ Bulletin, Singapore XX XIII (1980)

Johore — Mersing: H.M. Burkill & Shah 2508; Pulau Batu off Mersing: H.M.

Burkill & Shah 2526; Pulau Setindan: Corner 29760; Tg. Penawar: Cockburn

7646

Singapore — Henderson 35783, Hullett sn; Ridley 6054, 6531, 9564, sn in 1890,

sn in 1894, sn in 1906; Sinclair 39004, Maxwell 80-180

Further Distribution — India, Ceylon, Andaman Islands, Burma, Tonkin,

Banka Is., Borneo (fide Cogniaux).

10a. Memecylon edule Roxb. var. ovatum (Sm.) Cl. in Hk. f., Fl. Brit. Ind. II

(1879) 564. M. ovatum Smith, Rees Cyclop. 23 (1813). M. laxiflorum

Wall. ex Ridl., J. Str. Br. Roy. As. Soc. 79 (1918) 74 syn. nov. M.,

rhodophyllum Bakh. f., Med. Mus. Bot. Utrecht 91 (1943) 357 and Rec.

Trav. Bot. Neerl. 40 (1943-45) 357 syn. nov.

Tree up to 18 m tall, with a diameter up to 20 cm, or a shrub up to 5 m

iall, bark thin, finely fissured and flaking off in thin, broadly oblong pieces,

brownish; inner bark pale ochre, branchlets cylindric, often flattened and shallowly

grooved on 2 sides below the upper node, 1-2 mm thick, drying greyish to tan;

blades coriaceous, ovate, less commonly broadly elliptic or suborbicular, obtusely

acute to obtusely acuminate at the tip. broadly rounded to acute and slightly

decurrent at the base; venation pinnate, c. 15 pairs of nerves, very obscure to

invisible above, distinct, but rather faint below; intramarginal nerve obscure to

invisible below, c. | mm from the margin; midnerve sunken above, raised and

tapering below; dark, glossy green above, yellow/green below, drying dark brown

to blackish above, lighter brown below; 5-12 cm long, 3-6.5 cm wide; petiole

6-15 mm long, 1-2 mm thick; inflorescence from the axils of lower leaves and

more commonly from leafless nodes just behind the leaves; cymose, densely

flowered, often appearing glomerulate when the axes are short, 1-3 (exceptionally

4) cm long; primary axes 1-3 from each axil, vertically arranged, flattened and

4-angled or 2-grooved on 2 sides, 1-2 mm wide, glabrous, 5—15 (rarely 30) mm

long; secondary axes 5-8 (10) mm long, tertiary axes 0-2 (5) mm long, pedicels

3-4 mm long; calyx campanulate, widened above the ovary, margin with 4 shallow

undulations, or with 4 broadly rounded to acute lobes, sometimes truncate with

4 minute cusps; smooth outside, ridged internally, c. 1.5 mm long, 2 mm wide,

pink; corolla buds conical, apiculate, tips twisted; petals thin, venation visible,

broadly ovate to orbicular, broadly rounded and acute at the tip, truncate and

sometimes distinctly clawed at the base, midline and claw with a thicker texture;

2.5 mm long, 2.5 mm wide, pale to brilliant blue, often purplish; filaments 2.5

mm long, pale blue; anthers “J”? to ‘“‘U”’-shaped, c. 1 mm long; gland centrally

located, reddish black; stigma minute, style c. 3 mm long; ovules c. 8; fruit

globose, 8-10 mm diameter, calyx remnant raised, thin, areolus c. 2 mm wide;

exocarp green, turning yellow, red, pink, then purple-black when ripe, smooth

to slightly rugose and black when dry; often associated with keringga (red) ants,

pericarp at first thin, when ripe 0.5—1 mm thick, gritty.

Figure 10: a. calyx and style, b. petal, c. stamen. Figure 33.

habitat: open lowlands, commonly near the sea, recorded on limestone.

In general the blades of var. ovatum are larger and have longer petioles

than var. edule. The calyx in var. ovatum is usually broadly and shallowly

4-lobed while those of the typical variety are truncate and sometimes have 4

Revision of Memecylon L. 75

minute cusps. The bark of var. edule is narrowly fissured and does not

flake off while that of var. ovatum has wide fissures and flakes off. The blade

venation of var. edule is generally invisible, while those of var ovatum are faint,

but visible in most specimens, var. edule also has a shorter petiole and smaller

inflorescences which most frequently come from leaf axils. The petals of var.

edule are coriaceous, with a short claw, and white.

Memecylon campanulatum Cl. (q.v.) is very closely allied and differs in

having a nearly sessile inflorescence, truncate calyx, and blunt petals. As I have

not seen any intermediate forms linking the two taxa, I have maintained the two

as separate entities (q.v. discussion under M. campanulatum). The holotype of

M. laxiflorum Wall. ex Ridl. (Wallich 4472, from Singapore) is a fruiting specimen

which matches many similar specimens from throughout the peninsula. Con-

sidering the variation in the size of the leaves and inflorescence, Wallich 4472

is in no way variant and should be considered a synonym of M. edule Roxb.

var. ovatum (Sm.) Cl. This specimen was cited by Clarke and Cogniaux under

M. grande Retz., however King did not agree with this treatment and included

it under doubtful species. M. rhodophyllum Bakh. f., described from 5 sheets

collected by de Vriese (s.n.) from the Moluccas, only differs from M. laxiflorum

Wall. ex Ridl. in having slightly shorter (2-4 cm) infructescences on thinner

axes. All the isotypes are in fruit, however all of them are identical to var.

ovatum. The fruit of these specimens of M. rhodophyllum are smaller and have

a more prominent calyx remnant, but aside from this and the axes, significant

differences between this species and var. ovatum have not been found. There

are specimens at Singapore and Leiden which have smaller fruit and thinner axes

and these all fall well within the range of variation for var. ovatum, therefore

M. rhodophyllum has been reduced.

Thailand

Bang Son: Haniff & Nur 4245

Malay Peninsula

Perlis — Bukit -Lapi: Henderson 22803; Kaki Bukit: Kiah 35283

Kedah — Bukit Hantu: Chew 207: Gunong Kerriang, Alor Star: Kiah 35412;

Pulau Langkawi: van Balgooy 2319; Stone 6906, 6967; Chan 6800; Pulau Jerkam:

Corner sn on 7 Nov. 1941: Pulau Selang: Corner sn on 22 Nov. 1941; Pulau

Rawei: Ridley 15779; Pulau Tuba: Chen 41016

Penang — Pulau Jerjak: Ahmad 1004, 1012; Penang: Curtis 723 in Oct. 1885

(syntype M. laxifiorum Wall. ex Ridl.), Muka Head: Curtis 723 in March 1886

(syntype M. laxiflorum), Curtis 723 in May 1893 (syntype M. laxiflorum), Curtis

723 in 1893 (syntype M. laxiflorum); Gov.'t. Hill: Curtis 723 in May 1894; Batu

Ferengy: Curtis sn, May 1901; sine loc.: Forest Guard sn, Guard sn

Perak — Larut: King’s coll. 4175; Pangkor Island: Foxworthy 237, Corner

31656, Whitmore 3006

Trengganu — Pulau Kapas: Holttum 15215

Selangor — Ginting Simpah: Whitmore 4620; Gua Batu: Ridley 8279; Pulau

Angsa: H.M. Burkill & Shah 970; Pulau Selatan: H.M. Burkill & Shah 933

76 Gardens’ Bulletin, Singapore XX XIII (1980)

Pahang — Gunong Tahan: Ridley 16279; Kluang Terbang: Barnes sn; Kuantan:

I.H. Burkill & Haniff 17513; Pulau Chibeh: Corner sn on 19 Aug. 1935; Pulau

Sepoi: Corner 29846

Malacca — Malacca Pindah: Alvins sn

Johore — Kilat Estate: Ridley 15396 (syntype M. laxflorum Wall. ex Ridl.);

Minyak Buku: Ridley 11092 (syntype M. laxiflorum)

Singapore — Pulau Merambong: Corner sn on 29 Sept. 1935; Singapore: Ridley

9565, 9591; Wallich 4103B, 4!09B, 4472 (lectotype M. laxiflorum).

Further Distribution India, Ceylon, Andaman Islands, Burma, Tonkin;

Banka, Karimata, Billington Islands; Borneo (Indonesian part), Karimoen Djawa

Islands, Sumatra, Java (fide Cogniaux and Bakh. f.).

11. Memecylon excelsum Bl., Bijdr. Fl. Ned. Ind. 17 (1826) 1094; M. hetero-

pleurum Bl., Mus. Bot. Lugd.-Bat. 1:23 (1851) 362 syn. nov., M. heteropleurum

Bl. var. olivaceum King (‘‘olivacea’’), J. As. Soc. Bengal 69, II:1 (1900)

78 (Mat. Fl. Mal. Pen. III, 486), syn. in part by Furtado, Gard. Bull. Sing.

20 (1963) 120, non M. heteropleurum BI. sensu King l.c. 78; M. subtrinervium

Miq., Fl. Ned. Ind. Supp!. I, Sumatra (1860) 322 syn. nov.; M. maingayi

Cl. in Hk. f., Fl. Brit. Ind. II (1879) 567 syn. nov.; M. kurzii King lc. 77

syn. nov.; M. elmeri Merr., Univ. Calit. Publ. Bot. 15 (1929) 230 syn. nov.

Tree up to 18 m tall with a diameter up to 60 cm, bole straight, crown

spreading, branches long and deflexed, bark thin, finely fissured and flaking,

brownish-grey or greenish-brown; inner bark thin, pale brown, wood _ hard,

brownish; branches cylindric, often flattened and shallowly grooved on 2 sides

or sometimes slightly 4-lined or 4-angled below the upper node; 2—2.5 mm thick,

glabrous; blades coriaceous, elliptic to ovate, acute to obtuse at the tip, acute

at the base and not or only slightly decurrent on the petiole; venation pinnate,

15-22 pairs of veins, sunken above, slightly raised below, lower veins running

parallel to the midrib for 1-2 mm, then diverging directly towards the margin;

tertiary venation generally visible below; intramarginal nerve c. 2-4 mm from

the margin, nearly straight or looping, midnerve sunken above, raised and taper-

ing below; 12-26 cm long, 4-12 cm wide, glabrous; dark glossy green above,

duller green below, usually drying greenish-brown to dark brown above, olive-

green to brown below; petiole 3-4 mm long, 2-3 mm thick, glabrous; inflore-

scence of clustered cymes, appearing glomerulate, 6-10 mm long, sometimes

among the leaves, but usually just behind them, glabrous; primary axes several

from each leaf axil, usually on a tubercle, 1-4 mm long. flattened, c. 1 mm thick,

secondary axes not developed (glomerulate) or up to 3 mm long; tertiary axes,

when developed, up to 1 mm long; pedicels 1-3 mm long; bracts and bracteoles

ovate, acute, c. 0.5 mm long; calyx campanulate, thickened, truncate, smooth

outside, ridged internally, 3.5—-5 mm wide when mature, pink; petals coriaceous,

broadly elliptic to oblong, broadly rounded or truncate at the tip, truncate at

the base, 3-4 mm long, 3.5-5 mm wide, midnerve often raised on the dorsal

surface margins thinner, pale lilac to pink on the dorsal surface, white to cream

ventrally: filaments white, c. 2 mm long; bud anthers “‘J’’-shaped c. 2 mm long,

blue, with a narrow and elongated gland; style c. 4 mm long; ovules 6-8; fruit

oblong-elliptic, 8-20 mm long, 10-12 mm wide when ripe, calyx remnant raised,

aerolus c. 4 mm wide; exocarp green when immature, purple when ripe, drying

greenish to black with a rugose and often pustulate texture; pericarp 0.5-1 mm

thick; seed purple, containing a liquid.

Revision of Memecylon L. 77

Figure 11: a. inflorescence, b. calyx, c. petal, d. stamen, e. fruit,

f. areolus. Figure 34 a — e

The leaves are similar to M. oleifolium BI. (g.v.) (including M. ambiguum

BI and M. acuminatissum BIl.), but this species has a paniculate inflorescence at

least 4 cm long. M. paniculatum Jack (q.v.) has nearly sessile !eaves, a paniculate

inflorescence, smaller fruit, and winged branches.

The holotype of M. excelsum Bl. (Blume 208, from Java) at Leiden is in

poor condition and is without inflorescences and flowers. A specimen collected

by Bakh. Sr. (3092), also from Java, has identical branches and leaves as the

holotype. This specimen has a few loose buds which are identical in structure

to those on the isotype of M. heteropleurum Bl. (Korthals sn. from Sumatra).

The only distinction is that the blade venation of M. heteropleurum is more

prominent. Blume’s descriptions of these two species differ only with the blade

venation. Fruiting collections of M. heteropleurum, e.g. Bakh. Sr. 1777 (Sumatra)-

and Achmad 779 (P. Simeuloee); match those of Blume s.n. from Java which

has blades identical to those on the holotype of M. excelsum Bl. The majority

of specimens in the Leiden coliection of M. excelsum have the M. heteropleurum

- type blade venation, however since all other features between these two species

are the same it is necessary, due to priority, to reduce the latter to a synonym

of the former.

The holotype of M. subtrinervium Mig. (Teysmann 835, from Sumatra) at

Utrecht is without any axes, flowers, or fruit. Miquel (lc. 322) did note that

the species has few flowered inflorescences with a peduncle c. 7 mm long. The

young berries are said to be oviod, constricted at the top; and have a persistent,

4-lobed calyx limb. The branches on the holotype are cylindric and the blades

have a distinct intramarginal nerve; both of these features match the two types

of M. heteropleureum at Leiden — especially the smaliest leaf on the isotype.

The larger leaves on this isotype resemble M. excelsum s.s_ Since it is virtually

impossible to determine the structure of the inflorescence, and what the leaves

and fruit of M. subtrinervium looked like, it appears from the specimens avail-

able that it is the same as M. excelsum. It is obvious that many botanists have

been confused over M. subtrinervium and have, as an alternative, referred most

of their specimens to M. heteropleurum and M. excelsum. The isotypes of M.

elmeri Merr. (Elmer 21646, from Sabah) at Singapore and Leiden are identical

to the holotype of M. heteropleurum Bi. Both sheets are in bud, however the

leaves and flowers (especially the immature anthers) match the buds on the holo-

type of M. heteropleurum. They are both, therefore, synonyms of M. excelsum

Bl. Furtado reduced M. elmeri to a synonym of M. maingayi Cl. M. Kurzii King

(l.c. 77), originally described as M. subtrinervium Miq. var. grandiflorum Kurz

(‘grandiflora’) (J. As. Soc. Beng. (1876) 131), is recorded by King from the

Nicobar islands. The specimens of King’s coll. 509, cited by King under this

species are in poor condition, and the specimen from BM is without inflore-

scences and flowers. The specimen from Kew is somewhat better in that the

axes are intact and there are a few loose post-flowering specimens. The branches

and blades of these two sheets are very similar to those of M. excelsum BL.,

however the inflorescence is larger (up to 1 cm long) and the calyx is described

as being undulate with 4 broad teeth. This feature is not apparent with the

post flowering material. In any case, the calyx remnant on these immature fruit

is irregularly cracked and it is suspected that King misinterpreted these as lobes.

The size and shape of these pieces are very similar to those of M. maingayi.

78 Gardens Bulletin, Singapore XX XIII (1980)

Memecylon maingayi Cl. is ungestionably the same as M. excelsum and

M. heteropleurum. The holotype of M. maingayi (Maingay 814 (1422 ) has larger

buds than the holotype of M. heteropleurum, but this is merely a developmental

factor; the structure of the inflorescences, and flowers — especially the bud

anthers — of the two specimens are the same. ‘There is variation in the pro-

minence of the blade venation and texture, but with numerous collections of

M. excelsum and its synonyms available the general vein pattern is the same.

Nicobar Island — King’s collector 509 (type M. kurzii King)

Malay Peninsula

Kedah — Bukit Perak For. Res.: Chan 13120

Perak — Larut: King’s collector 4726; Lumut: Ridley 9474; Pondok Tanjong:

Sallih 9710; Tapa: Wray 1310 (lectotype M. heteropleurum BI. var. olivaceum

King); sine loc.: Scortechini 89, Yong 94670

Kelantan — Gunong Sitong: Nur & Foxworthy 12257; Kuala Krai: Ismail 104273

Trengganu — Bukit Kajang: Corner sn, 30 Oct. 1935; Gunong Tebu For. Res.:

Kochummen 2507; Sekayu For. Res.: Suppiah 11846, 11865; Gunong Padang:

Whitmore 12543, Belara: Wood 76072

Pahang — Aur For. Res.: Whitmore 3631; Pulau Tioman: Kadim & Noor 580;

Kemasul For. Res.: Temerloh: Kochummen 98586; Whitmore 0078: Kota

Glanggi: Henderson 22482; Lepar For. Res.: Suppiah 108998; Lesong For.

Res.: Ahmad & Shukor 450; Tahan River: Ridley 2240

Negri Sembilan — Seremban: Ismail 109424

Malacca — Selangor: Alvins 245, 430: sine loc.: Maingay 814 (1422) (type

M. maingayi Cl.)

Johore — Anbong, Endau: Kadim & Noor 416; Bekok For. Res.: Jumali 4100;

Bukit Tinjar Laut: Ngadiman 36931; Gunong Besar: Everett 14076; Kluang

Reserve: Kochummen 2824, Ng 97953; Kota Tinggi: Corner 28702; Maokil For.

Res.: Shing 68877; Mersing: Wyatt-Smith 76278; Sarom: Ridley sn in 1900;

Sungei Kayu: Corner 29245, 32246, 32504, 32759, Kiah 31981, 32311; Ulu Sungei

Segamat: Jumali 4496 (973), Samsuri & Shukor 727.

Further Distribution — Sumatra, Simeuloee Is., Mentawai Islands, Java,

Borneo (Sabah, Kalimantan) (fide Bakh. f.).

12. Memecylon floridum Ridl., J. Str. Br. Roy. As. Soc. 79 (1918) 73.

Tree 10-17 m tall with a diameter of up to 15 cm; bark brown-grey, finely

fissured, thin; slash inner bark thin, brown; slash wood yellow-orange, hard;

upper nodes thickened, 4-angled to 4-auriculate; upper internodes sharply 4-angled

or somewhat compressed and appearing 2-grooved, glabrous, c. 1.5 mm thick,

drying greyish to tan; older branches mostly cylindric with widened nodes, drying

greyish; blades subcoriaceous, lanceolate to elliptic, acuminate-rostrate at the

tip (acumen 1-2 cm long), sharply narrowed and decurrent at the base; mid-

nerve sunken above, raised and tapering below; secondary venation pinnate, faint;

6.5—-8.5 cm long, 2.5-3 cm wide; drying greenish, often with a brown hue above,

greenish below; petiole 5-6 mm long, c. 1.5 mm wide; inflorescence cymose,

Revision of Memecylon L. 19

from leafy and upper leafless nodes, 10-20 mm long, many flowered, glabrous:

bracts and bracteoles lanceolate, acute, c. 0.5 mm long, falling off early; primary

axes 1 or 2 per axil, flattened, 5-9 mm long with 1, less commonly 2, nodes;

secondary axes not developed, glomerulate at the tips of the primary axes,

pedicels c. 4 mm long; calyx campanulate, widened above the ovary, 1.5 mm

long, 2 mm wide, margin truncate to undulate with 4 broadly triangular lobes,

each c. 0.25 mm long, smooth outside, ridged internally, yellow-green; petals

broadly ovate, acute at the tip, truncate at the base, slightly thickened with

thinner margins, 3 mm long, 2.5 mm wide, white: filaments c. 2.5 mm _ long,

anthers axe-shaped, without a gland, purple; stigma minute, style slender, c.

4 mm long, immature fruit pryiform, becoming globose when mature, 10-13 mm

diameter, calyx remnant slightly raised, flattened, areolus c. 3 mm wide; exocarp

smooth, green turning yellow then pinkish, drying blackish; pericarp fibrous,

ec. 2 mm thick.

Figure 12: a. calyx, b. petal, c. stamen

vernacular: nipis kulit (Perak, Kelantan)

This species is very closely related to M. cinereum King (q.v.) in having

similar inflorescences and nearly identical flowers. The axes are slightly shorter

in M. cinereum and its blades are less acute at the base, have invisible venation,

and dry black. A major difference lies in the fruit viz. those of M. cinereum

are smaller and have a very thin, non-fibrous pericarp.

Memecylon malaccense (Cl.) Ridl. (g.v.) and M. lilacinum Z. & M. (q.v.)

also have similar inflorescences, flowers (especially the glandless anthers), but

have different leaves and fruit. M. acuminatum Sm. (q.v.) is readily distinguished

by having an anther gland, smalier fruit, larger and more complex inflorescence,

and cylindric branches.

Perak — Gunong Bubu: Everett 13922; Larut: King’s collector 3551 (type).

3870

Kelantan — Kuala Mersing: Ng 5460; Relai For. Res.: Cockburn 7414

Trengganu — Sekayu For. Res.: Suppiah 11867; Ulu Bendong, Kajang: Corner

30190; Ulu Besut: Cockburn 8288

Pahang — Raub: Renggol 20341; Tasek Bera: Chen & Noor 249

Negri Sembilan — Tampin For. Res.: Shing 17093

Singapore — Liew 37742, Ngadiman 36360, Ridley 6219

13. Memecylon fruticosum King, J. As. Sac. Bengal 69, II: 1 (1900) 74 (Mat.

Fl. Mal. Pen. III, 482): M. epiphyticum King l.c. 74 (lc. 482), syn. nov.;

M. tenuifolium Ridl., Fl. Mal. Pen. I (1922) 812, syn. nov.; M. depokkense

Bakh. f., Med. Mus. Bot. Utrecht 91 (1943) 339 and Rec. Trav. Bot. Neerl.

40 (1943-45) 339, syn. nov.

Shrub 1.5-2.5 m tall, often epiphytic, or a treelet up to 3 m tall; branches

sharply 4-angled to 4-winged, generally prominently so below the nodes giving

an auricled appearance, c. 1.5 mm thick, glabrous; blades subcoriaceous, glabrous,

lanceolate to ovate, acuminate at the tip (acumen up to 1 cm long), rounded

at the base, 5-13 (16) cm long, 1.5-6 cm wide; venation pinnate, 10-12 pairs

80 Gardens’ Bulletin, Singapore X X XIII (1980)

of nerves, sunken and obscure above, slightly raised or obscure below; intra-

marginal nerve distinct or obscure, 14 mm from the margin, broadly looping;

mid-nerve sunken above, raised and tapering below; light green when young,

turning yellowish/greenish, drying dark brown with a roughened texture above,

brown to greenish-brown below; petiole 2-3 mm long, 1-1.5 mm thick, glabrous;

inflorescence cymose, often in pairs from leaf axils, sometimes from leafless nodes,

1-1.5 cm long; primary axes flattened, 0.5-1 mm wide, 1-6 mm long with 1

or extended for 1 mm with a second node, secondary axes glomerulate or up

to 1 mm long, pedicels 1.5-5 mm long, clustered; bracts and bracteoles ovate,

acute, thin, up to 0.75 mm long, caducous; calyx funnelform, c. 1 mm _ long,

3 mm wide, smooth outside, ridged internally, truncate, or with 4 undulations,

or 4 broadly triangular, acute lobes, sinuses wide; petals oblong to broadly ovate,

tipped with a mucro, truncate at the base, 2 mm long, 1.25—1.5 mm wide, margins

thinner, mid-line thickened and slightly raised dorsally; filaments 2—2.5 mm long;

anthers “‘U’’-shaped with a distinct gland, c. 1 mm long; style cylindric, c. 3 mm

long, ovules 9-12; immature fruit oblong-pyriform, 8-10 mm long, 6-7 mm wide,

becoming globose, 8-10 mm diameter, when mature; calyx remnant distinctly

raised (c. 1 mm), areolus c. 4 mm wide; exocarp smooth, green turning light red

when ripe; pericarp c. 0.5 mm thick.

Figure 13: a. branch, leaf, and inflorescence, b. calyx, c. petal,

d. anthers

King notes that M. fruticosum and M. epiphyticum approach M. dichotomum

(Cl.) King (q.v.), but the latter has several major differences viz. 4-angled to

cylindric branches which are not winged, more prominent leaf venation; a much

shorter inflorescence with larger, thicker, and persistent bracts; papillose calyx

tube; and larger fruit with a thick pericarp. The anthers of the 3 species have

a similar ““U” shape. Unfortunately, all three syntypes and other collections of

M. fruticosum seen lack petals and mature fruit, however from its entirely different

inflorescence M. fruticosum is a distinct, albeit poorly known, species.

Memecylon epiphyticum King merely differs from M. fruticosum in having

a shorter (more reduced) inflorescence, fewer flowers, and smaller leaves. The

two syntypes (King’s coll. 5184, and Wray 2727) were collected after flowering

and the specimens are in immature fruit. These parts are identical to King’s

coll. 2971 which is the lectotype of M. fruticosum.

Memecylon tenuifolium Ridl. hardly differs from M. fruticosum and M.

epiphyticum except that its leaves are narrower and the inflorescence is shorter

(c. 5 mm). The holotype of M. tenuifolium (Kassim 0748) at Kew has 3 loose

fruit which are depressed globose, c. 10 mm diameter, and are without a raised

areolus. Since Ridley did not describe the fruit of this species, plus the fact

that these fruit differ from those of M. fruticosum, it is strongly suspected that

they have been attached to the sheet by mistake. It is not common to find

both flowers and mature fruit on the same specimen of Memecylon.

Memecylon depokkense Bakh. f., from Java, is another variant of M. fruti-

cosum which has shorter inflorescences (7-8 mm long), and differs from M.

tenuifolium in having more flowers and wider blades. Both species, in all other

respects, are identical to M. fruticosum.

Memecylon corticosum Ridl. (q.v.) is similar to M. fruticosum in the angled-

winged branches, and structure of the inflorescence; and it is possible that the

Revision of Memecylon L. 8]

mature stamens are also “U’’-shaped. M. corticosum differs in having larger,

lanceolate blades with prominent venation. As collections of either species that

have any transitional features have not been seen, the two species are maintained

as separate taxa in this paper.

Penang — Government Hill: Ridley sn, Mar. 1896; West Hill: Curtis 1773

Perak — Larut: King’s collector 2971 (lectotype), 3265 (syntype), 3425 (syntype),

5184 (syntype M. epiphyticum King); Kunstler 3265; Sungei Larut: Wray 2727

(lectotype M. epiphyticum King)

Trengganu — Bukit Kajang: Corner 30329

Selangor — Bukit Payong Kajang: Symington 24235

Pahang — Telok Mungkuang, Temerloh: Kassim 0748 (type M. tenuifolium Ridl.)

Negri Sembilan — Gunong Angsi: Sow 23712

Singapore — Ridley 5753

Further Distribution — Natoena Islands (west of Sarawak), Java (fide Bakh. f.).

14. Memecylon garcinioides Bl., Mus. Bot. Lugd.-Bat. 1:23 (1851) 358.

Shrub up to 5 m tall or a tree 6-15 m high; upper internodes cylindric,

sometimes slightly flattened and shallowly grooved immediately below the upper

node, c. 1 mm thick, drying greyish, older branchlets drying khaki-tan; blades

subcoriaceous, elliptic to ovate, acuminate at the tip (acumen 8-10 mm long),

narrowed and slightly decurrent at the base, midnerve sunken above, raised and

tapering below, secondary venation very obscure to invisible on both surfaces;

5.5-10 (13.5) cm long, 2-6 cm wide; drying dark brown to blackish or sometimes

greenish above, brown often mottled with a greenish hue below; petiole 24 mm

long, 1-1.5 mm thick; inflorescence from leaf axils, cymose, 1-3.5 cm long, many

flowered, axes flattened, glabrous; bracts and bracteoles lanceolate, acute, c. 0.5

mm long, falling off before the petals mature: primary axes 1-3 per axil, 0.4-1.3

(2) cm long with 1 or 2 nodes; secondary axes not developed or up to 4 mm

long; pedicels 2-4 mm long; calyx campanulate, slightly constricted above the

ovary, c. 2 mm long, and slightly wider, margin truncate with 4 minute, triangular

cusps, internal ridges prominent, drying with a slightly rugose-papillose texture;

bud petals dome shaped, acute; mature petals broadly ovate to suborbicular,

acute at the tip, truncate at the base, thickened with a keeled mid-rib, 1.5-2 mm

long, 1.5—-2 mm wide; anthers suborbicular, 0.75-1 mm long, with a thin, flat,

connective extension with a long shallow gland along its entire surface; filaments

c. 2 mm long; stigma minute, style slender, 3-3.5 mm long; fruit globose, 6-7

mm diameter, areolus 2-3 mm wide internal ridges distinct, calyx remnant slightly

raised very thin; exocarp drying black or greyish-green, with a slightly roughened

texture, c. 0.25 mm thick.

Figure 14: a. calyx, b. petal, c. stamen

The truncate calyx with 4 minute points and suborbicular anthers with a

tail-like extension (actually a remnant) of the connective distinguish this snecies.

The holotype of M. garcinioides Bl. var. elongatum BI. (lc. 358), which was

considered to be identical to M. intermedium by King (l.c. 86) and a synonym

82 Gardens’ Bulletin, Singapore XX XIII (1980)

of M. garcinioides by Bakh. f. (l.c. 363), has blades up to 13 cm long and 4.5

cm wide with inflorescenes 10-12 mm long. It is obvious, therefore, that the

lengths of the blades and inflorescences of all these taxa and specimens studied

at Lieden and Singapore do not correlate.

Furtado (Gard. Bull. Sing. 20 (1963) 119) annotated a set of specimens in

the Singapore collection as M. hepaticum Bl., which are actually M. garcinioides.

The type specimen of M. hepaticum (Blume s.n., from Sumatra) at Leiden is

without inflorescences, flowers, and mature fruit; therefore it is not certain what

this species really is since the description is inadequate for a critical study.

The specimens from Johore and Malacca all have lanceolate leaves, but the

inflorescences and flowers (especially the globose anthers) strongly resemble M.

garcinioides. For this reason this set of specimens has been cited under M.

garcinioides. No specimens from the Malay Peninsula have been seen with the

same kind of leaves as the type collection of M. hepaticum. The anthers of

M. intermedium BI. (q.v.), especially in bud, often resemble those of M. garci-

nioides. Good distinguishing features of M. intermedium include its smaller and

narrower leaves, and calyx with 4 triangular lobes.

Perak — Gunong Boobo: King’s collector 7394; Gunong Malacca: King’s collector

7152, Kunstler 7123; Jor Batang Padang: Henderson 10886; Larut: King’s

collector 1984, 2938; Maxwell’s Hill: Wray 2961, 3203; Sungai Seluang: Shah

& Shukor 3451; Ulu Bubong: King’s collector 10034

Kelantan — Sungai Renong: Nur & Foxworthy 12159

Selangor — Bukit Kulu: Goodenough 10812; Gunong Bunga Bua: Ng 1160;

Kuala Lumpur: Mohamet 2024; Raweang: Ridley 7333; Sungei Bulok: Hamid

1600, Symington 21077; Ulu Gombak: Strugnell 12738

Pahang — Gunong Senyum: Evans 13190; Taman Negara: van Balgooy 2584;

Tasek Bera; Stone 9472a, 9474

Negri Sembilan — Gunong Angsi, Loh 17304

Malacca — sine loc.: Alvins 1212; Mt. Ophir: Keng et al. 6312

Johore — Endau: Kadim & Noor. 314; Gunong Panti: Everett 13837, H.M.

Burkill 3189; Joh: Ridley 14167; Kota Tinggi: Corner 32491; Mawai: Corner

30882; Mawai — Jemaluang Road: Corner sn, on 9 Feb. 1935; Mawai — Kota

Tinggi Road: Corner 30976; Sungei Kayu: Corner 29251, 32476, Kiah 31961;

Sungei Kayu Ara: Corner 29481; Ulu Langat For. Res.: Whitmore & Wong

115654

Singapore — Ngadiman 34975, Rao sn; Ridley 4805,, 8118, 14171

Further Distribution — _ peninsular Thailand, Sumatra, Java, Borneo

(Kalimantan), Karimata Is. (fide Craib and Bakh. f.).

15. Memecylon globosum Bakh. f., Med. Mus. Bot. Utrecht 91 (1943) 355 and

Rec. Trav. Bot. Neerl. 40 (1943-45) 355.

Revision of Memecylon L. 83

Shrub up to 2 m tall or a tree up 5 mm ttall; branchlets cylindric, often

grooved immediately below the upper node on 2 sides, smooth, glabrous, 1.5—2

mm thick, drying tan; blades coriaceous; oblong, elliptic, or ovate; acuminate

at the tip (acumen up to 12 mm long), narrowed and shortly (1-3 mm) decurrent

at the base, midnerve distinct, intramarginal and secondary nerves obscure to

invisible above, more distinct below; 7-20 (23) cm long, 3-11.5 cm wide; drying

brownish above, lighter brown below; petiole flattened, dorsally grooved, 5-8 mm

long, 1.5—3.5 mm thick; inflorescence cymose, 1-1.5 cm long, from leaf axils,

sometimes terminal, few to many flowered, glabrous; bracts ovate, acute, c. 1.5

mm long, caducous; primary axes generally several per axil, flattened, 2.5-5 mm

long, with one node, lateral axes shorter; pedicels 3-4 mm long; calyx cam-

panulate, thickened, widened above the ovary, c. 3 mm long, 4 mm wide; margin

with 4 short, broadly rounded lobes, sinuses splitting in mature flowers, glabrous:

petals thick, (sub) orbicular, c. 4 mm long, 4 mm wide, obtuse at the tip, truncate

at the base, reflexed at maturity; filaments c. 2.5 mm long; anthers “J”? shaped,

c. 2 mm long, connective thick with a prominent gland; 14-16 mm diameter,

areolus flattened, c. 4 mm wide; exocarp greenish (immature), drying browaish-

tan with a slightly roughened texture; pericarp fibrous, c. 1 mm thick.

Figure 15: a. inflorescence and leaf, b. calyx, c. petal, d. stamen

The branches and leaves closely resemble those of M. oleifolium Bi. (q.v.)

which has, among other major differences, longer and more complex inflorescences,

and elliptic-ovate, pustulate fruit. M. globosum is easily distinguished from other

species of Memecylon by its short inflorescence, relatively large flowers, 4 broad

calyx lobes, and large, globose fruit.

The type collection of M. globosum (van Steenis 1265, from the Natoena

Islands west of Sarawak) consists of buds, and the mature flowers and fruit were

not previousiy described. Several other collections from Borneo, Johore, and

Singapore compare closely with this type collection, thus enabling a complete

description of this species. There is considerable variation in the size and shape

of the leaves, however the inflorescence, flowers, and fruit seem to be constant.

The species is apparently rare and ranges from southzrn Johore, Singapore, to

Borneo.

Johore — Gunong Blumut, Kluang: Shah-& Sanusi 2173

Singapore — Bukit Timah: Ridley 6768

Further Distribution — Natoena Islands, Borneo (fide Bakh. f.).

16. Memecylon hullettii King, J. As. Soc. Beng. 69, II: 1 (1900) 76 (Mat. FI.

Mal. Pen. III, 484).

Shrub or treelet up to 3 m tall; branchlets cylindric, often flattened and

shallowly grooved below the upper node, 1.5—-2 mm thick; blades coriaceous,

lanceolate, ovate, to oblong; acuminate at the tip (acumen up to 2 cm long),

broadly rounded and usually cordate at the base; venation pinnate, 15-20 pairs

of nerves, very obscure but visible on both surfaces; intramarginal nerve 1-2 mm

from the margin; 12-27 cm long, 3.5-8 cm wide; drying dark brown above,

lighter brown below: petiole 1-2 mm long, shorter than the basal lobes of the

blades, thus appearing sessile, 1.5-2 mm thick; inflorescence a compound umbel,

6-8 cm long, often reflexed, many flowered; axes muricate-papillose; primary

84 Gardens’ Bulletin, Singapore XX XIII (1980)

axes solitary from leafy nodes, 4-5 cm to the first node of the inflorescence,

often extending beyond the first node for 0.5-1.5 cm; secondary axes 5-7, ranging

in length from 2 mm (peripheral ones) to 1.5 cm (central one); pedicels 4-5

mm long, papillose, in terminal clusters ranging from 3-5 flowers (peripheral

axes) to c. 25 on the terminal axis; bracts ovate, acute, c. 0.75 mm long, appear-

ing clustered, caducous; calyx campanulate, flattening somewhat after maturity,

truncate with 4 minute cusps, widened abruptly above the ovary, papillose out-

side, ridged internally, 1.5 mm long, 3 mm wide; petals coriaceous, broadly ovate,

broadly round and apiculate at the tip, rounded and shortly clawed at the base,

slightly keeled in the upper 4 dorsally, margins thinner, 3 mm long, 4 mm wide,

white, filaments c. 2.5 mm long, anthers blue, c. 1 mm long, suborbicular in

outline, connective shortened and reduced to about the same size as the locules,

gland distinct, situated on the curved tip of the connective; stigma minute, style

slender c. 5 mm long; ovules c. 6; fruit globose, 7-9 mm diameter, capped by

the persistent calyx remnant, c. 0.5 mm high, with a thickened, undulate margin,

areolus c. 2-3 mm wide, internal ridges generally indistinct; exocarp smooth, c.

0.2 mm thick, white, then pinkish, finally blueish when ripe, drying blackish.

Figure 16: a. branch, leaf, and inflorescence, b. calyx and corolla bud,

c. petal, d. stamen

The broadly rounded or cordate, nearly sessile leaves; long primary axis,

and compound umbellate inflorescence distinguish this species from all others

in the region. The leaves of M. dichotomum (Cl.) King (q.v.) are often similar,

but its angled branchiets and glabrous inflorescence axes immediately distinguish

it from M. hullettii. The anthers of M. garcinioides Bl. (q.v.) appear similar,

but all other vegetative and floral characteristics of the two species differ con-

siderably.

Johore — Gunong Belumut: Holttum 10854; Gunong Muntajak: Nur 19976;

Gunong Pulai: King & Hullett 253 (lectotype), Mat 3739, Sinclair 39549; Khiang:

Holttum 9436; Kluang For. Res.: Alphonso, Sanusi, Sidek 210; Shah & Sanusi

2133; Kota Tinggi: Shah 454; Pelepah Kiri: Corner sn on 19 Oct. 1936; Sedenah:

Ridley 13501; Sungai Batu Pahat: Lake & Kelsall 4073 (syntype); Sungai Buloh

Kasap: Corner 29984; Sungai Kayu Ara: Corner 29234; Sungei Pelepoh Kiri:

Corner sn, on 26 June 1938; Virgin Jungle Res.: Ahmad 390

17. Memecylon intermedium Bl., Mus. Bot. Lugd.-Bat. I:23 (1851) 358.

Tree up to 25 m tall, with a diameter up to 1 m, buttressed in larger trees,

bark grey-brown, smooth, rugose, finely fissured and flaking off, inner bark

brownish; branches cylindric, often with a shallow groove on 2 faces which

usually disappears before the next node, I-1.5 mm thick, drying tan to brown;

blades subcoriaceous to coriaceous, glabrous, lanceolate to ovate, acuminate at

the tip (acumen up to | cm long), narrowed and decurrent at the base, mid-

nerve sunken above, raised below, secondary venation extremely obscure or in-

visible; 6-9 cm long, 2-3 cm wide, drying dark brown to blackish above, brown

below, with a roughened texture; petiole flattened, 3-4 mm long, 1.5—2 mm thick;

inflorescene from.leaf axils, cymose, 1.5—-2 cm long, many flowered, glabrous;

bracts and bracteoles lanceolate, c. 0.5 mm long, caducous; primary axes usually

solitary, flattened, 7-12 mm long usually with 1 node, seondary axes glomerulate

or up to 2 mm long, pedicels 1-2 mm long; calyx campanulate, 1.5-2 mm long,

2-3 mm wide, widened above the ovary, smooth and glabrous, flattening after

flowering, margin truncate with 4 cusps or short lobes, internal ridges prominent;

Revision of Memecylon L. 85

petals broadly ovate to sub-orbicular, 2.5-3 mm long, c. 3 mm wide, acute at

the tip, truncate at the base, thickened with a slightly raised midrib, margins

thinner, pinkish; filaments c. 2 mm long, pale blue; anthers “‘J’’-shaped, c. 2 mm

long, gland distinct; stigma minute, style cylindric c. 3 mm long; fruit globose.

6-8 mm wide, calyx remnant nearly flat or slightly raised, areolus c. 2 mm wide,

exocarp smooth, green ripening blue, drying black, pericarp c. 0.2 mm_ thick.

Figure 17: a. branch, leaves, and inflorescence;

b. calyx, c. petal, d. anthers

In terms of the inflorescence and flowers this species closely resembles M.

acuminatum Sm. (q.v.), however that species has larger fruit with a thicker, more

rugose pericarp, and thinner blades. The leaves and fruit of M. glomeratum

Bl. (known from Sumatra and Borneo) are very similar to those of M. intermedium,

however the former has much shorter (apparently nearly sessile) inflorescences.

Authentic inflorescences and flowers of M. glomeratum have not been seen,

thus further comparisons cannot be made here. Blume notes that M. intermedium

is close to M. garcinioides Bl. (q.v.), however I feel that its affinities lie with

M. edule Roxb. (q.v.). M. garcinioides has sub-orbicular anthers with a curious

appendage which can be used to immediately distinguish the species.

The anthers of M. intermedium Bl., especially in bud, are closest to those

of M. garcinioides Bl. (q.v.) which differs in having larger leaves and a minutely

apiculate calyx. The anthers of M. garcinioides have, in general, a more reduced

connective, thus giving the anther an orbicular rather than “J”? shape. Several

collections from Java known as M. intermedium Bl. var. longipes Bl. (l.c. 359),

and reduced to the typical variety by Bakh. f. (l.c. 363) have inflorescences up

to 6 cm long. Sepcimens of this species from the Malay Peninsula have inflore-

scences 1.5—2 cm long.

The typical variety of M. edule Roxb. (q.v.) is also close to M. intermedium

and can be distinguished by having inflorescences behind the leaves, blades which

dry greenish or brownish — never black, and 4-angled or 2 grooved branches.

Kedah — Gunong Jerai: Chelliah 9801, Evans & Gorden 95; Hou 798, 814;

Stone 8635; Stone & Mahmud 8505

Trengganu — Gunong Padang: Whitmore 12690

Penang — Penara Bukit: Curtis sn in June 1890

Perak — Slim Hills For. Res: Whitmore 0812

Kelantan — Bukit Brangkat: Shah & Ali 2880; Gunong Stong: Whitmore 12453

Selangor — Gunong Bunga Bua: Ng 1153, Whitmore 0345; Semangko For.

Res.: Whitmore 12556; Ulu Langat For. Res.: Whitmore 12175; sine loc.:

Kepong no. 85084

Pahang — Balok Game Res.: Kochummen 2095; Fraser’s Hill: Nur 11259;

Gunong Benom: Whitmore 3194; Gunong Lesong, Rompin: Shah & Shukor

3130: Kuala Kelepah: Shah & Noor 1767; Kuala Teku: Whitmore 4803;

Meranoh: Shing 17265

86 Gardens’ Bulletin, Singapore XX XIII (1980)

Malacca — Gunong Ledang: Ridley 3296

Johore — Bukit Chongkrak: Shah & Samsuri 3635; Kuala Palong For. Res:

Everett 14263; Rengam For. Res.: Suppiah 17770

Further Distribution — Sumatra, Java (fide Bakh. f.).

18. Memecylon kunstleri King, J. As. Soc. Beng. 69, II:1 (1900) 76 (Mat. FI.

Mal. Pen. III, 484).

Tree up to 20 m tall, diameter up to 45 cm; branches cylindric, striate, often

with two very shallow grooves below the upper node, becoming cylindric, c.

1 mm thick, drying tan; blades subcoriaceous, glabrous, lanceolate-ovate, shortly

acuminate (c. 5 mm) at the tip, rounded and often shallowly cordate at the base,

venation pinnate, 10-12 pairs of nerves, obscure on both surfaces in smaller

blades, distinct on larger ones, sunken above, slightly elevate below; intramarginal

nerve obscure or distinct, 2-2.5 mm from the margin; 6-13 cm long, 2.5-5 cm

wide; deep glossy green drying dull dark brown above, brown below; with a

roughened texture; petiole 2-3 mm long, 1-1.5 mm thick; inflorescence cymose,

24.5 cm long, glabrous, from leaf axils, many flowered, axes sharply 4-angled;

primary axes | or 2 per axil, 10-35 mm long with 1 or 2 nodes, secondary axes

3-15 mm long, pedicels 2-3 mm long; calyx campanulate, flattening afier flowering,

truncate or undulate at the margin, with 4 minute cusps, smooth outside, pro-

minently ridged inside, 1.25-1.5 mm long, 2 mm wide; petals thickened with

thinner margins, oblong to broadly ovate, rounded and apiculate at the tip, trun-

cate at the base, 1.5-2 mm long, 2—2.5 mm wide, waxy white; filaments c. 2.5

mm long, bud anthers ‘‘J’’-shaped, “‘U’’-shaped when mature, c. 1 mm _ long,

gland distinct; dark brown; style c. 3 mm long, ovules 12-15; immature fruit

ovoid, 7-9 mm long, 4-5 mm wide, crowned with the obscurely 4-toothed calyx

remnant, areolus c. 3 mm wide, waxy pale green, drying grey-greenish, pericarp

c. 0.2 mm thick.

Figure 18: a. branch, leaves, and inflorescence;

b. calyx and style, c. petal, d. stamen, e. areolus

The cylindric branches and glabrous inflorescence axes distinguish this species

from M. paniculatum Jack (q.v.), M. fruticosum King (q.v.), and M. corticosum

Ridi. (q.v.). The leaves of M. kunstleri closely resemble those of the type collec-

tions of M. appendiculatum Bl. and M. nudum Bl. — both of which were reduced

to synonyms of M. costatum Mig. by Bakh. f. (lc. 345), and now synonymous

with M. paniculatum. These species have similar inflorescences, but differ in

having 4-angled branches and pubescent inflorescence axes.

