CONTENTS

Vol. XXXIV

Pages

PART I — Ist June, 1981

J.F. MAXWELL: A Revision of Pternandra (Melastomataceace).................. 1—90

a An st) A mevysion of Petradmm (Rutacese) <2... 5 ke ee ee ee 91-131

ernst) ee) oe Pern Geaus Jectario i Malaya <5 2620.02). st ee ee 132-147

LIM-HO Chee Len: Experimental Findings of the Tissue Culture of Orchid Hybrids at

GE, Ma Oe ete ee ge a na ar a 148-160

K. SWARUPANANDAN, A.R.R. MENON. and K. BALASUBRAMANYAN: A New

ae AICI AEP MRCUICSOSAMMSIE or 5 ohh St, toate fed wee kee et eels et ee 161-169

A.G. PIGGOTT: Changes in the Fern Flora of Gunong Ulu Kali. ............... 170-176

B.C. STONE: A New Species of Pandanus (Pandanaceae) from Burma ............ 177-179

Benito C. TAN: Orthorrynchium elegans (Phyllogoniaceae): new record for the

eS eS ee aah atars emer e. eelea i a Bt e e 180-183

J.F. MAXWELL: Botanical Note: how to cite the name of Miss Joaquim’s orchid ..... 184-186

Book Review: Grasses in Malayan Plantations by P.R. Wycherley and Ahmad

ee RCN VISE a. Seen OL ah ee RE i oe a as a es, nt Kms s 187

PART II — 15th March, 1982

WONG Yew Kwan: Horticultural Notes on the Angsana (Pterocarpus indicus)........ 189-201

LIM-HO Chee Len: Tissue Culture of Curculigo latifolia (Hypoxidaceae)........... 203 —208

B.C. STONE: Contributions to the Flora of the Solomon Islands .............. 209-222

Nomenclature of Joinvillea (Joinvilleaceae)..................2.. 223—225

ln Memoriam: Dt Monte Gress Manuel... sw ee ee ee 227

Subash CHANDRA: Some Aspects of Interrelationships among Drynarioid Ferns. .... . 229—238

Jennifer NG: Jn-Vitro Pollen Germination of Cassia fistula ................... 239-242

IE So EER CITA 0, ent ty es oT le oe Me a Pana a 243-248

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> J.F. MAXWELL:

3 A Revision of Prernandra (Melastomataceae) RR ee o es 1-90 “

a T.G. HARTLEY:

\ A Revision of the Genus Tetradium (Rutaceae) ... hee bie 73 91-131

R.E. HOLTTUM:

‘N The Fern Genus Tectaria in Malaya ae eo es aA Wes 132-147

CL. Lin-Ho:

2 Experimental Findings of the Tissue Culture of Orchid yas at the

Singapore Botanic Gardens se x; = 148-160

y K. SWARUPANANDAN, A.R.R. MENON, AND K. BALASUBRAMANYAN:

r A New Key for Biological Identification _... A ae << “ 161-169

ae A.G. PIGGOTT:

Ay Changes in the Fern Flora of Gunong Ulu Kali _.... 7 we ae 170-176

- B.C. STONE:

y A New Species of Pandanus (Pandanaceae) from Burma “ae Je; 177-179

() 8B. TAN:

B\4 Orthorrhynchium elegans (Hook. f. et Wils.) Reichdt. Spee ceo maneee),

New Record for the Philippine Moss Flora 73 180-183

J.F. MAXWELL:

Botanical Note: How to Cite the Name of Miss Joaquim’s Orchid ... 184-186

Book Review: P.R. Wycherley and Ahmad Azli bin Mohd. Yusof,

Grasses in Malayan Plantations y 187

Published by the Botanic Gardens

Parks and Recreation Department

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cover. , end

THE GARDENS’ BULLETIN

SINGAPORE

Vol. 34: 1 1 June 1981

CON TEN TS

PAGES

J.F. MAXWELL:

A Revision of Pternandra (Melastomataceae) ‘f. Fr. me. 4 1- 90

T.G. HARTLEY :

A Revision of the Genus Tetradium (Rutaceae) ... 2 ¥. 2: 91-131

R.E. HOLTTUM:

The Fern Genus Tectaria in Malaya B° ss i - 7” 132-147

C.L. Lim-Ho:

Experimental Findings of the Tissue Culture of Orchid 3 she at the

Singapore Botanic Gardens “9 9 , 148-160

K. SWARUPANANDAN, A.R.R. MENON, AND K. BALASUBRAMANYAN :

A New Key for Biological Identification .... i zs 7 it 161-169

A.G. PIGGOTT :

Changes in the Fern Flora of Gunong Ulu Kali... % x #. 170-176

B.C. STONE:

A New Species of Pandanus (Pandanaceae) from Burma Py bie 177-179

B. TAN:

Orthorrhynchium elegans (Hook. f. et Wils.) Reichdt. cscrcarpecbate

New Record for the Philippine Moss Flora Y 180-183

J.F. MAXWELL:

Botanical Note: How to Cite the Name of Miss Joaquim’s Orchid... 184-186

Book Review: P.R. Wycherley and Ahmad Azli bin Mohd. Yusof,

Grasses in Malayan Plantations ‘a: 187

Published by the Botanic Gardens

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A REVISION OF THE GENUS PTERNANDRA (MELASTOMATACEAE)

J.F. MAXWELL

Botanic Gardens, Singapore

ABSTRACT

Pternandra (Melastomataceae) is revised; the genus Kibessia is reduced to synonymy,

as it differs only in one structural detail (the calyptrate calyx). Fifteen species and two

varieties are recognized in this revision, of which two species (P. angustifolia and P.

crassicalyx) are new. Two taxa are retained at the rank of variety for the first time (P.

azurea var. cordata (Korth.) Maxwell, and P. galeata var. elmeri (Merr.) Maxwell).

Thirteen previously described taxa of Pternandra and Kibessia are here reduced to

synonymy. Descriptions, morphological discussion, key to taxa, illustrations of some

taxonomically important features, and photographs of some type specimens have been

included.

INTRODUCTION

Revived interest in the Malaysian flora and the Flora Malesiana project subsequent to

the end of World War II has greatly increased both the quantity and quality of herbarium

specimens available for research. This has certainly been true for Pternandra (including

Kibessia), and both prompted and facilitated the completion of this revision. Nonethe-

less, the present paper is not really an attempted definitive treatment. The Philippine and

New Guinean materials are still not satisfactory and work remains to be done in collect-

ing as well as interpreting these plants in the eastern part of the Malay Archipelago. How-

ever, it is hoped that this paper will resolve some of the taxonomic confusion that has

plagued both foresters and botanists working with Pternandra since the time of William

Jack (1820). Comments on, additons to, and critiques of this revision will be greatly

welcomed by the author.

Acknowledgements: Prof. Hsuan Keng (National University of Singapore) provided a

constant source of advice and criticism, and Dr. Chang Kiaw Lan (Curator, Herbarium of

the Botanic Gardens, Singapore) graciously cooperated in my work by permitting me to

study and borrow the Gardens’ Herbarium specimens. In the period May—July 1977 |

visited the Rijksherbarium, Leiden, Netherlands, a trip arranged by Drs. R. Geesink,

who also arranged my accommodation; thanks are due both to him and his wife, Dia

Geesink M. D., for their hospitality and help. I wish also to thank Prof. Dr. C. Kalkman,

Dr. C. G. G. J. van Steenis, Dr. R. C. Bakhuizen van den Brink, Dr. W. Vink, and

especially Dr. H. Sleumer and Dr. J. F. Veldkamp at Leiden for preparing the Latin

diagnoses in this paper. Dr. D. G. Frodin (University of Papua New Guinea) and Dr. M. G.

Manuel (formely of the University of Malaya) read the manuscript and offered several

constructive comments which have been incorporated herein. | am especially grateful to

Dr. B. C. Stone (University of Malaya) for editing the manuscript and for his continued

interest in my research. For access to specimens I am indebted to the Keepers and

Curators of the Herbarium, Royal Botanic Gardens, Kew; the Herbarium of the National

Botanic Garden, Brussels; the Herbarium of the University of Utrecht, Netherlands; and

for loans of several type specimens to the British Museum (Natural History) and the

University of Florence. In addition, Mrs. Chiryayupin Chandraprasong formerly of the

Herbarium, Department of Agriculture, Bangkok (BK) and Dr. Tem Smitinand of the

2 Gard. Bull. Sing. 34:1 (1981)

Royal Thai Forest Herbarium (BKF) kindly made available materials for study. Mr.

Jumali Kafrawi (National University of Singapore herbarium technician) kindly redrew

several floral diagrams and Mrs. Lucia Sunderaj Krueger completed plates 5, 8,9 and 13.

HISTORY OF THE GENUS PTERNANDRA

Pternandra(Greek pterna = heel; aner, andros ='man) was established by Jack in 1822

with the type species P. coerulescens. The generic name refers to the heel-like extension

of the anther connective. Jack’s type material originated from Penang. In 1828, De

Candolle described Kibessia on the basis of Melastoma azurea Bl. Kibessia is distinguished

by the presence of a calyx cap and setose, often branched, bristles on the calyx tube and

cap. Ewyckia was described by Blume in 1831, with E. cyanea as type species. This was

listed immediately after Kibessia and differs in having a truncate calyx which is obscurely

4-lobed, and a much smoother tessellate calyx pattern. Apparently Blume was unaware

of Jack’s publication since P. coerulescens and E. cyanea are essentially the same (there

is no calyx cap and the calyx pattern is tessellate).

In 1849 Blume established two more genera which are considered to be synonyms

of Pternandra. The first, Rectomitra, based on Eywickia galeata Korth., differs from

Fywickia in having a calyx cap which splits into several lobes and a verrucose to tubercu-

late calyx pattern. Blume obviously considered the ornamentation of the calyx tube more

important than the cap in his generic distinctions, otherwise he would have placed E.

galeata in Kibessia. The second, Macroplacis, based on Kibessia cordata Korth., was

distinguished on the basis of vegetative characters from Kibessia; the floral features of this

species and of K. azurea (Bl.) DC. are the same.

Naudin (1852) reduced Pternandra to Eywickia,but maintained Kibessia, Rectomitra,

and Macroplacis as distinct genera. The last genus to be described on the basis of a Pter-

nandra specimen was Apteuxis Griff. (1854), based on a Burmese plant which Griffith

named A. trinervis. Miquel (1855) considered Pternandra and Rectomitra as synonyms of

Eywickia, but maintained Kibessia and Macroplacis. Bentham and Hooker (1865) re-

cognized Pternandra and reduced Rectomitra and Macroplacis to synonyms of Kibessia.

Triana (1871) maintained Pternandra, Kibessia, and Rectomitra, but combined

Eywickia with Pternandra and Macroplacis with Kibessia. The bases for his generic

distinctions were thus: Pternandra — no calyx cap; Kibessia — calyx cap falling off as a

unit; Rectomitra — calyx cap splitting into several lobes. Kurz (1877) accepted Pternan-

dra and listed only P. coerulescens Jack and P. capitellata Jack for his accout of the

Burmese forest flora. C. B. Clarke (1879) recognized both Pternandra and Kibessia in

his acount of the Melastomataceae in the Flora of British India, including three species

of Pternandra (all from the Indian subcontinent) and four species of Kibessia (all from

the Malay Peninsula). Of the latter, two are now regarded as synonyms of P. coerule-

scens. Kibessia simplex,as described by Clarke,is not Korthals’ species but is P. echinata

Jack; K. pubescens Decne. and K. acuminata Decne. are also now regarded as P. echinata;

and K. tuberculata (Korth.) Hk. f. ex Cl. is a distinct species now called P. tuberculata

(Korth.) Nayar (1978). Thus, in Clarke’s treatment only three species of Pternandra

continue to be recognized, and only P. coerulescens Jack is found in India. Clarke correct-

ly reduced Griffith’s Apteuxis trinervis to Pternandra coerulescens. Clarke distinguished

Pternandra from Kibessia by its lack of a calyx cap, small cymes, and smooth calyx

pattern. Clarke’s treatment does not include any taxa of Eywickia, Rectomitra, or Macro-

placis, so no information on the status or relationships of those taxa is present either.

Maxwell: Prernandra (Melastomataceae) 3

Baillon (1890) was the first botanist to combine Kibessia and Pternandra, but

Cogniaux (1891),in his monograph of Melastomataceae, continued to distinguish these

two genera, although he combined Rectomitra with Kibessia. Cogniaux divided Kibessia

into three sections, based on Kibessia sensu stricto (Eukibessia Cogn.), Rectomitra (as

Sect. Rectomitra (Bl.) Cogn.), and Macroplacis (as Sect. Macroplacis (Bl.) Cogn.). Krasser

(1893),in a treatment based on Cogniaux, kept both Kibessia and Pternandra.

In King’s (1900) treatment of the Malayan Melastomataceae, three species of

Pternandra (P. coerulescens, with three varieties; P. echinata, and P. griffithii King) were

recognized. Merrill (1921) enumerated four Bornean species of Pternandra and twelve

Bornean species of Kibessia; among these sixteen species twelve are maintained in the

present account as distinct taxa.

In Ridley’s (1922) Flora of the Malay Peninsula six species of Pternandra were

recorded, one of which (P. galeata (Korth.) Ridl.) had been included in Kibessia by

Cogniaux. Mansfield (1925) kept Kibessia and Pternandra as distinct genera in his list of

three taxa found in New Guinea. Burkill (1935), in his recombination of Kibessia azurea

Bl. to Pternandra azurea (Bl.) Burk., also noted that Kibessia DC. should be reduced to

a synonym of Pternandra Jack.

In the latest work on Malesian Mealstomataceae, Bakhuizen v. d. Brink (1943,

1943-5) also maintained both Pternandra and Kibessia on the traditional basis. He

discusses nine species and two varieties of Kibessia, and one species and two varieties of

Pternandra. Of these, seven species are now still regarded as distinct. The varieties in

Kibessia are not now kept; neither are the varieties in Pternandra.

Corner (1952) in his book Wayside Trees of Malaya lists Pternandra only, including

3 species.

Backer and Bakhuizen v. d. Brink (1963) in the Flora of Java list one species, P.

azurea (Bl.) Burk., and indicate in a footnote that they consider Kibessia as a synonym

of Pternandra.

Nayar (1978) transferred 18 taxa of Kibessia to Pternandra. Unfortunately, the

overall value of the publication is dubious since most of the nomenclatural changes were

made without any discussion. Of Nayar’s new combinations only seven are herein con-

sidered as denominating distinct taxa, including Pternandra cogniauxii Nayar, a new name

for Kibessia korthalsiana Cogniaux. Nine other recombined names are herein reduced to

synonyms. Pternandra ledermannii (Mansf.) Nayar is a dubious species with close affinities

to P. rostrata (Cogn.) Nayar. In my research studies at Leiden, Kew, and the British

Museum, I did not notice any of Nayar’s annotation labels on the relevant specimens

of Pternandra.Nayat proposed a new species, Pternandra moluccana which, however, is

based on a specimen in poor condition, that is unquestionably a species of Medinilla.

This specimen, at Leiden, does not have a calyx calyptra or 8 equal stamens without

‘appendages, as specified in Nayar’s description. Pternandra moluccana thus will become

a synonym under Medinilla when revisional work on that genus is carried out. Finally

it may be noted that Pternandra ‘birtella’ in Nayar’s paper should be P. hirtella (Cogn.)

Nayar.

GENERIC CIRCUMSCRIPTION OF PTERNANDRA

Since there is but one real structural difference which can be used to separate

Kibessia from Pternandra, that is the presence of a calyptra or calyx cap in the former

4 Gard. Bull. Sing. 34:1 (1981)

and its absence in the latter, an assessment of the importance of this character is required

and will determine the taxonomic result. In my opinion, this is not a feature that merits

generic distinction, nor should genera be separated on any single, isolated feature. In

the present treatment, therefore, Kibessia is submerged in Pternandra, the latter name,

being the oldest, thus being the correct name for this genus.

CONSTITUTION OF THE GENUS PTERNANDRA.

In this paper 15 species (with 2 varieties) of Pternandra are described, of which two

are newly proposed (P. angustifolia Maxw. and P. crassicalyx Maxw., both from Borneo).

Thirteen names are reduced to synonymy, four herbarium (in scheda) names are de-

valued, and one species attributed to Kibessia, and four species originally placed in

Pternandra, are removed to other genera. Three taxa (species) remain dubious, and a

fourth could not be evaluated herein because of unavailability of material for study.

TAXONOMIC TREATMENT

Pternandra Jack

W. Jack, Malay Misc. II (1822) 60; repr. Hook. Comp. Bot. Mag. I (1835) 157;

Calcutta J. Nat. Hist. 5: 15 (1843) 309. Bentham & Hooker f., Gen. Plant. 1: 2 (1865)

771. Triana, Trans. Linn. Soc. 28 (1871) 153. Kurz, For. Fl. Brit. Burma I (1877) 508;

J. As. Soc. Bengal 46: 2 (1877) 78 (Contr. Knowl. Burm. Fl., 30). Clarke in Hook. f.,

Fl. Brit. Ind. II (1879) 551. Baillon, Adansonia 12 (1876—9) 87; Hist. Pl. 7 (1880) 23,

62, part. Cogniaux in Boerlage, Handl. Fl. Ned. Ind. I (1890) 527; in DC., Monogr. Phan.

7 (1891) 1102. Krasser in Engler & Prantl, Die Nat. Pflanzenfam. 3: 7 (1893) 196. King,

J. As. Soc. Bengal 69, II: 1 (1900) 67 (Mat. Fl. Mal. Pen. 3, 475). Guillaumin in Le-

comte, Fl. Gen. Indo-Ch. 2 (1921) 923. Merrill, J. Str. Br. Roy. As. Soc. 86 (1921) 452.

Ridley, Fl. Mal. Pen. 1 (1922) 807. Mansfeld in Engler, Bot. Jahrb. 60 (1925) 137.

Craib, Fl. Siam. Enum.I; 4 (1931) 701. Bakuizen f., “Thesis” (1943) 323; Med. Mus.

Bot. Utrecht 91 (1943) 323; Rec. Trav. Bot. Neerl. 40 (1943—45) 323. Corner, Ways.

Trees Mal. ed. 2 (1952) 451. Backer & Bakhuizen f., Fl. Java 1 (1963) 371. Nayar, Bull.

Bot. Surv. India 17 (1978) 51, Keng, Gard. Bull. Sing. 31 (1978) 110.

Kibessia DC., Prodr. 3 (1828) 311; Mem. Melastom. (1828) 81. Endlicher, Ench. Bot.

(1841) 647, gen. 6265. Korthals in Temminck, Verh. Nat. Gesch. Bot. (1844) 253.

Blume, Mus. Bot Lugd.-Bat. 1 (1849) 7. Naudin, Ann. Sci. Nat. 3: 18 (1852) 262.

Miquel, Fl. Ned. Ind. 1 (1855) 569. Bentham & Hooker f., Gen. Plant. 1: 2 (1865)

771. Clarke in Hook. f., Fl. Brit. Ind. If (1879) 551. Cogniaux in Boerlage, Handl.

Fl. Ned. Ind. I (1890) 525; in DC., Monogr. Phan. 7 (1891) 1105. Krasser in Engler

& Prantl, Die Nat. Pfanzenfam. 3: 7 (1893) 195. Koorders-Schumacher, Syst. Verz.

2 (1910) 47. Merrill, J. Str. Br. Roy. As. Soc. 86 (1921) 450. Mansfeld in Engler,

Bot. Jahrb. 60 (1925) 137. Bakhuizen f., ““Thesis” (1943) 311; Med. Mus. Bot. Utrecht

91 (1943) 323; Rec. Trav. Bot. Neerl. 40 (1943—45) 311.

Ewyckia Blume, Flora 14 (1831) 524; Bijdr. Nat. Wet. 6 (1831) 265. Korthals, l.c. 254.

Walpers, Repert. Bot. Syst. 5 (1846) 724. Blume, l.c. (1849) 5. Miquel, l.c. 568.

Rectomitra Blume, l.c. (1849) 6. Naudin, l.c. 261. Triana, l.c. 23, 153.

Macroplacis Blume, |.c. (1849) 7. Naudin, l.c. 260. Miquel, l.c. 569.

Apteuxis Griffith, Notul. 4 (1854) 672.

Maxwell: Pternandra (Melastomataceae) 5

Trees to 25 m tall, diameter to 75 cm, less frequently shrubs. Upper branches

cylindric, angled, or winged; glabrous or pilose. Leaves opposite; blades chartaceous

to coriaceous, simple, entire, lanceolate to suborbicular, acute to acuminate, at base

narrowed, rounded, or cordate, 3—S-nerved from base, nerves sunken above, raised and

tapering below, intramarginal nerve thin, often merging with margin in apical part of

blade; petiole usually flattened; interpetiolar line or ridge distinct; stipules none. Inflore-

scence umbellate, cymose, paniculate, or nearly sessile and glomerulate, few to many-

flowered, from leafless or leafy nodes or terminal. Calyx tube campanulate, with an

external pattern (tessellate, umbonate, verrucose, or echinate plates, or setose appendages

which are barbed to branched); margin truncate and minutely 4-cusped, or with a

distinct calyx cap with or without tubercles or setae, which either falls off as one piece

or splits into 2—4 lobes. Petals 4, imbricate to the right, usually broad-ovate to sub-

orbicular, acute, at base truncate, thick or thin, glabrous. Stamens 8, equal, glabrous,

with flattened filaments and dolabriform to oblong, laterally flattened anthers with

antrorse aestivation, the connective thickened, straight, eglandular, usually with a minute

spur near the filament. Stigma subcapitate to cylindric, 4-grooved or -lobed; style slender;

ovary 4-locular, placentation parietal, ovules numerous. Fruit a berry, campanulate-

truncate, urceolate, or globose; the exocarp tessellate or echinate. Seeds numerous,

flattened, quadrangular-rhombic, glossy tan or light brown.

Type species: Pternandra coerulescens Jack

From other genera of Melastomataceae, this genus is distinguished by its woody

habit, tessellate to echinate calyx tube, calyptra or calyx cap in most species, small and

only minutely appendaged anthers, 4-locular ovary, and many-seeded fruit. The charac-

teristic leaf venation pattern (3—5 basal nerves) is usually a good field character, but

confusion on this basis with other plants such as Rhodamnia (Myrtaceae), Cinnamomum

(Lauraceae), and Strychnos (Loganiaceae), or with Memecylon of the same family, is

possible. Strychnos, however, is a genus of which the species are mostly woody climbers.

MORPHOLOGY

Habit. — Of the 17 taxa discussed herein, there are nine which consist exclusively of

trees, the remainder are either trees or shrubs. Species which are constantly or noticeably

arboreal are: P. coriacea, P. galeata, P. hirtella, P. tuberculata. | have seen four taxa in.

the field (P. coerulescens, P. echinata, P. galeata, and P. tuberculata), but data on the

other taxa has been gleaned from herbarium label notes. All taxa are interrestrial but one

species, P. cogniauxii,has apparently the capacity to grow as an epiphyte, as noted on two

specimens. P. echinata, though usually recorded as a tree, has twice been recorded as a

climber; however in Singapore I have seen this only in tree form. Certain taxa, as follows,

attain a considerable height and trunk diameter: P. azurea may reach 30 m and 75 cm;

P. galeata var. elmeri, 20 m and 45 cm; and P. tuberculata, 20 m and SO cm. Buttresses

are common in larger individuals and the bole is usually straight with a wide crown.

Bark and wood. — Typically the outer bark is thin (0.25 — 1 mm) with longitudinal

fissures or craks, is smooth, and peels off in narrow strips. In colour it ranges from grey-

brown to blackish, often mottled with various dark shades. The inner bark is 0.5 — 2 mm

thick, variable in pigmentation, white, reddish-brown, greenish-yellow, or dark red. The

wood is hard and usually yellowish or brownish. Browne ((Forest Trees of Sarawak and

Brunei, 1955) gives some useful information on the general features and wood anatomy.

Branchlets. — These are useful in identification, especially vegetative material; they

are either terete or flattened and grooved on two faces, or quadrangular below the upper

6 Gard. Bull. Sing. 34:1 (1981)

node and tapering to a terete condition, or quadrangular entirely, or even 4-winged,

with membranous undulate wings on the angles. Older branchlets and branches of all

taxa are terete and ungrooved, and without angles or wings.

The following taxa have entirely terete, cylindric internodes: P. crassicalyx, P. galeata

var. elmeri, and P. teysmanniana.

Five taxa have branchlets which are usually flattened and grooved, or are somewhat

quadrangular below the upper node: P cogniauxii, P. coriacea, P. galeata var. galeata, P.

multiflora, and P. rostrata.

Two taxa have quadrangular internodes: P. echinata, with the angles extending fully

from one internode to the next, and P. gracillis, with the internode flattened and grooved

on two faces below the upper node. In P. hirtella, the branchlets are basically quadrangular

but the angles are slightly winged.

Two taxa have distinctly winged branchlets: P. azurea var. cordata and P. tessellata.

There are six taxa with pubescent branchlets (P. azurea, both var. azurea and var.

cordata, P. crassicalyx, P. echinata, P. hirtella, and P. tessellata). All other taxa have

glabrous branchlets. P. echinata can be distinguished from all other taxa solely on the

basis of its 4-angled and pubescent branchlets.

The branchlet epidermis is thin, smooth, brownish or blackish, and may peel off (e.g.

P. coerulescens, P. multiflora) leaving a smooth, tan undersurface. The wings of the

branchlets in those taxa with this feature are fragile and may break off on the lower older

part of the stems, leaving an interrupted series of irregular rigdes, all of which however

eventually fall off, leaving smooth, cylindric, glabrous internodes. The upper internodes

are usually flattened, but fill out to cylindric form in age. Articulation zones at the base

of the petiole tend to coalesce and may be mistaken for stipular scars. Branch tips with

very young leaves show this feature clearly. The second node usually has a distinct inter-

petiolar ridge or line.

Leaves. — Usually these are simple, entire, glabrous or finely puberulent, 3—5-nerved

(except P. angustifolia, |-nerved) and may be chartaceous to coriaceous in texture. There

is always a pair of intramarginal nerves. Secondary venation is pinnate in plan from each

main nerve and is visible except in a few cases (the taxa with thickened leaf blades, e.g.

P. coerulescens and P. galeata var. elmeri). In shape the blades range from lanceolate to

suborbicular; the apex from rounded to acute, and the base from acute to cordate. The

petiole is usually flattened; it is glabrous except in those five taxa which have pubescent

branchlets. |

Some taxa are recognizable from their leaves: P. cogniauxii, with 5 main nerves and

a cordate base; P. coriacea, with small, coriaceous blades; P. teysmanniana, with blades

glaucous beneath; etc. The leaves of P. echinata, P. azurea var. azurea, and P. azurea var.

cordata are very similar. Also those of P. galeata, P. coerulescens; and P. rostrata, P.

tuberculata; and P. crassicalyx and P. hirtella are similar.

Inflorescences. — P. coerulescens has a wide range of structural variation in the

inflorescence, but most other taxa have definite inflorescence forms. That of P. multi-

flora is distinct in having the largest, most complex panicle (to 5 ramifications). Some

specimens of P. coerulescens may have inflorescences branched to the 4th degree, but

Maxwell: Pternandra (Melastomataceae) 7

most are less branched. Shortening of the ultimate axes result in a cyme-like or umbelli-

form appearance in some specimens of P. tessellata and P. coerulescens. In most taxa the

inflorescences are cymose and 3- or 1-flowered. Solitary flowers occur in P. azurea,

P. hirtella, and P. rostrata. In P. echinata such a transition may be seen, with the cymes

ranging from 9- to 3-flowered, or even 1-flowered. Some species have the inflorescence

condensed glomerulate, with few or solitary flowers, e.g. P. angustifolia, P. cogniauxii,

P. galeata, and P. gracilis.

Most species possess axillary inflorescences from leafy nodes, but in P. cogniauxii,

P. galeata var. elmeri, and P. multiflora, the nodes are leafless and thus the inflorescences

are ramuline. In P. coriacea the inflorescense axes are flattened and grooved, in P. rostrata

also but quadrangular, and in P. azurea, P. echinata, and P. multiflora they are clearly

quadrangular. All axes are glabrous except in the five taxa with pubescent branchlets.

The bracteoles and bracts are paired, connate, sheathing, and usually persistent. Only

those of P. echinata are manifest or variable enough to be of taxonomic importance.

Calyx. — In both flowering and fruiting material the calyx provides the most distinct

differences between he taxa. The structure of the calyx tube and the nature of the calyx

margin has allowed the taxonomic discrimination or, in contrast, the synonymous status,

of most of the taxa herein treated.

In P. galeata the calyx is urceolate and in the same species the variety elmeri is

campanulate. It may be ornamented externally with a distinctive pattern of 4-angled

flat or umbonate scales (tessellate pattern); or conical tubercles (tuberculate pattern);

or with flattened or cylindric appendages (echinate pattern) which may be entire or

branched and glabrous or pubescent. P. coerulescens always shows the tessellate pattern;

this also appears, sometimes, in P. galeata, which however can be distinguished by the

urceolate form of the calyx. The umbonate-tessellate pattern occurs in P. crassicalyx and

P. tessellata, while a more extreme form (conical-tuberculate) is common in e.g. P. cogni-

auxii, P. rostrata, and P. tuberculata. In P. coriacea and P. teysmanniana, for example,

the pattern nearer the pedicel is umbonate-tuberculate but the gradual elongation of the

umbos in the more distal scales reaches, at the margin, results in an echinate pattern.

A more distinctly echinate pattern, with appendages longer than wide, is common to

many taxa, though the shape of the appendage varies from setiform, as in P. azurea and

P. hirtella (setae often branched) to flattened in P. angustifolia and P. echinata (append-

agesentire).In P. azurea var. cordata the appendages are both flattened and branched.

The calyx margin is truncate, completely in P. coerulescens and P. multiflora. The

remaining 15 taxa have a calyx cap, or at least minute calyx lobes. Five taxa have a

smooth calyx cap which ultimately splits into 2, 3, or 4 persistent lobes: P. cogniauxii,

P. galeata, both varieties, P. teysmanniana, and P tuberculata. The remaining taxa have

a smooth calyx cap (P. crassicalyx and P. rostrata) or one with appendages near the

middle (P. azurea, P. hirtella) which falls off as a unit at the time of petal expansion.

Thus the mature flowers of these taxa appear to have a truncate (i.e. lobeless) calyx.

Examination of buds is: necessary to determine the difference, a crucial matter e.g. in

distinguishing P. multiflora from P. coerulescens. In P. tuberculata the calyx is 4-lobed,

a characteristic feature, and in P. rostrata the calyx is calyptrate, thus enabling distinction

of these quite similar species.

Internally the calyx shows 8 lines and 8 grooves corresponding in which the anthers

lie in the bud state. These correspond with the extraovarial chambers found in some

other genera of this family, i.e. those of the tribes Dissochaeteae and Oxysporeae.

8 Gard. Bull. Sing. 34:1 (1981)

Corolla. — The four petals are equal, symmetrical, usually broad-ovate to sub-

orbicular, acute, and truncate at the base. They are usually thin with visible veins

(P. gracilis) or thicke: with indistinct or invisible veins (e.g. P. coerulescens). As taxon-

omic features the petals are not especially useful and are often lacking in herbarium

specimens.

Stamens. — These are 8, glabrous, equal, with smooth, flat filaments in all taxa of

Pternandra. In contrast to most other genera of Melastomataceae, Pternandra has very

reduced and unspecialized anthers, similar in many ways to those of Astronia and (some)

Memecylon. The anthers are usually dolabriform, i.e. axe-shaped, or oblong, bilocular

(each opening by a longitudinal slit), and with a thickened, non-glandular connective

which often bears a spur near the filament. The stamens appear to be rather uniform in

the genus and are not very important in taxonomy.

Gynoecium and fruit. — They ovary is inferior, 4-locular; the style slender, glabrous,

and the stigma subcapitate to cylindric, at first 4-grooved and twisted, later 4-lobed. The

placentation is parietal, with numerous ovules (Fig. 1). The fruit is a berry, which tends

to retain its younger shape with a thickened, gritty pericarp and contains many flattened,

quadrangular-rhombic or cuneate seeds. In P. galeata the fruit (unlike the floral calyx) is

urceolate, but in all other taxa the fruit shape is pretty uniform. Seed morphology,

however, may play a promising taxonomic role and should be investigated more tho-

roughly.

DISTRIBUTION

P. coerulescens and P. echinata var. pubescens (Decne.) King occur in Thailand, the

former known from Chang Island in Trat Province, the latter (which is indistinguishable

from var. echinata) from throughout the southern peninsular part of Thailand. In the

Malay Peninsula and Singapore, four taxa occur: P. coerulescens, P. galeata, P. tuber-

culata, and P. echinata—the latter is apparently restricted to the Malay Peninsula and the

Riouw Archipelago. Ten taxa appear to be endemic to Borneo: P. cogniauxii, P. gracilis,

P. galeata var. elmeri, P. teysmanniana, are uncommon; P. azurea var. cordata, and

P. rostrata are common in Borneo, but also occur in Sumatra and Java, the latter also

in New Guinea. P. coerulescens, P. galeata s.s:,and P. tuberculata range from W. Malaysia to

New Guinea. Much more collecting must be done in Sumatra, Celebes, Moluccas, and

New Guinea to determine more precisely the taxa of Pternandra which they include.

In terms of relative abundance I have adopted five crude but suggestive categories:

RARE: P. angustifolia, P. coriacea, P. galeata var. elmeri, P. tessellata,

P. teysmanniana

NOT COMMON: P. azurea var. cordata, P. cogniauxii, P. crassicalyx, P. gracilis, P.

hirtella, P. multiflora

COMMON: P. azurea var. azurea, P. galeata var. galeata, P. tuberculata

VERY COMMON: | P. echinata, P. rostrata

MOST COMMON: P. coerulescens.

Maxwell: Pternandra (Melastomataceae)

Fig. 1. Pternandra

echinata Jack ovary, x-section, showing parietal placentae and ovules. Drawn

from Maxwell 76-757.

10 Gard. Bull. Sing. 34:1 (1981)

KEY TO THE TAXA OF PTERNANDRA

1. Calyx tube with setose to echinate scales which are distinctly longer than wide.

2. Scales branched near the tip, especially those near the calyx margin; calyx cap

or lobes tuberculate to echinate near middle.

3. Blades mostly narrowed or sometimes rounded, never cordate, at base;

petiole at least 3 mm long.

4. Branchlets and blades beneath densely red-brown pilose, the pube-

scence persistent; blades 9-17 cm long, 4.5—8 cm wide, coriaceous

Sx ark cheer eat eles Gendered | eee (12) P. hirtella (Cogn.) Nayar

4. Branchlets glabrous at maturity; nerves minutely pilose beneath near

petiole, glabrescent; blades generally thinner, smaller than as above

SF. pint. shel een nee ARE A ie, A (2) P. azurea (Bl.) Burk. var. azurea

3. Blades rounded and usually cordate at base; petiole up to 2mmlong .....

ithe se higant eles tha pale aera (3) P. azurea var. cordata (Korth.) Maxw.

2. All scales entire; calyx cap with or without tubercles or appendages.

5. Blades with 3 main nerves and 2 intramarginal nerves.

6. Blades beneath, branchlets, and inflorescence axes pubescent.

7. Scales flattened, never barbed; cap without tubercles or

appendages..N. 2). Sh auiiwreae (8) P. echinata Jack

7. Scales setose, the upper ones often barbed; cap with

tubergles, 3... Sa\2 se (2) P. azurea (B1.) Burk. var. azurea.

6. Blades beneath, branchlets, and inflorescence axes glabrous.

8. Blades coriaceous, glaucous beneath, generally

drying grey to whitish; inflorescence axes 1—1.5 cm

long; cap without tubercles... 7 rr vee ee

she pleats SHURE ale a RS (6) P. coriacea (Cogn.) Nayar

8. Blades chartaceous, green beneath; inflorescence axes

less than 5 mm long; cap with setose to echinate

appendages... on¥08 (11) P. gracilis (Cogn.) Nayar

5. Blades with 1 main nerve and 2 intramarginal nerves...........

ee Mere Yate (1) P. angustifolia Maxw.

1. Calyx scales tessellate, umbonate, verrucose, tuberculate; usually wider than long.

9. Scales tessellate or slightly umbonate near the margin.

Maxwell: Pternandra (Melastomataceae) 11

10.

Blades cordate at base, 5-nerved, 13—26 cm long, 7—13 cm wide; inflorescence mostly

hh RE ao eo a (4) P. cogniauxii Nayar

Blades narrowed or rounded at base, 3-nerved; inflorescence mostly from leafy

nodes; blades usually smaller than as above.

11. Calyx cap or lobes distinct; fruit urceolate or campanulate

12. Calyx tube and cap c. 0.25 mm thick, cap splitting into 4 persistent lobes;

fruit urceolate, the pericarp to 0.5 mm thick.

13. Blades subcoriaceous to coriaceous; calyx tessellate..............

sie ran hated hecWte Gece J 2 ae - (9) P. galeata (Korth.) Ridl. s.s.

13. Blades thick coriaceous; calyx umbonate to tuberculate...........

ee Patera 6 fi Bhi te sak +: (10) P. galeata var. elmeri (Merr.) Maxw.

12. Calyx tube and cap c. 1.5 mm thick, cap falling off in one piece; fruit

campanulate truncate, pericarp c. 1.5 mm thick....................

11. Calyx cap not developed; fruit globose.

14. Inflorescence 7—75 mm long; petals white, blue, or purple ......

AM £0tiwe tats dock dae aAVn emetic: (5) P. coerulescens Jack

15. Branchlets cylindric.

16. Blades subcoriaceous to coriaceous; cap smooth or tuberculate.

17. Blades broadly rounded to cordate at base, 5-nerved; inflorescences

mostly from behind the leaves; cap tessellate to tuberculate ........

Peek TEs da 5 ions VN cue phe Ke ets tina be (4) P. cogniauxii Nayar

17. Blades narrowed to rounded at base, 3-nerved; inflorescences mostly

from leafy nodes; cap smooth.

18. Inflorescence glomerulate, primary axes c. 1 mm long; pedicels

1—2 mm long; calyx tube 3—4 mm long; petals thin; fruit urceo-

late; leaf blades to 5 x 3 cm, petiole 1.5—2.5 mmlong. .........

| atte 5 Egat aie ae Pamaeae (16) P. teysmanniana (Cogn.) Nayar

12 Gard. Bull. Sing. 34:1 (1981)

18. Inflorescence umbellate, cymose, or of solitary flowers; primary axes and pedicels

generally over 2 mm long; calyx tube 4—6 mm long; petals thcik; fruit campanulate,

truncate; blades at least 6.5 cm long, 3 cm wide; petiole 2—8 mm long.

19. Cap splitting into 4 lobes; primary and secondary inflorescence axes puberulous

isan @ GAY Go EW aeh MOE Ge le Sh nee hee eae (17) P. tuberculata (Korth.) Nayar

19. Cap falling off in one piece; primary and secondary axes glabrous.

20. Inflorescence axes 1.5—2 mm thick; mature calyx 6 mm long, 5 mm wide;

leaf blades 6.5—21 cm long, 3—10 cm wide, drying light or dark brown

» “eMCAHIL becn.cas Ageia Sates eis OF Sposa (14) P. rostrata (Cogn.) Nayar

20. Inflorescence axes to 1 mm thick; mature calyx 4.5 x 4 mm; blades 4.5—6.5

cm long, 2.5—4 cm wide, usually drying greenish to yellowish beneath

tiation» “enrsh oon yale, ME EEREN Oo ete ee (6) P. coriacea (Cogn.) Nayar

16. Blades chartaceous.

21. Cap tuberculate; blades drying brownish or olive-green; inflorescence axes

glabrous: :\0'..- 'utdne eau oad tiene eee (11) P. gracilis (Cogn.) Nayar

21. Cap smooth; blades drying blackish to brown; inflorescence axes pilose ......

ort cart-ehye, equtot sos Real tesa et ine ee (17) P. tuberculata (Korth.) Nayar

LIST OF TAXA IN PTERNANDRA

1. P. angustifolia Maxw. — Sarawak, endemic.

2. P. azurea (Bl.) Burk. var. azurea. — Lingga Archipelago, Sumatra, Java, Borneo.

3. P. azurea var. cordata (Korth.) Maxw. — Borneo, Sumatra.

4. P. cogniauxii Nayar — Borneo, endemic.

5. P. coerulescens Jack — Thailand, W. Malaysia, Riouw Archipelago, Borneo, Celebes,

Moluccas, New Guinea, Lingga, Archipelago, Torres Strait.

6. P. coriacea (Cogn.) Nayar — Borneo, endemic.

7. P. crassicalyx Maxw. — Sarawak, endemic.

8. P. echinata Jack — Thailand, W. Malaysia and Singapore, Borneo.

9. P. galeata (Korth.) Ridl. var. galeata. W. Malaysia, Sumatra, Borneo, New Guinea.

10. P. galeata var.elmeri (Merr.) Maxw. — Borneo, endemic. |

11. P. gracillis (Cogn.) Nayar — Borneo, endemic.

12. P. hirtella (Cogn.) Nayar — Sarawak, endemic.

13. P. multiflora Cogn. — Sabah and Sarawak, endemic.

14. P. rostrata (Cogn.) Nayar — Borneo and Sumatra; one collection from New Guinea.

15. P. tessellata (Stapf) Nayar — Borneo, endemic.

16. P. teysmanniana (Cogn.) Nayar — Kalimantan, endemic.

17. P. tuberculata (Korth.) Nayar — W. Malaysia, Riouw Archipelago, Sumatra, Borneo,

New Guinea.

1. Pternandra angustifolia Maxw., sp. nov.

Rami cylindrici glabri, epidermide brunnescenti, deglubescenti; folia membranacea,

glabra lanceolata acuminata, basi acuta, uninervia, nervis intramarginalibus tenuibus

exceptis, lamina 55—75 mm longa, 12.5—20 mm lata, petiolis 1—2 mm longis. Inflore-

Maxwell: Pternandra (Melastomataceae) 13

scentia solitaria, axillaris, reducta uniflora, glabra, axi primario c. 5 mm longo, nodis 2,

pedicello c. 1 mm longo, calycis tubus echinatus glaber c. 4 mm longus; calyptra mammi-

formis laevis glabra tenuis apice acuminato, minutissime 2—4-dentato. Fructus ignotus.

Type: SARAWAK: Gunung Santubong, hillside, heath forest, alt. c. 200 m, Haron

21444 (L, holotype; isotypes in A, K, BO, KEP, MEL, and SING). Fig. 2.

Small tree. Upper branchlets cylindric, smooth, glabrous, c. 1 mm thick, epidermis

drying brown and peeling off leaving a smooth, tan coloured surface on older branchlets

and branches. Blades subcoriaceous, glabrous, lanceolate, acuminate at the tip (acumen

1—1.5 cm long), narrowed at the base; uninerved, midnerve sunken above, raised and

tapering below; intramarginal nerves faint, 1-2 mm from the margin and merging with

it in the upper part of the blade; secondary venation pinnate, c. 10—20 pairs, obscure

above, obscure to invisible below; 5.5—7.5 cm long, 1.25—2 cm wide, drying grey-green

above, brown below. Petioles 1—2 mm long, c. 1 mm thick, glabrous. Inflorescences of

solitary flowers from leafy nodes, axes slightly flattened, glabrous. Bracts and bracteoles

lanceolate, acute, c. 1 mm long, glabrous. Primary axes c. 5 mm long, with two nodes,

c. 0.5 mm thick at the base, pedicels c. 1 mm long. Calyx tube campanulate, densely

covered with glabrous, entire, echinate appendages with are shortest near the pedicel

(c. 1 mm) and longest (c. 2 mm) near the margin, c. 4 mm long, 4 mm wide, pale green

with a magenta hue near the base, drying brown; calyx cap mammiform, acute and 2—4

apiculate at the tip, smooth? thin, glabrous, 4—5 mm long, dehiscence unknown but

probably splitting into several pieces. Bud petals imbricating to the right, broadly ovate,

acute at the tip, truncate at the base, thickened with thinner margins, venation invisible,

c. 3.5 x 3.5 mm. Filaments c. 1 mm long; anthers c. 2 mm long. Stigma in bud cyllindric,

4-grooved, c. 1 mm long; style c. 5 mm long, glabrous. Fruit not seen.

Pternandra angustifolia Maxw. is readily distinguished from all other species of

Pternandra by its small, narrow blades which have a prominent midrib and two faint

intramarginal nerves. All other species of Pternandra have 3 or 5 nerves in addition toa

pair of intramarginal ones. The echinate calyx tube and smooth cap strongly resemble

P. echinata Jack, however the latter species is easily recognized by its thin, 3-nerved,

pilose blades; large bracts, and pilose, angled branchlets.

I have chosen the epithet augustifolia so that the most obvious distinguishing feature

of this species is indicated. P. angustifolia is in no way related to Kibessia angustifolia

Bl. which is probably the same as P. echinata Jack (q.v.). Fortunately Nayar, in his

relentless pursuit of botanical fame, overlooked a possible recombination of K. angusti-

folia to Pternandra in his recent uncritical paper (Bull. Bot. Surv. India 17 (1978) 51).

Fortunately, this oversight is insignificant when one considers all the confusion that

Nayar has created in his infuriating and inept meddlings in the Melastomataceae.

Fig. 2. A: calyx; B: bud petal; C: stamen.

Distribution:

Borneo

Sarawak — Gunong Santubong: Haron 21444 (holotype L, isotypes: K, SING, SAN,

A, BO, KEP, MEL) on a hillside of a heath forest at an altitude of c. 200 m.

Gard. Bull. Sing. 34:1 (1981)

Fig. 2. Pternandra angustifolia Maxw. A-C: Haron 21444 (holotype).

Maxwell: Pternandra (Melastomataceae) 15

2. Pternandra azurea (Bl.) Burk., Kew Bull. (1935) 319. Melastoma azureum Bl.,

Bijdr. Fl. Ned. Ind. 17 (1826) 1079. Kibessia azurea (Bl.) DC., Prodr. III (1828)

196; Cogniaux ex Winkler in Engl., Bot. Jahr. 48 (1912) 109: Merrill, J. Str. Br.

Roy. As. Soc. 86 (1921) 450; Bakhuizen f., ‘“‘Thesis” (1943) 314, Med. Mus.

Bot. Utrecht 91 (1943) 314, Rec. Trav. Bot. Neerl. 40 (1943—45) 314; Backer

& Bakhuizen f., Fl. Java I (1963) 371. Kibessia azurea (Bl.) DC. var. typica

Hochreutiner, Pl. Hochr. II (1925) 162; Heyne, De Nut. Pl. Ned. Ind. II (1927)

1205. Kibessia azurea Bl. var. subalata (Bl.) Miq., Fl. Ned. Ind. I (1855) 570;

Bakh. f. l.c. (1943, 1943—45) 316, syn. nov. Pternandra azurea (Bl.) Burk. var.

subalata (Bl.) Nayar, Bull. Bot. Surv. India 17 (1978) 52. Kibessia subalata Bl.,

Mus. Bot. Lugd.-Bat. I:] (1849) 8; Naudin, Ann. Sci. Nat. 3:18 (1852) 262;

Triana, Trans. Linn. Soc. 28 (1871) 152 (“‘subulata”’). Kibessia simplex Korth.

in Temm., Verh. Nat. Gesch. (1844) Bot. 253, Bakh. f.,1.c. (1943, 1943—45)317,

syn. nov. Pternandra simplex (Korth.) Baill., Hist. Pl. 7 (1880) 24, pro comb. non

fig. 37 et 38. Pternandra borneaensis Nayar, |.c. 52, nom. superf. (nom. nov.

for K. simplex Korth.). Kibessia simplex Korth. var. oblonga BI., |.c. (1849) 9,

syn. nov. Kibessia acuminatissima Merr., Univ. Calif. Publ. Bot. 15 (1929) 228,

syn. nov. Pternandra acuminatissima (Merr.) Nayar, |.c. 53.

var. aZurea.

Tree up to 30 m tall with a diameter up to 75 cm, often shrubby. Bark finely fissured,

soft, c. 0.5 mm thick, flaking and peeling off in strips 1—3 cm wide, brown; inner bark

brown to light yellowish, 2 mm thick; wood yellowish. Branchlets 4-angled, minutely

puberulous, glabrescent, brown-blackish when dry, peeling and cracking with a sinuate-

alate appearance, rapidly falling off leaving a cylindric, smooth branch drying grey-tan,

1—2 mm thick, nodes flattened. Blades chartaceous to subcoriaceous; lanceolate, elliptic,

or ovate; acuminate at the tip (acumen up to 1.5 cm long), narrowed and slightly de-

current at the base; immature blades minutely puberulous, especially below, becoming

glabrous, glabrous above; prominently 3-nerved from the base, nerves sunken above,

prominently raised and tapering below, lateral nerves slightly thinner, minutely puberu-

lous near the base below, glabrescent; intramarginal nerve faint, c. 1 mm from the margin;

secondary venation pinnate from each of the 3 main nerves, c. 20—30 pairs from each,

slightly raised below; 6.5—15 cm long, 3—6 cm wide, drying dark brown or olive-brown

above, lighter brown below. Petioles flattened, 2—4 mm long, 1.5—2 mm wide, at first

minutely puberulous, later glabrescent. Inflorescences from leafy nodes, frequently

terminal; flowers solitary or in 3-flowered cymes; axes 4-angled, minutely puberulous,

glabrescent, 10—30 mm long. Bracts lanceolate, acute, 1—1.5 mm long, minutely puberu-

lous. Primary axes 5—20 mm long with 1 or 2 nodes, c. 1 mm thick; secondary axes not

developed (solitary flowers) or up to 8 mm long; pedicels 3—6 mm long, slightly longer

and thicker in fruit. Calyx tube campanulate, densely covered with minutely puberulous

bristles, which are often curved or hooked, simple or branched, or with flattened, triangu-

lar, mostly barbed scales which are shorter near the pedicel and longer and usually more

barbed near the margin; bristles and scales often mixed together, longest ones 4 mm long

and 1 mm wide at the base; scales and bristles not or obscurely barbed in bud; margin

truncate, thickened, at maturity 8—10 mm long, 8 mm wide, minutely puberulous

throughout, glabrous with 16 internal lines; calyx cap conical, acute, c. 5S mm long,

distinctly tuberculed or with slender, frequently curved, spines near the middle, minutely

puberulous throughout, falling off as a unit, pale green. Petals slightly thickened, broadly

Ovate to suborbicular, obtuse to acute at the tip, broadly narrowed and truncate at the

base, margins thinner, c. 5 mm long, 5 mm wide, glabrous, pale pink, blueish-pink, or

dark blueish. Filaments flattened, c. 5 mm long; anthers c. 2 mm long. Stigma cylindric,

at first 4-grooved, later 4-lobed, 2.5—3 mm long. Style slender, 6—8 mm long, glabrous.

16 Gard. Bull. Sing. 34:1 (1981)

Fruit campanulate, truncate, bristles and/or scales frequently slightly more developed,

internal lines distinct, style often persisting; 8-10 mm long, 9-12 mm wide, minutely

puberulous externally; areolus glabrous; exocarp green then pale yellow, turning red, and

finally purplish when ripe; pericarp c. 1 mm thick, gritty. Seeds numerous, flattened,

rhombic-rectangular, glossy tan-khaki, c. 0.75 mm long.

Vernacular: siri siri (Tawau District, Sabah), siren siren (Tambunan District, Sabah),

polok (Iban); Java (Soend.): ipis koelit, ki boeroej, ki djambe, ki djeboeg.

Pternandra azurea (Bl.) Burk. var. azurea is easily distinguished from all other taxa

of Pternandra by the usually barbed calyx scales and acute blade bases. P. azurea var.

cordata (Korth.) Maxw. has more persistent, sinuate-alate wings on the branches and

cordate blades;and P. echinata Jack has thinner, pilose blades and branches, entire calyx

scales, and lacks tubercles on the calyx cap.

The two type specimens of K. simplex (Korthals s.n., from Borneo) at Leiden are,

unfortunately, in poor condition with only one small bud remaining on one sheet, while

the other collection is vegetative. Miquel, in his original description of this species, merely

included a few words about the leaves and that the calyx has simple setae with a sub-nude

cap. The bud specimen of the type collection is very immature and has simple bristles.

The branches, blades, and inflorescences of this collection of P. simplex match those of

the type specimens of P. azurea. In fact, I have not been able to distinguish the two

species vegetatively. P. azurea, according to Blume, has branched bristles on the calyx

and it is fortunate that the types of this species at Leiden are well preserved, since I have

been able to get a good idea about these two species. Blume notes for K. simplex that the

bristles are rarely branched. In addition to the numerous collections examined by Bak-

huizen f. at Leiden plus the many other newer collections of this species at Leiden, I

have been able to see that these two species are the same.