The dry blades and *“‘U” shaped anthers of M. kunstleri and M. fruticosum

are similar, however the winged branches and shorter inflorescences of the latter

are obvious differences.

Kedah — Sungai Terap, Selma: Henderson 35447

Penang — Penang Hill: Nauen sn, in April 1940

Perak — Changkat, Serdang: Wray 744; Gunong Dipang: King’s collector 8195

(syntype); Ulu Bubong: King’s collector 10419 (lectotype)

Selangor — Labu: Ridley 7334

Revision of Memecylon L. 87

19. Memecylon lilacinum Zoll. & Mor., Syst. Verzeich. (1845-46) 9; M. myrsinoides

BI. var. lilacinum (Zoll. & Mor.) King, J. As. Soc. Bengal 69, If: 1 (1900)

81 (Mat. Fl. Mal. Pen. III, 489); M. myrsinoides Bl., Mus. Bot. Lugd.-Bat.

1:23 (1851) 356 syn. nov.; M. laevigatum BI., |.c. 358 syn. nov.; M. laevigatum

BI. var. sulcicarpum Furtado in scheda, Herb. Singapore; M. pseudo-nigrescens

Bl., lc. 357 syn. nov.; M. pseudo-nigrescens Bl. var. acuminatum BI. in

scheda, Leiden; M. confine Bl., |.c. 357 (syn. by Bakh. f., Med. Mus. Bot.

Utrecht 91 (1943) 361 and Rec. Trav. Bot. Neerl. 40 (1943-45) 361).

Tree up to 25 m tali, diameter up to 40 cm, bole shorily fluted; less frequently

a shrub up to 3 m tall; bark pale grey-brown, thin, finely fissured and flaking;

slash inner bark thin, brown; slash wood dark yellow to white, wood dense and

sinks in water; upper nodes flattened, prominent after leaf fall, black when dry;

upper internodes often shortened (8-12 mm) or 2-3 cm long, 1-1.5 mm thick,

flattened and grooved on 2 sides, often somewhat 4-angled, becoming cylindric,

drying greyish to khaki-tan; blades coriaceous, glabrous, lanceolate to ovate,

acuminate at the tip (acumen 5-15 mm long), acute and decurrent at the base,

2.5-8.5 cm long, (1) 1.5—-4 cm wide, mid-nerve sunken above, raised below, other

venation invisible; dark glossy green drying dark brown to blackish above, dull

green drying brown to dark brown below; petiole 4-8 mm long, 1-1.5 mm thick,

glabrous; inflorescence from the axils of lower leaves or upper leafless nodes,

cymose and appearing glomerulate, many flowered, 5-8 mm long, glabrous; bracts

and bracteoles subulate, c. 0.25 mm long, falling off before the petals mature:

primary axes flattened, with 1 node, 1-1.5 mm long, secondary axes 3-5, glome-

rulate or c. 1 mm long each with 3 pedicels c. 1 mm long; calyx campanulate,

glabrous, smooth, 1.5 mm long, 1.25-1.5 mm wide, tan, margin truncate with

4 distinct triangular cusps each c. 0.25 mm high; corolla buds dome shaped,

acute, tips twisted; petals lanceolate-oblong, sharply acuminate at the tip, truncate

at the base, 1.5—2 mm long, c. 1 mm wide, slightly thickened with a keeled dorsal

mid-nerve, margins thinner; white, lilac, or blue; filaments c. 1.5 mm long, lilac;

anthers c. 1.5 mm long without a gland, whitish; style slender, 3-5 mm long,

lilac; ovules c. 8; fruit globose, 8-12 mm diameter, flattened at both poles, calyx

remnant minute, flattened in the plane of the fruit; areolus c. 1.5 mm wide not

sunken; generally with 4-8 shallow (often indistinct) vertical grooves, especially

near the pedicel; exocarp smooth when immature, often becoming somewhat

rugose in mature dry specimens, infrequently with minute whitish pustules, green

when immature, turning yellow to pale -orange, finally black when ripe, grey-

green or black when dry; pericarp 1-2 mm thick, gritty.

Figure 19: a. inflorescence, b. calyx and corolla bud, c. petal, d. stamen

Figure 31: bark

vernacular: delik (Selangor), nipis kulit (Kedah, Johore), tengading (Temuan)

The very compact cymes, calyx with 4 cusps, narrow and acuminate petals,

glandless anthers, and the depressed, grooved, globose fruit distinguish this species.

Memecylon edule Roxb. (q.v.) can be confused here, but it has no or extremely

minute calyx points, wider and blunter petals, an anther gland, and a smooth,

globose fruit with a thin pericarp and no grooves.

Some specimens may be confused with M. garcinioides Bl. (qv.) wh.ch has

cylindric branches without grooves, thinner blades, and different flowers (especially

the anthers) and fruit. The fruit on some specimens, e.g. Chan 6764, have dried

88 Gardens’ Bulletin, Singapore XX XIII (1980)

black and the exocarp has scattered, minute pustules. Everett 13821 has fruit

with a similar colour and texture, but the shape is more pyriform with a narrowed

tip, unequal base, and lacks grooves. The type specimens in Leiden of M.

myrsinoides (Blume 1374 from Java) all have relatively small blades, 3-4.5 cm

long and 1.5-2.5 cm wide on branches with short (8-12 mm) upper internodes.

The inflorescences on the specimen designated as the holotype are about 5 mm

long with numerous flowers. These specimens, at first glance, appear to differ

from the type specimens at Leiden of M. laevigatum Bl. collected by Blume

(s.n.) from Borneo. The blades are generally larger, 5.5-8.5 cm long, 2.5-4 cm

wide, and the upper internodes are all over 2 cm long. The inflorescences are

essentially the same except that the secondary axes are slighily less obscure.

The two type specimens of M. myrsinoides Bl. var. lilacinum (Zoll. & Mor.)

King in the Leiden collection (Zollinger 187) are intermediate between the two

species discussed above. The blades are 5-6.5 cm long and 2.5—3.25 cm wide

and the inflorescences are structurally the same, but with more flowers and longer

(c. 5 mm) styles. The upper nodes are from 1.5—2 cm long, that is intermediate

between the other two species. All three taxa have identical flowers which are

typified by the truncate margin with 4 distinct cusps; thin, oblong, acuminate

petals, and the uniquely shaped anthers without a gland. Fruit of the large

leaved M. laevigatum and the smaller leaved M. myrsinoides and various inter-

mediate specimens from several sheets at Singapore and Leiden are the same.

A specimen collected by van Steenis (12657) from Java has branches and

leaves which match the type specimens of M. myrsinoides and the inflorescences,

being very dense and having long styles, are the same as those on Zollinger’s

collections of var. lilacinum. Paie 13320, from Sarawak, has short internodes

characteristic of M. myrsinoides and larger blades which range in size for those

of var. lilacinum and M. laevigatum. Dilmy et al. 11, from east Java, has both

small and medium sized blades, relatively long upper internodes, and _ inflore-

scences similar to those of M. myrsinoides and M. laevigatum. Even on the

type specimens of M. myrsinoides and M. laevigatum the range of blade size

overlaps, and with all taxa the upper internodes are shallowly grooved on 2 sides

and dry a khaki-tan colour.

Unfortunately, variation in the length of the upper internodes and size of

the blades has caused some taxonomic confusion. Cogniaux (l.c. 1159, 1160)

separated the species mainly on the size of the inflorescences and considered

M. lilacinum as a synonym of M. myrsinoides. Bakh..f. (1.c. 335) maintained both

species and var. lilacinum as distinct taxa. Since all three taxa are structurally

idental and have intermediate traits which link all of them togetther, they are

considered as representing one species and in this paper have been lumped

under the oldest name viz. M. lilacinum Zoll. & Mor.

Memecylon cinereum King, M. floridum Ridl., and M. malaccense (C1.) Ridl.

have very similar inflorescences, calyx lobes, and identical glandless anthers. ‘The

blades, petals, and fruits differ (q.v. discussions under these species). The holo-

type of M. peudo-nigrescens (Blume s.n. from Sumatra) in the Leiden collection

has branches, blades; and most important, immature fruit which is depressed

globose, grooved, and with a thick, rugose pericarp. It is, undoubtedly, the

same as M. myrsinoides Bl. et al. Other specimens in the Leiden and Singapore

collections confirm this fact.. The blades in most specimens of M. pseudo-

nigrescens are ovate and fall within the upper size range of M. myrsinoides and

.—_— ——— |

Revision of Memecylon L. 89

match the smaller ones of M. laevigatum. ‘There are no flowers on the holotype

of M. pseudo-nigrescens, however, Lorzing 10163 from Sumatra has inflorescences

and flowers, especially the glandless anthers, similar to the holotype of ™.

myrsinoides. Ldorzing’s specimen has the calyx margin distinctly undulate with

4 triangular cusps. This feature differs slightly from the truncate calyx in the

holotype of M. myrsinoides BI.

There is no reason to maintain M. pseudo-nigrescens BI. as a distinct taxon

and since it cannot be adequately distinguished from M. myrsinoides et al. it

should be combined with M. lilacinum Zoll. & Mor.

A specimen collected by Korthals (s.n.) from Sakoembang, Borneo has M.

pseudo-nigrescens Bl. var. acuminatum written on the original label. This speci-

men is identical to the holotype of M. pseudo-nigrescens Bl. Bakh. f. (Lc. 362)

placed this specimen in his list of species under M. pseudo-nigrescens. 1 agree

with Bakh. f. (l.c. 361) in reducing M. confine BI. (l.c. 357) to a synonym of

M. pseudo-nigrescens. Cogniaux (l.c. 1160) combined M. pseudo-nigrescens and

M. confine under M. glomeratum BI. (l.c. 356), however the fruit of the holotype

of M. glomeratum Bl. at Leiden is globose with a smooth, thin pericarp which

dried black. The branches and leaves of M. glomeratum do, however, match

those of M. myrsinoides, M. pseudo-nigrescens, et. al. Flowers of M. glomeratum

have not been described and all the material of this species at Leiden lacks

flowers. It is not certain, therefore, what M. glomeratum really is, but at least

from the immature fruit is not the same as M. lilacinum Z. & M. M. oligoneurum

Bl. (q.v.) has similarly shaped, glandless anthers; but differs greatly in the

3-nerved blades, different calyx and petals, 2 ovules, and globose fruit. M.

minutiflorum Miq. (q.v.) has similarly shaped fruit, but differs in having larger

fruiting axes, a raised areolus, and blades which dry with lighter colours.

The fruit in some specimens, e.g. Goh 15701 and Chew 726 is obovate to

oblong, often with grooves; thus differing from the typically depressed globose

shape of most fruiting material. Furtado annotated Goh’s specimen as M.

laevigatum Bl. var. sulcicarpum Furtado var. nov. (in scheda Herb. Sing.). This

variety is not accepted here.

Perlis — Ginting Kabok: Ridley 15058; Kaki Bukit: Kiah 35281

Kedah — Bukit Enggang: Everett 13768; Bukit Perak For. Res.: Chan 13118;

Bukit Selambau: Meh 8977; Gunong Inan For. Res.: Whitmore 4678: 48th

mile Jeninag Road: Kiah 36152; Pulau Adang: Ridley 15778; Pulau Langkawi:

Batten-Pooll sn, Keng et al. 30, Stone 6934; Ulu Muda For. Res.: Chan 6764

Penang — Waterfall: I.H. Burkill 6893; Garden: Curtis 100 in’ Dec. 1884, Curtis

100 at Government Hill ino 1885; Curtis = 100 at the waterfall in March 1893,

1503, 2219, 3596, sn at the waterfall in June 1893 and June 1901, sn; King’s

collector 1457; Ridley 7962, sn at the waterfall in June 1893 and March 1915

Perak — Larut: King’s collector 3517, 3768, 5923; Wray 2258; Ulu Bubong:

King’s collector 10442; Ulu Kerling: King’s collector 8828; sine loc. Scortechini 87

Kelantan — Kulal Badong: Henderson 10395

Trengganu — Sungei Pelong: Suppiah 14877

90 Gardens’ Bulletin, Singapore XX XIII (1980)

Selangor — Sungei Buloh: H.M. Burkill sn on 24 Nov. 1956; H.M. Burkill

& Shah 1048: Kochummen 2568; Sungei Lalang Kejang: Symington 24224;

Ulu Langat: Gadoh 1137, 1295; Weld’s Hill: Ahmad 2477; CF 2477, 6419;

Kochummen 99517; Pawanchee 12902; coll. ? 6410

Pahang — Lesong For. Res.: Ahmad & Shukor 450A; Rompin: Goh (Soh)

15701; Gunong Benom, Ulu Krau: Yusoff 99102; Temerloh: Hamid 10678

Negri Sembilan — Gunong Angsi: Shing 17338; Gunong Tampin: Holttum 9570

Malacca — Bukit Senanau: Holttum 9669; Julutong: Goodenough 1767; Poloh

Diak: Alvins 163: Poloh Dulle: Alvins 2296; Pulau Rumbia: Seimund sn; sine

loc.: Alvins 69, Griffith 2328

Johore — Gunong Panti: Chew 726; Labis For. Res.: Ahmad & Shukor 523;

Pulau Tuiggi: I.H. Burkill 903; Sedili Ketchil: Corner 28562; Sungei Sedili:

Corner 25896, sn on 28 March 1932; Tana Runto: Ridley 2026; Tebrau River:

Ridley 13502 (aff.)

Singapore — Corner sn on 7 May 1937, Hullett 390, Jumali 943; Maxwell

76-790, 77-84, 78-56; Ridiey 1815, 3848, 4804, 6218, sn in 1892 and 1893;

Walker 176: Ahmad 1416, 1475

Further Distribution — Indo-China, Burma, Thailand, Sumatra, Java, Banka

Island, Borneo (Kalimantan), Karimata Is., Celebes (fide Craib and Bakh. f.).

20. Memecylon malaccense (Cl.) Ridl., J. Str. Br. Roy. As. Soc. 79 (1918) 73;

M. amabile Bedd. var. malaccensis Cl. in Hk. f., Fl. Brit. Ind. Il (1879) 555.

Shrub 1-4 m tall, branches widely 2-grooved and appearing 4-angled, slightly

winged below the upper nodes, becoming cylindric below, c. 1 mm thick; blades

coriaceous, broadly ovate to broadly lanceolate, acuminate at the tip, broadly

rounded and usually shallowly (c. 0.5 mm) cordate at the base, venation extremely

obscure to invisible on both surfaces, midnerve sunken above, raised and taper-

ing below, 4-7 cm long, 2.5-4 cm wide, drying brown above, lighter brown

below; petiole 0.5-1 mm long, | mm wide, basal lobes of cordate blades longer

than the petiole and appearing sessile; inflorescence cymose, 8-10 mm long, from

leaf axils, glabrous; bracts and bracteoles lanceolate-ovate, acute, c. 0.75 mm long,

falling off early; primary axes | or 2 per axil, cylindric, 1.5-3 mm long, secondary

axes glomerulate or c. 0.25 mm long, pedicels clustered at the tips, c. 1 mm

long, 4-8 per inflorescence; calyx campanulate, truncate with 4 broadly rounded

and minutely mucronate lobes, smooth outside, ridged internally, 1.5-2 mm long,

2-3 mm wide; petals thick with thinner margins, broadly ovate, acute at the

tip, clawed at the base, c. 3 mm long, 2.5 mm wide; anthers without a gland;

ovules c. 6; fruit globose, capped by the raised calyx remnant which often has

4 distinct points, areolus c. 2 mm wide, exocarp smooth, drying black, pericarp

c. 0.25 mm thick.

Figure 20: a. calyx and style, b. petal, c. stamen

M. malaccense Ridl. appears to be very closely related to M. lilacinum Z.

& M. (q.v.). The branches, inflorescence, calyx, and mainly the anthers are very

similar. The petals of M. malaccense are more ovate and shortly clawed at

the base. The blades of M. malaccense differ in having rounded and very

shallowly (c. 0.5 mm) cordate*bases. The fruit also differs in being globose with

a smooth, thin exocarp without grooves.

Revision of Memecylon L. 9]

I do not think that M. malaccense (Cl.) Ridl. should be united with M.

lilacinum Z. & M. (q.v.) since there are some important differences: mainly the

fruit and the leaves. Specimens of M. lilacinum with shallowly cordate blades

or smooth fruit have not been seen. More material of M. malaccense is needed

in order to properly understand its relation with M. lilacinum.

M. malaccense and M. cinereum King (q.v.) are essentially the same, except

that M. cinereum has larger blades which are acute and decurrent on a jonger

and thicker petiole; and a shorter, ic. more glomerulate, inflorescence. The

petals of M. lilacinum are generally narrower, the calyx lobes usually better

developed, and the fruit is much different.

M. floridum Ridl. (q.v.) has nearly identical flower morphology. however

the blades and fruit, namely the fibrous pericarp, differ significantly.

Malacca — Maingay 819 (2528) (syntype), 819 (2531) (lectotype).

21. Memecylon megacarpum Furtado, Gard. Bull. Sing. 20 (1963) 121; M.

pulchrum Cogn. in DC., Monogr. Phan. 7 (1891) 1141, nom. illeg. not Kurz,

For. Fl. Brit. Burma I (1877) 510 (= M. caeruleum Jack var. pulchrum

(Kurz) Cl. in. Hk. f., Fl. Brit. India [I (1879) 559); M. heteropleurum BI.

sensu King, J. As. Soc. Bengal 69, I1:1 (1900) 78 (Mat. FI. Mal. Pen.

III. 486).

Tree up to 15 m tall with a diameter up to 30 cm, or a shrub up to

3 m tall; bark smooth, dark grey to brown, thinly fissured and flaking; inner bark

brownish, yellow; slash wood pale brown; youngest branches slightly flattened

with a groove on 2 sides, becoming cylindric, c. 2 mm wide, drying tan to

brown; t-lades subcoriaceous, elliptic to ovate, acuminate at the tip (acumen

up to 1.5 cm long), gradually narrowed and somewhat rounded at the base;

glabrous; mid-nerve sunken above, raised and tapering below; secondary venation

pinnate with 20-30 pairs of nerves, straight, sunken above, slightly raised below;

intramarginal nerve 2-3 mm from the margin, straight; finer venation very obscure

to invisible; 15-20 cm long, 6-7.5 cm wide; drying greenish to dark brown above,

greenish to brown telow; petiole 1-3 mm long, 2—2.5 mm thick; inflorescence

of clustered cymes, from behind the leaves; 9-12 mm long, many flowered,

glabrous; axes flattened; bracts and bracteoles lanceolate, acute, 1-2 mm long, falling

off before the petals mature; primary axes clustered, often on raised tubercles,

1-2 mm long, 1.5 mm wide at the base, usually with 1 node; secondary axes

indistinct or up to 2 mm long; pedicels 2-3 mm long; calyx campanulate,

widened above the ovary, thick, margin truncate with 4 very shallow (best seen

in bud) undulations, each with a minute cusp, indistinguishable in mature flowers

where there are 4 minute thickenings below the rim, minutely papillose outside,

prominently ridged inside, c. 5 mm long, c. 5 mm wide, pink; petals broadly

ovate to sub-orbicular, glabrous, broadly and often irregularly rounded at the

tip, narrowed to a broad claw at the base, c. 5 mm long, 6 mm wide, very

thick with thinner margins, reflexed at maturity; filaments c. 2.5 mm long; anthers

“C” or “J-shaped, c. 2 mm long, gland very prominent; style c. 4 mm long,

ovules c. 10; fruit globose, 16-18 mm diameter, aerolus slightly (1 mm) raised,

c. 3 mm wide, exocarp green, drying olive to brown, with a roughened texture;

pericarp c. 3 mm thick; seed with a liquid inside.

Figure 21: a. calyx and style, b. petal, c. stamen, d. fruit (lateral view),

e. fruit (dorsal view) with areolus

92 Gardens’ Bulletin, Singapore XX XIII (1980)

Memecylon megacarpum Furtado is mostly closely related to M. dichotomum

(Cl.) King (q.v.), the latter having angled branches, smaller leaves, and fewer

flowers per inflorescence. Both species have large, globose fruit which are similar

to those of M. amplexicaule Roxb. (q.v.) which has thick, coriaceous, cordate

blades.

King misidentified several collections of M. megacarpum (e.g. Maingay 816

and Curtis 814) as M. heteropleurum Bl., which has been reduced to a synonym

of M. excelsum Bl. (q.v.) in this revision.

Memecylon excelsum has the calyx widened above the ovary, different blade

venation, and oblong to elliptic fruit. These distinctions are not obvious, except

in fruit, unless the type collections are examined.

While it is certain that the species described above is distinct from M.

dichotomum and M. excelsum, it is not certain that M. megacarpum is the correct

name. Cogniaux described M. pulchrum as having much larger leaves (30-35

cm long, 9-11 cm wide) and longer pedicels (3-5 mm), which have basal bracts.

By this it is assumed in this paper that the species lacks secondary axes. The

type collections of M. pulchrum (Beccari 585 and 1833, from Sarawak) were

not seen during this research. The epithet M. pulchrum Cogn. cannot be used

since it is a later homonym of M. pulchrum Kurz which was reduced to M.

caeruleum Jack. var.pulchrum (Kurz) Cl. by Clarke which is an entirely different

species from the Andamans.

Kedah — Bongsu For. Res.: Everett 14169; Bukit Enggang For. Res.: Whitmore

0405; Gunong Bintang: Haniff 21048; 33rd mile Jeniang Road: Kiah 35959;

Koh Mai For. Res.: Kiah 35153

Penang — West Hill: Curtis 814; sine loc.: Wallich 4102A

Perak — Gunong Batu Patch: Wray 1066, 1148; Gunong Bubu For. Res.: Hou

662; Larut: King’s collector 6621; Pangkor Island: Whitmore 3081; Slim Hills

For. Res.: Whitmore 0817; Sungai Ryah: King’s collector 1110; Ulu Kerling:

King’s collector 8589, 8689; Upper Perak: Wray 3425; Waterfall, Taiping: Burn-

Murdoch 162

Kelantan — Gunong Stong: Whitmore 12415; Ulu Sungei Lebir Ketchil: Cockburn

7115

Trengganu — Bukit Lanjut For. Res.: Loh 13459; Gunong Bubu Treng: Suppiah

11746; Gunong Tebu, Jabi; Shah, Shukor, Awang 3303; Ulu Telemong For. Res:

Loh 13445

Selangor — Bukit Lagong, Kepong: Suppiah 108881, Kuchummen 79112; Bukit

Tanggah: Everett 13781; Fraser’s Hill: IH. Burkill & Holttum. 7869; Gading

For. Res.: Chan 11239; Kuala Lumpur: Ridley 2053, Mat 2053; Ulu Gombak

For. Res.: Kochummen 2353, T. & P. 43 (2643), Yong 99002; Weld’s Hill:

Hamid 965, Jaamat 10266

Pahang — Batu Balain: I.H. Burkill & Haniff 15829; Bukit Beserah For. Res. :

1.H. Burkill & Haniff 16135, Whitmore 3751; Kadouchong, Pulau Tawar: Ridley

2242; Tahan River: Ridley -2340; Kemasul For. Res.: Ismail 98912; Kuala

Lumpat, Krau: Soepadmo 759; Lesong For. Res.: Suppiah 14891; Robinson’s

Revision of Memecylon L. 93

Falls, Cameron Highlands: Henderson sn; Raub: Sohadi 14661, Strugnell 20464,

Syed-Alli 23365; Sungei Lemoi: Jaamat 28188; Sungai Teku: Kiah sn, on 29

July 1936; Taman Negara: Everett 14458; Ulu Sungai: Shah & Noor 1743, 1843

Negri Sembilan — Bukit Tangga: Ridley sn, in Dec. 1920; Gunong Angsi For.

Res.: Nur 11699, Sohadi 14612

Malacca — Sungei Udang: Derry 582

Johore — Bunong Ma’okil Maur: Samsuri & Shukor 957; Gunong Panti: Corner

36290; Mawai-Jemalang Road: Corner 28683, 29016; Sedenah: Ridley 13507

Singapore — Henderson 35916: Maingay 815 (2746), 816 (3112); Ridley 3614a,

5092, 5928, 6215, 9210; Ahmad 1474

Further Distribution — Sumatra, Borneo (Sarawak) (fide Cogniaux and King).

22. Memecylon minutiflorum Migq., Fl. Ned. Ind. Suppl. I, Sumatra (1860) 323,

King, J. As. Soc. Beng. 69, II: 1 (1900) 80 (Mat. Fl. Mal. Pen. HI, 488).

Tree up to 25 m tall with a diameter up to 60 cm; bole straight, buttressed

in larger individuals; bark red-brown to brown-tan or blackish, finely fissured,

flaking, thin; slash inner bark pink to red, cambium whitish-purple; slash wood

orange-yellow or cream; youngest branches usually somewhat flattened with a

vertical groove between internodes on 2 opposite sides, often with 2 raised ridges

flanking each groove or 4-angled; older branches cylindric with little or no trace

of the grooves or ridges, smooth, tan, c. | mm thick; blades subcoriaceous,

glabrous, lanceolate, elliptic to broadly ovate, caudate-acuminate (acumen 1-1.5

cm long) at the tip, cuneate at the base and shortly decurrent on the petiole

(c. about half its length); venation pinnate, 8—10 pairs of nerves, invisible to very

obscure on both surfaces; intramarginal nerve invisible to very obscure, c. 1 mm

from the margin; mid-nerve distinct, sunken above, raised and tapering below;

5-10 cm long, 2—5.5 cm wide; dark green, drying light to dark brown above,

light brown or olive-greenish to yellowish below; petiole 3-6 mm long, 1-1.5 mm

thick, glabrous; inflorescence cymose from leaf axils, or less commonly just

behind the leaves, 7-15 (17) mm long, many flowered; axes flattened, grooved

on 2 sides, glabrous; bracts and bracteoles lanceolate, acute, up to 1.5 mm long,

caducous; primary axes usually solitary, 2-13 mm long, secondary axes glome-

rulate or up to 4 mm long; pedicels clustered, 1-1.5 mm long, minutely papillose;

calyx tube funnelform, constricted above the globose ovary, truncate with 4 cusps,

minutely papillose outside, tube 1-2 mm long, c. 2 mm wide, ovary 1 mm

diameter; petals broadly ovate to suborbicular, obtuse to acute at the tip, trun-

cate at the base, thin, midline slightly thickened, not keeled, c. 2-2.5 mm long,

1.5-2 mm wide, reflexed at maturity, pink; style c. 4 mm long; ovules 10-12;

fruit globose, often flattened at both poles, sometimes with 4 shallow longitudinal

grooves, or somewhat gibbous, 7-10 mm diameter, calyx remnant slightly (0.75

mm) raised, areolus c. 2 mm wide; exocarp green, drying yellow-green or grey-

green, with a roughened, almost mealy, texture, pericarp 1-2 mm thick, gritty

or not gritty.

Figure 22: a. branch and inflorescene, b. calyx and style, c. petal,

d. stamen

vernacular: kemuning tikus (Temuan)

94 Gardens’ Bulletin, Singapore XX XIII (1980)

This species is easily recognized by the papillose pedicels and calyx; bulbous

ovary abruptly widened into the funnelform, 4-cusped calyx; globose fruit with

a rugose-mealy texture, and blades which dry concolorous. The holotype of

M. minutiflorum at Utrecht (Diepenhorst 2337, from Sumatra) lacks inflorescences,

flowers, and fruit. The branches and blades, however, compare well with many

other specimens which have flowers and fruit that match King’s description,

e.g. Corner 30336, Mahmud 819, Haniff 15528, King’s coll. 6265). Reconstructing

an idea of what the holotype is has been done with the literature and other

specimens, therefore it is reasonably certain that the description above fits the

species.

It should be noted that the type collection of M. grande Retz. var. khasiana

Cl. (Griffith 2333, from the Khasia Mountains in East Bengal), which King

(I.c. 83) reduced to a synonym of M. celastrinum Kurz, has branches and leaves

identical to the holotype of M. minutiflorum. The fruit of Griffith’s collection

is smooth and has a larger calyx remnant.

Memecylon acuminatum Sm. (q.v.) is, in many respects, similar to M. minuti-

florum, but has cylindric branches, smooth calyx, and smooth fruit with a wider

and flat areolus. M. acuminatum Sm. var. flavescens Cl. (q.v.) is also closely

related but differs from both species by having a truncate calyx. The exact

relationship of var. flavescens to these two species is uncertain, and in this paper

King’s reduction of var. flavescens to a synonym of M. minutiflorum Miq. is not

accepted.

Memecylon minutiflorum has fruit which closely resembles those of M™.

lilacinum Z. & M. (q.v.). The two species can be easily distinguished by the

very short (appearing glomerulate) infructescence, flat areolus, and blades which

generally dry dark brown to blackish of the latter.

Memecylon constrictum Craib (Kew Bull. 1930, 324, only known from the

type collection of Haniff & Nur (2732), from peninsular Thailand) has a short

cymose inflorescence and a distinct constriction above the globose ovary which

is similar to that of M. minutiflorum. M. constrictum, however, has a smooth,

truncate calyx, and larger, thicker leaves. The type collection at Kew has a few

very immature buds and from the stamens and petals the species could be

related to M. minutiflorum. The species is poorly known and there is insufficient

material available to study the matter any further (Figure 29, a. — c.).

Thailand

Huey Mut: Kiah 24400; Puket: Haniff & Nur 2732 (type M. constrictum Craib)

Malay Peninsula

Kedah — Bukit Perak: Everett 13681; Gunong Baysu: Meh 17954; Gunong

Jerai: Stone & Mahmud 8540, Everett 13674; Jeniang Road: Kiah 35978; Sawa

For. Res.: Langkawi: Chelliah 6945; N. Pulau Dayang, Langkawi: Whitmore

15043; Machinchang For. Res.: Langkawi: Whitmore 11216, 15030; Langkawi:

Wilkinson 20776; Pulau Timon: Whitmore 15105; Sungei Batu Asap, Langkawi:

Haniff 15528; Sungai Lugong: Kiah 35086

Penang — West Hill Res.: Burn-Murdoch 28: Reservoir Line, West Hill: Curtis

816

Perak — Larut: King’s collector 5027, 6105, 6265; Maxwell’s Hill: Ridley 9312,

sn in June 1893, Wray 3240

Revision. of Memecylon L. 95

Trengganu — Bukit Kajang: Corner 30336, 30479; Sri Bangun: Sinclair & Kiah

39870; Ulu Sungei Trengganu: Cockburn 10509

Selangor — Bukit Peringkot: Kochummen 1502, Gadoh 1502, Gading For. Res. :

Shing 13357, Ginting Bidai: Ridley 7312; Ulu Gombak: Mahmud 819; Ulu

Selangor: Goodenough 10601; Weld’s Hill: Cubit 867

Pahang — Shah 172; Cameron Highlands: Everett 13664; Endau, Rompin:

Mahamud 15519; Gunong Seuyum: Henderson 22211; Jerantut: I-H. Burkill &

Haniff 16102; Kota Tongkat: Evans 13187; Kuala Tekam: Evans 13188; Lesong

For. Res.: Ahmad & Shukor 447, Samsuri & Shukor 447; Taman Negara:

Everett 14462

Negri Sembilan — Gemas: I.H. Burkill 4970; Gunong Sungei For. Res.: Osman

23759; Jelebu For. Res.: Suppiah 11285; Tebong For. Res.: Holttum 9641

Malacca — Bukit Panchor: I.H. Burkill 3046; Bukit Sidenau: Derry 127; Sungei

Udang: Alvins 21; sine loc.: Alvins 129, 2302, sn; Maingay 810 (2551)

Johore — Bukit Badok: Hassan & Kadim 92; Gunong Ma’okil: Samsuri &

Shukor 958; Gunong Panti West: Samsuri & Shukor 768; Kluang: Whitmore

8691; Kuala Telung Tinggi: Ridley 11078; Maokil For. Res.: Shing 6865, 6873;

Mawai-Jemulang Road: Corner 29021, 29044, 29450; Sungei Kayu: Kiah 32057;

Sungei Kayu Ara: Corner 28733, sn on 10 Feb. 1935; Sungei Sedili: Corner: 31942

Singapore — Kassim 261

Further Distribution — peninsular Thailand, Sumatra (fide Craib).

23. Memecylon oleifolium Bl. (“‘oleaefolium’’), Mus. Bot. Lugd.-Bat. 1:23 (1851)

359 (no. 877); M. ambiguum BI1., l.c. 359 (no. 879) syn. nov.; M. micranthum

BI., lc. 360; M. acuminatissimum Bl., |.c. 360 syn. nov.; M. horsfieldii Miq.,

Fl. Ned. Ind. I (1855) 572; M. grande Retz. var. horsfieldii (Miq.) Cl. in

Hk. f., Fl. Brit. Ind. II] (1879) 558; M. lampongum Migq., Fl. Ned. Ind.

Suppl. I, Sumatra (1860) 321; M. brandisianum Craib, Kew Bull. 1936, 323

syn. nov.

Tree up to 12 m tall with a diameter-up to 45 cm; bark thin, smooth,

fissured to slightly scaly, greyish to whitish; inner bark yellow; sap wood white

to brown; branches cylindric, 1-2 mm thick, drying greyish to khaki; blades sub-

coriaceous to coriaceous, glabrous, elliptic to ovate (often broadly so), acuminate

at the tip (acumen up to 17 mm long), acute and decurrent at the base; venation

invisible to distinct above; invisible, obscure, or prominent below; mid-nerve

sunken above, raised and tapering to the tip below; dark glossy green above,

pale green below; drying dark brown above, brown below; (6) 9-21 cm long,

(2.5) 3-10 cm wide; petiole (4) 6-9 mm long, 1.5—2.5 mm wide, glabrous; inflore-

scence from leafy or upper leafless nodes, cymose, 1.5—5 cm long, glabrous, axes

2-grooved to 4-angled; bracts and bracteoles lanceolate, acute at the tip, 0.5

mm long; primary axes 0,8-4 cm long, secondary 2-12 mm, tertiary axes generally

not developed or sometimes up to 2 mm long, pedicels 2.5-7 mm long; calyx

campanulate, truncate, 1.5-3 mm long, 2-4 mm wide, glabrous, cream coloured

with a purple hue; petals thickened, broadly ovate, acute and mucronate at the

tip, truncate at the base, 2-2.5 mm long, 2.5 mm wide, mid-nerve thickened and

slightly raised dorsally, margins thinner, blue to purple; filaments c. 2 mm long,

anthers ““C”’ or “‘J’’-shaped, purple, gland prominent; style 3-4 mm long; ovules

96 Gardens’ Bulletin, Singapore XX XIII (1980)

9-12; fruit elliptic-ovate, 8-10 mm long, 6-8 mm wide, pedicels 3-8 mm long,

calyx remnant raised, c. | mm high, areolus c. 2 mm wide; exocarp green turning

light yellow, black and smooth or with minute pustules when dry; pericarp c.

0.5 mm thick.

Figure 23: a. calyx and corolla bud, b. petal, c. bud stamen, d. mature

stamens, e. fruit

At first glance, the holotypes of M. oleifolium Bl. and M. acuminatissimum

Bl. appear to be very different; the former has smaller blades with indistinct or

invisible venation and the latter has larger blades with conspicuous venation.

The holotype of M. acuminatissimum (Korthals s.n., from Sumatra) lacks inflore-

scences and has only a few loose flowers buds. Comparison of this specimen

with other collections plus the original description has helped in reconstructing

an idea of what the species is. Cockburn 10839 and Whitmore 8773 are flowering

and fruiting specimens, respectively, which have branches and leaves similar to

the holotype of M. acuminatissimum. The inflorescences on Cockburn’s specimen

range from 2-5 cm long, with distinct secondary (infrequently with tertiary) axes,

pedicels 4-5 mm long, and a calyx 3 mm long and 4 mm wide. Whitmore’s

fruit are elliptic, c. 14 mm long, and 8 mm wide.

Distinguishing M. acuminatissimum from M. oleifolium, especially with nume-

rous collections of specimens which have been referred to either one, is difficult.

Some distinguishing features of M. acuminatissimum include the larger: leaves,

inflorescences, and calyx. Similar features include the structure of the inflore-

scence, flowers (especially the anthers), and the fruit. There are several specimens

which have blades with faint venation resembling those of M. oletfolium, e.g.

Cockburn 10839 and Chew 728, but both of these have the inflorescences and

calyx similar to M. acuminatissimum. Ng 97986 has leaves approaching the

size of M. acuminatissimum, but the venation is invisible. The inflorescence

is more similar to that of M. acuminatissimum. There are other specimens

which combine features of these two species, e.g. Whitmore 12320 (flowers) and

Kochummen 2618 (fruit) in which the inflorescence and fruiting axes are like

M. acuminatissimum and the leaves resembling both species. In at least two

other specimens, Jumali & Heaslett 3056, and Cockburn 10839, the larger leaves

resemble M. acuminatissimum and the smaller ones M. oleifolium. Since there

are no structural differences that can be used to separate these two species, plus

the fact that the leaves have intermediate forms, I feel that it is necessary to

lump M. acuminatissimum with M. oleifolium, thereby expanding the range of

variation of the latter even further.

Memecylon laurinum BI. (1.c. 359, no. 878), which Bakh. f. (l.c. 357) reduced

to M. oleifolium BI. var. laurinum (Bl.) Bakh. f. is only known from three

collection which include the two types (Korthals s.n., from Sumatra, and Korthals

s.n. from Borneo) and another specimen (without collector or number) from

Sumatra. Only the two specimens from Sumatra can be studied in detail since

the other type is without flowers or fruit.

The buds of var. laurinum differ from those of the typical variety in that

the calyx is quite large (c. 4 mm long) and apiculate, and the inflorescence is

shorter and more compact. The bud petals and anthers are of identical size

and shape to those of M. oleifolium. The branches and leaves are also the

same. The mature of the calyx differs from M. oleifolium and since there is

Revision of Memecylon L. 97

little similarity between the two taxa I do not feel confident in reducing the

variety to a synonym of the typical variety since its mature flowers and fruit

are still unknown.

Blume (lc. 359) notes under M. ambiguum Bl. that the inflorescence is

more dense, the blade venation is more prominent, and the base of the blades

is more rounded that with M. oleifolium. The type specimens of both species

at Leiden have been examined and while the vegetative features are somewhat

different, the structures of the inflorescence and the morphology of the flowers

are the same for both species.

Another species, M. micranthum BI. (l.c. 360) was reduced to a synonym

of M. oleifolium Bl. by Cogniaux (I.c. 1150). The type specimen of M. micranthum

Bl. at Leiden is exactly the same as the type specimen of M. ambiguum BI. in

all respects. Bakh. f. (lc. 353) combined M. micranthum with M. ambiguum,

but maintained M.. oleifolium as a distinct species. Cogniaux (l.c. 1143), how-

ever, considered M. ambiguum as a good species and included it in his mono-

graph in a different section from M. oleifolium. It is not understood how

Cogniaux could have done this since the leaves of M. ambiguum and M.

micranthum are exactly the same. This is not true with the leaves of M.

oleifolium.

In any case, the structure of the inflorescences and the flowers are the same

for all three species. This, therefore, is very good evidence for combining all

of them under one species viz. M. oleifolium which was published above the

other two. More evidence for this involves King’s coll. 4420 from Larut, Perak.

This specimen, cited by King (l.c. 83) as M. oleifolium combines features of

this species and M. ambiguum. The relatively small leaves and their texture

resemble M. ambiguum, however the acute base and the obscure venation are

like M. oleifolium. The inflorescence has many flowers on thicker axes which

is a M. ambiguum feature. The flower morphology matches both species. Bakh.

f. annotated this specimen as Memecylon spec. and it is easily understood why

since it is an apparent intermediate form between the two species.

In comparison to all the more typical specimens of M. oleffolium examined,

King’s coll. 4420 is very unusual. The specimens of M. ambiguum and M.

micranthum at Leiden, at first glance, appear to be distinct from M. oleifolium.

Similarly, most of the specimens of M. oleifolium match its type specimen. Un-

fortunately, there are only a few specimens of M. ambiguum and M. micranthum

at Leiden. thus I am not certain of any other variation within this group.

Koorders 24271, from Java, is the only specimen of M. ambiguum (including

M. micranthum) at Leiden that has fruit, and these match those of several collec-

tions of M. oleifolium.

From observations during this research and from the opinions of those

botanists mentioned above, it seems that these three species are the same. I have

little doubt that M. micranthum is the same as M. ambiguum and confidence is

maintained in combining both of them with M. oleifolium BI.

The holotype of M. lampongum Miq. (Teysmann 4281, from Sumatra) at

Utrecht is in very immature calyx bud, but from the branches, leaves, and structure

of the inflorescence this specimen matches Maingay 811 and to a lesser extent,

due to its having more mature flowers, Blume’s type collections of M. oleifolium.

98 Gardens’ Bulletin, Singapore XX XIII (1980)

Bakh. f. suggests that M. lampongum may be related to M. ambiguum, which

it certainly is, but the striking similarities with Maingay 811 and Blume’s material

is more obvious. I agree with King in reducing M. lampongum Migq. to a synonym

of M. oleifolium Bl. Agreement is also made with King (l.c. 83) in reducing

M. grande Retz. var. horsfieldii (Miq.) Cl. to M. oleifolium. A specimen at Leiden

(Maingayi 811) matches the types of the latter collected by Korthals (s.n.) from

Sumatra. This variety, with less prominent nerves, more acute buds, and shorter

inflorescences, lacks any structural differences from M. oleifolium. It cannot,

therefore, be considered distinct from M. oleifolium.

Memecylon pubescens (Cl.) King (q.v.) has flowers similar to those of M.

oleifolium, but the inflorescence of the former has many more flowers which are

congested in glomerules on the tips of the secondary axes. Also these axes are

pubescent and the venation of the blades, especially below, is much more prominent.

Transitional forms linking the two species have not been seen, therefore it is

better to maintain both of them as separate species.

The type specimens of M. brandisianum Craib (Kerr 14169 from peninsular

Thailand) match the type specimens of M. oleifolium Bl. at Leiden. The holotype

from Kew and the isotypes from BM and BK of M. brandisianum are in bud,

however the calyx, petals, and stamens match a bud specimen collected by Blume

(s.n.) from Sumatra. The leaves of Kerr’s specimens have dried with a green

colour on both surfaces in contrast to the typical brown colour of dried M.

oleifolium blades.

Craib, in his original description, notes that M. brandisianum is related to

M. garcinioides Bl. (q.v.), however it is obvious from the anthers of Kerr’s speci-

mens that the relation is only superficial. The stamens of M. garcinioides are

very unusual and quite distinct, and in no way, even in bud, resemble those of

M. brandisianum or M. oleifolium.

Thailand

Terutao: Kerr 14169 (type M. brandisianum Craib)

Malay Peninsula

Kedah — Miujol: Meh 17879

Perak — King’s collector 4420, 4439, 8198; Ulu Kerling: King’s collector 8571;

sine loc.: Scortechini 2069; Grik: Yong 94662

Kelantan — Bukit Yong: Shah & Shukor 3232; Ulu Sat For. Res.: Suppiah

104585

Trengganu — 34th mile Jerengau Road: Sinclair & Kiah 40939; Sungei Nipa:

Corner sn on 20 Nov. 1935; Ulu Brang: Moysey & Kiah 33632; Ulu Bendong:

Corner 30017; Ulu S. Loh: Cockburn 10839

Selangor — Gunong Simpah: Hume 8481, 8910, 9159; Sungei Lalang Kajang:

Symington 22644; Ulu Gombak: Kochummen 2718; Ulu Langat: Gadoh 1781;

sine loc.: Alvins 659

Revision of Memecylon L. 99

Pahang — Bukit Berelah, Kuantan: Shah, Sidek, Samsuri 3722; Kampong Aur:

Samsuri & Shukor 484; Kemasul For. Res.: Temerloh: Kochummen 98582,

Whitmore 0079; Lepar For. Res.: Suppiah 108952; Lompat Krau: Whitmore

3583; Pulau Tioman: Henderson 21651; Sabi Estate, near Bentong: Shah 223;

Ulu Kuantan: Craddock sn in 1903; Ulu Sungei Sat: Shah & Noor 1842

Negri Sembilan — Bukit Tangga: Ridley sn in Dec. 1920; Gunong Angsi: Nur

11679; Kuala Pilak: Tassin 4498

Malacca — Ayer Panas: Derry 1199; Bukit Besar: Ridley sn; Bukit Sutu, Sungei

Ujong: Alvins 1970; Bukit Tumiang: Alvins 2035; Sungei Ujong: Alvins sn;

sine loc.: Maingay 811 (1228)

Johore — Gua Riman, Kluang: Jumali & Heaslett 3056: Gunong Blumut: Whit-

more 8773; Gunong Ledang: Whitmore 12320, 12341; Gunong Panti: Ridley sn

in 1892, Holttum 18095, Corner sn on 20 Jan. 1936; Chew 728; Gunong Pulai:

Ridley 12179; Kluang For. Res.: Ng 97986; Labis For. Res.: Ahmad & Shukor

521, Kochummen 2618; Segamat: Shing 17155; Sungai Kayu Ara: Kiah 31994,

32079; Corner sn on 8 and 10 Feb. 1935; Virgin Jungle Reserve, Mersing:

Ahmad 376

Singapore — Ridley 6414A, 6416

Further Distribution — Sumatra, Java, Soemba Is. (fide Bakh. f.).

24. Memecylon oligoneurum Bl. Mus. Bot. Lugd.-Bat. I:23 (1851) 353.

Shrub up to 3 m tall, more commonly a tree 5—15 m tall with a diameter

of 15-30 cm; bark thin, finely fissured and flaking, light brown or grey; inner

bark thin, dull greyish-brown, sapwood hard, brownish-yellow; branches cylindric,

often a bit flattened and shallowly grooved on two faces below the upper node,

sometimes 4-angled on lower internodes, drying brown; blades coriaceous, elliptic,

often broadly so, acute to bluntly acuminate at the tip; narrowed and decurrent

at the base; main veins 3, from the base to the tip, depressed above, raised

(often prominently) below; secondary venation pinnate, very obscure to invisible

on both surfaces; intramarginal nerve obscure to invisible, 1-2 mm from the

margin, arching smoothly towards the tip, (5) 7-12 cm long, (2) 3.5-6.5 cm wide,

drying dark brown to blackish above, brown below: petiole 5-7 mm long, 1 mm

thick; inflorescence glomerulate, 3-5 mm long, glabrous; primary axes 0 -1.5

(exceptionally 2.5) mm long, flattened, usually several in the leaf axils, flowers

numerous; pedicels 1-2 mm long, crowded along the fused secondary axes; bracts

and bracteoles ovate, acute, 0.75 mm long; calyx campanulate, slightly to sharply

constricted above the ovary, often flattened when mature, c. 1 mm high, 1.5 mm

wide, glabrous, truncate with 4 obscure, broad, short, apiculate lobes, faintly

ridged inside, cream to whitish; petals thin, oblong to sub-orbicular, ventral surface

often concave at maturity, broadly rounded at the tip, narrowed or rounded to

a claw at the base, margins slightly thinner, mid-nerve slightly raised near the

tip of the dorsal surface; 1-1.5 mm long, I-1.25 mm wide, white, filaments 1.5—2

mm long; anthers rounded - cuneate to globose, 0.5-0.75 mm long, connective

several times to as long as the locules, without a gland, light yellow; stigma

capitate, minute; style slender c. 3 mm long; ovules 2; fruit globose, 5-7 mm

diameter, pedicels 3-4 mm long, calyx remnant raised and lobed, areolus c. 1.25

mm wide; often appearing didymous with 2 shallow, vertical grooves near the

pedicel extending half way up, two seeds have developed in this situation; exocarp

100 Gardens’ Bulletin, Singapore XX XIII (1980)

green when immature, turning blue and finally black when ripe; drying grey to

blackish, often mealy in texture; pericarp c. 0.5 mm thick.

Figure 24: a. leaf, b. inflorescence, c. calyx and style, d. petal,

e. stamen

The 3 basal veins, calyx with faint internal ridges, and glandless anthers

immediately distinguish this species from all others in the region. M. oligoneurum

Bl. is often confused with various species of Pternandra, especially P. galeata

(Korth.) Ridl., and P. coerulescens Jack. Pternandra in general, has thicker blades

with invisible secondary venation which dry black or very dark brown, larger

flowers, and many seeded fruit. The inflorescence of P. coerulescens, even when

it is very short, always has distinct primary axes, which are not on tubercles,

and sometimes secondary axes, fewer flowers, and a much larger, campanulate-

cyathiform, truncate calyx with an external tessellate pattern.

The epidermis on the branchlets of Pternandra coerulescens Jack is usually

blackish and in most instances peels off. This feature and that of the secondary

venation afford good distinguishing features with vegetative material. Confusion

is also possible with various species of Strychnos L. (Loganiaceae), however a

distinction can immediately be made by its climbing habit.

M. oligoneurum Bl. has, as far as can be determined, the fewest number of

ovules of all the species of Memecylon from the Malay Peninsula. The glandless

anthers are similar to those of M. lilacinum Z. & M. (q.v.), but that species has,

among other major differences, uninerved blades, and narrow petals.

Kedah — Gunong Bougan: Bosewell 18022

Penang — Curtis 1065, 1146, 2220, sn in March 189?; Forest Guard 2220

Perak — Larut: King’s collector 2513; Taiping: Shah & Sidek 1158; Tapah:

Ridley 14060; Ulu Bubong: King’s collector 10280, 10920; sine loc.: Burn-

Murdoch 191, Scortechini 1309

Kelantan — Sungai Lebir: Cockburn 115981; Ulu Sungei Aring: Whitmore 4490;

Selangor — Ginting Simpah: Hume 9309, 9687; Gombak For. Res.: van Balgooy

2125; Sepang: Corner sn on 27 Nov. 1941; Sungai Buloh: Nur 11881; Ulu

Gombak: Hume 9381

Pahang — Jengka For. Res.: Whitmore 0059; Kuala Lipis: Mat 4038; Pulau

Tioman: I.H. Burkill 1103; Rompin: Mahamud 17164; Taman Negara: Lim sn;

sine loc.: Nong 4038

Negri Sembilan — Gunong Angsi: Nur 11654

Malacca — Ayer Keroh: Ridley 10752; Bukit Buiang: Alvins 40

Johore — Labis For. Res.: Ogata 110396

Singapore — Kiah 34670

Further Distribution — Banka Is., Sumatra, Java, Philippines, Borneo (Kali-

mantan) (fide Bakh. f.).

ate ee

Revision of Memecylon L. 101

25. Memecylon paniculatum Jack, Mal. Misc. II (1822) 62; M. costatum Miq,.,

Verh. Kon. Ned. Inst. (1850) 29: M. appenduculatum Bl., Mus. Bot. Lugd.-

Bat. 1:23 (1851) 361; M. nudum BIL., |.c. 361; M. caloneuron Migq., Fl. Ned.

Ind. Suppl. I, Sumatra (1860) 321 (synonyms in Bakh. f., Med. Mus. Bot.

Utrecht 91 (1943) 345 and Rec. Trav. Bot. Neerl. 40 (1943-45) 345): Merrill,

J. Arn. Arbor. 33 (1952) 235; Furtado, Gard. Bull. Sing. 20 (1963) 121);

M. venosum Meir., Phil. J. Sci. 3 (1908) Bot. 154 syn. nov.; M. gigantifolium

Bim, Leaf. Phil. Bot.. 8 (1915)°2762 syn. nov.

Tree 10-30 m tall ,diameter up to 90 cm: bark thin, finely fissured and

flaking, brownish-grey, greyish-white; branchlets distinctly 4-winged or 4-angled,

less commonly 4-angled near the nodes and cylindric below, 3-4 mm_ thick,

glabrous and smooth, drying dark brown, epidermis peeling off leaving a smooth,

cylindric, light brown surface; blades subcoriaceous, glabrous, lanceolate, ovate,

obtuse or shortly and obtusely acuminate (acumen up to | cm long) at the tip,

broadly rounded or more frequently distinctly cordate (indentation 2—3 mm) at

the base; (8) 11-26 cm long, (3.5) 45-14 cm wide, venation pinnate, (8) 10-14

opposite to subopposite pairs, sunken and clearly visible above, prominently

raised below, arising at 45° from the mid-vein, straight or arching, intramarginal

nerve similar in appearance to the veins on both surfaces, 5-8 mm from the

margin, looping; mid-vein sunken above, prominently raised and tapering below;

drying dark brown above, lighter brown below, very brittle; petiole 2-3 mm long,

1—2 mm thick, glabrous; inflorescence cymose, c. 4 cm long, axes generally 4-angled,

minutely furfuraceus, becoming glabrous; primary axes I-—3, usually from leaf axils,

2-3 cm long, 1-2 mm thick; secondary axes 4-10 cm long, tertiary axes generally

glomerulate or up to 3 mm long, 4th axes glomerulate and rarely distinguishable:

pedicels c. 2 mm long, green; calyx funnelform-campanulate, truncate, often with

4 minute cuspus, 1-2 mm long, c. 3 mm wide, glabrous and smooth outside,

prominently ridged inside, green; petals slightly thickened, broadly ovate to sub-

orbicular, acute at the tip, truncate at the base, margins thinner, c. 2 mm long,

2 mm wide, white; anthers “‘J’’ shaped, c. 1 mm long, gland distinct, centrally

located; stigma minute, style c. 3 mm long; ovules c. 12; infructescence axes

generally not elongating, but becoming thicker, prominently angled to somewhat

4-winged, primary axes 3-4 mm thick, tertiary axes generally glomerulate, often

irregularly so due to the unequal lengths of the tertiary axes, up to 3 mm thick,

with numerous distinctly raised bract scars forming a rim about the pedicel scars;

pedicels c. 3 mm long; fruit globose, 6-8 mm diameter, capped by the persistent

calyx remnant; exocarp thin, whitish turning purple when ripe; or ovate, nipple-

like and crowned by the persistent calyx remnant at the tip; 10-13 mm long,

7-9 mm wide; yellow/green when immature, drying tan to blackish.

Figure 25: a. calyx, b. petal, c. & d. stamens

Memecylon beccarianum Cogn. (q.v.) is very close to, if not the same as,

M. paniculatum Jack. M. beccarianum differs from M. paniculatum in having

cylindric branchlets. As far as can be determined from the original description

and type specimens of M. beccarianum (from Borneo), this is the only major

difference. Unfortunately, Jack, in his original description, did not indicate whether

the branchlets are cylindric or angled to winged, however those specimens at

Singapore and Leiden which are considered in this paper as being M. paniculatum

all have angled to winged branchlets which become cylindric with age. Memecylon

beccarianum has been retained as a distinct species in this paper since it is not

certain that it is the same as M. paniculatum or one of its numerous synonyms.

102 Gardens’ Bulletin, Singapore XX XIII (1980)

Elmer 13548, from Mindanao, Philippines, which is the type material of

M. gigantifolium Elm., has the same leaves, inflorescences, and flowers as M.

paniculatum Jack and synonyms. Elmer’s isotype at Leiden has a lower branch

which is cylindric and with rather large leaves. In some specimens of M.

paniculatum seen the leaves are of this size and nearly sessile, however all other

features are distinctly those of M. paniculatum — including older branches which

are cylindric. M. gigantifolium does not differ from M. paniculatum sensu lat.

and should be reduced to a synonym of the latter.

Memecylon venosum Merr., according to Merrill, merely differs from M.

paniculatum in having smaller, more acuminate blades with prominent reticula-

tions on the undersurface; and a shorter inflorescence. A syntype of this specimen

at Leiden (Clemens, s.n., from Camp Keithley, Mindanao, in September 1906)

closely matches specimens annotated by Bakh. f. as M. costatum. ‘There are no

structural details in the inflorescences, flowers, or fruit which distinguish M.

venosum from M. costatum, which has been reduced to a synonym of M. pani-

culatum. ‘Two other collections identified as M. venosum Merr. at Leiden (Elmer

13666, 11752; both from Mindanao) match its syntype and confirm my belief

that M. venosum is the same as M. paniculatum.

Kedah — Gunong Raya, Langkawi: Haniff 15526; Koh Mai For. Res.: Kiah

35198

Penang — Penang Hill: Ng 1112

Perak — Chior For. Res.: Ng 5799; Dindings: Burn-Murdoch 255; Larut: King’s

collector 6945, 8305; Wray 3235; Sungai Seluang, Telok Anson: Shah & Shukor

3450

Kelantan —- Gua Nunik: Henderson 19711; Gunong Stong: Whitmore 12438;

Kuala Sepia: Whitmore 4318; Sungei Lebir: Cockburn 7029

Trengganu — Ulu Brang: Moysey & Kiah 33635

Pahang — Tahan River: Ridley 2237

Malacca — sine loc.: Maingay 813 (1567)

Johore — Kangka Sedili Ketchil: Corner 28603; Mawai: Ngadiman 34734; Panau

Sungei Sedili: Corner 37050; Panti River: Ridley 15399; Sungai Kayu: Kiah

32188

Singapore — Corner sn on 10 Sept. 1934; Sinclair 39658, 40234.

Further Distribution — Thailand (SE), Sumatra, Simeuloee Is., Java,

Karimata Is., Borneo (Kalimantan), Philippines, Celebes, Talaud & Soela Islands,

Moluccas (fide -Craib and Bakh. f.).

26. Memecylon pauciflorum Bl., Mus. Bot. Lugd.-Bat. 1:23 (1851) 356; M.

pulchellum Ridl., J. Str. Br. Roy. As. Soc. 61 (1912) 52, syn. in Ridl., FI.

Mal. Pen. I (1922) 816; M. pauciflorum Bl. var. obovatum Furtado in scheda

Herb. Sing.

var. pauciflorum

Revision of Memecylon L. 103

Shrub 2-3 m or a tree up to 8 m tall; bark thin, brown, longitudinally ridged:

branches somewhat flattened and deeply grooved on 2 sides, each groove bordered

by 2 sharp ridges, or sharply 4-angled, often with the grooves widened and

appearing winged near the upper nodes, | mm thick, glabrous; blades coriaceous,

ovate, bluntly acuminate at the tip, narrowed and not or only slightly decurrent

on the petiole, venation extremely obscure or more commonly invisible on both

surfaces, mid-nerve sunken above, raised and tapering below, 3-6 cm long, 1-3

cm wide, grey-green or glossy dull green above, yellow-green or olive-green below,

drying blackish or very dark brown above, brown below; peticle 1-3 mm long,

1 mm thick, glabrous; inflorescence from leaf axils, 5-7 mm long, umbellate or

a very contracted cyme which appears umbellate, primary axes | or 2 from each

axil, 4-angled, 1.5-4 mm long, often extended beyond the first node for up to

1 mm; secondary axes indistinct and reduced to tubercles c. 0.5 mm long; pedicels

2-2.5 mm long; flowers few (c. 5); bracts and bracteoles lanceolate, acuminate

at the tip, c. 0.25 mm long, cauducous; calyx campanulate, constricted above the

ovary, truncate with 4 distinct cusps, smooth outside, ridged internally, white:

corolla buds dome shaped, tips acuminate and twisted; petals thin, broadly ovate,

caudate-acuminate at the tip, truncate to shortly clawed at the base, midvein

slightly thickened, but not raised, c. 2 mm long, 1.5—2 mm wide, blue to purplish;

filaments blueish, anthers yellowish with a blue spur, gland distinct; fruit globose,

5-8 mm diameter, calyx remnant raised, areolus c. 1—-1.5 mm wide, exocarp

smooth and sometimes minutely warty, c. 1/5 mm thick, green turning pale

yellowish and tinged with pink, drying black or greenish.

Figure 26: a. calyx and corolla bud, b. petal. c. stamens

In many respects this species closely resembles M. lilacinum Z. & M. (q.v.),

however that species has a greater variation in leaf size, more flowers in each

inflorescence, and anthers without a gland. The fruit of M. lilacinum also is

larger, depressed globose, and has a thick, gritty pericarp. The anthers and

fruit provide the most reliable distinguishing features for the two species.

M. edule Roxb. var. edule (q.v.) very closely resembles M. pauciflorum,

however the former has a truncate calyx with or without 4 very small bumps

and a generally more complex inflorescence.

M. pulchellum Ridl. has more ovate ~-blades, slightly larger and narrower

calyx lobes, and shortly clawed petals. These features fall well within the range

of variation for M. pauciflorum, and, therefore, agreement is made with Ridley

in reducing M. pulchellum to a synonym of M. pauciflorum.

Memecylon pauciflorum Bl. var. obovatum Furtado, as noted on _ several

sheets in the Singapore collection (Henderson sn, from Selat Panchor, Langkawi

on 22 Nov. 1934; Corner sn, from Pulau Dayang Bunting, Langkawi on 13 Nov.

1941, etc.) differs from the typical variety in having obovate blades. Some

blades on, these collections are identical to those of typical M. pauciflorum, thus

Furtado’s unpublished variety is not worthy of further attention.

Kedah — Pulau Butong: Ridley 15828 (lectotype M. pulchellum Ridl.); Pulau

Dayang Bunting: Alphonso & Samsuri 179; Corner 37853, sn on 13 Nov. 1941;

Pulau Jerkam: Corner sn on 17 Nov. 1941; Pulau Langkawi: Haniff & Nur

7547, Holttum 17431; Sulau Sarang: Ridley 15829 (syntype M. pulchellum Ridl.);

Selat Panchor. Langkawi: Henderson 29090, sn on 22 Nov. 1934; Telok Udang:

Haniff 1071

104 Gardens’ Bulletin, Singapore XX XIII (1980)

Penang — Curtis 3434, King’s collector 1684, Nauen sn on Penang Hill in April

1940

Kelantan — Gua Musang: UNESCO 326; Gua Teja: Henderson 29680; Gunong

Brong Bertam: Shah & Ali 2886

Trengganu — S. Banum, Kemaman: Corner 25829

Selangor — Bukit Takun: Nur 34371; Gua Batu: Ridley sn in July 1897

Perak — Gunong Ginting: Ahmad & Sidek 579

Johore — Gunung Belumut: Holttum 16779; Kanga Sedili Ketchil: Corner 28565

Singapore — Lobb 296, Ridley 14178

Further Distribution — Burma, Laos, Thailand, Timor (type), New Guinea

(fide Craib and Bakh. f.).

26a. Memecylon pauciflorum Bl. var. brevifolium Craib, Fl. Siam. Enum. I:4

(Tosi 712.

Shrub 50-100 cm tail, branchlets flattened, deeply grooved on two sides,

each groove flanked by two sharp ridges; grooves often wider, thus the branchlets

appear somewhat 4-angled; older branches becoming cylindric, c. 1 mm _ thick;

blades coriaceous, ovate, obtuse at the tip, sometimes shallowly emarginate; acute

and shortly decurrent at the base; venation invisible on both surfaces, midvein

sunken above, slightly raised near the base and tapering to the tip below, 1.2-1.7

cm long, 6-8 (ex descr. 10) mm wide, drying dark brown above, brown below;

petiole 2 mm long, c. 0.5 mm thick; inflorescence, flowers, and fruit unknown.

Figure 35: isotype.

Craib notes that there is only one known collection of this taxon, and that

the specimen he examined was in fruit. The two specimens at Singapore are

vegetative and apparently only differ from the typical variety by the very small

leaves.

Thailand

Puket: small islands near Pulau Panji: Haniff & Nur 4070 (type).

27. Memecylon pubescens (Cl.) King, J. As. Soc. Bengal 69, II:1 (1900) 74

(Mat. Fl. Mal. Pen. HI, 482); M. grande Retz. var. pubescens Cl. in Hk.

f., Fl. Brit. Ind. If (1879) 558.

Tree up to 30 m tall, diameter up to 75 (exceptionally 2 m) cm; trunk

massive, twisted or straight, fluted to about 3 m high; crown with massive,

spreading limbs; bark slightly fissured, in thin, brittle, and elongate scales, dull

brown to grey-green; inner bark black-brown; sap wood yellow; branchlets

slightly flattened and grooved below the upper node, otherwise cylindric; 1.5 mm

diameter, smooth and glabrous, blades coriaceous, elliptic, acute or shortly acumi-

nate at the tip, acute and decurrent at the base; venation pinnate, 8-12 pairs of

nerves, sunken above; raised below; intramarginal nerve c. 3 mm from the

Revision of Memecylon L. 105

margin, looping; midnerve sunken above, prominently raised and tapering below;

8-15 cm long, 4-6.5 cm wide; dark, glossy green above, dull green below; drying

dark and often glossy brown above, dull brown below, entirely glabrous; petiole

8-10 mm long, 1.5—2 mm thick, glabrous; inflorescence from leaf axils or just

behind the leaves from leafless nodes, composed of an open panicle of cymes

or umbellate, 2.5-7 cm long; axes flattened, grooved on 2 faces and frequently

angled; minutely rufus hispid throughout; primary axes 1.5-5 cm long, usually

solitary, less frequently paired, with 1 or 2 nodes, 1.5-2 mm wide; secondary

axes 3-20 mm (exceptionally 25-35 mm) long, tertiary axes not developed or

up to 2 mm long, pedicels glomerulate, glabrous, c. 1 mm long; flowers numerous;

calyx campanulate, truncate, often with 4 minute points, widened above the

ovary, smooth outside, ridged inside, 1.25-1.5 mm long, 1.5 mm wide, petals

thickened, oblong to broadly ovate, obtuse with a mucro at the tip, truncate at

the base; mid-nerve slightly raised, but not keeled, dorsally, margins thinner,

c. 1.25 mm long, c. 1-1.5 mm wide; anthers “J” to ““C’’-shaped, gland prominent,

centrally located, connective thickened. c. 1 mm long: stigma minute, style c.

2 mm long; ovules 10-12; infructescence axes often glabrescent, pedicels 2 mm

long, fruit globose, 10-12 mm diameter, calyx remnant slightly raised, areolus

c. 2 mm wide; exocarp minutely muricate, pericarp c. 1 mm thick, gritty, green

when immature, purple to dark blue when ripe; drying black.

Figure 27: a. branch and inflorescence, b. calyx, c. petal, d. stamen,

e. fruit

The axes in M. hAullettii King are muricate-papillose and the inflorescence

is a compound umbel. M. beccarianum Cogn. has pubescent axes, however the

leaves in that species are larger, more rounded at the base, and the calyx has

4 minute points. Purseglove 5505 and Everett 13882, both from Johore, differ

from most other specimens of M. pubescens in having thicker blades. The

secondary veins are impressed above and are very faint to invisible below. The

finer venation is invisible. The inflorescence, flowers, and fruit of these collec-

tions appear to be the same as more typical specimens of M. pubescens.

Kedah — Koh Mai Res.: Kiah 35188; Ulu Muda For. Res.: Chan 6693, 6791,

6802

Perak — King’s collector 6089; Kinta Hills For. Res.: Zainuddin 99778; Slim

Hills For. Res.: Whitmore 0746

Trengganu — Ulu Brang: Whitmore 12539

Selangor — Semangko For. Res.: Whitmore 12554; Sungei Bulok For. Res.:

Kiai 8295, Sow & Tachon 16858

Pahang — Kemasul For. Res.: Hamid 10619; Pekan: I.H. Burkill & Haniff

17248, 17250; Temerloh: Pawanchee 13760

Negri Sembilan — Jelebu For. Res.: Suppiah 11353

Malacca — sine. loc.: Griffith 2336 (type)

Johore — Gunong Pulai: Purseglove 5505; Ulu Sungei Sedili Besar: Everett

13882

Singapore — Henderson 35911, 36437, Ridley 10390

Further Distribution — peninsular Thailand (fide Craib).

106 Gardens’ Bulletin, Singapore XX XIII (1980)

28. Memecylon wallichii Ridl., J. Str. Br. Roy. As. Soc. 79 (1918) 74; M.

heteropleurum Bl. var. olivaceum King (“‘olivacea’’), J. As. Soc. Bengal

69, Il: 1 (1900) 78 (Mat. Fl. Mal. Pen. III, 486), in part; M. longifolium

Ridl., Lc. 72 nom. illeg. non Cogn.; M. depressum Benth. ex Triana, Trans.

Linn. Soc. 28 (1871) 158; M. amplexicaule Roxb. sensu Clarke in Hk. f.,

Fi» Brit: ind.) 1 ((1879)-359:

Shrub up to 4 m tall or a tree up to 5 mm with a diameter of 15 cm; bark

greyish, finely fissured, thin; branchlets 4-angled, less commonly 4-winged, espe-

cially near the upper nodes, becoming cylindric with age, smooth, glabrous, light

brown-tan when dry, c. 2 mm thick; blades coriaceous, glabrous, lanceolate,

elliptic, to ovate; caudate-acuminate (acumen up to 2.5 cm long) at the tip;

broadly rounded to cordate at the base; venation pinnate, 14-18 pairs of nerves,

lower veins running parallel to the midvein for 1-2 mm, then diverging at 45°

towards the margin, straight or slightly arching, sunken and rather faint above,

prominently or less commonly slightly raised below; intramarginal nerve of similar

appearance, 6-9 mm from the margin, broadly looping; midvein sunken above,

prominently raised and tapering below; (12) 14-25 cm long, 3.5-10 cm wide;

drying olive-brown to dark brown above, brown or greenish below; petiole 1-2

mm long, 1.5-2.5 mm thick, glabrous; inflorescence cymose, 1—1.5 cm _ long,

glabrous, usually with about 10 flowers; primary axes 2 or 3 from leaf (oiten

terminal) axils, 2-8 mm long, 1-2 mm thick, 4-angled; secondary axes 1-2 mm

long, often clustered; tertiary axes generally glomerulate, less frequently up to

0.75 mm long; pedicels 2—2.5 mm long; bracts ovate, acute, thickened, c. 0.5—

0.75 mm long, fimbriate at the base inside; calyx campanulate, truncate, cracking

irregularly about the margin; smooth outside, ridged internally, glabrous, 3-4 mm

long, 3-4 mm wide, red; petals orbicular in outline, acute at the tip, truncate

at the base, thickened with thinner margins, 2.5-3 mm long, 3-4 mm wide; keel

slightly developed dorsally, white; filaments c. 2 mm long; anthers linear, locules

extending all along one side of the anther and curving around the distal tip,

connective smaller, with a distinct gland near the filament, 1.5—2 mm long, white;

stigma minute, style c. 4 mm long; ovules c. 8; fruit globose, 10-15 mm diameter,

smooth, capped by the raised calyx remnant, areolus c. 4 mm wide; pericarp c.

0.5 mm thick, pink to bright red when immature, dark purplish-red, then blackish

when ripe.

Figure 28: a. bracts and calyx, b. bud petal, c. stamens

The 4-angled to 4-winged branchlets and leaves are similar to M. paniculatum

Jack (q.v.), but that species has longer primary and secondary axes (total length

up to 4 cm), entirely different anthers, and smaller (globose or ovate) fruit.

M. excelsum BI. is also similar, but the inflorescence is behind the leaves, has

different anthers, and ovate fruit. M. corticosum Ridl. is nearly the same as

M. wallichii vegetatively, however that species has narrower blades, ramiflorus

inflorescences, and different anthers. The anthers of M. wallichii are unique

and are probably the most reliable structural feature that can be used to dis-

tinguish it from all related species.

Agreement is made with Furtado (Gard. Bull. Sing. 20 (1963) 122) in his

reduction of M. longifolium Ridl. to a synonym of M. wallichii Ridl. The

blades, inflorescence, and flowers (especially the anthers) of the holotype of M.

longifolium (Ridley 9475, from Perak) match those of Curtis 1294 — which is

a syntype of M. wallichii.

Revision of Memecylon L. 107

Memecylon longifolium Ridl. is a later homonym of a completely different

species described by Cogniaux (Monogr. Phan. 7 (1891) 1150) from New Guinea

and Sarawak. Ridley’s epithet, even though it has publication priority over

M. wallichii, cannot be used since it violates a nomenclatural rule.

The syntypes of M. heteropleurum BI. var. olivaceum King include specimens

which Furtado reduced to M. maingayi Cl. (Wray 1310) (now a synonym of

M. excelsum Bl.) and M. wallichii (King’s coll. 500). His observations are correct,

however I have not seen the other syntype of var. olivaceum (King’s coll. 10872)

to determine its proper identity.

Penang — 1.H. Burkill 1532, 2685, 3337, 6146, 6579; Curtis 457 (syntype); 965

from Experimental Nursery (syntype), 965 from Penang Hill, = 965 at Paya

Trobong in Aug. 1892, sn from Paya Trobury in Aug. 1892; Ridley sn in Mar.

1915; Wallich 4101, 4101C (lectotype)

Perak — Goping: King’s collector 500 (lectotype M. heteropleurum Bl. var.

olivaceum King); Gunong Bujang Melaka: Shah & Shukor 3406; Gunong Kele-

dang: Ridley 9628; Kali, Kuala Kangsar: Haniff 14949; Grik: Chelliah 98616;

Larut: King’s collector 2778 (syntype M. heteropleurum BI. var. olivaceum King);

3058 (syntype); Lumut: Curtis sn in Dec. 1902; Ridley 9475 (type M. longifolium

Ridl.); Piah For. Res.: Jaamat 39247; Taiping: Ridley 14687; Tapak: Ridley

14102; Waterfall: Wray 1964, 2326, 3278; Waterloo: Curtis 1294 (syntype); sine

loc.: Scortechini 231 (syntype), sn

Kelantan — Bertam: UNESCO 95

Malacca — Bujoing: Ridley 9526

Insufficiently Known Taxa

1. M. constrictum Craib, Kew Bull. 1930, 324. (q.v. M. minutiflorum Miq.).

Figure 29: a. calyx and corolla bud, b. petal, c. stamen.

2. M. lancifolium Ridl. Ml. Malay Pen. V suppl. (1925) 311.

Small tree, glabrous; blades coriaceous, lanceolate, acuminate at the tip

(acumen c. 1 cm long), narrowed and slightly decurrent at the base, midrib

sunken above, raised below; secondary venation pinnate with c. 15 pairs of nerves,

obscure to invisible on both surfaces; 15 cm long, 4 cm wide; drying olive-

greenish above, brown below; petiole c. 1 cm long, 2 mm thick; inflorescence

cymose, in pairs from leaf axils, axes slightly 4-angled to flattened, drying black;

primary axes c. 7 mm long, secondary axes 3, 7-8 mm long; tertiary up to 4 mm.

4th not developed or up to 1.25 mm long, pedicels c. 2 mm long; bracts and

bracteoles ovate, acute, c. 0.5 mm long; calyx campanulate, widened above the

ovary, smooth and glabrous outside, c. 2 mm long, c. 3 mm wide, margin with

4 broadly undulate lobes; petals broadly ovate, acute at the tip, truncate at the

base, very thick with thinner margins, in bud c. 1.5 mm long, 2 mm wide, blue:

anthers in bud ‘‘C’’-shaped, c. 1.25 mm long, gland distinct, centrally located;

fruit unknown.

Type: Ridley 3840, Sungei Morai, Singapore

108 Gardens’ Bulletin, Singapore XX XIII (1980)

The holotype of this species at Kew consists of one detached leaf and a

pile of broken inflorescences and flowers. Branchlets are not on the sheet. There

is not much to see and from the description I have been unable to study this

species in further detail. It is noted on this specimen that there is a duplicate

in the Singapore collection, however I could not find it there. The leaf does

not look like that of any species of Memecylon that I am familar with. There-

fore, since this species cannot be adequately reconstructed it has been included

in the section for inadequately known taxa.

Figure 30:. a. calyx, +b. petal, oc: stamen

3. M. pauciflorum BI. var. brevifolium Craib, Fl. Siam. Enum. 1:4 (1931) 712;

q.v. discussion under M. pauciflorum Bl. (26a.) Figure 35.

GENERAL CONCLUSIONS

In this treatment of Memecylon from the Malay Peninsula twenty eight

species and three variations (not including three imperfectly known taxa) are

discussed and twenty two taxa have been reduced to new synonyms. No new

taxa are described here, although there are several collections which I have not

been able to identify and these may be undescribed taxa. These specimens

will have to remain undetermined until more material is studied. Memecylon

dichotomum (Cl.) King var. rotundatum (Craib) Maxw. is the only new com-

bination included in this paper. Although specimens of M. beccarianum Cogn.

from the Malay Peninsula have not been seen, this Bornean species is presently

included in this treatment since it is likely that it occurs in the region.

While examining a large number of collections from the Malay Peninsula

and the Malay Islands at Singapore, Leiden, and Kew it became apparent that

several widely distributed species and varieties show a broad range of variation.

As a result a number of new synonyms which were formerly described as distinct

taxa from the Malay Islands are included here. The reduction of three and

five species to synonyms of M. oleifolium Bl. and M. excelsum BI., respectively;

appears, for this reason, to be justified.

Among King’s five new species and two new varieties of Memecylon from

the Malay Peninsula, one species and two varities have been reduced to synonyms

of other species. Among the remaining taxa which King included in his paper,

fourteen species and one variety have been accepted in this revision without

nomenclatural changes. In the meantime the present revision includes four reduc-

tion made by other authors on King’s work, plus two misinterpretations have

been found: M. heteropleurum Bl. and M. amplexicaule Roxb. — the former

being M. megacarpum Furtado, and the latter a mixture of M. amplexicaule

Roxb. and M. wallichii Ridl.

Among the thirty one species and three varieties of Memecylon from the

Malay Peninsula discussed by Ridley twenty species and one variety are included

as distinct taxa in this revision. The other thirteen taxa; including M. tenuifolium

Ridl., which was described as a new species; as synonyms of other species. In

general, Ridley followed King’s treatment quite closely, but did make some

changes which I have accepted: reducing M. microstomum Cl. to M. amplexicaule

Roxb.. and considering M. amabile Bedd. var. malaccensis Cl. as M. malaccense

(Cl.) Ridl. (King regarded this variety as a doubtful taxon). Ridley also followed

Revision of Memecylon L. 109

King in considering M. acuminatum Sm. var. flavescens Cl. as a synonym of

M. minutiflorum Mig. In this present work var. flavescens has been maintained

as a distinct, but poorly known taxon. The type collection of M. laxiflorum

Ridl., described in vol. V of his flora, is incomplete and I have been unable

to decide on its exact taxonomic status.

Craib’s three new species and two new varieties of Memecylon from peninsular

Thailand have been reconsidered in this revision and now includes only one

new species and one new variety — both of which are incompletely known and

perhaps synonyms of other taxa.

Bakhuizen van den Brink’s extensive work on Memecylon from the Malay

Archipelago includes twenty five taxa which are also found in the Malay Peninsula,

twelve of which are recognized in the present revision as distinct taxa and the

other thirteen as synonyms. At least four of his new species of Memecylon are,

in this paper, considered identical to previously described taxa which have been

recorded from the Malay Peninsula viz. M. depokkense Bakh. f. (= M. fruticosum

King), M. multiflorum Bakh. f. (= M. campanulatum Cl.), M. steenisii Bakh. f.

(= M. cantleyi Ridl.), and M. rhodophyllum Bakh. f. (= M. edule Roxb. var.

ovatum (Sm.) Cl. M. globosum Bakh. f. is the only new record of Memecylon

for the Malay Peninsula. His paper contains a tremendous amount of informa-

tion which also clarifies many taxonomic problems and has greatly improved our

knowledge of the family.

In regards to Corner’s treatment of Memecylon my conclusions differ from

his in that M. heteropleurum BI. is replaced with M. excelsum Bl., and M. ovatum

Sm. is reduced to M. edule Roxb. var. ovatum (Sm.) Cl.

Furtado’s paper has been thoroughly reviewed. The specimens listed under

M. campanulatum Cl. are all M. minutiflorum Mig. Furtado also discusses

M. hepaticum Bl. and cites several specimens from Johore as examples. The

type specimen of M. hepaticum Bl. (Blume s.n., from Sumatra) lacks inflore-

scences, flowers, and mature fruit; and the original description is inadequate for

detailed analysis. The specimen (at Leiden) does not resemble the collections

Furtado cites, and furthermore these specimens all have inflorescences and flowers

(especially the anthers) identical to those of M. garcinioides Bl. The leaves,

however, are longer and narrower than most collections of M. garcinioides BI.

from the Malay Peninsula and Malay Islands. Therefore, M. hepaticum BI. is

now considered to be an imperfectly known species which does not seem to

occur in the Malay Peninsula.

Furtado correctly recognized the syntypes of M. heteropleurum Bl. var.

olivaceum King as belonging to two other taxa: M. maingayi Cl. and M. wallichii

Ridl., however he did not reduce M. maingayi Cl. to M. excelsum Bl. as Blume’s

holotype (at Leiden) was not available to him. M. longifolium Ridl. was also

correctly reduced to a synonym of M. wallichii Ridl. A review of the salient

morphological characteristics of each taxon is outlined in Table 1. These features,

in several instances, can be used to distinguish individual taxa, while in others

several closely related taxa can be separated — their individual identities requiring

more detailed analyses.

Gardens’ Bulletin, Singapore XX XIII (1980)

110

‘Taxon

M. acuminatum

var. acuminatum

acuminatum var.

flavescens

amplexicaule

beccarianum

caeruleum

campanulatum

cantleyi

cinereum

corticosum

dichotomum vat.

dichotomum

dichotomum var.

rotundatum

TABLE 1. REVIEW OF SALIENT MORPHOLOGICAL CHARACTERS

Branchlets

cylindric

flattened-grooved

grooved-angled

flattened-grooved

winged

angled-winged

Calyx

4-lobed

truncate

truncate-undulate

truncate-apiculate

truncate

lobed

truncate-apiculate

truncate

Anther

Shape

(8!

triangular-

cuneate

Fruit

Shape

globose

ovoid

globose

ovoid

globose

ovoid

globose

Distinguishing Features

acuminate blades, long primary

axiS

poorly known

sessile, thick blades; glomerulate

inflorescence

Bornean species

thick blades, fruit

tall tree, thick blades, short

inflorescence

flattened calyx, fruit

acuminate petals, glandless

anthers, short inflorescence

winged branchlets; ramiflorus

inflorescence

bracts, inflorescence

longer inflorescence

WA sosn1 ‘OdIe]

111

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SJO[YOURIQ “SOABO] IOSIV]

sjoyyouvig ‘SoAvo] JO]]VUUS

Revision of Memecylon L.

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Gardens’ Bulletin, Singapore XX XIII (1980)

112

Taxon

minutiflorum

oleifolium |

oligoneurum

paniculatum

pauciflorum

pubescens

wallichii

Branchlets

flattened-grooved

cylindric

flattened-grooved

angled-winged

grooved-angled

cylindric-grooved

angled-winged

Calyx

cusped

truncate

truncate-apiculate

cusped

truncate-apiculate

truncate-cusped

truncate

©..1t0-

triangular-

cuneate

linear

Fruit

Shape

globose

ovoid

globose

globose,

ovoid

globose

Distinguishing Features

papillose calyx constricted above

the ovary

complex inflorescence, branchlets

drying tan, fruit

tri-nerved blades

blade venation, large

inflorescence

small: leaves, inflorescence

blade venation; complex and

pubescent inflorescence

axillary inflorescence and unique

anthers

Revision of Memecylon L. 113

INDEX TO COLLECTIONS FROM THE MALAY PENINSULA

Ahmad, 273 (9), 283 (8), 376 (23), 390 (16), 1004 (10a), 1012 (10a), 1015 (4),

1221 (6), 1416 (19), 1474 (21), 1475 (19), 2477 (19), 5099 (6)

Ahmad & Shukor 421 (9), 447 (22), 450 (11), 450a (19), 484 (4), 521 (23),

ae tZy, 525 -(19)

Ahmad & Sidek 579 (26)

Ahmat 2477 (19)

Allen & Kadim 494 (9)

Alphonso & Samsuri 179 (26)

Alphonso, Sanusi, Sidek 201 (16)

Alvins 21 (22), 24 (2), 40 (24), 69 (19), 119 (9), 129 (22), 163 (19), 245 (11),

280 (4), 430 (11), 659 (23), 765 (1), 1212 (14), 1925 (9), 1970 (23), 2035 (23),

2217 (9), 2296 (19), 2302 (22); sn G. Kalan, Malacca (4); sn Pang Kalan

Miniak, Malacca (4); sn Sungei Ujong (23): sn, s. loc. etc. (22); sn Malacca

Pindah, Malacca (10a); sn Sungei Ujong, Malacca (9); sn Malacca (10); sn,

Seremban, Negri Sembilan (6)

Annandale 1662(4), 1696 (10)

Awang-Lela 4578 (6)

van Baloogy 2125 (24), 2319 (10a), 2349 (10), 2479 (9), 2584 (14)

Barnes sn Kluang Terbang, Pahang (10a)

Batten-Pooll sn Langkawi, Kedah (19)

Best 13852 (7)

Bidin 15442 (10)

Bolinan 6787 (4)

Boswell 18022 (24)

Bray 11511 (2)

Brelkerata sn Kuala Trengganu (10)

Brown 52920 (9)

H.M. Burkill 3189 (14), sn Sungei Buloh, Selangor (19)

H.M. Burkill & Shah 253 (4), 970 (10a), 993 (10a), 1048 (19), 2508 (10), 2526

(10)

LH. Burkill 903 (19), 1103 (24), 1532 (28), 2655 (la), 2685 (28), 3046 (22),

3337 (28), 4477 (10), 4970 (22), 6146 (28), 6387 (9), 6579 (28), 6893 (19),

8888 (7); sn, Pulau Tioman (4)

114 Gardens’ Bulletin, Singapore XX XIII (1980)

I.H. Burkill & Haniff 15829 (21), 16102 (22), 16135 (21), 16219 (6), 16249 (9)

17091 (6), 17248 (27), 17250 (27), 17513 (10a)

1.H. Burkill & Holttum 7869 (21), 8697 (9), 8937 (10)

Burn-Murdoch 28 (22), 164 (9), 162 (21), 191 (24), 255 (25)

Cantley 148 (6); sn, Singapore (2)

(J.C.) Carrick 1639 (2569) (6)

C.F. 2477 (19), 6410 (19)

Chan 6693 (27), 6764 (19), 6791 (27), 6800 (10a), 6802 (27), 11239 (21), 13118

(19), 13120 (11), 13128. @), 13133 (2), 17539 (1); 17550 (2) Si fe2ee

17681 (4), 19815 (24), 19884 (4)

Chelliah 6945 (22), 9801 (17), 98204 (9), 98616 (28), 104387 (6)

Chew 138 (4), 207 (10a), 726 (19), 728 (23), 1463 (4), 41016 (10a)

Chew & Noor 249 (12)

Cockburn 7029 (25), 7103 (9), 7115 (21), 7414 (12), 7477 (2), 7646 (10), 8105 (5),

8288 (12), 10509 (22), 10839 (23), 115981 (24)

Corner 25765 (4), 25829 (26), 25896 (19), 28562 (19), 28565 (26), 28603 (25),

28683 (21), 28702 (11), 28733 (22), 29016 (21), 29021 (22), 29044 (22), 29234

(16), 29245 (11), 29251 (14), 29450 (22), 29459 (23), 29481 (14), 29760 (10),

29846 (10a), 29846 (4) in part; 29953 (2), 29984 (16), 30003 (2), 30017 (23),

30188 (2), 30190 (12), 30312 (9), 30327 (9), 20329 (13), 30336 (22), 30479

(22), 30781 (2), 30882 (14), 30976 (14), 31653 (10), 31654 (4), 31656 (10a),

31942 (22), 31961 (14), 32246 (11), 32476 (14), 32491 (14), 32504 (11),

32759 (11), 34657 (23), 36290 (21), 36295 (1), 36980 (6), 37050 (25), 37853

(26); sn, Gunong Panti, Johore (23); sn, Mawai-Jemalang Rd., Johore (14);

sn, Pelepah Kiri, Johore (16); sn, Sungai Kayu Ara, Johore (8 & 10 Feb.

1935 (23); sn, Sungai Kayu Ara, Johore 10 Feb. 1935 (22); sn, Sungai

Pelepoh Kiri, Johore (16); sn, Sungai Sedili, Johore (19); sn, Gunong Baling,

Kedah (9); sn, Pulau Dayang Bunting, Kedah (1), (26); sn, Pulau Jerkam,

Kedah, 7 Nov. 1941 (10a); sn, Pulau Jerkam, Kedah 17 Nov. 1941 (26);

sn, Pulau Selang, Kedah, 22 Nov. 1941 (4), (10a); sn, Tumpat, Kelantan

(10); sn, Bukit Kajang, Kemaman (11); sn, Sungai Nipa, Kemaman (23);

sn, Pulau Chibeh, Pahang (10a); sn, Pulau Duchong, Pahang (10); sn, Pulau

Tioman, Pahang (4); sn, Sepang, Selangor (24); sn, Singapore 10 Sept. 1934

(25); sn, Pulau Merambong, Singapore (10a); sn, MacRitchie Reservoir,

7 May 1937 (19)

Craddock sn, Ulu Kuantan, Pahang (23)

Cubitt 867 (22), 5109 (7), 6522 (9)

Cuming 2322 (4)

Curtis 54, Tanjong Bunga- Oct. 1884 (4); 54, Aug. 1889 (4); 54 (= 1507) (4);

100 garden, Penang (19); 100 Gov.’t Hill, Penang (19); = 100 (19), 457 (28);

723 Oct. 1885 (10a); 723 Muka Head, March 1886 (10a); 723 May 1893

Revision of Memecylon L. 115

(10a)27235 collioin .1893*(10a);5= 723 (10a); 766)(2); 814 (21), ‘815 -Ca),

816 (la), 816, Reservoir Line, West Hill, Penang (22); 965 experimental

nursery (28); 965 Penang Hill (28), = 965 (28), 1065 (24), 1146 (24), 1294

(2a 1295 (a), 1503: (19); 1507: (4)y01773*- (13), )2219; (19),, 2220 (24), 2338

(9), 2627 (10), 3434 (26), 3592 (2), 3596 (19), 3691 (10), sn, Langkawi, Kedah

(10); sn, Penang (2); sn, Paya Troburg, Penang (28); sn, Penara Bukit,

Penang (17); sn, Waterfall, June 1893 (19); sn, Penang, March 189? (24);

sn, Batu Ferengy, May 1901 (10a); sn, Waterfall, June 1901 (19); sn, Penang

(19); sn, Lumut, Perak, Dec. 1902 (28)

Derry 127 (22), 582 (21), 1028 (10), 1041 (1), 1184 (5), 3199 (23), 1240 (6)

Evans 13187 (22), 13188 (22), 13190 (14)

Evans & Gorden 95 (17)

Everett 13664 (22), 13674 (22), 13681 (22), 13684 (2), 13747 (2), 13768 (19),

13781 (21), 13812 (2), 13821 (sp.), 13837 (14), 13882 (27), 13922 (12), 13994

(2), 14076 (11), 14098 (2), 14169 (21), 14263 (17), 14458 (21), 14662 (22),

104909A (2)

Forest Guard 2220 (24); sn, Penang (10a)

Foxworthy 237 (10a), 1194 (1), 1736 (4)

Furtado 34854 (4)

Gadoh (anak Umbai) 959 (9), 1137 (19), 1295 (19), 1502 (22), 1661 (9), 1781 (23)

Goh 15701 (19)

Goodenough 1522 (4), 1767 (19), 10601 (22), 10602 (7), 10812 (14)

Griffith 2322 (4), 2324 (9), 2325 (5), 2325/1 (1), 2325/2 (la), 2326 (10), 2327

10); 2328.19), 2336. (27)

Guard sn, Penang (10a)

Hamid 965 (21), 1600 (14), 2959 (4), 9908 (7), 10619 (27), 10678 (19), 11615 (2)

Haniff 1071 (26), 1072 (10), 14949 (28), 14969 (9a), 15526 (25), 15528 (22),

21047 (2), 21048 (21)

Haniff & Nur 2732 (M. constrictum Craib), 4070 (26a), 4245 (10a), 7072 (10),

7479 (4), 7547 (26)

Haniff & Sallah 10475 (2)

Hassan & Kadim 92 (22)

~ Henderson 10395 (19), 10514 (7), 10565 (9), 10671 (6), 10704 (7), 10886 (14),

11515 (2). 18538 (4), 19581 (8), 19595 (9), 19711 (25), 21651 (23), 21772 (9),

22211 (22), 22403 (9), 22482 (11), 22803 (10a), 22881 (4), 23798 (9a), 24514

(6), 25064 (9), 29090 (26), 29125 (4), 29680 (26), 35447 (18), 35783 (10),

116 Gardens’ Bulletin, Singapore XX XIII (1980)

35911 (27), 35916 (21), 36357 (2), 36437 (27); sn, Selat Panchor, Kedah

(26); sn, Bukit Batu Papan, Kelantan (8); sn, Robinson Falls, Pahang (21)

Henderson & Nur 11142 (9), 11212 (2)

Hervey 2020 (4)

Holttum 9279 (9), 9436 (16), 9570 (19), 9574 (9), 9641 (22), 9669 (19), 10779

(1), 10854 (16), 15158 (10), 15215 (10a), 15216 (4), 16779 (26), 17431 (26),

18095 (23), 20535 (9), 24751 (6)

Hou 662 (21), 763 (10), 765 (5), 798 (17), 814 (17)

Hullett 390 (19); sn, Singapore (10)

Hume 8481 (23), 8910 (23), 9011 (7), 9159 (23), 9309 (24), 9381 (24), 9687 (24)

Ismail 95021 (2), 97802 (2), 98912 (21), 104273 (11), 104897 (2), 109424 (11)

Jaamat 10266 (21), 13908 (9), 28188 (21), 39247 (28)

Jumali 943 (19), 4100 (11), 4496 (973) (11), 6711 (4257) (9)

Jumali & Heaslett 3056 (23), 4008 (sp.)