Included among the specimens of this species that I have studied, the bristles on

calyx range from almost all being entire, entire below and branched near the margin,

and nearly all branched. The appendages range from setose to linear and in a few

collections the scales are flattened and triangular. There is also individual variation in

the degree of branching and in many specimens this appears to be a factor related to the

maturity of the flowers. Examples of some collections with almost entirely simple, setose

appendages include: Nahar 12700 and Luang 22383 (both from Sarawak) and Cockburn

& Saikeh 70033 (from Sabah). Some collections with branched bristles near the margin

are: Kostermans 7039, and Nooteboom & Chai (from Borneo); the latter collection also

has some calyx specimens that are all simple, thus there is variation in the nature of the

bristles even on the same plant. Finally, some collections with flattened, branched scales

include: Winkel 1415 (from Java), deVries 107 (from Sumatra); with Clemens 21565,

and Luang 21874 (both from Sarawak).

Transitional situations are common where there are simple and branched scales,

therefore there is nothing substantial that can be used to distinguish the two species on

all herbarium sheets. The petals and anthers from the few collections with mature flowers

that I have seen are the same, regardless of the nature of the calyx appendages. More

evidence lies in the development of the calyx bud which always appears to have simple

bristles when young. Maturity of the flowers also involves the development of the bristles

or scales, with the fruit having the most distinct appendages. Two sheets, Clemens 29367

(Sabah), and Paie 28437 (Sarawak) have buds identical to the one on the type specimen

of K. simplex and also include more mature buds with immature and mature fruit that

Maxwell: Prernandra (Melastomataceae) 17

have distinctly branched, and a few flattened, appendages. These more mature flowers

and fruits are identical to those on the type collections of P. azurea.

There is good reason to consider K. simplex as a synonym of P. azurea since not only

is the former species poorly definied, but it is extremely difficult to distinguish from

the latter. With the numerous variant intermediates, including Elmer 21310 (Sabah)

— type of K. acuminatissima Merr. — which has fruits covered with simple bristles except

near the margin where they are more flattened and branched; it is clear that P. azurea

is variable in the calyx ornamentation and that this factor cannot be determined unless

mature flowers and fruits are available.

King considered Kibessia simplex Korth. as a synonym of P. echinata Jack, however

the two taxa are easily distinguished by the calyx appendages and the absence of spines

on the cap of P. echinata.

Kibessia azurea (Bl.) DC. var subalata (Bl.) Mig. is clearly the same as P. azurea

and does not merit distinction as a separate taxon. The holotype of this variety (Korthals

s.n., from Sumatra) does indeed have winged branches which appear sinuate-alate due to

the peeling off of the epidermis. Beccari 688 (from Sumatra) shows all stages of peeling

from a smooth, cracked, sinuate-alate, to completely lost epidermis. The branches with

the epidermis lost are cylindric and are identical to those branches on the type collections

of K. simplex and P. azurea, plus most of the other collections of these two species at

Singapore and Leiden which have already lost their epidermis.

The calyx bristles in these specimens with an intact or peeling epidermis are almost

all entire (e.g. Lampangi 24078, Gibot 31294, and Brand 24547 (all from Borneo);

to flattened and branched (e.g. the holotype of the variety), Beccari 688, and Paie 28331

(from Sarawak). This not only shows that var. subalata is based on an inconsistent charac-

teristic, but also that these bristles are found on specimens that could be referred to both

K. simplex and K. azurea.

Waitz s.n., from Gunong Rantau, Kalimantan, the holotype of Kibessia simplex

Korth. var. oblonga Bl., has an abnormal inflorescence without flowers or fruits. From

the branches and leaves I think that the specimen is more like P. azurea rather than

P. coerulescens Jack as Bakhuizen f. indicates.

Nayar (1978), in an entirely cursory attempt to change as many taxa of Kibessia to

Pternandra as possible, renamed Kibessia simplex Korth. as Pternandra borneaensis

Nayar. Nayar obviously never examined any authentic material of K. simplex Korth.

otherwise he might not have renamed this taxon since it is the same as P. azurea var.

azurea. Pternandra simplex (Korth.) Baill., which Nayar assumed was a distinct taxon,

is not based on Kibessia simplex Korth. Nayar of course, did not comment on P. simplex

(Korth.) Baill. since this would have required more than a superficial attempt at taxon-

omic research which he apparently prefers to conscientiously avoid.

Baillon’s (1880) illustrations of Pternandra simplex (Korth.) Baill. are certainly not

based on Kibessia simplex Korth. since the diagrams show the calyx with a tessellate

and smooth pattern, not bristly as in P. azurea. The figures (37 and 38) are referrable to

P. rostrata (Cogn.) Nayar, the most likely choice, or P. tessellata (Stapf) Nayar. The

combination Pternandra simplex (Korth.) Baill. is a valid recombination, thus Nayar’s

Pternandra borneaensis Nayar is a superfulous name.

Fig. 3. A,B,C: calyx; D: mature petal; ), F: stamens; G: stigma; H: fruit; I: fruit scales; J: seeds.

18 Gard. Bull. Sing. 34:1 (1981)

Distribution:

Lingga Archipelago

Pulau Lingga — Pasir Pandjang: Bunnemeyer 6900.

Sumatra

Asahan — Loemban Ria: Boeea 7985.

Benkoelen — de Voogd 1143; Mt. Singlang: Schiffner 2309.

Borneo. Sabah — Anap, Bukit Mersing: Sibat 21874, 22383; Bombari: Gibot 18595;

Dallas: Clemens 29367, 30350; Merotai Besar: Gibot 31294, 31310; Mt. Kinabalu: Chew,

Corner, Stainton 7; Sipitang, Ulu Moyah: Wood 16663; Tawao: Elmer 21310 (Kibessia

acuminatissima Merr., isotype L); Ulu Sg. Kinabatangan: Gibot 32947; Ulu Sungei

Serudong: Aban 31163; Tenom-Kamabong-Katabu Track: Cockburn & Saikeh 70033;

Tambunan, Trusmadi For. Res.: Mikil 31450; Ulu Sungei Danum: Cockburn 85032.

Sarawak — Balleh,N.Sebatong, Ulu Mujong: Ashton 19923; Gat, upper Rejang River:

Clemens 21565; Kakus, Ulu Mayeng: Sibat 21736; Kalabit Highlands: Nooteboom

& Chai 02085; Kapit District, Bukit Raya: Soepadmo & Chai 28176; Sungei Membalua

For. Res.: Price 257; Miri, Bakam Road: Au & Chai 24125; Mostyn: Sinanggul 57191;

Mt. Murud: Nooteboom & Chai 01831; Semengoh For. Res.: Purseglove & Shah 4429

(galled); Serian, Sabal For. Res.: Nahar 12700; Sungei Balleh: Anderson & Paie 28331;

Ulu Balleh: Paie 28437.

Kalimantan — Amai Ambil: Hallier 3113; Balikapapan (Gunong Beratus): Koster-

mans 7507; Gunong Rantau: Waitz sn (Kibessia simplex Korth. var. oblonga Bl., holotype

L); Gunong Sahasi, Belajan River: Forman 428; Gunong Sakoembang: Korthals sn

(Kibessia simplex Korth., holotype L); Gunong Singgalang: Korthals sn (Kibessia azurea

Bl. var subalata (Bl.) Mig., holotype L); W. Koetai, Kombeng: Endert 5192; Kutei:

Kostermans 5535; near L. Petah: Endert 3195; near Mt. Kemul: Endert 3622; Mangapu,

Singkulirang: Walsh 715; Mt. Kenepai: Hallier 1763; Nunukan Island: Kostermans 9043,

Miejer 2252, Paymans 7; Tanjong Bangko: Kostermans 7039; exact location unknown:

Winkler 2321.

Java — Djakarta, Batavia-Bantam: van Steenis 11193; Batavia, Gunong Paniisan:

van Steenis 2328; Gunong Pangerango: Blume sn (Kibessia azurea Bl., holotype L);

Leugkong District: Kostermans 23820; Mt. Salak: Hochreutiner 761 (Kibessia azurea

Bl. var. typica Hochr., L); Palak: Danser 2481.

3. Pternandra azurea (Bl.) Burk. var. cordata (Korth.) Maxw., stat. nov. Pternandra

cordata (Korth.) Baill., Hist. Pl. 7 (1880) 23. Kibessia cordata Korth. in Temm.,

Verh. Nat. Gesch. (1844) Bot. 253 and tab. 66; Blume, Mus. Bot. Lugd.-Bat.

I: 1 (1849) 8; Naudin, Ann. Sci. Nat. 3: 18 (1852) 262; Miquel, Fl. Ned. Ind. I

(1855) 570; Triana, Trans. Linn. Soc. 28 (1871) 152; Cogniaux in Boerlage,

Handl. Fl. Ned. Ind. I (1890) 536; Cogniaux in DC., Monogr. Phan. 7 (1891)

1107; Krasser in Engler & Prantl, Die Nat. Pfl. Fam. 3:7 (1893) 196 and tab. 81,

D—G; van Steenis, Bull. Jard. Bot. Buitenz. 3: 13 (1) (1933) 47; Bakhuizen f.,

“Thesis” (1943) 316, Med. Mus. Bot. Utrecht 91 (1943) 316, Rec. Trav. Bot.

Neerl. 40 (1943—45) 316. Kibessia tetraptera Migq. |.c. 571; Triana, l.c. 152;

Cogniaux, l.c. (1890) 536 and l.c. (1891) 1107; Merrill, J. Str. Br. Roy. As.

Soc. 86 (1921) 451, Bakhuizen f., l.c. 316, pro syn.

Maxwell: Prernandra (Melastomataceae) 19

Tree 2—10 m tall, diameter up to 12 cm, or a shrub; outer bark smooth, grey-brown,

wood light yellow. Branchlets distinctly 4-angled and typically undulate winged, minutely

puberulous, 1.5—2 mm thick; epidermis brown, peeling and flaking off leaving irregular

and scattered remnants of the angles and wings; older branches without the epidermis,

smooth, cylindric, glabrous, drying greyish to khaki-tan. Blades chartaceous to sub-

coriaceous, lanceolate, elliptic, to ovate; acuminate at the tip (acumen 1—1.5 cm long),

broadly rounded and shallowly (1—2 mm) cordate at the base; 3-nerved from the base,

nerves sunken above, raised and tapering below; intramarginal nerves thin, c. 1 mm from

the margin;secondary venation pinnate from each of the 3 main nerves, c. 30—40 pairs,

slightly raised below; glabrous above, minutely puberulous on the nerves below; green

above, pale green below; drying olive-green to dark brown above, lighter brown below;

7—12 cm long, 3—6 cm wide. Petioles c. 2 mm long, flattened, blades appearing sessile,

minutely puberulous, c. 1.5 mm thick. Flowers solitary from leafy nodes or in a terminal

3-flowered umbel; axes 4-angled, minutely puberulous, c. 1 mm thick; bracts lanceolate,

acute at the tip, c. 1.5 mm long, minutely puberulous; primary axes 5—8 mm long with

usually 1 node, secondary axes not developed or up to 5 mm long, pedicels S—8 mm

long. Calyx tube campanulate, densely covered with setose to flattened bristles which are

shorter and simple or barbed in the lower half, longest (2—3 mm) and branched near the

margin, minutely puberulous; 6 mm long, 5 mm wide, green; calyx cap conical, acute,

with 1 or several whorls of simple, barbed, or branched appendages, minutely puberulous,

falling off in one piece, c. 5 mm long. Petals slightly thickened, broadly ovate to sub-

orbicular, acute and often mucronate at the tip, broadly narrowed at the base, c. 7-8 mm

long, 8—9 mm wide, purple or lilac. Filaments flattened, c. 2 mm long, yellow; anthers

with a thickened bulge on the connective near the filament; stigma cylindric, slightly

twisted to the right, 4-grooved and later 4-lobed, c. 2 mm long, yellow. Style slender,

c. 6 mm long. Fruits campanulate, truncate, scales persisting, c. 8 mm long, 10 mm wide;

areolus glabrous, internal lines distinct; pericarp c. 1 mm thick, gritty. Seeds numerous,

cuneate-quadrangular, flattened, c. 1 mm long, glossy khaki-tan. :

In general, the distinction between var. cordata and var. azurea is distinct viz. the

former has rounded to cordate blades, shorter petioles, and branchlets with more per-

sistent epidermal remnants in a sinuate-alate form. Ultimately, however, the epidermis

falls off leaving a smooth, greyish to khaki-tan surface common to both taxa.

As far as I can determine from the limited number of mature flowers available;

the petals, anthers, stigma, and fruits of var. cordata are the same as those of var. azurea.

There is less variation in the shape of the calyx appendages in var. cordata than in var.

azurea, and in young buds of several specimens of var. cordata (e.g. Boeea 9363, de Wilde

& de Wilde-Duyfjes 12829 (both from Sumatra), and Korthals s.n. (from Java)) are

indistinguishable from those on the type collections of K. simplex and P. azurea. The

branching of the scales occurs with the development of the flowers, and after flowering

and with mature fruit these appendages are readily observed.

There are some specimens, e.g. Reinwardt s.n., and Bunnemeyer 3443 (both from

Sumatra) that have some blades rounded or slightly cordate at the base and others that

are narrowed. Bakhuizen f. identified both of these collections as K. cordata, however a

distinction between the two taxa here is difficult to determine. The petioles in both

specimens are short, thus I tend to agree with Bakhuizen f. in his identifications.

Var. cordata is structurally very close to P. hirtella (Cogn.) Nayar which has acute

blade bases, cylindric to somewhat angled (not wingled) branches, and is covered witha

reddish-brown tomentum. Vegetatively var. cordata is very similar to P. tessellata (Stapf)

Gard. Bull. Sing. 34:1 (1981)

Fig. 3. Pternandra azurea (B1.) Burk. var. azurea

Mikil 31450; F, G: Luang 22383; H: Luan

- - A! Endert 3195; B: Luang 21736; C, D, E:

g 21874; I: Kostermans 9043; J: Ashton 19923.

;

Maxwell: Pternandra (Melastomataceae) 21

ie >

uar rae

Fig. 4. Pternandra azurea (B1.) Burk. var. cordata (Korth.) Maxw. - - A-D: de Wilde & de Wilde-

Duyfjes 12829; E, F: Batten-Pooll sn (Sobiga, Sumatra).

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Plate 1. Pternandra azurea (B1.) Burk. var. cordata (Korth.) Maxw. This is the holotype of Kibessia

tetraptera Miq. which Bakhuizen f. reduced to a synonym of Kibessia cordata Korth. in

1943. K. cordata was recombined by Baillon in 1880 as Pternandra cordata (Korth.) Baill.

and has been given varietal status in this revision. Photo: Rijksherbarium, Leiden.

Maxwell: Pternandra (Melastomataceae) 23

Nayar which has tessellate or tuberculate calyx scales. The winged branches and cordate

blades of both taxa are quite similar, however the intramarginal nerve in the latter species

is usually more prominent and often the venation appears to be 5-nerved. Furthermore,

the calyx cap in P. tessellata is smooth or has minute tubercles and the blades are usually

glabrous below.

Fig. 4. A, B: calyx; C: mature petal; D, E: stamens; F: stigma.

Plate 1. Holotype of Kibessia tetraptera Miq.

Distribution: (specimens examined in addition to those cited by Bakhuizen f., 1943

and 1943—45, p. 317)

Sumatra Aek Moente, Asahan: Boeea 9363; Gunung Leuser: de Wilde & de Wilde-

Duyfjes 12829; Gunong Malintang: Korthals sn (holotype U); Kapoeas: Teysmann 8655;

Loemban Ria, Asahan: Boeea 7604; Pajakumbua: Meijer & Vermeulen 5401; Palembang:

Dumas 1576; Pargam Biran, Asahan: Boeea 5739; Siberut Island: Boden-Kloss 14558;

near Sobaiga: Batten-Pooll sn; Taram: Meijer 6716, 6799; Tapanoeli: Opper Angkola:

Junghuhn sn (Kibessia tetraptera Miq., holotype U, isotype L); W. Indragiri: Meijer 4311.

Borneo Sarawak — Bergark: Haviland 107; Penkulu: Haviland 173.

Kalimantan — Gunong Kenepai: Hallier 1601.

4. Pternandra cogniauxii Nayar, Bull. Bot. Survey India 17 (1978) 52. Ewyckia cordata

Korth. in Temm., Verh. Nat. Gesch. (1844) Bot. 255; Walpers, Repert. 5

(1845—46) 724; Baillon, Hist. Pl. 7 (1880) 23; not Kibessia cordata Korth., |.c.

253 (= Pternandra azurea (Bl.) Burk. var. cordata (Korth.) Maxw.). Macroplacis

cordata (Korth.) Bl., Mus. Bot. Lugd.-Bat. I:1 (1849) 7 and fig. 3; Naudin, Ann.

Sci. Nat. 3:18 (1852) 260; Walp., Ann. 2 (1850-51) 612; Naudin, Ann. Sci.

Nat. 3:18 (1852) 260; Miquel, Fl. Ned. Ind. I (1855) 569. Rectomitra cordata

(Korth.) Triana, Trans. Linn. Soc 28:1 (1871) 153. Kibessia korthalsiana Cogn.

in Boerlage, Handl. Fl. Ned. I (1890) 537, nomen and in DC., Monogr. Phan. 7

(1891) 1111; Merrill, J. Str. Br. Roy. As. Soc. 86 (1921) 451; Schwartz, Mitt.

Inst. Bot. Hamburg 7:3 (1931) 257; Bakhuizen f., “Thesis” (1943) 319, Med.

Mus. Bot. Utrecht 91 (1943) 319. Rec. Trav. Bot. Neerl. 40 (1943-45) 319;

not Pternandra korthalsiana (Miq.) Triana, l.c. 153 (= Pternandra tuberculata

(Korth.) Nayar). Pternandra cardiophylla Ohwi, in scheda (L, BO, SING).

Shrub (epiphytic or hanging) up to 5 m tall, diameter up to 4.5 cm, or a tree up

to 5 m tall with a diameter of up to 7 cm. Bark c. 0.25 mm thick, grey to light brown;

inner bark c. 0.5 mm thick, white; branching nearly down to the base of the plant.

Branchlets somewhat flattened and grooved on two faces below the upper node, becom-

ing cylindric below, glabrous, c. 2—2.5 mm thick; epidermis brownish, flaking off and

leaving a smooth, greyish surface. Blades subcoriaceous, glabrous, narrowly to broadly

Ovate, acuminate at the tip (acumen 1—1.5 cm long), broadly rounded and usually

shallowly (2—5 mm) cordate at the base; distinctly 5-nerved from the base, nerves sunken

above, prominently raised and tapering below, lateral nerves less prominent than the mid-

vein, the two peripheral veins merging with margin in the upper 1/4 of the blade; intra-

marginal nerves thin, c. | mm from and merging with the margin in the lower 1/3 of the

blade; secondary venation pinnate from all 5 main veins, c. 20-30 pairs, distinct above,

slightly raised below, anastomosing and reticulate; drying brownish with an olive hue

above, brown below; 14—20 cm long, 5.5—10 (12) cm wide near the base. Petioles flat-

24 Gard. Bull. Sing. 34:1 (1981)

tened, glabrous, 1.5—3 mm long, 3—4 (6) mm wide. Inflorescences mostly from behind

the leaves, often on branches up to 1.5 cm thick, or in the axils of the lowest leaves,

glabrous, solitary or fascicled often on raised tubercles; axes cylindric, smooth, c. 0.75

mm thick. Bracts lanceolate to ovate, acute, c. 1—1.5 mm long. Primary axes 1—7, with

one or less commonly two nodes, 2—3 mm long, elongating 1 or 2 mm in fruit; pedicels

1—3 mm long, elongating to 5 mm in fruit. Calyx tube campanulate-cyathiform, glabrous,

c. 4.5 cm high, 3—4 mm wide, densely covered with tuberculate scales which are thicker

near the margin, red; glabrous inside with 16 fine vertical lines; cap dome-shaped, splitting

into 4 thin, broadly triangular, persistent lobes, each 5 mm long, 3.5 mm wide, covered

with thick tubercles externally, smooth inside, reflexed at maturity. Petals (ex Cogniaux

and Bakh. f.) obovate, obtuse, 3—3.5 mm long, 2.5 mm wide, pink, white, or whitish-

purple. Filaments flattened, 2.5—3 mm long, whitish; anthers hatchet-shaped, inappendi-

culate, 2.5 mm long, 1 mm wide, yellow. Stigma 4-lobed, c. 1 mm long, whitish-purple.

Style slender, 7—9 mm long, glabrous, often persisting in fruit, white. Fruits urceolate,

nearly smooth or sparsely tuberculate near the base, scales elongate, diamond-shaped,

not or slightly umbonate, more tessellate near the rim and neck, 7—8 mm long, 7-8 mm

wide, glabrous, pinkish-red, maroonish, and finally purplish when ripe; areolus 4—S mm

wide, smooth or with several faint lines, and glabrous inside; calyx lobes persisting; peri-

carp c. 0.5 mm thick. Seeds numerous, flattened, rhombic-cuneate, quadrangular, c. 0.75

mm long, glossy light brown.

Habitat: primary forests, secondary scrub.

Vernacular: puloh (Iban), timberas burung (Banjar-Malay) seri seri (Utan), pura saie

(Bidayuh).

Pternandra cogniauxii Nayar is easily distinguished from all other taxa of Pternandra

by its relatively large, S-nerved, cordate blades; solitary or fascicled flowers from behind

the leaves; and the urceolate fruit with a nearly smooth or slightly umbonate pattern and

persistent, tuberculate calyx lobes. The fruit resembles that of P. galeata (Korth.) Ridl.

in basic shape, however the scales in P. galeata are tesselate throughout, while in P.

cogniauxii the pattern is more raised; also the calyx lobes in P. galeata are smooth on

both sides.

Bakhuizen f. notes that Kibessia elmeri Merr. is similar to this species, however his

opinion seems to be based entirely on Merrill’s comment at the end of the original

description. This species, in my opinion, is P. galeata var. elmeri (Merr.) Maxw.

Due to confusion on the part of previous authors, it has been necessary to establish

a new name for this species. The original name, Ewyckia cordata Korth. (described from

material collected by Korthals, s.n., in Borneo, and maintained in the Lieden collection)

cannot be used because it is a later homonym of K. cordata Korth. (now P. azurea (B1.)

Burk. var. cordata (Korth.) Maxw.). Cogniaux, realizing this, changed the name to K.

korthalsiana Cogn. Unfortunately, P. korthalsiana (Miq.) Triana (based on Ewyckia

Korthalsiana Mig. (F1. Ind. Bat. Suppl. I (1860) 321) is not a proper name for this species

since, in my opinion, it is the same as P. tuberculata (Korth.) Nayar (specifically P.

griffithii King). Cogniaux and Bakhuizen f. both placed it under P. coerulescens Jack. In

any case, the epithet korthalsiana cannot be used for any other species of Pternandra

since P. korthalsiana was validly published by Triana, even though it is a completely

different species from K. korthalsiana Cogn. The epithet cogniauxii was selected in an

effort to clear the confusion and misunderstanding in this literature and various herbaria

concerning this species. P. cogniauxii Nayar, as far as | can determine, is endemic to

Borneo. This species has been named in honour of Celestin Alfred Cogniaux (1841

Maxwell: Pternandra (Melastomataceae)

Fig. 5. Pternandra cogniauxii Nayar - - A, D: Haviland (=174)B; B, C: Mamit 33396.

26 Gard. Bull. Sing. 34:1 (1981)

1916), a French botanist, who published a comprehensive monograph on the Melasto-

mataceae in 1891.

Fig. 5. A: calyx with calyptra; B: calyx with split calyptra; C: stamen; D: fruit.

Distribution:

Borneo. Sabah — Beaufort: Cuadra 1363; Mt. Dulit: Richards 2562; Mt. Kinabalu:

Clemens 26355, 26873.

Sarawak — Bau: Purseglove 4457; Bukit Temudok, Simanggang: Anderson T29;

Kampong Sadir: Mamit 33396; Kota For. Res.: Chai & Paie 31104; Penkalan Ampat:

Haviland sn in 1889; Rajang River, Kapit: Jacobs 5457; Stabut, Padawan: Mamit 29829;

Ulu Dapoi, Tinjar: Paie 23054; exact location unknown: Haviland (=174) B.

Kalimantan — Gunong Rantau: Korthals sn (Ewyckia cordata Korth., holotype L);

Ingaran: Nooteboom 1328; W. Koetai: Endert 2659, 2931, 4970; Mentawir, Balikpapan:

Sauvour K5, 133; Mt. Rikai: Hallier 1308; Mt. Trusmadi: Nooteboom 1474; Tarakan:

Meijer 2537; exact location unknown: Korthals sn (L).

5. Pternandra coerulescens Jack, Malay Misc. 2:7 (1822) 61; Hooker, Comp. Bot. Mag.

I (1835) 157, Calcutta J. Nat. Hist. 5:15 (1843) 309; Triana, Trans. Linn. Soc.

28 (1871) 153 (“caerulescens”); Cogniaux in Boerlage, Handl. Fl. Ned. Ind. I

(1890) 536; Cogniaux in DC., Monogr. Phan. 7 (1891) 1102; Krasser in Engler

& Prantl, Die Nat. Pfl. Fam. 3:7 (1893) 196; King, J. As. Soc. Bengal 69, II:1

(1900) 68 (Mat. Fl. Mal. Pen. 3, 476); Clarke in Schmidt, Fl. Koh Chang VI

(1902) 195; Gibbs, J. Linn. Soc. Bot. 42 (1914) 80; Merrill, J. Str. Br. Roy.

As. Soc. 86 (1921) 452 (“caerulescens”’); Ridley, Fl. Malay Pen. I (1922) 808;

Merrill, Ling. Sci. J. 13 (1934) 65 (“caerulescens”); Bakhuizen f., “Thesis”

(1943) 324, Med. Mus. Bot. Utrecht 91 (1943) 324, Rec. Trav. Bot. Neerl.

40 (1943—45) 324; Li, J. Arn. Arbor. 25 (1944) 40 (“caerulescens’’); Corner,

Way. Trees Mal. ed. 2 (1952) 452; Ho, Fl. Vietnam (1960) 337 (plate 121E,

“cerulescens’’), 342; Keng, Gard. Bull. Sing. 31 (1978) 110. Pternandra coerule-

scens Jack var. jackiana (Walp.) Cl. in Hk. f., Fl. Brit. India II (1879) 551; P.

jackiana (Walp.) Ridl., |.c. 808, Bakh. f., l.c. 325, pro syn.; Ewyckia jackiana

Walp., Repert. Bot. Syst. 5 (1846) 734. P. capitellata Jack, |.c. 60, P. coerule-

scens Jack var. capitellata (Jack) King, |.c. 69; Bakh. f., l.c. 326, syn. nov.;

Ewyckia capitellata (Jack) Walp., |.c. 724. P. paniculata (Miq.) Triana, |.c. 153.

P. coerulescens Jack var. paniculata (Miq.) King, l.c. 69. Ewyckia paniculata

Miq., Fl. Ned. Ind. Suppl. I, Sumatra (1860) 321; Bakh. f., 1.c. 325 pro syn.

P. paniculata Benth. ex Cl., l.c. 551; Cogniaux, l.c. (1891) 1104; King, l.c.

68, pro syn. and Bakh. f., l.c. 324, not sensu Ridley, |.c. 809 (= P. tuberculata

(Korth.) Nayar). P. coerulescens Jack var. cyanea (Bl.) Cogn., 1.c. (1891) 1104;

Bakh. f., l.c. 327. Ewyckia cyanea Bl., Flora 14 (1831) 525, syn. nov. Ewyckia

latifolia Bl., Mus. Bot. Lugd.-Bat. I (1849) 6. Pternandra latifolia (Bl.) Triana,

l.c. 153; Merrill, l.c. 452. Ewyckia latifolia Bl. var. membranacea Bl., |.c. (1849)

6, syn. of var. cyanea by Bakh. f., 1.c.327. P. coerulescens Jack var. mem-

branacea (B\.) Furtado in scheda Herb. Sing. Ewyckia medinilliformis Naud.,

Ann. Sci. Nat. 18 (1852) 261. Memecylon oligoneuron BI. var. maluense Mansf.

in Engler, Bot. Jahr. 60 (1925) 139, syn. nov.

Tree up to 20 m tall with a diameter up to 90 cm; less frequently a shrub up to 5 m

tall. Bark grey, greyish-red, brown, smooth, finely fissured, thin; slash inner bark fibrous,

Maxwell: Pternandra (Melastomataceae) 27

light brown, very thin; cambium white; sapwood cream to yellow, very hard. Branchlets

cylindric, smooth 1—2 mm thick, brown to blackish when dry; epidermis peeling off of

older branches, leaving a smooth brown surface. Blades extremely variable in texture,

size, and shape; chartaceous, subcoriaceous, to coriaceous; lanceolate and acute at both

ends; elliptic to ovate, acute or obtuse at the tip and narrowed or rounded at the base;

or very broadly ovate to suborbicular with acute tips and in most specimens of this sort

slightly narrowed to rounded at the base; 3-nerved from the base, nerves sunken above,

raised and tapering to the tip below; intramarginal nerves thinner, 1—6 mm from the

margin, arching to the tip; secondary venation pinnate from each of the 3 main nerves,

obscure to prominent, anastomosing and reticulate; smallest blades S—8 cm long, 2—3 cm

wide; largest ones 18—25 cm long, 11—15 cm wide; dark green above, green to yellow-

green below; drying dark brown to blackish or sometimes olive green above, brown to

khaki-green below; entirely glabrous. Petioles flattened, 5S—10 mm long, 1—3 mm wide,

glabrous. Inflorescences extremely variable in size and branching; glomerulate, cymose,

to paniculate; c. 7 mm to 7.5 cm long, glabrous, few to many-flowered; a’.es flattened

with a groove on each narrow side, green; primary axes 1—3, mostly fom leaf axils

or on swollen leafless nodes (common in specimens with large, coriaceous blades),

0.75—2.0 mm wide at the base, c. 1 mm — 6 cm long with 1—5 nodes, each primary, etc.

node subtended by a pair of ovate, acute, and often slightly keeled connate bracts,

0.5S—1 mm long, persistent; smooth or with a reticulate pattern of flattened or slightly

raised polygonal plates in a tessellate pattern; secondary axes not developed or up to

2.5 cm long, tertiary axes none or up to 7 mm long, 4th axes not developed or up to

4 mm long, pedicels 1—4 mm long, solitary on the tips of each ultimate ramification.

Calyx campanulate, tube with a tessellate, less frequently umbonate to somewhat tuber-

culate pattern; truncate or with 4 minute cusps on the margin; smooth internally and

without lines or ridges; 2.5—4 mm long, 3—3.5 mm wide; green, drying brown to black.

Petal buds conical, lobes imbricating to the right, c. 3 mm long; mature petals thick with

thinner margins, broadly ovate to suborbicular, acute to mucronate at the tip, rounded or

shortly clawed at the base, reflexed at maturity, 2-6 mm long, 2—4 mm wide; white,

lilac, or blue; margins white to purple. Filaments flattened, 2—4 mm long, often flexed

or curved at maturity, violet; anthers inflexed in bud, then elevating with their dorsal

surfaces at 90° to the filament, thus forming a flat topped ring about the style, finally

becoming erect (180° to the filament) with the locules facing the style, c. 2—3 mm long,

c. 1 mm wide, yellow or violet. Stigma clavate-cylindric, c. 1 mm long, yellow. Style

slender, straight, 4-7 mm long; fruiting axes often elongating a few mm and becoming

thicker. Fruits subglobose (slightly wider than long, and when immature appearing some-

what urceolate) or globose, 4-6 mm diameter, capped by the thinner margin of the

calyx, areolus 2—3 mm wide, style often persisting; exocarp in immature fruit tessellate,

becoming obscured or nearly smooth with maturity; green or yellow-green then purple

when ripe, drying black; pericarp c. 0.25 mm thick. Seeds numerous, flattened rectangular-

quadrate, c. 1 mm long, 0.5 mm wide, glossy tan.

Pternandra coerulescens Jack is easily distinguished from all other species of

Pternandra by the truncate calyx (no calyx cap), tessellate pattern on the calyx tube and

fruit, and subglobose to globose fruits. P. multiflora Cogn. has similar flora features, but

has a much larger inflorescences and yellow petals. P. coerulescens is often confused with

P. galeata (Korth.) Ridl., however the latter species has a calyx cap which splits into 4

persistent calyx lobes and urceolate fruit.

Vernacular: cursed shade, lidah katak, sial menahun, pako bonnot paya, sial menan,

nipis kulit, kayu kuku bami (Malacca), mempoyang (Selangor); sireh sireh (Borneo

Malay), puloh (Iban), kalombayng (Dusun), sari sari (Brunei), dulang dulang (Kedayan),

ubah meskalak (Sarawak); memeteng (Sumatra); New Guinea: mereiie (Arfak), merie

(Sidei).

28 Gard. Bull. Sing. 34:1 (1981)

Delimiting the range of variation of this species has never satisfied many botanists,

thus numerous species and varieties have been described as being distinct from Jack’s

original description. For the Malay Peninsula King includes four varieties (including

the typical variety), while Ridley considered all four of them as distinct species.

Bakhuizen f. lists all of these taxa under three varieties (including the typical variety) of

P. coerulescens.

King notes that the species is very variable and that some specimens are intermediate

between varieties. From the large collection of P. coerulescens in the Singapore and Leiden

herbaria it is quite obvious that the variation in the size of the leaves, length of the

inflorescences, and number of flowers confirms King’s observation. The variation is

so complex, and in many collections impossible to delemit, that I have decided to include

all related taxa as synonyms of P. coerulescens; thereby giving the species an extremely

wide range of variation in the leaves and inflorescence. This idea is completely justified by

the gradation of these variable characteristics in specimens from Thailand to New Guinea.

It is also necessary to note that none of these taxa are based on any structural features

and that the flowers and fruits of all the specimens examined are essentially the same.

Pternandra coerulescens Jack var. jackiana (Walp.) Cl. has short cymes (primary

axes 5-10 mm) with few flowers. Sessile or shortly peduncled (3 mm) inflorescences

with larger leaves (11—25 cm long, 9—15 cm wide) have been separated as var. capitellata

(Jack) King. Finally, specimens with a large, branching, many-flowered inflorescence

with many rudimentary leaves have been designated as var. paniculata (Miq.) King. This

variety is based on a specimen which has an abnormal (witches broom) inflorescence.

While there are many specimens in the Singapore and Leiden collections which can be

definitely placed in one of these varieties, there are many other collections which include

characteristics of two varieties, or are intermediate between them.

For the specimens with large leaves (aff. var. capitellata) there is Moysey & Kiah

33638 which has a subsessile inflorescence (primary axes 1—2 mm with 2—3 flowers

per axis); Sinclair 39635 has longer axes (S—10 mm) and more flowers (aff. var. jackiana);

and Ridley s.n. (Province Wellesley, June 1890) has a larger inflorescence which can

be considered as the typical variety (sensu Bakh. f.). From these three specimens it is

clear that the size of the leaves cannot be correlated with the length of the inflorescence,

vice versa and that var. capitellata includes characteristics shared by the original limits of

the other varieties.

Another series linking these varieties includes Suppiah 14820, which has the axes

ranging from 4—7 mm long; Murdoch 263, 8—20 mm, and T. & P. 76 (2676) with the

axes 6—8 mm long. The leaves of all three of these collections are small (7—9 cm long,

3—4 cm wide), except with T. & P. 76 (2676) which also has large var. capitellata-like

leaves.

Cantley 2948 and Ridley 2009b have the short var. jackiana kind of inflorescence,

while with Haniff 367 the axes (in fruit) are c. 2 cm long — intermediate between var.

jackiana and Ngadiman 34652 which could easily be included as the typical variety due

to the relatively large inflorescence. The leaves of all four of these sheets are the same

(6—9.5 cm long, 4—5 cm wide, and of similar texture). Finally, Kiah 37737 is few-

flowered with the primary axes c. 1 mm long which contrasts remarkably with Kiah

32429 which has the large, branching inflorescence of the typical variety. The branches

and leaves on these two sheets are identical.

Maxwell: Pternandra (Melastomataceae) 29

The type collection of Ewyckia cyanea Bl. (Moluccas) at Leiden has relatively thin

and narrow blades with short inflorescences. The type specimens of £. latifolia BI.

(Borneo) have slightly thicker and larger blades with identical inflorescences. It is quite

obvious that the two taxa are the same. From the limited number of collections available.

all the specimens of P. coerulesceris s.s. from the Moluccas and Celebes have relatively

thin, narrow E. cyanea-type blades. New Guinea has collections with both EF. cyanea

and £. latifolia-type leaves. This is also true in Borneo, Sumatra, the Malay Peninsula,

and Thailand where both are common. In many collections the thicker blades of E.

latifolia and short inflorescences are intermediate with those of what Bakhuizen f.

considers the typical variety of P. coerulescens, i.e. the inflorescences are larger. Haviland

(=2203)Y (Borneo), Singh 1079 (Johore, W. Malaysia), and Smitinand 5659 (SE. Thai-

land) are some examples which illustrate that var. cyanea (including F. Jatifolia) cannot

always be distinguished from typical P. coerulescens.

E. latifolia Bl. var. membranacea Bl. (Borneo) has larger and thinner blades, but

otherwise does not differ from F. latifolia. Vat. membranacea is matched by specimens

with both large and small blades, e.g. Amir 35638 (Sabah), which links it to FE. cyanea.

The blades and small inflorescences of E. latifolia merge with the typical variety

through Sanusi 5327 (Sarawak), and Mikil 37737 (Sabah) which are intermediate in the

size of the inflorescences. The blades are of both the thin and thicker types.

The collections from Sumatra also show that there is much variation of

P. coerulescens and that the numerous taxa that have been split from it can not always

be distinguished or even properly defined. Soepadmo 134 links EF. latifolia and E. cyanea

with Toroes 3835, which has a long inflorescence; and Alston 14331, which has a short

inflorescence. The variability of the leaves and inflorescences of the Sumatran material

is the same as that for Borneo; thus there is considerable mixing of traits in many

collections, while in others it is true that they match the various varieties that other

authors have described.

The very large blades of P. coerulescens from the Malay Peninsula, which usually

have a very short (var. capitellata) inflorescence, are not common in Borneo and I have

not found any in the collections from the Celebes and New Guinea. These larger blades,

as noted above, merge with smaller and thinner blades, which have small or large inflore-

scences and could be identified as var. cyanea or the typical variety.

Several collections, e.g. Hou 537 and 545, Paie 26926, and Brooke 10640 (all from

Sarawak) have small, thick blades with indistinct secondary venation; and short inflore-

scences (var. capitellata) appear to be distinct from all other variants of P. coerulescens.

These specimens can be linked to E. Jatifolia and the typical variety through other speci-

mens with various combinations of thinner blades (Bunnemeyer 7504, from the Lingga

Archipelago) or larger inflorescences (Soepadmo 134, Sumatra).

The gradation of the specimens from the Malay Peninsula and the Malay Archipelago

is quite striking, and from the examples noted above there is no doubt in my mind that

there is no solid basis for maintaining any of the taxa which have been split from P.

coerulescens. | have attempted to sort all the specimens according to leaf shape and size

and length of the inflorescence. As a result many groups include a bewildering assemblage

of contrasting characteristics, i.e. the variation of the leaves and inflorescence is so mixed

that it is virtually impossible to group many sheets beyond an individual basis.

30 Gard. Bull. Sing. 34:1 (1981)

As far as flowers are concerned there is some variation in the size and ornamentation

of the calyx and length of the filaments; and shape of the petals; but the shape of the

anthers and features of the stigma and style appear to be nearly the same. King notes that

3 or 4-anthers are imperfect, however in all of the flowers (including a liquid collection,

Maxwell 77—23, from Singapore) dissected all 8 of the stamens are of the same shape and

size.

The calyx in most specimens is ornamented with flat, smooth, tessellate plates. This

feature does not correlate in any way with leaf or inflorescence characteristics. In some

other specimens the ornamentation is of raised warts which appear umbonate or even

slightly tuberculate; and again, specimens in the Singapore and Leiden collections with

this type of pattern have variable leaf and inflorescence sizes. Symington & Kiah 28774

has a var. capitellata inflorescence and Maxwell 77—23 a larger one that could easily be

considered as the typical variety; the blades in the former are rather thin and narrow (aff.

E. cyanea), while in the latter they are coriaceous and considerably wider (typical variety).

The calyx is umbonate-tuberculate in both specimens.

A smooth, tessellate pattern is found in Ridley s.n. (Province Wellesley, June 1890)

and the specimen could easily be considered as being the typical variety; and with Moysey

& Kiah 33638 (var. capitellata), both with large leaves, and in the small-leaved specimen

collected by Suppiah (14820) which has a var. jackiana style inflorescence. Thus, the

tessellate calyx pattern is common to specimens of all the varieties.

In general, however, many of the var. paniculata specimens have a warty or tubercu-

late pattern and these have been properly identified as P. tuberculata (Korth.) Nayar.

In other specimens with the abnormal inflorescence the calyx pattern is tessellate (P.

coerulescens). Sometimes the pattern is extremely variable and inconsistent, that is leaf,

inflorescence, and calyx characteristics do not form a clear series in which correlation

of these traits is possible. In at least one specimen, Henderson 10012, (inflorescence =

var. capitellata) both the smooth and raised patterns are found.

One exceptional specimen, Derry 1159, has a tuberculate calyx very much re-

sembling P. tuberculata, but the absence of a calyx cap and the anthers distinguish this

as P. coerulescens. The inflorescence is rather large and, as far as this trait is concerned,

is easily included as the typical variety. This particular specimen and several others

were incorrectly annotated by Furtado (Herb. Sing.) as K. tuberculata since the tuber-

culate pattern of many is similar to this species. Dissection of flowers to examine the

stamens and presence of a calyx cap is necessary in order to distinguish many of these

specimens.

I do not agree with Bakhuizen f. in reducing Kibessia simplex Korth. var. oblonga

Bl. to a synonym of P. coerulescens. The holotype of the variety (Teysmann s.n., from

Borneo) lacks buds, but the leaves suggest close affinities with P. azurea (B1.) Burk. and

thin-bladed specimens of P. tuberculata.

Bakhuizen f. also reduced Kibessia angustifolia Bl. to a synonym of P. coerulescens.

The holotype (Waitz s.n., from Borneo) is without flowers or fruits, but the angled

branches; and thin, narrow leaves strongly resemble those of P. echinata Jack. These

vegetative parts are, in contrast to the puberulous indumentum of P. echinata, entirely

glabrous. ;

The holotype of Ewyckia paniculata Mig. at Utrecht (Teysmann s.n., from Bangka

Island) has abnormal inflorescences in which the axes and bracts are larger than normal

Maxwell: Prernandra (Melastomataceae) 31

and frequently with deformed buds. The holotype has some normal buds and these are

identical to those of many typical specimens of P. coerulescens Jack. I am certain of this

since the deformed buds lack calyx lobes or a calyx cap. Wallich cat. 4080, which

Bentham named as P. paniculata Benth. ex Cl. also has abnormal inflorescences similar

to those of P. paniculata (Miq.) Triana — the latter being reduced by Cogniaux to a

synonym of the former, and both species to P. coerulescens Jack by Bakhuizen f. King

considered P. paniculata Benth. ex Cl. as an abnormal form of P. coerulescens Jack.

Ridley was of the opinion that P. paniculata Benth. ex Cl. is a form of P. griffithii King

with a large, abnormal inflorescence. While the branches and leaves of the two respective

type specimens are similar, the flowers do not match. The calyx tube of P. griffithii

King is tuberculate and there is a distinct cap which splits into 4 lobes. In P. paniculata

Benth. ex Cl. the calyx tube is tessellate and there is no cap. P. griffithii King is, there-

fore, not related to, while P. paniculata Benth. ex Cl. is identical to P. coerulescens

Jack. P. griffithii King is, in my opinion, the same as P. tuberculata (Korth.) Nayar.

Furtado annotated several specimens in the Singapore collection as Pternandra

coerulescens Jack var. membranacea (Bl.) Furt. with the added comment that the taxon

is the same as var. cyanea (Bl.) Cogn. These specimens have the short var. jackiana in-

florescence, however the blades range in texture from subcoriaceous (Ridley 12113)

to coriaceous (Ridley 6217, 2009b). The leaves are all of the smaller (c. 8—12 cm long,

4—8 cm wide) size. As with all the other varieties described for P. coerulescens, var.

membranacea (Bl.) Furt. is neither distinct nor constant and, therefore, should be com-

bined with all other varieties of this species into a single taxon.

Ewyckia cyanea Bl. predates FE. latifolia Bl. var. membranacea Bl. and along with the

fact that Furtado’s new combination was never published, and is therefore invalid, var.

membranacea (B\.) Furt. should be ignored.

Ewyckia korthalsiana Mig., which Cogniaux considered the same as P. capitellata

and a synonym of var. cyanea by Bakhuizen f., is more closely related to P. tuberculata

(Korth.) Nayar (specifically P. griffithii King). As the holotype of E. korthalsiana is

without flowers or fruits, it is difficult to determine its exact relationship with P. coeru-

lescens and P. tuberculata. Miquel’s original description, however, indicates that the

calyx is 4-dentate with an areolate-verruculose pattern. This is good evidence to support

my belief that it is actually a synonym of P. tuberculata.

A specimen collected by Ledermann (s.n., from New Guinea) labeled Memecylon

oligoneuron var. maluense Mansf. in the Singapore collection is definitely P. coerulescens

Jack with a short inflorescence. Mansfield’s short description of this variety fits the speci-

men quite well, thus I am assuming that this specimen represents the taxon even though

I have not seen the type specimen (Ledermann 7007).

Fig. 6 A: inflorescence; B: flower bud; C: mature petal; D: stamen.

Plate 2. Holotype of Ewyckia paniculata Miq.

Distribution:

Hainan — Hung Mo Shan: Tsang & Fung 18034 (not seen, q.v. Merrill (1934) and Li

(1944) ).

Thailand. Trat: Chang Island: Klong Haat Sy Dang — Maxwell 74425, Klong Nayom

— Kerr 6827, 6828A; Takum: Kerr 17869.

Chumpon — sine. coll. 46.

ged Gard. Bull. Sing. 34:1 (1981)

aam + MM

Locule

onnec five

Fig. 6. Pternandra coerulescens Jack - - A-D: Sinclair 39635.

Maxwell: Pternandra (Melastomataceae) 33

Surat Thani — Kaw Pa-Ngan: Put 758; Langsuan, Tako: Put 1696; Nawng Wai:

Kerr 12270; Tung Luang: Kerr 12510.

Puket Island — Haniff 367.

Nakorn Sithammarat — Ban Natawn: Kerr 15650; Ban Plien: Kerr 15654; Tung

Song, Ban Pa Prek: Rabil 192.

Satun — Lakshnakara 346; Teratao Island: Kerr 14166.

Songkla — Hat Yai: Kerr 13538, Pradit 232; Natawee: Rabil 71,91.

Narathiwat — Bacho: B. Sangkhachand 141; Kao Re Chaw, Toh Moh: Lakshnakara

744; Nikomwang: Prayad 431.

W. Malaysia Kedah — Bukit Enggau: Everett 13752; Bukit Perak: Everett

13695; Bukit Sama Ganjah: Baba 21483; Bukit Selambau: Meh 8989; Gunong Bongau:

Harun 17710; Gunong Jerai: Meh 9040, 10160; Inchong Estate: Spare 3806; Kedah

Peak: Ridley sn, June 1893; Kochummen 16327, Stone, Mahmud, Sharif 8539; Langkawi

Is.: Haniff 15527;Mukim Sik: Dolman 21512; Pantai Acheh: Symington 37382; Pantai

Chichak: Meh 10182, Sungei Batu Asah: Ridley 15527; Sungei Rohan: Arshad 18029,

Symington 20875; Ulu Muda For. Res.: Bray 11508; Ulu Tawar Res.: Meh 12569;

sine loc.: Meh 21891.

Wellesley — Kriaui: Ridley 2009b, sn in June 1890; Nibong Tebal: Curtis 3470.

Penang — van Balgooy 2412; 1. H. Burkill 382, 3303; Chelliah 98142; Corner sn at

Balek Palu; Curtis 67, 879, sn at Government Hill; Muka Head: B. & K. Bremer 1770;

Penang Hill: King sn on 22 Aug. 1879, Nur 1235, Patt sn, Ridley sn in Dec. 1895 and

Jan. 1921, Selvaraj 99671, Shukor 104, Sidek 232, Sinclair 39033, H. Singh & Samsuri

196, Stone 6352; sine loc.: Wallich 4077 (neotype K), 4079, 4080, ? Wallich 3143;

Shimizu et al. 12969, 13160; Flippance sn on 27 Oct. 1932.

Perak — Batu Hampar, Taiping: Shah & Sidek 1159; Batu Tajoh: Wray 2156; Bikum

Sougkai: Murdoch 366; Bintan Hijan For. Res.: Everett 14505; Dindings: Murdoch 263,

Ridley sn on 12 March 1896; Goping: King’s collector 822, 839; Grik: I. H. Burkill

12391, Corner sn, Ismail 95009; Gunong Bubu: Cockburn 11940; Ipoh: Curtis 3159;

Keledang: Ridley 9685; Kota: Wray 3254; Larut: Wray 3975; Lumut For. Res.: Whit-

more 0975; Pondok Tanjong: For. Dep’t Fed. Mal. States 1141, Sallih 9721; Pulau

Lanang: Seimund sn on 22 Nov. 1925; Pulau Rumbia: Lowry 479; Sungai Krian Estate:

Spare 34458, 34566; Sungei Larut: Wray 2872; Taiping: Everett 13586, For. Dep't.

Fed. Mal. States 1155, Henderson 10012, Ridley 14686; Wray 1952, 2390; Telok Auson:

Haniff 14305; Waterfall: Fox 28; sine loc.: King’s coll. 864, Scortechini 43, Wray 3607.

Kelantan — Bukit Baka, Machang: Shah & Shukor 3192; Chabang Tongkat: Suppiah

108872; Kemahang: Whitmore 8884; Sungei Mering: Ng 5483; Sungei Perias: Whitmore

4109.

Trengganu — Bauk For. Res.: Chan 16878, Whitmore 3942; Kemaman, Ulu Ben-

dong: Corner 30026; Kemaman: Corner sn on 22 June 1932; Sekayu: Loh 13466,

Shing 13536; Sungei Loh: Cockburn 10720, 10735; Sungei Pelong: Suppiah 14815,

14820; Ulu Brang: Moysey & Kiah 33638, 33834; Ulu Sungei Trengganu: Cockburn

8499, 10552; Kemaman, north of Sungei Ayam: Whitmore 20183.

34 Gard. Bull. Sing. 34:1 (1981)

Selangor — Bangi For. Res.: Jaamat 10925; Batu Tiga: Ridley 12113; Ginting

Simpah: Stone 9568; Gunong Simpah: Strugnell 11226; Kepong: Ng 100014; Klang For.

Res.: I. H. Burkill 961; Kuala Lumpur: Curtis 2337, Mat 7034, Hamid 4981; Kuala

Selangor: T. & P. 211 (2811); Puchong: T. & P. 76 (2676); Raweang: Ridley 7329;

Rice Res. Inst. Exp. Station: Kochummen 97760; Sungei Bulok: Sow & Tachon 16865;

Singh, Samsuri, Sidek 381, 408; Sungei Pelok, Serang: Denny 4, 42; Sungai Tinggi; Nur

34132; Tasek: Ridley sn; Ulu Gombak; Hume 9229; sine loc.: Denny sn on 27 Nov.

1941.

Pahang — Aur For. Res.: Whitmore 3675; Balok For. Res.: Yeop 0842: Bukit

Cheraga: Sohadi 14728, Whitmore 20021; Chini For. Res.: Bray 11627, Cockburn

11054, 11077, Shing 17288; Jenderok: Kadim & Mohmud 59; Kuantan: Symington

& Kiah 28774; Lepar For. Res.: Suppiah 108954, 108979; Pekan: Ridley 1164, Maamat

16533; Taman Negara: Everett 14404; Tasek Bera: Henderson 24062; Titi Bungor:

Henderson 10624, Whitmore 15366.

Negri Sembilan — Seuawang For. Res.: For. Dep’t. Fed. Mal. States 1968; Tampai:

Ridley sn.

Malacca — Batang: Derry 1159; Bukit Bruang: Derry 14; Chabau: Alvins 2249;

Jus: Derry 15; Pengkalan Balak: Derry 31; Pulau Jarak: Seimund 78, 1142; Selandar

For. Res.: I. H. Burkill 1356; Sungei Udang: Alvins 20, Holmberg 863; Tebong: I. H.

Burkill 1334; sine loc.: Alvins 74, 648, 1151, 1284; Cuming 2316; Griffith 2273, sn;

Maingay 801 (1212), 802 (1508).