Kadim & Mahmood 27 (9), 50 (6)

Kadim & Noor 314 (14), 416 (11), 580 (11), 660 (4)

Kadim & Nur 27 (9)

Kalong 22411 (9)

Kassim 261 (22), 0748 (13), 1489 (4)

Keith 492 (8)

Keng et al. 30 (19), 126 (10), 6312 (14)

Kepong no. 85084 (17)

Kerr 14169 (23), 15408 (6), 15766 (9a), 21762 (4)

Kiah 24400 (22), 31740 (2), 31902 (9), 31961 (14), 31981 (11), 31994 (23), 32057

(21), 32079 (23), 32170 (6), 32188 (25), 32311 (11), 34670 (24), 35086 (22),

35153 (21), 35182 (2), 35188 (27), 35198 (25), 35280 (4), 35281 (19), 35283

(10a), 35412 (10a), 35959 (21), 35978 (22), 36152 (19); sn, Sungai Teku,

Pahang (21)

Kiah & Strugnell 24023 (9)

Kiai 8295 (27) .

Revision of Memecylon L. 117

King’s collector 500 (28), 509 (11), 1110 (21), 1457 (19), 1684 (26), 1687 (4),

1984 (14), 2513 (24), 2778 (28), 2938 (14), 2971 (13), 3058 (28), 3143 (7),

B2o9 47), 3205 (13), 3425 (13). 3458 (1), 3517 (19); 3551 (12). 3768 (19),

3870 (12), 4175 (10a), 4420 (23), 4439 (23), 4726 (11), 4972 (4), 5027 (22),

5036 (9), 5184 (13), 5297 (9), 5923 (19), 6089 (27), 6105 (22), 6265 (22),

6621 (21), 6754 (1), 6945 (25), 7125 (14), 7394 (14), 7635 (6), 8195 (18),

8198 (23), 8305 (25), 8571 (23), 8589 (21), 8689 (21), 8828 (19), 10034 (14),

10280 (24), 10419 (18), 10442 (19), 10588 (2), 10758 (7), 10783 (9), 10920 (24)

King & Hullett 253 (16)

Kloss 7027 (8); sn, Rawong, Selangor (5)

Kochummen 1502 (22), 2011 (9), 2095 (17), 2114 (5), 2353 (21), 2384 (6), 2484

(10), 2507 (11), 2568 (19), 2618 (23), 2718 (23), 2824 (11), 16520 (21), 16678

(9), 79112 (21), 98156 (9), 98582 (23), 98586 (11), 99517 (19)

Kunstler 3265 (13), 7123 (14)

Lake & Kelsall 4073 (16), 4074 (4)

Liew 37742 (12)

Lim sn from Taman Negara (24)

Lobb 296 (26) :

Loh 13369 (9), 13445 (21), 13459 (21), 17304 (14), 17310 (2)

Mahamud 0802°(2), 15519 (22), 17164 (24)

Mahmud 819 (22)

Mahmud bin Sider 13212 (10)

Maingay 809 (1423) (4), 810 (2551) (22), 811 (1228) (23), 812 (10),- 813 (1567)

(25), 814 (1422) (11), 815 (2746) (21), 816 (3112) (21), 817 (1688) (6), 818

(1424) (9), 819 (2528) (20), 819 (2531) (20), 820 (2598) (9), 821 (1565) (2)

Mat 2053 (21), 3739 (16), 3741 (1), 4038 (24); sn, Tahan River, Pahang (9)

Maxwell 76-790 (19), 77-84 (19), 77-372 (9), 78-56 (19), 80-180 (10)

Meh 8977 (19), 17879 (23), 17954 (22)

Mohamet 2024 (14)

Moysey & Kiah 33632 (23), 33635 (25), 33641 (6), 33738 (8), 33856 (8), 33908 (7)

Murdock 164 (9)

Nauen sn, Penang Hill, April 1940 (20); sn, Penang Hill, April 1940 (26): sn,

Penang Hill, May 1940 (6)

118 Gardens’ Bulletin, Singapore XX XIII (1980)

Ng 1112 (25), 1153 (17), 1160 (17), 5460 (12), 5799 (25), 97953 (11), 97986 (23),

98023 (5)

Ngadiman 34734 (25), 34975 (14), 36139 (5), 36360 (12), 36931 (11), 37009 (2)

Nong 4038 (24)

Noor & Samsuri 10 (4)

Nur 11259 (17), 11318 (10), 11572 (9), 11654 (24), 11679> (23), E1699 sige

11778 (1), 11881 (24), 19976 (16), 32862 (9), 34371 (26)

Nur & Foxworthy 12073 (9), 12257 (11), 12159 (14)

Ogata 110396 (24)

Osman 23669 (9), 23759 (22)

Pangi sn, Singapore (4)

Pawanchee 12902 (19), 13760 (27)

Poore 6133 (10)

Poore & Merton 1133 (10)

Purseglove 5505 (27)

Rao sn, Singapore (14)

Rao, Keng, Wee 77 (4)

Renggol 20341 (12)

Ridley 218 (4), 1027 (10), = 1027 (10), 1184 (5), 1815 (19), 2023 (2), 2026 (19),

2053 (21), 2237 (25), 2240 (11), 2241 (9), .2242. (21),. 2243 (6), 2340) (2h

2609 (9), 2938 (9), 3296 (17), 3297 (1). 3614a (21), 3840 (M. lancifolium

Ridl.), 3848 (19), 4574 (1), 4656 (1), 4804 (19), 4805 (14), 5092 (21), 5752

(9), 5753 (13), 5928 (21), 6054 (10), 6215 (21), 6218 (19), 6219 (12), 6411 (1),

6412 (1), 6413 (1), 6414 (23), 6414A (23), 6531 (10), 6788 (15), 7309

(1), 7312 (22), 7331 (9), 7333 (14), 7334 (18), 7962 (19), setae eee

8279 (102), 8615 (7), 9182 (2), 9210 (21), 9312 (22), 9474 (11), 9475 (28),

9526 (28), 9564 (10), 9565 (10a), 9591 (10a), 9628 (28), 9638 (9). 10390 (27),

10752 (24), 11078 (22), 11092 (10a), 12179 (23), 13012 (6), 13423 (9), 13501

(16), 13502 (19), 13507 (21), 14060 (24), 14102 (28), 14167 (14), 14171 (14),

14178 (26), 14687 (28), 14695 (9), 14696 (9), 15057 (4), 15058 (19), 15396 (10a),

15399 (25), 15570 (7), 15591 (1), 15618 (1) 15622 (2), 15693 (9), 15777 (4),

15778 (19), 15779 (10a), 15804 (10), 15828 (26), 15829 (26), 16279 (10a);

sn, Gunong Panti, Johore (23); sn, Sarom, Johore (11); sn, Sungai Bau,

Johore (1); sn, Tau, Kedah (4); sn, Yan, Kedah (4); sn, Kota Baru, Kelantan

(10); sn, Bukit Besar, Malacca (23); sn, Bukit Tangga, Negri Sembilan: 17,

22. 23 Dec. 1920 (9): sn, Bukit Tangga, Negri Sembilan, Dec. 1920 (21);

(23); sn, (= 2241), Kuala Tembeling, Pahang Aug. 1891 (9); sn, Kuala

Tembeling, Pahang in 1891 (9); sn, Praman Pekan, Pahang (10); sn, Gov.’t

Revision of Memecylon L. 119

Hill, Penang, Mar. 1896 (13); sn, Pangkor, Dindings (4); sn, Waterfall,

Penang in June 1893 (19); sn, Waterfall woods, Penang, Mar. 1915 (19);

sn, Penang, Mar. 1915 (28); sn, Dindings, Perak (4); sn, Maxwell’s Hill,

Perak, June 189? (9); sn, Maxwell’s Hill, Perak, June 1893 (22); sn, Gua

Batu, Selangor, July 1897 (26); sn, Singapore in 1890 (10); sn, Singapore in

1892 (21); sn, Singapore in 1893 (21); sn, Singapore in 1894 (10); sn,

Singapore in 1906 (10); sn, Bukit Jura, Wellelsey in Dec. 1895 (4)

Robinson 6321 (10), 6328 (4)

Saaid 20661 (6)

Sallin 9710 (11)

Samsuri & Shukor 447 (22), 450 A (19), 484 (23), 727 (11), 768 (22), 957 (21),

958 (22)

Scortechini 87 (19), 89 (11), 231 (28), 394 (7), 641 (4), 1027 (4), 1309 (24), 2035

(7); 2069 (23); sn, Perak (19); sn, Perak: (28)

Seimund sn, Pulau Rumbia, Malacca (10), (19), sn, Pulau Lalang, Perak (4)

Selvaraj 11189 (9), 99660 (4)

Shah 172 (22), 194 (9), 223 (23), 454 (16), 1535 (9), 1733 (6), 2484 (2), 2486 (9),

3666 (2)

Shah & Ali 2832 (sp.), 2880 (17), 2886 (26), 2891 (10)

Shah & Kadim 484 (2), 489 (9)

Shah & Noor 1743 (21), 1767 (17), 1777 (9), 1838 (8), 1842 (23), 1843 (21)

Shah & Samsuri 3635 (17), 4031 (10)

Shah & Sanusi 2133 (16), 2173 (15)

Shah & Shukor 2628 (2), 3130 (17), 3178 (2), 3232 (23), 3406 (28), 3450 (25),

3451 (14)

Shah, Shukor, Ahmad 2624 (9)

Shah, Shukor, Awang 3303 (21)

Shah & Sidek 1158 (24)

Shah, Sidek, Samsuri 3722 (23), 3777 (9)

Shing 6861 (2), 6865 (22), 6873 (22), 6877 (11), 13357 (22), 13538 (2), 17093 (12),

17155 (23), 17265 (17), 17338 (19), 17388 (2)

Shimizu & Stone 14452 (28)

Sidek 380 (6)

Sinclair 38992 (5), 39004 (10), 39045 (4), 39549 (16), 39658 (25), 40234 (25),

40726 (10)

120 Gardens’ Bulletin, Singapore XX XIII (1980)

Sinclair & Kiah 39870 (22), 39964 (6), 40939 (23), 40858 (8)

Soepadmo 759 (21), 837 (2)

Soh 15701 (19)

Sohadi 14612 (21), 14661 (21), 17910 (17)

Sow 23712 (13)

Sow & Tachon 16858 (27)

Stone 6906 (10a), 6934 (19), 6967 (10a), 8635 (17), 9472a (14), 9474 (14)

Stone & Mahmud 8505 (17), 8540 (22)

Strugnell 10988 (4), 12738 (14), 20464 (21), 22259 (9), 22472 (6), 22475 (6)

Suilliee 8888 (7)

Suppiah 11262 (9), 11258 (22), 11353 (27), 11408 (8), 11746 (21), 11846 (11),

11865 (11), 11867 (12), 14877 (19), 14891 (21), 17770 (17), 104585 (23),

104811 (6), 104957 (2), 108881 (21), 108952 (23), 108998 (11)

Syed-Alli 23365 (21)

Symington 21077 (14), 22644 (23), 24075 (9), 24199 (9), 24224 (19), 24235 (13),

37731 QQ)

T. & P. (Teo & Purseglove) 8 (2608) (6), 12 (2), 43 (2643) (21), 349 (2949) (10),

439 (3039) (2)

Tassin 4498 (23)

Turnau 741 (4)

Umbai see Gadoh (anak Umbai)

UNESCO 95 (28), 241 (9), 326 (26), 520 (9), 609 (8)

Walker 176 (19); sn, Singapore (4)

Wallich 4100E (4), 4101 (28), 4101C (28), 4102A (21), 4103B (10a), 4108 (10),

4109 (7), 4109B (10a), 4472 (10a)

Watson 16410 (4)

Whitmore 0059 (24), 0078 (11), 0079 (23), 0345 (17), 0405 (21), 0746 (27), 0812

(17), 0817 (21), 0925 (9), 3006 (10a), 3081 (21), 3194 (17), 3231 (9), 3583

(23), 3631 (11), 3683 (2), 3751 (21), 4039 (9), 4318 (25), 4490 (24), 4620

(10a), 4629 (9), 4678 (19), 4803 (17), 8691 (22), 8748 (9), 8773 (23), 8829

(2), 11216 (22), 12175 (17), 12320 (23), 12341 (23), 12415 (21), 12438 (25),

12453 (17), 12539 (27), 12543 (11), 12554 (27), 12556 (17), 12690 (17), 12984

(4), 15030 (22), 15043 (22), 15105 (22), 15868 (9), 20363 (22), 20569 (22)

Revision of Memecylon L. 121

Whitmore & Wong 115654 (14)

Wilkinson 20776 (22)

Wong 2566 (10)

Wood 76072 (11)

Wray 744 (18), 1066 (21), 1148 (21), 1310 (11), 1964 (28), 2258 (19), 2326 (28),

2727 (13), 2961 (14), 2989 (9), 3203 (14), 3235 (25), 3240 (22), 3278 (28),

3425 (21)

Wray & Robinson 5352 (10)

Wyatt-Smith 76278 (11)

Yeob 1097 (4)

Yong 94662 (23), 94670 (11), 99002 (20)

Yusoff 99102 (19), 99141 (6)

Zainuddin 99778 (27)

coll. ? 797 Nyalas, Malacca (2); 6410 Weld’s Hill, Selangor (19)

iZz Gardens’ Bulletin, Singapore XX XIII (1980)

INDEX TO TAXA (NUMBERS REFER TO EACH TAXON)

Memecylon acuminatissimum BI. ee coe an ne N E BS

. acuminatum Sm. var. acuminatum ae —u oe a eu J

. acuminatum Sm. var. flavescens Cl. as ae ee £4 ~ la

. amabile Bedd. var. malacensis Cl. At te ey: ye ste 20

. ambiguum BI. Pet ee e at eae ne ae on 23

. amplexicaule Roxb. te Sti _ sri Bo < Re 2

. amplexicaulis Roxb. Me aa a ae ae ee a 2

. appendiculatum Bl. ve, a ie Ay ee see a, aS

. beccarianum Cogn. “y om oe es ee She Aes 3

. brandisianum Craib the ee at ot m2 + 23

. caeruleum Jack - jae ae nid As aA ns a 4

. caeruleum Jack var. floribundum (Bl.) Kurz _... ial a £0. 4

. caeruleum Jack var. pulchrum (Kurz) Cl. sae soe iN ef. 21

. caloneuron Mig. a oe a ate as a une Se 25

M. campanulatum Cl. ... im a3 ~,) Hi: i at ae 5

M. cantleyi Ridl. oN oe oy ai no A, ae SY 6

M. celastrinum Kurz _... or: a one ee ‘4 ae te pe)

M. cinereum King Me v5 a aa Sa vt a Bat ¥;

M. confine Bl. ... ve et bee 1% at es vas ber 19

M. constrictum Craib... fr Py ee ? rat Hee a ae

M. cordatum Wall. ag ree ng a “a a a ~ 4

M. corticosum Ridl. i nF val ies - rT ay ve 8

M. costatum Maiq. or. sal reg Ae she 7 an Se omy)

M. curtisii Burk. & Hend. nol iy tes “A A: ie al 9a

M. depokkense Bakh. f. + ap Bae ces ae ny Art 13

M. depressum Benth. ex Triana “s tt ae pin Rts nai 28

M. dichotomum (Cl.) King var. dichotomum a Mo ay ne 9

_ dichotomum (Cl.) King var. rotundatum (Craib) Maxw. me eA 9a

Revision of Memecylon L.

. dichotomum (Cl.) King var. eugeniiflorum (Ridl.) Rid].

. dissitum Craib

edule Roxb. var. edule

edule Roxb. var. ovatum (Sm.) Cl.

elegans Kurz var. dichotoma Cl.

elmeri Merr.

. epiphyticum King

eugeniiflora Ridl.

. excelsum BI.

floribundum Bl.

floridum Ridl.

fruticosum King

garcinioides BI.

gigantifolium Elm. ...

globiferum Wall.

. globosum Bakh f.

glomeratum Bl.

gracilipes Rid.

gracilipes Ridl. var. rotundatum Craib

grande Rett. var. horsfieldii (Miq.) Cl.

grande Retz. var. khasiana Cl.

grande Reitz. var. pubescens Cl.

. hepaticum BI.

heteropleurum BI.

heteropleurum Bl. var. olivaceum King (“‘olivacea’’)

grande Retz. var. horsfieldii (Miq.) Cl.

. hullettii King

. Intermedium BI.

. intermedium Bl. var. longipes BI.

. kunstleri King

123

124

ee Ss Ss SF FS RS RR RR RR RR KR KR OS KR OS 8 See

Gardens’ Bulletin, Singapore XX XIII (1980)

. kurzii King

. laevigatum BI.

. laevigatum Bl. var. sulcicarpum Furtado

. lampongum Miq.

. lancifolium Rid.

. laurinum BI.

. laxiflorum Wall. ex Ridl.

. lilacinum Zoll. & Mor.

. longifolium Cogn.

. longifolium Ridl.

. maingayi Cl.

malaccense (Cl.) Ridl.

. Megacarpum Furtado

micranthum Bl.

microstomum Cl.

minutiflorum Miq.

multiflorum Bakh. f.

myrsinoides BI.

myrsinoides Bl. var. lilacinum (Zoll. & Mor.) King

nudum BI.

. oleifolium BI. (“‘oleaefolium’’)

. oleifolium Bl. var. laurinum (BI.) Bakh. f.

. oligoneurum BI.

ovatum Sm.

. paniculatum Jack

pauciflorum Bl. var. pauciflorum

pauciflorum Bl. var. brevifolium Craib

. pauciflorum Bl. var. obovatum Furtado

. perakense Merr.

. pseudo-nigrescens BI.

10a

26a -

26a

Revision of Memecylon L.

[se 6. ak ER EPR ER RS

. pseudo-nigrescens Bl. var. acuminatum BI.

. pubescens (Cl.) King

. pulchellum Ridl.

. pulchrum Cogn.

. pulchrum Kurz

. rhodophyllum Bakh. f.

. ridleyi Cogn. ex Ridl.

. steenisii Bakh. f.

. subtrinervium Mig.

. tenuifolium Ridl.

. umbellatum Wall. (in part)

. venosum Merr.

. wallichii Ridl.

125

Gardens’ Bulletin, Singapore XX XIII (1980)

REDUCED TAXA FROM THE MALAY PENINSULA

. brandisianum Craib, Kew Bull. 1930, 323 = M. oleifolium BI.

. dissitum Craib, Kew Bull. 1930, 325 = M. cantleyi Ridl.

. epiphyticum King, J. As. Soc. Beng. 69, II: 1 (1900) 74 (Mat. Fl. Mal. Pen.

III, 482) = M. fruticosum King

. eugeniiflora Ridl., J. Str. Br. Roy. As. Soc. 57 (1910) 48, and M. dichotomum

(Cl.) King var. eugeniiflorum (Ridl.) Ridl., Fl. Mal. Pen. I (1922) 812 =

M. dichotomum (Cl.) King

. gracilipes Ridl., J. Str. Br. Roy. As. Soc. 79 (1918) 72, and M. gracilipes Ridl.

var. rotundatum Craib, Fl. Siam. Enum. 1:4 (1931) 708 = M. dichotomum

(Cl.) King var. rotundatum (Craib) Maxw.

. heteropleurum Bl. var. olivaceum King, J. As. Soc. Beng. 69, II: 1 (1900) 78

(Mat. Fl. Mal. Pen. III, 486) = M. excelsum Bl. and M. wallichii Ridl.

. laxiflorum Wall. ex Ridl., J. Str. Br. Roy. As. Soc. 79 (1918) 74 = M. edule

Roxb. var. ovatum (Sm.) Cl.

. longifolium Ridl., J. Str. Br. Roy. As. Soc. 79 (1918) 72 = M. wallichii Ridl.

. maingayi Cl. in Hk. f., Fl. Brit. Ind. If (1879) 557 = M. excelsum BI.

. microstomum Cl. in Hk. f., Fl. Brit. Ind. II (1879) 557 = M. amplexicaule

Roxb.

. perakense Merr., Gard. Bull. Str. Settl. 8 (1935) 132 = M. dichotomum (C1.)

King var. rotundatum (Craib) Maxw.

. pulchellum Ridl., J. Str. Br. Roy. As. Soc. 61 (1912) 52 = M. pauciflorum BI.

. ridleyi Cogn. ex Ridl., J. Str. Br. Roy. As. Soc. 30 (1897) 85 = M. dichotomum

(Cl.) King

. tenuifolium Ridl., Fl. Mal. Pen. I (1922) 812 = M. fruticosum King

. umbellatum Wall. Cat. 4109, in part = M. cinereum King

Revision of Memecylon L. 127

O75 mm d 0-45 nm

Fig. 1. Memecylon acuminatum Sm. var. acuminatum a. calyx and corolla bud

b: (1) immature petal, (2) mature petal c. stamen in bud d. mature stamen

a.-d. Griffith 2325/1

O15 mm C O25 mm

Fig. la. Memecylon acuminatum Sm. var. flavescens Cl. a. calyx b. petal c. mature

stamen a. & c. Curtis 816, b. Griffith 2325/2 (isotype)

128 Gardens’ Bulletin, Singapore

CY Ge

erry

XX XIII (1980)

Ol EA Si

C O25 mm

Fig. 2. Memecylon amplexicaule Roxb

a. calyx b. petal c.stamen a.-c. Ismail 97802

Cy es

b LD sess OS mm

O.5mm

Fig. 3. Memecylon beccarianum Cogn. a. calyx and style b. petal c. stamen

a.-c. Beccari 1518 (syntype)

——E |

C O15 mm

Fig. 4. Memecylon caeruleum Jack a. inflorescence b. petal

seed a.-d. Noor & Samsuri 10

c. stamen d: fruit and

—_—

Revision of Memecylon L.

b 1 ee a

O75 mm

Fig. 5. Memecylon campanulatum Cl. a. inflorescence axes, calyx, and style b. petal

c. stamen a.-c. Griffith 2325 (holotype)

a Oloan.

Fig. 6. Memecylon cantleyi Ridl. a. inflorescence b. mature flower c. petal d. stamen

a. ee b. Henderson 24514, c. & d. Ridley 13012 (holotype)

129

130 Gardens’ Bulletin, Singapore XX XIII (1980)

(W:

C 0-75 mm

——__—_|

a 15mm

Fig. 7. Memecylon cinereum King a. calyx and style b. calyx c. bud petal d. mature

petal e. stamen a., c., e. Scortechini 2035 (syntype); b. & d. Wallich Cat. 4109 (M.

umbellatum Wall.)

dl y

—_—_——

O75 mim

Fig. 8. Memecylon corticosum Ridl. a. upper node and internode b. leaf c. flower

bud d. mature calyx e. petal f. stamen a.-c. Sinclair & Kiah 40858, d.-f. Kloss

7627 (holotype)

idan

Revision of Memecylon L.

Fig. 9. Memecylon dichotomum (Cl.) King var. dichotomum a. branch, leaf, and

internode b. bracts, calyx, and style c. petal

d. stamen a.-d. Maxwell 77-372

Fig. 9a. Memecylon dichotomum (Cl.) King var. rotundatum (Craib) Maxw. a. infore-

scence axes, bracts, and calyx b. bud stamens a. & b. Haniff 14969

131

‘32 Gardens’ Bulletin, Singapore XX XIII (1980)

©:25 mm d O.25 mm

Fig. 10. Memecylon edule Roxb. var. edule a. calyx and corolla bud b. mature calyx

and style c. petal d. & e. stamens a. & d. H.M. Burkill 2526; b. Hou 763; c. & e.

Brelkerta sn, from Kuala Trengganu-on 1 Nov. 1939

Fig. 10a. Memecylon edule ‘Roxb. var. ovatum (Sm.) Cl. a. calyx and style b. petal

c. stamen a.-c. Chan 6800

Revision of Memecylon L. 133

beat soe

f _——_—4

C21 nae

d 15m

Fig. 11. Memecylon excelsum Bl. a. inflorescence b. calyx cc. petal d. stamen

f. fruit (dorsal view) a.-d. Ahmad & Shukor 450, e. & f. Corner 32246

Fig. 12. Memecylon floridum Ridl. a. calyx and style b. petal c. stamen a.-c. King’s

coll. 3551 (holotype)

134 Gardens’ Bulletin, Singapore XX XII1 (1980)

Fig. 13. Memecylon fruticosum King a. branch, leaf, and inflorescence b. calyx

c. petal d. anthers a.-d. Kunstler 3265 (syntype)

- SES: O'S mm

b. \ nae

Fig. 14. Memecylon garciniaides Bl. a. calyx b. petal c. stamen a.-c. Korthals sn

(Sumatra) (holotype)

Revision of Memecylon L. 135

4 -

™

$f C 5 M mh

| mm J

d O75 man

Fig. 15. Memecylon globosum Bakh. f. a. inflorescence and leaf b. calyx c. petal

d. stamen a.-d. van Steenis 1265 (isotype)

Fig. 16. Memecylon hullettii King a. branch, leaf, and inflorescence b. calyx and

corolla bud c. petal d. stamen a.-d. Sinclair 39549

136 Gardens’ Bulletin, Singapore XX XIII (1980)

+———_4

\ | | 5 mm

Imm

Fig. 17. Memecylon intermedium Bl. a. branch, leaves, and inflorescence b. calyx

c. petal d. anthers a. Ng 1153, b.-d. Paie 27695

| mam

“a9

Cc |5 eam

Fig. 18. Memecylon kunstleri King a. branch, leaves, and inflorescence b. calyx and

style c. petal d. stamen e. areolus a.-e. King’s coll. 10419 (lectotype)

Revision of Memecylon L. 137

Fig. 19. Memecylon lilacinum Z. & M. a. inflorescence b. calyx and corolla bud

c. petal d. stamen a.-d. Blume sn (Java) (holotype)

Fig. 20. Memecylon malaccense (Cl.) Ridl. a. calyx and style b. petal c. stamen

a.-c. Maingay 2531 (819) (syntype)

138 Gardens’ Bulletin, Singapore XX XIII (1980)

lOmm

Fig. 21. Memecylon megacarpum Furtado a. calyx and style b. petal c. stamen

d. fruit (lateral view) e. fruit (dorsal view) a.-c. Curtis 814, d. & e. Burn-Murdoch 162

O:5 mm

Fig. 22. Memecylon minutiflorum Mig. a. branch and inflorescence b. calyx and style

c. petal d. stamen a. Cubitt 867, b.-d. Maingay 810 (2251)

Revision of Memecylon L. 139

J\\ af

b Tabak

E C es

| QImm

eae, coe

a J ove es San

Fig. 23. Memecylon oleifolium Bl. a. calyx and corolla bud b. petal c. bud stamen

d. mature stamen e. fruit a.-d. Blume sn (Sumatra) (isotype), e. Whitmore 0079

) b

30mm

Fig. 24. Memecylon oligoneurum Bl. a. leaf b. inflorescence c. calyx and style

d. petal e. stamen a.-e. Whitmore 0059

4 O-25 mm

140 Gardens’ Bulletin, Singapore XX XIII (1980)

as;

a | mm b O #5 mm C d ue 5

O Sear

Fig. 25. Memecylon paniculatum Jack a. calyx b. petal c. &d. stamens a.-c. Corner

37050, d. Moysey & Kiah 33635

ber Gage air

O5 mm aie

Fig. 26. Memecylon pauciflorum Bl. var. pauciflorum a. calyx and corolla bud b. petal

c. stamens a.-c. Corner 28565

+_—____

Simm

Fig. 27. Memecylon pubescens (Cl.) King a. branch and inflorescence b. calyx c. petal

d. stamen e. fruit a.-d. Kiah 35188, e. Everett 13882

Revision of Memecylon L. 141

a gun i

Crees

O cs mmr

Fig. 28. Memecylon wallichii Ridl. a bracts and calyx b. bud petal c. stamens

a.-c. Curtis 1294 (syntype)

©. 5 men

Fig. 29. Memecylon constrictum Craib a. calyx and corolla bud b. petal c. stamen

a.-c. Haniff & Nur 2732 (holotype)

YOhnen

OD D5 mm

Fig. 30. Memecylon lancifolium Ridl. a. calyx b. petal c. stamen a.-c. Ridley 3840

(holotype)

ir,

Reservo

Douglas Teo.

., mature tree at MacRitch

Photo by Mr.

Maxwell 76-790.

lon

—

—,

»<

Ped

<a)

BS}

1S)

Specimen :

1976.

Bark of Memecylon lilacinum Z. & M

18 Dec.

31.

OL MLOMPI IR Mie AOR Me Vea ‘hae %

OO OM ei ai HOP” apus

Ep ap haste. 0 Tage

Singapore;

Fig.

Revision of Memecylon L. 143

| - Piante pernessl

BECCAR

0, BECCARI. PIANTE BORNENSI. :

Con. S808

SARAWAK. F ) Jece ax eum

1568 é :

“3

a in “fee

LP FOR 6 Fl Mee a emt AI LS 7, oil st

Fig. 32. Syntype of Memecylon beccarianum Cogn. at Kew.

144

tty,

ts j

a Z »

a 4 g oe

a emacs fire ag

‘a — oe :

Vi ™ PGs B i‘: *& :

~ Us * ee (ta hn ol % ‘ ae ;

Bl Yj = fr ¢

z ~ FLORA GRIT. INDIA, we: Kee :

Z & Name by Mr. C. B, CLARKE, i=

= é f aati ic ,

po)

al “G4 lA

Bl Meemeylin

4 Li, AAs

ye ah

Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 33. Memecylon edule Roxb. var. ovatum (Sm.) Cl. from the Wallich collection

at Kew.

One of the oldest (1831) specimens of Memecylon collected in Singapore.

Originally distributed as M. laxiflorum by Wallich, later included under M. grande

Retz. by Triana (1871) and other botanists, described in 1918 by Ridley as M. laxiflorum

Wall. ex Ridl., and now a synonym of M. edule var. ovatum.

Revision of Memecylon L. 145

-Ly-bou Ly wy Hy

Cm TYPE

Mwds feed

EXC Ol ster b/

é atin a!

are!

Vie on tre. Le,

ick INPIBE.

Fig. 34. (a) a. Memecylon excelsum BI. (holotype) with four species included in this

revision as new synonyms; b. M. heteropleurum BBL. (holotype); c. M. subtrinervium

Mig. (holotype); > M. kurzii King (isotype).

(Vhrrttglin 44%

Det R.C. Bakhoizen Utrecht ~

v.d Brink fil

d. M. maingayi Cl. (holotype); and ec.

Photos a.-e. Rijksherbarium, Leiden

146 Gardens’ Bulletin, Singapore XX XIII (1980)

iy yy, Uj

wow DP SW NHN

St-bou Lp wy Hy

VOUT AT (Hote) Leto TYPE

iso- Syn.

E! Det. # C. Bakhuizen Utrecht Y, 19 yf :

pA v.d Brink i Se

ppt SOD 0)»

imbiiaasidliM-tebaatuaprsdusp ibaa

MANGE

Fig. 34. (b)

ULM EY III VV

147

f Memecylon L.

1Sion O

Rev

“UIs wolanicus

OO ee. NN mF 1

senbnninnecesnansconan

R.H. film /4 neg.2Q .

R.C

Der

—

VS,

‘

“~

;

Ri Ns

3 :

’

“~

148 Gardens’ Bulletin, Singapore XX XIII (1980)

Herbvariam of th- late A ©. MAIN

GAY, MOD

3 No, Od ly L

£2. Fe

7 j G4

Cee is Fe Fo

Disteitast Al at hoe Koya! Gardens, Kew. 1571.

bpp tt

nyo — ©

ise

“aay

97)

‘a

~©

i anedaednoomereoonnsunasaansnnce ermanenenne +

oeeeaenengmnanares een eaeNNRNNN IRE

a eamooeaannennenannsen ,

& ts 4

bod i Uf, Zz & ae a

ae vi cmesuibsanaaldumiaaiid

Wiles. ALMA AOA AAEM

AMON DAO AO OAT

Revision of Memecylon L.

Pion of the Andom in stands

Pe. tas ah ?

Iremccy (om flet3a . fing

Date 1S a. ife,

Herb. Lugd. Bat. :

150 Gardens’ Bulletin, Singapore XX XIII (1980)

‘LORA OF SIAM

LVR aa

SK . * : SN

* S Ss

S =

Fig. 35. Memecylon pauciflorium Bl. var. brevifolium Craib (isotype)

Photo by Dr. Ming Anthony

CERCOSPORA AND ALLIED GENERA OF SINGAPORE AND

THE MALAY PENINSULA

J. M. YEN! & G. Lim?

INTRODUCTION

The genus Cercospora and allied genera are represented by a great number

of species which cause leaf spot disease on a wide range of host plants. They

are responsible for much damage to economic plants such as cereals, grasses,

vegetables, forest trees and ornamentals and are important plant pathogens.

The diseased spots on the leaves vary in shape and size and are generally

coloured pale brown to brown. Under humid conditions, abundant conidia are

formed on the necrotic spots. Conidiophores appear as tufts or fascicles, arising

from the lesion surface or through the stomata in the diseased areas. The perfect

stage has not been encountered so far in the collections examined and described

here.

Since the genus Cercospora was established by Fresenius in 1863, numerous

species have been recorded and described by mycologists and phytopathologists.

The monograph by Chupp (1954) is a valuable reference source, and Deighton’s

many studies (1967, 1971, 1973) have added considerably to the elucidation of

this interesting group of fungi. More recently, Deighton (1976) has redescribed

a number of Cercospora-like species and estimated that some 2000 species names

have been published to date in the genus Cercospora, which is a heterogenous

genus and one of the largest of the Hyphomycetes.

The number of species described here totalled 98, of which 72 species have

been described and published previously by the authors as new Cercospora

species. This publication brings together their reports published in various

journals, which are now translated from French into English here. The recent

studies by Deighton (1976) make it necessary for certain revisions and changes

to the generic names of some species previously published as Cercospora species.

This communication updates the authors’ previous reports and facilitates access

to information on the prevalent leaf spot diseases affecting host plants in this

region caused by this group of fungal pathogens.

1 Research Associate in Botany, Museum of Natural History, Los Angeles,

California, USA.

2 Botany Department, University of Singapore.

152 Gardens’ Bulletin, Singapore XX XIII (1980)

1. Cercoseptoria balsaminicola (Yen & Lim) Yen, comb. nov. (Fig. 1, A,

B & E)

= Cercospora balsaminicola Yen & Lim, Cahiers du Pacifique 14:90,

1970a.

Leaf spots amphigenous, distinct, circular, pale brown to brown, dispersive

or confluent, 1-7mm in diam. Fruiting amphigenous, visible as minute black

pustules on both sides of the spots. Stromata well developed, black brown,

globose or subglobose, 20-60um in diam. Fascicles present, dense to very dense,

emerging through the stomata. Conidiophores olivaceous brown, erect or slightly

flexuous, continuous or I-septate, not geniculate, tip irregularly rounded, conidial

scars indistinct, 10-27.5 x 2.5-3.54m. Conidia very thin, acicular or filiform,

hyaline, usually curved, 1-8 septate, tip acute, base cylindrically truncate, 45-132

x 2-2.5um.

Habitat: On leaves of Impatiens balsamina (Balsaminaceae), Bukit Timah,

Singapore, leg. G. Lim, Mar. 12, 1969.

Note: This fungus differs from others (Cercospora fukushiana (Mats.)

Yamam. and C. balsaminiana Yen & Lim) by its very thin filiform conidia.

2. Cercoseptoria ixoriana (Yen & Lim) Yen, comb. nov. (Fig. 2)

= Cercospora ixoriana Yen & Lim, Bull. Soc. Mycol. Fr. 85:469, 1969.

Leaf spots amphigenous, circular, whitish gray, surrounded by brown border,

scattered, 5-16mm in diam., sometimes confluent. Fruiting strictly hypophyllous,

appearing as black minute pustules under the hand lens. Stromata globose, brown

to black brown, 30-48um in diam. Fascicles dense to very dense. External

hyphae strictly hypophyllous, emerging through the stomata, pale olivaceous

brown, septate, branched, 2.5—3.54m in width. Conidiophores either from fascicles

or arising from external hyphae, erect or flexuous, simple or sometimes branched,

olivaceous brown, O-2(-3) septate, 0-2 geniculate, tip irregularly rounded, coni-

dial scars indistinct, 16-40(-48) x 3-3.5um. Conidia strictly filiform, pale oliva-

ceous brown, straight or slightly curved, 3 septate, apex rounded, base cylindrically

truncate, 36-109 x 2.5-3.5um.

Habitat: On leaves of Jxora chinensis (Rubiaceae), Singapore, leg. G. Lim,

Aug. 2, 1969.

Note: Pseudocercospora ixoricola (Yen) Yen differs from this fungus by

its angular and strictly vein-limited leaf spots and its obclavate conidia; further-

more it has no external hyphae and developed stromata.

3. Cercoseptoria polygonigena (Yen) Yen, comb. nov. (Fig. 3)

= Cercospora polygonigena Yen, Revue de Mycol. 42:143, 1978.

Leaf spots distinct, soil-brown with an indefinite yellowish border, irregularly

circular, scattered. 2-7mm in diam., sometimes up to 20mm in length. Fruiting

amphiphyllous, but more abundant on the lower side of the leaf. Fascicles

poor to meagre (2-10 stalks), emerging through the stomata. Conidiophores

amphigenous, simple, rarely branched, straight or slightly tortuous, pale olivaceous

brown, 0-2 septate, 0-1 geniculate, tip slightly attenuate and truncate, conidial

Cerccspora and Allied Genera 133

scars inconspicuous, 10-65 x 3-3.5u4m. Conidia very thin, filiform, pale olivaceous

brown, straight or slightly curved, 3—7 septate, tip conic, base cylindrically truncate,

30-90 x 2-3um.

Habitat: On leaves of Polygonum sp. (Polygonaceae), Bukit Timah (Hwa

Chung College), Singapore, leg. Chuan-ling Yen, Apr. 15, 1973.

Note: Cercospora avicularis Winter and Cercospora polygonicola Kar &

Mana! are different from this species by their obclavate conidia.

4. Cercoseptoria rhododendricola (Yen) Deighton (Fig. 4)

= Cercospora rhododendricola Yen, Revue de Mycol. 31:137, 1966.

Leaf spots angular or irregular, vein-limited, with an indefinite border, dark

brown on upper side, yellowish brown on under side, 1.5-3mm in diam.,

frequently confluent and forming large patches. Fruiting strictly epiphyllous,

appearing as minute black pustules under the hand lens. Stromata black brown,

globular or irregularly globular, 30-60um in diam. Fascicles dense to very dense,

filling the stomatal opening. Conidiophores epigenous, simple, straight or slightly

flexuous, 0-1 geniculate, generally continuous, rarely 1-2 septate, tip irregularly

rounded or laierally shouldered, conidial scars inconspicuous, 18-26 x 2.4—3yum.

Conidia filiform or acicular, pale olivaceous brown, generally slightly curved,

3-6 septate, tip obtuse, base slightly attenuate and truncate, 54-96 x 2-2.5um.

Habitat: On leaves of Rhododendron sp. (Ericaceae), Botanic Gardens,

Singapore, leg. Mrs. Shu-hsien Yen (No. 112), Apr. 13, 1965.

Note: Cercospora handelii Bubak is distinct from this species by its bigger

and not vein-limited leaf spots (2-10mm in diam.) and its amphigenous fruiting.

5. Cercoseptoria tecomae-heterophyllae (Yen) Yen. comb. nov. (Fig. 5)

= Cerospora tecomae-heterophyllae Yen, Revue de Mycol. 31:143, 1966.

Leaf spots very small, circular, whitish and surrounded by a brown margin,

scattered or coalescing to form large patches, 0.2-Imm in diam. Fruiting amphi-

genous, more abundant on the upper side, appearing as minute black pustules

under the hand lens. Stromata generally epigenous, black brown, globose or

irregularly globose, 36-90um in diam. Fascicles dense to very dense, emerging

through rupture of the epidermis. External hyphae generally hypogenous,

emerging through the stomata, pale olivaceous brown, septate, branched, 1.5-3u.m

in width. Conidiophores showing two types: A, conidiophores coming from

fascicles, pale olivaceous brown, straight or flexuous, simple (never branched),

0-3 geniculate, 0-3 septate, unthickened small condial scars at the attenuate and

truncate tip, 25-48 x 3-4.5um; B, condiophores arising from external hyphae,

pale olivaceous brown, single, straight or slightly tortuous, 1-2 geniculate, 0-2

septate, tip attenuate and truncate 12-45 Xx 3—4,um. Conidia filiform or acicular,

very pale olivaceous brown, or subhyaline, slightly curved, 2—6 septate, tip obtuse

or rounded-obtuse, base slightly truncate and cylindrical, 36-72 x 1.5-2.5um.

Habitat: On leaves of Tecoma heterophylla (Bignoniaceae), Botanic Gardens,

Singapore, leg. Mrs. Shu-hsien Yen (No. 115), Apr. 13, 1965.

154 Gardens’ Bulletin, Singapore XX XIII (1980)

Note: This fungus is distinct from others (Cercospora tecomae Chupp

& Viegas and C. stenolobiicola Chupp) by its thin conidia.

6. Cercoseptoria tecomicola (Yen) Yen, comb. nov. (Fig. 6)

= Cercoseptoria tecomicola Yen, Revue de Mycol. 32:196, 1967a.

Leaf spots distinct, angular or irregularly angular, strictly vein-limited, brown

to dark brown, scattered, 0.5-4mm in diam. Fruiting amphigenous, visible as

minute black pustules on the upper surface of the leaf, appearing as gray velvet

on the lower side of the leaf spots. Stromata epigenous, at times lacking, dark

brown to black brown, globose, 30-554m in diam. Fascicles medium to dense

or very dense. External hyphae hypophyllous, emerging through the stomata,

pale brown, septate, branched, 2-3um in width. Conidiophores showing two

types: A, conidiophores coming from fascicles, pale brown, paler towards the

apex, erect or flexuous, simple or branched, 0-3 septate, 0-3 geniculate, tip

rounded or laterally shouldered, 21.5-49 x 3-4.8um; B, conidiophores arising

from the external hyphae, pale olivaceous brown, single, 0-1 septate, 0-1 genicu-

late, tip rounded or laterally shouldered, 6-19.5 x 2.5-4.5um. Conidia cylindrical

or acicular, pale olivaceous brown, slightly curved, 1-9 septate, tip subrounded,

base cylindrical and truncate, 18-79 x 2.5-3um.

Habitat: On leaves of Tecoma stans (Stenolobium stans) (Bignoniaceae),

Katong, Singapore, leg. Jo-min Yen (No. M.S. 771), Jan. 5, 1966.

Note: Cercospora stenolobiicola Chupp differs from this species by its hyaline

conidia and Cercoseptoria tecomae-heterophyllae (Yen) Yen is distinct from this

fungus by its very small (0.2-Ilmm in diam.) and not vein-limited leaf spots.

7. Cercospora abacopteridis Yen & Lim (Fig. 7, C-E)

Cahiers du. - Pacifique 17 :96;; 1973.

Leaf spots amphigenous, brown with an indefinite border, at first vein-limited,

then confluent. Fruiting hypophyllous, invisible. Stromata lacking, fascicles poor

to midly dense (2-18 stalks), emerging through the stomata. Conidiophores dark

brown, paler towards the apex, always simple, erect or slightly flexuous, 1-5

septate, with O-2 small geniculations, a large black brown conidial scar at the

rounded tip, 15-118 x 4-5um. Conidia acicular or filiform, hyaline, straight or

slightly curved, 1-35 septate, tip acute, base cylindrical-truncate with a black

brown scar at its end, 62-400 x 2—4ym. |

Habitat: On leaves of Abacopteris urophylla (Pteridophyte), Singapore, leg.

G. Lint (No, 71); Mata3i-ie 2.

8. Cercospora acori Yen (Fig. 8)

Revue de Mycol. 29:209, 1964.

Leaf spots indistinct, sometimes appearing as gray darkish specks. Fruiting

amphigenous, effuse, dark brown, Stromata lacking. External hyphae brown or

yellowish brown, branched, septate, 2.5-4.84m in width. Conidiophores as single

branches from procumbent threads, never fasciculate, brown or dark brown,

cylindrical, not geniculate, 2—6(—8) septate, tip rounded to conic, several small

brown scars on the upper part, 21-190 x 44.8um. Conidia pale olivaceous

brown, obclavate or obclavatoacicular, 4-9(-12) septate, tip acute, base obconically

truncate, 40-120 x 2.3um.

Cercospora and Allied Genera 155

Habitat: On leaves of Acorus calambus (Araceae), Botanic Gardens, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 7), Jan. 26, 1964.

Note: This fungus is parasitic on the uredospores of Uromyces aparganii

Clint. & Peck which is parasitic on the same host plant at the same time.

9. Cercospora apii Fres.

Beitr. Myk. 3:91, 1863.

Syn. Cercospora penicillata var. apii Fuckel, Hedwigia 2: 132, 1863.

Leaf spots subcircular to irregular, brownish to ashy gray, surrounded by

an indistinct greenish border, 1-Smm in diam., often confluent, Fruiting amphi-

genous, invisible. Stromata lackmg or very small. Fascicles very poor. Conidio-

phores olivaceous brown, simple, straight or slightly flexuous, 2-6 septate, not

geniculate, a large black brown scar at the rounded tip, 50-220 (-300) x 4-6um.

Conidia filiform, acicular, hyaline, straight or curved, multiseptate, tip acute to

subacute, base truncate, 35-210 (—300) x 2-3.5um.

Habitat: On leaves of Apium graveolans (Umbelliferae), Bukit Timah,

Singapore, leg. G. Lim, Apr. 25, 1969.

Distribution: N. America, Europe, India, Korea, Japan, Manuchuria and

Singapore.

10. Cercospora asystasiana Yen (Fig.9)

Revue de Mycol. 32:180, 1967a.

Leaf spots orbicular, brown with an indefinite border, scattered, 2-Smm in

diam., sometimes confluent. Fruiting strictly hypophyllous, invisible. Stromata

absent, sometimes present but very rudimentary. Fascicles very poor (2-5 stalks),

emerging through the stomata. Conidiophores hypogenous, simple, straight or

tortuous, dark brown, with 0-5 small geniculations, 0-5 septate, a large black

brown conidial scar at the rounded tip, several small scars on the upper part,

30-132 x 3-3.5um. Conidia hyaline, acicular or obclavato-cylindrical, mildly

curved, 5-20 septate, tip acute to obtuse, base subtruncate and with a black

brown scar at the end, 48-185 x 2.5—5yum.

Habitat: On leaves of Asystasia chelonoides (Acanthaceae), Botanic Gardens,

Singapore, leg. Jo-min Yen (No. M.S. 769), Jan. 5, 1966.

Note: This fungus is different from Pseudocercospora asystasiae (Yen) Yen

by its hypophyllous fruiting and hyaline conidia.

11. Cercospora balsaminiana Yen & Lm (Fig. 1, C & D)

Cahiers du Pacifique 14:91, 1970a.

Leaf spots none or indistinct. Fruiting epigenous, invisible. Stromata absent

or very rudimentary, consisting of few brown cells. Fascicles very poor (2-7

stalks), emerging through the stomata. Conidiophores pale brown, always simple,

cylindrical, erect, usually continuous, sometimes | septate, a large dark brown

conidial scar at the rounded tip, 15-40 x 4.5-6um. Conidia acicular, hyaline,

straight or slightly curved, 5-17 septate, tip acute, base truncate and with a black

brown scar at the end, 66-307 x 2.5-4.5ym.

156 Gardens’ Bulletin, Singapore XX XIII (1980)

Habitat: On leaves of /mpatiens balsamina (Balsaminaceae), Bukit Timah,

Singapore, leg. G. Lim, Mar. 12, 1969.

Note: Cercospora fukushiana (Mats.) Yamam. differs from this fungus by

its amphiphyllous fruiting and its 1-4 septate conidiophores.

12. Cercospora barleriae Yen & Lim (Fig. 10)

Cahiers du Pacifique 17:100, 1973.

Leaf spots amphigenous, angular, vein-limited, black on the upper side and

dark brown on the under side, usually confluent and forming large black brown

patch, sometimes extending over a great part of the leaf surface. Fruiting

strictly hypophyllous, effuse, visible as dark brown velvet on the under side of

spots. Stromata absent. Fascicles poor to mildly dense (2-18 stalks), emerging

through the stomata. Conidiophores olivaceous brown, paler towards the tip,

simple, rarely branched, flexuous, 1-7 septate, with 0-3 small geniculations, apex

slightly attenuate and truncate, with brown conidial scar at the end, 25-130 x

5—6um. Conidia pale olivaceous brown, obclavate or obclavato-cylindrical,

straight or slightly curved, 1-8 septate, tip rounded or subobtuse, base slightly

attenuate and truncate, 13-80 x 4-5Sum.

Habitat: On leaves of Barleria cristata (Acanthaceae), Singapore, leg. G.

Lam. (No. 8:U,"73); Jano oa o7t::

Note: Cercospora barlericola Payak and C. barleriae-cristatae Gov. & Thir.

are different from this fungus by their amphiphyllous fruiting and hyaline conidia.

13. Cercospora canavaliiana Yen & Lim (Fig. 11, A-F)

Cahiers du Pacifique 14:94, 1970a.

Leaf spots distinct, medium brown on the upper side, grayish brown on

the lower side, irregularly orbicular, 3-25 mm in diam. Fruiting amphigenous,

but chiefly hypophyllous. Stromata absent or very rudimentary. Fascicles poor

to medium (2-12 stalks), emerging through the stomata. Conidiophores oliva-

ceous brown, simple, cylindrical, always erect, not geniculate, continuous (rarely

1 septate), a large black brown conidial scar at the rounded and paler tip, 36—80

x 5-6%m. Conidia cylindrical or filamentous, hyaline, slightly curved. 3-10

septate, apex rounded, base truncate with a black brown scar at the end, 60-200

x 2.5-4.8um.

Habitat: On leaves of Canavalia gladiata (Leguminosae), Bukit Timah,

Singapore, leg. G. Lim, Apr. 5, 1969.

Note: This species differs from others (Cercospora canavaliae Syd. and

C. canavaliicola Saw. & Kats.) by its amphiphyllous fruiting and filiform conidia.

14. Cercospora canescens Ell. & Mart.

Amer. Nat. 16: 1003, 1882.

Syn. Cercospora vignicaulis Tehon, Mycologia 29:436, 1937.

Cercosporiopsis canescens (Ell. & Mart.) M. Miura, South Manch.

Railway Co. Agr. Rept. 27:529, 1928.

Cercospora and Allied Genera 157

Leaf spots amphigenous, suborbicular to irregular, reddish brown with an

indefinite border, at times vein-limited, 1.5-15mm in diam., sometimes confluent.

Fruiting amphigenous, more abundant on lower side, invisible. Stromata very

poor. Fascicles poor to dense (2-25 stlks), emerging through the stomata. Coni-

diophores may be divided into two kinds: A, conidiophores from fascicles

simple, erect or slightly flexuous, olivaceous brown, 0-7 septate, 0-3 geniculate,

a large black brown conidial scar at the rounded tip or somewhat laterally dis-

placed, 28.8-174 x 4-Sum: B, conidiophores arising from external hyphae, simple,

erect, olivaceous brown, continuous, a small conidial scar at the rounded tip,

2.4-12 x 3.6-S5um. Conidia hyaline, acicular, straight or curved, 3-18 septate,

tip acute, base truncate and a black brown scar at the end, 75-190 x 2.5-3.5um.

Habitat: On leaves of Phaseolus vulgaris (Leguminosae), Jurong, Singapore,

leg. Shu-ling Lin, Dec. 10, 1968.

Distribution: N. & S. America, Africa, India, China, Manchuria. Japan

and Singapore.

15. Cercospora capsici Heald & Wolf

Mycologia 3:15, 1911.

Leaf spots suborbicular, 1-5 up to 10mm in diam., at first brown, then

becoming grayish brown, raised on the upper surface, surrounded by a raised

dark brown border. Fruiting amphigenous, invisible. Stromata lacking or small,

brown 15-30um in diam. Fascicles poor to mildly dense (2-18 stalks), emerging

through the stomata. Conidiophores straight to slightly curved, simple, 1—3

abruptly geniculate, 1-2 septate, medium brown, paler towards the apex, a dark

brown conidial scar at the truncate tip, 30-60 x 4.5-5.54m. Conidia hyaline,

acicular, straight or slightly curved, 1-5 septate, tip subobtuse, base truncate,

30-125 x 3.5-4um.

Habitat: On leaves of Cansicum annum (Solanaceae), Bukit Timah, Singa-

pore, leg. G. Lim, Apr. 6, 1969.

Distribution: N. & S. America, China, Japan and Singapore.

16. Cercospora cassiae-nodosae Yen (Fig. 12)

Revue de Mycol. 43:97, 1979..

Leaf spots amphigenous, distinct, black, circular or subcircular, scattered,

12mm in diam., rarely 2-3 confluent. Fruiting strictly hypophyllous, visible

as minute black pustules on the under surface of the spots. Stromata globose

to irregularly globose, black brown, 25-40um in diam. Fascicles dense to very

dense, emerging through the stomata. Conidiophores hypophyllous, always simple.

erect or slightly tortuous, olivaceous brown, 2-5 septate, Q-2 geniculate, a dark

brown conidial scar at the rounded tip, 60-145 x 3-4um. Conidia obclavate,

olivaceous brown, curved, 3-7 (generally 5-6) septate, tip conic, base obconically

substruncate, with a dark brown scar at the end, 25-65 x 3-Sum.

Habitat: On leaves of Cassia nodosa (Leguminosae), Kuala Lumpur (For.

Res. Inst.), Malaysia, leg. Jo-min Yen (No. 71339), Sept. 28, 1971.

Note: This fungus is distinct from others by its black leaf spots, developed

stromata, coloured conidia and long conidiophores (60—-145pum).

158 Gardens’ Bulletin, Singapore XX XIII (1980)

17. Cercospora cassiae-occidentalis Yen (Fig. 13)

Revue de Mycol. 29:212, 1964.

Leaf spots indistinct, sometimes with many small gray reddish points on

both sides of the leaf. Fruiting amphigenous, invisible. Stromata lacking.

Fascicles very poor (2-6 stalks), filling stomatal opening. Conidiophores simple,

not branched, olivaceous brown, 2-6 septate, 1-5 geniculate, straight or tortuous,

paler and more narrow towards the tip, bulbiform at the base, a brown small

conidial scar at the attenuate tip, 60-128.5 x 4-5ym. Conidia pale olivaceous

brown, obclavato-cylindrical or cylindrical for the short ones, straight or slightly

curved, 3-6 septate, tip rounded or subrounded, base obconically truncate,

62.4-98.5 x 3.6-4.8um.

Habitat: On leaves of Cassia occidentalis (Leguminosae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No. 30) Apr. 25, 1964.

Note: The amphigenous fruiting and the unbranched conidiophores separate

this species from Pseudocercospora nigricans (Cooke) Deighton.

18. Cercospora citrullina Cooke

Grevillea 12:31, 1883.

Syn.: Cercospora cucurbitae Ell. & Ev., J. Mycol. 4:3, 1888.

Cercospora sechii Stevenson, P. Rico Ins. Exp. Sta. Dept. Agr. Ann.

Rept. (1947), b@: 1 3a bo19.

Cercospora chardoniana Chupp, Monogr. Univ. R. Rico. B. 2:245, 1934.

Cercospora momordicae Mendoza, Philipp. J. Sci. 75:173, 1941.

Cercospora momordicae Sawada, Descr. Cat. Formosan Fungi. 9: 173,

1943.

Leaf spots subcircular or angular to irregular, 2-Smm up to 10mm in diam.,

brownish with a dark brown to purplish raised border, centre becoming grayish.

Fruiting amphigenous but chiefly epiphyllous, invisible. Stromata lacking or

consisting of few brown cells. Fascicles poor to medium (2-10 stalks), emerging

through the stomata. Conidiophores simple, erect or slightly flexuous, usually

geniculate at the upper part, 2—8 septate, 1-6 geniculate, olivaceous brown, but

paler towards the apex, a large black brown conidial scar at the rounded tip,

40-240 x 3.5-S5um. Conidia acicular or filiform, sometimes cylindrical, hyaline,

straight or curved, 3-29 septate, tip acute, base truncate, 60-447 x 3-4um.

Habitat: On leaves of Momordica charantia (Cucurbitaceae), Bukit Timah, —

Singapore, leg. G. Lim, Jul. 17, 1969.

Distribution: N. & S. America, Europe, Africa, China, Japan and Singapore.

19. Cercospora coleiana Yen & Lim (Fig. 14)

Bull. Soc. Mycol. 85:463, 1969.

Cercospora and Allied Genera 159

Leaf spots amphigenous, circular to irregularly circular, brown to dark brown,

but centre becoming whitish and with an indefinite border, 1—-6mm in diam.

Fruiting amphigenous, but more abundant on the lower side, invisible. Stromata

lacking. Fascicles very poor (2-4 stalks), emerging through the stomata. Coni-

diophores pale brown to brown, simple or branched, erect or flexuous, 0-11

septate, 0-5 slightly geniculate, a small black brown conidial scar at the attenuate

apex, usually several (5-10) black brown conidial scars on the upper part of

the old ones, 40-181 x 5-7.2um. Conidia acicular, hyaline, straight or slightly

curved, 1-18 septate, tip acute, base truncate with a black brown scar at the

end, (40—) 80-276 x 2-4.8um.

Habitat: On leaves of Coleus sp. (Labiatae), Singapore, leg. G. Lim, Jul.

17,1969.

Note: Cercospora coleicola Chupp & Mull and C. colei Boed. are different

from this fungus by their conidiophores always simple, in dense to very dense

fascicles and especially by their epiphyllous fruiting.

20. Cercospora dioscorae-pyrifoliae Yen (Fig. 15)

Bull. Soc. Mycol. Fr. 84:5, 1968.

Leaf spots amphigenous, more distinct on the upper side, circular, 0.5-8mm

in diam., whitish in the centre, surrounded by a dark brown border, finally

forming “‘shot-hole”. Fruiting amphiphyllous, visible as short black hairs in

the centre of spots. Stromata lacking or very rudimentary, consisting of few

brown cells. Fascicles poor to dense (2-28 stalks), emerging through the stomata.

Conidiophores dark brown, simple, erect or slightly flexuous, 1-7 septate, 0-3

geniculate, 1-2 small black brown conidial scars at the rounded tip, 36-200 x

6-8.4um. Conidia acicular, hyaline, straight or curved. 3-18 septate, tip acute,

base truncate, with a black brown scar at the end, 45.6-168 x 3.5—6um.

Habitat: On leaves of Dioscorea pyrifolia (Dioscoreaceae), Jurong, Singa-

pore, leg. Jo-min Yen (No. M.S. 738), Dec. 17, 1965.

Note: This fungus differs from others (Cercospora ubi Rac. and Pseudo-

cerpospora ubicola (Yen) Deighton) by the “‘shot-hole” leaf spots and the acicular

and hyaline conidia.

21. Cercospora eupatorii-odorati Yen (Fig. 16, D-J)

Bull. Soc. Mycol. Fr. 84:10, 1968.

Leaf spots amphigenous, circular, more distinct on the upper side, brown

to dark brown, scattered, 2-6mm in diam., sometimes coalescing and forming

large patches. Fruiting strictly hypophyllous, invisible. Stromata absent or

small, dark brown, globose, 18-42um in diam. Fascicles poor to meagre (2-9

stalks), sometimes dense, emerging through the stomata. External hyphae hypo-

phyllous, also emerging through the stomata, pale olivaceous brown, septate,

branched, 2.5-5ym in width. Both the conidiophores from fascicles and arising

from external hyphae may be brown or olivaceous brown, straight or flexuous,

simple or branched, 0-10 septate, 0-2 geniculate, 1-2 small black brown conidial

scars at the shortly attenuate tip, 38.4-168 x 4.5-6um. Conidia cylindrical, pale

olivaceous brown, catenate or slightly curved, 1-4 (generally 3) septate, small

black brown conidial scars at both rounded ends, 22.8-57.6 x 3.6-Sum.

160 Gardens’ Bulletin, Singapore XX XIII (1980)

Habitat: On leaves of Eupatorium cdoratum (Compositae), Ipoh, Malaysia,

leg. Jo-min Yen (No. M.S. 728), Dec. 21, 1965.

22. Cercospora gloriosicola Yen & Lim (Fig. 17)

Bull. Soc. Mycol. Fr. 85:467, 1969.

Leaf spots distinct, oval or oblong, brown with an indefinite border, scattered

3-14 x 2-7mm. Fruiting amphigenous, appearing as minute black pustules.

Stromata very developed, black brown, globose, 48-70um in diam. Fascicles

dense to very dense, emerging through the stomata. Conidiophores pale brown,

simple or branched, tortuous, 3-6 geniculate, 4-6 septate, a large black brown

conidial scar at the rounded tip and several (3-5) black brown scars laterally

displaced on the upper part, 80-168 x 4.8-7.2um. Conidia hyaline, obclavato-

cylindrical, straight or slightly curved, 3-11 septate, apex rounded or subobtuse,

base truncate with a black brown scar at the end, 50-120 x 4—Sum.

Habitat: On leaves of Gloriosa superba (Liliaceae), Bukit Timah, Singapore,

leg. G. Lim, Aug. 2, 1969.

23. Cercospora henningsii Allesch.

Engler’s Pil; Ost-Afs: Tei, 535, a1 895:

Syn. Cercospora cassavae Ell. & Ev., Bull. Torrey Bot. Club, 22: 438,

1895.

Cercospora manihotis P. Henn. Hedwigia 41:18, 1902.

Cercospora cearae Petch, Ann. Roy. Bot. Gard. Paradeniya 3:10, 1906.

Septogloeum manihotis Zimm., Centralbl. f. Bakt. Abt. 2, 8:218, 1902.

Helminthosporium manihotis Rangel, Arch. Jard. Bot. Rio de Janeiro

2: tt aees

Leaf spots circular to subcircular, more distinct on the upper side of the

leaf, scattered, 3-12mm in diam. pale brown or tan dingy to dingy gray with

a yellowish brown margin, rather indefinite on the lower surface. Fruiting

amphigenous, visible as minute black pustules under the hand lens. Stromata

small, dark brown, globose, 20-40um in diam. Fascicles dense to very dense,

filling the stomatal opening. Conidiophores pale olivaceous brown, uniform in

colour and width, simple, straight or slightly flexuous, 0-2 midly geniculate, at

times sparingly septate, tip rounded with a small brown conidial scar, 10-50 x

3.5-5um, rarely as long as 100um. Conidia cylindrical, pale olivaceous brown,

straight or slightly curved, 2-8 septate, both ends bluntly rounded or base shortly

ohconically truncate, 30-60 x 4-6yum (rarely as large as 85 x 7m).

Habitat: On leaves of Manihot utilissima (Euphorbioceae), Bukit Timah,

Singapore, leg. G. Lim, Mar. 13, 1969.

Distribution: N. America, E. Africa, Barbados, Belgian Congo, Brazil,

Ceylon Colombia, Costa Rica, Gold Coast, Panama, Philippines, Puerto Rico,

San Domingo, Sierra Leone, Singapore, Taiwan, Tanganyika, Trinidad and Uganda.

Cercospora and Allied Genera 161

24. Cercospora holobrunnea Yen (Fig. 18)

Revue de Mycol. 31:112 1966.

Leaf spots indistinct, yellowish brown. Fruiting amphigenous, but mostly

hypophyllous. Stromata lacking. Fascicles lacking. External hyphae amphi-

genous, but mostly hypophyllous, olivaceous brown, emerging through the stomata,

septate, branched, 2—34m in width. Conidiophores as single branches from

exterrial procumbent threads, olivaceous brown, cylindrical, 1-2 septate, O-1

geniculate, a snfall brown conidial scar at the rounded tip, 24-36 x 2.4~-3.6um.

Conidia cylindrical, pale olivaceous brown, straight or slightly curved, 2—5 septate,

tip rounded, base truncate with a dark brown conidial scar at the end, 24-74.4

x 2.4-3um.

Habitat: On leaves of Hydrangea macrophylla (Hydrangeaceae), Bukit

Timah, Singapore, leg. Mrs. Shu-hsien Yen (No. 89), Dec. 4, 1964; on the same

host-plant, in Hwa Chung College, Singapore, leg. Chih-shao Lee & Mrs. Shu-

hsien Pen (No. 109), April 1965.

25. Cercospora hyalofilispora Yen (Fig. 19)

Revue de Mycol. 31:115, 1966.

Leaf spots none or indistinct. Fruiting hypophyllous, invisible. Stromata

lacking. Fascicles lacking. External hyphae hypophyllous, hyaline, emerging

through the stomata, septate, branched, very thin, 1-1.5um in width. Conidio-

phores arising as single branches from external threads, pale olivaceous brown,

0-1 septate, not geniculate, with zigzag membrane, a small brown conidial scar

at the attenuate tip, 16.8-28.8 x 3-3.6um; at times some conidiophores emerging

through the stomata, 30-50 x 3.64ym. Conidia filiform, hyaline, very thin,

slightly curved, 3-14 septate, tip acute, base truncate with a dark brown scar

at the end, 28-123 x 1-1.5ym.

Habitat: On leaves of Hydrangea macrophyila (Hydrangeaceae), Katong,

Singapore, leg. Mrs. Shu-hsien Yen (No. 95), Dec. 18, 1964.

26. Cercospora hydrangeae Ell. & Ev.

J. Mycol. 8:71, 1902.

Syn. Cercospora hydrangeicola Speg., Ann. Nac. Buenos Aires 20:426,

1910.

Cercospora hydrangicola Tharp, Mycologia 9:110, 1917.

Cercospora arboressentis Teh. & Dan., Mycologia 17:246, 1925.

Leaf spots subcircular, pale, brown with a black brown border, scattered,

37mm in diam., sometimes confluent. Fruiting amphigenous, but mostly epi-

phyllous. Stromata poor or small, brown, globose, 20-364m in doam. Fascicles

poor to dense (2-20 stalks), emerging through the stomata. Conidiophores

amphigenous, erect or flexuous, medium brown, not branched, 1-5 (rarely 8 or

9) septate, 1-5 geniculate, a large brown conidial scar at the rounded tip, 48-160

x 4.5-7um. Conidia acicular or obclavato-acicular, hyaline straight or curved,

5-11 septate, tip subacute, base truncate with a terminal dark scar,'\84-176 x

3.64.8m.

162 Gardens’ Bulletin, Singapore XX XIII (1980)

Habitat: On leaves of Hydrangea macrophylla (Hydrangeaceae), Bukit

Timah, Singapore, leg. Mrs. Shu-hsien Yen (No. 89), Dec. 4, 1964; on the same

host-plant, in Hwa Chung College, Singapore, leg. Chih-shao Lee & Mrs. Shu-

hsien Yen ( No. 96), Dec. 31, 1964.

Distribution: N. America, Argentine, Japan, Singapore and Taiwan.

27. Cercospora ipomveae-pes-caprae Yen & Lim (Fig. 20)

Bull. Soc. Mycol. Fr. 86:747, 1970b.

Leaf spots amphigenous, distinct, circular to irregular circular, pale brown

to brown, paler in the centre, with a thicker black brown border, scattered,

14mm in diam., sometimes confluent. Fruiting amphiphyllous, visible as black

minute pustules under the hand lens. Stromata substomatic, composed of finely

granulate membrane cells, black brown, globose, 19-38um in diam. Fascicles

poor to meagre (2-8 stalks), emerging through the stomata. Conidiophores

olivaceous brown, cylindrical, erect, simple, continuous or 1 septate, not geniculate,

a large black brown conidial scar at the rounded tip, 30-40 x 4-5ym. Conidia

obclavate, hyaline, straight or slightly curved, 3-14 septate, apex obtuse, base

truncate with a black brown scar at the end, 44-115 x 34.5ym.

Habitat: On leaves of Ipomoea pes-caprae (Convolvulaceae), Pulau Besar

island, Malaysia, leg. G. Lim, Jan. 10, 1970.

28. Cercospora mikaniicola Stev.

Trans. III. Acad. Sci. 10:213, 1917.

Syn. Cercospora mikaniae-cordiae Yen, Revue de Mycol. 30: 183, 1965.

Leaf spots amphigenous circular, at first very small, 0.5-Imm in diam.,

reddish brown with a whitish centre, then becoming larger, 3-Smm in diam.,

forming ‘‘Frog eyes’’ and surrounded by a brown border. Fruiting amphiphyllous,

but chiefly hypophyllous. Stromata lacking or very rudimentary. Fascicles very

poor (2-7 stalks), emerging through the stomata. Conidiophores simple, never

branched, erect, cylindrical, brown, paler toward the apex, 1-2 septate, 0-2

geniculate, a large black brown conidial scar at the rounded tip, some scars

laterally displaced, 52-102 x 5—7.2um. Conidia hyaline, abruptly obclavate, at

times somewhat fusiform, straight or curved, 3-9 septate, with a long whip-like

tip, base attenuate and truncate, with a black scar at the end, 89-138 x 7.2-12ym.

Habitat: On leaves of Mikania cordata (Compositae), Bukit Timah, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 68), Jul. 1, 1964.

Distribution: Puerto Rico and Singapore.

29. Cercospora malayensis Stev. & Solh.

Mycologia 23:394, 1931.

Leaf spots circular to irregular, tan to brown, usually with a purple or red

border, fairly broad, 5—25mm in length, frequently extending from the edge of

of the leaf to the midrib. Fruiting amphigenous, invisible. Stromata lacking

or rudimentary. Fascicles meagre to dense (5-20 stalks), emerging through the

Cercospora and Allied Genera 163

stomata or rupture of the epidermis. Conidiophores pale brown, simple, straight

or flexuous, apex rounded, 35-250 x 34um. Conidia subhyaline, acicular to

obclavato-acicular, multi-septate, tip acute, base truncate, 70-115 x 2.5-3um.

Habitat: On leaves of Hibiscus mutabilis (Malvaceae), Singapore, leg. G.

Lim (No. S.U. 48), Apr. 1970.

Distribution: N. & S. America, S. Africa, Brazil, Jamaica, Japan, Philippines,

Salvador, Singapore, Taiwan, Trinidad and Venezucla.

30. Cercospora nicotianae Ell. & Ev.

Proc. Acad. Sci. Phila. 45: 170, 1893.

Syn. Cercospora raciborskii Sacc. & Syd., Syll. Fung. 16: 1070, 1902.

Leaf spots amphigenous, circular to subcircular, larger ones concentrically

zonate, uniformly brown to dark brown, 1-15mm in diam., sometimes confluent.

Fruiting amphiphyllous, invisible. Fascicles poor (2-7 stalks), emerging through

the stomata. Stromata very rudimentary, consisting of a few brown cells. Conidio-

phores brown to dark brown, simple, sometimes branched, erect or flexuous,

0-4 geniculate, 2-8 septate. a large conidial scar at the rounded tip, 38-180 x

3.6-6um. Conidia acicular, hyaline, slightly curved, 3-15 septate, tip subacute,

with a black brown scar at the end, 45-165 x 34um.

Habitat: On leaves of Nicotiana tabacum (Solanaceae), Jurong, Singapore,

leg. Jo-min Yen (No. M.S. 723), Dec. 15, 1965.

Distribution: Africa, S. America, Europe, Japan, Peru, Singapore and

Taiwan.

31. Cercospora nicotianicola Yen (Fig. 21)

Revue de Mycol. 32:188, 1967a.

Leaf spots amphigenous, distinct, triangular or irregularly angular, vein-

limited, white and surrounded by a raised dark brown border, scattered, 1-3mm

in diam. Fruiting amphiphyllous, visible as black powder at the white centre

of leaf spots. Stromata small, dark brown, subglobose, 24-54.m in diam.

Fascicles poor to dense (2-18 stalks), emerging through the stomata. Conidio-

phores brown to dark brown, paler toward the tip, generally simple, at times

branched, 2-4(—5) septate, 0-2 geniculate, one or several small conidial scars at

the attenuate tip, 5-15 small brown scars laterally displaced on the upper part,

then appearing as denticulate form, 60-218 x 5—-6um. Conidia hyaline, filiform

or acicular, straight or slightly curved, 2-48 septate. tip rounded or obtuse, base

truncate and with a black scar at the end, 70-480 x 2.5—3.6um.

Habitat: On leaves of Nicotiana tabacum (Solanaceae), Jurong, Singapore,

leg. Jo-min Yen (No. M.S. 724), Dec. 18, 1965.

Note: The small, white leaf spots, the attenuate apex conidiophores and the

filiform conidia separate this species from Cercospora nicotianae Ell. & Ev.

32. Cercospora peristrophes-acuminatae Yen (Fig. 22)

Revue de Mycol. 29:230, 1964.

164 Gardens’ Bulletin, Singapore XX XIII (1980)

Leaf spots none or indistinct, sometimes as dark, brown areas but not distinct.

Fruiting hypophyllous, rarely amphiphyllous. Stromata lacking. Fascicles poor

to meagre (2-8 stalks), filling stomatal opening. Conidiophores hypophyllous,

rarely amphigenous, simple, erect, brown to dark brown, paler and more narrow

towards the tip, a small, black brown conidial scar at the tip and several small,

black-brown scars laterally displaced at the upper part, 3-12 septate, not geni-

culate, 156-295 x 6-7.2um. Conidia clavate or long-oblong, pale olivaceous

brown, 1-3 septate, straight, tip rounded, base attenuate and truncate, 32-45 x

8—8.5.m.

Habitat: On leaves of Peristrophe acuminata (Acanthaceae), Katong, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 20), Apr. 20, 1964.

33. Cercospora psophocarpicola Yen (Fig. 23)

Bull. Soc. Mycol. Fr. 83:339, 1967b.

Leaf spots circular or irregular, brown with an indefinite border, scattered,

14mm in diam., frequently confluent. Fruiting hypogenous, usually mixed with

that of Pseudocercospora psophocarpi (Yen) Deighton. Stromata lacking or very

rudimentary, black brown, irregularly globose, 15-20u.m in diam. Fascicles poor

to meagre (2-7 stalks), emerging through the stomata. Conidiophoyes strictly

hypophyllous, dark brown, simple, never branched, erect or slightly flexuous,

1-4 septate, 1-4 geniculate, a large, black brown conidial scar at the rounded

tip, 22.8-260 x 4.8-6.m. Conidia acicular, sometimes filiform, hyaline, straight

or slightly curved, 4-17 septate, apex acute, base truncate with a black brown

scar at the end, 54-234 x 2.4-3.6yum.

Habitat: On leaves of Psophocarpus tetragonolobus (Leguminosae), Jurong,

Singapore, leg. Jo-min Yen (No. M.S. 717bis), Dec. 31, 1965.

34. Cercospora pudicae Yen (Fig. 24)

Revue de Mycol. 29:234, 1964.

Leaf spots none or indistinct. Fruiting amphiphyllous, effuse, whitish on

both surfaces of the leaf. Stromata lacking. Fascicles rather dense or meagre,

emerging through the stomata. Conidiophores pale olivaceous brown, simple,

sometimes branched, erect or tortuous, O-1 septate, O-1 geniculate, medium

brown conidial scar at the rounded apex, 20.4-36 x 4.8-6um. Conidia hyaline

or subhyaline, long fusiform, 1-4 septate, straight or slightly curved, tip rounded

or subconic, base attenuate, 31.2-46 x 3-4.5yum.

Habitat: On leaves of Mimosa pudica (Leguminosae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No. 42), Mar. 15, 1964. |

Distribution: Burma, Cuba, India, Jamaica, Mauritius, Puerto Rico, Singa-

pore and Venezuela.

35. Cercospora sesamigena Yen & Lim (Fig. 25)

Cahiers du Pacifique 17: 103, 1973.

Cercospora and Allied Genera 165

Leaf spots amphigenous, brown, irregularly angular, scattered, 0.3-4mm in

diam., sometimes confluent on the edges of leaf. Fruiting amphiphyllous, in-

visible. Stromata absent or very rudimentary. Fascicles usually poor to meagre

(2-8 stalks), emerging through the stomata. Conidiophores pale brown, simple

or rarely branched, erect or flexuous, 0-8 septate, 0-3 geniculate, a large black

brown conidial scar at the rounded tip, several small brown scars laterally dis-

placed at the upper part, 25-185 x 4-5.5um. Conidia acicular, hyaline, straight

or slightly curved, 3-32 septate, tip acute, base truncate with a black brown scar

at the end, 75-490 x 3-4u.m.

Habitat: On leaves of Sesamum orientalis (Pedaliaceae), Singapore. leg.

G. Lim (No. S.U. 74), Feb. 12, 1972.

36. Cercospora thunbergiana Yen (Fig. 26)

Revue de Mycol. 30:198, 1965.

Leaf spots amphigenous, circular, brown with a dark brown border scattered,

2-7mm in diam., finally forming “‘shot-holes’’. Fruiting amphigenous, invisible.

Stromata lacking or very rudimentary, consisting of few brown cells. Fascicles

poor (2-7 stalks), emerging through the stomata. Conidiophores either coming

from fascicles or arising from the external hyphae, always simple, erect or slightly

tortuous, 2—9 septate, 0-2 geniculate, dark brown, paler towards the apex, a large,

black brown conidial scar at the truncate tip, 70-228 x 6-8.5um. Conidia acicular,

hyaline, straight or slightly curved, 7-34 septate, tip acute, base truncate with

a black brown scar at the end, 95-360 x 5—7um.

Habitat: On leaves of Thunbergia alata (Acanthaceae), Bukit Timah (Hwa

Chung College), Singapore, leg. Mrs. Shu-hsien Yen (No. 74), Aug. 3, 1964.

Note: This fungus differs from others (Cercospora thunbergiae Boed. and

Pseudocercospora thunbergiicola (Yen) Deighton by its amphigenous fruiting and

specially by its hyaline, acicular conidia.

37. Cercospora tithoniae Baker & Dale

Mycol. Papers 33:106, 1951.

Syn. Cercospora tithoniae Chidd., Mycopath. appl. Myc. 17:79, 1962.

Cercospora tithonicola (Chidd.) Yen, Revue de Mycol. 31: 144, 1966.

Leaf spots distinct, brownish, angular or irregular, vein-limited, small, scat-

tered, 0.3-2mm in diam., at last coalescing and covering the whole surface of

the leaf. Fruiting amphigenous, invisible. Stromata brownish, globose or irre-

gularly globose, 16-50ym in diam. Fascicles meagre to very dense (4-35 stalks),

emerging through the stomata. Conidiophores amphiphyllous, pale olivaceous

brown, unicolour simple, erect or slightly tortuous, 0-3 septate, 0-1 geniculate,

a large, black brown conidial scar at the rounded tip, 74-112 x 4-5um. Conidia

obclavato-cylindrical, pale olivaceous brown, straight or midly curved, 1-6 septate,

tip rounded, catenulate, then each end of the conidia holding a conidial scar,

base slightly attenuate and truncate with a black brown scar at the end, 33.6—85

x 3.6-Syum.

166 Gardens’ Bulletin, Singapore XX XIII (1980)

Habitat: On leaves of Tithonia diversifolia (Compositae), Botanic Gardens,

Singapore, leg. Mrs. Shu-hsien Yen (No. 113), Apr. 13, 1965.

Distribution: India and Singapore.

38. Cercospora triseptospora Yen (Fig. 27)

Revue de Mycol. 31:146, 1966.

Leaf spots none or indistinct, appearing as brownish areas. Fruiting hypo-

genous, rarely amphiphyllous, invisible. Stromata lacking or very small, black

brown, globose, 16-24um in diam. Fascicles poor to medium (2-11 stalks),

emerging through the stomata. Externai hyphae hypogenous, emerging through

the stomata, pale olivaceous brown, septate. branched, 1.5—2.54m in width. Coni-

diophores either coming from fascicles or arising from external hyphae, always

simple, pale brown, straight, cylindrical, 1-3 septate, O0u2 geniculate, several black

brown conidial scars displaced on the upper part, then the membrane becoming

denticulate, 25.5-70 x 3-4.5um. Conidia cylindrical or obclavato-cylindrical, pale

olivaceous brown, straight or slightly curved, with a finely granulate membrane,

3 septate, tip rounded, base slightly attenuate and subtruncate, 24-66 x 2.4—3um.

Habitat: On leaves of Hydrangea macrophylla (Hydrangeaceae), Katong,

Singapore, leg. Mrs. Shu-hsien Yen (No. 95), Dec. 18, 1964.

39. Cercospora volkameriae Speg.

Revista del Museo de la Plata 15:47, 1908.

Leaf spots amphigenous, angular, strictly vein-limited, reddish brown to gray

brown, scattered, 0.5-12mm in diam., more distinct on the upper side. Stromata

lacking or very rudimentary. Fruiting amphiphyllous, invisible. Fascicles poor

to medium (3-12 stalks), emerging through the stomata. Conidiophores always

simple, erect or flexuous, pale brown to olivaceous brown, I-11 septate, 0-5

geniculate, a large black brown scar at the rounded tip, 40-261 x 4-7.2ym.

Conidia acicular, hyaline, straight or slightly curved, 3-30 septate, apex acute,

base truncate with a black brown scar at the end, 82-278 x 3.6-7.2um.

Habitat: On leaves of Clerodendron fragrans (Verbenaceae), Changi, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 146), Dec. 31, 1966.

Distribution: Brazil and Singapore.

40. Cercospora zinniae Ell. & Mart.

J. Mycol. 1:20, 1885.

Syn. Cercospora atricinctata Head & Wolf, Mycologia. 3:14, 1911.

Cercospora zinniae Takah. & Yosh., Plant Protect (Shoku-butsu-boeki)

TELAT, Wed:

Leaf spots circular to angular or irregularly angular, more or less vein-limited,

at first brown, later becoming gray or white, with a dark brown or purplish

black margin, scattered, 2-14mm in diam.. but often coalescing and forming large

patches. Fruiting amphigenous, but mostly epiphyllous, visible as minute black

hairs under the hand lens. Stromata black brown, globose or subglobose, 20-42ym.

Fascicles poor to dense (2-20 stalks), emerging through the stomata. Conidio-

Cercospora and Allied Genera 167

phores pale brown to brown, simple, never branched, straight or tortuous, 0-10

septate, 14 geniculate, a large black brown conidial scar at the rounded tip

(sometimes tip shouldered), 48-300 x 4.8-7.2um. Conidia acicular or filiform,

hyaline, very long, curved or slightly undulate, 14-25 septate, tip obtuse, base

truncate with a terminal black brown scar, 150-328 x 3-33.6ym.

Habitat: On leaves of Zinnia el2gans (Convolvulaceae), Bukit Timah, Singa-

pore, leg. Lee Chih-shao (No. 14), Aug. 20, 1965.

Distribution: N. & S. America, Japan, Guam, Colombia, Guatemala, Puerto

Rico, San Domingo, Sa!vador, Singapore, Taiwan, Trinidad, Uganda and Venezuela.

41. Cercosporidium personatum (Berk. & Curt.) Deighton Mycol. Papers

112:71, 1967.

= Cladosporium personatum Berk. & Curt., Grevillea 3:106, 1875.

Cercospora personata (Bert & Curt.) Ell. & Ev., J. Mycol. 1:63, 1885.

Passalora personata (Berk. & Curt.) Shakil A. Khan & Mamal, Pakistan

J. sci. Res. ,13(4): 188, -1961.

Septogloeum arachidis Racib. Z. Pflanzenkr. 8:66, 1898.

Cercospora arachidis P. Henn., Hedwigia 41:18, 1902.

Perfect state: Mycosphaerella berkeleyii W.A. Jenkins, J. agric. Res.

56:330, 1938.

Leaf spots orbicular, dark brown, scattered, 0.5-3mm in diam. Fruiting

hypophyllous (never epiphyllous), visible as black pustules under hand _ lens.

Fascicles always. very dense. Stromiata well developed, pale brown, subglobose,

35-70nm in diam., but sometimes extending to 150um long. Conidiophores

strictly hypogenous, emerging through the stomata, olivaceous brown, simple,

erect or tortuous, continuous or | septate, with 0-3 small geniculations, a large

black brown conidial scar at the rounded tip, 35-65 x 5-7ym. Conidia fusiform

or obclavato-cylindrical, at times obclavate, straight or very slightly curved, 3-7

septate, tip rounded, base rounded or slightly subtruncate, with a large black

brown scar at the end, 42-73 x 6-8um.

Habitat: On leaves of Arachis hypogbea (Leguminosae), Kuala Lumpur,

Malaysia, leg. Jo-min Yen (No. 71139bis), Sept. 28, 1971.

Distribution: N. America, Argentina, Australia, Brazil, China (Kwangtung),

Congo, Hong-Kong, India, Jamaica, Malaysia, Nepal, New Guinea, Nigeria, W.

Pakistan, Paraguay, Philippines, Sarawak, Sierra Leone, Singapore, Trinidad and

Turkey.

Note: According to Chupp (1954), Cercospora arachidicola Hori and

Cercospora personata (Berk. & Curt.) Ell. & Ev. (= Cercosporidium personatum

(Berk. & Curt.) Deighton) always have amphiphyllous fruiting. But this material

always presents hypogenous fruiting.

168 Gardens’ Bulletin, Singapore XX XIII (1980)

42. Pseudocercospora abacopteridicola (Yen & Lim) Yen, comb. nov. (Fig.

7, A B).

= Cercospora abacopteridicola Yen & Lim, Cahiers du Pacifique 17:97,

ie gee

Leaf spots amphigenous, brown, with an indefinite border, at first vein-

limited, later confluent. Fruiting hypogenous, invisible. Stromata lacking. Fas-

cicles absent. External hyphae hypophyllous, emerging through the stomata, pale

olivaceous brown, septate, branched, 1.5-2.5um in width. Conidiophores hypo-

phyllous, simple, erect, 0-1 septate, not geniculate, tip rounded or slightly attenuate,

conidial scars inconspicuous, 6-16 x 2.5-3um. Condia pale olivaceous brown,

cylindrical or obclavato-cylindrical, slightly curved, 1-7(-9) septate, tip rounded,

base truncate, 32-80 x 2-3yum.

Habitat: On leaves of Abacopteris urophylla (Pteridophyte), Singapore, leg.

G. Lim (No. 7l1bis), Mar. 3, 1972.

43. Pseudocercospora abelmoschi (Ell. & Ev.) Deighton Mycol. Papers

140:139, 1976.

= Cercospora abelmoschi Ell. & Ev., J. Inst. Jamaica 1:347, 1893.

Syn. Cercospora hibisci-manihotis P. Henn., Hedwigia 43:146, 1904.

Cercospora hibisci-cannabini Sawada, Descr. Cat. Formosan Fungi 2: 153,

1922:

Leaf spots indistinct or small, angular, dark gray, more distinct on lower

side of the leaf, vein-limited, sometimes extending over the entire leaf surface.

Fruiting amphigenous, invisible. Stromata lacking or small, brown, globose,

21-32ym in diam. Fascicles poor to medium (2-15 stalks), emerging through

the stomata. Conidiophores olivaceous brown, simple or branched, erect or

slightly flexuous, not geniculate, 0-3(-4) septate, tip rounded or attenuate and

truncate, 32-100 x 4.8-6.m. Conidia pale olivaceous brown, attenuate on both

ends, then appearing as long fusiform, usually curved, crescent, 1-7 septate

(generally 3-5), tip obtuse, base attenuate and truncate, 36-100 x 4.5—6ym.

Habitat: On leaves of Hibiscus esculentus (Malvaceae), Bukit ‘Timah,

Singapore, leg. Koh Choy-kuan, May 25, 1969.

Distribution: N.C. & S. America, Australia, Japan, Singapore and Taiwan.

44. Pseducercospora acalyphigena (Yen) Yen, comb. nov. (Fig. 28)

= Cercospora acalyphicola Yen, Revue de Mycol. 31:109, 1966.

Leaf indistinct, at first irregularly subcircular, yellowish with indefinite border,

then confluent, yellowish brown or brownish gray. Fruiting amphigenous, invisible.

Stromata lacking or consisting of a few olivaceous brown cells. Fascicles poor

to medium (2-12 stalks). Conidiophores amphiphyllous, emerging through the

stomat1, pale oliviceous brown, erect or slightly flexuous, generally simple,

sometimes branched, 0-3 septate, O-1 geniculate, constricted at the septa, tip

attenuate and truncate, the base cells much larger (8.5-9.6um in diam.), 18-50

x 3.6-5u4m. Conidia cylindrical or obclivato-cylindrical nale olivaceous brown,

straight, sometimes slightly curved, 3-10 septate, tip conic, base attenuate and

truncate, 34-135 x 2.5-4.8ym.

Cercospora and Allied Genera 169

Habitat: On leaves of Acalypha indica (Euphorbiaceae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No. 40), May 16, 1964.

Note: Pseudocercospora acalyphicola (Petr.) Deighton differs from _ this

fungus by its shorter and hyaline conidia.

45. Pseudocercospora allamendae (Yen) Yen, comb. nov. (Fig. 29, A-F)

= Cercospora allamendae Yen, Revue de Mycol. 30: 166, 1965.

Leaf spots none or indistinct, sometimes appearing as indistinct gray black

areas. Fruiting hypophyllous, invisible. Stromata lacking. External hyphae

emerging through the stomata, hypogenous, pale olivaceous brown, septate,

branched, 2-2.5um in width. Conidiophcres hypogenous, in fascicles poor to

meagre (2-8 stalks), may be classified into two kinds: A, conidiophores coming

from fascicles, emerging through the stomatal opening, olivaceous brown, paler

and more narrow towards the tip, simple or branched, erect or tortuous, 0-2

geniculate, 14 septate, 21.640 x 3.5-4..8um, conidial scars inconspicuous; B,

conidiophores as single branches from procumbeni threads, simple, olivaceous

brown, 0-1 geniculate, 0-2 septate, tip rounded or slightly attenuate, condial scars

indistinct, 9.6-22 x 3-3.6um. Conidia cylindrical or obclavato-cylindrical, pale

olivaceous brown, straight or slightly curved, 2-5 septate, tip conic rounded, base

subtruncate, 31.2-54 x 2.4~-3.6um.

Habitat: On leaves of Allamanda cathartica (Apocynaceae), Katong, Singa-

pore, leg. Mrs. Shu-hsien Yen, (No. 22), Apr. 25, 1964.

Note: Pseudocercospora byliana (Syd.) Yen is different from this fungus

by its very developed stromata and its very dense fascicles.

46. Pseudocercospora aranetae (Borl. & Rold.) Yen, comb. nov. (Fig. 30)

= Cercospora aranetae Borl. & Rold., Araneta J. Agr. 11:181, 1964.