Johore — Alor Bukit: H. Singh 561; Bukit Paloh Estate: Shah & Kadim 436; Bukit

Panjang: Everett 14112; Endau: Kadim & Noor 294, Singh & Samsuri 1031; Gunong

Belumut: Holttum 10770; Pahang: Singh & Samsuri 1079; Gunong Pulai: Maxwell

81—13; Kota Tinggi: Ridley sn in Dec. 1892, Teruya 409; Kuala Sedili: H. M. Burkill

1866; Kuala Teuba: Ridley 11080; Palai: Ridley 12167; Peugbury: Foxworthy 1176;

Sungei Kayu: Kiah 32119, 32429; Singei Sedili Ketchil: Corner sn in June 1934; Virgin

Jungle Reserve: Ahmad 359.

Singapore — Baker sn on 4 Nov. 1917; Cantley 2948, sn; Corner sn in July 1933;

Hullett 320; Jumali 4414; Kiah 37737; Maxwell #7—23, 77-138; Ngadiman 34652,

34683, 36487; Nur sn on 30 Sept. 1915; Ridley 279, 2004, 6217, sn on 10 Feb. 1898,

sn in May 1905, sn in 1911; Sinclair 39635; Keng & Jumali 4043, Ahmad 1366, 1464,

Hib 135.

Riouw Archipelago — Neth. Ind. For: Serv. 27671, 28613, 30001, 30029, 30089.

Lingga Archipelago — Bunnemeyer 7025.

Sumatra — Aek Moente, Asahan: Boeea 9314; Bangka Island: Kostermans & Anta

120, 381, 1001, 1344; Teysmann 3396 H. B. (Ewyckia paniculata Miq., holotype U,

isotype BO); Damoeli, Koealoe: Toroes 1509; Goenoeng Si Papan: Toroes 3835; Gunung

Leuser: de Wilde & de Wilde Duyfjes 13971; Gunong Sengalan: Matthew sn in 1913;

Koealoe: Bartlett 6869, 6913; Muara Sakai: Alston 14331; Penjengat: Neth. Ind. For.

Serv. 21525; Tantau Parapat: Toroes 2139; Tapanoeli, Sibolga: Neth. Ind. For. Serv.

31021; Upper Rhauw, Pakanbaru: Soepadmo 79, 134; W. Indragiri, Taluk: Meijer 4078;

sine loc.: Buwalda 6292. °

Maxwell: Pternandra (Melastomataceae) 35

Borneo Sabah — Bayong, Mempakul: Tandom 4117; Binoni, Papar: Mikil 37737;

Bukit Kuku, Sandakan: Meijer 20560; Bukit Padang, Jesselton: Meijer 19942, Tikau

33732; Gunong Dajak: Aet 200; Hindian For. Res., Beaufort: Singh 24384; Kibilo,

Sandakan: Valera 1870; Lumat, Beaufort: Cuadra 1336, Kadir 1690; Lupak, Beaufort:

Bunaar 25836, Kinted 15889; Meruba, Beaufort: Karim 80300; Sandakan: Amir 35638,

Ampuria 32628, Creagh sn, Elleh 35596, Elmer 20194, Kadir 939, 3569; Puasa 4183;

Sepilof For. Res.: Charington 20853, Patrick 42922; Subuga: Sam 34678; Sionggau:

Brain 1291; Sungei Kapur: Meijer 22904, Singh 22883; Tawao: Elmer 20839, 21622.

Brunei — Bangawan For. Res.: Rahim 444, Alston 115; Bukit Gaharu, Serian: Paie &

Ashton 16660; Kota Belud, Kappal: Gibbs 4342.

Sarawak — Bako National Park: Brunig 7662, Carrick & Enoch 34, Chai & Paie

17850, Hou 537 and 545, Purseglove 4895, 5006, 5536, 5543; Stone 6854, 13418;

Baram District, Senada 4104; Berakas For. Res.: Anderson 4873; Kalaka: Haviland sn

on 24 April 1893; Kerengga Padang: Carrick 1488; Kuching: Haviland 68, =177, 2203,

=2203, (=2203)Y, sn on 17 May 1893, sn on 23 May 1893; Lambir Hills For. Res.:

Dan 4375; Limbrang: Haviland 2202, 2291 (2202); Lokapas Kudat: Orolofo: 18251;

Merintaman For. Res., Sipitang: Lantoh 72326; Miri: Luang 24141. Luang & Paie 24737;

Naman For. Res., Sibu: Sanusi 5327; Rejang River: Haviland & Hose 3636; Sabal For.

Res.: Paie 17031; Samapdi For. Res.: Paie 26180, 26926; Sungei Pirian Lundu: Chai

18523; Tanjong Po: Brooke 10640; Triso, Simanggang: Anderson 12852; exact location

unknown: Beccari 700, 742, 1962, 3141, 3275, 3363; Lave sn; Native collector 97,

307, 550.

Kalimantan — Bengkoka: Balajadia 2841, Shea & Minjulu 76132; Doessoen Lands:

Korthals sn (Ewyckia latifolia Bl. var. membranacea Bl., holotype L); Gunong Pamatton:

Korthals sn (Ewyckia latifolia Bl., holotype L); E. Kutei: Kostermans 5429; W. Koetai:

Endert 1545; Loa Dajanan, Samarinda: Kostermans 6706; Lubak Duaya: Meijer 23636;

Maslhi For. Res.: Krukoff 4021; Mensalong: Melegrito 2506; Mt. Kenepai: Hallier 1630;

Nununkan Island: Kostermans 9021, 9128; Meijer 2366; Pembilangan: Amdjah 804;

Sam Pajau: Aet 661; Samarinda: Reksodihardjo 101;Sampit: Kostermans 4709; Soengai

Kenepai, Hallier 2159; Sungei Menubar: Kostermans 5083; Sungei Susuk: Kostermans

5637; Tapang Padai: Omar 68; Tarakan Island: Meijer 2469; exact location unknown:

Winkler 2249, 2408.

Celebes — Manado: Eyma 3399, 4020; Moena: Neth. Ind. For. Serv. 21607.

Moluccas — W. Ceram: Kuswata & Soepadmo 212; Soela Island: Neth. Ind. For. Serv.

28834, 29817; sine loc.: Zippel sn (Ewyckia cyanea B1., holotype L).

New Guinea — Babo: Aet 690; Beriat: Schram 6007; Versteegh 4900; Daswa: Brass

5959; Hollandia: Neth. Ind. For. Serv. 5715, 25715; Ingembit: Henty, Ridsdale, Galore

31810; Ridsdale, Henty, Galore 31929; Reksodihardjo 291 , 343; Job Is.: Schram 14915;

Laem Morobe: Hartley 11913; Oriomo River: McVeigh 8292; Sepik-Gebiet: Ledermann

6536; 7788; Sidai: Koster 6784; Schram 1743; Sorong: Pleyte 704; Wersar: Versteegh

4969; sine loc.: Ledermann sn (Memecylon oligoneuron Bl. var. maluense Mansf., isotype ?

SING) (Ledermann 7007, type, not seen).

Australia. Queensland — Bamaga, Torres Strait: Hyland 3990.

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PPE

Plate 2. Pternandra coerulescens Jack. Ewyckia paniculata Miq. is based on a specimen with an

abnormal (witches broom) inflorescence described by Miquel in 1860 which Triana renamed

as Pternandra paniculata (Miq.) Triana in 1871. Bentham provided the name Pternandra

paniculata Benth. in the Wallich Catalogue (no. 4080) in 1831 which was not validly published

until 1879 by Clarke as P. paniculata Benth. ex Cl. King reduced this species to a synonym

of P. coerulescens Jack in 1900, while Ridley considered P. paniculata a synonym of P.

griffithii King (=P. tuberculata (Korth.) Nayar). King’s opinion has been confirmed in this

revision. This is the holotype of Ewyckia paniculata which was collected by Teysmann on

Bangka Island in 1871. Photo: Rijksherbarium, Leiden.

Maxwell: Prernandra (Melastomataceae) 37

6. Pternandra coriacea (Cogn.) Nayar, Bull. Bot. Survey India 17 (1978) 53. Kibessia

coriacea Cogn. in DC., Monogr. Phan. 7 (1891) 1109; Merrill, J. Str. Br. Roy.

As. Soc. 86 (1921) 450.

Tree up to 10 m tall with a diameter up to 60 cm. Bark smooth, thin, dark brown;

inner bark yellowish; sapwood light brown; heartwood dark brown. Branchlets slightly

flattened, becoming cylindric, 1—1.5 mm thick, epidermis dark brown, flaking and

peeling off leaving a smooth, tan coloured surface. Blades subcoriaceous, glabrous, elliptic

to ovate, acuminate at the tip (acumen c. 5 mm long), acute to somewhat rounded and

decurrent (1—2 mm) at the base; 3-nerved from the base, nerves sunken above, promi-

nently raised and tapering below; intramarginal nerves frequently appearing as a second

set of basal nerves (thus 5-nerved), but thinner and parallel, 1—1.5 mm away from the

margin; secondary venation pinnate from the 3 main nerves, c. 20 pairs, anastomosing

and reticulate; 4.5—6.5 cm long, 2.5—4 cm wide; glaucous below when fresh; drying dark

brown to olive-green above, grey-whitish or grey-greenish to light brown below. Petioles

flattened, 2—S mm long, 1—1.5 mm wide, glabrous. Inflorescences from leaf axils, com-

posed or solitary flowers and or 3-flowered cymes, up to 1.5 cm long; axes obscurely

4-angled to cylindric, glabrous. Bracts and bracteoles lanceolate, acute, c. 1 mm long.

Primary axes solitary or clustered, 2—4 mm long, with 1 or 2 nodes; secondary axes not

developed in solitary flowers, in the cymes 3—4 mm long, pedicels 2—3 mm long in the

cymes, 5—8 mm long with solitary flowers, usually puberulous. Calyx tube campanulate,

4 mm long, 3 mm wide, covered with entire, glabrous projections which are umbonate to

tuberculate in the lower part, longer and more echinate near the margin; cap conical,

acuminate, smooth, glabrous, 4—5 mm long, 3—4 mm wide, falling off in one piece. Petals

broadly ovate, 7-8 mm long, 6—8 mm wide, thin, white to very pale mauve, reflexed

at maturity. Filaments flattened, 3—3.5 mm long; anthers usually with a small extension

of the connective near the filament, 2—2.5 mm long. Stigma subcapitate to cylindric,

grooved, c. 1 mm long. Style slender, 7—9 mm long. Fruits urceolate, with a tessellate to

triangular-echinate pattern, 7-8 mm long, 6—7 mm wide, green then turning purple

when ripe. Seeds numerous, flattened, quadrangular-cuneate, smooth, glossy tan coloured,

c. 1 mm long.

Habitat: lowland swamp forests, peat swamps

Vernacular: ubah pandang (Malay)

Pternandra coriacea (Cogn.) Nayar is easily distinguished from all other species by

its relatively small, glaucous blades; solitary and short inflorescence axes; and the um-

bonate to triangular calyx scales which are never barbed.

Pternandra teysmanniana (Cogn.) Nayar differs from P. coriacea in having thick

coriaceous, 5-nerved blades with an obtuse to rounded tip, and fascicled to subsessile

flowers. P. coriacea is in many respects similar to P. tuberculata (Korth.) Nayar in leaf

size and texture, inflorescence, calyx pattern, and fruits. P. tuberculata, however, has a

calyx cap which splits into 4 lobes and leaves which often become considerably larger

than any collections of P. coriacea that I have seen. P. rostrata (Cogn.) Nayar differs in

having thicker inflorescence axes, larger flowers with an acute to rostrate cap, and larger

leaves.

P. coriacea is only known from Borneo and is apparently a relatively rare species.

More collections are desired in order to understand this species more completely.

Fig. 7. A: calyx; B: mature petal; C: stamen.

Plate 3. Lectotype of Pternandra coriacea (Cogn.) Nayar.

38 Gard. Bull. Sing. 34:1 (1981)

Fig. 7. Pternandra coriacea (Cogn.) Nayar - - A, C: Beccari 3324 (lectotype); B: Haviland (=972)B.

;

aw | JUA*as

Pternandra coriacea (Cogn.) Nayar, lectotype at Florence. Cogniaux listed two syntypes for

this species, both of which were collected by Beccari in Sarawak. This collection was chosen

as the lectotype for this revision. Cogniaux queried the name “‘Kibessia glauca Triana”’ in

his note on this specimen which, as far as I can determine, is an in scheda name. Photo:

Rijksherbarium, Leiden.

“ied

ran

40 Gard. Bull. Sing. 34:1 (1981)

Distribution:

Borneo. Sarawak — Garai: Haviland sn (548) in 1891; Kuching: Haviland =972B,

1753; Lake Kwang: Forest Department Sarawak 2606; Marop: Beccari 3164 (syntype

FI), 3324 (lectotype FI, isolectotypes K, P); Naman For. Res., Sibu: Ahmad 14335,

Anderson 3361/3; Sungei Mas: Bujang 20878.

Brunei — Liang: van Niel 4621; Ulu Sigei: Brunig 1006.

7. Pternandra crassicalyx Maxw., sp. nov.

Ramuli cylindrici, furfuracei, epidermide in sicco brunnescenti, deglubenti. Folia

subcoriacea oblonga rarius elliptica 75-150 mm longa, 30—75 mm lata, acuminata,

basi sive acuta, secus que petiolum decurrente, sive rotunda, supra fuscescentia usque

nigrescentia, infra (sicco) brunnescentia, subtriplinervis, nervis furfuraceis, glabrescenti-

bus, intramarginalibus tenuioribus; petiolis 40-60 mm longis furfuraceis glabrescenti-

bus. Inflorescentia axillaris, terminalisve, 15—40 mm longa, rhachi floribus solitariis

vel 3-umbellatis, quadrangularibus vel cylindricis, furfuraceis. Bracteae lanceolatae,

I15—50 mm longae acutae. Pedicelli 2—3.5 mm longi. Calyx late campanulatus tessellatus

vel umbonatus 6 mm longus, 8 mm latus, 1.5 mm crassus, furfuraceus; calyptra tholiformis

glabra laevis 7 mm longa, 1.5 mm crassa, tota integre decidua. Petala in alabastro crassa,

late rhombica, 6 mm longa, 8 mm lata, acuta. Fructus campanulatus truncatus, pericarpio

1,.5—2 mm crasso.

Type: SARAWAK: Kapit, Bukit Raya, Au 23944 (L, holotype; isotypes: K, SING,

SAN, A, BO, KEP, MEL, MOSC, FHO, CGE, P, SAR). Fig. 8.

Tree 5—12 (23?) m tall, diameter up to 15 cm. Bark smooth, soft, light grey-green

or light to dark brown, often mottled; inner bark thin, red, cambium red, sapwood white

to yellow-pink. Branchlets slightly flattened below the upper node, cylindric below,

epidermis drying dark brown, nearly glabrous or minutely brown furfuraceous, usually

peeling off on the second internode leaving a smooth, glabrous, grey-khaki surface when

dry, 1.5—2 mm thick; nodes slightly swollen. Blades subcoriaceious, oblong, less fre-

quently elliptic, accuminate at the tip (acumen up to 1 cm long), somewhat rounded or

acute and decurrent at the base; 3-nerved from the base, nerves sunken above, raised

and tapering below, intramarginal nerves thin, c. 1 mm from the margin, disappearing in

the upper part of the blade; secondary venation pinnate from each main nerve, c. 15—20

pairs, anastomosing and reticulate; glabrous above, glabrous or red-brown furfuraceous

on the nerves below; 7.5—15 cm long, 3—7.5 cm wide, drying dark brown to blackish,

frequently with a greyish hue, above; brown below. Petioles 4—6 mm long, 3 mm wide,

glabrous or red-brown furfuraceous. Inflorescences from leaf axils or terminal, 1.5—4 cm

long; flowers solitary or in 3-flowered umbels; axes slightly 4-angled to cylindric, red-

brown furfuraceous. Bracts and bracteloes lanceolate, acute, 1.5—5 mm long, red-brown

furfuraceous. Primary axes solitary (3) 8—22 mm long with 1 or 2 nodes, 0.5—1.5 mm

thick at the base; secondary axes not developed in solitary flowers, or up to 5 mm long,

pedicels 2—3.5 mm long. Calyx campanulate, with a tessellate to umbonate pattern of

scales which are smaller near the pedicel and longest about the margin, minutely fur-

furaceous, mature buds 6 mm long, 8 mm wide, 1.5 mm thick, purple; cap dome-shaped,

smooth, glabrous, 7 mm long, 1.5 mm thick, falling off in one piece, pale pink. Bud

petals imbricating to the right, broadly rhombic, acute and often asymmetric at the tip,

truncate at the base, thick with thinner margins where the venation is visible, 6 mm long,

8 mm wide, light violet, light blue, to purple. Filaments in bud c. 2 mm long, flattened,

Maxwell: Pternandra (Melastomataceae) 4]

J man &

map OF

Fig. 8. Pternandra crassicalyx Maxw. - - A: Au 23944 (holotype); B, C, D: Wright & Othman 32303:

E: Ampuria 32707.

42 Gard. Bull. Sing. 34:1 (1981) .

white; anthers in bud c. 2 mm long, connective with a minute spur near the filament.

Stigma in bud cylindric, twisted to the right, 2.5 mm long, very light violet when mature;

style slender, 4 mm long is bud. Mature fruits campanulate,truncate, 9-10 mm long,

10—12 mm wide, with a tessellate to umbonate pattern, scales often widely spaced,

margin truncate, areolus c. 7 mm wide, internal lines distinct, stigma often persisting,

epidermis red or violet, drying dark brown; pericarp 1.5—2 mm thick, gritty. Seeds

numerous, flattened, quadrate-cuneate, c. 0.75 mm long, glossy tan coloured.

Vernacular: sireh-sireh, siri siri; puloh (Iban), empulis (Iban).

Pternandra crassicalyx Maxw. is readily distinguished from all other species of

Pternandra by its large, thick, tessellate to umbonate calyx tube; large, thick, and smooth

cap; and the thick pericarp. Pternandra tuberculata (Korth.) Nayar has generally smaller,

glabrous leaves and flowers, and a calyx cap which splits into 4 lobes. Pternandra rostrata

(Cogn.) Nayar has similarly shaped fruits, however the calyx and fruit patterns are um-

bonate to tuberculate; with a thinner, more acute cap. *

_ The red-brown indumentum on the branchlets, blades (only in some specimens),

and inflorescences recall that of P. hirtella (Cogn.) Nayar, however the latter differs

greatly in having setose, branched calyx appendages; a setose cap, and entirely different

fruits.

Fig: 8. A, calyx; B: mature petal; C: stamen; D: stigma; E: fruit.

Distribution:

Borneo. Sabah — Beaufort: Mikil 28089; Beaufort Hill: Madius 49299; Keningan

District, Crocker Range: Ag. Nordin 85971; Kinabakan District, Tangkulap Kechil:

Madani 33239; Mt. Kinabalu: Chew, Corner, Stainton 7; Papar: Lajangah 32197; Tawau

District, Kalabakan: Gibot 30582.

Sarawak — Baram District, Entyout River: Hose 396; Batanson: Ampuria 32707;

Bintulu, Ulu Segan: Wright 27967; Bukit Iju: Jugah 23701; Bukit Raya, Kapit: Au 23944

(holotype L, isotypes: K, SING, SAN, A, BO, KEP, MEL,.MOSC, SAR, FHO, CGE, P).

Chai 18919, Wright 24713; Kota For. Res.: Chai & Paie 27931; Limbang, Ulu Medamit:

Wright & Othman 32303; Long Kerangan: Tong 35020; Long Palu: Fuchs 21214; Milinau

Gorge: Chew 437; Mt. Tawai For. Res.: Dewol & Alxius 88399; Miri — Nyabau: Luang

24557; Pelagus Rapids For. Res.: Anderson & Hou 502; Temburong, Bangar: Ashton

492; Ulu Singei Salimpupon: Bakar 26859, Dan 26859.

8. Pternandra echinata Jack, Mal. Misc. 2:9 (1822) 3; King, J. As. Soc. Bengal 69, II:1

(1900) 69 (Mat. Fl. Mal. Pen. 3, 477); Ridley, Fl. Mal. Pen. I (1922) 809;

Corner, Way. Trees Mal., ed. 2 (1952) 452 and fig. 149; Keng, Gard. Bull. Sing.

31 (1978) 110. Ewyckia echinata (Jack) Walp., Repert. Bot. Syst. 5 (1846)

724. Pternandra echinata Jack var. pubescens (Decne.) King, |.c. 70 (l.c. 478);

Craib, Fl. Siam. Enum. 1:4 (1931) 702, syn. nov. Kibessia pubescens Decne.,

Ann. Sci. Nat. 3:5 (1846) 318; Triana, Trans. Linn. Soc. 28 (1871) 152; Clarke

in Hk. f., Fl. Brit. India II (1879) 552; Cogniaux in DC., Monogr. Phan. 7 (1891)

1108. Pternandra echinata Jack var. bracteata Ridl., l.c. 809, syn. nov. Kibessia

echinata (Jack) Cogn., l.c. 1108; Merrill, J. Str. Br. Roy. As. Soc. 86 (1921)

451. Kibessia acuminata Decne., l.c. 316; Triana, l.c. 153; Cogniaux, l.c. 1108;

get)

Maxwell: Pternandra (Melastomataceae) 43

King, |.c. 70 (1.c. 478) pro syn.; Bakh. f., “Thesis” (1943) 319, Med. Mus. Bot.

Utrecht 91 (1943) 319, Rec. Trav. Bot. Neerl. 40 (1943—45) 319. Pternandra

acuminata (Decne.) Nayar, Bull. Bot. Survey India 17 (1978) 52. Kibessia cupu-

laris Decne., Deles. Ic. Pl. V (1846) tab. 5 and l.c. 317. Kibessia simplex auct.

non Korth.: Blume, Mus. Bot. Lugd.-Bat. I:1 (1849) 9; Triana, I.c. 152; Clarke,

lc. 552; Cogniaux, l.c. 1108; King, l.c. 70 (l.c. 478). Kibessia angustifolia

Bl., l.c. (1849) 9; Walpers, Ann. 2 (1850-51) 611; Miquel, Fl. Ned. Ind. I

(1855) 571; Triana, l.c. 153; Cogniaux, l.c. 1109; Merrill, 1.c. 450 (aff.).

Tree up to 20 m tall with a diameter (dbh) up to 60 cm, bole straight, infrequently

a shrub or climber (probably incorrect). Bark smooth, finely fissured, thin; grey, yellow-

brown, brown-grey, brown-black; inner bark brown, red, orange; wood yellow, white.

Branchlets distinctly and often sharply 4-angled or slightly 4-winged, becoming cylindric

with age; nearly glabrous to densely, but minutely, pilose with red-brown, less commonly

light yellowish, hairs; becoming glabrous or not with age; 1.5—2.5 mm thick; nodes

slightly thicker, with a distinct interpetiolar, stipule-like ridge in an inverted “‘U”-shape

and uniting the angles on the internodes, becoming thickened with age. Blades thin,

lanceolate to elliptic, acuminate at the tip (acumen 1—1.5 cm long) narrowed and de-

current for a few mm or somewhat rounded at the base; 3-nerved from the base, nerves

sunken above, raised and tapering below; intramarginal nerves very faint, 0.5—1 mm from

the margin, looping; secondary venation pinnate from each main nerve, anastomosing

and reticulate; glabrous above, glabrous to densely, but minutely (especially on the 3

main nerves) pilose with red-brown, less frequently light yellowish, hairs; 7—12 cm long,

2.5—4.5 cm wide; dark green above, green to green-brown below; drying brownish to

dark olive-green above, lighter brown to light olive-green below. Petioles flattened and

grooved, 3—6 mm long, 1 mm thick; glabrous to densely, but minutely pilose as the

branchlets and blades. Inflorescences varying from solitary flowers to cymes with 3—9

flowers (i.e. umbel with 3 flowers or simple cyme with 5—9 flowers), (2)3—6 cm long,

from leaf axils and often terminal; axes quadrangular, sparsely to densely pilose as

described above with the branchlets, etc., 1—1.5 mm thick; green, drying brown or tan;

primary axis solitary, 1-2 cm long with 2—3 nodes; secondary axes not developed, thus

the flowers are solitary; or present as 3 axes, each with one flower, i.e. appearing as

an umbel, or cymose with the middle axis having tertiary axes, each with one flower

(inflorescence 5-flowered); 1—1.5 mm long; maximum development as a 9-flowered

inflorescence with each secondary axis having 3 tertiary axes, each terminated by one

flower; tertiary axes 7—14 mm long; pedicels 4—8 mm long. Bracts of the inflorescence

and individual ramifications extremely variable in size, ranging from linear-lanceolate-

ovate to elliptic-broadly ovate; generally acute to acuminate at the tip, narrowed at the

base, distinctly 3-nerved, thin, glabrous to pilose as the branchlets, leaves, etc.; 5-20 mm

long, 2—9 mm wide, green, drying dark brown or olive-greenish; persisting in fruit.

Calyx tube campanulate, acutely tuberculate near the pedicel, becoming echinate near

the margin with triangular, obtuse to acuminate appendages, 2—S mm long, frequently

twisting or curling; minutely papillose throughout; tube 6—7 mm long; 7—8 mm wide,

smooth internally with 16 slightly raised lines opposite and alternating with the filaments,

green; calyx cap acute to acuminate, 6—8 mm long, 6—8 mm wide at the base, usually

minutely papillose throughout, green; falling off as a unit with the expansion of the

petals. Petals thick, broadly ovate to oblong, truncate to broadly rounded with an acute

to acuminate cusp at the tip; broadly narrowed at the base; 5—13 mm long, 3—9 mm

wide, erect to spreading at maturity, blue, whitish-lilac, white. Filaments thick, flattened

with a slightly raised ridge on one side, c. 2 mm wide, 3—4 mm long, whitish-lilac; bud

anthers introrse, i.e. dorsal surface facing the style, later becoming horizontal, i.e. dorsal

surface facing the style, later becoming horizontal, i.e. dorsal surfaces in a flat plane 90°

44 Gard. Bull. Sing. 34:1 (1981)

to and encircling the style, later becoming erect, i.e. dorsal surfaces and locules parallel

to the style; subreniform, at first with the flap-like margins of the connective near the

filament folded down, later becoming erect forming a flat, ovate, obtuse crest, 1—1.5

mm long; without other projections; 3—3.5 mm long, 2—2.5 mm wide; connectives

yellow, locules reddish. Stigma cylindric, often slightly twisted to the left, 2.5—5 mm

long, c. 1 mm thick, persisting in fruit, yellowish. Style S—9 mm long, whitish. Fruits sub-

globose (slightly wider than high) or globose, 7-10 mm high, 8—15 mm wide; areolus

6—9 mm diameter, internal ridges obscure to distinct; appendages on the calyx tube,

especially near the margin, up to 9 mm long; epidermis green and turning pale yellow,

purplish when ripe, brown-tan when dry; pericarp 1—1.5 mm thick, gritty. Seeds

numerous, flattened, rhombic-quadrangular, c. 0.75 mm long, testa glossy tan.

Habitat: primary and secondary forests

Vernacular: kayu kaki kura (Temuan in Selangor), lemak ketam (Kedah), pako anun

and sial-mi-na-on (Malacca), mempoyan and serri menwan (Pahang), sial menaon and

sial menahun, cursed shade (Selangor and Pahang).

Pternandra echinata Jack is easily distinguished from P. tuberculata (Korth.) Nayar

by the former having larger flowers and fruits, minutely pilose branchlets and undersides

of the chartaceous blades, and the distinctly echinate appendages near the margin of the

calyx tube in both flower and fruit.

In addition to obvious differences in flower morphology, P. azurea (Bl.) Burk. has

similar branchlets which, however, tend to crack and peel at the angles leaving a sinuate-

alate apperance, and less pubescent baldes.

King notes that P. echinata has a truncate calyx with 4 narrow, acuminate teeth.

All the specimens with flowers of this species in the Singapore collection, including some

that he collected himself in Singapore, have a distinct calyx cap, or in those specimens

where the cap has fallen off and in fruiting specimens, the calyx tube is truncate and

without any teeth. It is possible that King mistook some of the external, echinate

appendages for calyx lobes.

Other authors have not agreed on the exact status of P. echinata Jack. The species

was considered a synonym of K. azurea (Bl.) DC. by Moritzi (in Zollinger’s Syst. Verz.

(1845—46) p. 11), and identical to K. simplex Korth. by Cogniaux (Boerlage, Handl.

Fl. Ned. Ind. I (1891) 1108). Burkill (Kew Bull. (1935) 319) considered the species as

being P. azurea (Bl.) Burk. Bakhuizen f. (1943, 1943—45) was of the opinion that P.

echinata Jack is allied to P. coerulescens Jack and that the former “is an up to now un-

known species of that genus.” Finally, Merrill (J. Arn. Arbor. 33 (1952) 241) suspected

that the species is K. azurea (Bl.) DC.

After having examined specimens of these species and having seen living material

of P. echinata it is quite apparent that the description of K. acuminata Decne. corres-

ponds with P. echinata. It is certain that K. acuminata is not the same as K. azurea (B1.)

DC., K. simplex Korth., or P. coerulescens Jack. Clarke (Fl. Brit. Ind. II (1879) 552)

and Cogniaux list specimens collected by Walker (304) and Maingay (804) (both from

Singapore) as representative of Kibessia acuminata Decne. Both specimens are un-

questionably P. echinata. King also notes that K. acuminata is not distinct and sub-

sequently reduced it to P. echinata. I fully agree with his treatment. Bakhuizen f.

considered K. acuminata Decne. as a distinct species and listed two collections after his

description viz. Bunnemeyer 6424, which is P. echinata, and Hallier 3113, which is

P. azurea (B1.) Burk. var. azurea.

Maxwell: Pternandra (Melastomataceae) 45

There is considerable justifiation, therefore, in assuming that Jack’s short descrip-

tion: “pedunculis axillaribus terminalibusque, calyxibus ovariisque echinatus” matches

the most salient characteristics — the echinata calyx tube — of the specimens that I

have identified as P. echinata Jack.

Kibessia cupularis Decne. in Delessert’s Icones is identical to P. echinata, thus con-

firming the opinions of Cogniaux and King on the synonomy of the former with the

latter species.

I have not hesitated to reduce P. echinata Jack var. pubescens (Decne.) King to a

synonym of P. echinata Jack since the pubescence of the branchlets, leaves, and axes

of the inflorescences vary considerably. No distinction can be made between those

specimens that are glabrous or slightly pilose and those which have “much minute rusty

pubescence”. King also had his doubts about the distinction of this variety and comments

that Wallich did not accept this variety and considered it as being “true” P. echinata

Jack.

Ridley designated P. echinata Jack var. bracteata Ridl. on the basis of “large, ovate-

lanceolate, persistent bracts”. Indeed, there are several specimens, in addition to the

types of the variety (Ridley, sn, from Bukit Panjang, Singapore), e.g. Spare 1045, King

& Jumali 6298, Maxwell 76—757; that have these kind of bracts. However, some of the

older collections including, King s.n. (Singapore, 7 September 1879), Curtis 270, Hullett

661, and Ridley 9189 have bracts similar to the isotype. In fact, Curtis 270 is annotated

“P. echinata var. pubescens King”. Considering the fact that there are specimens with

either intermediate sized bracts (Goodenough 2010 from Singapore, and Lambok 2723

from Rompin, Pahang) or have both small and large bracts on the same specimen (often

the same inflorescence) (C. F. Kedah 20788 from Gunong Jerai, Kedah; Santiago sn,

from Kepong; and I. H. Burkill 446, from Malacca) it is quite possible that King did

not consider the larger bracts as being unusual. His description merely indicates “...

flowers with one or more pairs of curved, linear-oblong bracteoles”. Both Ridley and

Bakhuizen f. did not include descriptions of the bracts of this species, thus I am

uncertain of the size limits for the typical variety.

I feel that there is no good reason to maintain var. bracteata Ridl. as distinct from

the typical variety of P. echinata Jack since there are some specimens that cannot be

definitely distinguished. As far as all other structural and vegetative features are concern-

ed, var. bracteata does not differ from those specimens with small bracts.

It is interesting to note that all specimens of P. echinata Jack from Singapore and

most of the collections from Johore have large bracts. Specimens from southern Thailand

to Johore mostly have smaller bracts. Craib does not record var. bracteata Ridl. for

Thailand. It is not surprising, therefore, that four of the specimens with intermediate or

mixed-sized bracts come from north of Johore and the other from Singapore since the

centers of distribution probably include the northern part of the Malay Peninsula for

small bracts and Singapore — Sumatra for large bracts. Ridly also notes that the distri-

bution of var. bracteata includes Singapore, Sumatra, and Java, however I have not seen

any specimens of this species from the latter locale. It is probable that P. echinata may be

found in Sumatra.

The holotype of Kibessia? angustifolia Bl. at Leiden (Waitz s.n., from Borneo)

merely consists of a short branch and a few leaves. These branches are 4-angled and

the blades are thin and narrow. These parts differ from P. echinata in being entirely

glabrous, otherwise they are indistinguishable. This species is not, as Bakhuizen f. claims,

the same as P. coerulescens Jack.

46 Gard. Bull. Sing. 34:1 (1981)

Fig. 1; Ovary, x-section.

Fig.9. A: inflorescence and leaf; B: flower; C: mature petal; D: mature stamen.

Plate 4. A and B: flowers; C: branchlet with mature fruits; D: mature trunk and bark.

Distribution:

Thailand. Nakorn Sithammarat, Thung Song: Geesink & Santisuk 5362; Trang,

Khao Chong: Phusomsaeng 58; Satun, Klawng Ton: Kerr 14575; Songkla, Klawng Yai:

Kerr 15884; Pattani, Betong: Kerr 7430; Narathiwat, Waeng: B. Sangkhachand 853;

Toh Moh, Kao Re Chaw: Lakshnakara 737.

W. Malaysia. Kedah — Bukit Perak For. Res.: Chan 13137; Gunong Bongsu For.

Res.: Spare 3775; Gunong Jerai: Meh 10159, Everett 13672; Kedah: Meh 21884;

Ulu Mada For. Res.: Chan 6732.

Penang — Curtis 270, = 270 in Sept. 1890 and March 1892; Ridley sn on 9 July

1890; Wallich Cat. 4078 (neotype K), 4078A (P. echinata Jack var. pubescens (Decne.)

King, lectotype K).

Perak — Gunong Bubu For. Res.: Everett 13962, Suppiah 11933; Larut: King’s

collector 3518, 3578; Chan 13195, Everett 13581 at Maxwell’s Hill; Taiping: Wray

2354; sine loc.: Kunstler 87; Scortechini 1643, 1894 (1824?).

Kelantan — Kampong Gobek: Shah & Kadim 565; Kuala Lebir: Ridley sn in Feb.

1917; Sungai Jenal: Cockburn 7446; Sungai Lebir Kechil: Cockburn 7138.

Trengganu — Bukit Langut For. Res.: Suppiah 11430; Bukit Rambai: Loh 13418.

Selangor — Batu Tiga: Mahamet sn in April 1890; Berembun near Bukit Tangga:

Ismail 104877; Bukit Kulu: Goodenough 10604; Bukit Lassong For. Res.: Kochummen

93466; Kepong: Ng 118166, Santiago sn; Klang Gates: Hume 7077; Petaling Jaya:

Woods 145; Rawang: Kloss sn; Semangkok: Ridley sn; Sungei Buloh: H. M. Burkill &

Shah 1033, 1095; Ulu Gombak: Carrick 1482 (2412), Hume 8873; Ulu Langat: Gadoh

Umbai 880, 1361, 1769; Weld’s Hill: Cubitt 811.

Pahang — Benka Road: Soh 15089; Chini For. Res.: Bray 11632; Everett 14559;

Gunong Benom, Ulu Krau: Ismail 97845; Whitmore 3153; Gunong Tapis: Cockburn

10907; Lepar For. Res.: Suppiah 108979; Lubok Paku: Ngadiman 16122; Raub: I. H.

Burkill & Haniff 16241, 16868; Sohadi 14715; Rompin: Lambok 2723; Taman Negara,

Latah Berkoh: Keng, Wee, & students 3; Sungei Tahan: Kiah 31912; Tahan Woods:

Whitmore 4788; Taman Negara: Everett 14456; Shah & Shukor 2672; Ulu Tembeling:

Henderson 22130.

Negri Sembilan — Bukit Taugga: Nur 11831; Gunong Angsi: Sohadi 14585; Gunong

Tampin: Ridley sn on 16 Jan. 1917; Jelebu For. Res.: Suppiah 11297; Johol: Ridley sn

on 18 Jan. 1917; Kuala Klawan: Franck 1129; Menyala For. Res.: Ng 1822; Hou 728;

Tampin For. Res.: Shing 17057.

Malacca — Batang: Derry 17; Bukit Bruang: Curtis =270 in May 1900, Goodenough

1346; Bukit Sedaman For: Res.: Holttum 9672; Kesang Batu: Hervey 2010b; Selundun;

Alvins 249; Sungei Baru For. Res.: I. H. Burkill sn on 13 Feb. 1914; Sungei Ujong:

Maxwell: Pternandra (Melastomataceae) 47

Connective -

Locule?

—

WY }

Fig. 9. Pternandra echinata Jack - - A: I. H. Burkill & Haniff 16808; B, C, D: Maxwell 76-757.

Vy,

Be Shy

Y My)

Le Wy

Vee

* Plate 4. Pternandra echinata Jack A: flower, front view; B: flower and bracts, side view; C. branchlet

with mature fruits; D: mature trunk with bark. Photo A taken at the Botanic Gardens “‘Jungle”’

on 11 January 1981; B and C from Maxwell 76-757 collected at the Bukit Timah Nature

Reserve, Singapore on 7 December 1976; and D at the Bukit Timah Nature Reserve on

14 September 1980. Photos: A, Hugh Tan; B and C, Douglas Teo; and D, Dr. Chang Kiaw

Lan. Scales in cm.

Maxwell: Pternandra (Melastomataceae) 49

Alvins 2148; sine loc.: Alvins 586, 727, 1246, sn; Cuming 2337; Griffith 2272, sn:

Maingay 803 (1213).

Johore — Bukit Pakat: Ridley sn in 1900; Bukit Tangga Tujoh: Shah, Ahmad, Noor,

2057; Castlewood: Ridley 9189; Endau: Singh (& Samsuri) 1028; Gunong Lambak:

Whitmore 15547; Gunong Panti: Bain 6015, Chan 17667; Gunong Pulai: Maxwell

(observed); Kota Tinggi: T. & P. 648 (3248); Kuala Palong For. Res.: Everett 14272;

Mt. Ophir: Shah & Ahmad 3658; Pulau Tinggi: I. H. Burkill sn in June 1915; Dungei

Tukong Estate: Spare 1045. ~

Singapore — Anderson 62; Cantley 184, 2900; Goodenough = (Ridley) 352, 2010;

Gilliland 5075; Hardial 624; Henderson 1372; Hullett 661; Kasim 577; Keng & Jumali

522, 3071 (K242), 6298; King sn on 7 Sept. 1879; Maingay 804 (3333); Maxwell

76—757; Panki 769; Ridley 352, sn at Bukit Panjang (P. echinata Jack var. bracteata

Ridl. (holotype K, isotype SING); Walker 304; Botanic Gardens: Noor sn on 26 April

1921, Nur sn on 24 April 1924 and 19 Nov. 1929, Furtado sn on 19 Nov. 1929, Desmakh

sn on 10 Sept. 1921; Hill 1103; Ahmad 1442; Ali sn at Bukit Timah on 18 April 1974.

Riouw Archipelago. Pulau Dompak: Bunnemeyer 6424.

Borneo. sine loc.: Waitz sn (aff.) (Kibessia angustifolia Bl., holotype L).

9. Pternandra galeata (Korth.) Ridl., Fl. Mal. Pen. I (1922) 808. Ewyckia galeata

Korth. in Temm., Verh. Nat. Gesch. (1844) Bot. 254 and tab. 67; Miquel, FI.

Ned. Ind. I (1855) 569. Rectomitra galeata (Korth.) Bl., Mus. Bot. Lugd.-Bat.

I:1 (1849) 6 and fig. 1; Naudin, Ann. Sci. Nat. 3:18 (1852) 261; Triana, Trans.

Linn. Soc. 28 (1871) 153. Kibessia galeata (Korth.) Cogn. in Boerlage, Hand. Fl.

Ned. Ind. I (1890) 537 (nomen) and in DC., Monogr. Phan. 7 (1891) 1110;

Merrill, J. Str. Br. Roy. As. Soc. 86 (1921) 451; Bakhuizen f. ““Thesis” (1943)

320, Med. Mus. Bot. Utrecht 91 (1943) 320, Rec. Trav. Bot. Neerl. 40

(194345) 320. Kibessia galeata (Korth.) Cogn. var. pluriflora Mansf. in Engler,

Bot. Jahr. 60 (1925) 138, syn. nov. Pternandra forbesii E. G. Baker, J. Bot. 42

(1924) suppl. no. 744, 41.

var. galeata

Tree 7—22 m tall with a diameter up to 30 cm. Bark smooth, pinkish to pale brown,

shallowly fissured and rugose, 0.5 mm thick; inner bark pale brown, 2 mm thick; wood

pale brown. Branchlets flattened at and 1—2 mm below the upper node, otherwise

cylindric, smooth, 1—2 mm thick, upper nodes 3—4 mm wide. Blades subcoriaceous to

coriaceous, elliptic to ovate, acuminate at the tip (acumen 0.5—1.5 cm long), narrowed

and decurrent for a few mm at the base; prominently 3-nerved from the base, veins

sunken above, raised and tapering below; intramarginal nerves very faint, 0.5—1.5 mm

from the margin, looping; secondary venation pinnate, distinct but faint from each of

the 3 main nerves, anastomosing and reticulate; 9-14 cm long, 3.5—8 cm wide, drying

brown on both surfaces in thinner blades, olive-tan in thicker ones. Petioles flattened,

3—5 mm long, 1.5—3 mm wide, shorter and broader in thicker blades. Inflorescences

glomerulate, often from thickened nodes behind the leaves, c. 1 cm long, several to many

-flowered; primary axes 1—5 mm long, secondary axes not developed or up to 1 mm long,

pedicels usually solitary on the tip of each ultimate axis, 1—2 mm long. Bracts connate

and sheathing, ovate, acute at the tip, c. 1 mm long. Calyx tube campanulate, ornamented

with flat or slightly raised (tessellate, umbonate, to slightly verruculose) polygonal plates

outside, smooth with 16 faint vertical lines internally, c. 4 mm long, 3—3.5 mm wide;

P 4 *

50 Gard. Bull. Sing. 34:1 (1981)

clayx cap dome-shaped, apiculate, splitting into 2 or 4 persistent lobes, each c. 2.5 mm

long, 3 mm wide, smooth and without scales or plates on both surfaces. Petals thick,

broadly ovate to suborbicular, broadly rounded and acute at the tip, tapering to a broad

base, 3 mm long, 4 mm wide, reflexed at maturity, blue to pinkish-white. Anthers sub-

reniform, c. 2 mm long, connective shortly spurred; filaments flattened in bud, flexed-

sinuate later becoming straight, 2—3 mm long. Stigma broadly conical, 4-lobed, c. 1 mm

long. Style 8—9 mm long, elongating up to 15 mm and persisting in fruit. Fruit urceolate,

5 mm long, 7—9 mm wide, narrowed in a 1—2 mm long neck above the globose body,

lobes often persisting, areolus 3—4 mm wide; exocarp roughened, obscurely to distinctly

tessellate, drying tan to black; pericarp c. 0.2 mm thick. Seeds numerous, linear, acute,

c. 1 mm long, khaki-tan with a black ridge.

Vernacular: lidah batak, lagis pukuan hutan, kemasulan, pasoeian.

Pternandra galeata (Korth.) Ridl. var. galeata is easily separated from specimens of

Pternandra coerulescens Jack with short inflorescences by the presence of a calyx cap

and the 1—2 mm long neck on the fruits. P. galeata differs from P. tuberculata (Korth.)

Nayar by having a shorter inflorescence and tessellate, never tubercled, calyx tube, and

urecolate fruits. P. cogniauxii Nayar has similar fruits, but has 5-nerved, cordate blades

and a tuberculate calyx tube and lobes.

According to the original description, Kibessia galeata (Korth.) Cogn. var. pluriflora

Mansf. merely differs from var. galeata in having many flowers which are mostly in groups

of 3, ovate buds, and conical calyx caps. From the specimens of P. galeata in the Singa-

pore and Leiden collections, this variety is not distinct and falls within the range of

variation for var. glaeata.

It should be noted that there are one or two primary axes in variety pluriflora

(Ledermann 8585, from New Guinea) from the leaf axils, each c. 1 mm long, with 2 or

3 secondary axes each c. 3 mm long, and pedicels c. 1.5 mm long. The ovate, acute bracts

at each node are about 1 mm long and persistent. The inflorescence in this specimen has

longer axes than specimens from the Malay Peninsula and Borneo. In respect to the

leaves, inflorescence, and calyx pattern var. pluriflora resembles P. coerulescens Jack,

however the presence of a distinct cap and lobes immediately distinguishes P. galeata

from the latter.

The two isotypes of P. forbesii E. G. Baker at Leiden (Forbes 3219, from Sumatra)

are unquestionably the same as P. galeata since the fruits are distinctly urceolate. This

collection was determined as K. galeata Cogn. by Hallier in 1916.

Fig. 10. A: inflorescence; B: flower; C: mature petal; D: stamen; E: mature infructescence.

Distribution:

W. Malaysia Pahang — Auwik River: Burn-Murdoch 196; Endau, Rompin: Mahamud

15506; Pianggu, Endau: Evans sn.

Johore — Sungei Sedili: Corner sn on 27 March 1932, Maxwell 78—260; Kwala

Sembrang: Lake & Kelsall 4073; Kwala Tehing Tinggi: Ridley sn.

Sumatra — Bangka Island: Kostermans & Anta 111, 201; Indragiri: Buwalda 6827;

Penassa Siak: Ridley 9021; Sungei Bigin Telok, Palembang: Forbes 3219 (P. forbesii

E. G. Baker (holotype K. isotypes L); Upper Riauw, Pakanbaru: Soepadmo 29.

Maxwell: Pternandra (Melastomataceae)

Fig. 10, Pternandra galeata (Korth.) Ridl. var. galeata - - A-D: Corner sn, Sungei Sedili, Johore on

27 March 1932; E: Maxwell 78-260.

52 Gard. Bull. Sing. 34:1 (1981)

Borneo. Sabah — Keningau: Cockburn 73016.

Brunei — Ulu Damit: Ashton 932.

Sarawak — Segan For. Res., Bintulu: Paie 15589; Sungei Arang, Baram: Yakup

11209; Ulu Sungei Karap, Baram: Anderson 30709.

Kalimantan — Balikpapan: Kostermans 4456, Neth. Ind. For. Serv. 24637 (aff.);

Bandjermasin: Korthals sn (holotype L, isotype L), Motley 200; Berau: Kostermans

21540; Danau Menjeban: Main 2162; loa Haur, Samarinda: Kostermans 6868, 9916;

W. Koetai: Neth. Ind. For. Serv. 29231; Kuala Kuajan, Sampit: Kostermans 4721, 8066;

Lampit: Buwalda 7935; Mentawir, Balikpapan District: Kostermans 9706; Pelawan

Besar, Sangkoelirang: Walsh 748.

New Guinea — Manikiong: Koster 1212 (aff.); Sepik-Gebiet: Ledermann 8585

(Kibessia galeata (Korth.) Ridl. var. pluriflora Mansf., isotype L); Vogelkop Peninsula:

van Royen 4006 (aff.); Warsamson Valley, Sorong: Schram 12887; Western District,

Kiunga: Katik 46796.

10. Pternandra galeata (Korth.) Ridl. var. elmeri (Merr.) Maxw., stat. nov. Pternandra

elmeri (Merr.) Nayar, Bull. Bot. Survey India 17 (1978) 53. Kibessia elmeri

Merr., Univ. Calif. Publ. Bot. 15 (1929) 228.

Tree up to 20 m tall with a diameter up to 45 cm. Bark smooth to scaly, brown,

peeling off in 1 mm long strips; inner bark pale white-brown, 2 mm thick, white near

the cambium; sapwood white. Branchlets cylindric, smooth, c. 3 mm thick, drying tan

to blackish. Blades thick coriaceous, glabrous, broadly ovate, acuminate at the tip

(acumen c. 1 cm long), broadly rounded and slightly decurrent at the base; 3-nerved from

the base, nerves sunken above, raised and tapering below; intramarginal nerves thin, 1—2

mm from the margin, disappearing in the upper part of the blade; secondary venation

distinct, pinnate from each main nerve, anastomosing and reticulate; 12—16 cm long,

8—10 cm wide; drying olive-greenish to brown above, greenish to light brown (black in

juvenile blades) below. Petioles 8-10 mm long, 3—4 mm thick, glabrous. Inflorescences

in fasciles from leafy or leafless nodes, of 3—S flowered cymes c. 1 cm long or of solitary

flowers; axes glabrous, slightly flattened. Bracts and bracteoles ovate, acute at the tip,

1—1.5 mm long, persistent. Primary axes up to 5 mm long with 1—2 nodes, secondary

axes usually not developed, pedicels 0.5—0.75 mm long. Calyx campanulate, glabrous,

with a tessellate or slightly umbonate pattern, scales smaller near the pedicel, largest

near the margin; 4—5 mm long, 4—5 mm wide; cap dome-shaped, smooth and glabrous,

splitting into 4 triangular lobes each c. 4 mm long, falling off as the petals mature. Bud

petals broadly ovate to suborbicular, obtuse to acute at the tip, narrowed and truncate

at the base, thickened, venation faint, white. Bud filaments flattened, c. 2 mm long; bud

anthers 2—2.5 mm long. Stigma c. 1 mm long. Style slender c. 5 mm long in bud, elonga-

ting to 8 mm in fruit. Fruits urceolate, 7-8 mm long, 7—9 mm wide, tessellate, areolus

3—4 mm wide; white, drying brown to blackish; pericarp c. 0.5 mm thick. Seeds numer-

ous, flattened, rhombic-quadrangular-cuneate, c. 0.75 mm long, glossy tan to light brown.

Kibessia elmeri Merr., which Merrill placed in section Macroplacis Cogn. where the

calyx cap opens into 4 lobes and the anthers lack a connective spur, is similar to P. galeata

var. galeata in the structure of the inflorescence, calyx pattern, petals, and fruits. In

several specimens of P. galeata, e.g. Kostermans 9916, from Kalimantan, the calyx splits

into 4 lobes and the anthers have a small spur near the filament. Elmer 21337, an isotype

of K. elmeri, has only a few loose buds and immature fruits, and the anthers are too

Maxwell: Pternandra (Melastomataceae)

o SMM

Fig. 11. Pternandra galeata (Korth.) Ridl. var. elmeri (Merr.) Maxw. -

- A-C: Elmer 21337 (isotype).

54 Gard. Bull. Sing. 34:1 (1981)

immature to determine whether or not such a spur is present. Singh 30042, from Sabah,

has leaves which match Elmer 21337, and also has one inflorescence with some loose

fruits which are urceolate and have a tessellate pattern. These fruits match those of

several specimens of P. galeata, e.g. Soepadmo 29 (Sumatra) and Paie 15589 (Sarawak),

however these two specimens have smaller and thinner blades which do not resemble

those of var. elmeri. I still have not seen mature flowers of var. elmeri, however from the

specimens available it is apparent that var. elmeri is very closely related to P. galeata

due to the tessellate calyx pattern, cap which splits into 4 lobes, and the urceolate fruits.

Important differences between the two taxa include the larger, thicker baldes, and the

larger and more branched inflorescences of var. elmeri.

The generally larger inflorescences of var. elmeri are identical in structure to some

specimens of P. rostrata (Cogn.) Nay. with large leaves, e.g. Hose 462 (Sarawak); and

Kadir 2726, Singh 22369 (both from Sabah); however the latter taxon differs in usually

having thinner blades, umbonate to tuberculate calyx tube pattern, calyx cap which

falls off in one piece, and campanulate-truncate fruits.

- Pternandra congniauxii Nayar, the type of section Macroplacis Cogn. (Kibessia korthal-

siana Cogn.), is similar in the splitting of the calyx cap and urceolate fruits, but differs

in having a tuberculate calyx pattern and 5-nerved, cordate blades.

Referable here is a vegetative specimen collected by the Neth. Ind. Forest Service

(16214, from Kalimantan) which has leaves similar to those of var. elmeri. This specimen

also resembles large-leaved collections of P. coerulescens Jack which often has nearly

sessile or shortly cymose inflorescences. P. coerulescens is easily distinguished by its lack

of a calyx cap and globose fruits.

Nayar’s recombination of Kibessia elmeri Merr. to Pternandra was done without

having seen any relevant material, thus P. elmeri (Merr.) Nayar is merely another indica-

tion of Nayar’s inability to do critical botanical research.

Fig. 11. A:calyx; B: stamens@sinnt.

Distribution:

Borneo. Sabah — Tawau: Elmer 21337 (isotypes L, SING), Baradaya For. Res.:

Singh 30042.

Kalimantan — central Kutei: Kostermans 10325; W. Koetai: Neth. Ind. For. Service

16214 (aff.).