Leaf sports amphigenous, distinct, angular to irregular, small, with an in-

definite border, more or less vein-limited, scattered, 14mm in diam., sometimes

confluent and forming large patches. Fruiting amphiphyllous, but more abundant

on the lower side, appearing as gray or dark gray velvet on the lower surface

of the spots. Stromata absent or small, brown, subglobose, 20-30.m in diam.

Fascicles poor to dense (4-26 stalks), emerging through the stomata. Conidio-

phores amphigenous, chiefly hypophyllous, olivaceous brown, erect or flexuous,

simple or branched, 1-4 septate, with 0-1 geniculate, tip attenuate, 39.6-68.4 x

4-6.m. Conidia obclavate, pale olivaceous brown, straight or slightly curved,

3-6 septate, tip obtuse, base attenuate and truncate, 42-62.4 x 3.6-4.8um.

Habitat: On leaves of Psophocarpus tetragonolobus (Leguminosae), Jurong,

Singapore, leg. Jo-min Yen (No. M.S. 717), Dec. 31, 1965.

Distribution: Philippines and Singapore.

47. Pseudocercospora asystasiae (Yen) Yen, comb. nov. (Fig. 31)

= Cercospora asystasiae Yen, Revue de Mycol. 32:178, 1967a.

170 Gardens’ Bulletin, Singapore XX XIII (1980)

Leaf spots distinct, circular, but more clear on the upper surface, yellowish

brown or brown, with a black brown border, scattered, 3-10mm in diam., some-

times confluent. Fruiting amphigenous, invisible. Stromata lacking or very small,

dark brown, globose, 14-30ym in diam. Fascicles poor to dense (3-16 stlks),

emerging through the stomata. Conidiophores amphiphyllous, straight or slightly

flexuous, simple or branched, olivaceous brown, unicolour, 0-1 geniculate, O—5

septate, tip rounded or truncate, conidial scars inconspicuous, 24-126 x 3.6—5ym.

Conidia obclavate, but cylindrical for the short ones, pale olivaceous brown,

straight or mildly curved, 3-8 septate, tip obtuse or subrounded, base slightly

attenuate and truncate, 37.2-110 x 3.6—Sum.

Habitat: On leaves of Asystasia coromandeliana (Acanthaceae), Botanic

Gardens, Singapore, leg. Jo-min Yen (No. M.S. 731), Jan. 5, 1966.

48. Pseudocercospora bauhiniae (H. & P. Syd.) Deighton Mycol. Papers

140: 140, 1976.

= Cercospora bauhiniae H. & P. Syd. Ann. Mycol. 12:202, 1914.

Leaf spots circular to anguglar, 2-9mm in diam., grayish brown to pale

brown, with a dark brown line margin. Fruiting chiefly hypophyllous. Stromata

dark brown, globose to subglobose, 15—-40um in diam. Fascicles dense to very

dense. Conidiophores pale brown to medium brown, paler and more narrow

towards the tip, simple, 0-1 septate, straight to mildly curved, not geniculate,

unthickened small conidial scar at the bluntly rounded tip, 10-20 x 3-4.5ym,

sometimes as long as 40um. Conidia subhyaline to pale olivaceous brown, cbcla-

vate, short ones may be cylindrical, almost straight, 3-9 septate, tip subobtuse,

base obconically truncate, 30-125 x 2.5—Syum.

Habitat: On leaves of Bauhinia malabarica (Leguminosae), Botanic Gardens,

Singapore, leg. Mrs. Shu-hsien Yen (No. 114), May 13, 1965.

Distribution: Colombia, Ethiopia, Gold Coast, Singapore, Philippines,

Transvaal and Venezuela.

49. Pseudocercospora byliana (Syd.) Yen, comb. nov.

= Cercospora byliana Syd., Ann. Mycol. 22:433, 1924.

Leaf spots angular to irregular, gray to reddish tinge with narrow reddish

brown line border, 2-8mm in diam. Fruiting amphigenous, more abundant on

the upper side, appearing as minute black pustules in the centre part of the spots.

Stromata globular to irregular, 50-85y4m in length, pale brown. Fascicles dense

to very dense. Conidiophores pale olivaceous brown, paler towards the tip, simple,

never branched, 0-1 septate, not geniculate, 8-24 x 3-4um. Conidia obclavate,

pale olivaceous brown, straight or slightly curved, multiseptate, tip subobtuse,

base obconically truncate, 40-12 x 3.5-Sym.

Habitat: On leaves of Allamanda cathartica (Apocynaceae), Katong, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 45), May 19, 1964.

Distribution: Singapore, Transvaal and Tzaneen.

Cercospora and Allied Genera 171

50. Pseudocercospora cassiae-alatae (Yen & Lim) Yen, comb. nov. (Fig. 32,

D et E)

= Cercospora cassiae-alatae Yen & Lim, Cahiers du Pacifique 17:99,

1973.

Leaf spots amphigenous, irregularly circular, black brown with an indefinite

border, scattered, 15mm in diam. Fruiting amphiphyllous, invisible. Stromata

lacking. Fascicles absent. External hyphae amphigenous, emerging through the

stomata, septate, branched, pale olivaceous brown, 1.5—2.5um in width. Conidio-

phores arising from the external hyphae, pale olivaceous brown, simple, erect or

slightly curved, 0-6 septate, 0-2 geniculate, apex irregularly rounded or truncate,

conidial scars inconspicuous, 3-45 x 2.5-3.54m. Conidia cylindrical, very thin,

pale olivaceous brown, usually curved, 1-10 septate, tip rounded or obtuse, base

slightly attenuate and truncate, 15-90 x 1.5—2um.

Habitat: On leaves of Cassia alata (Leguminosae), Singapore, ieg. G. Lim

Pha. 0. 66), Jan, 5.1971.

Note: This fungus differs from others by only the presence of external

hyphae and solitary conidiophores.

51. Pseudocercospora cassiigena (Yen & Lim) Yen, comb. nov. (Fig. 32,

A-C).

= Cercospora cassiigena Yen & Lim, Cahiers du Pacifique 17:98, 1973.

Leaf spots angular to irregular, brown, more or less vein-limited, scattered

1-4mm in diam., sometimes confluent. Fruiting amphiphyllous, invisible. Stro-

mata pale brown to brown, globose, 25—40um in diam. Fascicles dense to very

dense, emerging through the rupture of epidermis. Conidiophores pale brown,

always simple, erect, continuous, not geniculate, apex rounded or slightly atten-

uate and truncate, conidial scars inconspicuous, 6-11 x 2-3u4m. Conidia pale

olivaceous brown, obclavate, or obclavato-cylindrical, straight or slightly curved,

strictly 3 septate, tip rounded, base slightly obconically truncate, 22-36 x 2-2.5um.

Habitat: On leaves of Cassia alata (Leguminosae), Singapore, leg. G. Lim

(No. S.U. 66bis), Jan. 5, 1971.

Note: This species is different from others by its continuous conidiophores

and its strictly 3 septate conidia.

52. Pseudocercospora centrosematicola (Yen & Lim) Yen, comb. nov. (Fig.

= Cercospora centrosematicola Yen & Lim, Cahiers du Pacifique

PY 5108,..1975.

Leaf spots irregularly angular, brown, with an indefinite yellowish border,

more distinct on the upper side, scattered, 14mm in diam. Fruiting amphi-

genous, invisible. Stromata usually forming inside the epidermal cells, sometimes

under the epidermal cells, dark brown, globose or irregularly globose, 20-45um

in diam. Fascicles dense, emerging through rupture of the epidermal cells. Ex-

ternal hyphae hypophyllous, emerging through the stomata, pale olivaceous brown,

septate, branched, 1.5-2.54m in width. Conidiophores may be divided into two

72 Gardens’ Bulletin, Singapore XX XIII (1980)

kinds: A, conidiophores arising from fascicles, amphigenous, olivaceous brown,

simple when young, branched when old, continuous or 1-7 septate, 0-3 geniculate,

apex irregularly rounded or shortly attenuate, a small unthickened conidial scar

at the tip, 12-65 x 3.5-4um; B, conidiophores arising from the external hyphae,

hypogenous, olivaceous brown, always simple, erect or slightly curved, generally

continuous, sometimes | septate, apex slightly attenuate and truncate, 6-30 x

3.5-4um. Conidia pale olivaceous brown, filiform or obclavato-cylindrical, straight

or slightly curved, apex subobtuse, 3-12 septate, base obconically truncate, 50-127

X 2-3 um.

Habitat: On leaves of Centrosema pubescens (Leguminosae), Sembawang,

Singapore, leg. ‘G-’Lim ‘(Nor SU. 72).4 Deco sae

Note: Cercospora cylindrospora Stev. & Solh. and C. centrosemae Chupp

& Muller are different from this species by their strictly hypophyllous fruiting.

Pseudocercospora bradburyae (Young) Deighton and Cercospora clitoria Atk.

differ from this fungus by their strictly epiphyllous fruiting.

53. Pseudocercospora chrysanthemicola (Yen) Deighton (Fig. 34)

Mycol. Papers 140:141, 1976.

= Cercospora chrysanthemicola Yen, Revue de Mycol. 29:215, 1964.

Leaf spots circular, sometimes oblong up to 30mm, generally scattered,

3-10mm in diam., at first yellowish on upper side and grayish brown on the

lower side, then becoming dark brown with brown margin on the under surface.

Fruiting amphiphyllous invisible. Stromata amphigenous, substomatal, dark

brown, globular, 22—60um in diam. Fascicles poor to dense (2-20 stalks), filling

stomatal opening. Conidiophores amphiphyllous, simple, not branched, oliva-

ceous brown, cylindrical, erect or tortuous, an unthickened scar at the rounded

tip, O-S septate, 0-2 geniculate, 20-110 x 3.6-6um. Conidia pale olivaceous

brown, obclavate or obclavato-cylindrical, straight or slightly curved, 1-5 septate,

tip rounded, base obconically truncate, 31.2-92.4 x 3.6-Sum.

Habitat: On leaves of Chrysanthemum sinense (Compositae), Katong,

Singapore, leg. Mrs. Shu-hsien Yen (No. 17), Apr. 10, 1964.

Note: Cercospora chrysanthemi Heald & Wolf differs from this fungus

by its hyaline and acicular conidia.

54. Pseudocercospora columnaris (Ell. & Ev.) Yen, comb. nov.

= Cercospora columnaris Ell. & Ev., Proc. Acad. Nat. Phila. 46: 380,

1894.

Syn. Isariopsis griseola Sacc., Michelia 1:273, 1878.

Graphium laxum Ellis, Torrey Bot. Club 8: 64-66, 1881.

Arthrobotryum puttemansii P. Henn., Hedwigia 417309) L0ze

_ Cercospora stuhlmanni Py Henn., Bot. Jahrb.’ v. Engler, 33: 40, 1904.

is tiaies suathe griseola (S1cc.) Feri?! Ann. Mycol. aA 273, 1909.

Cercospora and Allied Genera 173

Leaf spots distinct, angular brown to grayish brown, strictly vein-limited,

2-Smm in diam. Fruiting amphigenous or only hypophyllous, often effuse, appear-

ing as short black brown hairs or a dark gray velvet. Stromata none or very

rudimentary. Fascicles dense to very dense. Conidiophores olivaceous brown,

simple, cylindrical, or filamentous, 1-9 septate, not geniculate, tip rounded or

attenuate and truncate, conidial scars inconspicuous, 220-410 x 3.6—-6.m. Conidia

crescent, attenuate at both ends, pale olivaceous brown, generally 3 septate (some-

times 5—6 septate), with constrictions at septa, apex rounded-conic, base obconico-

truncate, 30-60 x 4-7yum.

Habitat: On leaves of Phaseolus vulgaris (Leguminosae), Jurong, Singapore,

leg. Lin Shu-ling, Dec. 10, 1968.

Distribution: Africa, N. America, China, Europe, Hawaii, Japan, Puerto

Rico, Singapore and Trinidad.

55. Pseudocercospora cordiicola (Yen) Yen, comb. nov. (Fig. 35)

= Cercospora cordiicola (Yen) Yen, Revue de Mycol. 32:182, 1967a.

Syn. Cercospora cordiae Yen, Revue de Mycol. 29:216, 1964.

non Cercospora cordiae Chupp, Monogr. Univ. P.R. Ser. B, 2:245, 1934.

Leaf spots indistinct, sometimes appearing as small angular dark brown

speckles. Fruiting amphiphyllous, invisible. Stromata lacking or very rudimen-

tary. Fascicles poor to meagre (2-9 stalks), filling stomatal openings. External

hyphae pale olivaceous brown, emerging through the stomata, septate, branched,

2—2.5u4m in width. Conidiophores may be classified into two kinds: A, conidio-

phores coming from fascicles, simple or branched, tortuous, olivaceous brown,

paler and more narrow towards the tip, 1-3 septate, 0-2 geniculate, with un-

thickened conidial scar at the tip, 22.4-32.4 x 3.6-4.8um; B, conidiophores arising

from external hyphae, olivaceous brown, 0-2 septate, 0-1 geniculatee, 13-24 x

3.6-4.8um. Conidia filiform or obclavato-acicular, hyaline, curved, 4-8 septate,

tip acute, base obconically truncate, 79.2-126 x 1.5—2.4um.

Habitat: On leaves of Cordia cylindristachya (Boraginaceae), Katong,

Singapore, leg. Mrs. Shu-hsien Yen (No. 28), Apr. 24, 1964.

56. Pseudocescorpora crotalaricola (Yen) Yen, comb. nov. (Fig. 36)

= Cercospora crotalaricola Yen, Revue de Mycol. 29:220, 1964.

Leaf spots none or indistinct. Fruiting amphigenous, invisible. Stromata

lacking. Fascicles poor to meagre (2-8 stalks), filling stomatal openings. External

hyphae pale olivaceous brown, also filling stomatal openings, septate, branched,

2.4-4um in width. Conidiophores may be divided into two kinds: A, conidio-

phores coming from fascicles, straight or tortuous, single or branched, olivaceous

brown, 1-5 septate, 0-2 geniculate, with constrictions at septa, tip rounded,

thickened conidial scars inconspicuous, 36-149 x 4.8-7.8um; B, conidiophores

as single branches from procumbent threads, olivaceous brown, 1-3 septate, not

geniculate, tip rounded, 36-48 x 4.5-7um. Conidia acicular, cylindrical or

obclavato-cylindrical, pale olivaceous brown, straight or undulate, 5-12 septate,

tip conic or rounded, base obconically truncate, 10-134 x 3.5-Sym.

174 Gardens’ Bulletin, Singapore XX XIII (1980)

Habitat: On leaves of Crotalaria striata (Leguminosae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No. 33), Apr. 25, 1964.

57. Pseudoercospora cruenta (Sacc.) Deighton (Fig. 11)

Mycol. Papers 140:142, 1976.

= Cercospora cruenta Sacc., Michelia 2:149, 1880.

(Mycosphaerella cruenta (Sacc.) Latham, Mycologia 26:516, 1934).

Cercospora phaseolorum Cooke, Grevillea 12:30, 1883.

Cercospora vignae Ell. & Ev., J. Mycol. 3:19, 1887.

Cercospora dolichi Ell. & Ev., J. Mycol. 68:72, 1889.

Cercospora vignae Raciborski, Zeit. Pflanzenkr. 3:66, 1898.

Cercospora lusoniensis Sacc., Ann. Mycol. 12:314, 1914.

Cercospora raciborskii Mats. & Nag., J. Plant Protect. (Byochugai-zashi)

18:714-722, 1931.

Cercospora vignae-sinensis Tai & Wei, Sinensia 4:126, 1933.

Cercospora neovignae Yamam., Trans. Sapporo Nat. Hist. Soc. 13: 142,

1934.

Cercospora vignae-sinensis Sawada, Descr. Cat. Formosan Fungi 8:125,

1934.

Leaf spots circular to irregular, 1-l0mm in diam., yelowish to rusty brown

or blood red, sometimes concentrical with gray centre and somewhat darker

margin, often confluent. Fruiting amphigenous, but mainly hypophyllous, some-

times distinctly effuse. Stromata rudimentary or small, filling stomatal openings,

olivaceous brown. Fascicles medium to dense, emerging through the stomata.

Condiophores simple, straight or subflexuous, pale olivaceous brown, 0-3 septate,

1-3 geniculate, an unthickened small conidial scar at the conic tip, 10-75 x 3—Syum.

Conidia cylindrical or oblcavato-ylindrical, pale olivaceous brown, straight or

slightly curved, 4-14 septate, tip rounded, base sharply obconic and subtruncate,

25-50 x 2-5ym.

Habitat: On leaves of Vigna sinensis (Leguminosae), Taiping, Singapore, leg.

Jo-min Yen, Dec. 30, 1965; on V. sinensis, Jurong, Singapore, leg. Lin Shu-ling,

Dec. 10, 1968; on V. sinensis, Singapore, leg. G. Lim, Apr. 11, 1969.

Distribution: N. & S. America, Ceylon, China, Europe, Japan and Singapore.

58. Pseudocercospora daturina (Yen) Deighton (Fig. 37)

Mycol. Papers 140: 143, 1976.

= Cercospora daturina Yen, Revue de Mycol. 30:169, 1965.

Leaf spots none or indistinct. Fruiting amphigenous, invivsible. Stromata

lacking. Fascicles poor to meagre (2-8 stalks), emerging through the stomata.

Conidiophores amphiphyllous, pale olivaceous brown, single or branched, erect

Cercospora and Allied Genera 175

or tortuous, 1-4 septate, 1-3 geniculate, an unthickened small conidial scar at

the attenuate tip, 30-80 x 4-6um. Conidia cylindrical, pale olivaceous brown,

straight or slightly curved, 3-10 septate, tip subconic, base obconic truncate,

51-123 x 3.5-Sum.

Habitat: On leaves of Datura alba (Solanaceae), Botanic Gardens, Singapore,

leg. Mrs. Shu-hsien Yen (No. 70), Aug. 4, 1964.

59. Pseudocercospora delonicis (Yen) Yen, comb. nov. (Fig. 38)

= Cercospora delonicis Yen, Revue de Mycol. 29:223, 1964.

Leaf spots none or indistinct. Fruiting amphigenous, sparingly effuse. Stro-

mata lacking. External hyphae amphigenous, filling stomatal openings, pale oliva-

ceous brown, septate, branched, 2-3.6u.m in width. Conidiophores as solitary

branches from procumbent threads, olivaceous brown, cylindrical, 0-3 septate,

0-1 geniculate, tip rounded, conidial scars inconspicuous, 6-31 x 2.5—3.5um.

Conidia cylindrical, pale olivaceous brown, straight or slightly curved, 1-6 septate,

tip rounded, base attenuate and truncate, 40.5-61.2 x 2.5-4.5um.

Habitat: On leaves of Delonix regia (Leguminosae), Katong, Singapore, leg.

Mrs. Shu-hsien Yen (No. 41), May 15, 1964.

60. Pseudocercospora ervatamiae (Yen & Lim) Yen, comb. nov. (Fig. 39)

= Cercospora ervatamiae Yen & Lim, Bull. Soc. Mycol. Fr. 86:745,

1970b.

Leaf spots irregular, brown to dark brown, with an indefinite yellowish border,

scattered, 0.5-5mm in diam., sometimes confluent. Fruiting amphiphyllous,

appearing as black minute pustules under hand lens. Stromata well developed,

black brown, globose, 15-40 (—60)um in diam. Fascicles dense to very dense,

emerging through the stomata. Conidiophores pale olivaceous brown. very short,

simple, sometimes branched, erect, rarely with one geniculation, continuous, some-

times one septate, apex attenuate and truncate, conidial scars inconspicuous,

12-25 x 3-4ym. Conidia cylindrical, pale olivaceous brown, straight or slightly

curved, 1-6 spetate, apex rounded, base attenuate and truncate, 28-80 x 34m.

Habitat: On leaves of Ervatamia coronaria (Apocynaceae), Singapore, leg.

G. Lim, Mar. 11, 1970.

Note: Pseudocercospora tabernaemontanae (H. & P. Syd.) Deighton some-

times resembles this fungus, but it is different from this species by its big “‘shot-

hole” leaf spots and especially by its branched and septate conidiophores.

61. Pseudocercospora eupatorii-formosani (Saw.) Yen, comb. nov. (Fig. 16,

A-C)

= Cercospora eupatorii-formosani Sawada, Taiwan Agr. Res. Inst.

86:169. 1943.

Leaf spots brown, angular, vein-limited, small, scattered, 24mm in diam.,

rarely confluent. Fruiting amphiphyllous, invisible. Stromata black brown, glo-

bose or subglobose, 18-57.5.m in diam. Fascicles dense to very dense (10-30

stalks), emerging through the stomata. Conidiophores amphigenous, short, pale

176 Gardens’ Bulletin, Singapore XX XIII (1980)

brown, simple, slightly flexuous, continuous, rarely one-septate, tip rounded or

shortly attenuate and truncate, conidial scars inconspicuous, 18-26.5 x 3-4.8um.

Conidia cylindrical or filiform, pale olivaceous brown, slightly curved, 3-10 a

tate, tip conic, base slightly obconic- truncate, 34-110 x 2.5-3.5um.

Habitat: On leaves of Eupatorium odoratum (Compositae), Ipoh, Malaysia,

leg. Jo-min Yen (No. M.S. 728),"Dec*" 2h” 1965:

Distribution: Malaysia and Taiwan.

62. Pseudocercospora fici-chartaceae (Yen & Lim) Yen, comb. nov. (Fig. 40)

= Cercospora fici-chartaceae Yen & Lim, Cahiers du Pacifique 14:97,

1970a.

Leaf spots circular with an indefinite border, orange brown with a black

brown centre on the upper side, pale brown to yellowish brown on the lower

side, 2-8mm in diam. Fruiting amphigenous, but chiefly epiphyllous. Stromata

epigenous and endogenous within an epidermal cell, globose or subglobose,

21.5-40um in diam. Fascicles poor to dense (2-20 stalks), emerging through

rupture of epidermal cells. Conidiophores usually epigenous, but sometimes

hypogenous and emerging through the stomata, simple, erect, dark brown, 2-7

septate, not geniculate, tip rounded and paler, conidial scars indistinct, 60-124

x 4.5-6.m. Conidia obclavate, sometimes _obclavato- cylindrical, pale olivaceous

brown, straight or curved, at times crescent, 3-5 septate, tip rounded, base obconic-

truncate, 39.5- 70 x 3.5-Syum.

Habitat: On leaves of Ficus chartacea (Moraceae), Bukit Timah, Singapore,

leg. G. Lim, Apr. 4, 1969.

63. Pseudocercospora formosana (Yamam.) Deighton (Fig. 41)

Mycol. Papers 140:144, 1976.

= Cercospora formosana Yamam., J. Soc. Trop. Agr. 6:600, 1934.

Syn. Cercospora lantana-aculeatae Yen, Revue de Mycol. 31:124, 1966.

Leaf spots small, angular, vein-limited, black brown on the upper side,

brown on the lower side, scattered, 0.5-2mm in diam. Fruiting hypophyllous,

invisible. Stromata lacking. External hyphae emerging through the stomata,

pale olivaceous brown, septate, branched, 1.5—2.54m in width. Conidiophores

may be divided into two kinds: A, conidiophores arising from poor fascicles

(2-6 stalks), simple or branched, pale olivaceous brown, straight or flexuous,

0-2 geniculate, 1-3 septate, sometimes with constrictions at septa, tip attenuate

and truncate, sometimes shouldered, conidial scars inconspicuous, 12-36 x 3.5—5um;

B, conidiophores as solitary branches from external hyphae, not branched, pale

olivaceous brown, straight or slightly flexuous, 0-2 septate, 0-2 geniculate, tip

attenuate and truncate, 5-28 x 2.5-3.6ym. Conidia filiform, pale olivaceous

brown, slightly curved, 2-6 septate, tip obtuse, base obconic-truncate, 36-82 x

2—3.m.

Habitat: On leaves of Lantana aculeata (Verbenaceae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No. 86), Sept. 20, 1964.

Distribution: Singapore and Taiwan.

Cercospora and Allied Genera 177

64. Pseudocercospora globosae (Yen) Deighton (Fig. 42)

Mycol. Papers 140: 144, 1976.

= Cercospora globosae Yen, Revue de Mycol. 29:224, 1964.

Leaf spots none or indistinct. Fruiting amphigenous, effuse, dark gray.

Stromata amphigenous, brown, in shape of dumb-bell, each end globular, 15.5-

33.54m in diam. Fascicles dense, filling stomatal openings. Conidiophores simple

or branched, tortuous, olivaceous brown, 0-3 geniculate, 0-6 septate, condial

scars inconspicuous, tip attenuate and truncate, 31-111.5 x 3.5-6u.m. Conidia

acicular or filiform, pale olivaceous brown, straight or curved, 2-12 septate, tip

conic, base obconic-truncate, 62.5-150 x 2.5-3.5um.

Habitat: On leaves of Gomphrena globosa (Amaranthaceae), Katong, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 19), Apr. 20, 1964.

Note: Cercospora gomphrenicola Speg. resembles this fungus by the in-

distinct leaf spots, but differs from it by its mostly one septate and constricted

conidia.

65. Pseudocercospora hiratsukana (Tog. & Kats.) Deighton

Mycol. Papers 140:34, 1976.

= Cercospora hiratsukana Tog. & Kats., J. Jap. Bot. 28:286, 1953.

Leaf spots subcircular or irregular, brown with an indefinite border, fre-

quently confluent and forming large patches. Fruiting amphigenous, but chiefly

hypophyllous. Stromata amphigenous, olivaceous brown, globose, 24-64um in

diam. Fascicles dense to very dense. Conidiophores very short, pale olivaceous

brown, continuous, tip attenuate and truncate, conidial scars unthickened, 6-10

x 2.5-3um. Conidia subhyaline or very pale olivaceous brown, cylindrical,

straight or midly curved, 2—5 septate, tip rounded or subobtuse, base slightly

attenuate and truncate, 31-62 x 1.5—2.5um.

Habitat: On leaves of Dioscorea sp. (Dioscoreaceae), Botanic Gardena,

Singapore, leg. Mrs. Shu-hsien Yen (No. 54bis), Jun. 7, 1964.

Distribution: Japan and Singapore.

66. Pseudocercospora ipomoeae-purpureae (Yen) Yen, comb. nov. (Fig. 43)

= Cercospora ipomoeae-purpureae Yen, Revue de Mycol. 30:173, 1965.

Leaf spots circular to angular, dark brown, more distinct on the upper side,

scattered, 0.5-3mm in diam. Fruiting amphiphyllous, but more abundant on

the lower surface. Stromata amphigenous, small, brown or dark brown, globose,

18-40um in diam. Fascicles medium to very dense (6-30 stalks), emerging through

the stomata. Conidiophores amphiphyllous, simple or branched, straight or flex-

uous, olivaceous brown or pale olivaceous brown, 0-1 geniculate, 0-2 septate,

with constrictions at the septa, small unthickened condial scars at the rounded

or attenuate and truncate tip, 22-35 x 3.5-4.8um. Conidia cylindrical or obclavato-

cylindrical, pale olivaceous brown, straight or slightly curved, 3-11 septate, tip

rounded or subconic, base obconic-truncate, 40-120 x 3-4um.

178 Gardens’ Bulletin, Singapore XX XIII (1980)

Habitat: On leaves of Ipomoea purpurea (Convolvulaceae), Botanic Gardens,

Singapore, leg. Mrs. Shu-hsien Yen (No. 72), Apr. 4, 1964.

67. Pseudocercospora ixoricola (Yen) Yen, comb. nov. (Fig. 44)

= Cercospora ixoricola Yen, Revue de Mycol. 31:119, 1966.

Leaf spots distinct, angular or irregular, with an indefinite border, reddish

brown on upper side, yellowish brown on lower side, centre whitish and zonate,

frequently vein-limited, scattered, 3-14mm in diam., sometimes confluent. Fruit-

ing hypophyllous, invisible. Stromata lacking or very poor. Fascicles poor to

meagre (2-8 stalks), emerging through the stomata. Conidiophores hypophyllous,

brown, unicolour, simple or branched, erect or slightly flexuous, 0-2 geniculate,

3-5 septate, with constrictions at the septa, tip rounded or attenuate-truncate,

conidial scars inconspicuous, 38.5—78 x 3.5-5um. Conidia obclavate, pale oliva-

ceous brown, straight or slightly curved, 3-10 septate, tip obtuse to acute, some-

times rounded, base obconic-truncate, 44-138 x 3.5—-Sum.

Habitat: On leaves of /xora javanica (Rubiaceae), Katong, Singapore, leg.

Mrs. Shu-hsien Yen (No. 117), May 12, 1965.

Note: Pseudocercospora ixorae (Solh.) Deighton differs from this species

by its amphiphyllous fruiting and cylindrical conidia.

68. Pseudocercospora jasminicola (Muller & Chupp) Deighton

Mycol. Papers 140:74, 1976.

= Cercospora jasminicola Muller & Chupp, Arch. Inst. Biol. Veg. Rio

de Janeiro 3:93, 1936.

Leaf spots circular, brown with a reddish brown border, more distinct on

the upper side, scattered, 1-3mm in diam., sometimes coalescing and forming

large patches. Fruiting amphigenous, but chiefly hypogenous, visible as minute

black pustules under hand lens. Fascicles poor or mildly dense (3-16 stalks),

emerging through the stomata. Conidiophores olivaceous brown, simple, never

branched, straight or flexuous, 1-2 geniculate, 2-6 septate, with constrictions at

septa, tip attenuate and truncate, sometimes shouldered, conidial scars indistinct,

5-50 x 2-4um. Conidia obclavato-cylindrical, pale olivaceous brown, straight

or slightly curved, 1-4 septate, tip obtuse, base obconically subtruncate, 24-60 x

2—3ym.

Habitat: On leaves of Jasminum grandiflorum (Oleaceae), Katong, Singa-

pore, leg. Jo-min Yen (No. M.S. 732), Jan. 18, 1966.

Distribution: Brazil, India, Singapore and Taiwan.

69. Pseudocercospora jatrophae-curcas (Yen) Deighton (Fig. 45)

Mycol. Papers 140: 146, 1976.

= Cercospora jatrophae-curcas Yen, Revue de Mycol. 30:176, 1965.

Cercospora and Allied Genera 179

Leaf spots none or indistinct, fruiting amphiphyllous, but more abundant

on the lower surface, invisible. Stromata lacking or very small, consisting of

few dark brown cells. Fascicles poor to dense (2-20 stalks), emerging through

the stomata. External hyphae pale brown, filling the stomatal openings, septate,

branched, 2-2.5um in width. Conidiophores either coming from fascicles or

arising from external hyphae, pale brown, simple or branched, generally tortuous,

0-4 septate, 0-3 geniculate, with constrictions at septa, tip attenuate and truncate,

(S—) 18-48 x 3-S5ym. Conidia cylindrical, pale olivaceous brown, slightly curved,

3-7 septate, tip rounded, base obconic-truncate, 42-102 x 2.5-3.5um.

Habitat: On leaves of Jatropha curcas (Euphorbiaceae), in Botanic Gardens,

Singapore, leg. Mrs. Shu-hsieri Yen (No. 55), Jun. 7, 1964.

70. Pseudocercospora katongensis (Yen) Yen, comb. nov. (Fig. 46)

= Cercospora katongensis Yen, Revue de Mycol. 31:120, 1966.

Leaf spots none or indistinct, sometimes mixed with spots of Cercospora

hydrangeae Ell. & Ev. Fruiting hypogenous, invisible. Stromata poor or small,

black brown, globular, 13.5-36um in diam. Fascicles poor to very dense (2-32

stalks), emerging through the stomata. Conidiophores hypophyllous, pale oliva-

ceous brown, simple or branched, straight or mildly flexuous, not geniculate, 0-3

septate. sometimes with constrictions at septa, tip rounded, conidial scars in-

conspicuous, 25-36 x 2.5-3.6um. Conidia cylindrical or oblcavato-cylindrical,

pale olivaceous brown, straight or slightly curved, sometimes with a geniculation,

3-8 septate, tip subrounded, base obconic truncate, 36-90 x 2.5-3.6um.

Habitat: On leaves of Hydrangea macrophylla (Hydrangeaceae), Katong,

Singapore, leg. Mrs. Shu-hsien Yen (No. 95), Dec. 18, 1964; on the same host

plant, leg. Lee Chih-shao & Mrs. Shu-hsien Yen (No. 109), Apr. 18, 1965.

Note: Cercospora hydrangeae Ell. & Ev. differs from this species by its

amphigenous fruiting and hyaline conidia.

71. Pseudocercospora lyciicola (Yen) Yen, comb. nov. (Fig. 47)

= Cercospora lyciicola Yen, Revue de Mycol. 30:180, 1965.

Leaf spots indistinct, sometimes appearing as yellow areas on the upper side

and dark gray areas on the lower side. Fruiting amphigenous, invisible. Fascicles

poor to very dense (2-30 stalks), emerging through the stomata. Stromata sub-

stomatous, globular or irregular, dark brown, 30-46m in diam. Conidiophores

amphiphyllous, pale brown to brown, paler towards the tip, simple cr branched,

straight or tortuous, 1-4 septate, with constrictions at septa, unthickened small

condial scar at the attenuate and truncate tip, 23-78 x 3.5-5.5um. Conidia

cylindrical or obclavato-cylindrical, pale olivaceous brown, straight or slightly

curved, 3-13 septate, tip rounded, base obconic truncate, 33.5—148 x 3.6-6um.

Habitat: On leaves of Lycium chinense (Solanaceae), Geylang, Singapore,

leg. Mrs. Shu-hsien Yen (No. 65), Jun. 28, 1964.

Note: Cercospora chengtuensis Tai and C. lycii Ell. & Hals. are different

from this fungus by their strictly hypogenous fruitings.

180 Gardens’ Bulletin, Singapore XX XIII (1980)

72. Pesudocercospora macarangae (H. & P. Syd.) Deighton (Fig. 48)

Mycol. Papers 140:47, 1976.

= Cercospora macarangae H. & P. Syd., Ann. Mycol. 12:575, 1914.

Syn. Cercospora macarangae Yen & Lim, Bull. Soc. Mycol. Fr. 86:748,

1970b.

Leaf spots distinct, circular, brown to dark brown, with an indefinite border,

more distinct on the lower side, scattered, 2-7mm in diam., sometimes confluent.

Fruiting epiphylious, visible as black minute pustules under hand lens. Stromata

globose, black brown, usuaily above the stomata, 25-40um in diam. Fascicles

poor to dense (2-25 stalks), emerging through the stomata. Conidiophores brown,

paler toward the tip, simpie, never branched, erect or flexuous, 1-10 septate,

apex rounded or slightly truncate, 20-200 x 4-5um. Conidia obclavate, pale

olivaceous brown, straight or slightly curved, 2-6 septate, tip obtuse or subacute,

base obconically truncate, 22-72 x 4-Syum.

Habitat: On leaves of Macaranga indica (Euphorbiaceae), Singapore, leg.

G. Lim, Nov. 28, 1969.

73. Pseudocercospora mori (Hara) Deighton

Mycol. Papers 140: 148, 1976.

= Cercospora mori Hara, J. Sericult. Assoc. Japan 27:227, 1918.

Syn. Cercospora mori Mar. & Stey., Bull. Soc. Roy. Bot. de Belg. 61

(N.S. 11): 166, 1929.

Leaf spots none or indistinct. Fruiting strictly hypophyllous, scantily effuse,

olivaceous gray to dark brown. Stromata lacking or consisting of few brown

cells. Fascicles lacking or very poor (2-6 stalks), emerging through the stomata.

External hyphae hypogenous, filling stomatal openings, olivaceous browa, septate,

branched, 2-2.54m in width. Conidiophores (either arising from fascicles or

from external hyphae) simple or branched, erect or slightly flexuous, 1-2 geni-

culate, 14 septate (occasionally 7-8 septate), brown, paler toward the apex, tip

sharply attenuate, conidial scars inconspicuous, 19-71 x 3.5—5.5y4m (occasionally

to 148m long). Conidia obclavate, but obclavto-cylindrical for the short ones,

pale olivaceous brown, 3-8 septate, straight or midly curved, tip acute or obtuse,

base obconically truncate, 20-80 (-108) x 3.5—Sym.

Habitat: On leaves of Morus alba (Moraceae), Bukit Timah, Singapore, leg.

Jo-min Yen (No. M.S. 648), Nov. 25, 1965.

Distribution: N. America, Belgium Congo, Japan, W. Pakistan, Singapore

and Taiwan.

74. Pseudocercospora muntingiae (Petr. & Cif.) Deighton

Mycol. Papers 140: 148, 1976.

= Cercospora muntingiae Petr. & Cif., Ann. Mycol. 30:324, 1932.

Cercospora and Allied Genera 181

Leaf spots distinct, angular to irregular, reddish brown with a yellowish

margin, scattered, 0.5-5mm in diam. Fruiting epiphyllous, invisible. Stromata

dark brown, globose or subglobose, small, 10-254m in diam. Fascicles medium

to dense. Conidiophores pale olivaceous brown, uniform in colour aad in width,

sparingly septate, not geniculate not branched, undulate or bent variously, un-

thickened conidial scars at the rounded to conic tip, 10-35 x 24um. Conidia

obclavato-cylindrical, pale olivaceous brown, straight or curved, 2-8 septate, tip

rounded, base subtruncate to obconically truncate, 20-65 x 2—4um.

Habitat: On leaves of Muntingia calabura (Elaeocarpaceae), Bukit Timah,

Singapore, leg. G. Lim, Apr. 4, 1969.

Distribution: N. America, San Domingo and Singapore.

75. Pseudocercospora peltophori (Yen) Yen, comb. nov. (Fig. 49)

= Cercospora peltophori Yen, Revue de Mycol. 29:228, 1964.

Leaf spots none or indistinct. Fruiting hypophyllous, invisible. Stromata

lacking. Fascicles lacking or very poor (2-4 stalks), emerging through the stomata.

Conidiophores hypogenous, either coming from fascicles or arising from procum-

bent threads, pale olivaceous brown, simple or branched, erect or tortuous, 0-4

septate, 0-2 geniculate, tip conic, conidial scars inconspicuous, 10-36 x 3—Sym.

External hyphae hypogenous, pale olivaceous brown, filling stomatal openings,

septate, branched, 2—3.54m in width. Conidia pale olivaceous brown, obclavato-

cylindrical or subacicular, slightly curved or undulate, 3-6 septate, tip conic,

base obconically trunate, 31-72 x 2.5-3.5ym.

Habitat: On leaves of Peltophorum pterocarpum (Leguminosae), Katong,

Singapore, leg. Mrs. Shu-hsien Yen (No. 37), May 7, 1964.

76. Pseudocercospora pergulariae (Yen & Lim) Yen, comb. nov. (Fig. 50)

= Cercospora pergulariae Yen & Lim, Cahiers du Pacifique 14: 100,

1970a.

Leaf spots angular or irregular, more distinct on the lower side, vein-limited,

black brown on the upper side, black on under side, scattered, 14mm in diam.

Fruiting strictly hypophyllous, appearing as black carpet on the lower surface

of the spots. Strom1ta black brown, globose, 24-40um in diam. Fascicles dense

to very dense, emerging through the stomata. Conidiophores hypophyllous, dark

brown, simple, not branched, erect or flexuous, 0-4 septate, 0-2 geniculate, tip

attennate and truncate, conidial scars inconspicuous, 30-91 x 5-6um. Conidia

cylindrical or obclavato-cylindrial, pale olivaceous brown to brown, straight or

slightly curved, with a finely granulated membrane, 1-5 septate, tip rounded, base

obconierlly truncate, 40-91 x 4.5—5.5um.

Habitat: On leaves of Pergularia minor (Asclepiadaceae), Bukit Timah,

Singanore, leg. G. Lim, Apr. 11, 1969.

77. Pseudocercosnora phyllanthi-niruri (Yen) Yen, comb. nov. (Fig. 51).

= Cercospora phyllanthi-niruri Yen, Revue de Mycol. 32:192, 1967a.

Syn. Cercospora phyllanthicola Yen, Revue de Mycol. 30:186, 1965.

non Cercospora phyllanthicola Shakil & Kamal, Indian Phytopath.

15:296, 1962.

182 Gardens’ Bulletin, Singapore XX XIII (1980)

Leaf spots none or indistinct. Fruiting hypophyllous, effuse, gray, often

extending and covering the whole lower surface of the leaf. Stromata lacking.

Conidiophores olivaceous brown, simple or branched, erect or flexuous, 1-5

septate, 1-3 geniculate, at times with constrictions at septa, unthickened small

conidial scar at the rounded tip, 54-126 x 4.5-64m. Conidia crescent shaped,

pale olivaceous brown, generally 3 septate, tip rounded, base slightly attenuate

and truncate, 45-60 x 4.5—6ym.

Habitat: On leaves of Phyllanthus niruri (Euphorbiaceae), Katong, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 18), Apr. 1, 1964.

Note: This fungus is different from Cercospora phyllanthicola Shakil &

Kamal and Pseudocercospora phyllanthi (Chupp) Deighton by its crescent shaped

and 3 septate conidia.

78. Pseudocercospora piperina (Yen) Yen, comb. nov. (Fig. 52)

= Cercospora piperina Yen, Revue de Mycol. 31:131, 1966.

Leaf spots distinct, circular, dark brown with grayish brown centre on the

upper side, surrounded by a raised border, olivaceous brown with an indefinite

margin on the lower side, scattered, 2-10mm in diam. Fruiting amphiphyllous,

but more abundant on the upper side, appearing as minute black pustules in

the centre of leaf spots. Fascicles dense to very dense. Stromata black brown,

globular to irregularly globular, 36—76um in diam.. Conidiophores amphiphyllous,

always simple, cylindrical, straight, continuous (rarely one septate), laterally

shouldered at the tip, pale olivaceous brown, tip generally rounded, conidial scars

inconspicuous, 21-30 x 2.5-6um. Conidia cylindrical, pale olivaceous brown,

generally straight, at times slightly curved, 3—6 septate, tip rounded, base trun-

cate, 38.5-73.5 x 3-4.5yum.

Habitat: On leaves of Piper muricatum (Piperaceae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No. 105), Feb. 15, 1965.

Note: Cercospora cartanthes P. Henn. resembles somewhat this fungus, but

is different from this species by its hypogenous fruiting and obclavate conidia.

79. Pseudocercospora piperis-muricati (Yen) Yen, comb. nov. (Fig. 53)

= Cercospora piperis-muricati Yen, Revue de Mycol. 31:134, 1966.

Leaf spots circular or irregularly circular, dark brown, with an indefinite

border on the upper side, pale brown on the lower side, scattered, 2-8mm in

diam. Fruiting strictly hypogenous, invisible. Stromata lacking or very rudi-

mentary. Fascicles hypophyllous, poor to dense (5-28 stalks), emerging through

the stomata. Conidiophores olivaceous brown or pale olivaceous brown, simple

or branched, straight or tortuous, 1-3 septate, 0-1 geniculate, tip long attenuate

and truncate, conidial scars inconspicuous, 30-56.5 x 3.5-4.8um. Conidia cylin-

drical, generally straight, sometimes slightly curved, pale olivaceous brown, 1-5

septate, tip rounded, base slightly attenuate and truncate, 31-65 x 2-3.5um.

Habitat: On leaves of Piper muricatum (Piperaceae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No, 103bis), Feb. 15, 1965; on same host plant, Bukit

Timah, Singapore, leg. Mrs. Shu-hsien Yen (No. 105bis), Mar. 19, 1965.

Cercospora and Allied Genera 183

Note: Cercospora piperata Asth. & Mah. resembles somewhat this species,

but differs from it by its continuous and unbranched conidiophores and especially

by its obclavate conidia.

80. Pseudocercospora psophocarpi (Yen) Deighton (Fig. 30, D-G)

Mycol. Papers 140:151, 1976.

= Cercospora psophocarpi Yen, Bull. Soc. Mycol. Fr. 83:338, 1967b.

Leaf spots distinct, circular with an indefinite border, brown on upper side,

olivaceous brown on lower side, scattered, 3-10mm in diam., sometimes confluent.

Fruiting amphigenous, mostly epiphyllous. visible as minute black pustules under

hand lens. Stromata black brown, irregularly globose, 24-57.5.m in diam. Fas-

cicles dense to very dense. Conidiophores pale olivaceous brown, simple, short,

erect or slightly tortuous, continuous, not geniculate, tip shortly attenuate, 13-28

x 34pm. Conidia cylindrical, pale olivaceous brown, curved, 1-6 septate, some-

times with slight constrictions at septa, tip rounded, base attenuate and truncate,

25.5-54 x 4-6um.

Habitat: On leaves of Psophocarpus tetragonolobus (Leguminosae), Jurong,

Singapore, leg. Jo-min Yen (No. 717bis), Dec. 31, 1965.

Note: This species differs from Pseudocercospora aranetae (Borl. & Rold.)

Yen by its well developed stromata and continuous conidiophores.

81. Pseudocercospora pterocarpicola (Yen) Yen, comb. nov. (Fig. 54)

= Cercospora pterocarpicola Yen, Revue de Mycol. 42:145, 1978.

Leaf spots distinct, orbicular, yellowish brown, 1-2mm in diam. Fruiting

strictly hypophyllous, invisible. Stromata rather developed, substomatic, globose

or irregularly globose, 25—-40um in diam. Fascicles dense to very dense, emerging

through the stomata. Conidiophores hypophyllous, always simple, erect or flex-

uous, brown, paler toward the apex, 1-16 septate, 0-3 geniculate, tip rounded,

conidial scars inconspicuous, 35-203 x 44m. Conidia obclaviform, pale olivaceous

brown, generally curved, 3—7 septate, tip conic or conic rounded, base obconically

Habitat: On leaves of Pterocarpus indicus (Leguminosae), Kuala Lumpur

(For. Res. Inst.), Malaysia, leg. Jo-min Yen (No. 71341), Sept. 28, 1971.

82. Pseudocercospora puerariicola (Yamam.) Deighton (Fig. 55, A, C)

Mycol. Papers 140:151, 1976.

= Cercospora puerariicola Yamam., Phytopath. Lab. Taithoku Imp.