11. Pternandra gracilis (Cogn.) Nayar, Bull. Bot. Survey India 17 (1978) 53. Kibessia

gracilis Cogn. in DC., Monogr. Phan. 7 (1891) 1110; Merrill, J. Str. Br. Roy. As.

Soc. 86 (1921) 451; Schwartz, Mitt. Inst. Bot. Hamburg 7:3 (1931) 256;

Bakhuizen f., “Thesis” (1943) 323 (name), Med. Mus. Bot. Utrecht 91 (1943)

323 (name), Rec. Trav. Bot. Neerl. 40 (1943—45) 323 (name). Pternandra

viridula Ohwi, in scheda (L, BO).

Shrub 2—5 m tall or a tree up to 10 m high, diameter up to 15 cm. Bark smooth,

pale grey-brown, outer and inner bark brownish, cambium pale whitish, sapwood brown-

ish. Brachlets 4- angled, frequently flattened and grooved on two faces below the upper

node, c. 1 mm thick, glabrous (rarely setose); epidermis light brown, peeling off leaving

Maxwell: Prernandra (Melastomataceae) 55

a smooth, cylindric, glabrous khaki-tan coloured surface when dry. Blades chartaceous,

glabrous, lanceolate to elliptic, acuminate at the tip (acumen c. 1 cm long), narrowed

and decurrent for 1—2 mm at the base; 3-nerved from the base, nerves sunken above,

prominently raised and tapering below; intramarginal nerves thin, c. 1 mm from the

margin; secondary venation pinnate from each of the 3 main nerves, c. 20—30 pairs; ana-

stomosing and reticulate; drying brownish, often with an olive hue, above; lighter brown

below; 5—15 cm long, 2.5—S (8) cm wide. Petioles flattened, glabrous, 3—4 mm long,

1—2 mm wide. Inflorescences of solitary flowers or in clusters, from leaf axils; primary

axes 1—3 mm long with one or two nodes, each with one pedicel, pecicels c. 1 mm long.

Bracts ovate, acute, up to 1 mm long; axes and bracts glabrous. Calyx tube campanulate,

glabrous, densely tuberculate in the lower part, appendages becoming longer (up to

1 mm), thicker, and often somewhat flattened (muricate to narrowly echinate) in the

upper half, especially near the margin; 3—S mm long, 4—5 mm wide, pink to red; cap

dome-shaped, rounded at the tip, glabrous, 4-grooved, c. 1.5 mm long, up tc the middle

provided with appendages similar to those about the margin, splitting into + persistent,

triangular, acute lobes, each c. 1.5 mm long. Petals thin, glabrous, broadly ovate to sub-

orbicular, broadly rounded at the tip, clawed at the base, c. 4 mm long, 3 mm wide;

filaments flattened, 2—4 mm long; anthers c. 2 mm long, 1 mm wide, pale pink. Stigma

oblong, cylindric, c. 1 mm long, 4-lobed, yellow. Style slender, S—8 mm long, elongating

to 10 mm after flowering, pale pink. Fruits globose, appendages similar to those in

flower, often less densely spaced or nearly smooth with little trace of the scales, 6—8 mm

wide, orange turning red; calyx lobes persisting, green or yellow; pericarp c. 0.2 mm

thick. Seeds numerous, rectangular-quadrate, often flattened, c. 1 mm long, glossy

khaki-tan coloured.

Habitat: primary, frequently dipterocarp, forests

Vernacular: puloh (Iban).

Pternandra gracilis (Cogn.) Nayar is easily distinguished from all other taxa of

Pternandra by its solitary flowers on each primary axis, axes which are never more than

5 mm long, tuberculate-echinate calyx tube, cap which also has appendages and splits

into 4 persistent lobes; and the chartaceous, glabrous blades which are narrowed at both

ends.

Pternandra tuberculata (Korth.) Nayar is sometimes confused with this species, but

the former has larger and longer inflorescneces, larger flowers with an umbonate to

tuberculate pattern, smooth cap, thicker blades, and larger fruits.

Clemens 21580, from Sarawak, differs from all other specimens of P. gracilis examin-

ed in having black, minutely setose branches. The leaves and fruits are identical to those

of other specimens of P. gracilis.

Fig. 12. A,B: calyx;C: Mature petal; D: stamen; E: immature fruit.

Plate 5. Holotype of Pternandra gracilis (Cogn.) Nayar.

Distribution:

Borneo.Sabah — Gunong Balapau: Richards 2414;Gunong Lumaku: Nooteboom 1176;

Lumat, Beaufort: Madani 35095; Mt. Kinabalu: Carr 26691; Chew, Corner, Stainton 97,

1201; Clemens 28071, 29592, 31285, 32134, sn in May 1933; Mt. Trusmadi: Nooteboom

1368; Sungei Tuban: Ashton 18371; Usun Arau: Asah ak Luang 22684.

Gard. Bull. Sing. 34:1 (1981)

Fig. 12. Pternandra gracilis (Cogn.) Nayar - - A, C, D: Beccari 3172 (holotype); B: Ashton 18371;

E: Ashton 720.

Aohtuns! hitte nichts aut

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PS7T9TT/OG

der bel der Siec

; 0, BECCARL PIANTE BORNENSI.

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i RAGIATO DI SARAWAK. MAX Laute ball...

Plate 5. Pternandra gracilis (Cogn.) Nayar, holotype at Florence, which was collected by Beccari

in Sarawak in March 1867. Photo: Rijksherbarium, Leiden.

58 Gard. Bull. Sing. 34:1 (1981)

Brunei — Andulau For. Res.: Ashton 385; Kuala Temorong, Machang: Ashton 720.

Sarawak — Bukit Iju: Luang 23248; Bukit Mentagai: Luang 23263; Bukit Salong,

Kapit: Paie 25858, 25862; Gat, upper Rejang River: Clemens 21580; Lanko Battu:

Beccari 3172 (holotype (4244) FI; isotypes (4244A) FI, BR).

Kalimantan — W. Koetai near Mt. Kemoel: Endert 3866.

12. Pternandra hirtella (Cogn.) Nayar, Bull. Bot. Survey India 17 (1978) 52 (“birtella”’).

Kibessia hirtella Cogn. in DC., Monogr. Phan. 7 (1891) 1107; Merrill, J. Str. Br.

Roy. As. Soc. 86 (1921) 451; Bakhuizen f., “Thesis” (1943) 313, Med. Mus.

Bot. Utrecht 91 (1943) 313, Rec. Trav. Bot. Neerl. 40 (1943—45) 313.

Tree up to 12 m tall, diameter up to 15 cm. Outer bark smooth, thin, very dark grey

to black, longitudinally cracked; inner bark dark red; wood moderately hard, sapwood

pale yellow, heartwood darker and staining reddish after exposure. Branchlets 4-angled,

sometimes narrowly undulate-winged, densely red-brown pilose, c. 2 mm thick, epidermis

flaking off leaving a smooth, cylindric, glabrous, tan-khaki coloured surface. Blades

subcoriaceous to coriaceous, elliptic to ovate, acuminate at the tip (acumen 0.5—1.5 cm

long); narrowed, less frequently rounded, and shortly (1—2 mm) decurrent at the base;

3-nerved from the base, nerves sunken above, prominently raised and tapering below;

intramarginal nerves thin, 1—1.5 mm from the margin, sometimes giving the blades a

5-nerved appearance at the base; secondary venation pinnate from each of the 3 main

nerves, c. 15—25 pairs, slightly raised below, anastomosing and reticulate; blades glabrous

above, sparsely to densely red-brown pilose, especially on the main nerves, below; 9.5—20

cm long, (3) 4.5—8.5 cm wide, dark green above, paler beneath, glossy on both sides,

drying olive-brownish to dark brown above, brown below. Petioles flattened, 4-8 mm

long, 2—3 mm wide, densely red-brown pilose. Inflorescences of solitary flowers from

leafy nodes or in terminal, 3-flowered umbels; axes 4-angled, densely red-brown pilose.

Bracts and bracteoles lanceolate, acute at the tip, 4-7 mm long, 1—2 mm wide, densely

red-brown pilose, persisting. Primary axes 0.5—1.5 cm long, 1.5—2 mm thick, solitary

flowers with 2—4 nodes, the umbels with one node; secondary axes not developed in

solitary flowers or up to 4 mm long, pedicels 2—4 mm long. Calyx tube campanulate,

6—8 mm long, 5—7 mm wide, densely covered with setaceous bristles which are shorter

and simple near the base and longest (2—3 mm) and barbed near the rim, frequently curved

or hooked, densely red-brown pilose; calyx cap conical, acuminate, c. 6—7 mm long,

5—7 mm wide, with a whorl of bristles near the middle; densely red-brown pilose through-

out; falling off as a single unit, often remaining attached to one side of the rim in fruit,

up to 1.5 cm long. Mature flowers not seen. Bud petals slightly thickened, broadly ovate,

acute at the tip, truncate at the base, 6—7 mm long, 5—6 mm wide, margin thinner,

white. Bud anthers inappendiculate, c. 2 mm long, 1 mm wide, pale mauve; filaments

flattened. Stigma cylindric, slightly twisted to the right, later splitting into 4 lobes,

2.5 mm long. Style slender, c. 6 mm long, glabrous, elongating to 12 mm in fruit. Fruits

subglobose, truncate, covered with pilose bristles, 8—10 mm long, 11—13 mm wide,

areolus 7—8 mm wide, internal lines distinct, 16, glabrescent; exocarp greenish, reddish,

then dark violet when ripe; pericarp c. 1.5 mm thick, gritty. Seeds numerous, flattened,

rectangular-cuneate, glossy khaki-tan.

Habitat: primary lowland forests

Vernacular: merkatak (Kuching, Sarawak); puloh, pulu (Iban)

Fig. 13. A: calyx; B: anther; C: stigma.

Maxwell: Pternandra (Melastomataceae)

O- S

? mm

“ie Les ae

Fig. 13. Pternandra hirtella (Cogn.) Nayar. A: Ridley 12281; B, C: Carrick & Enoch 349.

60 Gard. Bull. Sing. 34:1 (1981)

As far as the basic structure of the inflorescence and calyx bristles are concerned,

P. hirtella closely resembles P. azurea (Bl.) Burk. var. azurea and var. cordata (Korth.)

Maxw. There are several specimens at Singapore and Leiden of both varieties of P. azurea

which have setose bristles which are mostly unbranched (e.g. Endert 3195. Kostermans

9043, and Nahar 12700 — all from Borneo); however with other specimens of P. azurea

there is considerable variation in the density, branching, and thickness of the calyx

appendages. The bristles in P. hirtella are constant, i.e. the lower part of the calyx tube

consists mostly of curved and barbed appendages, while those near the margin are

branched. This species is quite constant in its characteristics and I have seen that it differs

little in pubescence, leaf texture and shape, and the flowers.

P. hirtella is easily distinguished from all other species of Pternandra by its dense,

red-brown pubescence which persists on the branches, blade undersurfaces, petioles,

inflorescence axes, flowers, and fruits. The fruits differs from that of P. azurea in being

subglobose and truncate, rather than campanulate and truncate. The blades in P. hirtella

are also larger and thicker than in P. azurea.

Cogniaux (1891) lists two syntypes for this species viz. de Vriese sn and Beccari

17 — both from Borneo. I have not seen de Vriese’s specimen which Cogniaux indicates

is at Leiden, however Beccari 17, at Florence, was examined on loan and has been de-

signated as the lectotype of Pternandra hirtella (Cogn.) Nayar.

Distribution:

Borneo Sarawak - Bako National Park: Ashton 17815; Carrick & Encoh 349;

Purseglove 5043; Batu Anam: Dauk & Tachun 35601, 35604; Bukit Empan Ra’a, Serian:

Paie 28069; Bukit Iju, Ulu Arip, Balingian: Luang 23610; Gunong Santubong, Kuching:

Bujang 13698; Kuching: Banyeng & Sibat 24913; Beccari 17 (lectotype, FI); Haviland sn

on 14 April 1893; Lambir Forest, Miri: Morshidi 24090; Penkulu Ampat: Haviland 175;

Sampadi For. Res.: Paie 27694; Santubong, Kuching: Ridley 12281; Santulang: Ridley

sn in Jan. 1918; Selang For. Res., Kuching: Paie 8452; Semengoh For. Res., Kuching:

Anderson 14643, 26854; Bojeng 14643; Brain 5368; Brown 2290; Galau 15703; Setapuk

For. Res., Kuching: Bojeng 9304; Sundu: Bishop (Hose) & Haviland 230 (inflor.

abnormal); Sungei Sabal, Tapang, Serian: Sinclair 10277; Ulu Kakus, Anap: Haron

29993; Ulu Kuyong, Anap: Chai 19317.

13. Pternandra multiflora Cogn. in DC., Monogr. Phan. 7 (1891) 1104; Merrill, J. Str. Br.

Roy. As. Soc. 86 (1921) 452.

Tree up to 15 m tall with a diameter up to 1 m. Bark smooth, grey or mottled with

brown; inner bark greyish to yellow, sapwood greyish-brown. Branchlets flattened and

deeply grooved on the flattened faces, often 4-angled, becoming cylindric, 1.5—2 mm

thick, epidermis brown to black, peeling off leaving a tan-khaki coloured surface; nodes

flattened and swollen. Blades subcoriaceous, glabrous, elliptic to ovate, acuminate at the

tip (acumen 1—1.5 cm long), narrowed and frequently shortly decurrent at the base;

prominently 3-nerved from the base, nerves sunken above, prominently raised and

tapering below; lateral veins less prominent than the midvein; intramarginal nerves thin,

1—2 mm from the margin; secondary venation pinnate from each of the 3 main nerves,

c. 15—25 pairs equaly prominent on both surfaces, slightly raised, anastomosing and

prominently reticulate; 12—22 cm long, 5.5--7.5 cm wide; drying olive-brown above,

brown below. Petioles flattened, 5-10 mm long, 2—3 mm wide. Inflorescences paniculate

from behind the leaves and on thicker branches, 15—30 cm long, very loose and open,

many-flowered; axes grooved near the nodes, otherwise 4-angled, smooth, glabrous,

Maxwell: Prernandra (Melastomataceae)

Connechve

Locule

Fig. 14. Pternandra multiflora Cogn. - - A-D: Purseglove 5149: E: Brain 5369.

62 Gard. Bull. Sing. 34:1 (1981)

drying dark brown to blackish; primary axes generally solitary at each node, less fre-

quently paired, 10—25 cm long with 3—5 nodes; secondary axes 2 or 4 from each node

of the primary axis, peripheral axes less developed, 3—8 cm long with 1—3 nodes; 3d

axes c. 1--2.5 cm long, 4th 5- 10 mm, Sth not developed or up to 4 mm long, pedicels

1—2 mm long. Bracts and bracteoles lanceolate, acute, 0.5—2 mm long. Calyx tube cam-

panulate-cyathiform, scales few, tessellate to umbonate, rather large, often indistinct

giving a rugose texture, 2.5--3 mm long, 2 mm wide, smooth and glabrous; internally

with faint lines, margin often appearing 4-lobed with the extension of 4 large scales

alternating with 4 smaller ones at the margin, pale yellow. Petals yellow, ovate to sub-

orbicular, broadly rounded at the tip, broadly rounded and truncate to broadly clawed at

the base, 1.5—2 mm long, 1.5—2 mm wide, thin with visible venation. Filaments flatten-

ed, c. 1.5 mm long, 0.5 mm wide; anthers c. 1 mm long, yellow, connective thick, often

papillose, ridge-like with a short spur extending beyond the filament. Stigma capitate,

4-srooved, c. 0.75 mm long. Style somewhat flattened, slender, 5—6 mm long. Fruits

(sub) globose, truncate with remnant scales at the tip, tessellate-rugose and minutely

lepidote externally, 5—8 mm diameter; areolus 4 mm wide, style often persisting, exocarp

ripening orange to light brown, drying khaki-tan; pericarp c. 0.2 mm thick. Seeds

numerous, rhombic-cuneate-quadrangular, c. 0.5 mm long, tan to light brown, glossy.

Pternandra multiflora Cogn. is easily distinguished from all other species of Pternan-

dra, especially P. coerulescens Jack, by its long, complex, many-flowered inflorescence;

tessellate to umbonate calyx pattern, and absence of a cap. The yellow petals and fruits

also appear to be distinct. The closest species is P. coerulescens which has smaller in-

florescences; thick white, blue, or purple petals; and fruits which dry black.

Cogniaux notes in his description that the fruits of P. multiflora are shallowly

4-grooved (leviter 4-sulcata), however I have not seen this character on any of the fruiting

specimens of this species in the Singapore or Leiden collections.

Fig. 14. A: inflorescence; B: flower; C: mature petal; D: stamen; E: fruit.

Distribution:

Bomeo Sabah — Beaufort District: Madani 35246; Beaufort Hill: Mikil 31979;

Sun Suran Trail, Penampang District: Mikil 37755.

Sarawak — Bukit Lambar, Miri: Luang & Nulong 24464; Gunong Matang, Kuching:

Chai 19758; Kuching: Bojeng 9428; Lambir Forest, Miri: Morshidi 22892; Lebang For.

Res., Bintulu: Paie 15836; Limbang, Kunaeang: Haviland 68 (544); Matang: Beccari 1551

(lectotype FI), 2981 (syntype FI); Haviland sn on 24 Aug. 1888; Miri: Dan 1907

(S1458), Hose 598; Miri River: Hose 71; Niah River, Miri District: Wright 29135; Pelagus

For. Res.: Chai 33177; Penkulu Ampat: Haviland 476; Selang For. Res.: Paie 8484;

Semengoh For. Res.: Brain 3569; Sungei Tau: Purseglove 5149; Tegora: Haviland 2019;

exact location unkown: native collector 1139 (distribution Manila).

14. Pternandra rostrata (Cogn.) Nayar, Bull. Bot. Survey India 17 (1978) 53. Kibessia

rostrata Cogn. in DC., Monogr. Phan. 7 (1891) 1110; Merrill, J. Str. Br. Roy.

As. Soc. 86 (1921) 451. Kibessia verrucosa Merr., J. Str. Br. Roy. As. Soc. 86

(1922) 341, syn. nov. Pternandra verrucosa (Merr.) Nayar, l.c. 53; Kibessia

verruculosa Merr., Univ. Calif. Publ. Bot. 15 (1929) 227 (sphalm). Kibessia

affinis Merr., 1.c. (1929) 227, syn. nov. Pternandra affinis (Merr.) Nayar, l.c. 53.

Maxwell: Pternandra (Melastomataceae) 63

Kibessia tuberculata (Korth.) Hk. f. var. grandifolia Bakh. f., ‘“Thesis’’ (1943)

322, Med. Mus. Bot. Utrecht 91 (1943) 322, Rec. Trav. Bot. Neel. 40 (1943—45)

322, syn. nov. Pternandra grandifolia (Bakh. f.) Nayar, 1.c. 53.

Tree 8—15 m tall, diameter up to 25 cm, or a shrub up to 3 m tall; buttressed in

older individuals. Bark smooth, grey-brown, thin, finely fissured; inner bark red-brown,

thin; sapwood pale yellow. Branchlets cylindric, often somewhat flattened or obscurely

4-angled below the upper node, 1.5—2.5 mm thick, smooth, drying grey to brownish.

Blades subcoriaceous to coriaceous, glabrous, elliptic to ovate (often broadly so),

acuminate at the tip (acumen 0.5—1 cm long), narrowed or broadly rounded and usually

decurrent at the base; prominently 3-nerved from the base, nerves sunken above, raised

and tapering below; intramarginal nerves faint, 1—2 mm from the margin, disappearing

in the upper half of the blade; secondary venation pinnate from each main nerve, con-

spicuous, anastomosing and reticulate; 6.5—21 cm long, 3—10 cm wide; dark, often

glossy, green above; green below; drying dark brown to olive-brown above, brown to

tan-brown or with a greenish hue below. Petioles flattened, 2—8 mm long, 1.5—4 mm

thick, glabrous. Inflorescences of axillary or terminal cymes of 3 or of solitary flowers,

1—2.5 cm long, axes somewhat flattened to 4-angled, glabrous; primary axes usually

several from leaf axils, with 1—3 nodes, solitary flowers generally having two nodes and

two pairs of bracts as the secondary axes have not developed, c. (1) 5—10 mm long;

secondary axes not developed or 3—5 mm long; pedicels (0.5) 1—3 mm long. Bracts and

bracteoles lanceolate to ovate, acute, 1—2 mm long, persistent, green. Calyx tube cam-

panulate, ornamented with raised tubercles (umbonate to tuberculate), glabrous, 4—6

mm long, 3—5 cm wide at maturity; cap conical, thick, acute to rostrate, falling off in

one piece, 3—8 mm long, 4—5 mm wide; green; tube smooth and obscurely 16-lined

internally, lines opposite and alternating with the filaments. Petals thick, broadly ovate

to suborbicular, often asymmetric, broadly rounded and apiculate at the tip, shortly

clawed at the base, 6—8 mm long, 8—10 mm wide, reflexed at maturity, lilac. Filaments

flattened, 3—4 mm long, lilac; anthers subreniform, connective with a dorsal extension

near the filament, c. 2 mm long, yellow. Stigma clavate-cylindric, 1—2 mm long, yellow.

Style 5—6 mm long, whitish, elongating to about 10 mm and persisting in fruit. Fruits

campanulate, truncate at the top, 7—9 mm long, 10—12 mm wide, areolus 7—9 mm wide,

internal ridges and lines distinct; exocarp green, umbonate to distinctly tuberculate;

pericarp c. 1.5 mm thick, gritty. Seeds numerous, rectangular-quadrate, often appearing

cuneate, c. 1.0 mm long, testa glossy tan.

Vernacular: nipis kulit, tahi kerlan, talinga basing (Brunei), sandi-sandi (Dusun-Kina-

batangan).

Pternandra rostrata is very variable in the size and texture of the leaves, pattern on

the calyx tube and fruits, and shape of the calyx cap. This species differs from P. tuber-

culata (Korth.) Nayar which has a more slender and more branched inflorescence, and a

short calyx cap which splits into 4 lobes. The leaves of the two species are usually in-

distinguishable.

The type collection of P. rostrata (Beccari 979, from Sarawak) has a rostrate calyx

cap which apparently falls off in one piece. Other specimens with a rostrate cap, e.g.

Haviland 972, Kadir 2726, and Gansau 46311; vary in the size and texture of the leaves,

and length of the inflorescences. The calyx pattern ranges from umbonate to tuberculate

and in many other collections several different patterns can be found on the same speci-

men, that is umbonate, verruculose, and tuberculate patterns. These patterns do not

correlate with the size or shape of the leaves.

64 Gard. Bull. Sing. 34:1 (1981)

Pternandra verrucosa Merr. has the same leaves as Beccari 979, and with the few

loose fruits on paratype specimens (Ramos 1191, from Sabah), Merrill’s description of

the inflorescence and comparison with many specimens of P. rostrata at Singapore and

Leiden, I cannot find any constant differences between the two taxa. Mature flowers

are sparse, and from the internal structures from buds I cannot make any distinctions

either.

The isotype of P. affinis Merr. (Elmer 21478, from Sabah) has leaves which could

easily be confused with several specimens of P. galeata (Korth.) Ridl., but the loose

fruits have a tuberculate pattern. Although the shape of these fruits is urceolate, as with

P. galeata, the specimen is actually P. rostrata. Confirming evidence lies in the fact that

many immature fruits of P. rostrata are urceolate with a verrucose to tuberculate pattern

(not tessellate as with P. galeata) and in some specimens, e.g. Gansau 46311, fruits

identical to Elmer 21478 plus buds with a rostrate cap are on the same specimen. Aside

from the fact that the pattern on the fruits differ, the presence of a cap which falls off

in one piece (in contrast to the caps which split into 4 lobes with P. galeata) is sufficient

proof for me to consider Kibessia affinis as a synonym of P. rostrata.

Several collections from Borneo (e.g. Meijer 37999, Gibot 60613, Mikil 38496,

Latangah 33144, etc.) collected at elevations above 1300 m. have much smaller leaves

and more obtuse calyx caps than most other specimens of P. rostrata that I have seen.

These specimens will require further study with more flowering and fruiting material in

order to determine their exact taxonomic status. I have included them in this section

since they do not resemble any other species of Pternandra that I have studied.

Kibessia affinis Merr., K. verrucosa Merr., and K. tuberculata var. grandifolia Bakh.

f. are, in my opinion, all synonyms of P. rostrata. Even with type material of all these

taxa, deciding on their true taxonomic status has been difficult since these specimens are,

in general, not only incomplete, but also in poor condition. Reconstructing various

aspects of each one of them has been by comparison with other specimens at Singapore

and Leiden which match each type, plus the original descriptions.

All of these taxa have shortly cymose (up to 2 cm) inflorescences; campanulate

calyx tubes with umbonate to verrucose scales; smooth and acute to rostrate caps; thick

and broadly ovate to suborbicular (often asymmetric) petals; and campanulate fruits

with a truncate apex and an umbonate to verruculose pattern. The different taxa have

leaves which vary in size and, combined with the various stages of flower or fruit develop-

ment for each specimen, has resulted in considerable taxonomic confusion.

The isotype of K. affinis (Elmer 21478, from Sabah) has several fruits in various

stages of development, most of which are slightly constricted below the margin giving a

somewhat urceolate appearance. The pattern on these fruits is identical to that on the

only mature fruit (which is crushed) included with Rutten 537 — the holotype of var.

grandifolia. The fruits of Ramos 1191, a paratype of K. verrucosa, differ slightly in being

more umbonate and less verrucose, and are not, at least in most fruits, urceolate, i.e.

there is little, if any, construction below the margin. There are, however, some fruits

with Ramos 1191 which match those of K. affinis.

Merrill notes that K. affinis is closely related to (Kibessia) Pternandra galeata

(Korth.) Ridl., however the latter species is easily distinguished by its tessellate to some-

what umbonate calyx pattern, and the distinctly urceolate fruits (neck 1—2 mm) with

a much smoother pattern, and often with persistent calyx lobes.

Maxwell: Pternandra (Melastomataceae) 65

Griffith 2272/1 has a few calyx specimens which have identical scales to those of

var. grandifolia. The leaves on the two sheets show a transition from small (P. griffithii)

to moderately large (var. grandifolia). As noted above, the mature fruits of var. grandi-

folia the same as those of K. affinis which also has individual variations which match

those of K. verrucosa. The shape and venation of the blades in all of these taxa are

essentially the same. Large blades with prominent venation of K. verrucosa are matched

by those of var. grandifolia, which are slightly smaller; then progressively smaller leaves

from K. affinis, P. griffithii, and P. tuberculata.

It is obvious from other collections in Singapore and Leiden that all of these taxa are

the same since many specimens with small to large leaves have buds, flowers, or fruits

which match those fragments from one or more of the type collections discussed above.

In other words, these taxa have been described from individual specimens which were

collected at a particular stage in the development of the individual flower or fruit. By

comparing specimens collected in other stages of maturity, it is obvious that leaf size is

not a constant factor since all specimens can be linked by the distinct calyx pattern and

by the truncate fruits.

Fig. 15. A—F: G: mature petal; H: stamen.

Distribution:

Sumatra Exact location unknown: Beccari 179.

Borneo Sabah — Balat Camp, Lamag District: Lassan & Fox 72828; Balung, Tawao:

Clemens 32474, Gibot 32474; Batu Lima, Sandakan: Agama 463 (Kibessia verrucosa

Merr., paratype L), Villamil 147 (Kibessia verrucosa Merr., paratype L), Ramos 1722

(Kibessia verrucosa Merr., isotypes L, SING); Beram Bangan, Kudit District: Brand

30862; Camp Sepilok For. Res., Sandakan: J. Singh 22369; Columbon River: Clemens

34102; Dulit Ridge: Richards 1665; Elopura, Sandakan: Cuadra 2190, 2224; Taloon

10767; Garson, Tawau District: Mujin 40583; Gaya Island: Mujin 33573, Creagh sn: in

May 1895, Aug. 1895, July 1896; Gunong Alab: Nooteboom 959; Kabili-Sepilok For.

Res.: Keith 7123, 7175; Puasa 7012; Kalumpang Tawao, Lahad Datu District: Chai

29822, 31692; Keratam Camp, Sandakan: Wing 33963; Kuala Daramakud, Kinabatangan

District: Kadir 16867; Kundasong, Ranau District: Singh 27494; Loro For. Res., Kudat:

Siranggol 51101; Mendulong: Nooteboom 1071; Mostyn, Lahad Datu District: Agam

& Aban 40855; Mt. Dulit: Richards 1108; Mt. Kinabalu: Clemens 30515, 30812, sn;

Sadau 42826; Nanga Pelagos: Daud & Tachun 35669; Pock For. Res., Sempora District:

Gansau 46311; Pulau Balem Bangan: Stone & Anderson 86751, 86804; Stone 13177;

Pulau Gaya: Ampuria 41397, Meijer 20488; Ranau District: Leopold, Gary, Dewol

73815, Baldek 32349; Sapa Tali, Lemag: Ampuria 36418; Sandakan: Elmer 20203,

Sam 21797, Tangulon 1565; Sepilok For. Res., Sandakan: Brand 30997, Charington

20870, For. Dep’t. Borneo 1148; Sekong Camp, Sandakan: WIng 34921; Sekong Kechil,

Sandakan: Wing 38987; Sepilok For. Res., Kennedy Bay: Siranggol 39909; Sipanggar

Island: Ampuria 41333; Siptiang: Madani 35127; Sosopodon For. Res.: Gibot 60613,

Lajangah 33144, Meijer 37999, Mikil 38496, Mujin 33777; Sungei Dusun, Sandakan:

Elleh 34360; Sunsuran, Penampang District: Binideh & Talip 65301; Supu For. Res.,

Kinabangan: Puasa 10071; Tawao: Elmer 21478 (Kibessia affinis Merr., isotypes L,

SING), Gansau 47803, Gibot 30431, 34125; Tandom 8802, Wood 16459 (aff.); Tidok,

Sandakan District: Hepburn 23722; Ulu Mujong, Balleh: Ashton 13975; Ulu Sungai

Kalumpang: Gibot 30467.

Brunei — Bukit Lambir: Ashton 37, Fuchs 21297.

66 Gard. Bull. Sing. 34:1 (1981)

Fig. 15. Pternandra rostrata (Cogn.) Nayar - - A, G: Haviland 972; B, H: Hose 462; C: Kadir 2726;

D: Jacobs 5302; E: Siranggol 51101; F: Brooke 10010.

Maxwell: Pternandra (Melastomataceae) 67

Sarawak — Bettotan: Boden-Kloss 19057, Kloss 19027; Bukit Mentagai: Luang

22821; Entoyut River, Baram District: Hose 462; Gat, upper Rejang River: Clemens

21564, 21575; Gunong Mulu, Baram: Paie 15091; Gunong Rara For. Res.: Chow 75670;

Gunong Sahari: Forman 498; Kuching: Haviland 972; Lawas: Brooke 10010; Lema’as

For. Res.: Chong 33666; Long Kerangan, Barma District: Tong 34998; Matang: Haviland

176; Miri River, Baram District: Hose 653; Mt. Matang: Clemens 20937; Mt. Singglio:

Haviland =176; Piningiao: Beccari 979 (holotype FI); Puman Besar: Geh & Samsuri 708;

Rejang River, Kapit District: Jacobs 5302; Semengoh For. Res., Kuching: Ghazalli &

Asah 13662; Sungei Ukong, Miri: Haron 21395; Ulu Mujong, Balleh: Ashton 13975;

Ulu Sungei Bukong: Benang 24406, Luang 24454; Ulu Sungei Mamut Bakong, Miri:

Luang 24377; Ulu Sungei Sekaloh, Miri: Luang 27855, Wright 29129; Ulu Tubau: Ashton

17717; exact location unknown: Becari 301.

Kalimantan — Amai Ambit: Hallier 3208; Batu Pondong, Sangkoelirang District:

Kostermans 13664; Berau: Kostermans 21573; Boeloengan Beraoe: Rutten 537 (Kibessia

tuberculata (Korth.) Hk. f. ex Cl. var. grandifolia Bakh. f., (holotype U); Gunong Ke-

tapong, Sangkoelirang: Walsh 443; Hunukan: Meijer 2148; Indonesia Islands: Kostermans

21115; W. Koetai: Endert 2596, 4980; Liang Gagang: Hallier 2736; Loa Djanan, Sama-

rinda: Kostermans 6440; Muara Muntai Kutei: Meijer 496; Soengai Sambas: Hallier 1082:

Sungei Wain, Balikapan: Kostermans 4154, 4238.

New Guinea. Sepikgebiet: Ledermann 8717a (aff.) (Kibessia ledermannii Mansf.,

isotype SING).

15. Pternandra tessellata (Stapf) Nayar, Bull. Bot. Survey India 17 (1978) 53. Kibessia

tessellata Stapf, Trans. Linn. Soc. 4:2 (1894) 162; Gibbs, J. Linn. Soc. Bot. 42

(1914) 81; Merrill, J. Str. Br. Roy. As. Soc. 86 (1921) 451. Pternandra tessellata

Ohwi, in scheda (L, BO).

Shrub or tree 3—12 m tall, diameter up to 15 cm. Branchlets distinctly 4-undulate-

winged, minutely pilose, 1—1.5 mm thick, epidermis dark brown when dry, flaking off

(including wings) leaving a smooth, cylindric, glabrous, tan-khaki coloured surface.

Blades chartaceous to subcoriaceous; lanceolate, elliptic, to ovate; acuminate at the tip

(acumen up to 10 mm long), broadly rounded and shallowly (1—2 mm) cordate at the

base; prominently 3-nerved from the base, intramarginal nerves 1—2 mm from the margin,

less prominent, but often giving the blades a 5-nerved appearance; nerves sunken above,

prominently raised and tapering below; secondary venation pinnate from each of the 3

main nerves, c. 20 pairs, slightly raised below, reticulate; juvenile blades puberulous on

the main nerves near the petiole, adult ones glabrous; drying olive-brown to blackish

above, lighter brown below, very brittle; 7.5—12 cm long, 3—5.5 cm wide. Petioles c. 2

mm long, c. 2 mm thick, at first minutely puberulous, later glabrous. Inflorescences from

leaf axils, often terminal, composed of solitary flowers or 2—3-flowered umbels;-axes

flattened, minutely puberulous, c. 1 mm thick. Bracts and bracteoles lanceolate, acute,

c. 1.5 mm long, minutely puberulous; primary axes 6—15 mm long, secondary axes

not developed (solitary flowers) or up to 10 mm long; pedicels 3—S mm long. Calyx tube

campanulate, covered with tessellate-umbonate-tuberculate scales, lower ones smaller and

flatter increasing in size and projection to the margin where they are triangular-tuber-

culate; 4-5 mm long, 4—5 mm wide, minutely puberulous, glabrescent, green; calyx cap

conical, acuminate, larger buds with a few minute tubercles in the middle, smaller ones

smooth, minutely puberulous throughout, falling off as one piece. Bud petals broadly

Ovate, acute at the tip, narrowed to the truncate base, 5-6 mm long, 7—8 mm wide,

thick, lilac-blue to dark red-violet. Bud filaments flat, c. 1 mm long, inappendiculate,

c. 2 mm long, yellow. Bud stigma 4-lobed, c. 1.5 mm long; style c. 4 mm long. Fruits

Fig. 16. Pternandra tessellata (Stapf) Nayar - -

Gard. Bull. Sing. 34:1 (1981)

24 4

fe ee ees es a

A-E: Haviland 1383 (isotype).

Maxwell: Pternandra (Melastomataceae) 69

campanulate, truncate, scales flatter and more tessellate, triangular and erect near the

margin, glabrous, areolus with conspicuous lines; style persisting, 8-9 mm long, 10—12

mm wide, green, drying dark brown to black; pericarp c. 1 mm thick, gritty. Seeds

numerous, flattened, quadrangular-cuneate, glossy khaki-tan, c.0.75 mm long.

Vegetatively, with its sinuate-winged branchlets and cordate blades, this species re-

sembles P. azurea (Bl.) Burk. var. cordata (Korth.) Maxw., however there are differences

which include the setose and often branched calyx scales of the latter. Immature buds

of P. tessellata frequently resemble those of P. rostrata (Cogn.) Nayar when the tubercles

on the cap are not apparent. The rostrate tip and calyx scales are also similar.

The calyx scales of P. tessellata also resemble those of P. tuberculata (Korth.) Nayar,

however the cap in that species is without any trace of tubercles. Unfortunately, flower-

ing material of P. tessellata is sparse, thus I cannot be certain of the morphology of the

mature internal flower parts. Stapf notes that his species is close to Kibessia simplex

Korth. (=P. echinata Jack) and to Curtis 953 from Penang which is a syntype of

P. griffithii King (=P. tuberculata). P. tessellata differs in many significant ways from

P. echinata Jack, e.g. the length of the scales, winged branchlets, shape and pubescence of

the leaves, etc.

P. tessellata appears to be a distinct species without any intermediate forms linking

it to other species. More material, especially flowers, is required to better understand the

species.

Stapf lists the type collection as being Haviland 1303 from Koung, Tampassuk River,

Mt. Kinabalu, Sabah; however I believe that the actual number is 1383 since specimens of

this species in the Singapore, Leiden, and Kew collections collected by Haviland at this

locality in 1892 are labeled as K. tessellata Stapf. I have not seen Haviland 1303 in any

collections of Melastomataceae that I have studied.

Fig. 16. A,B: calyx;C: bud petal; D: stamen; E: fruit.

Distribution:

Borneo Sabah — Bukit Tibang: Anderson & Paie 28628; Long Bah: Brooke 9037;

Mt. Kinabalu: Clemens 26798, 26906, 30349; Gibbs 3969; Haviland 1383 (holotype

K, isotypes L, SING). Quoin Hill, Tawau: Brand 24547; Tambato, Tambunan: Puasa-

Angian 3988; Tambunan Hill: Alphonso 57.

Kalimantan — Central East, W. Koetai: Endert 2961, 4561.

16. Pternandra teysmanniana (Cogn.) Nayar, Bull. Bot. Survey India 17 (1978) 53.

Kibessia teysmanniana Cogn. in DC., Monogr. Phan. 7 (1891) 1109; Merrill,

J. Str. Br. Roy. As. Soc. 86 (1921) 451; Bakhuizen f., “Thesis” (1943) 323

(name),Med. Mus. Bot. Utrecht 91 (1943) 323(name),Rec. Trav. Bot. Neerl.

40 (1943—45) 323 (name). Kibessia pachyphyllum Bakh. f., 1.c. (1943, 1943—

45) 318, syn. nov. Pternandra pachyphylla (Bakh. f.) Nayar, l.c. 52, syn. nov.

Tree (other field notes lacking). Branchlets cylindric, glabrous, 1.5 mm thick, drying

blackish; upper nodes flattened, c. 3 mm thick. Blades coriaceous, glabrous, elliptic,

ovate, to obovate, rounded at the tip, narrowed to rounded at the base; 3-nerved, veins

sunken above, raised and tapering below; intramarginal nerves invisible; secondary vena-

tion pinnate, reticulate, distinct above, invisible to obscure below; 3—5 cm long, 1.5—3

70 Gard. Bull. Sing. 34:1 (1981)

cm wide, drying greenish to brown above, tan to lighter brown below. Petioles 1.5—2.5

mm long, 1.5—2 mm thick, glabrous. Inflorescneces glomerulate, on tubercles from leaf

axils or just below them, many-flowered, glabrous; primary axes usually several per axil,

c. 1 mm long, each with a solitary pedicel 1—2 mm long. Bracts and bracteoles ovate,

acute, 0.5—1 mm long, glabrous. Calyx campanulate, glabrous, pattern closely tuberculate

in the lower part, with larger and longer tubercles near the margin; 3—4 mm long, 2.5—3

mm wide; cap conical, acute, glabrous, smooth, thin, c. 2.5 mm long, splitting into several

(4?) caducous lobes. Petals broadly ovate, acute at the tip, narrowed and truncate at the

base, thickened with thinner margins where the venation is visible, 3—5 mm long, 2—2.5

mm wide. Filaments flattened, 2 mm long; anthers 2—2.5 mm long, 1 mm wide, with a

connective spur near the filament; stigma cylindric, 1 mm long. Style slender, 3—4 mm

long, elongating to 6 mm in fruit. Fruits urceolate, 5 mm long, 5 mm wide, pattern

tessellate to slightly umbonate, exocarp drying brown; pericarp c. 0.2 mm thick. Seeds

numerous, flattened, cuneate-quadrangular, 1—1.25 mm long, glossy brown.

The relatively small (the smallest for the genus), thick, blades with a rounded tip;

fascicled inflorescences, small flowers, and urceolate fruits with a tessellate pattern

distinguish this apparently rare species from all others.

The inflorescences and fruits are similar to those of P. galeata (Korth.) Ridl. and

P. cogniauxii Nayar, however the former has a tessellate pattern on the calyx and the

latter is easily recognized by its large, cordate, 5-nerved blades. Pternandra teysmanniana

is not, as Bakhuizen f. notes under Kibessia pachyphyllum, related to kibessia acuminata

Decne., which is, in my opinion, related to Pternandra echinata Jack. The calyx pattern

of K. acuminata is more tuberculate to echinate and is probably and underdeveloped

form of P. echinata where the scales are flattened-echinate.

Kibessia pachyphyllum Bakh. f., only known from the type collection, is undoubted-

ly identical to P. teysmanniana. The holotype of the former (Hallier 2223, from Kaliman-

tan) is without inflorescences or fruits, however comparison of its leaves and buds with

those of two collections of P. teysmanniana confirms its synonomy. Bakhuizen f. did not

describe K. teysmanniana in his monograph. Nayar’s recombination of K. pachyphyllum

Bakh. f. to Pternandra was done without having examined any relevant material, thus

indicating his failure to contribute to or understand the taxonomy of Pternandra.

Fig. 17: A: inflorescence; B, C: calyx; D: mature petal; E: stamen.

Plate 6. Holotype of Pternandra teysmanniana (Cogn.) Nayar.

Distribution:

Borneo Kalimantan — Mdjang: Teysmann 8165 (holotype FI, isotype BR); Sei

Kenepai Singang: Hallier 2223 (Kibessia pachyphylla Bakh. f., holotype L, isotype BO);

Soengai Djemila: Hallier 2247; exact location unknown: Jaheri 1893.

Maxwell: Pternandra (Melastomataceae)

Fig. 17. Pternandra teysmanniana (Cogn.) Nayar - - A, D, E: Teysmann 8165 (holotype); B, C: Hallier

2223 (Kibessia pachyphyllum Bakh. f., holotype).

+ L beugt “IY He

So hw NN = OF

Plate 6. Pternandra teysmanniana (Cogn

after J. E. Teysmann (1808-1882

Archipelago. This specimen is from

Leiden.

Neo. SI bi es

Communic, ex. Heth. Hort. Bot. Bog,

Ly Ce

Archipel. Lad.

pre : f

Borneo. - é flag,

Le ve

AOE T pp.

oy GM Fey

ig Uittey. Ce

Teyemann.

) Nayar, holotype at Florence. Collected by and named

, a Dutch botanist who collected extensively in the Malay

Borneo and was collected in 1876. Photo: Rihksherbarium,

Y sets a

Maxwell: Pternandra (Melastomataceae) 73

17. Pternandra tuberculata (Korth.) Nayar, Bull. Bot. Surv. India 17 (1978) 53; Keng,

Gard. Bull. Sing. 31 (1978) 110 (@ tuberculata (Korth.) Maxw.). Ewyckia

tuberculata Korth. In Temm., Verh. Nat. Gesch. (1844) Bot. 255. Rectomitra

tuberculata (Korth.) Bl., Mus. Bot. Lugd.-Bat. I:1 (1849) 7. Kibessia tuberculata

(Korth.) Hk. f. ex Cl in Hk. f., Fl. Brit. India II (1879) 553; Cogniaux in DC.,

Monogr. Phan. 7 (1891) 1110; Koorders-Schumacher, Syst. Verz. 2 (1910) 47;

Bakhuizen f., “Thesis” (1943) 322, Med. Mus. Bot. Utrecht 91 (1943) 322,

Rec. Trav. Bot. Neerl. 40 (1943—45) 322. Pternandra griffithii King, J. As. Soc.

Bengal 69, II:1 (1900) 70 (Mat. Fl. Mal. Pen. 3, 478), syn. nov.; Pternandra

griffithii King var. cordata King, |.c. 71 (l.c. 479), syn. nov. Pternandra panicu-

lata Benth. ex Ridl., Fl. Mal. Pen. I (1922) 809 (including P. griffithii King).

Ewyckia korthalsiana Miq., Fl. Ned. Ind. Suppl. I, Sumatra (1860) 124

(nomen), 321. Pternandra korthalsiana (Mig.) Triana, Trans. Linn. Soc. 28

(1871) 153, syn. nov.

Tree up to 20 m tall with a maximum diameter of 50 cm. Bole straight, not buttress-

ed; outer bark 0.5 mm thick, grey to brown with many shallow fissures, smooth and peel-

ing off; inner bark white, wood yellowish. Branchlets terete, smooth, glabrous, drying

blackish, c. 2 mm thick, nodes flattened and swollen, c. 4 mm wide. Blades subcoriaceous

to coriaceous, glabrous, broadly lanceolate to broadly ovate, acuminate at the tip

(acumen 6—10 mm long), narrowed and decurrent or rounded to cordate at the base;

3-nerved from the base, nerves sunken above, raised and tapering below; secondary

venation pinnate, c. 15—20 pairs, reticulate; 5.5S—12 (19) cm long, 3—6 (9.5) cm wide,

dark green above, green below; drying olive to blackish above, tan to dark brown below.

Petioles 4—5 mm long, 1—2 mm wide, glabrous. Inflorescences cymose, from leaf axils, less

commonly leafless nodes, often terminal, 1—2.5 cm long, several to many flowers per

axil; axes 4-angled, puberulous. Bracts and bracteoles lanceolate to ovate, obtuse to acute

at the tip, 2—3 (S) mm long, larger ones concave, minutely puberulous. Primary axes | to

several per axil, 4-20 mm long, with 1—3 nodes, up to 1 mm thick; secondary axes 2—4

mm long, pedicels 2—3 mm long. Calyx campanulate, 4—S mm long, c. 4 mm wide, green,

covered with minutely puberulous, often appressed, tubercles which are shorter near the

pedicel and longest (c. 2 mm) near the rim; cap mammiform, papillose-puberulous, smooth,

c. 2 mm long, splitting into 4 triangular lobes. Bud petals imbricating to the right, when

mature thickened, broadly ovate to suborbicular, acute at the tip, truncate at the base,

4—5S mm long, 4—5 mm wide, lilac. Filaments flattened, c. 2 mm long, lilac; anthers c. 2

mm long, yellow. Stigma 4-lobed, c. 2 mm long, yellow. Style slender, glabrous, 4-5 mm

long, whitish. Fruits campanulate, truncate at the margin, 7—8 mm long, 7—8 mm wide,

pattern umbonate-verruculose; exocarp grey-green, drying black. Seeds numerous,

flattened, cuneate-quadrangular, c. 0.5 mm long; testa glossy tan.

The type collection of P. tuberculata (Korthals s.n., from Sumatra) has some in-

florescences and young buds. From the descriptions of this species it is apparent that it

has a calyx cap which splits into 4-lobes. The leaves of the type material match those

on Griffith 2272/1 — which is the lectotype of P. griffithii. This specimen and the syn-

type collection (Curtis 953) have buds and mature flowers. The nature of the calyx cap

and inflorescnece is different from those of P. rostrata which has shorter, thicker, and

glabrous flowering axes; an acute to rostrate cap which falls off in one piece; and gene-

rally larger blades. This last point has many exceptions, e.g. Koster 1481, from New

Guinea, has leaves larger than the type collection of P. rostrata (Beccari 979); and

Meijer 37999 (P. rostrata) which has leaves smaller than Griffith 2272/1. 1 have not been

able to separate the two species on the basis of vegetative characters since the leaves vary

considerably for each.

74 Gard. Bull. Sing. 34:1 (1981)

The type collections of P. griffithii are probably the same as P. tuberculata. I have

attempted to construct an idea of P. tuberculata using P. griffithii as a basis. Cogniaux

and Bakhuizen f. adopted this approach and considered P. griffithii King as a synonym,

however Ridley combined P. griffithii with P. paniculata Benth. ex C1. which is based

on a specimen with an abnormal, witches broom inflorescences. I agree with King and

Bakhuizen f. in considering P. paniculata Benth. ex Cl. as a synonym of P. coerulescens

Jack. Another possibility is that the two species are different and that P. coriacea (Cogn.)

Nayar is the same as P. tuberculata. The basic problem lies in trying to determine what

the nature of P. tuberculata really is.

The blades of the type collection of P. tuberculata closely resemble those of Beccari

3164 and 3324 — both types of P. coriacea. The leaves on these collections are always

small, in contrast to those of P. griffithii which often become rather large. The basic

structure of the inflorescences of P. tuberculata, P griffithii, and P. coriacea are similar

and the only distinction I can find between the taxa lies with the calyx cap which falls

off as a unit in P. coriacea. It is due to the nature of the cap plus its small leaves that I

have followed Cogniaux in considering P. coriacea distinct from P. tuberculata. Both

species are uncommon and more flowering and fruiting material are needed to fully

understand these taxa.

The holotype of Ewyckia korthalsiana Miq. at Utrecht (Teysmann sn, from Bangka

Island) is without flowers or fruits, however there are two short parts of two inflore-

scences remaining. &. korthalsiana is written on the original label and on another more

recent tag P. capitellata Jack is noted. The primary axes on this specimen are between 8

and 10 mm long, thus they are longer than the maximum (3 mm) generally considered for

P. capitellata. Bakhuizen f., therefore, included E. korthalsiana under P. coerulescens

Jack var. cyanea (Bl.) Bakh. f. Cogniaux, however, included it under P. capitellata. Since

the specimen lacks flowers and fruits I cannot be absolutely certain that it is P. coerule-

scens.

The blades and remnants of the two inflorescences on this specimen, in my opinion,

more closely resemble Griffith 2272/1 — which is the lectotype of P. griffithii King. The

shape, texture, and colour of the dry leaves match, and the inflorescence axes are more

like those of P. griffithii than of P. coerulescens. Miquel’s otiginal description indicates

that the calyx is “obtuso-4-dentatus, areolato-verrucellosus”. This is not a feature of

P. coerulescens which has a truncate calyx with a tessellate or somewhat umbonate

pattern. Teysmann’s specimen, from its general appearance, more closely resembles

several other collections of P. tuberculata than P. coerulescens which tends to dry black.

Wray 1994, lectotype of Pternandra griffithii var. cordata, has slightly cordate blades.

However, this feature is only developed on some blades and in others the indentation is

more like that of an auricle. Most specimens of P. tuberculata in the Singapore and

Leiden collections have acute bases, but some of the blades in Robinson & Kloss 6148 are

rounded, and in fact look like some of the blades in Wray 1994. The texture of

the blades in Wray 1994 is very similar to many specimens of P. tuberculata (Ridly s.n.

from Biodi Tepah, Nov. 1908 — which has rounded blades, and Nauen 35879). Some of

the blades in these specimens are broadly rounded and some appear to be almost cordate.

The distinction of var. cordata, in my opinion, does not merit a separate taxon, therefore

[I have united it with P. tuberculata.

Maxwell 77—22, from Singapore, differs from all other flowering specimens of

P. tuberculata that I have seen in having a thin, mammiform cap which falls off in one

piece. The collection in all other respects matches other collections of P. tuberculata.

Maxwell: Pternandra (Melastomataceae) 75

er) 3

Fig. 18. Pternandra tuberculata (Korth.) Nayar - - A, G: Buwalda 6239; B, E: Griffith 2272/1

(Pternandra griffithii King, lectotype); C, D: Maxwell 77-22; F: Koster 1481.

16 Gard. Bull. Sing. 34:1 (1981)

Fig. 18. A,B: calyx;C: calyptra; D: petal; E: stamen; F: style in bud; G: mature style.

Plate 7. Holotype of Pternandra tuberculata (Korth.) Nayar.

Plate 8. Lectotype of Pternandra griffithii King.

Distribution:

W. Malaysia. Kedah — Serdang: Nauen 35879; sine loc.: Meh 17863.

Penang — Curtis 453 (syntype P. griffithii King var. cordata King, not seen), 953

(syntype P. griffithii King), 2784, 2786, sn in June 1890.

Perak — Bidoi Tapah: Ridley sn in Nov. 1908; Larut: Wray 1994 (P. griffithii King

var. cordata King, lectotype SING); Pondok Tanjong For. Res.: For. Dept’t. Fed. Mal.

States 1147; Taiping: Ridly 14685 (abnormal).

Trengganu — Ulu Brang: Moysey & Kiah 33646 (abnormal).

Selangor — Batu Tigai: Ridley 11936; Genting Highlands Rd., 9th mile: Kochummen

023063; Semangkok Forest Reserve: Chan 13270.

Malacca — sine loc.: Griffith 2272/1 (P. griffithii King, lectotype K).

Singapore — Ahmad 1054, 1365; Anderson 63, Maxwell 77—22; Ridley 3918,

4729.

Riouw Archipelago. Bukit Tanjong: Buwalda 6239.

Sumatra. West Coast — Gunong Koerintgi: Korthals sn (Ewyckia tuberculata Korth.,

holotype L); Kuala Belilas: Buwalda 6693.

Bangka Island — Soengailiat: Teysmann sn (Ewyckia korthalsiana Miq., holotype U,

isotypes (?) U, BO).

Borneo Sabah — Sosopodon: Mikil 38499,

New Guinea llolo: Streimann & Kairo 26184; Japen Island: Koster 11178; Kaloal:

Koster 1481.

18. ABNORMAL AND UNIDENTIFIED PTERNANDRA.

There are several collections of Pternandra in the Singapore and Leiden herbaria

that have abnormal inflorescences with witches broom deformities or are vegetative (Plate

2). With many of these specimens I am reasonably certain as to their identities (indicated

by aff.) however with others I cannot be absolutely certain what species they are.

Since the vegetative characteristics of P. echinata Jack and P. azurea (Bl.) Burk. are often

similar I have, in most instances, been able to sort these specimens on the basis of collect-

ing locale, that is P. echinata is mostly restricted to the Malay Peninsula and P. azurea to

the Malay Islands. A more difficult problem lies with abnormal or vegetative collections

of what closely resembles P. coerulescens Jack and P. tuberculata (Korth.) Nay. from the

Malay Peninsula, and these two species plus P. galeata (Korth.) Ridl. and P. rostrata

(Cogn.) Nay. from the Malay Islands. The branchlets and leaves of these four species

Woh GN «67

- 8 Bou v7 wy HY

Lf of FES

i y I i s

(ef. & * es. bia Lined ttebala

& ers BEE j j Ranks,

a Oat

RAGLSGLa Feet

ATE. 5 Ede

Det. RC. Bakhuizen Utrecht “* 19 y2

v.d° Brink fil

tae ]

Herb, Lugd. Batay.

nal —— a

- hip kn 2

* *

- ~ ae 9 int

Plate 7. Holotype of Pternandra tuberculata (Korth.) Nayar at Leiden. This species was originally

described by Korthals in 1844 as Ewyckia tuberculata Korth. and was recombined by Blume

as Rectomitra tuberculata (Korth.) Bl. in 1849. It was changed to Kibessia tuberculata

(Korth.) Hk. f. ex Cl. in 1879 and finally to P. tuberculata (Korth.) Nayar in 1978. Photo:

Rijksherbarium, Leiden.

Plate 8.

Lectotype of Pternandra griffithii King at Kew which has been reduced to a synonym ot P.

tuberculata (Korth.) Nayar in this revision. Pternandra griffithii King, from Malacca, was

named by King in 1900 in honour of William Griffith (1810-1845), a British surgeon-

botanist, who collected extensively and published detailed accounts of the flora of British

India and the Malay Peninsula. Photo: Rihksherbarium, Leiden.

Maxwell: Pternandra (Melastomataceae) 79

often resemble each other, therefore without flowers or fruits I have not ventured

to guess what species these specimens are. Hopefully collectors will take notice of

abnormal Pternandra inflorescences and make an effort to collect normal specimens from

the same tree so that future workers can study the degree and variability of such de-

formities which can be positively identified.

In several specimens, e.g. Ridley 391 (8?); Ridley sn, from Bidoi Tapah, Perak in

November 1908; Nauen 35879; and Moysey & Kiah 33646; the inflorescences are abnormal

in that they are 8—15 cm long, with a few rudimentary leaves and enlarged, lanceolate,

acuminate bracts which often resemble the leaves; and with or without a few, often de-

formed, flowers. These specimens, in my opinion, are P. tuberculata (Korth.) Nay. since

what I can see of the calyx resembles this species more than that of P. coerulescens

Jack.

Ridley was of the opinion that P. paniculata (Miq.) Triana is a panicl:d form of

P. griffithii King where the inflorescence is “abnormal or converted into a nass of small

leaves, with a few abnormal flowers intermixed.” Bakhuizen f. considered P. paniculata

as a synonym of the typical variety of P. coerulescens Jack. Moreover, he notes that

P. paniculata is based on a specimen with a witches broom deformity of the inflore-

scence. Apparently this abnormal condition affects both P. coerulescens and P. tubercu-

tata, however it is extremely difficult to tell the species apart without normal flowers or

fruits.

The calyx tube in some specimens, e.g. Kostermans 13664 (Borneo) and others with

this abnormal inflorescence, is tuberculate with a distinct and smooth cap. These speci-

mens are most likely P. rostrata or P. tuberculata.

Other specimens, e.g. Teysmann sn, from Bangka Island — the holotype of

P. (Ewyckia) paniculata (Mig.) Triana; Winkler 2408, Haviland =177 (both from Borneo)

and others have a tessellate calyx and no cap. These collections are clearly P. coerulescens

Jack.

Kibessia simplex Korth. var. oblonga Bl. (Mus. Bot. Lugd.-Bat. I:1 (1849) 9) is

described from a specimen (Waitz sn, from Borneo) which has an abnormal inflorescence

which resembles those of P. azurea and P. coerulescens. Unfortunately, the specimen

lacks buds, thus I cannot be absolutely certain of what species it really is; however from

the leaves and branches var. oblonga more closely resembles the type collections of

P. azurea (Bl.) Burk. and Kibessia simplex Korth. — which I consider a synonym of the

former. I do not agree with Bakhuizen f. in reducing var. oblonga to a synonym of

P. coerulescens Jack.

Craib (F1. Siam. Enum. 1:4 (1931) 702) notes that several collections of P. coeru-

lescens Jack from southern Thailand have abnormal inflorescences. He also indicates that

several species of Diospyros (Ebenaceae) and Maesa (Myrsinaceae) are also affected in

this manner.

Specimens Examined

W. Malaysia Kedah — Kedah Peak: Robinson & Kloss 6148.

Penang — |. H. Burkill 6145 (aff. 5).

Perak — Ayer Larut: Wray sn; Dindings: H. M. Burkill & Shah 185 (aff. 5); Pangkor:

Ridley sn; sine, loc.: Scortechini 248, sn.

80 Gard. Bull. Sing. 34:1 (1981)

Singapore — Murton 92 (aff. 8), Ridley sn at Chan Chu Kang.

Sumatra. Bangka Island: Kostermans & Anta 85.

Borneo. Sarawak — Bukit Lambir, Baram District: Chew 961; Segan For. Res::

Wright 27105; Sungei Lebang, Bintulu: Ashton 16299.

New Guinea. Warsamon Valley, Sorong: Moll 11601, 11670 (aff. 9, 17).

Sine Loc.: Boerlage 6688 (aff. 2).

APPENDIX

1. Kibessia ? angustifolia Bl., Mus. Bot. Lugd.-Bat. I, 1 (1849) 9. This is probably

P. echinata Jack.

2. Pternandra discolor Cogn. in DC., Monogr. Phan. 7 (1891) 1194. An isotype at

Brussels, Balansa 1145, from Tonkin, is Rhodamnia dumetorum (DC.) Merr. & Perry,

of the Myrtaceae.

3. Pternandra ? cordifolia Cogn., \.c. 1105. An isotype at Leiden and the holotype at

Brussels, Beccari 570 from Sarawak, consist of leaves of Medinilla muricata Bl.

4. Kibessia galeata var. latifolia Ohwi, Bot. Mag. Tokyo 57 (1943) 18. The

type (Kanehira & Hatusima 13146, New Guinea) has not been seen.

5. Pternandra moluccana Nayar, Bull. Bot. Surv. India 17 (1978) 53. The holotype

at Leiden (Pleyte 253 from Halmaheira) is Medinilla sp.

6. Pternandra parasitica Finl. & Wall. (Wall. cat. 4086A) is Pogonanthera pulverulenta

(Jack) BI.

7. Kibessia celebica Koord., Fl. van Celebes II (1898) 466 is Medinilla myrtiformis

(Naud.) Triana.

8. Pternandra simplex (Korth.) Baill., Hist. Pl. 7 (1880) 24, f. 32, 33. The two figures

cited do not represent this species (the correct name of which is Pternandra azurea

var. azurea) and it is most likely that they were based on P. rostrata (Cogn.) Nayar

or perhaps P. tessellata (Stapf) Nayar.

9. Pternandra ledermannii (Mansf.) Nayar, Bull. Bot. Survey India 17 (1978) 53.

Kibessia ledermannii Mansf. in Engler, Bot. Jahr. 60 (1925) 138.

This species, from New Guinea, supposedly differs from P. coriacea (Cogn.) Nayar

in having larger leaves, shorter pedicels; and smaller, flatter calyx tubercles. P. coriacea

also has glaucous blade undersurfaces which dry whitish to greyish, and are more ovate

and less lanceolate-elliptic than those of P. ledermannii. The tubercles in P. coriacea are

more acute and in P. ledermannii more flattened and hexagonal.

I believe that P. ledermannii is closer to P. rostrata (Cogn.) Nayar, however I cannot

be certain of this until more New Guinea material becomes available. Mansfield lists

Ledermann 8717a, which I have examined at Singapore, as possibly being equal to the

Maxwell: Pternandra (Melastomataceae) 81

holotype: Ledermann 8711 — which I have not seen. The collection at Singapore consists

of a few flowering axes and one flower bud. The calyx pattern and the thickened, acumi-

nate cap, which apparently falls off in one piece, resemble P. rostrata more than any

other taxon. It is definitely not P. tuberculata (Korth.) Nayar or P. glaeata (Korth.) Ridl.

var. galeata both of which have caps have caps that split into four lobes.

Nayar, without having examined any authentic material of Kibessia ledermannii

Mansf., merely transferred this species to Pternandra without any discussion. Needless

to say, such pseudo-academic pursuits, however beneficial to their author, merely create

more unnecessary confusion and certainly detract from one’s taxonomic credibility —

and in the case of Nayar, an already dubious one.

| INDEX TO TAXA

(numbers refer to the accepted taxon)

Peas caniiciata (Jack) Waipie LeU) ee. 4

(alias tte ASABE MINS Se ee len a: OP? Love tae i an aii ag ea aa 5

ee ee Cray aiig tat) Oar is ACL ee oe ee a RS Le 8

pereere ramet terete alias ct ib ude Saran SS oe. 9

Eerste cpg in a gtd oN Ce eG Sy Soa oN ow esis 0 5

INES re. ve cn ee. Le anes Ok ee ge wad ee i7

aE ee oes ee ee care eee tle oe ees te 5

nm nla yar, membranace”a Bl: . : 2... eee Pies Vi idk ee et 5

PT ee ot Oe cia nce a al iMie ae ee 5

Ee oe ote hin atage chiles os see s,* = =.

i be eh GR eet ee Me yc were acs es re

unmnmmrmnrriisreerne. 1 2 2G) jh Ree Sista a sok fa 2 Get eg ee ss 8

RL a I a aie ae ea ae 2

INE eee ek Tobe ace veces ea Per se te he AT Sys ava-e “pe ws 14

TSP l tis an eh Ss ols fine bank ese mene sop 8

DIRE Re hae Caer ng Att, ORES ee Ck Ske Fae SS 7

Bee a wet aubalai (BLY Nig.. |: 25. ee ee Z

mrememire Te wars iypien HOchr.<)) 2 20s 2 Pe OP ae 2

TOS REISS Sing LA SA a GL de ge ae a are Sa appendix

nN Ut tt Uo h ae a Sof e st IS FT a ono eed. ee 3

aT 8 aE TG ii Rien: 5 Se ner pe, dR, a a be a 6

EI Seen Pa ee a vic teh apg as ee 8

I 8 | Bk ot a eek ee a ee 8

ee ea A Ge eee ay ee Be ee os ela ve <p 10

REET ROE rh, SS Gyo ap fe gu es <4 is sce ncergee Boek 9

nee, Wal emir lis LUWI ee eee se es w s appendix

K. galeata (Korth.) Cogn. var. pluriflora Mansf. ..............005-. 9

eB Fa sR WES aS pa aia Se 6 eee ea 1]

eg de 00 Sy Fly week ase akpicin e 6 Skt 8 es 12

II SO sn Pe ety we ea wre ds mee 8 tee 4

PD oS et a noe a kw Beye: preyep ee appendix

ee ee ee g visca Wal b wie Bw a we eters 6 wens 16

ne te aay A ahs nts Wises os 6 se hes a 8

eee, fy aa gs bx sc bobo a po eens 8 e pomee at 14

I Et Cr nck es a ote wee a ee eo ee 8 2

A a ee are 8

TELE OMIGTIEd Bl... ek ee ee we ws 2

82 Gard. Bull. Sing. 34:1 (1981)

Ko stibalata Bh. i coin ce ok yale bn pn bd ag ne ace 2 .

K. subulata Bl. (sphaim. Triana) 2. oe. 2 ces a 2

K tesseliata' Stapf... . 02. oh ees 0 A ae 15

K. tetraptéra Mig... 2 eee ae ae ge 3

K. teysmanniana Cogn. 20.00 a 16

K, tuberculata (Korth.) Hk. f. ex Cl. «9. ie. 9 ss oe 17

K. tuberculata (Korth.) Hk. f. ex Cl. var. grandiflora Bakh.f. .......... 14

K. verrucosa Metr. .'. oe ai ee on een 14

K. verrucuiosa# Metr. 2.) 5.) a Pe 14

Macroplacis cofdata (Korth.) Bl... ee? 4

Nelastomaazureum Bi... a ee ee eee oo ae

Memecylon oligoneuron Bl. var. maluense Mansf. .................. 5

Pternandra acuminata (Decne.) Nayar. .. ova... v cae 8

P. acuminatissima (Metr.) Nayar oo... 5 2% « «sia + pe 2

Paffinis (Merr.) Nayar... 0 0 ee ee © 14

P: angustifolia Maxw. . . sje ds es Ste ee eee |

P. azurea (Bl.)-Burk. var. azuréa’:. 2. 0 a on ee ee eee Z

P. azurea (Bl\.) Burk. var. cordata (Korth, )(Maxw. S.A 5

P.-birtella (Cogn.) Nayar (sphalms)" St Oe ee ys

P. borneaensis.Nayat .. 2 6260.00 ca oe olla a Z

P. capitellata Jack 20.0 0... ace 4 a oe oleate 2 5

P. cardiophylla ORW1 3.3 eo oO PS ee eee 4

P. coerulescens Jack |... Se Sa eee 5

P. coerulescens Jack var. capitellata. Jack) Kine. 5... ..4 .s.3 ee 5

P. coerulescens Jack var. cyanea (BI) Cogn. S25... 2) oe Sa ee bid

P. coertulescens Jack var. jackiang’ Cl, A ee eee 5

P. coerulescens Jack var. membranacea (Bl.) Furtado. .............. 5

P. coerulescens Jack var. paniculata (Miq:) King”. 22.0. ee ee 5

P. ecogniauxit Nayar’. 22 8 SP OO ae ee eee 4

P. cordata (Korth.) Baill. 2°." WSS st ee ee 3

P.) cordifolia Cogn). so Se Pek a ee ee eee appendix

P. coriacea (Cogn.). Nayar 0) ).0 Sid es ee eee 6

P. crassicalyx-Maxw) «3 624s oa Pe ee eee 7

Ps.dwscalor Cogn. ©. os S295 Ge ke ie Oe ee ee en appendix

P-echinata Jacko. ek, ea ee eee 8

P. echinata Jack var .:bracteaia Rid. fh en aes oa eee 8

P. echinata Jack var. pubescens (Deene.) King: . 2... 5. oss ee 8

P. elmeri (Metr.) "Nayar ..82 Po SSS oe eee 10

P. forbesit E-Gi Bakker 20d = I RS TIN ate VR cy enh eae 9

P, galeata (Korth.) Ridli vat. galeata° >) oo a eee 9

P. galeata (Korth.) Ridl. var elmeri (Merr.) Maxw................0. 10

P. gracilis (Cogn) Naga | heodt iat tei je « fale OE 1]

P. grandifolia (Bakth f7¥ Nayar is:.60..5'. sal see ee 14

P, griffithii King 30.0. 0 eh oe ee eee 17

P. griffithii King var. cordata King. ...:. 220... 22) eae A eee 17

P-hirtella (Cogn.) Nayar... io .20/ th ea ed oe ie

P. jackiana (C1,) Ridl. . os... oa ae 5

P, korthalsiana (Miq.) Triana... 0...) 2. 7 nt 2 17

F.. latifolia (Bl.) Triana) 23S a Ge Ol ee 5

P. ledermannii (Mansf.) Nayar .. 2. . 3. 5.252. 2 appendix

P. moluccana Nayar... .-.s.. ys a0. s sh appendix

P, multiflora Cogn... 6 ee ad ee 13

Maxwell: Pternandra (Melastomataceae) 83

niemilnaMhiay 1895-4) 4 i ia..tpalst, Maccd PER oe ee 17

IS ORE 2 Gy Seat diasdy. otents Oo... 26S bee J 5

err eanes we ee SIS 2 Cac). AES besarte re lgdat were ore appendix

een Cemurmd ) AOU. 10) NS. £2 ).00. 1). EGE WALL SE we. 14

renee remit. 325 -O CE 1d Sall 22} Iba 25k, ea. - 2, appendix

IME Pr ft tenn et eae Aiadaccc ital. Ee) a 15

GR oS 2 eae 15

TES eT a ee ee 16

eens Gta) NAVAN) =). Oe toc eee RIA UET beelive b)K 17

P. verrucosa (Merr.) Nayar ...... age a AEE DE ROIRt. £63.80 224 14

ee CUB POSS By i Te 8) Se Pe ae 1]

mnumr crmiaie (erin) Iniana Oe Pol tS... ee ea 4

PPeMIDE Gy 3) 208. Joe oo). Se eL Sahl. (RIA ae ie. - 9

eenremmrmimo) 18k INe Gees. ok SELL ee 2.eG) Bid: 17

INDEX TO COLLECTIONS

(excluding all collections listed by Bakhuizen f. (1943, 1943-45)

except types or unless an identification change has been made)

Aban 31163 (2)

Aban & Saikeh 79379 (5)

Aet 200 (5S), 661 (5) 690 (5S)

Agama 436 (14)

Agama & Aban 40855 (14)

Ahmad 14335 (6)

S. Ahmad 359 (5), 1054 (17), 1365 (17), 1366 (5), 1442 (8), 1464 (5)

Ali sn Bukit Timah, Singapore on 18 April 1974 (8)

Alphonso 57 (15)

Alston 14331 (5)

Alvins 20 (5), 74 (5), 249 (8), 586 (8), 648 (5), 727 (8), 1151 (5), 1246 (8), 1284 (5),

2148 (8), 2249 (5); sn, Malacca -{8)

Amdjah 804 (5)

Amir 35638 (5)

Ampuria 32628 (14), 32707 (7), 36418 (14), 41333 (14), 41397 (14)

Anderson T29 (4), 62 (8), 63 (17), 3361/3 (6), 4873 (5), 12852 (5), 14643 (12), 26854

(12), 30709 (9)

Anderson & Hou 502 (7)

Anderson & Paie 28331 (2), 28628 (15)

Arshand 18029 (5)

Asah see Luang

Ashton 37 (14), 115 (5), 385 (11), 492 (7), 720 (11), 932 (9), 13975 (14), 16299 (18),

17717 (14), 17815 (12), 18371 (11), 19923 (2)

Au 23944 (7)

Au & Chai 24125 (2)

Baba 21483 (5S)

Badak 32349 (14)

Bain 6015 (8)

Bakar 26859 (7)

Baker sn, Singapore 4 Nov. 1917 (5)

Balajadia 2841 (5)

van Balgooy 2412 (5)

84 Gard. Bull. Sing. 34:1 (1981)

Banyeng & Sibat 24913 (12)

Bartlett 6869 (5), 6913 (5)

Batten-Pooll sn, near Sobaiga, Sumatra (3)

Beccari 17 (12), 179 (14), 301 (14), 700 (5), 742 (5), 968 (14), 979 (14), 1551 (13),

1962 (5), 2918 (13), 3141 (5), 3164 (6), 3172 (11), 3275 (5), 3324 (6), 3363 (5S)

Benang 24406 (14)

Bidin 84376 (5)

Binideh & Talip 65301 (14)

Bishop (Hose) & Haviland 230 (12)

Boden-Kloss 14558 (3), 19057 (14)

Boeea 5739 (3), 7604 (3), 7985 (2), 9314 (5), 9363 (3)

Boerlage 6688 (18)

Bojeng 9304 (12), 9428 (13), 14643 (12)

Brain 1291 (5), 5368 (12), 5369 (13)

Brand 24547 (15), 30862 (14), 30997 (14)

Brass 5959 (5)

Bray 11508 (5), 11627 (5), 11632 (8)

B. & K. Bremer 1770 (5)

Brooke 9037 (15), 10010 (14), 10640 (5)

Brown 2290 (12)

Brunig 1006 (6), 7662 (5)

Bujang 13698 (12), 20878 (6)

Bunaar 25836 (5)

Bunnemeyer 6424 (8), 6900 (2), 7025 (5)

H. M. Burkill 1866 (5)

H. M. Burkill & Shah 185 (18), 1033 (8), 1095 (8)

I. H. Burkill 382 (5), 961 (5), 1334 (5), 1356 (5), 3303 (5), 6145 (18), 12391 (5);

sn, Pulau Tinggi, Johore (8); sn, Sungei Baru, Malacca (8)

I. H. Burkill & Haniff 16241 (8), 16868 (8)

Burn-Murdoch 196 (9)

Buwalda 6239 (17), 6292 (5), 6693 (17), 6827 (9), 7935 (9)

Cantley 184 (8), 2900 (8), 2948 (5); sn, Singapore (5)

Carr 26691 (11)

Carrick 1482 (2412) (8), 1488 (5)

Carrick & Enoch 34 (5), 349 (12)

Chai 18523 (5), 18919 (7), 19317 (12), 19758 (13), 29822 (14), 31692 (14), 33177 (13)

Chai & Paie 17850 (5), 27931 (7), 31104 (4)

Chan 6732 (8), 13137 (8), 13195 (8), 13270 (17), 16878 (5), 17667 (8)

Charington 20853 (5), 20870 (14) :

Chelliah 98142 (5)

Chew 437 (7), 961 (18)

Chew, Corner, Stainton 7 (2), 97 (11), 1201 (11)

Chong 33666 (14)

Chow 75670 (14)

Clemens 20937 (14), 21564 (14), 21565 (2), 21575 (14), 21580 (11), 26355 (4), 26798

(15), 26842 (4), 26906 (15), 27873 (4), 28071 (11), 29367 (2), 29592 (11), 30349

(15), 30350 (2), 30515 (14), 30812 (14), 31285 (11), 32134 (11), 32474 (14),

34102 (14); sn, Mt. Kinabalu, May 1933 (11); sn, Mt. Kinabalu (14)

Cockburn 7138 (8), 7446 (8), 8499 (5), 10552 (5), 10720 (5), 10735 (5), 10907 (8),

11054 (5), 11077 (5), 11940 (5), 73016 (9), 85032 (2)

Cockburn & Saikeh 70033 (2)

Corner 30026 (5); sn, S. Sedili, Johore 27 March 1932 (9); sn, S. Sedili Kechil June 1934

(S); sn, Kemaman 22 June 1932 (5); sn, Balek Palu, Penang (5); sn, Grik, Perak (5);

sn, Singapore, July 1933 (5)

Maxwell: Pternandra (Melastomataceae) 85

Creagh sn, Gaya, Borneo, May 1895 (14); sn, Gaya, Borneo, Aug. 1895 (14); sn, Gaya,

Borneo, July 1896 (14); sn, Sandakan, Borneo (5)

Cuadra 1148 (14), 1336 (5), 1363 (4), 2190 (14), 2224 (14)

Cubitt 811 (8)

Cuming 2316 (5), 2337 (8)

Curtis 67 (5), 270 (8); =270, Bukit Bruang, Malacca May 1900 (8); =270, Penang Sept.

1890 and Mar. 1892 (8); 879 (5), 953 (17), 2337 (5), 2784 (17), 2786 (17), 3159

(5), 3470 (5); sn. Penang, June 1890 (17); sn, Gov.’t. Hill, Penang (5)

Dan 1907 (S1458) (13), 4375 (5), 26859 (7)

Dariser 2481 (2)

Daud & Tachun 35601 (12), 35604 (12), 35669 (14)

Denny 4 (5), 42 (5); sn, Selangor, 27 Nov. 1941 (5)

Derry 14 (5), 15 (5), 17 (8), 31 (5), 1159 (S)

Desmakh sn Singapore on 10 Sept. 1921 (8)

Dewol & Alxius 88399 (7)

Dewol & Karim 77865 (5)

Dewol & Termiji 78363 (13)

Dolman 21512 (5)

Dumas 1576 (3)

Elleh 34360 (14), 35596 (5)

Elmer 20194 (5), 20203 (14), 20839 (5), 21310 (2), 21337 (10), 21478 (14), 21622 (5)

Endert 1545 (5), 2596 (14), 2659 (4), 2931 (4), 2961 (15), 3195 (2), 3622 (2), 3866

(11), 4561 (15), 4970 (4), 4980 (14), 5192 (2)

Evans sn, Pianggu, Pahang (9)

Everett 13581 (8), 13586 (5), 13672 (8), 13695 (5), 13752 (5), 13962 (8), 14112 (5),

14272 (8), 14404 (5), 14456 (8), 14505 (5), 14559 (8)

Eyma 3399 (5), 4020 (5)

Flippance sn Penang on 27 Oct. 1932 (5)

Forbes 3219 (9)

Forest Dep.’t Borneo 577 (14), 1148 (14)

Forest Dep.’t Fed. Malay States 1141 (5), 1147 (17), 1155 (5), 1968 (5)

Forest Dep.’t Sarawak 2606 (6)

Forman 428 (2), 498 (14)

Fox 28 (5)

Foxworthy 1176 (5)

Franck 1129 (8)

Fuchs 21214 (7), 21297 (14)

Furtado sn Singapore on 19 Nov. 1929 (8)

Gadoh 880 (8), 1361 (8), 1769 (8)

Galau 15703 (12)

Gansau 46311 (14), 47803 (14)

Geesink & Santisuk 5362 (8)

Geh & Samsuri 708 (14)

Ghazalli & Asah 13662 (14)

Gibbs 3969 (15), 4342 (5)

Gibot 18595 (2), 30431 (14), 30467 (14), 30582 (7), 31294 (2), 31310 (2), 32474 (14),

32947 (2), 34125 (14), 60613 (14), 70493 (5)

Gilliland 5075 (8)

Goodenough =(Ridley’s) 352 (8), 1346 (8), 2010 (8), 10604 (8)

Griffith 2272 (8), 2272/1 (17), 2273 (5); sn, Malacca (5); sn, Malacca (8)

Hallier 1082 (14), 1308 (4), 1601 (3), 1630 (5), 1763 (2), 2049 (5), 2159 (5), 2223 (16),

2247 (16), 2311 (23217) (2), 2736 (14), 3113 (2), 3208 (14)

Hamid 4981 (5)

Haniff 367 (5), 14305 (5), 15527 (5)

Hardial 624 (8)

86 Gard. Bull. Sing. 34:1 (1981)

Haron 21395 (14), 21444 (1), 29993 (12)

Hartley 11913 (5)

Harun 17710 (5)

Haviland 68 (5), 68 (544) (13), 107 (3), 173 (3), (=174)B (4), 175 (12), 176 (14),

= 176 (14), =177 (5), 476 (13), 972 (14), =972B (6), 1383 (15), 1753 (6), 2019 (13),

2202 (5), 2203 (5), = 2203 (5), (=2203)Y (5), 2691 (2202) (5); sn, Kalaka, Sarawak

(5); sn, Kuching, Sarawak 14 April 1893 (12); sn, Kuching, Sarawak 17 May 1893,

23 May 1893 (5); Mantang, Sarawak (13); sn, Penkalan Ampat, Sarawak (4); sn,

Garai, Sarawak (6)

Haviland & Hose 3636 (5)

Henderson 1372 (8), 10012 (5), 10624 (5), 22130 (8), 24062 (5)

Henty, Ridsdale, Galore 31810 (5)

Hepburn 23722 (14)

Hervey 2010b (8)

Hill 1103 (8), 1135 (5)

Hochreutiner 761 (2)

Holmberg 863 (5)

Holttum 9672 (8), 10770 (5)

Hose 71 (13), 396 (7), 462 (14), 598 (13), 653 (14)

Hou 537 (5), 545 (5), 728 (8)

Hullett 320 (5), 661 (8)

Hume 7077 (8), 8873 (8), 9229 (5)

Hyland 3990 (5)

Ismail 95009 (5), 97845 (8), 104877 (8)

Jaamat 10925 (5), 16533 (5)

Jacobs 5302 (14), 5457 (4), 9531 (9)

Jaheri 1893 (16)

Jugah 23701 (7)

Jumali 4414 (5)

Kadim & Mohmud 59 (5)

Kadim & Noor 294 (5)

Kadir 939 (5), 1690 (5), 2726 (14), 3569 (5), 16867 (14)

Karim 80300 (5)

Katik 46796 (9)

Kasim 577 (8)

Keith 7123 (14), 7175 (14)

Keng & Jumali 522 (8), 3071 (K 242) (8), 4043 (8), 6295 (8)

Keng, Wee, & students 3 (8)

Kerr 6827 (5), 6827A (5), 7430 (8), 12270 (5), 12510 (5), 13538 (5), 14166 (5), 14575

(8), 15650 (5), 15654 (5), 15884 (8), 17869 (5)

Kiah 31912 (8), 32119 (5), 32429 (5), 37737 (5)

King sn, Penang Hill, Penang (5); sn, Singapore (8)

King’s collector 822 (5), 839 (5), 864 (5), 3518 (8), 3578 (8), 4856 (5)

Kinted 15889 (5)

Kloss 19027 (14); sn, Rawang, Selangor (8)

Kochummen 16327 (5), 023063 (17), 93466 (8), 97760 (5)

Korthals sn, Bandjermasin, Borneo (9); sn, G. Koerintgi, Sumatra (17); sn, G. Singgalang,

Sumatra (2); sn, Borneo (4)

Koster 1212 (aff. 9), 1481 (17), 6784 (5), 11178 (17)

Kostermans 4154 (14), 4238 (14), 4456 (9), 4709 (5), 4712 (9), 5083 (5), 5429 (5),

5535 (2), 5637 (5), 6440 (14), 6706 (5), 6868 (9), 7039 (2), 7507 (2), 8066 (9),

9021 (5), 9043 (2), 9128 (5), 9706 (9), 9916 (9), 10325 (10), 13664 (14), 21115

(14), 21540 (9), 21573 (14), 23820 (2)

Maxwell: Pternandra (Melastomataceae) 87

Kostermans & Anta 85 (18), 111 (9), 120 (S), 201 (9), 381 (5), 1001 (5), 1344 (5)

Krukoff 4021 (5)

Kunstler 87 (8)

Kuswata & Soepadmo 212 (5)

Lajangah 32197 (7), 33144 (14)

Lake & Kelsall 4073 (9)

Lakshnakara 346 (5), 737 (8), 744 (5)

Lambok 2723 (8)

Lampangi 24078 (2)

Lantoh 72326 (5), 73506 (4), 79946 (5), 83211 (2)

Lassan & Fox 72828 (14)

Lave sn, Borneo (5)

Ledermann 6536 (5), 7478 (5), 7788 (5), 8585 (9), 8717a (aff. 14); sn, New Guinea (5)

Leopold, Gary, Dewol 73815 (14)

Loh 13418 (8), 13466 (5)

Lowry 479 (5)

Luang 21736 (2), 21874 (2), 22383 (2), 22684 (11), 22821 (14), 23248 (11), 23263

(11), 23610 (12), 24141 (5), 24377 (14), 24454 (14), 24557 (7), 26131 (7), 27855

(14)

Luang & Nulong 24464 (13)

Luang & Paie 24737 (5)

Madani 33239 (7), 35095 (11), 35127 (14), 35246 (13)

Madius 49299 (7)

Mahamud 15506 (9)

Mahamet sn, Batu Tiga, Selangor (8)

Main 2162 (9)

Maingay 801 (1212) (5), 802 (1508) (5), 803 (1213) (8), 804 (3333) (8)

Mamit 29829 (4), 33396 (4)

Mat 7034 (5)

Matthew sn, Gunong Sengalan, Selangor (5)

Maxwell 74—425 (5), 75—854 (8), 76—757 (8), 77—22 (17), 77—23 (5), 77—138 (5),

78—260 (9), 81—13 (5)

McVeigh 8292 (5)

Meh 8989 (5), 9040 (5), 10159 (8), 10160 (5), 10182 (5), 12569 (5), 17863 (17),

21884 (8), 21891 (5)

Meijer 496 (14), 2148 (14), 2252 (2), 2366 (5), 2469 (5), 2537 (4), 4078 (5), 4311 (3),

6716 (3), 6799 (3), 19942 (5), 20488 (14), 20560 (5), 22904 (5), 23636 (5S), 37999

(14)

Meijer & Vermeulen 5401 (3)

Melegrito 2506 (5)

Mikil 28089 (7), 31450 (2), 31979 (13), 37737 (5), 37755 (13), 38496 (14), 38499 (17)

Moll 11601 and 11670 (aff. 9, aff. 17)

Morshidi 22892 (13), 24090 (12)

Motley 200 (9)

Moysey & Kiah 33638 (5), 33646 (17 or 18), 33834 (5)

Mujin 33573 (14), 33777 (14), 40583 (14)

Murdoch 263 (5), 366 (5)

Murton 92 (8)

Nahar 12700 (2)

Native collector 99 (5), 307 (distribution Manila) (5), 550 (distr. Manila) (5), 1139

(distr. Manila) (13)

Nauen 35879 (17)

88 Gard. Bull. Sing. 34:1 (1981)

Netherlands Indies Forest Service 5715 (5), 16214 (aff. 10), 21525 (5), 21607 (5),

24637 (aff. 9), 25715 (5), 27671 (5), 28613 (5), 28834 (5), 29231 (9), 29817 (5),

30001 (5), 30029 (5), 30089 (5), 31021 (5)

Ng 1822 (8), 5483 (5), 100014 (5), 118166 (8)

Ngadiman 16122 (8), 34652 (5), 34683 (5)

Noor sn, Singapore on 26 April 1921 (8)

Nooteboom 959 (14), 1071 (14), 1176 (11), 1328 (4), 1368 (11), 1474 (4)

Nooteboom & Chai 01831 (2), 02085 (2)

Nordin 85971 (7)

Nur 1235 (5), 11831 (8), 34132 (5); sn, Singapore 30 Sept. 1915 (5), sn, at Botanic

Garden on 24 April 1924 (8) and 19 Nov. 1929 (8)

Omar 68 (5)

Orolofo 18251 (5)

Paie 8452 (12), 8484 (13), 15091 (14), 15589 (9), 15836 (13), 17031 (5), 23054 (4),

25858 (11), 25862 (11), 26180 (5), 26926 (5), 27694 (12), 28069 (12), 28437 (2)

Paie & Ashton 16660 (5)

Panji 769 (8)

Patrick 42922 (5)

Patt sn, Penang Hill, Penang (5)

Phusomsaeng 58 (8)

Pleyte 704 (5)

Poore 1127 (18)

Pradit 232 (5)

Prayad 431 (5)

Price 257.(2)

Puasa 4183 (5), 7012 (14), 10071 (14)

Puasa-Angian 3988 (15)

Purseglove 4457 (4), 4895 (5), 5006 (5), 5043 (12), 5149 (13), 5536 (5), 5543 (5)

Purseglove & Shah 4429 (2)

Put 758 (5), 1696 (5)

Rabil 71 (5), 91 (5), 192 (5)

Rahim 444 (5)

Ramos 1191 (14)

Reksodihardjo 101 (5), 291 (8), 343 (8)

Richards 1108 (14), 1665 (14), 2414 (11), 2562 (4)

Ridley 279 (5), 352 (8), 1164 (5), 2004 (5), 2009b (5), 3918 (17), 4729 (17), 6217 (5),

7329 (5), 9021 (9), 9189 (8), 9685 (5), 11080 (5), 11936 (17), 12113 (5), 12167

(5), 12281 (12), 14685 (17 or 18), 14686 (5), 15527 (5); sn, Santulang, Borneo

Jan. 1918 (12); sn, Batu Pakat, Johore in 1900 (8); sn, Kota Tinggi, Johore Dec.

1892 (5); sn, Kwala Tehing Tinggi, Johore (9); sn, Kedah Peak, Kedak June 1893 (5);

Kuala Lebir, Kelantan Feb. 1917 (8); sn, Gunong Tampin, Negri Sembilan 16 Jan.

1917 (8); sn, Johol, Negri Sembilan 18 Jan. 1917 (8); sn, Tampai, Negri Sembilan

(5); sn, Gov.’t Hill, Penang 9 July 1890 (8); sn, Penang Hill, Penang Dec. 1895 (5);

sn, Penang Hill, Penang Jan. 1921 (5); Bidoi Tapah, Perak Nov. 1908 (17); sn, Din-

dings, Perak 12 March 1896 (5); sn, Pangkor, Perak (18); sn, Semangkok, Selangor

(8); sn, Tasek, Selangor (5); sn, Bukit Panjang, Singapore (8); sn, Chan Chu Kang,

Singapore (18); sn, Singapore 10 Feb. 1898 (5); sn, Singapore May 1905 (5); sn,

Singapore in 1911 (5); sn, Kriaui, Wellelsey June 1890 (5); sn, Kriaui, Wellelsey (5)

Ridsdale, Henty, Galore 31929 (5)

Robinson & Kloss 6148 (18)

Rutten 537 (14)

Sadau 42826 (14)

Saikeh 72099 (4)

Maxwell: Pternandra (Melastomataceae) 89

Sallih 9121 (5)

Sam 21797 (14), 34678 (5)

B. Sangkhachand 141 (5), 853 (8)

Santiago sn, Kepong, Selangor (8)

Sanusi 5327 (5)

Sauvour K5 (4), 133 (4)

Schram 1743 (5), 6007 (5), 12287 (9), 14915 (5)

Scortechini 43 (5), 248 (18), 1643 (8), 1894 (1824?) (8); sn, Perak (18)

Seimund 78 (5), 1142 (5); sn, Pulau Lalang, Perak (5)

Selvaraj 99671 (5)

Seuada 4104 (5)

Shah & Ahmad 3658 (8)

Shah, Ahmad, Noor 2057 (8)

Shah & Kadim 436 (5), 565 (8)

Shah & Shukor 2672 (8), 3192 (5)

Shah & Sidek 1159 (5)

Shea & Minjulu 76132 (5)

Shimizu et al. 12969 (5), 13160 (5)

Shing 13536 (5), 17057 (8), 17288 (5)

Shukor 104 (5)

Sibat see Luang

Sidek 232 (5)

Sinclair 10277 (12), 39033 (5), 39635 (5),

H. Singh 561 (5)

H. Singh & Samsuri 196 (5), 1028 (8), 1031 (5), 1079 (5)

H. Singh, Samsuri, Sidek 381 (5), 408 (5)

J. Singh 22369 (14), 22562 (14), 22883 (5), 24348 (5), 27494 (14), 30042 (10)

Siranggul 39909 (14), 51101 (14), 57191 (2)

Soepadmo 29 (9), 79 (5), 134 (5)

Seopadmo & Chai 28176 (2)

Soh 15089 (8)

Sohadi 14585 (8), 14715 (8), 14728 (5)

Sow & Tachon 16865 (5)

Spare 3775 (8), 1045 (8), 3775 (8), 3806 (5), 34458 (5), 34566 (5S)

Stone 6352 (5), 6854 (5), 9568 (5), 13177 (14), 13418 (5)

Stone & Anderson 86751 (14), 86804 (14)

Stone, Mahmud, Sharif 8539 (5)

Streimann & Kairo 26184 (17)

Strugnell 11226 (5)

Sundaling 80757 (9), 80808 (5)

Suppiah 11297 (8), 11430 (8), 14815 (5), 14820 (5), 11933 (8), 108979 (8), 108872

(5), 108954 (5), 108979 (5)

Symington 20875 (5), 37382 (5)

Symington & Kiah 28774 (5)

T. & P. (Teo & Purseglove) 76 (2676) (5), 211 (2811) (5), 648(3248)(8)

Taloon 10767 (14)

Tandom 4117 (5), 8805 (14)

Tangulon 1565 (14)

Tarmiji & Dewol 84192 (14)

Teruya 409 (5)

Teysmann 3396 (5), 8165 (16), 8655 (3)

Tikau 33732 (5)

Tong 34998 (14), 35020 (7)

90 Gard. Bull. Sing. 34:1 (1981)

Toroes 1509 (5), 2139 (5), 3835 (5)

Valera 1870 (5)

van Niel 4621 (6)

van Royen 4006 (aff. 9)

van Steenis 11193 (2)

Versteegh 4900 (5), 4969 (5)

Villamil 147 (14)

de Voogd 1143 (2)

Waitz sn, Borneo (aff. 8); sn, G. Rantau, Borneo (2)

Walker 304 (8)

Wallich 3143 (5), 4077 (5), 4078 (8), 4078a (8), 4079 (5), 4080 (5)

Walsh 443 (14), 715 (2), 748 (9)

Whitmore 0975 (5), 3153 (8), 3675 (5), 3942 (5), 4109 (5), 4788 (8), 8884 (5), 15366

(5), 15547 (8), 20021 (5), 20183 (5)

Wiakabu & Simaga 70214 (5)

v. Wijk 49 (9)

de Wilde & de Wilde-Duyfjes 12829 (3), 13971 (5), 15574 (5)

Wing 33963 (14), 34921 (14), 38987 (14)

Winkler 2249 (5), 2321 (2), 2408 (5)

Wood 16459 (aff. 14), 16663 (2)

Woods 145 (8)

Wray 1952 (5), 1994 (17), 2156 (5), 2354 (8), 2390 (5), 2872 (5), 3254 (5), 3607 (5),

3975 (5); sn, Ayer Larut, Perak (18)

Wright 24713 (7), 27105 (18), 27967 (7), 29129 (14), 29135 (13)

Wright & Othman 32303 (7)

Yakup 11209 (9)

Yeob 0842 (5)

sine coll. 46 (Chumpon, Thailand) (5)

A REVISION OF THE GENUS TETRADIUM (RUTACEAE)

T. G. HARTLEY

Herbarium Australiense

C.S.I.R.O. Division of Plant Industry

Canberra, Australia 2601

ABSTRACT

The genus Tetradium, which has traditionally been placed in the synonymy of the

rutaceous genus Euodia, consists of nine species and is distributed from the Himalayas

east to Japan and south to Java and Sumbawa. The genus is described and its taxonomic

history and relationships to other genera are discussed. The nine species are keyed,

described, and their interrelationships discussed. New combinations are made for the

names of seven species, and one species, Tetradium sumatranum, is described as new.

Among the Rutaceae of the southeast Asian-Pacific region is a group of trees and

shrubs that is distinctive in having opposite, pinnately compound leaves in combination

with terminal inflorescences, follicular fruits, and seeds that remain attached in the de-

hisced follicle. These plants were first described in 1790 when Loureiro based the genus

Tetradium on a Vietnamese species, Tetradium trichotomum Lour. Although Loureiro’s

description of this plant was reasonably adequate and a representative specimen was

lodged at the British Museum, the identity of Tetradium in the Rutaceae was not

definitely established until the publication of the first volume of Bentham and Hooker’s

Genera Plantarum (1862). In the meantime, Sprengel (1824) transferred Tetradium

trichotomum to the simaroubaceous genus Brucea J. F. Mill., DeCandolle (1825)

recognised Tetradium as a distinct genus and placed it between Brucea and Ailanthus

Desf., while Jussieu (1825) and Endlicher (1840) listed it in the Rutaceae as an insuffi-

ciently known genus. Also during this period, three additional genera, Philagonia Bl.,

from Java, Boymia A. Juss., from mainland China, and Megabotrya Hance ex Walpers,

from Hong Kong, were based on plants that are congeneric with Tetradium trichotomum.

Bentham and Hooker placed Tetradium, along with Philagonia, Boymia, and

Megabotrya, in the synonymy of the genus Euodia J. R. & G. Forst. They were essentially

followed in this by Engler (1896, 1931), in the standard major work on the southeast

Asian-Pacific Rutaceae, who treated Tetradium as a section of Euodia and listed Phila-

gonia, Boymia, and Megabotrya in its synonymy. This is a remarkably conservative treat-

ment, considering the morphologic differences between the type species of Euodia,

Euodia hortensis J. R. & G. Forst., from the New Hebrides, and the plants typifying

Tetradium, Philagonia, Boymia, and Megabotrya, but its correctness has apparently never

been questioned.

In connection with the present study I have examined type or otherwise authentic

specimens of nearly all of the 200 or so species that have been placed in Euodia sensu

Engler, and it is evident that those with opposite, pinnately compound leaves and

follicular fruits with persistent seeds form a natural group around the type species of

Tetradium and stand well apart, morphologically, from a group of species centered

around the type species of Euodia. It is also evident that the majority of the species that

have been placed in Euodia in this classification are more correctly placed in the genus

Melicope J. R. & G. Forst. than in either Tetradium or Euodia.

9) Gard. Bull. Sing. 34:1 (1981)

In view of this, the formulation of a more natural classification of these species

reinstating Tetradium (including Philagonia, Boymia, and Megabotrya) at the rank of

genus and redefining Euodia and Melicope. These three genera are thus characterized as

follows:

Tetradium. Leaves opposite, pinnately compound; inflorescences terminal or ter-

minal and from the axils of the uppermost pair of leaves; flowers unisexual, 4—5-merous;

stamens the same number as petals; carpels 1- or 2-ovulate; fruits follicular; seeds shiny,

smooth, remaining attached in the dehisced follicle. Nine species; distributed from the

Himalayas east to Japan and south to Java and Sumbawa.

Euodia. Leaves opposite, trifoliolate or unifoliolate; inflorescences axillary; flowers

bisexual, 4-merous; stamens the same number as petals; carpels 2-ovulate; fruits follicular;

seeds more or less dull and roughened, discharged when the follicle dehisces. About six

species; distributed from New Guinea and northeastern Australia east to Samoa.

- Melicope. Leaves opposite, trifoliolate, unifoliolate, or simple; inflorescences axillary,

lateral, or rarely (only in a few species from Madagascar) terminal; flowers bisexual or

unisexual, 4-merous; stamens the same number or twice as many as the petals; carpels

2-ovulate; fruits follicular to capsular; seeds shiny, smooth, remaining attached in the

dehisced fruit. Probably about 150 species; distributed from Madagascar to India and

southern China, throughout Malesia, Micronesia, Melanesia, and Polynesia to the Society

Islands, and south to southeastern Australia and New Zealand.

Of the species that are considered here to belong in the genus Melicope, the flowers

of those that were misplaced in Euodia are 4-staminate whereas those of the type species

of Melicope, Melicope ternata J. R. & G. Forst., from New Zealand, and the 50 or so

species that center around it, morphologically, are 8-staminate. This is the only consistent

difference between these plants and it is clearly not a sign of generic distinctness, as has

traditionally been thought, because it cuts across many lines of obvious relationships

between species.

The main differences between Tetradium, Euodia, and Melicope, as these genera

are interpreted here, are given in the following key:

Seeds more or less dull and roughened, discharged when the follicle dehisces; leaves

trifoliolate or unifoliolate; inflorescences axillary................... Euodia

Seeds shiny, smooth, remaining attached in the dehisced fruit.

Leaves pinnately compound; inflorescences terminal or terminal and from the

axils of the uppermost pair of leaves:°..°>5 bn MVR eee Tetradium

Leaves trifoliolate, unifoliolate, or simple; inflorescences axillary, lateral, or

rarely terminal). . 20 Ath, Pe A ee eee Melicope

(Euodia and Melicope are being revised. The necessary nomenclatural changes for

species of Melicope that were originally described in Euodia will be made in a future

publication.)

Besides the distinguishing characters noted above, Tetradium differs from Euodia and

Hartley: Tetradium (Rutaceae) 93

Melicope in having strong sexual dimorphism in the gynoecium. In its carpellate flowers

the carpels are contiguous, forming a 4- or 5-lobed ovary, whereas in its staminate flowers

the rudimentary carpels are divergent, finger-like structures which are about as long as

the functional carpels. There is no similar dimorphism in Euodia or in Melicope. The

flowers of the former are always bisexual and, in species of the latter with unisexual

flowers, the non-functional gynoecium is merely a reduced replica of the functional

gynoecium.

Strikingly similar sexual dimorphism to that found in Tetradium is exhibited by the

genus Phellodendron Rupr. (Rutaceae), which consists of about eight species and is

distributed from southwestern China northeast to southeastern U.S.S.R. and Japan and

south, discontinuously, to Taiwan. Phellodendron differs rather markedly from Tetra-

dium in having a syncarpous functional gynoecium that develops into a drupaceous fruit.

Its rudimentary carpels are distinct, however, and otherwise appear to be identical with

those produced in Tetradium. In other features, as well, these two genera are so alike that

were it not for a difference in the position of the axillary bud — which is sunken in the

base of the petiole in Phellodendron and exposed in the leaf axil in Tetradium — it

would seem impossible to distinguish one from the other in sterile or staminate herba-

rium material.

While it may seem unlikely that a genus with syncarpous, drupaceous fruit would be

a close relative of one with apocarpous or subapocarpous, follicular fruit — and indeed,

on the basis of this fruit difference Engler (1896, 1931) placed Phellodendron and

Tetradium (as a section of Ewodia) in separate subfamilies of the Rutaceae, the Todda-

lioideae and Rutoideae, respectively — I think there is little doubt that Phellodendron

and Tetradium are, in fact, closely related. Such a relationship is not unusual in the non-

aurantioid Rutaceae. Acronychia J. R. & G, Forst., for example, which has syncarpous,

drupaceous fruit, appears to be closely related to Melicope, most species of which have

apocarpous or subapocarpous, follicular fruits (Hartley, 1974), and Zanthoxylum L.

(including Fagara L.), which has apocarpous or subapocarpous, follicular fruit, appears to

be the closest relative of Toddalia Juss., which has syncarpous, drupaceous fruit (Hartley,

unpubl.). A particularly close evolutionary proximity of the apocarpous and syncarpous

conditions appears to be evidenced in Phellodendron where the presence of separate

rudimentary carpels in a genus with a syncarpous functional gynoecium would seem to

indicate immediate apocarpous ancestry. I think it is likely that this ancestor was a

species of Tetradium.

Tetradium also appears to be closely related to Zanthoxylum, which consists of

about 200 species and is pantropical and ranges north into temperate latitudes of eastern

Asia and North America. Zanthoxylum differs from Tetradium mainly in having alternate

leaves. The two genera are consistently similar in having apocarpous or subapocarpous,

follicular fruits with smooth, shiny seeds that remain attached in the dehisced follicle. In

most other relevant characters Zanthoxylum is variable, but many of its species, includ-

ing several from southeast Asia (see Huang, 1957, and Hartley, 1966) are similar to

Tetradium in having pinnately compound leaves, terminal inflorescences, unisexual,

4- or 5-merous flowers, and divergent, finger-like, rudimentary carpels, among other

characters.

Zanthoxylum, Tetradium, and Phellodendron appear to be related to one another in

a linear sequence. The following outline shows the apparent interrelationships of the three

genera as indicated by various morphologic features.

94 Gard. Bull. Sing. 34:1 (1981)

Leaves alternate; functional gynoecium apocarpous or subapocarpous (or sometimes

reduced to a single carpel); fruits follicular. ...... 01. Ss ayes Zanthoxylum

Leaves opposite; functional gynoecium apocarpous or subapocarpous; fruits follicu-

RAE cine np0id ute uw, 050, 0 0.9 kha. 9 Ske ns, yo (el ene eee Pee: Tetradium

Leaves opposite; functional gynoecium syncarpous; fruits drupaceous. ..........

Pee ee OR a i OOS et Phellodendron

Toddalia, which is mentioned above as probably being a close relative of Zanthoxy-

lum, does not seem to be part of this sequence.