Univ. Contr. 26: 142, 1934.

Leaf spots subcircular to angular, at first pale brown, then becoming brown

to dark brown, scattered, 1-3mm in diam., sometimes confluent. Fruiting amphi-

phyllous, chiefly epiphyllous. Stromata dark brown, globose, 26—5Oum in diam.

Fascicles dense to very dense, emerging through the stomata. Conidiophores

olivaceous brown, simple, continuous, straight or slightly tortuous, unthickened

conidial scars at the attenuate and truncate tips, 13-24 x 3.5-4.8um. Conidia

obclavato-cylindrical, pale olivaceous brown, midly curved, 3-6 septate, tip sub-

obtuse, base obconically truncate, 40-66 x 3-4.5um.

184 Gardens’ Bulletin, Singapore XX XIII (1980)

Habitat: On leaves of Pueraria phaseoloides (Leguminosae), Tanglin, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 51), May 29, 1964.

Distribution: N. America, Japan, Singapore and Taiwan.

83. Pseudocercospora puerariina (Yen) Yen, comb. nov. (Fig. 55, B, D-H).

= Cercospora puerariina Yen, Revue de Mycol. 30:189, 1965.

Leaf spots more distinct on upper surface of the leaf, at first very small

(0.5mm in diam.), gray, irregular or polygonal, vein-limited, then becoming yellow

brown and more heavily coloured in the centre, 1-2mm in diam., sometimes

becoming large brown areas. Fruiting amphigenous, more abundant on the lower

side. Stromata lacking. Fascicles poor to meagre (2-12 stalks), emerging through

the stomata. External hyphae pale brown, filling stomatal openings, septate,

branched, 2—2.54m in width. Conidiophores may be divided inio two kinds: A,

conidiophores coming from fascicles pale brown, paler at the tip, simple or

branched, generally tortuous, 1-3 geniculate, 1-6 septate, with constrictions at

septa, unthickened small conidial scar at the shortly attenuate tip, 30-53 x 4-6ym;

B, conidiophores as solitary branches from procumbent threads, pale oblivaceous

brown, never branched, cylindrical, 0-2 septate, O-1 geniculate, tip attenuate,

15-27.5 x 3.5-5um. Conidia obclaviform or acicular, pale olivaceous brown,

straight or slightly curved, 6-12 (—15) septate, tip acute, base obconically trun-

cate, 90-200 x 3.5—Sym.

Habitat: On leaves of Pueraria phaseoloides (Leguminosae), Katong, Singa-

pore, leg. Mrs. Shu-hsien Yen (No. 44), Mar. 18, 1964.

Note: Pseudocercospora puerariicola (Yamam.) Deighton is different from

this fungus by its continuous and always unbranched conidiophores.

84. Pseudocercospora salicina (Ell. & Ev.) Deighton (Fig. 56)

Mycol. Papers 140:94, 1976.

= Cercospora salicina Ell. & Ev., J. Mycol. 3:19, 1887.

Syn. Cercospora babylonicae Saw. Formosan Agr. Res. Inst. Rept. 87:79,

1944.

Cercospora minutipes Yen, Revue de Mycol. 31:128, 1966.

Cercospora Ssalicis-babylonicae Yen, Rsvue de Mycol. 31:139, 1966.

Leaf spots amphigenous, subcircular or irregularly angular, more distinct

on the upper side, dark brown, with an indefinite border, scattered, 0.5-4mm

in diam., sometimes confluent. Fruiting amphigenous, visible as minute black

pustules under hand lens, but sometimes invisible, even under hand lens. Stromata

subepidermic, black brown, globose, 30-60um in diam., but sometimes lacking.

Fascicles poor to very dense. Conidiophores amphiphyllous, brown to dark

brown, short, simple, generally continuous, at times 1-2 septate, tip rounded or

attenuate and truncate, conidial scars inconspicuous, 8.5-17 x 3-4.5u4m. Conidia

obclavato-cylindrical or fusiform for the short ones, pale olivaceous brown, straight

or slightly curved, 1-6 septate, tip rounded, base obconically truncate, 18-60 x

2.5-3.5um.

Cercospora and Allied Genera 185

Habitat: On leaves of Salix babylonica (Salicaceae), Katong, Singapore leg.

Mrs. Shu-hsien Yen (No. 92), Dec. 14, 1964.

Distribution: N. America, India, Japan, Singapore and Taiwan.

85. Pseudocercospora scopariicola (Yen) Deighton (Fig. 57)

Mycol. Papers 140:152, 1976.

= Cercospora scopariicola Yen, Bull. Soc. Mycol. Fr. 84:11, 1968.

Leaf spots none or indistinct. Fruiting amphigenous, effuse, visible as a

thin gray velvet on the underside. Stromata lacking or very small, consisting

of few brown cells. Fascicles meagre to dense (3-18 stalks), emerging through

the stomata. External hyphae hypophyllous, very abundant, filling stomatal open-

ings, pale brown, septate, branched, 2.5-3.54m in width. Conidiophores (either

arising from fascicles or from external hyphae), pale brown, simple or branched,

0-2 geniculate, 1-5 septate, with constrictions at septa, tip rounded or slightly

attenuate and truncate, conidial scars inconspicuous, 12-48 x 2.5-5um. Conidia

cylindrical or obclavato-cylindrical, sometimes filiform, pale olivaceous brown,

slightly curved, 3-13 septate, tip rounded or obtuse, base slightly obconically

truncate, 67-165 x 2.5—3.6um.

Habitat: On leaves of Scoparia dulcis (Scrophulariaceae), Taiping, Singapore,

leg. Jo-min (No. M.S. 730), Dec. 30, 1965.

Note: Cercospora scopariae Lagy & Thir. differs from this fungus by the

presence of leaf spots and absence of external hyphae and especially by the

hyaline conidia.

86. Pseudocercospora singaporensis (Yen) Yen, comb. nov. (Fig. 58)

= Cercospora singaporensis Yen, Revue de Mycol. 29:237, 1964.

Leaf spots distinct, subcircular, yellowish gray to brownish gray, vein-limited

on the upper side, scattered, 0.5-4mm in diam. Fruiting amphiphyllous, invisible.

Stromata lacking. Fascicles poor to medium (2-10 stalks), emerging through the

stomata. Conidiophores olivaceous brown, simple or branched, erect or tortuous,

0-2 geniculate. 1-4 septate, a small unthickened conidial scar at the conic tip,

31-77 x 4.5-5.5um. Conidia cylindrical or obclavato-cylindrical, pale olivaceous

brown, straight or slightly curved, generally 3 septate (rarely 1 or 4 septate),

tip rounded base obconically truncate, 30-67 x 3.5m.

Habitat: On leaves of Cassia occidentalis (Leguminosae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No. 16), Apr. 2, 1964.

Note: This fungus is different from Pseudocercospora nigricans (Cooke)

Deighton by its distinct leaf spots and 3 septate conidia.

87. Pseudocercospora stahlii (F.L. Stev.) Deighton

Mycol. Papers 140:82, 1976.

= Helminthosporium stahlii F.L. Stev., Trans. Ill. Acad. Sci. 10: 208,

1917-°° |

186 Gardens’ Bulletin, Singapore XX XIII (1980)

Syn. Cercospora stahlii (Stev.) Subramanian, J. Indian Bot. Soc. 35: 460,

1956.

Cercospora passiflorae-foetidae Yen, Revue de Mycol. 29:226, 1964.

Leaf spots hypophyllous, angular or irregular, vein-limited, dark gray on

lower side, scattered, |-6mm in diam. Fruiting amphiphyllous, but mostly hypo-

phyllous, effuse, dark gray. Stromata small, pale brown, globose, 20-40um in

diam. Fascicles meagre to dense (2-18 stalks), filling stomatal openings. Coni-

diophores brown to dark brown, branched, 3-7 septate, 1-4 geniculate, apex

long attenuate as sharp point, 78-153 x 5-7um. Conidia clavate, pale olivaceous

brown, paler toward the base, straight, 1-3 septate, tip rounded, base attenuate

and subtruncate, 20.5-43 x 4-7yum.

Habitat: On leaves of Passiflora foetida (Passifloraceae), Katong, Singapore,

leg. Mrs. Shu-hsien Yen (No. 23), Apr. 25, 1964.

Distribution: Africa, N. America, Brunei, India, Malaysia, New Guinea,

Puerto Rico, Sabah, Singapore, Taiwan and Trinidad.

88. Pseudocercospora stephanotidis (Yen) Yen, comb. nov. (Fig. 59)

= Cercospora stephanotidis Yen, Revue de Mycol. 32:193, 1967a.

Leaf spots amphigenous, angular or irregular angular, more distinct on the

lower side, vein-limited, brown to dark brown, with an indefinite border, scattered,

2—6mm in diam., sometimes confluent. Fruiting amphiphyllous, but chiefly hypo-

phyllous, visible as black velvet on the lower surface of the spots. Stromata

black brown globose, 30-60um in diam. Fascicles dense to very dense, emerging

through the stomata. Condiophores brown or dark brown, paler toward the

apex, simple or branched, erect or flexuous, 0-5 septate, 0-2 geniculate, tip

rounded, membrane finely granulated, conidial scars inconspicuous, 30-80 x

6-7.5u4m. Conidia obclavate or obclavato-cylindrical, brown or dark brown,

straight or slightly curved, membrane finely granulated, 1-5 septate, tip rounded,

base obconically truncate, 37-92 x 4.5—7yum.

Habitat: On leaves of Stephanotis floribunda (Asclepiadaceae), Serangoon

Garden, Singapore, leg. Jo-min Yen (No. M.S. 729), Nov. 28, 1965.

89. Pseudocercospora stigmaphyllicola (Yen) Deighton (Fig. 60)

Mycol. Papers 140:153, 1976.

= Cercospora stigmaphyllicola Yen, Revue de Mycol. 30:192, 1965.

Leaf spots amphigenous, at first irregular, brown on upper side, but dark

brown on lower side, at last coalescing and covering the whole surface of the

leaf. Fruiting hypophyllous, invisible. Stromata lacking. Fascicles poor to

medium (2-11 stalks), filling the stomatal openings. External hyphae hypogenous,

abundant, emerging through the stomata, septate, branched, pale olivaceous brown,

1.5-2.54m in width. Conidiophores hypophyllous, either arising from fascicles

(poor to meagre, 2-6 stalks) or from external hyphae, simple (sometimes branched),

straight or tortuous, 0-2 septate, 0-2 geniculate, apex attenuate and truncate or

Cercospora and Allied Genera 187

laterally shouldered, conidial scars inconspicuous, 8.5-36 x 3.5-4.5um. Conidia

obclavato-cylindrical or cylindrical, pale olivaeous brown, straight or slightly

curved, 2-7 (rarely 1 or 8) septate, tip obtuse, base obconically truncate, 38-113

x 2-3yum.

Habitat: On leaves of Stigmaphyllon ciliatum (Malpighiaceae), Botanic

Gardens, Singapore, leg. Mrs. Shu-hsien Yen (No. 57), Jun. 7, 1964.

90. Pseudocercospora tabernaemontanae (H. & P. Syd.) Deighton

Mycol. Papers 140:154, 1976.

= Cercospora tabernaemontanae H. & P. Syd., Philipp. J. Sci. (Bot.)

8:507, 1913.

Leaf spots distinct, circular, 3-10mm in diam., pale tan or dingy gray with

a narrow brown border, often forming “‘shot-holes”. Fruiting amphigenous,

invisible. Stromata globose to elongate, bark brown, 20-5O0um in diam. Fas-

cicles dense to very dense. Conidiophores pale olivaceous brown, short, not

septate, not geniculate, unthickened, small conidial scar at the conic tip, 10-35

x 24um. Conidia cylindrical or obclavato-cylindrical, pale olivaceous brown,

straight or slightly curved, multiseptate, tip subobtuse to conic, base obconically

truncate, 15-65 x 2-3.5ym.

Habitat: On leaves of Tabernaemontana malacaensis (Apocynaceae), Tam-

pines, Singapore, leg. Jo-min Yen (No. M.S. 726), Dec. 30, 1965.

Distribution: Burma, India, W. Malaysia, Philippines, Sabah and Singapore.

91. Pseudocercospora tabernaemontanicola (Yen) Yen, comb. nov.

= Cercospora tabernaemontanicola Yen, Bull. Soc. Mycol. Fr. 84:14,

1968.

Leaf spots circular, brown, with an indefinite yellowish brown border on

the upper side, scattered, 2-5mm in diam. Fruiting hypophyllous, visible as

minute black pustules on the lower side. Stromata lacking or consisting of few

brown cells. Fascicles poor to medium (3-15 stalks), emerging through the

stomata. External hyphae hypogenous. filling stomatal openings, pale brown,

septate, branched, 2.5-4.5ym in width. Conidiophores strictly hypophyllous, either

arising from fascicles or arising from external hyphae, simple or branched, erect

or flexuous, 0-4 septate. 0-2 geniculate, apex attenuate and truncate, conidial

scars inconspicuous, 18-79 x 3.5-Sum. Conidia cylindrical (rarely obclavato-

cylindrical), pale olivaceous brown, curved, straight for the short ones, 3-8 sep-

tate, tip rounded or subobtuse, base slightly attenuate and truncate, 50-130 x

3-4.5um.

_ Habitat: On leaves of Tabernaemontana coronaria (Apocynaceae), Botanic

Gardens, Singapore, leg: Jo-min Yen (No. M.S. 725), Nov. 26, 1965.

Note: Pseudocercospora tabernaemontanae (H. & P. Syd.) Deighton is

different from this:fungus by its ‘“‘shot-hole” leaf spots and its simple and con-

tinuous condiophores.

188 Gardens’ Bulletin, Singapore XX XIII (1980)

92. Pseudocercospora thunbergiicola (Yen) Deighton (Fig. 61)

Mycol. Papers 140:154, 1976.

= Cercospora thunbergiicola Yen, Revue de Mycol. 30:1965, 1965.

Leaf spots amphigenous, circular to angular, or irregular, with a whitish

centre, surrounded by a dark brown border, scattered, 0.5-3mm in diam. Fruit-

ing amphiphyllous, invisible. Stromata rudimentary or small, brown, subglobose,

20-30um in diam. Fascicles meagre to dense (2-22 stalks), emerging through the

stomata. External hyphae amphigenous, also emerging through the stomata, pale

brown, septate, branched, 2—34m in width. Conidiophores either arising from

fascicles or arising from external hyphae, olivaceous brown, paler and narrower

towards the apex, generally simple, sometimes branched, erect or tortuous, 0-4

septate, 0-2 geniculate, tip attenuate and truncate, conidial scars inconspicuous,

12-80 x 3.5-5um. Conidia cylindrical or obclavato-cylindrical, pale olivaceous

brown, straight or slightly curved, 4-11 septate, tip subacute, base attenuate and

truncate, 67-174 x 3.5—Sym.

Habitat: On leaves of Thunbergia alata (Acanthaceae), Botanic Gardens,

Singapore, leg. Mrs. Shu-hsien Yen (No. 60), Jun. 20, 1964.

Note: Cercospora thunbergiae Boel. is different from this fungus by its

hypogenous fruiting and continuous and non-septate conidiophores.

93. Pseudocercospora timorensis (Cooke) Deighton (Fig. 50)

Mycol. Papers 140: 154, 1976.

= Cercespora timorensis Cooke, Grevillea 12:38, 1883.

Syn. Cercospora batatae Zimm., Ber. Land. — Forstw. Deutsch. Ostafr.

2:28, 1904.

Cercospora batatae P. Henn., Bot. Jahrb. von Engler 38:118, 1907.

Leaf spots circular to irregular, at first yellowish brown with an indefinite

border, later becoming grayish in the centre, scattered, 6-10mm in diam. Fruit-

ing amphigenous, but mostly hypophyllous, invisible. Stromata lacking, or very

rudimentary. Fascicles poor to medium (3-12 stalks), emerging through the

stomata. Conidiophores pale olivaceous brown, always simple, cylindrical, erect

or slightly flexuous, continuous, sometimes 1-2 septate, tip attenuate and truncate,

14-50 x 3-S5ym. Conidia cylindrical, pale olivaceous brown, slightly curved,

3-7 septate, tip rounded, base very slightly attenuate and truncate, 40-103 x

3.5-4.5um.

Habitat: On leaves of Ipomoea batatas (Convolvulaceae),.. Bukit Timah,

Singapore, leg. G. Lim, Apr. 25, 1969..

Distribution: Africa, China, Ecuador, Japan, Panama, Philippines and

Singapore.

Cercospora and Allied Genera 189

94. Pseudocercospora trematicola (Yen) Deighton (Fig. 62)

Mycol. Papers 140:154, 1976.

= Cercospora trematicola Yen, Bull. Soc. Mycol. Fr. 86:752, 1970b.

Leaf spots distinct on the lower side, but indistinct on the upper sider, gray,

angular, very small, strictly vein-limited, scattered, 0.2-Imm in diam., often coa-

lescing and forming gray large patches. Fruiting strictly hypophyllous, visible

as gray velvet under hand lens. Stromata lacking. Fascicles occasionally present,

Very poor (2-6 stalks), emerging through the stomata. External hyphae hypo-

phyllous, emerging through the stomata, pale olivaceous brown, septate, branched,

2.5—3.5um in width. Conidiophores either arising from fascicles or from external

hyphae, simple or branched, erect or flexuous, not geniculate, 0-5 septate, tip

attenuate and truncate, 12-57.5 x 3.5-7um. Conidia obclavato-cylindrical, but

cylindrical for the short ones, pale olivaceous brown, straight or slightly curved,

2-10 septate, tip rounded, base obconically truncate, 36-96 x 4—5yum.

Habitat: On leaves of Trema orientalis (Urticaceae), Jurong, Singapore, leg.

Jo-n0ny en, (No: M:S..727),. Dec., 17,1965.

95. Pseudocercospora trematis-cannabini (Yen & Lim) Deighton (Fig. 63).

Mycol. Papers 140: 154, 1976.

= Cercospora trematis-cannabini (‘tremae-canabinae’) Yen & Lim, Bull.

Soc. Mycol. Fr. 86:749, 1970b.

Leaf spots amphigenous, circular to irregularly circular, pale brown, with

an indefinite border, scattered, 0.5-5mm in diam., sometimes confluent. Fruit-

ing strictly epiphyllous, visible as minute black hairs under hand lens. Stromata

dark brown, globose, 18-30um in diam. Fascicles medium to dense, emerging

through the stomata. Conidiophores olivaceous brown, paler toward the apex,

simple, never branched, filamentous, 2-14 septate, 0-2 geniculate, tip rounded

or occasionally truncate, 45-170 x 3-4ym. Conidia obclavato-cylindrical or

cylindrical, pale olivaceous brown, usually curved, 5—12 septate, tip rounded or

subobtuse, base shortly attenuate and truncate, 50-115 x 34ym.

Habitat: On leaves of Trema cannabina (Urticaceae), Johor Baru, Malaysia,

leg. G. Lim, Nov. 28, 1969.

Note: This fungus differs from others by its strictly epigenous fruiting and

long filamentous conidiophores.

96. Pseudocercospora ubicola (Yen) Deighton (Fig. 64).

Mycol, Papers 140:155, 1976.

Leaf spots amphigenous, irregular, brown, at first small and vein-limited,

then confluent and forming large patches with an indefinite border. Fruiting

strictly hypogenous, invisible. Stromata lacking. Fascicles poor to medium

(2-15 stalks) emerging through the stomata. External hyphae hypophyllous,

also emerging through the stomata, 2-5 in fascicles, pale, olivaceous brown, sep-

tate, branched, 1.2—2.5um in width.. Conidiophores either arising from fascicles

or from external hyphae, generally simple, erect or tortuous, 0-3 septate, 0-2

190 Gardens’ Bulletin, Singapore XX XIII (1980)

geniculate, sometimes with constrictions at septa, pale olivaceous brown, tip atten-

uate and truncate, 6-30 x 3.5-5ym. Conidia very thin, acicular or filiform,

subhyaline, straight or slightly curved, 3-8 septate, tip conic, base slightly obconic

truncate, 56-101 x 2-2.5um.

Habitat: On leaves of Dioscorea sp. (Dioscoreaceae), Botanic Gardens,

Singapore, leg. Mrs. Shu-hsien Yen (No. 54), Jun. 7, 1964.

97. Pseudocercospora viticicola (Yen & Lim) Yen, comb. nov. (Fig. 65).

= Cercospora viticicola Yen & Lim, Cahiers du Pacifique 17: 104, 1973.

Leaf spots irregularly circular to irregularly angular, grayish brown, with an

indefinite black brown border, scattered, 14mm in diam., distinct on the upper

side, but strictly indistinct on the lower side, sometimes confluent. Fruiting

amphiphyllous, visible as black minute pustules under hand lens. Stromata

epigenous, forming within the epidermal cell, black brown, globose, 24-44um

in diam. Fascicles dense to very dense, emerging through rupture of epidermal

cells. External hyphae strictly hypophyllous, emerging through the stomata, pale

olivaceous brown, septate, branched, 2—34m in width. Conidiophores may be

divided into two kinds: A, conidiophores arising from fascicles, always epi-

genous, pale olivaceous brown, simple, never branched, continuous or one septate,

erect or slightly tortuous, apex irregularly rounded or slightly attenuate and

truncate, 8-35 x 2-4.5um; B, conidiophores arising from external hyphae, always

hypogenous, pale olivaceous brown, simple, erect, continuous, tip attenuate and

truncate, 2-15 x 2-3.5um. Conidia obclavato-cylindrical or obclavate, pale oliva-

ceous brown midly curved, 3-7 septate, tip subobtuse, base obconically truncate,

33-70 x 2.5-3.5ym.

Habitat: On leaves of Vitex pubescens and V. trifolia (Verbenaceae), Singa-

pore, leg. G. Lim (No.-S:U.: 75))) Mar, 34972.

Note: Cercospora weberi Chupp differs from this species by its epigenous

fruiting and strictly epiphyllous external hyphae.

98. Stenella fagraeae Yen

Revue de Mycol. 32:200, 1967a.

Leaf spots amphigenous, very distinct, circular, pale brown but heavily

coloured in the centre, later forming “‘Frog eyes’’, with fine concentric lines in

the centre and surrounded by a raised and thick border, cattered, 1-3mm in

diam. Fruiting amphiphyllous, appearing as minute black pustules under hand

lens. Stromata generally small, sometimes well developed, globose or irregularly

globose, dark brown to nearly black, 40-90um in diam. Fascicles medium to

dense 2-2) stalks), emerging through the stomata or through the rupture of

the epidermis. External hyphae emerging through the stomata, pale brown to

brown, septate, branched, 3.5-4.54m in diam. Conidiophores amphiphyllous,

either arising from fascicles or arising from external hyphae, always simple,

cylindrical, straight, pale brown to brown, 0-1 septate, not geniculate, with a

finely granulated membrane, tip rounded or irregularly truncate, 1-2 brown coni-

dial scars at the apex, 28-43 x 3.5-4.5ym. Conidia cylindrical or obclavato-

cylindrical, pale brown, straight or slightly curved, with a finely granulated mem-

brane, 1—5 septate, tip rounded, base obconically truncate, 26-48 x 3.5—4.2um.

Habitat: On leaves of Fagraea fragrans (Loganiaceae), Bukit Timah, Singa-

pore, leg. Chih-shao Lee (No. 13), Aug. 20, 1965.

Cercospora and Allied Genera 191

Index of fungi

Cercoseptoria (6 sp.) ipomoeae-pes-caprae (27)

balasaminicola (1) mikaniicola (28)

ixoriana (2) malayensis (29)

polygonigena (3) nicotianae (30)

rhododendricola (4) nicotianicola (31)

tecomae-heterophyllae (5) peristrophes-acuminatae (32))

tecomicola (6) psophocarpicola (33)

pudicae (34)

Cercospora (34 sp.)

sesamigena (35)

b teridis (7

abacopteridis (7) thunbergiana (36)

i (8

acori (8) tithoniae (37)

= (9

apii (9) triseptospora (38)

asystasiana (10) volkameriae (39)

balsaminiana (11)

barleriae (12)

zinniae (40)

canavaliiana (13) Cercosporidium (1 sp.)

canescens (14) personatum (41)

capsici (15)

cassiae-nodosae (16) Pseudocercospora (56 sp.)

cassiae-occidentalis (17) abacopteridicola (42)

citrullina (18) abelmoschi (43)

coleiana (19) acalyphicola (44)

dioscoreae-pyrifoliae (20) allamendae (45)

eupatorii-odorati (21) aranetae (46)

gloriosicola (22) asystasiae (47)

henningsii (23) bauhiniae (48)

holobrunnea (24) byliana (49)

hyalofilispora (25) cassiae-alatae (50)

hydrangeae (26) cassiigena (51)

192

centrosematicola (52)

chrysanthemicola (53)

columnaris (54)

cordiicola (55)

crotalaricola (56)

cruenta (57)

daturina (58)

delonicis (59)

ervatamiae (60)

eupatoriil-formosani (61)

fici-chartaceae (62)

formosana (63)

globosae (64)

hiratsukana (65)

ipomoeae-purpureae (66)

ixoricola (67)

jasminicola (68)

jatrophae-curcas (69)

katongensis (70)

lyciicola (71)

macarangae (72)

mori (73)

muntingiae (74)

peltophori (75)

Gardens’ Bulletin, Singapore XX XIII (1980)

pergulariae (76)

phyllanthi-niruri (77)

piperina (78)

piperis-muricati (79)

psophocarpi (80)

pterocarpicola (81)

puerariicola (82)

puerariina (83)

salicina (84)

scopariicola (85)

singaporensis (86)

stahlii (87)

stephanotidis (88)

stigmaphyllicola (89)

tabernaemontanae (90)

tabernaemontanicola (91)

thunbergiicola (92)

timorensis (93)

trematicola (94)

trematis-cannabini (95)

ubicola (96)

Viticicola (97)

Stenella (1 sp.)

fagraeae (98)

aoe

Cercospora and Allied Genera

193

Index of fungal synonyms

Arthrobotryum (1 sp.)

puttemansii (54)

Cercospora (93 sp.)

abacopteridicola (42)

abelmoschi (43)

acalyphicola (44)

allamendae (45)

arachidis (41)

aranetae (46)

arboressentis (26)

articinctata (40)

asystasiae (47)

balsaminicola (1)

batatae (93)

bauhiniae (48)

byliana (49)

-cassavae (23)

cassiae-alatae (50)

cassiigena (51)

cearae (23)

centrosematicola (52)

chardoniana (18)

chrysanthemicola (53)

columnaris (54)

cordiae (55)

cordiicola (55)

crotalaricola (56)

cruenta (57)

cucurbitae (18)

daturina (58)

delonicis (59)

dolichi (57)

ervatamiae (60)

eupatoril-formosani (61)

fici-chartaceae (62)

formosana (63)

globosae (64)

hibisci-cannabini (43)

hibisci-manihotis (43)

hiratsukana (65)

hydrangicola (26)

ipomoeae-purpureae (66)

ixoriana (2)

ixoricola (67)

jasminicola (68)

jatrophae-curcas (69)

katongensis (70)

lantanae-aculeatae (63)

lusoniensis (57)

lyciicola (71)

macarangae (72)

manihotis (23)

mikaniae-cordiae (28)

minutipes (84)

momordicae (18))

mori (73)

muntingiae (74)

neovignae (57)

passiflorae-foetidae (87)

peltophori (75)

penicillata var. apii (9)

pergulariae (76)

personata (41)

phaseolorum (57)

phyllanthi-niruri (77)

phyllanthicola (77)

piperina (78)

piperis-muricati (79)

polygonigena (3)

psophocarpi (80)

pterocarpicola (81)

puerarlicola (82)

puerariina (83)

raciborskii (30, 57)

rhododendricola (4)

salicina (84)

sechii (18)

salicis-babylonicae (84)

scopariicola (85)

singaporensis (86)

stahlii (87)

stephanotidis (88)

stigmaphyllicola (89)

stuhlmanni (54)

tabernaemontanae (90)

tabernaemontanicola (91)

tecomae-heterophyllae (5)

tecomicola (6)

Gardens’ Bulletin, Singapore XX XIII (1980)

thunbergiicola (92)

timorensis (93)

tithoniae (37)

tithonicola (37)

trematicola (94)

trematis-cannabini (95)

ubicola (96)

vignae (57))

vignae-sinensis (57)

vignicaulis (14)

viticicola (97)

zinniae (40)

Cercosporiopsis (1 sp.)

canescens (14)

Cladosporium (1 sp.)

personatum (41)

Graphium (1 sp.)

laxum (54)

Helminthosporium (2 sp.)

manihotis (23)

stahlii (87)

Isariopsis (1 sp.)

griseola (54)

Passalora (1 sp.)

personata (41)

Phaeoisariopsis (1 sp.)

grisoela (54)

Septogloeum (2 sp.)

arachidis (41)

manihotis (23)

Cercospora and Allied Genera 195

Index of host plants

Abacopteris urophylla (7, 42)

Acalypha indica (44)

Acorus calambus (8)

Allamanda cathartica (45, 49)

Apium graveolens (9)

Arachis hypogaea (41)

Asystasia chelonoides (10)

A. coromandeliana (47)

Barleria cristata (12)

Bauhinia malabarica (48)

Canavalia gladiata (13)

Capsicum annum (15)

Cassia alata (50,51)

C. nodosa (16)

C. occidentalis (17, 86)

Centrosema pubescens (52)

Chrysanthemum sinense (53)

Clerodendron fragrans (39)

Coleus sp. (19)

Cordia cylindristachya (55)

Crotalaria striata (56)

Datura alba (58)

Delonix regia (59)

Dioscorea pyrifolia (20)

Dioscorea sp. (65, 96))

Ervatamia coronaria (60)

Eupatorium odoratum (21, 61)

Fagraea fragrans (98)

Ficus chartacea (62)

Gloriosa superba (22)

Gomphrena globosa (64)

Hibiscus esculentus (43)

H. mutabilis (29)

Hydrangea macrophylla (24, 25, 26, 38,

70)

Impatiens balsamina (1, 11)

Ipomoea batatas (93)

I. pes-caprae (27)

I. purpurea (66)

Ixora chinensis (2)

I. javanica (67)

Jasminum grandiflorum (68)

Jatropha curcas (69)

Lantana aculeata (63)

Lycium chinense (71)

Macaranga indica (72)

Manihot utilissima (23)

Mikania cordata (28)

Mimosa pudica (34)

Momordica charantia (18)

Morus alba (73)

Muntingia calabura (74)

Nicotiana tabacum (30, 31)

Passiflora foetida (87)

Peltophorum pterocarpum (75)

Pergularia minor (76)

Peristrophe acuminata (32)

Phaseolus vulgaris (14, 54)

Phyllanthus niruri (77)

196 Gardens’ Bulletin, Singapore XX XIII (1980)

Piper muricatum (78, 79) Tabernaemontana coronaria (91)

Polygonum sp. (3) T. malacaensis (90)

Psophocarpus tetragonolobus (33, 46, 80) Tecoma heterophylla (5)

Pterocarpus indicus (81) T. stans (6)

Pueraria phaseoloides (82, 83) Thunbergia alata (36, 92)

Rhododendron sp. (4) Tithonia diversifolia (37)

Salix babylonica (84) Trema cannabina (95)

Scoparia dulcis (85) T. orientalis (94)

Sesamum orientalis (35) Vigna sinensis (57)

Stenolobium stans (6) Vitex pubescens (97)

Stephanotis floribunda (88) V. trifolia (97)

Stigmaphyllon ciliatum (89) Zinnia elegans (40)

Cercospora and Allied Genera } 197

REFERENCES

CHUPP, C. (1954). A monograph of the fungus genus Cercospora. Ithaca, N.Y.

DEIGHTON, F.C. (1967). Studies on Cercospora and allied genera. II Passalora, Cercos-

poridium and some species of Fusicladium on Euphorbia. Mycol. Pap. 112, C.M.I.

Kew, Surrey.

DEIGHTON, F.C. (1971). Studies on Cercospora and allied genera. III Centrospora.

Mycol. Pap. 124, C.M.I. Kew, Surrey.

DEIGHTON, F.C. (1973). Studies on Cercospora and allied genera. IV Cercosporella

Sacc., Pseudocercosporella gen. nov. and Pseudocercosporidium gen. nov. Mycol. Pap.

133, C.M.I. Kew, Surrey.

DEIGHTON, F.C. (1976). Studies on Cercospora and allied genera. VI Pseudocercospora

Speg., Pantospora Cif. and Cercoseptoria Petr. Mycol. Pap. 140, C.M.I. Kew, Surrey.

FRESENIUS, G. (1863) Beitrage zur Mikologie. Frankfurt.

YEN, J.M. (1964). Etude sur les champignons parasites du sud-est asiatique. I. Premiere

note sur quelquees nouvelles especes de Cercospora de Singapour. Rev. Mycol. 29

(3), 209-240.

YEN, J.M (1965). Etude sur les champignons parasites du sud-est asiatique. III. Deuxieme

note sur quelques nouvelles especes de Cercospora de Singapour. Rev. Mycol. 30

(3), 166-204.

YEN, J.M. (1966). Etude sur les champignons parasites du sud-est asiatique. IV. Troisieme

note sur quelques nouvelles especes de Cercospora de Singapour. Rev. Mycol. 31

(2), 109-149.

YEN, J.M. (1967a). Etude sur les champignonos parasites du sud-est asiatique. VII.

Quatrieme note sur quelques Cercospora et Stenella de Singapour (Malaisie). Rev.

Mycol. 32 (3), 177-202.

YEN, J.M. (1967b). Etude sur les champignons parasites du sud-est asiatique. IX. Cin-

quieme note sur les Cercospora de Singapour (Malaisie). Bull. trimest. Soc. mycol.

Fr. 83 (2), 336-341.

YEN, J.M. (1968). Etude sur les champignons parasites du sud-est asiatique. X. Sixieme

note sur les Cercospora de Malaisie. Bull. trimest. Soc. mycol. Fr. 84 (1), 5-18.

YEN, J.M. (1978). Etude sur les champignons parasites du sud-est asiatique. XXVII.

Onzieme note sur les Cercospora de Malaisie. Rev. Mycol. 42 (2), 143-147.

YEN, J.M. (1979). Etude sur les champignons parasites du sud-est asiatique. XXVIII.

Douxieme note sur les Cercospora de Malaisie. Rev. Mycol. 43 (1), 97-100.

YEN, J.M. and LIM, G. (1969). Etude sur les champignons parasites du sud-est asiatique.

XIV. Huitieme note sur les Cercospora de Malaisie. Bull. trimest. Soc. mycol. Fr.

85 (4), 459-474.

YEN, J.M. and LIM, G. (1970a). Etude sur les champignons parasites du sud-est asiatique.

XII. Septieme note sur les Cercospora de Malaisie. Cah. Pacifique 14, 87-104.

YEN, J.M. and LIM, G. (1970b). Etude sur les champignons parasites du sud-est asiatique.

XV. Neuvieme note sur les Cercospora de Malaisie. Bull. trimest. Soc. mycol. Fr.

86 (3), 745-753.

YEN, J.M. and LIM, G. (1973). Etude sur les champignons parasites du sud-est asiatique.

XX. Dixieme note sur les Cercospora de Malaisie. Cah. Pacifique 17, 95-114.

198 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 1. Cercoseptoria balsaminicola (Yen & Lim) Yen (A, B & E): A, stromata and

fascicle of young conidiophores; B, old conidiophores; E, conidia. Cercospora

balsaminiana Yen & Lim (C & D): C, fascicle of condiophores; D. conidia.

Cercospora and Allied Genera 199

Fig. 2. Cercoseptoria ixoriana (Yen & Lim) Yen: A, fascicle of young conidiophores;

B & D, old conidiophores; C & F, external hyphae and unbranched conidiophores;

E, conidia.

200 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 3. Cercoseptoria polygonigena (Yen) Yen: A, fascicle of young conidiophores;

B & C, fascicles of old conidiophores; D, conidia.

Cercospora and Allied Genera 201

gee aser

¢ #,

St St

TASH

eae e

Fig. 4. Cercoseptoria rhododendricola (Yen) Deighton: A & B, fascicles of conidio-

phores and stromata; C, conidia.

202

Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 5. Cercoseptoria tecomae-heterophyllae (Yen) Yen: A, B & C, fascicles of

E &

conidiophores and stromata; D,

phores; F, conidia.

, external hyphae and unbranched condio-

Cercospora and Allied Genera 203

Fig. 6. Cercoseptoria tecomicola (Yen) Yen: A & B, fascicles of young conidicphores

and stromata: C. D & E, fascicles of old conidiophores without stromata; F, conidia,

G, external hyphae and unbranched conidiophores.

204

Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 7. Cercospora abacopteridis Yen & Lim (C-E): C, fascicle of conidiophores;

D, germination of conidia; E, conidia.

Pseudocercospora abacopteridicola (Yen & Lim)

Yen (A & B): A, conidia; B, external hyphae and conidiophores.

Cercospora and Allied Genera 205

Fig. 8. Cercospora acori Yen: A, B & C, young conidiophores; D, old conidiophores:

E, conidia; F-K, penetration by Cercospora hyphae via the germinative pore of

uredospores.

206 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 9. Cercospora asystasiana Yen: A, conidia; Aa, germination of conidia; B & C,

fascicles of conidiophores; D, single conidiophore; E, rudimentary stromata.

207

and Allied Genera

ospora

208 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 11. Cercospora canavaliiana Yen & Lim (A-F): A & C, fascicles of conidio-

phores; B, unbranched conidiophore; D & E, rudimentary stromata; F, conidia.

Pseudocercospora cruenta (Sacc.) Deighton (G & H): G, stromata and conidiophores;

H, conidia.

Cercospora and Allied Genera 209

Fig. 12. Cercospora cassiae-nodosae Yen: A, fascicle of conidiophores; B, ends of

conidiophores; C, conidia.

210 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 13. Cercospora_ cassiae-occidentalis Yen: A, fascicle of young conidiophores;

B, fascicle of old conidiophores; C, rudimentary stromata; D, conidia.

Cercospora and Allied Genera 211

Fig. 14. Cercospora coleiana Yen & Lim: A, fascicle of old conidiophores; B,

unbranched conidiophores; C, fascicle of young conidiophores; D, conidia.

212 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 15. Cercospora dioscoreae=pyrifoliae Yen: A, B & C, conidiophores with abnormal

globular swellings; D, fascicle of young conidiophores; E, fascicle of old conidiophores;

F, conidia.

Cercospora and Allied Genera 213

Fig. 16. Cercospora cupatorii-odorati Yen (D-J): D & F, fascicles of old conidio-

phores and stromata; E, external hyphae and unbranched conidiophores; G, fascicle

without stromata; H, fascicle of young conidiophores; I, conidia; J, germination of

conidia. Pseudocercospora eupatorii-formosani (Saw.) Yen (A-C): A, fascicle of young

conidiophores; B, fascicle of old conidiophores; C, conidia.

214 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 17. Cercospora gloriosicola Yen & Lim: A, fascicle of conidiophores; B, germina-

tion of conidia; C, conidia.

Cercospura and Allied Genera

215

Fig. 18. Cercospora holobrunnea Yen: A, external hyphae and unbranched conidio-

phores; B, conidia; C, germination of conidia.

216 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 19. Cercospora hyalofilispora Yen: A & B, external hyphae and unbranched

conidiophores; C, formation of conidia; D, conidia.

-_—

Cercospora and Allied Genera 217

Bik, da

Fig. 20. Cercospora ipomocac-pes-caprae Yen & Lim: A, fascicle of old conidio-

phores; B & C, fascicle of young conidiophores; D, conidia.

218

Gardens’ Bulletin, Singapore XX XIII (1980)

Sa eee? ve a =——— : |

[ j

Fig. 21. Cercospora nicotianicola Yen: A, conidia; B, fascicle of old conidiophores

and stromata; C & Ca, branched conidiophores; D, e 4s of conidiophores.

Cercospora and Allied Genera 219

Fig. 22. Cercospora peristrophes-acuminatae Yen: A & C, fascicles of conidiophores;

B, tip of conidiophore; D, conidia.

220 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 23. Cercospora psophocarpicola Yen: A & B, fascicles of conidiophores; C, old

single conidiophore; D, conidia.

Cercospora and Allied Genera 221

Fig. 24. Cercospora pudicae Yen: A-C, fascicles of conidiophores; D, conidia.

Cercospora and Allied Genera 223

Fig. 26. Cercospora thunbergiana Yen: A-C, fascicles of conidiophores; D & E,

formation of stromata; F, conidia.

224

Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 27. Cercospora triseptospora Yen: A & C, external hyphae and unbranched

conidiophores; B & D, fascicles of conidiophores and stromata;; E, conidia.

Cercospora and Allied Genera 225

”

oe)

Su eee A

AION RE fi

Fig. 28. Pseudocercospora acalyphigena (Yen) Yen: A, young conidiophores; B, old

and branched conidiophores; C, rudimentary stromata; D, conidia.

Cercospora and Allied Genera

221

Fig. 30. Pseudocercospora aranetae (Borl. & Rold) Yen (A-C): A & B, fascicles of

conidiophores; C, conidia. Pseudocercospora psophocarpi (Yen)

D-F, fascicles of conidiophores and stromata; G, conidia.

Deighton (D-G):

228 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 31. Pseudocercospora usystasiae (Yen) Yen: A, conidia; B, fascicle of old

conidiophores; C, fascicle of young conidiophores; D, stromata; E, absence of stromata.

Cercospora and Allied Genera

Fig. 32. Pseudocercospora cassiae-ala

hyphae and unbranched conidiophores; E, co

& Lim) Yen (A-C): A, fascicle of young conidi

tae (Yen & Lim) Yen (D & E): D, external

nidia. Pseudoce spora cassiigena (Yen

ophores; B, old conidiophores; C, conidi:

229

230 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 33. Pseudocercospora centrosematicola (Yen & Lim) Yen: A, fascicle of old

conidiophores and stromata; B, fascicle of young conidiophores and stromata; C. conidia.

D, branched conidiophores; E & F, external hyphae and single conidiophores; G, ger-

mination of conidia; H, secondary conidia.

Cercospora and Allied Genera 231

Fig. 34. Pseudocercospora chrysanthemicola (Yen) Deighton: Ae ae of young

: conidiophores and stromata; B, conidia; C & D, fascicle of old conidiophores.

pepe Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 35. Pseudocercospora cordiicola (Yen) Yen: A & C, fascicle of conidiophores;

B, conidia; D & E, external hyphae and unbranched conidiophores.

Cercospora and Allied Genera 233

Fig. 36. Pseudocercospora crotalaricola (Yen) Yen: A, fascicle of young conidiophores;

B, old conidiophores and external hyphae; C & E, external hyphae and unbranched

conidiophores; D, branched conidiophores; F, conidia.

934 Gardens’ Bulletin, Singapore X XXIII (1980)

Fig. 37. Pseudocercospora daturinad (Yen) Deighton: A, fascicle of young conidio- ,

phores; B, C & E, fascicle of old conidiophores; F, conidia; G, branched conidiophores.

Cercospora and Allied Genera 235

Fig. 38. Pseudocercospora delonicis (Yen) Yen: A, B, D, E & F, external hyphae

and unbranched conidiophores; C, conidia; G & H, germination of conidia; I, infection

by germinative tube through leaf stoma.

236 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 39. Pseudocercospora ervatamiae (Yen & Lim) Yen: A, fascicle of old conidio-

phores and stromata; B & C, fascicle of young conidiophores; D, formation of conidia;

E, conidia.

Cercospora and Allied Genera

Fig. 40. Pseudocercospora fici-chartaceaeue (Yen & Lim) Yen: A & C, conidiophores

emerging through stomata; B, fascicle of yaung conidiophores emerging through rupture

of epidermis; D, fascicle of old conidiophores; E, conidia.

237

238 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 41. Pseudocercospora formosana (Yamam.) Deighton: A & C, fascicle of old

conidiophores; B & D, external hyphae and unbranched conidiophores; E, conidia.

Cercospora and Allied Genera 239

Fig. 42. Pseudocercospora globosae (Yen) Deighton: A, fascicle of old conidiophores

and stromata; B, fascicle of young conidiophores; C, conidia.

740 Gardens’ Bulletin, Singapore XX XIII (1980)

ZB = ~— —— ; -

ee tS eee

CS eo 3 2

=) I

Age

Fig. 43. Pseudocercospora ipomoeue-purpureae (Yen) Yen: A, fascicle of young

conidiophores and stromata; B & C, fascicle of old conidiophores; D, conidia and

germination of conidia; E, formation of conidia. al

Cercospora and Allied Genera 241

Fig. 44. Pseudocercospora ixoricola (Yen).Yen: A & B, fascicles of old conidiophores;

C, fascicle of young conidiophores; D, conidia; E, germination of conidia.

242 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 45.

Pseudocercospora jatrophae-curcaus (Yen) Deighton:

conidiophores;

E, conidia;

F, fascicle of young conidiophores.

A & B, fascicles of old

C & D, external hyphae and unbranched conidiophores;

Cercospora and Allied Genera 243

Fig. 46. Pseudocercospora katongensis (Yen) Yen: A, B & D, meagre fascicles;

C & F, dense fascicles of conidiophores and stromata; E, conidia; G, geniculate conidia.

244 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig.. 47. Pseudocercospora lyciicola (Yen) Yen: A & B, fascicles of old conidiophores

and stromata; C, formation of stromata; D, germination of conidia; E, anastomosis of

conidia; F, conidia.

«~

a aeeetiaeniion

Cercospora and Allied Genera 245

Fig. 48. Pseudocercospora macarangae (Syd.) Deighton: A & B, fascicles of old

conidiophores and stromata; C, fascicle of young conidiophores; D, formation of

conidia; E, conidia.

247

Cercospora and Allied Genera

peste |

fitter

Aa]

A beled,

Fig. 50. Pseudocercospora pergulariae (Yen & Lim) Yen (A-E): A, fascicle of coni-

diophores and stromata; B, fascicle of young conidiophores; C, old conidiophores;

Pseudocercospora timorensis (Cooke) Deighton (F-H): F, old conidio-

D & E, conidia. bp.

phores; G, fascicle of young conidiophores; H, conidia.

248 Gardens’ Bulletin, Singapore XX XIII (1980)

an’

were

ori

Chew,

nae

aiaeg

Sgt ahe

Fig. 51. Pseudocercospora phyllanthi-niruri (Yen) Yen: A, fascicle of young conidio-

phores; B & C, fascicles of old conidiophores; D, conidia.

Cercospora and Allied Genera 249

Soviet

}

Fig. 52. Pseudocercospora piperina (Yen) Yen: A & C, fascicles of conidiophores;

B, conidia; D, formation of stromata.

250 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 53. Pseudocercospora piperis-muricati (Yen) Yen: A-C, fascicles of conidiophores;

D, old conidiophores; E, conidia.

Cercospora and Allied Genera 251

Fig. 54. Pseudocercospora pterocarpicola (Yen) Yen: A, conidia; B, germination of

conidia; C, fascicle of conidiophores and stromata.

252 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 55. Pseudocercospora puerariicola (Yen) Yen (A & C): A, fascicle of conidio-

phores and formation of conidia; C, fascicle of old conidiophores. Pseudocercospora

puerariina (Yen) Yen (B & D-H): B & E, fascicles of conidiophores; D, F & H, external

hyphae and unbranched conidiophores; G, conidia.

Cercospora and Allied Genera 253

Fig. 56. Pseudocercospora salicina (Ell. & Ev.) Deighton: A, formation of stromata;

& C, fascicles of young conidiophores and stromata; D, old conidiophores and

formation of conidia; E, conidia; F, branched conidiophores and formation of conidia.

Gardens’ Bulletin, Singapore XX XIII (1980)

254

A & D, fascicles of conidio-

B & F, external hyphae and

ola (Yen) Deighton:

conidiophores;

Cercospora and Allied Genera 255

Fig. 58. Pseudocercospora singaporensis (Yen) Yen: A-D, fascicles of conidiophores;

E, conidia.

Gardens’ Bulletin, Singapore XX XIII (1980)

256

A & B, fascicles of conidiophores

; C-E, conidia; F & G, old branched conidiophores.

rcospora stephanotidis (Yen) Yen:

Fig. 59. Pseudoce

and stromata

Cercospora and Allied Genera Zot

1 eed tees

ae | OEY

a ae

Rp cen ) Serr

Fig. 60. Pseudocercospora stigmaphyllicola (Yen) Deighton: B & E, fascicles of coni-

diophores; A, C & D, external hyphae and unbranched conidiophore; F, conidia.

258 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 61. Pseudocercospora thunbergiicola (Yen) Deighton: A & B, fascicles of coni-

diophores; E, old conidiophores; C, F & H, external hyphae and unbranched conidio-

phores; D, anastomosis of conidia; G, conidia.

Cercospora and Allied Genera 259

Fig. 62. Pseudocercospora trematicola (Yen) Deighton: A, B & C, fascicles of conidio-

phores and external hyphae; D, external hyphae and formation of unbranched conidio-

phores; E, conidia.

Poe

Gardens’ Bulletin, Singapore XX XII (1980, a

260

‘

a ¢ ~~.

Va . f ‘ :

itd al? 4 9

ry < 4

a «

7 S

»éa: 7 .

* ;

B, C & D, ends of conidiophores”

‘

of conidiophore.

ores and stromata;

conidia; F, base

Fig. 63. Pseudocercospora trematis-cannabini (Yen & Lim) Deighton:

young conidioph

of conidia;

Cercospora and Allied Genera 261

Fig. 64. Pseudocercospora ubicola (Yen) Deighton: A & F, fascicles of conidiophores;

B, C & D, external hyphae and unbranched conidiophores; E, conidia.

262 Gardens’ Bulletin, Singapore XX XIII (1980)

Fig. 65. Pseudocercospora viticicola (Yen & Lim) Yen: A, fascicle of young conidio-

phores and stromata; B, fascicle of old onidiophores; C & D, external hyphae and

b hed dioph E idia; mination of. conidia.

Cercospora and Allied Genera 263

Fig. 66. Stenella fagraeae Yen: A & D, fascicles of conidiophores and stromata;

B, stromata and formation of external hyphae; C, external hyphae and unbranched

conidiophores; E, conidia.

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Gardens’ Bulletin Singapore

Index to

vol. 33: 1 & 2 (1980)

taxa in BOLD type are described or discussed in detail. italicised names are

synonyms, basionyms, or taxa from outside the Malay Peninsula which are not

discussed in detail

Abacopteris lineata (Bl.) Ching i23

menisciicarpa (Bl.) Holtt. 23

multilineata (Hook.) Ching var. malayensis Holtt. 22

peltochlamys (C. Chr.) Holtt. 20

rubicunda (v.A.v.R.) Holtt. 24

salicifolia (Hook.) Holtt. 24

triphylla (Sw.) Ching 23

var. parishii (Bedd.) Holtt. 24

urophylla (Hook.) Ching 23, 154, 168

Abelmoschus esculentus Moen. 334

moschatus Medic. 334

Abutilon indicum (L.) Sweet 334

Acalypha godseffiana Mast. 354

hispida Burm. f. 354

indica L. 169, 354

siamensis Oliv. ex Gage 355

wilkesiana M.-A. 355

Acorus calamus L. 155

Acronychia porteri Hk. f. 343

Actephila excelsa M.-A. var. javanica (Miq.) P. & H. = 355

Aegle marmelos Corr. 343

Aglaia cordata Hiern. 348

glabriflora Hiern. 348

odorata Lour. 348

odoratissima BI. 348

oligophylla Miq. 348

palembanica Miq. 348

trichostemon DC. 348

Agrostistachys longifolia Benth. 355

sessilifolia P. & H. 355

Agyneia bacciformis L. 367

Alchornia javensis M.-A. 355

rugosa M.-A. 355

villosa M.-A. 355

Aleurites moluccana Willd. 355

Allamanda cathartica L. 169, 170

Amoora aphanamixis Schul. 349

cucullata Roxb. 349

rubescens Hiern. 349

rubiginosa Hiern. 349

Ampelopteris prolifera (Retz.) Copel. 3

Amphineuron immersum (BI.) Holtt. 28

opulentum (Kaulf.) Holtt. 28

terminalis (Hook.) Holltt. 27

Anplectrum laeve Ohwi 315

Antidesma alatum Hook. 356

coriaceum Tul. 355

cuspidatum M.-A. 336

neurocarpum Miq. 356

velutinosum BI. 356

Aphanamixis rohituka Pierre 349

Apium graveolens L. 155

Aporusa benthamiana Hk. f. 336

bracteosa P. & H. 356

confusa Gage

falcifera Hk. f.

frutescens BI.

lunata (Mig.) Kurz

maingayi Hk. f.

microstachya M.-A.

nervosa Hk. f.

nigricans Hk. f.

prainiana King ex Gage

symplocoides (Hk. f.) Gage

Arachis hypogea L.

Arthrobotrym puttamansii P. Henn,

Aspidium aridum D. Don

callosum BI.

ciliatum Benth.

crassifolium Bl.

flaccidum Bl.

glandulosum BI.

gracilipes BI.

heterocarpon BI.

hispidulum Decne,

immersum BL.

latebrosum Mett.

menisciicarpon BL.

opulentum Kaul.

Aspidium pennigerum BI.

polycarpum BI.

subpubescens BI.

Asystasia chelonoides

cormandeliana Nees

Austrobuxus nitidus Miq.

Averrhoa bilimbi L.

carambola L.

Azadirachta indica Juss.

Baccaurea bracteata M.-A.

griffithii Hk. f.

hookeri Gage

kingii Gage

kunstleri King ex Gage

lanceolata M.-A.

macrophylla M.-A.

maingayi Hk. f.

minor Hk. f.

motleyana King

parviflora M.-A.

pyriformis Gage

racemosa M.-A.

reticulata Hk. f.

summatrana M.-A.

Barleria cristata L.

Bauhinia malabarica

Blumeodendron tokbrai J. J. Sm.

Breynia discigera M.-A.

Boletus albo-ater Schw.

atripurpureus Corn.

dictyotus

griseipurpureus Corn.

longipes Mass.

var. albus —

malaccensis (Pat. & Bak.) Corn.

mucosus Corn.

nanus Mass.

obscurecoccineus Hoehn.

358

156

170

358

294

290

292

290, 294

290-296

291292

295

294

phaeocephalus Pat. & Bak. 294

rarus Corn. 295

ridiculus Corn. subgen. Boletellus 294

rubiicolor Corn. 295

tristis Pat. & Bak. 290-296

Breynia coronata Hk. f. 358

reclinata Hk. f. 358

rhamnoides M.-A. 358

vitis-idaea C. E. C. Fisch. 358

Bridelia pustulata Hk. f. 359

stipularis BI. 359

tomentosa BI. 359

Brownlowia argentata Kurz 330

lanceolata Benth. 330

tersa (L.) Kosterm. 330

Brucea javanica Merr. 345

sumatrana Roxb. 345

Byttneria maingayi Mast. 338

Camellia lanceolata (Bl.) Seem. 289

sinensis O.K. 306, 307

Canarium commune L. 346

grandiflorum Benn. 346

grandifolium (Ridl.) Lam 346

kadondon Benn. 346

littorale BI. 346

nitidum Benn. 346

patentinervium Miq. 346

pilosum Benn. 346

purpurascens Benn. 346

rufum Benn. 346

secundum Benn. 346

vulgare Leenh. 346

Canavalia gladiata (Jacqg.) DC. 156

Capsicum annum L. 157

Carapa moluccensis Lmk. 351

obovata BI. 351

Cassia alata L. 171

nodosa B.-H. ex Roxb. 157

occidentalis L. 158, 185

Ceiba pentandra (L.) Gaertn. 336

Centrosema pubescens Benth. 172

Cercoseptoria balsaminicola (Yew & Lim) Yew 152

ixoriana (Yen & Lim) Yen 1?

polygonigena (Yen) Yen 152

rhododendricola (Yen) Deigh. 153

tecomae-heterophyllae (Yen) Yen 153

tecomicola (Yen) Yen 154

Cercospora abacopteridis Yen & Lim 154

abacopteridicola Yen & Lim 168

Cercospora abelmoschi Ell. & Ev. 168

acalyphicola Yen 16%

acori Yen 154

allamendae Yen 169

apii Fres. [55

arachidicola Hori 167

arachidis P. Henn. 167

aranetae Borl. & Rold. 169

arboressentis Teh. & Dan. 161

asystasiae Yen

asystasiana Yen 155

atricinctata Head & Wolf 166

avicularis Wint. ty

habylonicae Saw. ¢

balsaminiana Yen & Lim

balsaminicola Yen & Lim

barleriae Yen & Lim

barleriae-cristate Gov. & Thir.

barlericola Payak

batatae P. Henn.

batatae Zimm.

byliana Syd.

canavaliae Syd.

canavaliiana Yen & Lim

canavalticola San. & Kats.

canescens Ell. & Mart.

capsic Heald & Wolf

cartanthes P. Henn.

cassavae Ell. & Ev.

cassiae-alatae Yen & Lim

cassiae-nodosae Yen

cassiae-occidentalis Yen

cearae Petch

centrosemae Chupp & Mull.

centrosematicola Yen & Lim

chardoniana Chupp

chengtuensis Tai

chrysanthemi Heald & Wolf

chrysanthemicola Yen

citryllina Cooke

clitoria Atk.

colei Boed.

coleiana Yen & Lim

coleicola Chupp & Mull.

columnaris Ell. & Ev.

cordiae Chupp

cordiae Yen

cordiicola (Yen) Yen

crotalaricola Yen

cruenta Sacc.

cucurbitae Ell. & Ev.

cylindrospora Stev. & Solh.

daturina Yen

delonicus Yel.

dioscorae-pyrifoliae Yen

dolichii Ell. & Ev.

ervatamie Yen & Lim

eupatorii-odorati Yen

eupatorii-formosani Sav.

fici-chartaceae Yen & Lim

formosana Yam.

fukushaiana (Mats.) Yam.

elobosae Yen

gloriosicola Yen & Lim

gomphrenicola Speg.

handelii Bubak

hiratsukana Teg. & Kats.

henningsii All.

hibisci-cannabini Saw.

hibisci-manihotis P. Henn.

holobrunnea Yen

hyalofilispora Yen

hydrangeae Ell. & Ev.

hydrangeicola Speg.

hydrangicola Tharp

ipomoeae-pes-caprae Yen & Lim

ipomoeae-purpureae Yen

152, 155

152

156

188

170

156

157

182

160

171

157

158

160

172

171

172

158

172

173

152, 156

ixoriana Yen & Lim

ixoricola Yen

jasminicola Muell. & Chupp

jatrophae-curas Yen

katongensis Yen

lantana-aculeatae Yen

lusoniensis Sacc.

lychii Ell. & Hals.

lyciicola Yen

macarangae H. & P. Syd.

macarangae Yen & Lim

malayensis Stev. & Solh.

manihotis P. Henn.

mikaniae-cordiae Yen

mikaniicola Stev.

momordicae Mend.

momordicae Saw.

mori Hara

mori Mar. & Stey.

muntingiae Petr. & Cif.

minutipes Yen

neovignae Yam.

nicotianae Ell. & Ev.

nicotianicola Yen

passiflorae-foetidae Yen

peltophori Yen

penicillata var. apii Fuck.

pergulariae Yen & Lim

peristrophes-acuminatae Yen

personata (Berk. & Curt.) Ell. & Ev.

phaseolorum Cooke

phyllanthicola Shak. & Kam.

phyllanthicola Yen

phyllanthi-niruri Yen

piperata Asth. & Mah.

piperina Yen

piperis-muricata Yen

polygonicola Kar & Manal

polygonigena Yen

psophocarpi Yen

psophocarpicola Yen

pterocarpicola Yen

pudicae Yen

puerariicola Yam.

puerariina Yen

raciborskii Mats. & Nag.

raciborskii Sacc. & Syd.

rhododendricola Yen

salicina Ell. & Ev.

salicis-babylonicae Yen

scopariicola Yen

sechii Stev.

sesamigena Yen & Lim

singaporensis Yen

stahlii (Stev.) Sub.

stenolobiicola Chupp

stephanotidis Yen

stigmaphyllicola Yen

stuhlmanni P. Henn.

tabernaemontanae H. & P. Syd.

tabernaemontanicola Yen

tecomae Chupp & Viegas

tecomae-heterophyllae Yen

tecomicola Yen 154 ~

thunbergiae Boel. 188

thunbergiana Yen 165

thunbergiicola Yen 188

timorensis Cooke 188

tithoniae Bak. & Dale 165

tithoniae Chidd. 165

tithonicola (Chidd.) Yen 165

trematicola Yen 189

trematis-cannabini Yen & Lim 189

triseptopora Yen 166

ubi Rac. 159

vienae Ell. & Ev. 174

vignae Racib. 174

vignae-sinensis Tai & Wee 174

vienicaulis Tehon 156

viticola Yen & Lim 190

volkameriae Speg. 166

weberi Chupp 190

zinniae Ell. & Mart. 166

zinniae Takah. & Yosh. 166

Cercosporidium personatum (Berk. & Curt.) Deigh. 167

Cercosporiopsis canescens (Ell. & Mart.) Miura 156

Chingia peprigida (v.A.v.R.) Holtt. 15

sakayensis (Zeill.) Holtt. yi

Chisocheton divergens BI. 349

erythrocarpus Hiern. 349

macrophyllum King 349

patens BI. 349

paucifiorus King 349

pentandrus Merr. 349

spicatus Hiern. 349

Christella arida (D. Don) Holtt. 25

dentata (Forsk.) Brow. & Jermy 27

hispidula (Decne.) Holtt. 26

papilio (Hope) Holtt. 25

parasitica (L.) Lev. 26

subpubescens (BI.) Holtt. 26

Chrysanthemum sinense Sab. ex Sweet 172

Cicca acida Merr. 365

Citrus aurantifolia Swing. 343

grandis Osb. 343

hystrix DC. 343

limon Burm. f. 343

medica L. 343

microcarpa Bunge 343

paradisis Macf. 343

reticulata Blanco 343

sinensis Osb. 343

Cladosporum personatum Berk. & Curt. 167

Claoxylon indicum Hassk. 359

longifolium Endl. ex Hassk. 359

Clausena excavata Burm. f. 343

lansium Skeels 343

Cleistanthus heterophyllus Hk. f. 359

hirsutulus Hk. f. 359

macrophyllus Hk. f. 359

malaccensis Hk. f. 359

myrianthus Kurz 359

sumatranus (Mig.) M.-A. 359

Clerodendron fragrans (Vent.) Willd. 166

Codiaeum variegatum (L.) BI. 360

Coeladepas glanduliferum P. & H. 363

preyy fyoe

Coelostegia griffithii Benth.

Coleus

Commersonia bartramia Merr.

Connaropsis griffithii PI.

Corchorus acutangulus Lam.

Cordia cylindristachya

Coryphopteris arthroytricha Holtt.

badia (v.A.v.R.) Holtt.

gymnopoda (Bak.) Holtt. var. bintangensis Holtt.

gymnopoda var. gymnopoda

gymnopoda var. humilis Holit.

hirsutipes (Cl.) Holtt.

pectiniformis (C. Chr.) Holtt.

pectiniformis var. hirsuta Holtt.

tahanensis Holtt.

unidentata (Bedd.) Holtt.

viscosa (Bak.) Holtt.

Creochiton anomala (King & Stapf ex King) Veldk.

ledermannii Mansf. var. turbinata Maxw.

Crotalaria striata DC.

Croton argyratus BI.

caudatus Geisel.

griffithii Hk. f.

heterocarpus M.-A.

heteropetalum Ridl.

laevifolius BI.

Cyclosorus aridus (D. Don) Ching

contiguus (Rosen.) Ching

dicranogramma (v.A.v.R.) Holtt.

ecallosus Holtt.

extensus (Bl.) Ching

ferox (Bl.) Ching

glandulosus (Bl.) Ching

gongylodes (Schk.) Link

heterocarpus (Bl.) Ching

heterocarpus var. glaucostipes (Bedd.) Holtt.

interruptus (Willd.) Ching

interruptus (Willd.) Ito

larutensis (Bedd.) Ching

latipinna (Hk.) Tard.

megaphyllus (Mett.) Ching

papilio (Hope) Ching

parasiticus (L.) Farw.

polycarpus (Bl.) Holtt.

porphyricola (Copel.) Ching

stipellatus (Bl.) Ching

subpubescens (Bl.) Ching

sumatranus (v.A.v.R.) Ching

toppingii (Copel.) Ching

truncatus (Poir.) Far.

unitus (L.) Ching

Dacryodes laxa (Benn.) Lam.

rostrata (Bl.) Lam

Dapania racemosa Korth.

Daphniphyllum laurinum (Benth.) Baill.

Dalenia beccariana (Cogn.) Ridl. ex Nayar

beccariana var. matangensis Nayar

furfuracea Ridl.

Datura alba Nees

Delonix regia (Boj. ex Hk.) Raf.

Dioscorea

Dioscorea pyrifolia Kunth.

Diplectria anomala (King) Veldk.

, 190

beccariana (Cogn.) O. Ktz. 392

glabra (Merr.) Nayar ssp. glabra 313

glabra var. glabra 313

glabra ssp. kinabaluensis Veldk. 313

glabra var. kinabaluensis (Veldk.) Maxw. 313

glabra var. micrantha (Veldk.) Maxw. 313

glabra var. papuana (Mansf.) Maxw. 313

micrantha Veldk. 313

papuana Mansf. 313

viminalis (Jack) O. Ktz. 372

Dissochaeta sect. Omphalopus (Naud.) Baill. 313

alstoniit Nayar 318

annulata Hk. f. ex Triana

var. griffithii (Nayar) Maxw. 325

var. johannis-winkleri (Schwiz.) Maxw. 3135,°825

densiflora Ridl. 319

fallax (Jack) BI. 313

glandiformis Maxw. 313, 314

hirsuta Hk. f. ex Triana 319

intermedia BI.

var. leprosa (Bl.) Maxw. 315

var. sagittata (Bl.) Maxw. 315

johannis-winkleri Schwtz. 313

laeve Ohwi ex Maxw. 315,316

leprosa (Bl.) BI. 315

macrosepala Stapf 320

malayana Furt. 320

marumioides Cogn. 321

marumioides Furt. 321

microplectrosa Maxw. 316, 317

porphyrocarpa Ridl. 32]

pulchra (Korth.) Maxw. 318

reticulata BI. 32]

rostrata Korth.

var. alstonii (Nayar) Maxw. 318

var. densiflora (Ridl.) Maxw. 319

var. esetosa Maxw. 318, 319

var. floccosa Maxw. 319, 320

var. hirsuta (Hk. f. ex Triana) Maxw. 319

var. horrida (Bakh. f.) Maxw. 320

var. macrosepala (Stapf) Maxw. 320

var. malayana (Furt.) Maxw. 319, 320

var. porphyrocarpa (Ridl.) Maxw. 32]

var. setosa (Schwtz.) Maxw. 321, 326

sagittata BI. 315

sarawakensis (Nayar) Maxw. 321

setosa Schwtz. 321

spectabilis Maxw. 321

velutina Bl. var. reticulata (Bl.) Maxw. 321

Dryopteris badia v.A.v.R. 10

chlamydophora C. Chr. 12

falcinella v.A.v.R. , 21

norrisii Rosen. 21

pectiniformis C. Chr. 10

peltata v.A.v.R. 23

peltochlamys C. Chr. 20

perrigida v.A.v.R. 15

porphyricola Copel. 18

pterospora V.A.V.R. 19

sumatrana V.A.N.R. 26

toppingii Copel. 21

trichopoda C. Chr. 12

Dysoxylum acutangulum Miq. 350

cauliflorum Hiern.

costulatum Miq.

flavescens Hiern.

macrothyrsum Miq.

thyrsoideum Griff.

tubinatum King

Drypetes pendula Rid.

Durio griffithii (Mast.) Bakh.

oblongus Corn. non Mast.

singaporensis Rid.

zibethinus L.

Elaeocarpus ferrugineus Steud.

floribundus BI.

ganitrus Roxb.

eriffithii Mast.

hullettii King

jackianus Wall.

littoralis Kurz

macrocerus Merr.

masteri King

nitidus Jack

obtusus BI.

paniculatus Wall.

parvifolius Wall.

pedunculatus Wall.

petiolatus Wall.

polystachyus Wall.

rugosus var. singaporensis Ridl.

salicifolius King

sphaericus K. Sch.

stipularis Bl.

Embelia officinalis Gaertn.

Entoloma bicolor (Mass.) Hor.

burkillae Mass.

curtipes (Mass.) Hor.

discophorum Corn. & Hor.

flavidum (Mass.) Corn. & Hor.

limosum

lineum

longipes (Mass.) Hor.

sercellum (Fr.) Kum.

stylophorum (B. & Br.) Sacc.

tricolor (Mass.) Hor.

Erythroxylum cuneatum Kurz

nova-granatense Hiern.

Ervatamia coronaria

Euodia glabra BI.

robusta Hk. f.

roxburghiana Benth. ex Hk. f.

Eupatorium odoratum L. f.

Euphorbia antiquorum L.

atoto Forst. f.

hirta L.

milii Moul.

nerrifolia L.

pulcherrima Willd.

splendens Bo}.

thymifolia L.

tirucalli L.

Eurycoma longifolia Jack

Excoecaria agallocha L.

bicolor Hask.

Fagraea fragrans Roxb.

, 176

Fahrenheitia pendula (Hassk.) A.-S. 361

Ficus chartacea 176

Furtado, Caetano Xavier 302

Galearia fulva (Tul.) Mig. 361

Gelonium glomerulatum Hassk. 367

Glochidion brunneum Hk. f. 361

desmocarpum Hk. f. 362

glomeratum Boerl. 362

hypoleucum Boerl. 362

laevigatum Hk. f. 362

leiostylum Kurz 362

littorale BI. 362

microbotrys Hk. f. 362

perakense Hk. f. 362

rubrum BI. 362

sericeum Hk. f. 362

singaporense Gage 362

superbum Baill. 362

wallichianum M.-A. 362

zeylanicum Juss. 362

Gloriosa superba L. 160

Glycosmis chiorosperma Spr. 344

citrifolia Ridl. 344

lanceolata Spr. 344

malayana Ridl. 344

Gomphrena globosa L. 177

Goniopteris aspera Presl 22

repanda Fée 23

Gordonia acuminata Wail. 270

acutifolia (War.) Keng 310

alpestris (Krug. & Urb.) Keng 308

angustifolia (Brit. & Wils.) Keng 308

benitoensis (Brit. & Wils.) Keng 308

brandegeei Keng 310

curtyana (A. Rich.) Keng 309

ekmanii (Schm.) Keng 309

fruticosa (Schrad.) Keng 310

haemotoxylon Swartz 309

humboldtii Keng 310

lasianthus Ell. 305

moaensis (Marie-Vic.) Keng 309

obovata (Waw.) Keng 310

planchonii Keng 310

portoricensis (Krug. & Urb.) Keng 309

pubescens Cavan. 310

robusta (Kob.) Keng 31]

samuelssonii (O.C. Schm.) Keng 309

singaporeana Wal. ex Ridl. 306, 307

spathulata (Kob.) Keng 31]

speciosa Choisy 310

tomentosa (Mart. & Zucc.) Spreg. 31]

urbanii (O.C. Schm.) Keng 309

villosa Macf. 309

wrightii (Griseb.) Keng 309

Gossypium barbadense L. 334

Graphium laxum Ell. 172

Grewia acuminata Juss. 333

blattaefolia Corn. 332

hirsuta (Korth.) Kochum. 333

latifolia auct. non. Muell. 332

omphacarpa Miq. 333

umbellata Roxb. 333

Gyroporus 295

—)

Haemocharis alpestris Krug. & Urb.

angustifolia Britt. & Wils.

benitoensis Britt. & Wils.

integerrima (Mig.) Koord. & Val.

semiserrata (Nees) Camb. var. acutifolia Waw.

tomentosa Mart. & Zucc.

Helminthosporum manihotis Rang.

Stahlii F.L. Stev.

Hemiontis prolifera Retz.

Heritiera borneensis (Merr.) Kosterm.

elata Rid.

littoralis Dry. ex W. Ait.

simplicifolia (Mast.) Kosterm.

Hevea brasiliensis (HBK) M.-A.

Hibiscus abelmoschus L.

esculentus L.

mutabilis L.

rosa-sinensis L.

schizopetalus (Mast.) Hk. f.

surattensis L.

syriacus L.

tiliaceus L.

Hydrangea macrophylla (Thunb.) Ser. ex DC.

Hygrophorus firmus B. & Br.

Impatiens balsamina L.

Indorouchera griffithiana (PI.) Hall.

Ipomoea batatas (L.) L.

pes-caprae (L.) R. Br.

purpurea

Irvingia malayana Oliv. ex Benn.

Isariopsis griseola Sacc.

Ixonanthes icosandra Jack

reticulata Jack

Ixora chinensis Lmk.

javanica (Bl.) DC.

Jasminum grandiflorum L.

Jatropha curcas L.

hastata Jacq.

multifida L.

pandurifolia And.

podagrica Hk.

Kleinhovia hospita L.

Koilodepas longifolium Hk. f.

Lansium domesticum Corr.

Lantana aculeata L.

Laplacea curtyana A. Rich.

ekmanii O. C. Schm.

fruticosa (Schr.) Kob.

grandis T. S. Brand.

moaensis Marie-Vic.

pubescens Pl. & Lind.

robusta Kob.

samuelssonii O. C. Schm.

semiserrata (Nees) Camb. var. obovata Waw.

speciosa HBK

speciosa Kunth.

spathulata Kob.

urbanii O. C. Schm.

wrightii Griseb.

Lastrea ridleyi Bedd.

robinsonii Ridl.

unidentata Bedd.

Longetia malayana P. & H.

Lycium chinense Mill.

ait,

5.09%

BYE ioe

. Ol 166, G79

8 i:

7363

Macaranga conifera M.-A.

cornuta Corn.

gigantea M.-A.

griffithiana M.-A.

heynei I1.M. John.

hosei King ex Hk. f.

hullettii King ex Hk. f.

hypoleuca M.-A.

indica

javanica Hk. f.

maingayi Hk. f.

populifolia M.-A.

pruinosa M.-A.

punticulata Gage

recurvata Gage

trichocarpa M.-A.

triloba M.-A.

Macrolenes ciliatiloba (Baker f.) Bakh. f.

dimorpha (Craib) Maxw.

echinulata {Naud.) Bakh. f. var. esetosa

(Craib) Maxw.

griffithii Nayar

hirsuta (Cogn.) Maxw.

horrida Bakh. f.

rufolanata (Ridl.) Maxw.

stellulata (Jack) Bakh f.

var. ciliatiloba (Baker f.) Maxw.

subulata Maxw.

Macrothelypteris torresiana (Gaud.) Ching

Magistostigma malaccense Hk. f.

Mallotus albus (Roxb.) M.-A.

macrostachyus M.-A.

paniculatus M.-A.

Malvaviscus arboreus Cav.

Manihot esculenta Cr.

glaziovii M.-A.

utilissima Pohl

Marumia dimorpha Craib

hirsuta Cogn.

rhodocarpa Cogn. var. esetosa Craib

rufolanata Ridl.

Maxburretia furtadoana Dran.

Medinilla spirei Guill.

Melia azedarach L.

indica L.

Melochia corchorifolia L.

Melodendron montana BI.

montanum BL.

Memecylon acuminatissimum BI.

acuminatum Sm.

var. acuminatum

var. flavescens Cl.

amabile Bedd. var. malaccensis Cl.

ambiguum BI.

amplexicaule Roxb.

appendiculatum BI.

beccarianum Cogn.

hbrandisianum Craib

caeruleum Jack

var. pulchrum (Kurz) Cl.

caloneuron Mig.

campanulatum Cl.

cantleyi Rid.

. 365

ot Slade

capitellatum L.

cinereum King

confine BI.

constrictum Craib

cordatum Wall.

curtisii Burk. & Hend.

depokkense Bakh. f.

depressum Benth. ex Triana

dichotomum (Cl.) King

var. dichotomum

var. eugeniiflorum (Ridl.) Rid.

var. rotundatum (Craib) Maxw.

dissitum Craib

edule Roxb.

var. edule

var. ovatum (Sm.) Cl.

elegans Kurz var. dichotoma Cl.

elmeri Merr.

epiphyticum King

eugeniiflora Ridl. (“‘eugeniflora’’)

excelsum BI.

floribundum BI.

floridum RidI.

fruticosum King

garcinioides BI.

var. elongatum BI.

gigantifolium Elm.

globiferum Wall.

globosum Bakh. f.

gracilines Ridl.

var. rotundatum Craib

grande Retz.

var. horsfieldii (Miq.) Cl.

var. khasiana Cl.

var. pubescens Cl.

hepaticum BI.

heteropleurum BI.

var. olivaceum King

horsfieldii Miq.

hullettii King

intermedium BI.

kunstleri King

kurzii King

laevigatum BI.

var. sulcicarpum Furt. (in scheda)

lampongum Miq.

lancifolium Ridl.

laxiflorum Wall. ex Ridl.

lilacmum Zoll. & Mor.

longifolium Ridi.

maingayi Cl.

malaccense (Cl.) Ridl.

megacarpum Furt.

micranthum Bl.

microstomum Cl.

minutiflorum Mia.

multiflorum Bakh. f.

myrsinoides BI.

var. lilacinum (Zoll. & Mor.) King

nudum BI.

oleifolium BI. (‘‘oleaefolium’’)

oligoneuron BI.

ovatum Sm.

94. 107

106

101

104

76, 91

76, 106

107

101

paniculatum BI. 101

pauciflorum BI.

var. brevifolium Craib 104, 108

var. obovatum Furt. (in scheda) 102 |

var. pauciflorum 102

perakense Merr. 71

pseudo-nigrescens BI. 87

var. acuminatum Bl. (in scheda) 87

pubescens (Cl.) King 104

pulchellum Ridl. 102

pulchrum Kurz 9]

pulchrum Cogn. 91

rhodophyllum Bakh. f. 74

ridleyi Cogn. ex Ridl. 69

steenissi Bakh. f. 64

subtrinervium Maiq. 76

tenuifolium Ridl. 79

umbellatum Wall. 66

venosum Mert. 101

wallichii Rid. 106

Meniscium salicifolium Hk. 24

triphyllum Sw. 23

var. parishii Bedd. 24

Menisorus 2

Merope angulata Sw. 344

Mesophlebion beccarianum (Ces.) Holit. 12

chlamydophorum (C. Chr.) Holtt. 12

crassifolium (Bl.) Holtt. 13

motleyanum (Hk.) Holtt. 12

trichopodium (C. Chr.) Holtt. 12

Metathelypteris dayi (Bedd.) Holtt. 6

decipiens (Cl.) Ching 6

flaccida (Bl.) Ching 5

gracilescens (Bl.) Ching 6

Micrococca mercurialis Benth. 365

Mikania cordata (Burm. f.) B.L. Rob. 162

Mimosa pudica L. 164

Momordica charantia L. 158

Morus alba L. 180

Muntingia calabura L. 181, 333

Murraya exotica L. 344

koenigii Spr 344

paniculata Jack 344

Mycosphaerella berkeleyii W. A. Jenk. 167

cruenta (Sacc.) Lath. 174

Neesia altissima BI. 337

malayan Bakh. 337

Neodissochaeta sarawakensis Nayar 321

Nephrodium beccarianum Ces. 12

dayi Bedd. 6

glaucostipes Bedd. 19

gracilescens var. hirsutipes Cl. 8

gymnopodium Bak. 9

larutense Bedd. 17

motleyanum Hk. 12

papilio Hope 25

pennigerum Hk. 18

var. malayense Bedd. 21

polytrichum Bak. 12

sakayensis Zell. 14

terminalis Hk. 27

viscosum Bak. 9

Nicotiana tabacum L. 163

Omphalopus fallax (Jack) Naud. 313

—

var. novoguineensis Mansf.

Ostodes macrophyllus Benth.

Oxalis barrelieri L.

corniculata L.

corymbosa DC.

Paramignya egriffithii Hk. f.

longispina Hk. f.

scandens Craib. var. ridleyi Sw.

Parathelyteris beddomei (Bak.) Ching

glanduligera (Kuntze) Ching

nipponica (Fr. & Sav.) Ching

Passalora personata (Berk. & Curt.) Shak.

Passiflora foetida L.

Pedilanthus tithymaloides Poit.

Peltophorum pterocarpum (DC.) Back.

Pentace triptera Mast.

Pergularia minor

Peristrophe acuminata

Phaeoisariopsis griseola (Sacc.) Ferr.

Phaseolus vulgaris L.

Phegopteris rubicunda v.A.v.R.

Phyllanthus acidus (L.) Skeels

arnarus Sch.

emblica L.

niruri L.

pulcher Wall. ex M.-A.

urinaria L.

Pimeleodendron griffithianum Benth.

Piper muricatum BI.

Pneumatopteris callosa (Bl.) Nakai

ecallosa (Holtt.) Holtt.

truncata (Poir.) Holtt.

Polygala brachystachya BI.

paniculata L.

Polypodium dentatum Forsk.

paludosum BI.

parasiticum L.

pennigerum Forst.

rectangulare Zoll.

truncatum Poir.

unitum L.

Polygonum

Polyspora axillaris (Roxb. ex Ker) Sw.

Polystichum torresianum Gaud.

Porphyrellus gracilis (Peck) Sing.

Pronephrium affine (BI.) Holtt.

asperum (Presl) Holtt.

glandulosum (BI.) Holtt.

lineatum (BI.) Presl

menisciicarpum (BI.) Holtt.

X parishii (Bedd.) Holtt.

peltatum (v.A.v.R.) Holtt.

repandum (Fée) Hoiit.

rubicundum (v.A.v.R.) Holtt.

salicifolium (Hk.) Holtt.

triphyllum (Sw.) Holtt.

Pseudocercospora abacopteridicola (Yen & Lim) Yen

abelmoschi (Ell. & Ev.) Deigh.

acalyphicola (Petr.) Deigh.

acalyphigena (Yen) Yen

allamendae (Yen) Yen

aranetae (Borl. & Rold.) Yen

asystasiae (Yen) Yen

bore 175

182, 365

169

169, 183

155, 169

bauhiniae (H. & P. Syd.) Deigh. 170

bradburyae (Young) Deigh. 172

byliana (Syd.) Yen 169, 170

cassiae-alatae (Yen & Lim) Yen 171

centroseinaticola (Yen & Lim) Yen 171

chrysanthemicola (Yen) Deigh. 172

columnaris (Ell. & Ev.) Yen 172

cordiicola (Yen) Yen 173

crotalaricola (Yen) Yen 173

cruenta (Sacc.) Deigh. 174

daturina (Yen) Deigh. 174

delonicus (Yen) Yen 175

ervatamiae Yen & Lim 175

eupatorii-flormosani (Saw.) Yen 175

fici-chartaceae (Yen & Lim) Yen 176

formosana (Yam.) Deigh. 176

globosae (Yen) Deigh. 177

hiratsukana (Tog. & Kats.) Deigh. 177

ipomoeae-purpureae (Yen) Yen 177

ixorae (Solh.) Deigh. 178

ixoricola (Yen) Yen 152, 178

jasminicola (Muell. & Chupp) Deigh. 17%

jatrophae-curcas (Yen) Deigh. 178

katongensis (Yen) Yen 179

lyciicola (Yen) Yen 179

macarangae (H. & P. Syd.) Deigh. 180

mori (Hara) Deigh. 180

muntingiae (Petr. & Cif.) Deigh. 180

nigricans (Cooke) Deigh. 158, 185

peltophori (Yen) Yen 18]

perguiariae (Yen & Lim) Yen 181

phylianthi (Chupp) Deigh. 182

phyllanthi-niruri (Yen) Yen 181

piperina (Yen) Yen 182

piperis-muricati (Yen) Yen 182

psophocarpi (Yen) Deigh. 164, 183

pterocarpicola (Yen) Yen 183

puerariicola (Yam.) Deigh. 183, 184

puerariina (Yen) Yen 184

salicina (Ell. & Ev.) Deigh. 184

scopariicola (Yen) Deigh. 185

singaporensis (Yen) Yen 185

stahlii (F.L. Stev.) Deigh. 185

stephanotidis (Yen) Yen 186

stigmaphyllicola (Yen) Deigh. 186

tabernaemontanae (H. & P. Syd.) Deigh. 175. 187

tabernaemontanicola (Yen) Yen 187

thunbergiicola (Yen) Deigh. 188

timorensis (Cooke) Deigh. 188

trematicola (Yen) Deigh. 189

trematis-cannabini (Yen & Lim) Deigh. 189

ubicola (Yen) Deigh. 159, 189

viticicola (Yen & Lim) Yen 190

Pseudodissochaeta spirei (Guill.) Veldk. & Maxw. 324

Pseudophegopteris paludosa (Bl.) Ching 5

rectangularis (Zoll.) Holtt. 5

Psophocarpus tetragonolobus DC. 164, 169, 183

Pteris interrupta Willd. 13

Pternandra coerulescens Jack 100 :

galeata (Korth.) Ridl. 100 |

Pterocarpus indicus Willd. 183

Pterococcus corniculatus P. & H. - 366

glaberrimus Hassk. 366

Pterospermum diversifolium BI. 338

—

=e ara

javanicum Jungh.

Ptychopyxis caput-medusae Ridl.

costata Miq. var. oblanceolata A.-S.

Pueraria phaseoloides (Roxb.) Benth.

Pyrenaria acuminata Pl. ex Choisy

garrettiana auct. non Craib

johorensis Keng

Kunstleri King

kunstleri auct. non King

lanceolata T. & B.

lasiocarpa Korth.

masocarpa Korth.

mindanaensis Merr.

oidocarpa Korth.

pahangensis Keng

parviflora Ridl.

serrata BI.

var. kunstleri (King) Keng

var. masocarpa (Korth.) Keng

var. serrata

tawauensis Keng

villosula Miq.

viridifolia Sym.

wrayi King

Quassia indica (Gaertn.) Noot.

Rhododendron

Ricinus communis L.

Roucheria griffithana PI.

Ruta graveolens L.

Salix babylonica L.

Salomonia cantoniensis Lour.

Samadera indica Gaertn.

Sandoricum indicum Cav.

koetjape (Burm. f.) Merr.

Santiria apiculata Benn.

eriffithii (Hk. f.) Engl.

laevigata Bl.

laxa Benn.

Sapium discolor M.-A.

Sarcotheca griffithii Hall. f.

Sauropus albicans Bl.

androgynus Merr.

Scaphium affine Pierre

linearicarpum Pierre

macropodum (Miq.) Beu. ex Heyne

Scoparia dulcis L.

Sebastiania chamaelea M.-A.

Septogloeum arachidis Racib.

manihotis Zimm.

Sesamum orientalis L.

Sida acuta Burm.

carpinifolia L.

cordifolia L.

rhombifolia L.

Sloanea javanica Szy. ex K. Sch.

Sphaerostephanos asplenioides J. Sm.

hendersonii Holtt.

heterocarpus (BI.) Holtt.

larutensis (Bedd.) C. Chr.

latebrosus (Mett.) Holit.

norrisii (Rosen.) Holltt.

peltochlamys (C. Chr.) Holtt.

penniger (Hk.) Holtt.

Ps po

5 Os

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th?

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iA pe

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W206, Z

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y2Te,

284,

272

288

306. 307

ae |

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, 286

, 288

polycarpus (Bl.) Copel.

porphyricola (Copel.) Holtt.

pterosporus (v.A.v.R.) Holtt.

var. altilobus Holtt.

var. pterosporus

toppingit Copel.

unitis (L.) Holtt.

Sphaerostylis malaccense P. & H.

Stenella fagraeae Yen

Stenolobium stans (L.) Seem.

Stephanotis floribunda (R. Br.) Bron.

Sterculia bicolor Mast.

coccinea Jack

cordata BI.

elongata Ridl.

foetida L.

javanica R. Br.

laevis Wall.

macrophylla Vent.

nobilis Smith

parviflora Roxb.

rubiginosa Vent.

Stigmaphyllon ciliatum

Strobilomyces

Strychnos

Suregada multiflora Baill.

Swietenia macrophylla King

mahogani Jacq.

Synostemon acciformis G. L. Webs.

Tabernaemontana coronaria Willd.

malaccensis

Tarrietia simplicifolia Mast.

Tecoma heterophylla

stans (L.) HBK

Ternstroemia macrophylla Wall.

Thea lanceolata Pierre

Thelypteris arida (D. Don) Mort.

aspera (Presl) Reed

badia (v.A.v.R.) Ching

beccariana (Ces.) Reed

brunnea (Wall.) Ching

callosa (Bl.) Iwats.

chlamydophora (C. Chr.) Ching

ciliata (Benth.) Ching

crassifolia (Bl.) Ching

dayi (Bedd.) Nay. & Kaur

dentata (Forsk.) E. St. John

ecallosa (Holtt.) Reed

flaccida (Bl\.) Ching

gracilescens (Bl.) Ching

herbacea Holtt.

heterocarpa (Bl.) Mort.

hirsutipes (C\.) Ching

hispidula (Decne.) Reed

immersa (Bl.) Ching

interrupta (Willd.) Iwats.

larutensis (Bedd.) Reed

latebrosa (Mett.) Reed

malayensis (C. Chr.) Reed

megaphylla (Mett.) Iwats.

menisciicarpa (Bl.) Iwats.

motleyanum (Hk.) Holtt.

opulenta (Kaulf.) Fosb.

oppositipinna (v.A.v.R.) Ching 5

paleata (Copel.) Holtt. 12

paludosum (BI.) Iwats. 5

papilio (Hope) Iwats. 25

parasitica (L.) Iwats. 26

pectiniformis (C. Chr.) Ching 10

peltata (v.A.v.R.) Reed a3

peltochlamys (C. Chr.) Reed 20

pennigera (Forst.) H.H. All. 19

polycarpa (BIl.) Iwats. 18

porphyricola (Copel.) Ching 18

prolifera (Retz.) Reed 13

pterospora (v.A.v.R.) Reed 20

rectangularis (Zoll.) Nay. & Kaur 5

rubicunda (v.A.v.R.) Iwats. 24

sakayensis (Zeill.) Reed 14

salicifolia (Hk.) Reed 24

singalanensis 6

subpubescens (Bl.) Iwats. 26

terminalis (Hk.) Tag. & Iwats. 27

torresiana (Gaud.) Alst. 5

triphylla (Sw.) Iwats. 23

var. parishii (Bedd.) Iwats. 24

truncata (Poir.) Iwats. 15

uliginosa (Kunze) Ching 5

unidentata (Bedd.) Holtt. 8

unita (L.) Mort. 18

viscosa (Bak.) Ching is

Thespesia populnea (L.) Sol. ex Corr. 336

Thunbergia alata Boi. ex Sims 165, 188

Tithonia diversifolia (Hemsl.) A. Gray 166

Trema cannabina Lour. 189

orientalis (L.) BI. 189

Trigoniastrum hypolecum Mig. 352

Trigonochlamys grandifolia Ridl. 346

eriffithii Hk. f. 347

Trigonospora ciliata (Benth.) Holtt. 11

Trigonostemon longifolius Baill. 367

Triomma malaccensis Hk. f. 347

Triphasia trifolia P. Wils. 344

Triumfetta bartramia L. 333

pseudocana Spr. & Craib 3553

rhomboidea Jacq. 333

Turraea breviflora Rid. 351

Urena lobata L. 333

Uromyces aparganii Clint. & Peck 155

Viena sinensis (L.) Savi ex Hassk. 174

Vitex pubescens Vahl 190

trifolia L. 190

Wikstroemia fruticosa Schr. 310

Xanthophyllum affine Korth. 352

discolor Chod. 352

ellipticum Korth. 352

eriffithii var. curtissi (King) Ng 352

kingit Chod. 352

maingayi Benn. 352

obscurum Benn. 352

palembanicum Mig. 352

Xylocarpus granatum Koen. 351

moluccensis (Lmk.) Roem. 351

Zanthoxylum hirtellum Ridl. 344

nitidum DC. 344

Zinnia elegans Jacq. 167

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