This revision is based on herbarium specimens. The contributing herbaria, with abbre-

viations from Holmgren and Keuken’s Index Herbariorum, Part 1, ed. 6 [Reg. Veg. 92

(1974)], are as follows: Arnold Arboretum of Harvard University, Cambridge (A); State

Herbarium of South Australia, Adelaide (AD); Auckland Institute and Museum, Auckland

(AK); Botanisches Museum, Berlin-Dahlem (B); Bernice P. Bishop Museum, Honolulu

(BISH); British Museum (Natural Histroy), London (BM); Herbarium Bogoriense, Bogor

(BO); Queensland Herbarium, Brisbane (BRI); Botanical Survey of India, Calcutta (CAL);

C.S.I.R.O. Herbarium Australiense, Canberra (CANB); Botany Division, D.S.I.R., Crist-

church (CHR); Forest Research Institute and Colleges, Dehra Dun (DD); Royal Botanic

Garden, Edinburgh (E); Conservatoire et Jardin Botaniques, Geneve (G): Gray Herbarium

of Harvard University, Cambridge (GH); Royal Botanic Gardens, Kew (K); Rijksher-

barium, Leiden (L); Komarov Botanical Institute, U.S.S.R. Academy of Sciences, Lenin-

grad (LE); National Herbarium of Victoria, Melbourne (MEL); University Herbarium,

University of Michigan, Ann Arbor (MICH); National Herbarium of New South Wales,

Sydney (NSW); New York Botanical Garden, New York (NY); Museum National

d’Histoire Naturelle, Paris (P); Institute of Botany, Academia Sinica, Peking (PE); Depart-

ment of Botany, Academy of Natural Sciences, Philadelphia (PH); Botanic Gardens,

Singapore (SING); Department of Botany, University of Tokyo (TI); Botanical Museum

and Herbarium, Utrecht (U); Herbarium of the University of California, Berkeley (UC);

National Museum of Natural History (Department of Botany), Smithsonian Institution,

Washington D. C. (US); Naturhistorisches Museum, Wien (W).

Tetradium Lour., Fl. Cochinch. (1790) 91, non Dulac, 1867. Type species: Tetradium

trichotomum Lour.

Philagonia Bl., Cat. Pl. Buitenz. (1823) 20. Bijdr. (1825) 250. Type species:

Philagonia sambucina Bl.

Boymia A. Juss., Mem. Mus. Hist. Nat. 12 (1825) 507,, Mem. Rutac. (1825) 124.

Type species: Boymia rutaecarpa A. Juss.

Megabotrya Hance ex Walpers, Ann. Bot. Syst. 2 (1852) 258. Type species: Mega-

botrya meliaefolia Hance ex Walpers.

Xanthoxylum subgenus Oxyactis Benn., Ann. Mag. Nat Hist. ser. 3, 10 (1862) 201.

Type species: Xanthoxylum daniellii Benn.

Evodia section Tetradium Lour., Engl. in Engler & Prantl, Nat. Pflanzenfam. III: 4

(1896) 121; ibid.ed.2, 19a (1931) 228. Type species: Evodia trichotomum

(Lour.) Pierre.

Hartley: Tetradium (Rutaceae) 95

Evodia section Evodioceras Dode, Bull. Soc. Bot. France (1908) 55, (1909) 706.

Type species: Evodia daniellii (Benn.) Hemsl.

Evodia section Oxyactis (Benn.) Rehd. & Wils. in Sargent, Pl. Wils. 2 (1914) 133,

nomen illegit., based on the same type as Evodia section Evodioceras Dode.

Evodia section Subtrigonospermum Huang, Acta Phytotax. Sin. 6 (1957) 116. Type

species: Evodia subtrigonosperma Huang.

Small to large trees or occasionally shrubs; dioecious or rarely occasional plants

monoecious; evergreen or deciduous; indumentum of simple trichomes. Buds naked,

pubescent, the lateral buds axillary. Leaves opposite, imparipinnate or rarely occasional

leaves paripinnate; lateral leaflets mostly petiolulate; terminal leaflet on an extension of

the rachis; leaflet blades usually oil dotted, venation pinnate, margin often crenulate with

an oil dot at the sinus of each crenulation. Inflorescences corymbose or subcorymbose or

occasionally grading to paniculate, spreading or rarely rather compacted, several- to

many-flowered, terminal or terminal and from the axils of the uppermost pair of leaves.

Flowers unisexual or rarely occasional flowers bisexual, 5- or 4-merous, globose, broadly

ellipsoid, or obovoid in bud; sepals 5 or 4, ovate to triangular, basally connate, valvate,

persistent in fruit; petals 5 or 4, elliptic to ovate-elliptic, usually hooked adaxially at the

apex, distinct, erect, narrowly imbricate in bud, deciduous in fruit; stamens the same

number as petals, distinct, opposite the sepals; functional stamens elongating to about 1.5

times the length of the petals, filaments usually more or less villous from about the middle

to the base, adaxially, otherwise glabrous, linear-subulate, curved inward basally, other-

wise straight and more or less erect, anthers ovoid, obtuse, dorsifixed; rudimentary sta-

mens (produced in carpellate flowers) much shorter than the petals or sometimes ob-

solete, ligulate, persistent in fruit; disc intrastaminal, in staminate flowers conical to

cylindrical or occasionally barrel-shaped, in carpellate flowers pulvinate to barrel-shaped,

persistent in fruit as a short stipe; gynoecium 5- or 4-carpellate, together with the sub-

tending disc about as long as the petals; functional carpels free to the base or connate

toward the base, otherwise contiguous and forming a 5- or 4-lobed, 5- or 4-loculate,

subglobose to obovoid ovary, placentation axile, ovules 2 or 1 per locule, style apical,

straight, composed of 5 or 4 more or less contiguous stylar elements; stigma 1, peltate

and usually shallowly 5- or 4-lobed; rudimentary carpels (produced in staminate flowers)

finger-like, free to the base or connate toward the base, divergent and sometimes con-

torted. Fruits of 1-5 or 14 1- or 2-seeded follicles, carpels not developed into follicles,

persistent; follicles free to the base or connate toward the base, dehiscing adaxially,

apically, and partially down the abaxial surface, epicarp dry at maturity, subwoody, the

outer surface more or less glandular-punctate, endocarp cartilaginous abaxially grading to

pergamentaceous adaxially, persistent in the dehisced follicle. Seed(s) shiny, smooth,

black to dark reddish brown, persistent in the dehisced follicle; endosperm copious,

fleshy; embryo straight, cotyledons broadly elliptic, plano-convex, hypocotyl terminal.

Trichomes are uniformly simple and unicellular in Tetradium and in all species the

new growth of vegetative and floral shoots is clothed with fine, short indumentum. As

these shoots mature, this juvenile vestiture may be partially or entirely lost or may be

retained and is often further developed. As a result, mature leaves, branchlets of the

present year’s growth, and axes and branches of inflorescences vary from glabrous to

densely pubescent. For the most part this variability is taxonomically insignificant.

A rather unusual surface feature occurs in Tetradium austrosinense and T. suma-

tranum, and to a lesser extent in a few specimens of T. fraxinifolium. The lower surface

96 Gard. Bull. Sing. 34:1 (1981)

of the leaflets of these plants is glaucous and minutely papillate. The papillae appear to

be globules of wax.

The terminal inflorescence in Tetradium originates from the middle bud of the three

found between the uppermost pair of leaves of the shoot. The two lateral buds of the

three may form upper axillary inflorescences or they may remain dormant during the

flowering period and later form vegetative shoots.

Inflorescences range from corymbose to subcorymbose to paniculate and from

spreading to rather compacted. This variability is not taxonomically significant.

Tetradium is remarkably variable (for a rutaceous genus) in ovule number and in

several characteristics of the seeds and their mode of attachment in the dehisced follicle.

This variability, summarized in the following outline, provides a basis for a natural classi-

fication of the species.

A. Carpels 2-ovulate.

B. Seeds 2 per follicle (follicles appearing to be 1-seeded in T. sambucinum).

C. Both seeds functional; outer testa spongy, bounded externally by a shiny,

crustaceous pellicle; inner testa bony.

D. Seeds hemispheric, the two coherent and appearing to represent a

single, globose or subglobose seed, attached in the dehisced follicle to

an adaxial strip of pergamentaceous endocarp tissue.

1. ZT. sambucinum.

D. Seeds subtrigonous, separate, attached in the dehisced follicle to a

funicular aril.

2. T. fraxinifolium.

C. One seed sterile (of almost normal size — containing what appears to be

endosperm but lacking an embryo); seeds ovoid to ellipsoid, coherent,

attached in the dehisced follicle to an adaxial strip of pergamentaceous

endocarp tissue; testa bony in-the fertile seed, thin and brittle in the sterile

seed, bounded externally by a shiny pellicle, without spongy tissue.

3. T. daniellii.

4. T. calcicolum.

B. Seed 1 per follicle (paired with an aborted seed). subglobose to ovoid to broadly

ellipsoid, attached in the dehisced follicle to an adaxial strip of pergamentaceous

endocarp tissue; outer testa spongy, bounded externally by a shiny, crustaceous

pellicle; inner testa bony.

5. T. glabrifolium.

6. T. trichotomum.

7. T. ruticarpum.

A. Carpels 1-ovulate.

Hartley: Tetradium (Rutaceae) 97

E. Seed ellipsoid to subreniform, attached in the dehisced follicle to

an adaxial strip of pergamentaceous endocarp tissue; outer testa

spongy, bounded externally by a shiny, crustaceous pellicle; inner

testa bony.

8. T. austrosinense.

E. Fruits and seeds not known.

9. T. sumatranum.

Regarding the relative evolutionary levels of these characters, it is probable that the

one-ovulate condition is more highly derived than the two-ovulate and that in the two-

ovulate species the one-seeded condition is more highly derived than the two-seeded.

The levels of the other characters are more difficult to judge, but it seems likely, mainly

because of dissimilarity to Zanthoxylum, the supposed ancestor of Tetradium, that

the arillate condition in 7. fraxinifolium and the absence of spongy outer testa in

T. daniellii and T. calcicolum represent derived states from the non-arillate condition

and presence of spongy outer testa.

Regarding the interrelationships of the species, Tetradium daniellii and T. calcicolum

are very closely related and stand well apart from the other species of the genus. The

same is true of 7. austrosinense and T. sumatranum. Tetradium glabrifolium, T. tricho-

tomum, and T. ruticarpum are less closely interrelated, but clearly comprise a natural,

fairly isolated group. The remaining two species, 7. sambucinum and T. fraxinifolium,

are not at all closely related and are markedly distinct (especially the latter) from the

other species of the genus.

As is noted in the generic synonymy, three sections of Euodia have been based

on species of Tetradium, namely, section Tetradium, based on Euodia trichotoma

(= Tetradium trichotomum); section Evodioceras, based on Euodia daniellii (= Tetradium

daniellii); and section Subtrigonospermum, based on Euodia subtrigonosperma

(= Tetradium fraxinifolium ). In view of the interrelationships of the species, these

sectional names could be used to designate natural species groupings: section Tetradium

for T. glabrifolium, T. trichotomum, and T. ruticarpum; section Evodioceras for T.

daniellii and T. calcicolum; and section Subtrigonospermum for T. fraxinifolium. |

prefer not to use them, however, because of the small size of the genus and also because

doing so would necessitate describing two additional sections, one for 7. sambucinum

and one for T. austrosinense and T. sumatranum.

Economically, Tetradium is of rather minor importance as an ornamental tree

(notably T. daniellii, which is hardy at cold temperate latitudes) while the fruit of one

species, T. ruticarpum, is reported (as Boymia ruticarpa and Euodia officinalis) to have

medicinal properties (Siebold and Zuccarini, 1837; Rehder and Wilson, 1914).

Chemically, Tetradium is reported to yield a variety of secondary metabolites, and

from this standpoint the genus may have as yet unrealized economic value. These reports

are as follows:

1. The isoquinoline alkaloid berberine is reported from Tetradium glabrifolium (as

Euodia glauca and EF. meliifolia) by Price (1963) and Hegnauer (1973).

98 Gard. Bull. Sing. 34:1 (1981)

2. Several quinazoline alkaloids are reported from Tetradium ruticarpum (as Euodia

ruticarpa) by Price (1963), Hegnauer (1973), and Waterman (1975).

3. An amide is reported from Tetradium daniellii (as Euodia hupehensis) by Waterman

(1975).

4. The flavanoids diosmin and hesperidin are reported from Tetradium daniellii (as

Euodia daniellii, E. hupehensis, and E. velutina) and T. ruticarpum (as Euodia officinalis

and £. ruticarpum) by Hegnauer (1973).

5. Three limonoids are reported from Tetrodium ruticarpum (as Euodia ruticarpa) by

Hegnauer (1973).

6. Two triterpenes are reported from Tetradium fraxinifolium (as Euodia fraxinifolia)

by Hegnauer (1973).

The reports of the isoquinoline alkaloid berberine from Tetradium glabrifolium were

based on earlier studies and are considered to be doubtful by Hegnauer (1973) and

Waterman (1975); both of these authors point out that isoquinoline alkaloids, which are

of wide occurrence in families of the Ranales sensu lat., are probably restricted, in the

Rutaceae, to Zanthoxylum (including Fagara), Phellodendron, and Toddalia. The close

mutual relationship between Zanthoxylum, Tetradium, and Phellodendron, proposed in

the present study, suggests that species of Tetradium may well synthesize these com-

pounds. It would thus be of interest to re-examine material of T. glabrifolium.

KEY TO FLOWERING MATERIAL

1. Perianth, androecium, and gynoecium predominantly 5-parted (4-parted in occa-

sional flowers)... <5. cs uc aveuleid we sym idea ee Boe sane ge ia ee 2.

2. Leaflets minutely papillate below; ovules 1 per carpel................. Et

3. Leaflets below sparsely to densely pubescent with spreading or ascending

hairs, main veins 9-12 pairs; petals 2.5-3 mmlong. ... 8. T. austrosinense.

3. Leaflets below puberulent with minute, appressed hairs, main veins 13-16

pairs; petals 32°35 Gaim. Jone. n.\.20. cn seein on ee 9. T. sumatranum.

2. Leaflets not papillate; ovules 2 per carpel. °/.. 7... 2 2, ee 4.

4. Petals purple, drying dull purplish red, sparsely to rather densely

appressed pubescent abaxialiy”. .) as «. ee 4. T. calcicolum.

4. Petals green to yellow to white, drying brown or whitish, glabrous or

nearly so abaxially. oo... ss us, chejovd apes sual co tay ee >

5. Lower surface of leaflets usually conspicuously (10x) oil-dotted,

usually pubescent over the entire surface; ovary anc rudimentary

carpels usually glabrous; 2) 7. T. ruticarpum.

5. Lower surface of leaflets usually inconspicuously oil-dotted,

usually entirely glabrous or with pubescence restricted to the

Hartley: Tetradium (Rutaceae) 99

midrib toward the base (rarely pubescent over the entire surface);

ovary pubescent between the carpels and often on the exposed

surface; rudimentary carpels usually pubescent, at least toward the

eee regis. oO. onc Sesit Sisnse sbive gad... 2... 6.

Leaflets below usually obviously glaucous (dried material),

ultimate veinlets clearly visible (10x), densely reticulate;

sepals about 0.5 mm long; ovules collateral or subcollateral;

rudimentary carpels usually pubescent toward the base and

glabrous toward the apex (rarely entirely glabrous). .......

SUE AE Me Eg Se che ee Se 5. T. glabrifolium.

Leaflets below seldom obviously glaucous, ultimate veinlets

usually inconspicuous; sepals 0.5-1.5 mm long; ovules super-

posed; rudimentary carpels uniformly villous. . 3. 7. daniellii.

1. Perianth, androecium, and gynoecium predominantly 4-parted (S-parted in occa-

sional flowers)

se aS, See ee) ee el oe oe eT. e. ae” Soe a Ya ee Se ies ot he Oe “ais “ae ‘oe 6 ©: 0 “ee 6-8 ©

7. Petals 2.5-3 mm long; disc sparsely to rather densely pube-

scent; main veins of leaflets departing from the midrib at

an angle of 45-50 degrees. ........ 1. 7. sambucinum.

7. Petals 3-6.5 mm long; disc glabrous; main veins of leaflets

departing from the midrib at an angle of 65-80 degrees.

8. Leaflets with large oil dots at the margin and smaller

dots scattered elsewhere on the blade, margin crenu-

late or rarely entire, main veins 13-22 pairs.......

PAA ee es op Peis h a ER Ak 2. T. fraxinifolium.

8. Leaflets with oil dots all of approximately the same

size, margin entire, main veins 11-14 pairs. .......

ee eae le ie o 6. T. trichotomum.

KEY 1O FRUITING MATERIAL

1. Follicles 2-seeded (appearing to be 1-seeded in T: sambucinum). ............ 2

2. Seeds 4-4.5 mm long, separate, firmly attached in the dehisced follicle to a

ERIC RS SPORE IVINS hee eee AR ty 205 WN ei 2. T. fraxinifolium.

2. Seeds 1.5-4 mm long, coherent, loosely attached in the dehisced follicle to an

adaxial strip of pergamentaceous endocarp tissue.................0-: 3

3. Fruits 4-carpellate,’ all four carpels usually developing into follicles;

follicles 3-4 mm long and about as wide; seeds collateral, hemispheric, the

two together appearing to represent a single globose or subglobose seed.

nn ete ek ACS okey) Tri. pak ee ches « vocaia 1. 7. sambucinum.

1. In all species of Tetradium the carpel number can be determined in fruiting material since carpels

that do not develop into follicles are persistent.

100 Gard. Bull. Sing. 34:1 (1981)

3. Fruits predominantly 5-carpellate (occasional fruits 4-carpellate), 2-5 of

the carpels developing into follicles; follicles 5-11 mm long, definitely longer

than wide; seeds superposed, ovoid to ellipsoid. ................. 4

4. Follicles beaked, sparsely to rather densely pubescent with spreading

hairs, 5-11 mmilongy!s:2.>. 2 tel oe earls ee 3. T. daniellii.

4. Follicles not beaked, sparsely to densely appressed-pubescent, about

5.mm long. - 0) .632)s issna ls ee eee ee 4. T. calcicolum.

1. Follicles l-seeded. . .. 0. 0000 c een ne cee ee hn J

5. Follicles about 3.5 mm long, developed from 1-ovulate carpels

and thus not containing an aborted seed; leaflets minutely papillate

below) G5 2.5) ae 8. T. austrosinense.”

5. Follicles 3.5-7 mm long, developed from 2-ovulate carpels and thus

containing an aborted seed (usually flattened against the upper

adaxial surface of the developed seed); leaflets not papillate.... 6.

6. Fruits predominantly 4-carpellate (occasional fruits 5-carpel-

late), each composed of 1-4(-5) follicles... .... 7. 2. eee

wole SRT S RPA oy eee hoe te Ceres 6. T. trichotomum.

6. Fruits predominantly 5-carpellate (occasional fruits 4-carpel-

late), each composed of 1-5 follicles, .. 0. s..cyee See fs

7. Follicles densely appressed-pubescent laterally, glabrous

abaxially; endocarp sparsely to rather densely pubescent;

leaflets usually glabrous or nearly so, ultimate veinlets

clearly visible (10x) on the lower surface, densely reticu-

late’ sector 4 Ae Se RR ee ok 5. T. glabrifolium.

7. Follicles glabrous or occasionally with sparse hairs late-

rally and/or abaxially; endocarp glabrous; leaflets sparsely

to densely pubescent below, ultimate veinlets usually

inconspicuous.) Sys Res -. eeas 7. T. ruticarpum.

1. Tetradium sambucinum (BI.) Hartley, comb. nov.

Philagonia sambucina Bl., Cat. Pl. Buitenz. (1823) 21; Bijdr. (1825) 250. Lectotype

(chosen here): Blume, October 1822 (carpellate flowering material), Java,

Salak.

Philagonia procera Bl. ex Nees, Flora 8 (1825) 125, nomen illegit., based on Phila-

gonia sambucina Bl.

2. Tetradium sumatranum, for which fruits are not known, is closely related to 7. austrosinense

(both species have 1-ovulate carpels and minutely papillate leaflets). Vegetative differences

between the two are given in the key to flowering material.

Hartley: Tetradium (Rutaceae) 101

Evodia sambucina (Bl.) Hook. f. ex Koorders & Valeton, Meded. Lands Plantentuin

17 (1896) 216.

Philagonia sambucifolia Dode, Bull. Soc. Bot. France (1908) 55, (1909) 701, sphalm.,

= Philagonia sambucina Bl.

Evodia sambucifolia Dode, ibid., sphalm., = Philagonia sambucina Bl.

Medium to large tree to 34 m high; branchlets puberulent to finely pubescent when

young, becoming glabrous or nearly so. Leaves 17-36 cm long; rachis puberulent,

especially adaxially; leaflets (3-)4-5(-6) pairs; petiolules of lateral leaflets obsolete to 10

mm long, terminal leaflet on an extension of the rachis 10-20 mm long; blades of leaflets

chartaceous to subcoriaceous, inconspicuously oil-dotted, below drying pale green to

brown, usually with appressed puberulence or short spreading hairs on the midrib and

main veins, otherwise glabrous or nearly so; above usually with appressed puberulence or

short spreading hairs on the midrib, otherwise glabrous or nearly so; elliptic-oblong or

less often elliptic, ovate, lanceolate, or oblanceolate, 6-18 cm long, 2-5 cm wide, base

in lateral leaflets acute to obtuse, usually rather oblique, in terminal leaflet cuneate, main

veins 13-16 pairs, departing from the midrib at an angle of 45-50 degrees, ultimate vein-

lets usually rather conspicuous (10x), rather loosely reticulate, margin crenulate, at least

toward the apex, inconspicuous oil dots associated with the crenulations, apex acuminate,

the acumen 10-20 mm long. Inflorescences 13-25 cm long, rachis puberulent, branches

puberulent to finely pubescent, pedicels puberulent to finely pubescent, obsolete to 1.5

mm long. Flowers 4-merous; sepals puberulent to glabrous, usually ciliolate, 0.5-0.8 mm

long; petals yellowish green, drying brown, abaxially glabrous or rarely sparsely puberu-

lent, adaxially sparsely to rather densely villous, 2.5-3 mm long; disc sparsely to rather

densely pubescent; ovary appressed pubescent, carpels free to the base abaxially, connate

toward the base adaxially, ovules 2 per carpel, collateral; rudimentary carpels more or

less villous toward the base, otherwise glabrous, free to the base. Fruits 4-carpellate, all

four carpels usually developing into follicles; follicles puberulent, trigonous, not beaked,

3-4 mm high and about as wide, free to the base abaxially, connate toward the base

adaxially, endocarp sparsely hairy. Seeds black, 2 per follicle, collateral, hemispheric,

coherent and appearing to represent a single globose or subglobose seed, 1.5-2 mm long,

attached in the dehisced follicle to an adaxial strip of pergamentaceous endocarp tissue;

outer testa spongy (bounded externally by a crustaceous, shiny pellicle); inner testa bony.

Distribution. W. Malaysia, Sumatra, Java, and Sumbawa; recorded from primary

and secondary, usually poorly-drained, rain forests, 30-1400 m.

W. Malaysia. Trengganu: G. Mandi Angin, Whitmore FRI 12139 (L).

Sumatra. Res. Atjeh: Simaloer Island, Achmad 607 (BO), 775 (BO, L), 1022 (BO,

L), 1142 (BO, L, SING), 1662 (BO, L), 1736 (BO, L); Gajo Loeeus, Penosan (G. Geroe-

pal), Anonymous bb 22358 (BO, L, SING); Gajoe Loeéus, G. Agosan, Anonymous bb

22426 (A, BO, BRI, L, NY, SING). West Coast: Soeliki, Anonymous bb 6536 (BO, L),

de Boer 25 (bb 7428) (BO); Fort de Kock, Theunissen 2 (BO, L); Mt. Sago, near Paja-

koemboeh, Maradjo 25 (L); Malalak, Anonymous bb 7421 (L); Pariaman, Diepenhorst

HB 1399 (BO, U), HB 2560 (BO), HB 2917 (BO, L, U); Sawahloento, Ham, 29 May 1911

(BO, L); Lasi, Anonymous bb 13104 (BO); without definite locality, Anonymous bb

6621 (BO). East Coast: Sibaulangit, Lorzing 435] (BO), 5282 (BO); Langkat, Timbang,

Lesger 255 (L, SING).

102 Gard. Bull. Sing. 34:1 (1981)

Java. West: G Kendeng, Backer 25908 (BO, L); G. Salak, Blume, October 1822

(L, lectotype of Philagonia sambucina Bl.; L. PR, duplicates of lectotype), Koorders

241568 (BO); G. Batoe, Backer 25786 (BO); Sanggrawa, Koorders 69088 (BO), 69098

(BO), 69106 (BO, P); Preanger, Koorders 26356 (BO); G. Kapal, Koorders 110968 (BO).

Central: Pringombo, Koorders 69118 (BO), 379068 (BO). Without definite locality:

Blume (A, BO, CANB, GH, L, NSW, U, US).

Lesser Sunda Islands. Sumbawa: Mt Batulante, Kostermans 18864 (A, BO); without

definite locality, Kostermans 18716a (L).

Blume’s original description of Philagonia sambucina was based on staminate and

carpellate material. The lectotype is chosen from the latter.

2. Tetradium fraxinifolium (Hook.) Hartley, comb. nov.

Philagonia fraxinifolia Hook., Ic. Pl. (1845) t. 710. Lectotype (chosen here): Wallich,

1821 (carpellate flowering material), Nepal.

Evodia fraxinifolia (Hook.) Benth., Fl. Hongkong (1861) 59.

Evodia impellucida Hand.-Mazz., Symb. Sin. 7 (1933) 626. Type: Handel-Mazzetti

9393, China, Yunnan Province, prope fines Tibeto-Birmanicas, in faucium

Naiwanglong.

Evodia poilanei Guill., Bull. Soc. Bot. France (1944) 91, (1945) 214. Syntypes:

Poilane 12927, Vietnam, Tonkin, O. de Chapa; Poilane 26800, Tonkin, N. de

Phong-ho.

Evodia subtrigonosperma Huang, Acta Phytotax. Sin. 6 (1957) 118.t31. Type:

Yu 20510, China, Yunnan Province, in silvis ad Muhconga, Kiukiang.

Evodia impellucida var. macrococca Huang, ibid. 117. t. 30. Type: Tsai 56591,

China, Yunnan Province, Shang-pa-hsien (not seen).

Evodia robusta Huang, ibid. 119, non Hook. f., 1875. Type: Tsai 56776, China,

Yunnan Province, Lu-si-hsien (not seen).

Small to medium tree to 12 m high; branchlets glabrous, glabrate, or occasionally

finely pubescent. Leaves 23—67 cm long; rachis glabrous, glabrate, or occasionally finely

pubescent; leaflets 2—7 pairs; petiolules of lateral leaflets obsolete to 10 mm long,

terminal leaflet on an extension of the rachis 12—35 mm long; blades of leaflets charta-

ceous to subcoriaceous, usually conspicuously oil-dotted, below drying whitish, green, or

brown, often glaucous and sometimes minutely (10x) subpapillate, glabrous to sparsely

or rarely densely pubescent, above glabrous to puberulent, lanceolate or occasionally

ovate, elliptic-oblong, or elliptic, 9-25 cm long, 3.5—8.5 cm wide, base in lateral leaflets

acute to rounded, often oblique, in terminal leaflet cureate, main veins 13—22 pairs,

departing from the midrib at an angle of 65—80 degrees ultimate veinlets usually in-

conspicuous, margin crenulate or rarely entire, marginal oil dots larger than those

scattered elsewhere on the blade, apex acuminate, the acumen 7—25 mm long. Inflore-

scences 7—24 cm long, rachis, branches, and pedicels puberulent to pubescent, pedicels

1—6 mm long. Flowers predominantly 4-merous (occasional flowers 5-merous); sepals

finely pubescent to glabrous, ciliolate, 0.5—1.5 mm long; petals pale yellow to green,

Hartley: Tetradium (Rutaceae) 103

drying brown, abaxially glabrous, adaxially villous to glabrous, 3.5—6.5 mm long; disc

glabrous; ovary appressed-pubescent to glabrous, the carpels connate toward the base

abaxially, free to the base adaxially, ovules 2 per carpel, subcollateral; rudimentary

carpels sparsely to rather densely appressed-pubescent toward the base, otherwise gla-

brous, connate toward the base. Fruits predominantly 4-carpellate (occasional fruits

5-carpellate), 1—4 of the carpels developing into follicles; follicles sparsely pubescent to

glabrous, compressed-subglobose, not beaked, 5.5—10 mm long, connate toward the base

abaxially, free to the base adaxially, endocarp sparsely pubescent or glabrous. Seeds

reddish-brown to brownish-black, 2 per follicle, subcollateral, separate, subtrigonous,

equal or slightly unequal in size, 4-5 mm long, firmly attached in the dehisced follicle

to a rather thick, fleshy, funicular aril; outer testa spongy (bounded externally by a shiny,

crustaceous pellicle); inner testa bony.

Distribution. Nepal east to northern Vietnam and southwestern China; recorded from

well-drained forests and thickets, 750—3000 m.

Nepal. Annapura Himal, Lati Khola, Stainton, Sykes, & Williams 6679 (BM); north

of Pokhara, Stainton, Sykes, & Williams 5064 (A, BM); Ganesh Himal, Stainton 3685

(BM); Likhu Khola, Stainton 4546 (BM); Tamur Valley, southeast of Terhathum,

Williams & Stainton 8465 (BM); Mewa Khola, Stainton 6838 (BM); Pha Khola, Williams

1019 (BM); Yampodin-Ghatte, Kanai, Murata, & Togashi 6301109 (BM); without definite

locality, Wallich Cat. No. 8521 (BM, GH, L, P, W), Wallich, 1820 (BM), Wallich, 1821

(K, lectotype of Philagonia fraxinifolia Hook.).

Sikkim. Suriel, Wilson, 8 September 1921 (A); without definite locality, Clarke

26635B (L), Hooker (BRI, GH, L, MEL, NY, US, W), King 4868 (CAL), Herb. Kuntze

6930 (NY), Thomson, 3 May 1857 (L, US), Treuther 139 MEL).

Bhutan. Shali, Cooper 4473 (BM).

India. West Bengal: Darjeeling, Clarke 27327 (BM), 35229C (BM); Darjeeling District,

Takdah, Chakrabutti 132D (DD), Lace 2205 (CAL). Assam: Garo Hills, Panigrahi 22427

(L); Shillong, Clarke 44335 (US); Khasi Hills, Chand 5377 (L), 6160 (MICH), 8294

(L, MICH), Clarke 7312 (BM), 45921B (US), Hooker & Thomson (BRI, GH, L, MEL,

NY, U, W), Kanjilal 2634 (DD), Koelz 23190 (MICH), 28383 (MICH), 29441 (MICH),

30358 (MICH), 33721 (MICH); Khasi and Jaintia Hills, Kanjilal 4553 (CAL, DD); Khasi

Hills and Bhramaputra Plains, Herb Kurz (BM); Subansiri Frontier Division, Palin to Amji,

Sastry 40950 (L); Naga Hills, Koelz 25374 (MICH), 26051 (L, MICH), 261718 (MICH),

Prain’s Collector 953 (A, W); Lushai Hills, Koelz 27485 (MICH); without definite

locality, Biswas 4060 (A), King’s Collector (L).

Burma. Valley of the Nam Tamai, Kingdon-Ward 13088 (BM); North Triangle,

Kingdon-Ward 21150 (A); hills around Htawgaw, Forrest 25070 (E); Hkamhkam-

Htawgaw Road, Kermode 17346 (DD); Kachin Hills, Mokim, March 1898 (A); without

definite locality, Rodger 139 (CAL).

Thailand. Northern: Doi Angka, Pa Ngein, Garrett 644 (L), Kerr 644 (L).

Vietnam. Tonkin: N. de Phong-ho, Poilane 26800 (P, holosyntype of Euodia poilanei

Guill.); O. de Chapa, Poilane 12927 (P, holosyntype of Euodia poilanei Guill.).

China. Yunnan Province: Prope fines Tibeto-Birmanicas in convalle fluvii Djeou-

djian in pluviisilva frondosa subtropica faucium Naiwanglong, Handel-Mazzetti 9393

104 Gard. Bull. Sing. 34:1 (1981)

(W, holotype of Euodia impellucida Hand.-Mazz.; A, isotype); Shweli-Salween Divide,

Forrest 24115 (BM, K); Shweli Valley, Forrest 8687 (E); three days south of Teng-yueh

[Teng-chung] , Forrest 26470 (E, US); Kiukiang Valley north of Muhconga, Yu 20510

(PE, holotype of Euodia subtrigonosperma — only a photograph seen; E, isotype); Kiu-

kiang, Taron, Chiengen, Yu 1945] (E); without definite locality, Forrest 15803 (K),

16079 (K), 17743 (K), 17859 (K), 18592 (K), 25250 (K).

Cultivated. India: Madras, Nilgiri Hills, Brandis 1855 (A).

The type sheet of Philagonia fraxinifolia consists of two flowering branchlets, one

staminate and one carpellate, and one separate carpellate inflorescence. Because the

species is dioecious with unisexual flowers, these specimens would have been taken from

two or perhaps three different plants. The carpellate flowering branchlet is chosen as the

lectotype.

In the plate accompanying the original description of Philagonia fraxinifolia, figure 1

depicts a bisexual flower, although in the legend it is referred to as a female flower. The

same drawing is on the type sheet. This is obviously an error because the female flowers

in this species have ligulate staminodes rather than fully developed stamens as are shown

in the drawing.

In the original description of Philagonia fraxinifolia, Hooker gives the following

synonymy:

Tetradium? fraxinifolium Wall. in Herb. Hook. 1821 [Wallich’s question mark].

Rhus fraxinifolium? Don, Prodr. Fl. Nep. p. 248 [Hooker’s question mark] .

The Wallich name, which also appears (with the question mark) on the label of the

type sheet of Philagonia fraxinifolia, was never published, and Hooker merely questions

the conspecificity of Rhus fraxinifolium and Philagonia fraxinifolia. In a short discussion

following the description he states: “It /Philagonia fraxinifolia/ may be the Rhus above

quoted of the late Mr. Don, though assuredly the style and stigma do not correspond with

that genus.” I have not seen the type of the Don plant (it is apparently not in the British

Museum — R. J. Henderson, pers. comm.), but it was described as having hermaphroditic

flowers and serrate leaves, neither of which characters agrees with material I have seen of

Tetradium fraxinifolium.

The type collections of Euodia impellucida, EF. poilanei, and E. subtrigonosperma,

and a paratype (Forrest 8687) of E. impellucida vat. macrococca, the type of which I

have not seen, all fall well within the range of variability of the Himalayan and Indian

collections that center, morphologically, around the type of Philagonia fraxinifolia. None

of the authors of the former taxa mention a possible relationship with the latter species,

and I suspect they did not compare it with their new taxa.

I have not seen any authentic material of Euodia robusta Huang (a later homonym of

E. robusta Hook. f., described from Malaya), but judging from the description of its

seeds (two per locule, subtriangular, reddish-brown, 4—5 mm long), it is certainly allied

to Tetradium fraxinifolium, and on other characters falls reasonably well within its range

of variability.

In the original description of Euodia impellucida, Handel-Mazzetti described the

leaflets as “ . . . glandulis pellucidis marginalibus tantum.” In the type collection of this

Hartley: Tetradium (Rutaceae) 105

species the leaflets are glaucous and minutely subpapillate below with conspicuous

marginal oil dots. Smaller oil dots are, in fact, present elsewhere on the blade but are

mostly obscured by the waxy bloom.

3. Tetradium daniellii (Benn.) Hartley, comb. nov.

Xanthoxylum daniellii Benn., Ann. Mag. Nat. Hist. ser. 3:10 (1862) 201, ¢. 5.

Type: Daniell, 1861, China, Liaoning Province, Talien-whan.

Zanthoxylon bretschneideri Maxim., Bull. Acad. Imp. Sci. Saint-Petersbourg 29

(1884) 73; Melanges Biol. Bull. Phys.-Math. Acad. Imp. Sci. Saint-Petersbourg

9 (1884) 655. Type: Bretschneider 1328, China, Hopeh Province, Shang-fang-

shan.

Evodia daniellii (Benn.) Hemsl., J. Linn. Soc. Bot. 23 (1886) 104.

Ampacus daniellii (Benn.) Kuntze, Rev. Gen. Pl. 1 (1891) 98.

Evodia delavayi Dode, Bull. Soc. Bot. France (1908) 55, (1909) 707. Type: Delavay

4526, China, Yunnan Province, Ta long tan pres de Ta pin tze.

Evodia henryi Dode, ibid. 706. Syntypes: Farges 413, China, Szechwan Province,

distr. de Tchen-kéou-tin; Henry 6712, China, Hupeh Province, Ichang; Wilson

3183, China, Hupeh Province, Hupeh occidental (not seen).

Evodia hupehensis Dode, ibid. 707. Type Henry 2939, China, Hupeh Province,

Ichang.

Evodia labordei Dode, ibid. 707. Type: Laborde & Bodinier 2729, China, Kweichow

Province, Mt. de Kaopo (not seen).

Evodia sutchuenensis Dode, ibid. 705. Type: Farges 1284 p.p. staminate, China,

Szechwan Province, distr. de Tchen-kéou-tin.

Evodia baberi Rehd. & Wils., p.p., in Sargent, Pl. Wils. 2 (1914) 131. Syntypes:

Wilson 1164, July 1908 (staminate), China, Szechwan Province, Wa-shan (=

Tetradium daniellii); Wilson 1164, October 1908 (carpellate), China, Szechwan

Province, Wa-shan (= Tetradium ruticarpum).

Evodia henryi var. villicarpa Rehd. & Wills., ibid. 134. Type: Wilson 3571, China,

Szechwan Province, west of Romi-chango.

Evodia velutina Rhed. & Wils., ibid. 134. Type: Wilson 994, China Szechwan Pro-

vince, west and near Wen-chuan Hsien.

Evodia vestita W. W. Smith, Notes Roy. Bot. Gard. Edinburgh 10 (1917) 38. Type:

Forrest 11091, China, Yunnan Province, on the Tong Shan in the Yangtze bend.

Evodia daniellii var. delavayi (Dode) Huang, Acta Phytotax. Sin. 6 (1957) 128, ¢.

36, fig A

Evodia daniellii var. henryi (Dode) Huang, ibid. 129; t. 36, fig. C.

106 Gard. Bull. Sing. 34:1 (1981)

Evodia daniellii var. hupehensis (Dode) Huang, ibid. 131; t. 36, fig. H.

Evodia daniellii var. labordei (Dode) Huang, ibid. 130; 't. 36, fig. B.

Evodia daniellii var. villicarpa (Rehd. & Wils.) Huang, ibid. 128; t. 36, fig. G.

Shrub or small to rather large tree to about 20 m high; branchlets puberulent to

finely pubescent, becoming glabrous or nearly so. Leaves 15—44 cm long; rachis pube-

scent to glabrous; leaflets 2-4(-5) pairs; petiolules of lateral leaflets obsolete to 8 mm

long, terminal leaflet on an extension of the rachis 6—35 mm long; blades of leaflets

chartaceous to subcoriaceous, sparsely and usually inconspicuously oil-dotted, below

drying pale grey-green and occasionally somewhat glaucous, usually glabrous except for

rather long hairs along the midrib toward the base (often restricted to the axils of main

veins toward the base) grading to entirely glabrous or rather densely pubescent, above

glabrous or nearly so, broadly ovate to lanceolate or rarely elliptic or elliptic-oblong,

5—18.5 cm long, 2.5—10.5 cm wide, base in lateral leaflets acute to subtruncate, sub-

rounded, or subcordate, usually slightly oblique, in terminal leaflet acute to cuneate or

occasionally rounded, main veins 7—14 pairs, departing from the midrib at an angle of

45—75 degrees, ultimate veinlets usually inconspicuous, margin subentire to crenulate,

often with rather large oil dots associated with the crenulations, apex acuminate, the

acumen 3—20 mm long. Inflorescences 3.5—19 cm long, rachis, branches, and pedicels

puberulent to pubescent, pedicels obsolete to 5 mm long. Flowers predominantly

5-mérous (occasional flowers 4-merous); sepals pubescent to glabrate, ciliolate, 0.5—1.5

mm long; petals white or whitish, drying whitish or pale brown, abaxially glabrous,

adaxially sparsely or occasionally densely pubescent or villous, (2.5—)3—5 mm long;

disc glabrous; ovary pubescent between the carpels, otherwise rather densely hairy to

glabrous, the carpels free to the base, ovules 2 per carpel, superposed; rudimentary

carpels densely or occasionally rather sparsely hirsute, free to about the base. Fruits

predominantly 5-carpellate (occasional fruits 4-carpellate), 2—5 of the carpels developing

into follicles; follicles sparsely to rather densely pubescent with spreading hairs, narrowly

pyriform, tapering apically-adaxially into a narrow beak 0.7—4.5 mm long, including the

beak 5—11 mm long, free to the base, endocarp glabrous. Seeds black, 2 per follicle,

superposed, ovoid to ellipsoid, coherent, the upper seed fertile, 2.5—4 mm long, the lower

seed sterile (containing what appears to be endosperm but no embryo), usually 1.5—3.5

mm long, the pair of seeds attached in the dehisced follicle to an adaxial strip of

pergamentaceous endocarp tissue; testa in fertile seed bony, without spongy outer testa,

in sterile seed thin, brittle, also without spongy outer testa.

Additional illustrations. Huang, Acta Phytotax. Sin. 6 (1957) t. 35 & t. 37, figs.

D, E, & F (as Euodia daniellii); ibid. t. 34 (as Euodia sutchuenensis); ibid. t. 33 (as

Euodia vestita).

Distribution. Southwestern China (extreme south-eastern Tibet and Yunnan Pro-

vince) northeast to Korea; recorded from woods, margins, and open slopes; sea level to

3200 m.

China. Southeastern Tibet: Mekong-Salween Divide, Forrest 14299 (E, K). Yunnan

Province: Mekong-Salween Divide, Forrest 14772 (E, K), Rock 10270 (A, US); between

Yung-ning and Yung-peh [Yung-sheng] , Schneider 1663 (A, E); Chien-ch’uan-Mekong

Divide, Forrest 22294 (A, K); Tong-shan, in the Yangtze bend, Forrest 11091 (E, holo-

type of Fuodia vestita W. W. Smith; A, BM, K, isotypes); Ta-long-t’an pres de Ta-pin-tze

[Ta-p’ing-tzu], Delavay 4526 (P, holotype of Euodia delavayi Dode; K, isotype; A,

Hartley: Tetradium (Rutaceae) 107

photograph of isotype); valleé de Ta-ngai-tong, Maire 3361 (UC); without definite locality,

Forrest 16484 (K), 16926 (K). Szechwan Province: west of Romi-chango [Tan Pa],

Wilson 3571 (A, holotype of Euodia henryi var. villicarpa Rehd. & Wils.; BM, US, iso-

types); Wa-shan, Wilson 1164, July, 1908 (A, holosyntype of Euodia baberi Rehd. &

Wils.), 3570 (US); Monkong [Hsiao-chin] Ting, Wilson 3569 (A, US); Pao-hsing, Chu

3419 (BM, E, K, W); Li-fan Hsien [Li Hsien] , Wang 21610 (A); west and near Wen-ch’uan

Hsien, Wilson 994 (A, holotype of Euodia velutina Rehd. & Wils.; BM, US, isotypes);

Omei-shan, Fang 7855 (BM), Sun 687 (US); district de Tchen-kéou-tin [Chien-ko] ,

Farges 413 (NY, isosyntype of Euodia henryi Dode), 1284 (P, holotype of Euodia sut-

chuenensis Dode; K, isotype); without definite locality, Bock & Rosthorn 1884 (A),

Farges 49 (W). Shensi Province: T’ai-pai-shan, Purdom 1038 (A, US), Wang 1518 (A);

Miao-wan, Fr. Hugh [Scallan], 1899 (A); Tsinling-schan [Tsin’lin Shan] inter Mei [Mei-

hsien] et Liu-pa, Fenzel 683 (W); Ning-shan Hsien, Kung 3051 (A); Ichun Hsien, Meyer

1929 (A); Huan-tow-shan, Fr. Hugh [Scallan], July 1899 (A, BM); Kusan [Ku Shan],

Fr. Hugh [Scallan] 75 (A, BM); Yuian-ch’ti District, Ye-cho-shan, Smith 6445 (A); with-

out definite locality, Giraldi, 1897 (A). Shensi Province; Ja Siu, Ren 6122 (UC). Hopeh

Province: Peking Mts., Bretschneider 152 (GH); Shang-fang-shan [Shan-fang-shan] ,

Bretschneider 1328 (LE, holotype of Zanthoxylum bretschneideri Maxim.), Liu 2217

(A, NY); Eastern Tomb, Li 10032 (NY); Kuan Tso Ling, Liu 1225 (A, UC). Liaoning

Province: Taliewhan [Ta-lien Wan], Daniell, 1861 (BM, holotype of Zanthoxylum

daniellii Benn.); New Town, Port Arthur [Lu-shan] , Wilson 8800 (A). Shantung Province:

Taitsinggong, prope Tsingtao [Ch’ing-tao], Licent 13367 (W); Lao Shan, Chiao 2708

(A, B, NY, UC, US), 2925 (A, NY, UC, US), 2926 (NY, US), 2942 (A, B, NY, US),

Meyer 278 (A); without definite locality, Meyer 262 (A). Honan Province: Hweihsien

[Hui-hsien] , Chunghu, Hers 739 (A); Chengchow [Cheng-hsien] , Hao 3508 (A); Sung-

hsien, Sankuan Miao, Hers 543 (A); Lushih, Lao Kiun Shan, Hers 1157 (A); without

definite locality, Wilson leg. Hers 3 (A). Kiangsu Province: Hwang San Yu, Ren & Tao

Univ. Nanking 12579 (UC). Hupeh Province: western Hupeh, Wilson 2183 (A, K, NY,

US, W), 2183a (K, NY), 2415 (A, K, NY, W), 2695 (K); Fang Hsien, Wilson 324 (A,

BM, GH, US, W), 4628 (A); Pa-tung Hsien, Henry 2555 (K), Wilson 3576 (A, BM, US, W);

Hsing-shan Hsien, Henry 6485 (K), Wilson 387 (US), 387a (A), 3574 (A, US); Chang-lo

Hsien [Wu-feng] , Wilson 387b (A); Pao-k’ang Hsien, Wilson 2693a (A, K); Nan-t’o and

mountains to northward, Henry 4534 K); I-ch’ang, Henry 2939 (BM, US, isotypes of

Euodia hupehensis Dode), 6712 (CAL, GH, US, isosyntypes of Euodia henryi Dode);

Ch’ang-yang Hsien, Wilson 3575 (A, US); Ch’ang-yang, Wilson 2693 (A, K, NY, W);

Heh Ya Tsze, Ching 3985 (A), Chun Univ. Nanking 4330 (GH); Liang Sung Kou, Chun

3783 (A), Univ. Nanking 4129 (US); Ta-gon, Chun leg. Huang 3969 (A); without definite

locality, Henry 4482 (CAL, NY), 4534A (MEL).

North Korea. Pingyang, Jack, 18 September 1905 (A, GH), Wilson 10568 (A, BM).

South Korea. Seoul, Jack, 24 September 1905 (A); Chemulpo [Inch’on], Carles

107 (BM, K); Province Keiki, Keijyo, Wislon 10555 (BM, US), Province Keiki, Hakusen

and on Kanghwa Island, Wilson 10612 (A, BM, US); Mt. Chii, Koidzumi, 1933 (MICH);

Shinsyu [Shinsu-do] , Keisyonando, Uno 2598 (A, NY, PH).

Cultivated. South Korea: Seoul, Chung 1169 (MICH), Faurie 468 (A). New Zealand:

Botanic Garden Christchurch, Sykes CHR 124917 (CHR), CHR 124998A (CHR), CHR

124998B (CHR). United States of America: California, Davis, McCaskill & Reagan 684

(AK); Massachusetts, Jamaica Plain, Arnold Arboretum, Hornby 4276 (MICH), 6898

(MICH), Merrill, August 1925 (NSW), Palmer, 4 August 1942 (AD, MICH, UC), Sorensen

6672 (AK); Maryland, Glendale, Plant Introduction Station, Meyer, 13 October 1967

108 Gard. Bull. Sing. 34:1 (1981)

(CHR); Pennsylvania, Reading, Hampden Heights, Shappell Garden, Brumbach 7238

(CHR), 7288 (AK, CHR). Great Britain: Edinburgh, Royal Botanic Garden, Anonymous

C6858 (CHR); Kew, Royal Botanic Gardens, Anonymous, 16 July 1961 (US, W), Boom

10347 (L). Netherlands: Baarn, Cantonspark, Anonymous 5537 (U), 6026 (U); Wagenin-

gen, Rijksarboretum, Boom 8456 (L); Den Haag, Zuiderpark, Boom 13136C (L). West

Germany: Darmstadt, Hortus, Boom 30080 (L), 34597 (L).

As interpreted here, Tetradium daniellii is quite variable, especially in petiolule

length (obsolete to 8 mm), vestiture of leaflet lower surface (glabrous to rather densely

pubescent), leaflet shape and size (broadly ovate to lanceolate or rarely elliptic or elliptic-

oblong, 5—18.5 cm long, 2.5—10.5 cm wide), vestiture of the follicle (sparsely to rather

densely pubescent), length of the follicle beak (0.7—4.5 mm), and overall length of the

follicle (6é—11 mm).

The type material of most of the names here placed in synonymy differs somewhat

from the type collection of Zanthoxylum daniellii, which has rather short- to medium-

length petiolules, ovate, medium-sized leaflets with vestiture restricted to the midrib

below, and sparsely pubescent, rather large follicles with medium-length beaks of

Euodia delavayi has subsessile leaflets that are pubescent along the midrib and main

veins below; that of E. henryi has nearly glabrous leaflets and unusually long-beaked

follicles; that of E. hupehensis has leaflets tending to.be lanceolate; that of

FE. sutchuenensis has unusually large, ovate to broadly elliptic leaflets; that of E. henryi

var. villicarpa has rather densely pubescent, long-beaked follicles; that of FE. velutina

has leaflets that are rather densely pubescent below and small, short-beaked follicles;

that of E. vestita has leaflets that are rather densely pubescent below and rather large,

long-beaked follicles. Euodia labordei, the type of which I have not seen, is described as

having subsessile to short-petiolulate leaflets that are sparsely pubescent on the nerves

below and rather small, villous follicles.

This may appear to be an overly conservative interpretation of this species, but in

the study of the large number of herbarium specimens of obviously closely inter-related

plants here assigned to it, I have not found sufficient morphologic discontinuity in any

of the variable characters, or sufficiently distinct combinations of various character

states to warrant recognition of more than a single taxon.

Zanthoxylum bretschneideri, the type of which closely matches that of Zanthoxy-

lum daniellii, was placed in the synonymy of Euodia daniellii by Rehder & Wilson (1914)

and Huang (1957).

One of the two syntypes of Euodia baberi (Wilson 1164, July 1908) closely matches

the type of Zanthoxylum daniellii. The other, as is noted in the synonymy, is Tetradium

rutaecarpum.

4. Tetradium calcicolum (Chun ex Huang) Hartley, comb. nov.

Evodia calcicola Chun ex Huang, Acta Phytotax. Sin. 6 (1957) 120.4. 32 . Type:

Ching 6306, China, Kwangsi Province, south of Nee Bai, Kweichow border.

Large shrub or small tree to 6 m high; branchlets puberulent, becoming glabrous or

nearly so. Leaves 9—19 cm long; rachis puberulent to glabrate; leaflets 2 pairs; petiolules

of lateral leaflets 1-2.5 mm long, terminal leaflet on an extension of the rachis 5—15

mm long; blades of leaflets subcoriaceous, sparsely and inconspicuously oil-dotted, below

Hartley: Tetradium (Rutaceae) 109

drying pale grey-green, somewhat glaucous, pubescent along the midrib toward the base,

otherwise glabrous, above entirely glabrous or puberulent on the midrib toward the base,

ovate to elliptic, 5—9.5 cm long, 2.3—4.5 cm wide, base in lateral leaflets obtuse to

rounded, usually slightly oblique, in terminal leaflet obtuse to rounded, main veins

10—14 pairs, departing from the midrib at an angle of 60—85 degrees, ultimate veinlets

usually inconspicuous, margin entire, apex acuminate, the acumen S—15 mm long. In-

florescences 5.5—13 cm long, rachis and branches finely pubescent, pedicels finely

appressed-pubescent, 1—3.5 mm long. Flowers 5-merous; sepals puberulent, ciliolate,

0.7—1 mm long; petals purple, drying dull purplish-red, abaxially sparsely to rather

densely appressed pubescent, adaxially villous, 3.5—4 mm long; disc glabrous, ovary

sparsely to densely appressed pubescent, the carpels free to the base, ovules 2 per carpel,

superposed. Fruits 5-carpellate, all of the carpels developing into follicles; follicles gla-

brate abaxially, densely appressed pubescent laterally, pyriform, not beaked, about

5 mm long, free to the base, endocarp glabrous. Seeds black, 2 per follicle, superposed,

ellipsoid, coherent, the upper seed fertile, about 3 mm long, the lower seed sterile (con-

taining what appears to be endosperm but no embryo), about 2 mm long, the pair of

seeds attached in the dehisced follicle to an axile strip of pergamentaceous endocarp

tissue; testa in fertile seed bony, without spongy outer testa, in sterile seed thin, brittle,

also without spongy outer testa. Staminate flowers not seen.

Distribution. Southwestern China, Kwangsi Province and (according to Huang, loc.

cit.) Yunnan Province; recorded from woods and thickets, 690—800 m.

China. Kwangsi Province: south of Nee Bai, border of Kweichow, Ching 6306 (A,

NY, UC, isotypes of Euodia calcicola Chun ex Huang); Pin-lam, Ko 55618 (A).

5. Tetradium glabrifolium (Champ. ex Benth.) Hartley, comb. nov.

Boymia glabrifolia Champ. ex Benth. in Hooker, J. Bot. Kew Gard. Misc. 3(1851)

330. Type: Champion 382, China, Hong Kong.

Megabotrya meliaefolia Hance ex Walpers, Ann. Bot. Syst. 2 (1852) 259. Type:

Hance, China, Hong Kong.

Evodia meliaefolia (Hance ex Walpers) Benth., Fl. Hongkong. (1861) 58.

Evodia glauca Mig., Ann. Mus. Bot. Lugd.-Bat. 3(1867)23. Syntypes (4

collections): Oldham 130, Japan, Nagasaki; Buerger, Mohnike, and FPierot,

Japan, without further locality.

Ampacus meliaefolia (Hance ex Benth.) Kuntze, Rev. Gen. Pl. 1 (1891) 98.

Evodia ailantifolia Pierre, Fl. For. Cochinch. 3 (1893) t. 287, fig. b. Type: Herb.

Pierre 3862, Vietnam, Austro-Cochinchina, in Prov. Tay nuih.

Eurycoma dubia Elmer, Leafl. Philipp. Bot. 2 (1908) 481. Type: Elmer 10120,

Philippines, Prov. Negros Oriental, Cuernos Mts.

Evodia balansae Dode, Bull. Soc. Bot. Fance (1908) 55, (1909) 705. Type: Balansa

4042, Vietnam, Tonkin, Tu-phap.

Evodia fargesii Dode, ibid. 704. Syntypes: Farges, China, Szechwan Province, distr.

de Tchen-kéou-tin; Wilson 1930 and 221]0a, China, western Hupeh Province.

110 Gard. Bull. Sing. 34:1 (1981)

Evodia yunnanensis Huang, Acta Phytotax. Sin. 6 (1957)104, 4.26 Type: Wang

80688, China, Yunnan Province, Cheun-yueh-hsien (only photograph seen).

Phellodendron burkillii van Steenis, Gard. Bull. Sing. 17 (1960) 357, fig. 1 Type:

Anonymous KEP 78904, W. Malaysia, Kedah, Enggang Forest Reserve.

Large shrub or small to medium tree to 20 m high; branchlets finely pubescent to

glabrate when young, becoming glabrous or nearly so. Leaves 14—38 cm long; rachis

finely pubescent to glabrous; leaflets (1—)2—9 pairs; petiolules of lateral leaflets 3—15

mm long, terminal leaflet on an extension of the rachis 10—35 mm long; blades of leaflets

chartaceous to subcoriaceous, sparsely and inconspicuously oil-dotted or without evident

dots, below drying whitish or pale green or less often brownish-green, usually obviously

glaucous, glabrous or with appressed to spreading hairs on the midrib, especially toward

the base and, to a lesser extent, on the main veins; above glabrous or with appressed to

spreading hairs on the midrib, especially toward the base, broadly ovate to lanceolate or

less often elliptic to elliptic-oblong, often strong unequal-sided, 4—15 cm long, 1.7—6 cm

wide, base in lateral leaflets acute to subrounded or subtruncate, usually oblique, in

terminal leaflet acute to cuneate, main veins 8—18 pairs, departing from the midrib at

an angle of 60—80 degrees, ultimate veinlets conspicuous (10x), densely reticulate,

margin entire or more or less crenulate, rarely with conspicuous oil dots associated with

the crenulations, apex acuminate, the acumen 5—30 mm long. Inflorescences 9-19 cm

long; rachis, branches, and pedicels finely pubescent to glabrous, pedicels 0.3—4 mm long.

Flowers predominantly 5—merous (occasional flowers 4-merous); sepals finely pubescent

to glabrous, ciliolate, about 0.5 mm long; petals green to yellow to white, drying whitish

to brown, abaxially glabrous or occasionally with sparse, appressed hairs, adaxially villous

to nearly glabrous, 2.7—4 mm long; disc glabrous; ovary rather densely pubescent be-

tween the carpels, otherwise glabrous or finely pubescent toward the apex, carpels free

to the base or connate toward the base abaxially, ovules 2 per carpel, collateral or sub-

collateral; rudimentary carpels sparsely to densely pubescent toward the base or occa-

sionally entirely glabrous, connate basally. Fruits predominantly 5-carpellate (occasional

fruits 4-carpellate), 4—S (or occasionally 1—3) of the carpels developing into follicles;

follicles sparsely to densely appressed-pubescent laterally, otherwise glabrous, trigonous,

not beaked, 3.5—S mm high and about as wide, free to the base or connate toward the

base abaxially, endocarp sparsely to rather densely pubescent. Seed black, | per follicle,

paired with an aborted seed, subglobose to broadly ellipsoid to ovoid, 2.5—4 mm long,

attached in the dehisced follicle to an adaxial strip of pergamentaceous endocarp tissue;

outer testa spongy (bounded externally by a shiny, crustaceous pellicle); inner testa

bony.

Additional illustrations. Huang, Acta Phytotax. Sin. 6 (1957) t. 25, figs. A, B, &C

(as Euodia glauca, E. meliifolia, and E. fargesii); Li, Woody Fl. Taiwan (1963) 370, t. 133

(as Euodia meliifolia); Chang in Li et al., FI. Taiwan 3 (1977) 516, t. 708 (as Euodia

meliifolia).

Distribution. Sikkim and northeastern India east through Indochina and southern

China to Taiwan and southern Japan, and south to W. Malaysia, Sumatra, and the

Philippines; recorded from well-drained forests, thickets, and open places; sea level to

1200 m.

Sikkim. Without definite locality, Kurz (BO); Thomson (BO).

India. West Bengal: Darjeeling Division, Division Forest Officer 1384 (DD); Tista

Valley, Haines BB873 (A, DD). Assam: Goalpara, Long Sung Block, Kanjilal 5081 (US);

Hartley: Tetradium (Rutaceae) 111

Darrang, Charduar Reserve, Kanjilal 4969 (A, DD); Khasi Hills, Chand 6002 (L), Koelz

30662 (L), Native Collector of Bot. Gard. Calcutta (BO); Khasi and Jaintia Hills, Anony-

mous 4586 (NSW); without definite locality, Biswas 3724 (A), Jenkins (BO, DD).

Burma. Maymyo, Maung Ba Pe 1576 (DD, MEL); Maymyo Plateau, Lace 5847

(CAL); South Tenasserim, Lace 715 (DD).

Thailand. Northern: Lampang, Che Sawn, Kerr 4762 (UC); Phitsanulok, Phusom-

saeng et al. BKF 46522 (L). Peninsular: Chumphon, Kuring, Kerr 11438A (L); Surat

[Surat Thani], Kanchanadit, Kerr 13035 (L); Krabi, Sangkhachand BKF 36894 (L).

Vietnam. Tonkin: Quonbi, Balansa 1171 (L); Tu-phap, Balansa 4042 (P, holotype

of Euodia balansae Dode; K, isotype); Dam Ha, Sai Wong Mo Shan, Tsang 30475 (A,

BO, L). Annam; Hue & vicinity, J. & M. S. Clemens 3286 (A, MEL, NY, PH, U, UC, US,

W); Province Tay nuih [Tayninh] , Herb. Pierre 3862 (P, holotype of Euodia ailantifolia

Pierre).

China. Yunnan Province: Che-li Hsien, Wang 75621 (A), 75796 (A, PE), 77601 (A),

77770 (A), 77809 (A), 77841 (A), 77859 (A), 77955 (A), 78669 (A); Jenn-yeh Hsien

[Cheun-yueh-hsien] , Meng-la, Wang 80688 (PE, holotype of Euodia yunnanensis Huang

— photograph seen); without definite locality, Forrest 12954 (K). Szechwan Province:

Tchen-keou-tin [Chien-ko] , Farges (BM, NY, US, isosyntypes of Euodia fargesii Dode);

Wan-hsien, Hwa 57 (A, UC); Wu-shan, Wilson 1930, May 1907 (W). Anhwei Province:

Chiuhwashan [Chiu-hua-shan], Ching 2773 (A, UC), Fan & Li 200 (K), Sun 1202 (A,

NY); 1300 (A, NY); Huang-shan, Chien 1129 (W); Si Liu, Chemen, Ching 3098 (A, K,

UC). Hupeh Province: Western Hupeh, Wilson 1930, June 1907 (A, E, isosyntypes of

Euodia fargesii Dode), 1930a (A, K, NY, W), 2210a (A, E, NY, W, isosyntypes of Euodia

fargesii Dode), 241 5a (K, W); Pa-tung Hsien, Wilson 247 (A, BM, US), 1930, July 1907

(K), 1930b (K), 2270 (A, K, US, W), 2322 (K, W), 3579a (A); Hsing-shan Hsien, Wilson

381 (A, US, W); Pakang [Pao-k’ang Hsien], Wilson 2415b (K); Nan-t’o and mountains

to northward, Henry 4577 (BM, GH, K); I-ch’ang, Henry 13 (K, US); Ch’ang-yang-hsien,

Wilson 253 (A, BM, US), 1930, August 1907 (NY), 3579 (A, BM, GH, US, W); without:

definite locality, Cheo Univ. Nanking 18389 (BO), Henry 6157 (NY), Silvestri 1218

(A, K). Chekiang Province: T’ien-mu-shan, Law 1277 (K), 1339 (K); Tai-pai-shan, Keng

1134 (A, UC); Tung-yang Hsien, Keng 914 (A, UC), 921 (A, UC); Yen-tang Shan, Chiao

Univ. Nanking 14748 (K, NY, UC); King Yuan [Ch’ing-yuan] , Ching 2559 (A, K, UC,

US, W); between P’ing-yang and Tai-shan, Ching 21/63 (A, BM, K, NY, UC, US, W);

Chen Chiong, 40 miles south of Siachu, Ching 1805 (A, BM, UC, W). Kweichow Province:

Ta Ho Yen, Fan Ching Shan, Kiangkou [Chiang-k’ou] Hsien, Steward, Chiao, & Cheo

329 (A, L, NY, US, W); Lang-tai, Tsiang 9567 (NY). Hunan Province: Pinkiang [P’ing-

chiang] , Hsiung 5726 (A); Sinning [Hsin-ning] Hsien, Fan & Li 556 (A, BM, BO, L, W);

Sintien [Hsin-t?ien] Hsien, Fan & Li 377 (A, BM, L, W). Kiangsi Province: Kiukiang

[Chiu-chiang] , Wilson 1584 (A, NSW, US); Kuling [Lu-shan], Wilson 1583 (A, BM,

US), 1585 (A, NSW), Hu 2358 (A), Tsiang 10718B (NY); Lushan Mts., Chung & Sun 255

(A, NY), 317 (A, NY), 329 (A, NY),685 (A, NY); Lu Shan, Cheo 3 (K, US), 169 (K), Chiao

Univ. Nanking 18766 (US), Steward Univ. Nanking 4668 (A, GH, UC, US); Feng-ch’eng,

Tsiang 10352 (NY); Lung-nan Hsien, Lau 4780 (A, BM, US). Kwangsi Province: Bako

Shan, W. Poseh [Pai-se] , Ching 7644 (NY, UC); Seh-fang Dar Shan, S. Nan-ning, Ching

7950 (A), 8317 (A, NY, UC); Shap Man Taai Shan, southeast of Shang-sze [Shang-ssu] ,

Tsang 24189 (A, NY); without definite locality, Ching 7679 (UC), Liang 69916 (A),

70120 (A). Kwangtung Province: Fang-ch’eng District, Kung P’ing Shan, Tsang 26685

(A, K); Kochow [Kao-chou] District, Kou Liang Ling, Tsiang 1017(A, K, NY, UC);

Lo-fou Shan, Merrill 10773 (A, NY, UC), Tsang Lingnan Univ. 9938 (US), Tsiang 1643

112 Gard. Bull. Sing. 34:1 (1981)

(A, MEL), 1720 (A, K, US); Canton [Kwang-chou] & vicinity, Levine CCC 1143 (A, GH,

US), CCC 1775 (A, GH), leg. Ah To CCC 1854 (A, GH, US), Sampson leg. Asui, 25

August 1885 (K); Hwei-yang District [Hui-yang Hsien], Lin Fa Shan, Tsang 25921 (A);

Poon Yue District, Levine CCC 3244 (A, GH); without definite locality, McClure Lingnan

Univ. 19709 (A, NY). Hainan Island: Ching Mai [Ch’eng-mai] District, Lei 9 (B, K, L,

NY, UC, US, W), 832 (A, B, BO, K, L, NY, US, W); Nodoa [Tan-hsien] & vicinity, Chun

Univ. Nanking 5687 (UC), Univ. Nanking 5766 (UC), Tsang Lingnan Univ. 15585 (A,

K, NY, UC, US); Dung Ka to Wen Fa Shi, Chun & Tso 43761 (A, L, NY, US, W), 43795

(BISH, L, NY, W); Dung Ka to Mo San, Chun & Tso 43564 (NY); Ch ‘ang-kiang [ Ch’ang-

chiang] District, Ngo Ko Shan, Lau 1930 (A, NY); Five Finger Mountain [Wu-chih

Shan], Chun Univ. Nanking 7056 (UC), Fenzel 222 (W); Kan-en [Kan-ch’eng] District,

Chim Fung Mountain, Lau 5346 (A); Manning [Wan-ning] , How 73189 (BO); Lo-tung,

Lau 27468 (A); Hainan, Liang 63392 (NY), 64634 (NY), 65135 (A, NY), Wang 33111

(NY), 34072 (NY, UC, US), 35680 (NY, US), 36445 (NY); Po Teng [Pao-t’ing] &

vicinity, Chu 29 (UC), How 72749 (BISH, BM); Yaichow [Yai-ch’eng], How 71094

(A, B, NY, US), How & Chun 70139 (B, NY, US), Liang 62250 (A, NY), 62345 (NY),

63026 (NY, US); Fan Yah, Chun & Tso 44010 (A, BISH, L, NY, W); Taam-chau District,

Tsang Lingnan Univ. 16247 (A, K, NY, UC); Loi area, Hung Mo Shan, McClure Lingnan

Univ. 18274 (A, B, K, NY, UC, US), Tsang & Fung Lingnan Univ. 18274 (B, MICH);

Tai Tsing, McClure CCC 7751 (A, BM, K, NY); without definite locality, Chun Univ.

Nanking 5899 (UC), Univ. Nanking 7121 (UC), Fenzel 24 (W), Henry 8706 (K). Hong

Kong: Kowloon, Liou 720 (NY); Chung Chi College, Hu 5567 (US), 5596 (US); Tai-o,

Chun 3099 (A, NY); Tai P’o, McClure CCC 13300 (A); Saigon, Chun 6864 (A, BO, UC);

Lantau Island, Tsang Lingnan Univ. 16637 (A, NY, UC, US, W); Mt. Victoria, Tang 257

(A); Wan-tsai Gap Road, Sampson, 5 November 1886 (K); Bowen Road, Lau 159 (A);

Happy Valley, Lamont, October 1874 (L), Sargent, 5 November 1903 (A); Little Hong

Kong Woods, Lamont 116 (L); without definite locality, Champion 382 (K, holotype of

Boymia glabrifolia Champ. ex Benth.), Chun 5225 (UC), 6562 (UC), 7477 (NY), 40010

(K, NY), Herb. Forbes 68 p. p. (PH), Ford, 17 August 1893 (A), Gibbs Herb. Hongkong

7456 (NSW), Hance (K, holotype of Megabotrya meliifolia Hance ex Walpers), Herb.

Hance 311 (BM, W). Fukien Province: Chuanchow [Ch’uan-chou], Chung 1050 (UC).

Without definite locality: Parkes, 1841 (K).

Taiwan. Taipei, Kangu, Keng, 26 October 1950 (A, L, US); Taihoku, Sinten, /to,

24 September 1923 (BM); Nan-t’ou, Lake Candidius [Jih-yueh T’au] , Wilson 997] (A);

Nan-t’ou, Mt. Bigen, Kawakami & Mori 3196 (A); Heng-ch’un, Chang 2418 (NY); Bankin-

sing, Henry 831 (A, K, NY), 1562 (A, NY); in montibus Okaseki, Faurie 34 (A, BM);

South Cape, Henry 932 (A), 974 (A, BM), 1296 (A, K, US).

Ryukyu Islands. Amami O Shima, Hosoyamada, 29 July 1927 (A); Okinawa, Hatu-

sima 18236 (US), 18241 (US), Kurata & Nakaike 2302 (U), Sonohara 8 (US), 67 (L);

Ishigaki, Smith 120 (US); Yonaguni, Walker & Tawada 6837 (US).

Japan. Kyushu Island: Nagasaki, Maximowicz, 1863 (BM, BO, K, L, MEL, NY, US,

W), Oldham 130 (L, holosyntype of Euodia glauca Miq.; GH, K, W, isosyntypes); Isahaya,

Hiroe 247] (UC); Province Satsuma, Masamune, 26 September 1922 (NY); Province

Ohsumi, Satamura, Hatusima 14525 (A); without definite locality, Wilson 6112 (A).

Without definite locality: Buerger (L, holosyntype of Euodia glauca Miq.; U, isosyntype),

Mohnike (L. holosyntype of Euodia glauca Miq.), Pierot (L, holosyntype of Euodia

glauca Miq.), Teysmann (BO).

W. Malaysia. Kedah, Enggang Forest Reserve, Anonymous KEP 78904 (L, holotype

of Phellodendron burkillii van Steenis; K, isotype).

Hartley: Tetradium (Rutaceae) 113

Sumatra. Central, Tandjoeng Ampaloe, Koorders 104108 (BO).

Philippines. Luzon: Mountain Province, Ifugao, Banaue, Conklin & del Rosario PNH

72715 (L); Benguet Subprovince, Panai, Santos BS 31968 (A, P, US). Negros: Negros

Oriental, Dumaguete (Cuernos Mts.) Elmer 10120 (A, BISH, K, L, NY, US, W, isotypes

of Eurycoma dubia Elmer).

Cultivated. India, Dehra Dun, New Forest, Raizada, July 1939 (DD), September

1939 (DD).

The type collections of Boymia glabrifolia and Megabotrya meliifolia, both from

Hong Kong, are clearly conspecific and the two names have long been considered to be

synonymous. Assuming that their respective dates of publication as given in the present

synonymy are correct, the former name has priority. Apparently only Seemann (Bot.

Voy. Herald (1857) 370), who placed the latter name in synonymy under the former,

considered this to be the case. Bentham (loc. cit.), in publishing the new combination

Euodia meiiifolia, listed Boymia glabrifolia in its synonymy and _has been followed by

subsequent authors.

The leaves of Tetradium glabrifolium, as the species is interpreted here, are quite

varaible in length (14—38 cm) and in number of leaflets [(1—)2—9 pairs]. The larger

leaves of the syntypes of Euodia glauca, from Japan, and the type of Euodia ailantifolia,

from Vietnam, are about twice as long (up to 38 cm) and have about twice as many

leaflets (up to 9 pairs) as those of the type collection of Boymia glabrifolia. In other

characters, these collections do not differ significantly, and among other collections of

Tetradium glabrifolium, especially from Kwangtung Province and Taiwan, a complete

range of intermediate leaf sizes and leaflet numbers may be found.

The type collection of Eurycoma dubia, from the Philippines, is in young flower

bud, but is almost certainly conspecific with the type of Boymia glabrifolia. Merrill,

Philipp. J. Sci. 14 (1919) 409, placed the Philippine species in the synonymy of Euodia

meliifolia.

The type of Euodia balansae, from Vietnam, is similar to that of Euodia ailantifolia,

differing mainly in having leaves with fewer leaflets (4—5 pairs). Dode, sometime after

his publication of the former species, recognized the two as being conspecific. An un-

dated annotation slip on the holotype sheet of F. ailantifolia reads: ‘“‘Evodia balansae

mihi Dode Specimen a me visum post speciem descriptam.”

The syntypes of Euodia fargesii, from the Chinese provinces of Szechwan and Hupeh,

have leaflets that are pubescent along the lower midrib toward the base. As Tetradium

glabrifolium is interpreted here, this is a common feature in collections from the northern

part of its range. Most collections from the southern part of the range, including the type

of Boymia glabrifolia, have glabrous leaflets, but occasional collections from southern-

most China, Indochina, Taiwan, and the Philippines have similarly pubescent leaflets.

The collection number of one of the syntypes of Euodia fargesii, Wilson 1930,

represents four collections made from different localities on different dates. They are as

follows: May 1907, Szechwan (W); 7 June 1907, Western Hupeh (A, E); 1 July 1907,

Hupeh, Pa-tung (K); and August 1907, Hupeh, Chang-yang (NY). Dode, in the original

description of FE. fargesii, cited only the collection from Western Hupeh.

114 Gard. Bull. Sing. 34:1 (1981)

I have seen only a photograph of the type of Euodia yunnanensis, but all nine of the

Wang collections cited above from Yunnan Province, Che-li Hsien, are paratypes. These

collections differ from most of the other material I have placed in Tetradium glabrifolium

mainly in having leaflets with the lower surface pubescent on the midrib and main veins.

This is only a minor variation which also occurs, sporadically, in collections from Assam,

Thailand, Vietnam, and Kwangsi Province. The paratypes of £. yunnanensis also differ

from most of the other material of this species in having leaflets that are not noticeably

glaucous on the lower surface. Glaucousness appears to be variable in most parts of the

range of the species.

The type of Phellodendron burkillii, from W. Malaysia, is clearly conspecific with

that of Boymia glabrifolia. In an unpublished note distributed with reprints of the original

description of the Malayan speciies, van Steenis points out that since its publication it has

been found to be conspecific with Euodia meliifolia.

The leaflets of Tetradium glabrifolium are distinctive in the genus in having densely

reticulate ultimate veinlets that are clearly visible on the lower surface with about 10x

magnification. This is a difficult feature to describe quantitatively, but once recognized

provides a reliable “spot” character for the identification of the species.

6. Tetradium trichotomum Lour., Fl. Cochinch. (1790) 91. Type: Loureiro, Vietnam,

Cochinchina.

Brucea trichotoma (Lour.) Spreng., Syst. 1 (1825) 441.

Ampacus trichotoma (Lour.) Kuntze, Rev. Gen. Pl.1 (1891) 98.

Evodia viridans Drake, J. Bot. (Morot) 6 (1892) 273. Syntypes: Balansa 3669,

Vietnam, Tonkin, Mont Bavi; Balansa 4038, Tonkin, village de Tchion-tao a la

base du Mont Bavi; Balansa 4043, Tonkin, Tu-phap.

Evodia trichotoma (Lour.) Pierre, Fl. For. Cochinch. 3 (1893) t. 287, fig. a.

Evodia colorata Dunn, Kew Bull.(1906)2. Syntypes: Henry 12137, 12137A, 12137B,

and 12137C, China, Yunnan Province, Szemao.

Evodia hainanensis Merr., Philipp. J. Sci. 21 (1922) 346. Type: McClure CCC 8449,

China, Hainan Island, Five Finger Mt.

Evodia lenticellata Huang, Acta Phytotax. Sin. 6 (1957) 98,t.24. Type: Hung,

Chang, & Tsiang 33600, China, Szechuan Province, Mt. Omei (not seen).

Evodia trichotoma var. pubescens Huang, Acta Phytotax. Sin. 16 (1978) 83. Type:

Chang 10748, China, Kwangsi Province, Lung-lin Autonomous District (not

seen).

Shrub or small tree to 8 m high; branchlets finely pubescent to glabrate. Leaves 12—

37 cm long; rachis finely pubescent to glabrate; leaflets (1—)2—5(—6) pairs; petiolules of

lateral leaflets obsolete to 5.5(—10) mm long, terminal leaflet on an extension of the

rachis 10—20(—30) mm long; blades of leaflets chartaceous to subcoriaceous, oil-dotted,

below usually drying green or brownish-green, glabrous or with fine pubescence mainly

on the midrib and main veins, above glabrous or with fine pubescence on the midrib,

Hartley: Tetradium (Rutaceae) 115

elliptic, elliptic-oblong, lanceolate, or occasionally ovate, 3—16 cm long, 1.3—5 cm wide,

base in lateral leaflets cuneate to subrounded, often oblique, in terminal leaflet cuneate,

main veins 11—14 pairs, departing from the midrib at‘an angle of 65—85 degrees, ultimate

veinlets inconspicuous, margin entire, apex acuminate, the acumen 5—15 mm long.

Inflorescences 5—30 cm long, rachis finely pubescent to glabrate, branches and pedicels

finely pubescent, pedicels 0.5—2.5 mm long. Flowers predominantly 4-merous (occa-

sional flowers 5-merous); sepals finely pubescent to glabrate, ciliolate, 0.5—1 mm long;

petals green to yellow to white, drying brown to whitish, abaxially glabrous, adaxially

sparsely subvillous to glabrous, 3—4 mm long; disc glabrous; ovary glabrous or with a few

short hairs basally between the carpels, carpels connate toward the base abaxially, free to

the base adaxially, ovules 2 per carpel, collateral; rudimentary carpels sparsely hairy or

glabrous, connate toward the base. Fruits predominantly 4-carpellate (occasional fruits

5S-carpellate), 1—4 of the carpels developing into follicles; follicles entirely glabrous or

with sparse pubescence laterally toward the base, subglobose to obovoid, not beaked,

4—7 mm long, connate toward the base abaxially, free to the base adaxially, endocarp

glabrous. Seed balck, 1 per follicle, paired with an aborted seed, subglobose to broadly

ovoid, 3.7—6 mm long, attached in the dehisced follicle to an adaxial strip of perga-

mentaceous endocarp tissue; outer testa spongy (bounded externally by a shiny, crusta-

ceous pellicle); innter testa bony.

Additional illustration. Huang, Acta Phytotax. Sin. 6 (1957) 4. 23 (as Euodia

trichotoma).

Distribution. Thailand, Laos, and Vietnam north and east to the Chinese provinces

of Shensi, Hupeh, and Kwangtung; recorded from well-drained forests and thickets;

300—1900 m.

Thailand. Northern: Chiang Mai, Doi Chang, Rock 1778 (A, US).

Laos. Phongsaly,, Poilane 25989 (A, US).

Vietnam. Tonkin: Laichau, Poilane 29658 (L); Chapa, Petelot 2483 (A), 3124

(A, UC), 5692 (A, US); Mont Bavi, Balansa 3669 (G—DC, holosyntype of Euodia viridans

Drake); village de Tchion-tao a la base du Mont Bavi, Balansa 4038 (G—DC, holosyntype

of Euodia viridans Drake; K, LE, isosyntypes); Tu-phap, Balansa 4043 (G—DC, holosyn-

type of Euodia viridans Drake); route de Hanoi a Hoa Binh, Pételot 2599 (A); Dam-ha,

Sai Wong Mo Shan, Tsang 30357 (A, E, L); Cho Ganh, Pételot 1165 (UC); Hacoi, Taai

Wong Mo Shan, Tsang 27026 (A, E), 29300 (A, E), 29454 (A, BO. E, L). Annam: In

montibus Cochinchtnae, Loureiro (BM, holotype of Tetradium trichotomum Lour.).

China. Yunnan Province: Fo-hai [Meng-hai] , Wang 73987 (A), 74237 (A), 74893

(A), 77145 (A), 77294 (A); between Muang-hun and Muang-hai, Rock 239] (A, UC, US);

Che-li [Ching-hung] Hsien, Wang 75835 (A), 75942 (A), 77980 (A), 79663 (A); Szemao

[Fu-hsing-chen] , Henry 12137 (K, holosyntype of Euodia colorata Dunn; A, US, isosyn-

types), 12137A (K, holosyntype of Euodia colorata Dunn; A, NY, isosyntypes), 12]37B

(K, holosyntype of Euodia colorata Dunn; A, US, isosyntypes), /2/37C (K, holosyntype

of Euodia colorata Dunn; NY, isosyntype), 12237 (A, K, NY), 12237A (A, K), 12237B

(NY); I-wu, Henry 13577 (K); zw. Yangwu & Schangyentang n. von Yuenkiang, Wiss-

mann 120 (W); Meng-tsze [Meng-tzu] , Henry 1095] (A, K, NY, US); Ping-pien Hsien,

Tsai 61062 (BO), 62045 (BO); Nan-chiao, Wang 75/11 (A), 75116 (A). Szechwan Pro-

vince: Tienchuan [T’ien-ch’tian] , Kuan & Wang 3333 (K); Hung-ya Hsien, Wilson 3573

(A, GH, US, W); Kiating Fu [Lo-shan], Wilson 3572 (A, US), 4568 (A); An Hsien &

Vicinity, Wang 22187 (A); without definite locality, Chu 3997 (W), Fang 3376 (E, NY).

116 Gard. Bull. Sing. 34:1 (1981)

Shensi Province: Young-hsien, Koo 2033 (PE). Hupeh Province: Fang [Fang-hsien] ,

Wilson 2578 (K); Hsing-shan Hsien, Wilson 791 (A, US). Kwangsi Province: North Hin

Yen, Tsin Hung Shan, Ching 7010 (A, NY). Kwangtung Province: Fang-ch’eng District,

Kung-p’ing-shan, Tsang 26720 (E); Fang-ch’eng District, Na Leung, Tsang 26637 (A, K).

Hainan Island: Hung Mo Shan, Tsang & Fung Lingnan Univ. 18218 (A, K, NY), Tsang,

Tang, & Fung Lingnan Univ. 17744 (A, K, NY, UC, US); Tai Pin, Gressitt 1061 (BM,

UC), 1118 (A, BM, UC); Liamui, Gressitt 1186 (A); Fan Yah, Chun & Tso 44140 (A, B,

NY, US); Five Finger Mountain [Wu-chih Shan] , McClure CCC 8449 (A, US, isotypes of

Euodia hainanensis Metrr.); Pao-ting, How 71981 (A), 72442 (A), 73470 (A, BO); without

definite locality, Henry 8605 (K).

Euodia viridans, E. colorata, and E. hainanensis, here considered (on the basis of type

and syntype collections) to be conspecific with Tetradium trichotomum, have previously

been placed in synonymy under Euodia trichotoma — see Rehder and Wilson (1914),

Merrill, Comm. Lour. Fl. Cochinch. (1935) 219, and Huang (1957).

I have not seen the type collection of Euodia lenticellata, from Szechwan Province,

but four of the collections cited above, Wilson 3572, Wang 22187, and Chu 3997, from

Szechwan Province, and Ko 2033, from Shensi Province, are paratypes. As Tetradium

trichotomum is interpreted here, these specimens merely represent a northern variant

with reduced leaves (down to 12 cm long with leaflets as small as 3 cm long and 1.3 cm

wide) and flowers (petals about 3 mm long); among other collections of the species there

is a complete series of intermediates with the larger-leaved, larger-flowered plants, such as

represented by the type of T. trichotomum, that tend to occur in the southern part of

the range.

Huang (1957), in distinguishing between Euodia lenticellata and E. trichotoma in

a key, notes that they differ, respectively, in lenticel prominence (convex vs. not convex)

and vestiture of the lower surface of the leaflets (covered with long, soft hairs vs. glabrous

or nearly so). In the study of the above-mentioned paratypes of FE. lenticellata and

collections from the northern part of the range that match them closely (Wilson 791 and

2578, from Hupeh Province, Fang 3376, from Szechwan Province, and Henry 13577 and

Wang 75116, from Yunnan Province), I have found that the lenticels are variable in pro-

minence and that the lower surface of the leaflets varies from pubescent with spreading

hairs on the midrib and main veins to nearly glabrous with short, appressed hairs on the

midrib only. Similar variability in these characters also can be found among collections

of more typical Tetradium trichotomum from the southern part of the range.

7. Tetradium ruticarpum (A. Juss.) Hartley, comb. nov.

Boymia rutaecarpa A. Juss., Mem. Mus. Hist. Nat. 12 (1825) 507. t. 25, fig. 39,

Mem. Rutac. (1825) 124; ¢. 25, fig. 39. Type: d’Incarville, China, sub nomine

vernaculo Ou-Tchou-Yu (not seen).

Evodia rutaecarpa (A. Juss.) Benth., Fl. Hongkong. (1861) 59.

Ampacus rutiecarpa (A. Juss.) Kuntze, Rev. Gen. Pl. 1 (1891) 98

Evodia bodinieri Dode, Bull. Soc. Bot. France (1908) 55, (1909) 703. Syntypes:

Bodinier 1689, China, Kouy-tchéou [Kweichow] Province, mont de Lout Song

Koan (not seen); Ford (ex Herb. Hong Kong Bot. Gard) 343, China, Kouang-

tourng [Kwangtung Province] (not seen).

Hartley: Tetradium (Rutaceae) 117

Evodia officinalis Dode, ibid. Syntypes: Farges 305, China, Szechwan Province,

Sutchuen oriental, district de Tchen-Keéou-tin (not seen); Henry 6199, China,

Hupeh Province, Ichang;.Henry 1802 (not see) and 6136, China, Hupeh Province,

Patung District; Henry 6549, China, Hupeh Province (not seen); Wilson 1309,

China, Western Hupeh Province, Chien-shih Hsien.

Evodia baberi Rehd. & Wils., p. p., in Sargent, Pl. Wils. 2 (1914) 131. Syntypes:

Wilson 1164, July 1908 (staminate), China, Szechwan Province, Wa-shan (=

Tetradium daniellii); Wilson 1164, October 1908 (carpellate), China, Szechwan

Province, Wa-shan (= Tetradium ruticarpum).

Evodia rugosa Rehd. & Wils., ibid. 132. Type: Henry 10245, China, Yunnan Province

Mengize.

Evodia hirsutifolia Hayata, Ic. Pl. Formosa 6 (1916)5. Type: Inaba, August 1911

Formosa, Rinkiho.

Evodia compacta Hand.-Mazz., Symb. Sin. 7 (1933) 627. Type: Handel-Mazzetti

12320, China, Hunan Province, monte Yiin-shan bei Wu-kang.

Evodia compacta var. meionocarpa Hand.-Mazz.,ibid. Type: Chien 1029, China

Anhwei Province, Hwang-shan.

Evodia rutaecarpa vat. bodinieri (Dode) Huang, Acta Phytotax. Sin. 6 (1957) 113.

t. 27, figs. E & F.

Evodia rutaecarpa forma meionocarpa (Hand.-Mazz.) Huang, ibid. 112.

Evodia rutaecarpa var. officinalis (Dode) Huang, ibid. 114; t. 27, fig. A.

Shrub or small tree to 9 m high; branchlets puberulent to finely pubescent when

young, usually becoming glabrous or nearly so. Leaves 15—40 cm long; rachis finely

pubescent or occaisonally nearly glabrous; leaflets (1—)2—6(—7) pairs; petiolules of

lateral leaflets obsolete to 9 mm long, terminal leaflet on an extension of the rachis

10—30 mm long; blades of leaflets chartaceous, usually conspicuously oil-dotted, below

drying brown to greenish brown or rarely pale green and slightly glaucous, rather sparsely

to densely pubescent, above puberulent to densely pubescent on the midrib, otherwise

rather sparsely pubescent to glabrous, elliptic to ovate or less often lanceolate, oblanceo-

late, or obovate, often rather unequal-sided, 4.5—17 cm long, 2—8 cm wide, base in

lateral leaflets acute to obtuse or less often rounded, subtruncate, or cuneate, oblique or

not, in terminal leaflet cuneate or rarely attenuate, main veins 9—17 pairs, departing from

the midrib at an angle of 60—85 degrees, ultimate veinlets rather conspicuous (10x) or

not, rather loosely reticulate, margin entire or rather irregularly crenulate, inconspicuous

oil dots associated with the crenulations, apex acuminate, the acumen 7—20 mm long.

Inflorescences 2.5—18 cm long, rachis and branches sparsely to densely pubescent,

pedicels sparsely to densely pubescent, puberulent, or nearly glabrous, obsolete to 3.5

mm long. Flowers predominantly 5-merous (occasional flowers 4-merous); sepals sparsely

to densely pubescent, puberulent, or nearly glabrous, ciliolate, 0.5—1.2 mm long; petals

green to yellow to white, drying brown or whitish, abaxially glabrous or with sparse,

short hairs, adaxially villous to nearly glabrous, 3—5 mm long; disc glabrous; ovary

glabrous or sparsely hairy, the carpels connate toward the base abaxially, free to the base

adaxially, ovules 2 per carpel, collateral or subcollateral; rudimentary carpels glabrous or

118 Gard. Bull. Sing. 34:1 (1981)

rarely sparsely hairy, connate toward the base. Fruits predominantly 5-carpellate (occa-

sional fruits 4-carpellate), 1—5 of the carpels developing into follicles; follicles glabrous or

occasionally sparsely hairy, subglobose, not beaked, 3.5—6 mm long, connate toward the

base abaxially, free to the base adaxially, endocarp glabrous. Seed black, 1 per follicle,

paired with an aborted seed, ovoid or occasionally ellipsoid or subglobose, 3.5—5.5 mm

long, attached in the dehisced follicle to an adaxial strip of pergamentaceous endocarp

tissue; outer testa spongy (bounded externally by a crustaceous, shiny pellicle); inner

testa bony.

Additional illustrations. Siebold and Zuccarini, Fl. Jap. 1 (1837 or 1838) 50, t. 2/

(as Boymia ruticarpa); Huang, Acta Phytotax. Sin. 6 (1957) ¢. 27, figs. B, C, & D, & t. 28

(as Euodia ruticarpa).

Distribution. Nepal east to east-central China (Kiangsu and Chekiang Provinces) and

Taiwan; recorded from well-drained forests, thickets, and open places; 120—3000 m.

Nepal. Samela, Polunin, Sykes, & Williams 483 (BM); Gurjakhani, Stainton, Sykes,

& Williams 3549 (BM); Lumsum, Stainton, Sykes, & Williams 2930 (A, BM).

Sikkim. Lachung, Smith & Cave 2620 (MEL); without definite locality, Hooker

(GH, L, W), Prain’s Collector 281 (CAL).

Bhutan. Punakha, Cooper 3048 (E).

India. Assam: Khasi Hills, Kanjilal 7048 (CAL).

Burma. Adung Valley, Kingdon-Ward 9496 (A, BM).

China. Yunnan Province: Valli Doyonlumba ad fluvium Lu-djian (Salween), Handel-

Mazzetti 9605 (A, W); Chien-chuan-Mekong Divide, Forrest 21513 (K, US); Yunnan

Bor.-occident., Pe-yen-tsin, Ten 396 (E); Tali Range, Forrest 11668 (BM, K), 27966

(BM); vicinity of Tengyueh [T’eng-ch’ung] , Forrest 7592 (K), 11849 (BM, K, UC, W);

Mengze [Meng-tzu] , Henry 10245 (A, holotype of Euodia rugosa Rehd. & Wils.; K, NY,

isotypes); Ngaza, drainage basin of the Yangtze west of Likiang [Li-chiang] , Rock 10610

(A, US); Kiang-ti [Chiang-ti] , Maire 247 (E); Yungpeh [Yung-sheng] Mountain, Forrest

15105 (E, K); mountains of Yang-p’i River, Rock 6235 (A, US); without definite locality,

Forrest 13468 (K), Ten 271 (W). Szechwan Province: Huang-ya-hsien, Fang 8801 (A, K);

Omei Hsien, Chien 6055 (UC); Omei Shan, Faber 26 (A, K), 233 (K, NY), Fang 2412

(A, NY), 12553 (A, BM), Lee 3050 (US), Wang 23163 (A), Wilson 4772 (A, K); Omei-

shan, Fang 7444 (A, K, NY); Mo-ting-ling, Wang 22511 (A); western Szechwan, Wa-shan,

Wilson 1164, October 1908 (US, isosyntype of Euodia baberi Rehd. & Wils.); Nan-

ch’uan Hsien, Fang 1119 (A); vicinity of Wan-hsien, Hwa 83 (A, UC). Anhwei Province:

Chiu-hua Shan, Ching 2779B (A), Fan & Li 117 (K, US), 190 (K), Sun 1280 (A); Hwang-

shan, Chien 1029 (W, holotype of Euodia compacta var. meionocarpa Hand.-Mazz.);

Huang Shan, Ling Univ. Nanking 9623 (UC); Wang Shan, Ling Univ. Nanking 7732 (UC);

near Sujen, McClure Lingnan Univ. 15232 (A, UC); without definite locality, Chang

5102 (K). Kiangsu Province: I-hsing, Ling Univ. Nanking 12447 (UC). Hupeh Province:

Chien-shih Hsien, Chow 1714 (A, NY), Wilson 882 (K), 1309 (A, K, NY, US, W, isosyn-

types of Euodia officinalis Dode); western Hupeh, Wilson 2212 (K, W); Fang [Fang-

hsien], Henry 6569 (GH, K); Pa-tung, Chow 714 (A, NY), Henry 6136 (E, K, MEL,

NY, US, isosyntypes of Euodia officinalis Dode), Wilson 251 (A, BM, US); Chang-lo

[Wu-feng] , Wilson 3577 (A, BM); Nan-t’o and mountains to northward, Henry 2077

Hartley: Tetradium (Rutaceae) 119

(GH, K), 4525 (K); Ch’ang-yang, Henry 6199A (GH, K, US), Wilson 366a (A); I-ch’ang,

Henry 924 (K), 1676 (GH, K), 2259 (K), 2616 (K), 6199 (BM, E, GH, isosyntypes of

Euodia officinalis Dode), Wilson 366 (A, BM, GH,‘US); without definite locality, Henry

6136A (GH, K), Silvestri, August 1906 (A). Chekiang Province: Anh-ki [Chiu-an-chi] ,

Ching 4908 (A); Mo-kan Shan, Cheo & Wilson Univ. Nanking 12807 (GH, UC); Mo-kan-

shan, Read 1204 (BM); Tien-mu-shan [Hsi-t’ien-mu-shan] , Ching 5102 (A), Law 1347

(K), 1389 (K); Ch’ang-hua-hsien, Keng 631 (A, UC); Ning-po Mts., Faber 95 (A, K),

1717 (K); Tsing-tien-hsien [Ch’ing-t’ien-hsien] , Keng 47 (A), 96 (A); region of King

Yuan [Ch’ing-yuan], Ching 2493 (A, UC, US, W); without definite locality, Liou 380

(NY). Kweichow Province: Tsunyi [Tsun-i] Hsien, Liang-feng-yah, Steward, Chiao, &

Cheo 115 (L, NY, US, W); Tungt’ze [T’ung-tzu] , Tsiang 4919 (NY); environs du Kouy-

yang [Kuei-yang], Bodinier 1689 bis (P), Tsiang 8655 (NY); Kweiting [Kuei-ting] ,

Tsiang 5447 (NY); Tu-yun, Tsiang 5870 (NY); inter Nganschan et Nganping, Handel-

Mazzetti 10440 (A, W); inter Duyun et Gudong, Handel-Mazzetti 10690 (A, W). Hunan

Province: monte Yiin-schan bei Wu-kang, Handel-Mazzetti 12320 (W, holotype of Euodia

compacta Hand.-Mazz.; A, isotype); ad flumen Tsi-djiang supra Hsin-hwa, Handel-Maz-

zetti 12564 (A, W); Ch’ang-ning Hsien, Yang-shan, Fan & Li 180 (BM, BO, L, W). Kiangsi

Province: Kiukiang, Bullock 10 (US), Shearer, 1873 (K); Ti-ping-shan, Hsiung 5315 (A);

Kuling Lu-shan, Chiao Univ. Nanking 18585 (NY), Univ. Nanking 18618 (NY, US),

Chung 4382 (W); Lu Shan, Chung & Sun 614 (A, NY), Jp Univ. Nanking 1083 (UC);

Nan-ch’ang, Chung 607 (A), 693 (A); Kan River about 60 li south of Nan-ch’ang, Chung

28 (A), 31 (A); Kao-an, Hua-ling-shan, Tsiang 10406 (NY), 10419 (NY); Kiennan

[Ch’uan-nan] District, Tung Lei Village, Lau 4128 (A, BM); without definite locality,

Anonymous CCC 51919 (A). Kwangsi Province: Ling-yun Hsien, Loh Hoh Tsuen,

Steward & Cheo 463 (A, BM, NY, W); San-chiang Hsien, Ling-wang Shan, Steward &

Cheo 1016 (A, BM, NY, W); Tou-ngok Shan, near T’ung-chung Village, Tsang 23111

(A. W). Kwangtung Province: Lok-chong [Lo-ch’ang], Tso 20688 (NY), 20886 (NY);

Yu-yuen [Ju-yiian] , Ko 52936 (A); Ying Tak [Ying-te] , Tai Tsan, Tsang & Wong Lingnan

Univ. 14845 (A, UC), CCC 14188 (A, UC); Sin-fung [Hsin-feng] District, Taam 694 (A);

Wung-yuen District, Lau 2462 (A); Lung-t’au Shan, Anonymous CCC 12776 (UC, US,

W), To & Tsang 12776 (A, BM); Lok-fan Anonymous Lingnan Univ. 9933 (NY, US, W).

Fukien Province: Fan Ka Cha, Dunn 112 (A).

Taiwan. Taipingshan [T’ai-p’ing], Chang 4882 (NY); Kagi Province, Arisan [A Lan

Shan], Wilson 10880 (A); Nantou Hsien, Horisha, Sasaki 42 (UC), Wilson 9955 (A);

Nantou, Kawakami & Mori 3397 (A); Jitsugetsu Tan [Jih-yuen T’an] , Tanaka 468 (A,

BM, NY, US); Taiheisan (Mt. Taihei), between Taiheisan Club and Minamoto, Bartlett

6059 (MICH, US); Rinkiho, Inaba, August 1911 (TI, holotype of Euodia hirsutifolia

Hayata); without definite locality, Suzuki-Tokio 11705 (A).

Cultivated. China. Hunan Province: Ping-kiang [P’ing-chiang] , Hsiung 5723 (A).

Kwangsi Province: Ling-yun Hsien, Loh Hoh Tsuen, Steward & Cheo 703 (BM, BO, NY,

W); North Lin Yen, Tsin Lung Shan, Ching 6894 (A, UC, US). Japan. Honshu Island:

Yokohama, Maximowicz, 1862 (BM, BO, GH, L, US, W); Province Chiba, Konodai,

Santo 399 (US); Kyoto Prefecture, Kitaserakawa, Muroi 6942 (A); Koyoto Prefecture

Yase, Muroi 4985 (A); Koyasan, Faurie 95 (W). Kyushu Island: Nagasaki, Maximowicz,

1863 (BM), Oldham 131 (GH, K, L); Province Ohsumi, Mt. Kirishima, Masamure, 27

September 1924 (NY). Without definite locality: Herb. Buerger (U), Mohniki (L), Pierot

281 (L), Siebold, 1829 (GH, L).

I have not seen the type of collectin of Boymia ruticarpa and it is apparently not

among the specimens of the Jussieu herbarium photographed on IDC microfiches (L.

Pedley, pers. comm.). Nevertheless, on the basis of Jussieu’s original description and

accompanying illustration, I am reasonably certain of its identity.

120 Gard. Bull. Sing. 34:1 (1981) ©

I have included a rather wide range of specimens in this species among which I have

not been able to find any constant or correlating characters enabling me to recognise

more than a single taxon. Some of the morphologic variation I have accepted includes:

leaves 15—40 cm long; leaflets (1—)2—6(—7) pairs, 4.5—17 cm long, 2—8 cm wide,

sparsely to densely pubescent on the lower surface; inflorescences 2.5—18 cm long,

compact to spreading; petals 3—S mm long; fruits 3.5—6 mm long; and seeds 3.5—5.5

mm long.

Of the names placed in synonymy, the type material of Euodia officinalis has rather

large, spreading inflorescences and small flowers; that of E. rugosa has rather small,

sparsely pubescent leaves (the leaflets are more or less wrinkled, but I do not think this

is taxonomically significant in this case) and small, compact infructescences; that of

E. hirsutifolia has rather long leaves with small leaflets and small fruits; and that of E.

compacta and E. compacta var. meionocarpa have small, compact infructescences.

I have not seen either of the syntype collections of Euodia bodinieri, Bodinier 1689

and Ford 343, but it seems likely that one of the specimens cited above, Bodinier 1689

bis (P), which is labelled as Euodia bodinieri, was considered by Dode to represent that

species. It has rather small, sparsely pubescent leaves and small flowers.

Euodia hirsutifolia was placed in the synonymy of Euodia ruticarpa by Li, Woody

Fl. Taiwan (1963) 371 and Chang in Li et al., Fl. Taiwan 3 (1977) 517.

8. Tetradium austrosinense (Hand.-Mazz.) Hartley, comb. nov.

Evodia austrosinensis Hand.-Mazz., Sinensia 5 (1934) 1. Type: Ching 6214, China,

Kwangsi Province, Wang Tung, N. Luchen, near Shan-fang.

Small to medium tree 7—20 m high; branchlets pubescent when young, becoming

glabrous or nearly so. Leaves 20—35 cm long; rachis sparsely to densely pubescent;

leaflets 3—5 pairs; petiolules of lateral leaflets 2—4 mm long, terminal leaflet on an exten-

sion of the rachis 15—30 mm long; blades of leaflets chartaceous, sparsely and inconspicu-

ously oil-dotted or without evident oil dots, below glaucous, minutely (10x) papillate,

sparsely to densely pubescent with spreading or ascending hairs, above sparsely puberu-

lent to nearly glabrous, broadly elliptic or occasionally elliptic-oblong, ovate, ovate-

lanceolate, or, in some terminal leaflets, obovate to suboblanceolate, 5.5—14.5(—17) cm

long, 2.5—7(—8.5) cm wide, base in lateral leaflets obtuse to nearly rounded, often

oblique, in terminal leaflet cuneate, main veins 9—12 pairs, departing from the midrib

at an angle of 55—70 degrees, ultimate veinlets usually rather conspicuous (10x), rather

loosely reticulate, margin entire, apex acuminate, the acumen 4—8 mm long. Inflore-

scences 11—18 cm long, rachis and branches rather sparsely to densely pubescent, pedi-

cels sparsely pubescent, 1.5—2 mm long. Flowers predominantly 5-merous (occasional

flowers 4-merous); sepals sparsely pubescent, about 0.5 mm long; petals green or greenish

—yellow, drying brown, abaxially sparsely puberulent to glabrous, adaxially sparsely to

densely villous, 2.5—3 mm long; disc glabrous; ovary glabrous, the carpels connate toward

the base, ovule 1 per carpel; rudimentary carpels villous toward the base, otherwise

glabrous, connate toward the base. Fruits predominantly 5-carpellate (occasional fruits

4-carpellate), 1—5 of the carpels developing into follicles; follicles glabrous or rarely with

a few scattered hairs, trigonous or subtrigonous, not beaked, 3.5 mm long and about as

wide, connate toward the base, endocarp glabrous. Seed black, 1 per follicle, ellipsoid to

subreniform, 2.3—2.5 mm long, attached in the dehisced follicle to an adaxial strip of

pergamentaceous endocarp tissue; outer testa spongy (bounded externally by a crusta-

ceous, shiny pellicle); inner testa bony.

Hartley: Tetradium (Rutaceae) 121

Distribution. Northern Vietnam and adjacent southwestern China; recorded from

forests (apparently well-drained); 360—1500 m.

Vietnam. Tonkin: Chapa, Pételot 4892 (NY, US); Chapa prope Laokai, Fenzel 50

(W); Route de Chapa a Laokai, Pételot 4375 (NY).

China. Kwangsi Province: Wang-tung, North Luchen, near Shan-fang, Ching 6214

(W, holotype of Euodia austrosinensis Hand.-Mazz.; A, NY, isotypes); Yung [Jung] Hsien,

Ta Tseh Tsuen, Steward & Cheo 878 (A, BO, NY, W); North Lin-yen, Tsin-lung Shan,

Ching 7026 (NY). Kwangtung Province: Lung-t’au Mt., near lu, To, Tsang, & Tsang CCC

12483 (A, K, NY, UC, US, W); Taai-yeung Shan, McClure 6556 (A, K, NY).

9. Tetradium sumatranum Hartley, sp. nov.

Arbor mediocris vel magna usque 42 m alta; ramulis puberulis; foliis (19—) 26—48

(—55) cm longis; rhachidi puberula; foliolis in paribus (3—)4—6 (—7); petiolulis folio-

lorum lateralium obsoletis vel usque 3 mm longis, rhachidi ad apicem extensa 8 — 20 mm

longa foliolum terminale ferente; laminis chartaceis vel subcoriaceis, sparse et incon-

spicue pelluciodo-punctatis vel sine punctatione distincta, subtus glaucis, minute pa-

pillatis, minute adpresse puberulis, supra praeter costa puberulis glabris, elliptics, ellip-

tico-oblongis vel interdum late lanceolatis, 6.5—13.5 cm longis, 3.5—5.5 cm latis, basi

obtusis vel rotundatis et plerumque inaequilateralis in foliolis lateralibus, actuis vel

cuneatis in foliolo terminali, venis primariis utrinsecus costa 13—16, sub angulo 55 — 70

abeuntibus, venulis ultimis inconspicuis, margine integris, apice acuminatis, acumine

7—10 mm longo; inflorescentiis 10—17.5 cm longis, rhachidi et ramis puberulis vel

subtiliter pubescentibus, pedicellis subtiliter pubescentibus, 0.5—2 mm longis; floribus

pentameris; sepalis dense adpresse pubescentibus, ca. 0.5 mm longis; petalis viridulis,

in sicco brunneis, abaxialiter subtiliter adpresse pubescentibus vel fere glabris, adaxia-

liter villosis, 3.2—4.5 mm longis; disco glabro; ovario aequaliter vel inaequaliter adpresse

pubescenti, carpellis basin versus connatis, ovulo in quoque carpello solitario; carpellis

rudimentariis basin versus villosis, aliter glabris, basin versin versus connatis,; fructibus

non visis.

Holotypus: Krukoff 4248 (NY).

Medium to large tree to 42 m high; branchlets puberulent. Leaves (19—)26—48(—5S5)

cm long; rachis puberulent; leaflets (3—)4—6(—7) pairs; petiolules of lateral leaflets

obsolete to 3 mm long, terminal leaflet on an extension of the rachis 8—20 mm long;

blades of leaflets chartaceous to subcoriaceous, sparsely and inconspicuously oil-dotted

or without evident oil dots, below glaucous, minutely (10x) papillate, puberulent with

minute, appressed hairs, above glabrous except for puberulent midrib, elliptic, elliptic-

oblong, or occasionally broadly lanceolate, 6.5—13.5 cm long, 3.5—5.5 cm wide, base in

lateral leaflets obtuse to rounded, usually oblique, in terminal leaflet acute to cuneate,

main veins 13—16 pairs, departing from the midrib at an angle of 5S—70 degrees, ultimate

veinlets inconspicuous, margin entire, apex acuminate, the acumen 7—10 mm long. In-

florescences 10—17.5 cm long, axis and branches puberulent to finely pubescent, pedicels

finely pubescent, 0.5—2 mm long. Flowers 5-merous; sepals densely appressed-pubescent,

about 0.5 mm long; petals greenish, drying brown, abaxially finely appressed-pubescent

or nearly glabrous, adaxially villous, 3.2—4.5 mm long; disc glabrous; ovary evenly or

unevenly appressed pubescent, the carpels connate toward the base, ovule 1 per carpel;

rudimentary carpels villous toward the base, otherwise glabrous, connate toward the base.

Fruits not seen.

Distribution. Endemic to Sumatra; recorded from well-drained, primary rain forest;

400 m.

122 Gard. Bull. Sing. 34:1 (1981)

Sumatra. East Coast: Pematangsiantar, Lorzing 16614 (L), 16938 (L); Asahan,

Masihi Forest Reserve, Krukoff 4248 (NY, holotype; A, BRI, L, NY, SING, US, iso-

types); Asahan, Pargambiran, Rahmat Si Boeea 5783 (L).

INSUFFICIENTLY KNOWN OR OTHERWISE EXCLUDED NAMES

Boymia martinicensis (Lam.) G. Don, Gen. Hist. 1 (1831) 805 [based on Fagara

martinicensis Lam., Tabl. Encycl. 1 (1792) 334] = Zanthoxylum martinicense

(Lam.) DC., Prodr. 1(1824) 726.

Evodia? chaffanjoni Lév\. in Feddes, Repert. Spec. Nov. Regni. Veg. 13 (1914) 265

= Euscaphis japonica (Thunb.) Dippel according to Rehder, J. Arnold Arb. 15

(1934) 2.

Evodia meliaefolia var. celebica Koorders, Versl. Minahasa (1898) 371; Koorders-

Schumacher, Syst. Verzeich. 3 (1914) 59; Koorders, Suppl. Fl. Celebes 3 (1922)

12, t 22. Syntypes: Koorders 187548, 187558, and 187576, Celebes, Prov.

Minahassa. The only fertile material I have seen of this taxon is staminate. While

undoubtedly belonging in Tetradium, its further identity cannot be determined

with certainty without seeing carpellate material. The lower surface of the

leaflets is minutely papillate, as in 7. austrosinense and T. sumatranum, and, to a

lesser extent, in some specimens of 7. fraxinifolium. It seems probable that it is

most closely related to 7. sumatranum. Relevant collections in the Index to

Exsiccatae are identified as “aff. 9” (Tetradium sp. aff. sumatranum).

Evodia mollicoma Hu & Chen, Acta Phytotax. Sin. 1 (1951) 225 = Zanthoxylum

molle Rehd. according to Huang, Acta Phytotax. Sin. 6 (1957) 43.

Evodia odorata Lévl. in Feddes, Repert. Spec. Nov. Regni. Veg. 9 (1911) 458

[Zanthoxylum odoratum (Levl.) Levi. ibid. 13 (1914) 266] = Zanthoxylum

rhetsoides Drake according to Huang, Acta Phytotax. Sin. 6 (1957) 47.

-Evodia parviflora Craib, Kew Bull. (1915) 425 = Turpinia parviflora (Craib) Craib,

Fl. Siam Enum. I (1926) 339.

Evodia tonkinensis Engl. in Engler & Prantl, Nat. Pflanzenfam. III: 4 (1896) 121 (in

key; no specimen cited) = Evodia ailantifolia Pierre according to Engler, ibid.

ed. 2, 19a (1931) 228. Probably = Tetradium glabrifolium (Champ. ex Benth.)

Hartley.

Tetradium amarissimum (Lour.) Poir.,Encycl.Suppl. 5 (1817) 291 [based on Gonus

amarissimus Lour., Fl. Cochinch. (1790) 658 | = Brucea javanica (L.) Merr.

according to Nooteboom, FI. Males. ser. 1: 6 (1962) 211.

3. The Euodia names included here are only those representing plants that belong in Tetradium or

could be confused with it (in having pinnately compound leaves). Euodia, in the sense of its type

species and as interpreted in the present study, has unifoliolate or trifoliolate leaves.

Hartley: Tetradium (Rutaceae) 123

ACKNOWLEDGEMENTS

I wish to thank the directors and curators of the herbaria listed in the introduction

for making specimens in their care available to me. Thanks are also extended to Roy

Pullen, C.S.I.R.O. Division of Plant Industry, who translated into English portions of

Huang’s paper on Chinese Rutaceae, and to Rod Henderson, the Australian Botanical

Liaison Officer at Kew in 1979, who assisted in having the type of Tetradium trichoto-

mum loaned to me.

LITERATURE CITED

Bentham, G., and J. D. Hooker (1862). Gen. Pl. 1, 296.

DeCandolle, A. P. (1825). Prodr. 2, 88.

Endlicher, S. L. (1840). Gen. Pl., 1149.

Engler, A. (1896). Rutaceae. In Engler & Prantl, Nat. Pflanzenfam. III: 4, 95—201.

. (1931). Rutaceae. Nat. Pflanzenfam. ed. 2. 19a, 187—358.

Hartley, T. G. (1966). A Revision of the Malesian Species of Zanthoxylum (Rutaceae). J. Arnold

Arb. 47, 171—221.

(1974). A Revision of the Genus Acronychia (Rutaceae). J. Arnold Arb. 55, 469—523,

§25—567.

Hegnauer, R. (1973). Rutaceae. Chemotaxonomie der Pflanzen 6, 174—239. Basel. Birkhauser.

Huang, C. C. (1957). Preliminary Study on Chinese Rutaceae (1) [in Chinese]. Acta Phytotax. Sinica

6. 1—143:

Jussieu, A. (1825). Mem. Rutac., 138.

Price, J. R. (1963). The Distribution of Alkaloids in the Rutaceae. In: Chemical Plant Taxonomy,

T. Swain, Editor, 429—452. Academic Press, London and New York.

Rehder, A. and E. H. Wilson (1914). Rutaceae. In Sargent, Pl. Wils. 2, 121—151.

Siebold, P. F. and J. G. Zuccarini (1837 or 1838). Fl. Jap. 1, 50.

Sprengel, K. (1824). Syst. Veg. 1, 441.

Waterman, P. G. (1975). Alkaloids of the Rutaceae: Their Distribution and Systematic Significance.

Biochem. Syst. & Ecol. 3, 149—180.

124 Gard. Bull. Sing. 34:1 (1981)

INDEX TO COLLECTIONS

The numbers in parentheses refer to the corresponding species in the text.

Achmad 607, 775, 1022, 1142, 1662, 1736 (1).

Anonymous 4586 (5); 5537, 6026, C6858 (3).

Backer 25786, 25908 (1).

Balansa 117] (5); 3669, 4038 (6); 4042 (5); 4043 (6).

Bartlett 6059 (7).

Biswas 3724 (5); 4060 (2).

Bock & Rosthorn 1884 (3).

Bodinier 1689 bis (7).

de Boer 25 (1).

Boom 8456, 10347, 13136c, 30080, 34597 (3).

Bosschen buitengewesten series (bb), Netherlands Indies Forest Service; the follwoing

by anonymous collectors: bb 6536, bb 6621, bb 7421, bb 13104 (1); bb 17104,

bb 17158, bb 21321 (aff.9); bb 22358, bb 22426 (1); bb 24508, bb 24988,

bb 26271, bb 26282, bb 28232 (aff. 9).

Brandis 1855 (2).

Bretschneider 152, 1328 (3).

Brumbach 7238, 7288 (3).

Bullock 10 (7).

Canton Christian College series (CCC); the following by anonymous collectors:

CCC 12776, CCC STIID- Cy .

Carles 107 (3).

Celebes series, Forest Research Institute, Buitenzorg; the following by an anonymous

collector: Cel II-467 (aff. 9).

Chakrabutti 132D (2).

Champion 383 (5).

Chand 5377 (2); 6002 (5); 6160, 8294 (2).

Chang, C. E., 2418 (5); 4882 (7).

Hartley: Tetradium (Rutaceae) 125

Chane, RB. 5102 (7).

Cheo 3, 169, Univ. Nanking 18389 (5).

Cheo & Wilson Univ. Nanking 12807 (7).

Chiao 2708, 2925, 2926, 2942 (3); Univ. Nanking 14748 (5); Univ. Nanking 18585,

Univ. Nanking 18618 (7); Univ. Nanking 18766 (5).

Chien 1029 (7); 1129 (5); 6055 (7). 3

Ching 1805, 2163 (5); 2493 (7) 2559, 2773 (5); 2779B (7); 3098 (5); 3985 (3); 4908,

5102 (7); 6214 (8); 6306 (4); 6894 (7); 7010 (6); 7026 (8); 7644, 7679, 7950,

53173):

Chow 714, 1714 (7).

Chu, K, L., 3419 (3); 3997 (6).

Chu, V. M., 29 (5).

Chun 7477 (5).

Chun, N. K., 40010 (5).

Chun, W. Y., 3099 (5); 3783, leg. Huang 3969, Univ. Nanking 4129, Univ. Nanking 4330

(3); 5225, Univ. Nanking 5687, Univ. Nanking 5766, Univ. Nanking 5899, 6562,

6864, Univ. Nanking 7056, Univ. Nanking 7121 (5).

Chun & Tso 43564, 43761, 43795, 44010 (5); 44140 (6).

Chung, H. H., 28, 31, 607, 693 (7); 1050 (5); 4382 (7).

Chung, T. H., 1169 (3).

Chung & Sun 255, 317, 329 (5); 614 (7); 685 (5).

Clarke 7312, 26635B, 27327, 35229C, 44335, 45921B (2).

Clemens, J. & M.S., 3286 (5).

Conklin & del Rosario PNH 72715 (5).

Cooper 3048 (7); 4473 (2).

Delavay 4526 (3).

Diepenhorst HB 1399, HB 2560, HB 2917 (1).

Division Forest Officer 1384 (5).

Dunn /12 (7).

126 Gard. Bull. Sing. 34:1 (1981)

Elmer 10120 (5).

Fabet26, 95,293, 1717 (7).

Fan & Li 117, 180, 190 (7); 200, 377, 556 (S).

Fang 1119, 2412 (7); 3376 (6); 7444 (7); 7855 (3); 8801, 12553 (7).

Farges 49, 413, 1284 (3).

Faurie 34 (5); 95 (7); 468 (3).

Fenzel 24 (5); 50 (8); 222 (5); 683 (3).

Herb. Forbes 68 (5).

Forrest 7592 (7); 8687 (2); 11091 (3); 11668, 11849 (7); 12954 (5); 13468 (7); 14299,

- 14772 (3); 15105 (7); 15803, 16079 (2); 16484, 16926 (3); 17743, 17859, 18592

(2); 21513 (7); 22294 (2); 24115, 25070, 25250, 26470 (2); 27966 (7).

Garrett 644 (2).

Gibbs Herb. Hong Kong 7456 (5).

Gressitt 1061, 1118, 1186 (6).

Haines BB 873 (5).

Herb: Hancets 171.65).

Handel-Mazzetti 9393 (2); 9605, 10440, 10690, 12320, 12564 (7).

Hao 3508 (3).

Hatusima 14525, 18236, 18241 (5).

Henry 13, 831 (5); 924 (7); 932, 974, 1296, 1562 (5); 1676, 2077, 2259(7), 2595 ar

2816 (7); 2939, 4482 (7); 4525 (7); 4534, 4534A (3); 4577 (5); 6136, 6136A (7);

6157 (5); 6199, 6199A (7); 6485 (3); 6569 (7); 6712 (3); 8605 (6); 8706 (5); 10245

(7); 10951, 12137, 12137A, 12137B, 12137C, 12237, 122374, 1224798

Hers 543, 739,'1157 (3).

Hiroe 247] (5).

Hornby 4276, 6898 (3).

How 71094 (5); 71981, 72442 (6); 72749, 73189 (5); 73470 (6).

How & Chun 70139 (5).

Hsiung 5315, 5723 (7); 5726 (5).

Hartley: Tetradium (Rutaceae) 127

Pri tt.,306 (5).

me £0007, 5596 (5).

Rev. Fr. Hugh [Scallan] 75 (3).

Hwa 57 (5); 83 (7).

Ip. Univ. Nanking 1083 (7).

Kanai, Murata, & Togashi 6301109 (2).

Kanjilal 2634, 4553 (2); 4969, 5081 (5); 7048 (7).

Kawakami & Mori 3196 (5); 3397 (7).

Keng, Y. L., 47, 96, 631 (7); 914, 921, 1134 (5S).

Kepong series (KEP), Malaya Forestry Department; the following by an anonymous

collector: KEP 78904 (5).

Kermode / 7346 (2).

Kerr 644 (2); 4762, 11438A, 13035 (5).

King 4868 (2).

Kingdon-Ward 9496 (7); 13088, 21150 (2).

Ko 52936 (7); 55618 (4).

Koelz 23190, 25374, 26051, 26118, 27485, 28383, 29441, 30358 (2); 30662 (5):

33721 (2).

Foo 2033 (6).

Koorders 69088, 69098, 69108, 69116 (1); 104106 (5); 110966 (1); 187548, 187556,

187576 (aff. 9); 241568, 26356, 37906B (1).

Kostermans /871]6a, 18864 (1).

Krukoff 4248 (9).

Kuan & Wang 3333 (6).

Kung 305] (3).

Kurata & Nakaike 2302 (5).

Lace 715 (5); 2205 (2); 5847 (5).

Lamont /16 (5).

128 Gard. Bull. Sing. 34:1 (1981)

Lau, S. K., 1930 (5); 2462, 4128 (7); 4780, 5346, 27468 (5).

150, ¥./' 3, 9900)

Law 1277, 1339 (5); 1347, 1389 (7).

Lee 3050 (7).

Lei 9, 832 (5).

Lesger 255 (1).

Levine CCC 1143, CCC 1775, leg. Ah To CCC 1854, CCC 3244 (5S).

Li 10032 (3).

Liang 62250, 62345, 63026, 63392, 64634, 65135, 69916, 70120 (5).

Licent 13367 (3).

Ling Univ. Nanking 7732, Univ. Nanking 9623, Univ. Nanking 12447 (7).

Lingnan University Herbarium: the following by an anonymous collector: Lingnan Univ.

9933 (7).

Liou 380 (7); 720 (5).

bi 1225,. 2217 (3):

Lorzing 4351, 5282 (1); 16614, 16938 (9).

Maire 247 (7); 3361 (3).

Maradjo 25 (1).

Maung Ba Pe 1576 (5).

McCaskill & Reagan 684 (3).

McClure 6556 (8); CCC 7751 (5); CCC 8449 (6); CCC 13300 (5); Lingnan Univ. 15232

(7); Lingnan Univ. 18274, Lingnan Univ. 19709 (5).

Merrill, E. D., 10773 (5).

Meyer, F.N., 262, 278, 1929 (3).

Muroi 4985, 6942 (7).

Oldham 130 (5); 131 (7).

Panigrahi 22427 (2).

Petelot 1165, 2483, 2599, 3124 (6); 4375, 4892 (8); 5692 (6).

Hartley: Tetradium (Rutaceae)

Phusomsaeng et al. BKF 46522 (5).

Pierot 28] (7).

Herb. Pierre 3862 (5).

Poilane 12927 (2); 25989 (6); 26800 (2); 29658 (6).

Polunin, Sykes, & Williams 483 (7).

Prain’s Collector 28] (7); 953 (2).

Purdom /038 (3).

Rahmat Si Boeea 5783 (9).

Read 1204 (7).

Ren 6/22 (3).

Ren & Tao Univ. Nanking 12579 (3).

Rock 1778, 2391 (6); 6235 (7); 10270 (3); 10610 (7).

Rodger 139 (2).

Sangkhachand BKF 36894 (5).

Santos BS 31968 (5).

Sasaki 42 (7).

Sastry 40950 (2).

Sato 399 (7).

Schneider 1663 (3).

Silvestri 12/8 (5).

Smith, A., 720 (5).

Smith, H., 6445 (3).

Smith & Cave 2620 (7).

Sonohara 8, 67 (5).

Sorensen 6672 (3).

Stainton 3685, 4546, 6838 (2).

Stainton, Sykes, & Williams 2930, 3549 (7): 5064, 6679 (2).

129

130

Steward Univ. Nanking 4668 (5).

Steward & Cheo 463, 703 (7); 878 (8); 1016 (7).

Steward, Chiao, & Cheo //5 (7); 329 (5).

Sun, C. L., 687 (3).

Sun, S. C., 1202 (5); 1280 (7); 1300 (5).

Suzuki-Tokio 11705 (7).

Sykes CHR 124917, CHR 124998A, CHR 124998B (3).

Taam 694, 797 (7).

Tanaka 468 (7).

ane 257 (3).

Ten271,-3 96.7):

Theunissen 2 (1).

To & Tsang-12776 (7).

To, Tsang, & Tsang CCC 12483 (8).

Treuther 139 (2).

Tsai 61062, 62045 (6).

Tsang, W. T., Lingnan Univ. 9938, Lingnan Univ. 15585, Lingnan Univ. 16247, Lingnan

Univ. 16637 (5); 23111 (7); 24189, 25921 (5); 26637 (6); 26685 (5); 26720, 27026,

29300, 29454, 30357 (6); 30475 (5).

Tsang & Fung Lingnan Univ. 18218 (6); Lingnan Univ. 18274 (5).

Tsang, Tang, & Fung Lingnan Univ. 17744 (6).

Tsang & Wong CCC 14188, Lingnan Univ. 14845 (7)...

Tsiang 1017, 1643, 1720 (5); 4919, 5447, 5870, 8655 (7); 9567, 10352 (5); 10406,

10419 (7); 10718B (5).

Tso 20688, 20886 (7).

Uno 2598 (3).

Walker & Tawada 6837 (5).

Wallich Cat. No. 8521 (2).

Wang, C., 33111, 34072, 35680, 36445 (5).

Gard. Bull. Sing. 34:1 (1981)

Hartley: Tetradium (Rutaceae) 131

Wang, C. W., 73987, 74237, 74893, 75111, 75116 (6); 75621, 75796 (5); 75835, 75942,

77145, 77294 (6); 77601, 77770, 77809, 77841, 77859, 77955 (5); 77980 (6);

78669 (5); 79663 (6); 80688 (5).

Wang, F. T., 21610 (3); 22187 (6); 22511, 23163 (7).

Wang, T. P., 1518 (3).

Whitmore FRI 12139 (1).

Williams 7019 (2).

Williams & Stainton 8465 (2).

Wilson 3 (3); 247 (5); 251 (7); 253 (5); 324 (3); 366, 366a (7); 381 (5); 387, 387a,

387b (3); 791 (6); 882 (7); 994 (3); 1164 — Oct 1908 (7); 1164 — July 1908 (3):

1309 (7); 1583, 1584, 1585, 1930, 1930a, 1930b (5); 2183, 2183a (3); 2210,

2210a (5); 2212 (7); 2322 (5); 2415 (3); 2415a, 2415b (5); 2578 (6); 2693, 2693a,

Peete S37 SDSL (35). 35/4, JJ7) (6); 3974, 3575, 3576. 3); 3577 (7);

3579, 3579a (5); 4568 (6); 4628 (3); 4772 (7); 6112 (5); 8800 (3); 9955 (7): 9971

(5); 10555, 10568, 10612 (3); 10880 (7).

Wissmann /20 (6).

Yu 19451, 20510 (2).

132

THE FERN GENUS TECTARIA CAV.IN MALAYA

R. E. Holttum

Royal Botanic Gardens, Kew

SUMMARY

A new key to the species of Tectaria in Malaya is presented, with information sup-

plementary to that in Holttum, Ferns of Malaya (1955; second edition, with Appendix

Il, 1968). Earlier names for six species described in 1955 are cited, with a fuller synony-

my, and six additional species are described, namely T. herpetocaulos Hottum, T. melano-

caula (B1.) Copel., T. decurrens (Presl) Copel., T. tricuspics (Bedd.) Copel., T. simonsii

(Bedd.) Ching, 7. fauriei Tagawa, and T. cherasica Holttum sp. nov. Six of these addi-

tional species are still only known in Malaya from single collections.

INTRODUCTION

The present paper is compiled from information gathered during a recent attempt

to survey all species of Tectaria described from specimens originating in Asia, Malesia,

and the Pacific, based mainly on specimens and literature available at Kew, also on a

study of Presl’s herbarium at Prague (1967) and on notes made in the herbaria at Singa-

pore and Bogor in 1978.

The material is presented as an addition to that recorded in Holttum, Ferns of

Malaya 1955 (issued in January 1955 though the printed date is 1954). Earlier names

have been discovered for six of the species described in 1955 and much additional in-

formation on synonymy. Descriptions published in works on the floras of neighbouring

countries are cited, including fuller references to Beddome who included Malaya in

his Handbook to the ferns of India (1883 and Supplement 1892); one species named

and described by Beddome in 1892 (T. tricuspis) was overlooked when my book of

1955 was prepared because the type collection is not represented in the Singapore her-

barium. A fuller statement on known distribution is provided for all the species dealt

with in 1955.

A second edition of my book was published in 1968 (the printed date is 1966)

containing, in Appendix II (pp 629-638), further information, especially concerning

additional species discovered by Mrs. Betty Molesworth Allen; three of these additional

species belong to the genus Tectaria, namely T. decurrens, T. herpetocaulos, and T. me-

lanocaula, These three species, briefly characterized in 1968, are here more fully des-

cribed, also a further species (7. simonsii) also discovered by Mrs. Allen, another (T.che-

rasica) recently collected by Mrs. A.G. Piggott in Pahang, the forgotten species named by

Beddome, and another species, 7. fauriei, which I also overlooked in preparing my

book. A new key covering all species now known in Malaya is provided.

Of the additional species, only 7. decurrens is represented by several earlier col-

lections made in Malaya (all in Perak). 7. herpetocaulos, T. melanocaula, and T. simonsii

are still only knwon in Malaya from specimens collected in the forest near the road to

the Cameron Highlands at altitudes between 2000 and 3000 feet. So far as I know

T. tricuspis has not been collected again in Malaya since Kunstler found it nearly one

hundred years ago, though specimens (given another name) have since been found in i

western Sarawak. Mrs. Piggott’s specimen from limestone on Bukit Cheras in Pahang

Holttum: Tectaria in Malaya 133

appears to be unique. Under 7. amplifolia is an indication that there is perhaps a dis-

tinct allied species on limestone in Pahang and Kelantan still not named. All this shows

how much more there is to be known about Tectaria in Malaya. More field work by

trained observers is needed.

The drawings published in Holttum 1955 show the kind of details which need

to be observed when learning to recognize these plants in the field. For most purposes

a hand lens is sufficient, though to see indusia clearly a higher magnification is necessary.

Indusia shrivel on old sori and young fronds should be looked for which show them

more clearly. Scales at the base of stipes may yield distinctive characters, but many

herbarium specimens do not show them. Buds at the bases of pinnae or at the bases of

some main veins in the terminal lamina only develop on older fronds (in T. siifolia

apparently only on sterile ones), so it is important to look for these.

There is no clear indication of the occurrence of natural hybrids in Malaya, though

such may occur; one is recorded in Ceylon. Experimental hybridization might yield

interesting results. Of six species examined cytologically by Prof. Manton from Malayan

plants, only one (7. barberi) proved to be tetraploid (see Holttum 1955, p.626) but a

tetraploid form of T. devexa has been found in New Guinea.

Synonymy. The synonyms are arranged in paragraphs, in each of which all names

are dependent on one type specimen, the type and its location being stated at the end

of the paragraph. In most cases I have seen holotypes or isotypes; those which I have

not seen are so marked. In the case of Fée, whose specimens were dispersed, I have seen

none of those which he himself studied, but in most cases here dealt with he cited collec-

tions which are represented by isotypes at Kew, and from his descriptions and his excel-

lent figures there is no doubt as to his meaning. I have examined microfiche photo-

graphs of Willdenow’s types; these give adequate information for the species here dealt

with. Some authors in the 19th century attempted to distinguish species which have

peltate indusia from those with reniform indusia; the latter were placed ina genus Sagenia.

Most species have more or less distinctly reniform indusia and it does not appear that

those with truly peltate indusia form a natural group, so that the genus Sagenia is now

not recognized. Fée and Presl both attempted to distinguish genera on vein patterns;

it is possible that some of these names may be useful for subdivisions of the genus, but

some are not clearly defined and all are generally now disregarded.

KEY TO THE MALAYAN SPECIES OF TECTARIA

DE ADV ee ORE we RE, ON eng ee 1. T.ingens.

1. Veins anastomosing, at least to form costal areoles

2. Veins forming areoles along costae and a variable number of other areoles;

free included veinlets in areoles few except in 7. barberi, none in costal areoles

3. Lamina hairy between veins on upper surface

4. Veins anastomosing in costal and costular areoles only... .2. 7.devexa

4. Veins more amply anastomosing ............... 3.7. coadunata

134 Gard. Bull. Sing. 34:1 (1981)

3. Lamina not hairy between veins on upper surface

5. Sori in a single row on each side of costae or costules; indusia

distinct

6. Fertile fronds much contracted; lamina to 25 cmlong ......

ve th a ak ep douse gee ge ween aie ag 4. T. brachiata

6. Fertile fronds not or little contracted, to60cmlong.......

re RR Ree ER 5. T. griffithii

5. Sori scattered, small, with inconspicuous indusia ... 6. T. barberi

2. Veins forming many areoles, the largest ones with included free veinlets, often

branched; free veinlets also present in costal areoles

7. --Fronds simple; entire’. <<"... 22272, 2 eee ee 7. T. singaporeana

7. Fronds deeply pinnatifid or pinnate or bipinnate

8. Sori small, very irregularly arranged and shaped, some much elongate

wd sok aoe Eg DG A a 8. T. semipinnata

8. Sori uniform or nearly so, none greatly elongate

9. Fronds pinnatifid to a wing along the rachis; stipe also winged

10. Lower lobes of frond much smaller than middle lobes

fh be bale SUG VRE. hie ele SAL Sy eee 9. T. grandidentata

10. Lower lobes of frond not reduced

11. Sori in two rows between main veins, immersed .......

BE, NA An A Aer hcg ce eee 10. 7: decurrens

11. Sori in more than two rows, not immersed

12. Indusia conspicuous, persistent, hairy on upper suface

ope ah Deeg tae at ete 11.7. tricuspis

12. Indusia small, caducous, not hairy .... 12.T. vasta

9. Fronds of mature plants with at least one pair of free pinnae

13. Sori in one row on each side of main veins, sometimes with additional

ones where pinnae are deeply lobed

14. Sori large, mostly on free veins in areoles

15. Basal pinnae lacking free pinnules..., . a5 see 13. T. crenata

15. Basal pinnae of well-grown plants bearing at least one pair of

free pinnules

Holttum: Tectaria in Malaya 135

16. Fronds dimorphous; lower surface quite glabrous .........

er ae a Se aie TSM 4.T. brachiata

16. Fronds little or not dimorphous; lower surface of costae

emer ee geet ee. 14. T. cherasica

14. Sori not on free veinlets in areoles, mostly small

13.

17. Fronds dimorphous; indusia inconspicuous; buds present at bases of sterile

LL Ec ea heerlen erat be aie a a ig lh eee eh a 15. T. siifolia

17. Fronds not dimorphous; indusia conspicuous; no buds... 16. T. amplifolia

Sori scattered or in several rows between main veins

18. Pinnae and pinnules c. 10 times as long as wide ..... 17. T. semibipinnata

18. Pinnae and pinnules proportionately much wider

19,

ty:

Stipe and rachis almost black; no buds at base of veins on apical lamina

20. Pinnae deeply lobed; sori indusiate......... 18.7. melanocaula

20--Piinae not lobed: Son exmdusiate os os 19. T. simonsii

Stipe and rachis not black; or, if black, buds present at bases of veins

of apical lamina

21. Middle pinnae broadly rounded at base on basiscopic side

22. Caudex erect; buds present at bases of some veins on apical

CEL ULE oe Sei, Intel ifie i les call sa abe in Sa mai A 20. 7. angulata

22. Caudex long-creeping; no buds on apical lamina ..........

on Ee Frere LR ee 21.7. herpetocaulos

21. Middle pinnae narrowly cuneate on both sides at base

23. Pinnae not decurrent at their bases to form rachis-wings;

no buds present at bases of pinnae ....... 22. T. oligophylla

23. Pinnae decurrent to form narrow rachis-wings which almost

reach the next lower pinnae; buds present at the bases of

pinnae ae beer eee ee 2. 25. 7. fauriel

1. Tectaria ingens (Atkinson) Holttum 1955: 503, fig.296. Nephrodium ingens Atkin-

son ex Clarke, Trans. Linn. Soc. II Bot. 1 (1880) 526, pl. 73.

Type: C. B. Clarke 9295, Darjeeling (K).

Nephrodium setulosum Bak., Journ. Bot. 28 (1890) 265 .—Ctenitopsis setulosa (Bak.)

C. Chr., Notul. Syst. 7 (1938) 87; Ching, Bull. Fan Mem. Inst. Biol. Bot. 8 (1938) 321;

Tard. & C. Chr. in Fl. Gen. Indochine 7: 2 (1941) 352. — Type: Balansa 1836, Tonkin

(K, isotype).

136 Gard. Bull. Sing. 34:1 (1981)

Aspidium cardieri Christ, Journ. de Bot. 19(1905)62.—Tectaria cadieri (Christ) C.

Chr., Ind. Fil. Suppl. III (1934) 177. — Type: Cadiere 85, Annam, valley of river Song

Gianh, 19° 17’ N. (K, isotype).

Dryopteris raciborskii v.A.v.R., Handb. Malayan Ferns (1908) 197,816,new name for

Aspidium sagenioides sensu Racib. F1. Btzg.1(1898)179, non Mett.—Type: Raciborski

s.n., G. Salak, Java (BO; isotype K).

Tectaria matthewii Ching, Bull. Fan Mem. Inst. Biol. Bot. 2(1931)199, t.13.—Ctenit-

opsis matthewii Ching, ibid. 8 (1938) 319. — Type: C. G. Matthew s.n. 12 Dec. 1907,

Kwangtung, Lienchow, Mong-si Hap, in a ravine in deep shade (K).

In Malaya this species is still only known from two collections made on Gunong

Brinchang, Cameron Highlands, at about 1500 m, in the forest. I believe that the types

of all the above cited species are conspecific with Clarke’s original; they indicate a wide

distribution but, except near Darjeeling, few specimens apart from the types have been

collected. As it occurs in Java, it may be expected also on mountains in Sumatra.

Dr W.A. Sledge (Kew Bull. 27 (1972) 415), stated that in his opinion this species

does not differ significantly from Tectaria paradoxa (Fée) Sledge of Sri Lanka except

in its larger size. But the two differ also in the character of the scales at the bases of

stipes; in JT. ingens these are more than 10 mm long, their marginal cells not thin-walled,

but in 7. paradoxa the scales are shorter and have a distinct margin of thin-walled cells.

Dr Sledge also included T. fuscipes (Wall.) C. Chr. as a synonym of T. paradoxa, in

spite of Clarke’s comments on its distinctive characters. I agree with Clarke that its

very dark scales and narrower fronds are sufficient to establish its distinctness. It is

distributed from Sikkim to southern Burma (also in Thailand, Vietnam, and Hainan)

and might occur in northern Malaya.

2. Tectaria devexa (Kunze ex Mett.) Copel., Philip. J. Sci. 2C (1907) 475; Holttum

1955: 505, fig. 297. — Aspidium devexum Kunze, Bot. Zeit. 6 (1848) 255, nom. nud.;

Kunze ex Mett., Ann. Mus. Bot. Lugd. — Bat. 1 (1864) 237, in obs. — Aspidium inter-

medium Mett., Farngatt. IV (1858) 119, non Willd. 1810. — Type: Zollinger 2717,

Java (LZ, destroyed; isotype not seen).

Aspidium giganteum var. minor Hook., Spec. Fil. 4(1862)50.—Sagenia gigantea var.

minor Bedd., Ferns S. India (1864) 81, t. 243. — Type: Thwaites CP. 1358, Ceylon

(K; BM). )

Aspidium membranaceum Hook., Spec. Fil. 5(1864)105 .—Pleocnemia membranacea

(Hook.) Bedd., Ferns Br. India Suppl. (1876) 15. — Lectotype (selected here): Cuming

s.n. (wrongly labelled 277), Philippines (K).

Christensen regarded Aspidium devexum Kunze 1848 as a valid name, but it was

not. The first description of the species was published by Mettenius in 1858 under the

name Aspidium intermedium which was illegitimate, being a later homonym; he cited

A, devexum as a synonym. In 1864 however he evidently recognized the error and stat-

ed that the species described was A. devexum Kunze, which name thus acquired validity.

But in 1864 Hooker also redescribed the same species, mainly from a Philippine speci-

men, under the name A. membranaceum. As it is uncertain which name was earlier,

I prefer to retain the epithet devexa, which has been current for many years.

Holttum: Tectaria in Malaya Ka

3. Tectaria coadunata (J.Sm.) C. Chr., Contr. U.S.Nat. Herb. 26 (1931) 331; Ching,

Sinensia 2(1931) 18, pl. 1 & 2; Sledge, Kew Bull. 27 (1972) 418. — Sagenia coadunata

J.Sm. in Hook. J. Bot. 4 (1841) 184, new name for Aspidium coadunatum Hook. &

Grev., Ic. Fil. (1831) t. 202, non Kaulf. 1824. — Sagenia macrodonta Fée, Gen. Fil.

(1852) 313, t. 24, nom. nov. superfl.. — Tectaria macrodonta (Fée) C. Chr., Ind. Fil.

Suppl. III (1934) 181; Holttum 1955: 505. — Type: Wallich 377, Nepal (K).

Aspidium pinfaense Christ, Bull. Geogr. Bot. Mans.20(1909) Mem. 169.—Type: Cava-

lerie 872, Kweichow, Pin-fa (K, isotype).

This species is widely distributed from N.E. India to S.W. China and southwards

to Burma and Thailand. As noted in Holttum (1955) there are only a few small specimens

from the north of Malaya. Those at Kew were growing on limestone, which might

account for their small size: Henderson 21383 (Langkawi) and Kiah 35315 (Perlis).

Christ identified Philippine specimens with this species (Philip. J. Sci. 2C: 187) but they

are distinct and were later named T. christii Copel.

4. Tectaria brachiata (Zoll. & Mor.) Morton, Contr. U.S. Nat. Herb. 38 (1973) 217. —

Aspidium brachiatum Zoll. & Mor., Nat. en Geneesk. Archief Ned. Ind. 1 (1844) 399;

v.A.v.R. Handb. (1908) 252. — Lectotype (Morton): Zollinger 655 (G, L).

Aspidium zollingerianum Kunze, Bot. Zeit.4(1 846) 462: Bedd., Ferns Br. India (1867)

t. 251. — Nephrodium zollingerianum Bak., Syn. Fil. (1867) 298. — Type: as A. brachia-

tum.

Aspidium variolosum Wall. ex Hook., Spec. Fil. 4(1862)51; Bedd., Suppl. Ferns Br.

India (1876) t. 365 & Handb. (1883) 216; v.A.v.R.Handb. (1908) 248. — Tectaria

variolosa (Hook.) C. Chr., Contr. U.S.Nat.Herb. 26 (1931) 289: Ching, Sinensia 2 (1931)

21, pl. Ill; Backer & Posth., Varenfl. Java (1939) 76; Holttum 1955: 506, fig. 298: De-

Vol, Fl. Taiwan 1 (1975) 346. — Lectotype (Ching): Wallich 379, N.E.India (K).

Aspidium immersum Hook., Spec. Fil. 4(1862)58, non B1. 1828.—Type: Wallich s.n.,

without locality (K).

__ Phlebigonium impressum Fée, Gen. Fil. (1852)314, t.24A, fig.2.—Type:Griffith 34,

India (not seen).

Distribution: N.E. India to S. China and Taiwan; Thailand, Vietnam, northern

Malaya, Java.

Fertile fronds are much contracted as compared with sterile ones, and have simpler

venation, with one sorus-bearing vein in ‘each areole (see figure in Holttum, 1955). The

species is adapted to a climate with a seasonal dry period; its distribution is matched

by many angiosperms which are similarly adapted.

As pointed out by Morton, the epithet brachiata has precedence over variolosa,

though Backer & Posthumus placed the former as a synonym of the latter. Van Alder-

werelt treated them as two species, but his descriptions only differ in the texture of the

fronds which is not clearly definable; he had seen no type specimens.

5. Tectaria griffithii (Bak.) C. Chr., Ind. Fil. Suppl. III] (1934) 180; Tard. & C. Chr.

in Fl. Gen. Indoch. 7: 2 (1941) 411; Holttum 1968:636. — Nephrodium griffithii Bak.,

Syn. Fil. (1867) 300. —Sagenia griffithii (Bak.) Bedd., Ferns Br. India (1870) t. 337. —

Aspidium griffithii (Bak.) Bedd. ibid. Suppl. (1876) 15. — Type: Griffith s.n. Burma (K).

138 Gard. Bull. Sing. 34:1 (1981)

Nephrodium multicaudatum Clarke, Trans. Linn. Soc. Bot. 1 (1880) 540, t.77. —

Aspidium multicaudatum (Clarke) Bedd., Handb. (1883) 222; v.A.v.R., Handb. (1908)

250. — Tectaria multicaudata (Clarke) Ching, Sinensia 2 (1931) 20; Holttum 1955: 507,

fig. 299. — Type: Clarke 18427, Sylhet (K).

Aspidium malayense Christ, Philip. J. Sci. 2C (1907) 187. — Tectaria malayensis

(Chr.) Copel. ibid. 416 & Fern Fl. Philip. (1960) 305. — Several Philippine specimens

cited, no lectotype.

Distribution: Assam, Burma, Thailand, Vietnam, Malaya, Sumatra, Borneo, Philip-

pines.

6. Tectaria barberi (Hook.) Copel., Philip. J. Sci. 2C (1907) 414; Holttum 1955: 508,

fig. 300. — Polypodium barberi Hook., Spec. Fil. 5 (1864) 100. — Dictyopteris barberi

(Hook.) Bedd., Ferns Br. India (1869) t.322 & Handb. (1883) 515; v.A.v.R., Handb.

(1908) 515. — Phegopteris barberi (Hook.) Mett. in Kuhn, Linnaea 36 (1869) 125. —

Type: Wallich s.n., Penang (K).

Aspidium kunstleri Bedd., Handb. Suppl. (1892) 44. — Type: Kunstler 405, Gopeng,

Perak (K).

_ Phegopteris polycarpa Mett. in Kuhn, Linnaea 36 (1869) 124. — Aspidium kuhnii C.

Chr., Ind. Fil. (1905) 78, nom. nov. (not A.polycarpum B1.). — Polypodium polycarpum

(Mett.) Bak., Syn. Fil. ed. 2 (1874) 506. — Dictyopteris polycarpa (Mett.) Bedd., Ferns

Br. India Suppl. (1876) 20 & Handb. (1883) 300. — Type: Jagor s.n., Malacca (B, seen

by C. Chr.)

Distribution: Malaya, Borneo, Sumatra (specimens seen in Herb. BO).

Manton showed that this species is tetraploid (all others examined in Malaya are

diploid); see Holttum 1955: 626. It may thus be of hybrid origin, with doubling of

choromosomes, but I cannot suggest a likely parentage. Among species which have no

branched free veinlets in the areoles, it is peculiar in the number of its simple free vein-

lets, all excurrent. Hooker and Beddome did not see the indusia.

7. Tectaria singaporeana (Wall. ex Hook. & Grev.) Copel., Sarawak Mus. Journ. 2

(1917) 368; Holttum 1955: 512. — Aspidium singaporeanum Wall. ex Hook. & Grev.,

Ic. Fil. (1827) t. 26; Bedd., Ferns Br. India (1866) t. 168; v.A.v.R., Handb. (1908)

235. — Type: Wallich 374, Singapore (K).

Distribution: Malaya, Sumatra, Borneo.

Christensen has described a fern from West Kalimantan as Tectaria pandurifolia.

It differs from T. singaporeana in having the lamina of the fronds abruptly narrowed at

the base but decurrent as a narrow wing almost to the base of the stipe, also in the

presence of short hairs on the upper surface of the midrib which in T. singaporeana is

quite glabrous; I have not seen specimens.

8. Tectaria semipinnata (Roxb.) Morton, Contr. U.S. Nat. Herb. 38 (1974) 286. —

Acrostichum semipinnatum Roxb., Calcutta J. Nat. Hist. 4 (1844) 480. — Type: Rox-

burgh 2367 (or 2337), ‘“‘Malay Islands”, probably Penang (BM, BR).

Holttum: Tectaria in Malaya 139

Polypodium semipinnatum Roxb., Calcutta J. Nat. Hist. 4 (1844) 486; Morton,

Contr. U.S. Nat. Herb. 38 (1974) 360. — Type: W. Roxburgh Jr. s.n., probably from

Penang (BR).

Gymnogramma maingayi Bak., Syn. Fil. ed 2 (1874) 517. — Selliguea maingayi

(Bak.) Bedd., Ferns Br. India Suppl. (1876) 24 & Handb. (1883) 392. — Hemionitis

maingayi (Bak.) Ridl., J. Mal. Br. R. Asiatic Soc. 4 (1926) 106. — Aspidium maingayi

(Bak.) Holttum, Gard. Bull. S.S. 5 (1931) 207, fig. 1—8. — Tectaria maingayi (Bak.) C.

Chr., Ind. Fil. Suppl. III (1934) 182; Holttum 1955: 513, fig. 302. — Type: Maingay

1809, Malacca (K).

Polypodium heterosorum Bak., Syn. Fil. ed. 2 (1874) 506. — Dictyopteris heterosora

(Bak.) Bedd., Ferns Br. India Suppl. (1876) 20. — Type: Griffith s.n., Malacca, “Pulo

Bissar” = Pulau Besar (K).

Phegopteris subdecurrens Luerss., Bot. Centralbl. 11 (1882) 30. — Aspidium subde-

currens (Luerss.) C. Chr., Ind. Fil. (1905) 94 & Gard. Bull. S.S. 4 (1929) 393. — Type:

Kehding 2960, Singapore, Pulau Ubin (seen by C. Chr.)

Campylogramma trollii Goebel, Flora 125 (1931) 282, fig. 1-4. — Type: W. Troll,

Sumatra, Pulau Berhala, cult. Munchen (M).

Distribution: Malaya, Sumatra, Borneo, Anamba Islands.

The irregular form and distribution of the sori suggests hybridity, but the only

plant examined cytologically was diploid.

9. Tectaria grandidentata (Cesati) Holttum 1955: 514. — Polypodium dilatatum var.

grandidentatum Cesati, Atti. Acad. Napoli 7, no. 8 (1876) 27. — Polypodium grandi-

dentatum (Cesati) Bak., Ann. Bot. 5 (1891) 479; v.A.v.R., Handb. (1908) 661. — Pleo-

peltis grandidentata v.A.v.R., Handb. Suppl. (1917) 597; — Type: Beccari s.n., Jan.

1866, Sarawak, G. Matang (FI, K).

Distribution: Sarawak, Malaya, Lingga Archipelago, Sumatra.

This species agrees with T. singaporeana in having fronds decrescent towards their

bases, and in completely glabrous fronds. It is very local in occurence, in low country,

usually on rocks but not by streams.

10. Tectaria decurrens (Presl) Copel. in Elmer, Leafl. Philip. Bot. 1(1907) 234; Ching,

Sinensia 2 (1931) 22; Backer & Posth., Varenfl. Java (1939) 73; Copel., Fern Fl. Philip.

(1960) 315; Molesworth Allen, Gard. Bull. Sing. 22 (1967) 177 with phot.; Sledge,

Kew Bull. 27 (1972) 420.—Aspidium decurrens Presl, Rel. Haenk. (1825) 28; Bedd.,

Handb. (1883) 219; v.A.v.R., Handb. (1908) 247; Holttum, Novit. Bot. Inst. Bot. Univ.

Carol. Prag. 1968 (1969) 9. — Segenia decurrens (Presl) Moore, Ind. Fil. (1857) 86;

Racib., Fl. Btzg. 1 (1898) 247. — Type: Haenke s.n., Luzon (PRC).

Aspidium pteropus Kunze, Bot. Zeit. 4 (1846) 462; Mett., Farngatt. IV (1858)

120. — Sagenia pteropus (Kunze) Moore, Ind. Fil. (1858) 89; Bedd., Ferns S. India

(1863) t.82. — Type: Cuming 148, Luzon (LZ destroyed; isotype K).

Cardiochlaena alata Fée, Gen. Fil. (1852) 315. — Type: Cuming 148, Luzon.

140 Gard. Bull. Sing. 34:1 (1981)

Aspidium alatum Ridl., Trans. Linn. Soc. Bot. 9 (1916) 256, non Brack. 1854. —

Aspidium ridleyanum v.A.v.R., Handb. Suppl. (1917) 505, nom. nov..— Tectaria rid-

leyana (v.A.v.R.) C. Chr., Ind. Fil. Suppl. III (1934) 184. — Type: Kloss s.n., W. New

Guinea, Mt.Carstensz Exp., Canoe Camp (K).

Caudex erect. Stipe winged almost to the base, wing at the top 5-10 mm wide

each side, narrowing downwards; scales on abaxial side many, to 10 mm long, 1.5—2

mm wide at base, firm, entire. Lamina to 60 cm long consisting of 2—7 pairs of lobes

joined by a wing 5—10 mm wide each side of the rachis; lobes entire or sinuous or with

oblique lobules at ends of main veins, somewhat narrowed at the base, acuminate; basal

lobes sometimes forked near their bases; apex of lamina wider than the lateral lobes

and more narrowed towards its base; small veins distinct, forming areoles with free

included veinlets; lower surface of midribs and main veins bearing short hairs, upper

surface glabrous. Sori mostly in free veins in areoles, in a row on each side of main

veins, in depressions in the surface which form convexities on the upper surface; indusia

large, glabrous.

Distribution: very wide in mainland Asia, throughout Malesia and in the Pacific

eastwards to Tahiti. In Malaya, only collected in Perak: Scortechini without locality;

Matthew s.n., 25 Dec. 1911, Batu Kurau; Molesworth Allen 4486, Gopeng, on rocky

wet ground near stream, locally abundant.

11. Tectaria tricuspis (Bedd.) Copel., Sarawak Mus. Journ. 2 (1917) 369. — Aspidium

tricuspe Bedd., Handb. Suppl. (1892) 44. — Type: Kunstler 975, Perak, Gopeng (K).

Tectaria celemensiae Copel., Brittonia 1 (1931) 73. — Type: M.S. Clemens 22078,

Sarawak, Lundu, Mt Gadin (UC, K, NY).

Caudex thick, erect or suberect. Frond trilobed with a wing extending halfway

down the stipe (in a Sarawak specimen one pair of unlobed pinnae is connected to the

trilobed apical lamina by a winged rachis); stipe 35—40 cm long with a wing 2—8 mm

wide each side in its upper part; scales at base of stipe 10 x 1.5 mm, firm and dark

with distinct pale edges of thin-walled cells. Midlobe of apical lamina to at least 30 x

10 cm (sterile) or 23 x 6.5 cm (fertile), rather evenly elliptic with short acuminate apex

and entire margins; lateral lobes 22 x 8 cm (sterile) or 13 x 4.5 cm (fertile), shape as

apical lobe but widened on basiscopic side of midrib; main veins of sterile lateral lobes

10 mm apart, of fertile lobes 6 mm, at 45° to midrib, almost straight except near margin;

crossveins almost straight, smaller veins between them very slender, forming 2—3 rows

of irregular areoles containing branched free veinlets; lower surface of midrib and main

veins bearing very short erect hairs, upper surface densely covered with thick hairs,

other parts of surfaces glabrous; a bud present at the base of each lateral lobe. Sori

in two rows between the crossveins (4—S irregular rows between the main veins), on

veins which surround areoles; indusia rather large, apparently peltate, bearing many

hairs c. 0.3 mm long on the upper surface.

In Malaya, this species is still only known from the original collection, which is

not represented in the Singapore herbarium, for which reason I overlooked it when

writing my book. Apart from the indsia and the buds at the base of the main lobes,

this species is distinguished from T. vasta by the elliptic shape of the lateral lobes of

the lamina; in 7. vasta these lobes are little narrowed at their bases.

12. Tectaria vasta (B1.) Copel., Philip. J. Sci. 2C (1907) 411; Backer & Posth., Varen-

fl. Java (1939) 74; Holttum 1955: 512. — Aspidium vastum B1., Enum. P1. Jav. (1828)

—_-

Holttum: Tectaria in Malaya 141

142; Beddome, Handb. (1883) 212; v.A.v.R., Handb. (1908) 247. — Type: Blume, Java

(L).

Aspidium alatum Wall. ex Hook. & Grev., Ic. Fil. (1831) t.184; Hook., Spec. Fil. 4

(1862) 47. — Type: Wallich 378, Sylhet (K).

Distribution: Assam, Burma, Nicobar Islands, Thailand, Vietnam, Malaya, Sumatra,

Java, Borneo, Celebes, Moluccas, New Guinea.

13. Tectaria crenata Cav., Descr. Pl. Lec. publ. 1801 (1802) 250; C. Chr., Dansk Bot.

Ark. 9:3 (1937) 14 & Bishop Mus. Bull. 177 (1943) 102; Holttum 1955: 510; Copel.,

Fern F1. Philip. (1960) 312. — Type: Nee, Mariana Islands (MA, seen by C. Chr.)..

Aspidium repandum Willd., Sp. P1. 5 (1810) 216; Mett., Farngatt. IV (1858) 126;

v.A.v.R. Handb. (1908) 238. — Type: no collector cited, Philippines (B. Herb. Willd.

no. 19734).

Aspidium pachyphyllum Kunze, Bot. Zeit. 6 (1848) 259; Hook., Spec. Fil. 4 (1862)

56; v.A.v.R. Handb. (1908) 252. Sagenia pachyphylla (Kunze) Moore, Ind. Fil. (1857)

86; Racib., F1. Btzg. 1 (1898) 195. — Type: Zollinger 580, Java (not seen).

Distribution: Throughout Malesia and extending to Samoa. 7. dimorpha St John

(Occ. Pap. Bishop Mus. 21 (1954) 185, fig. 4), described from a specimen from Rotuma

Island, is at least very closely allied. T.crenata is closely allied to T. decurrens and the

two appear to hybridize in the Philippines.

14. Tectaria cherasica Holttum, sp. nov. forma et statura T. coadunato (J. Sm.) C. Chr.

similis, ab ea differt pagina supra inter venas glabra; venulis liberis brevibus multis in

areolis costalibus venarum inclusis, venulis liberis furcatis in areolis aliis etiam interdum

praesentibus; indusiis supra pilis tenuibus brevibus praeditis; paleis ad basin stipitum

non cellulis pallidis marginatis. Typus: A.G. Piggott 2027, Bukit Cheras, Panching,

Pahang, on limestone ridge, 24 Jan. 1977 (K).

Caudex stout, short-creeping. Stipe 27 cm long, stramineous or slightly flushed

with red above the base, copiously minutely hairy at base, glabrous upwards apart from

short hairs in the groove, basal scales to c. 7 x 1.5 mm, thin and translucent, light cast-

aneous, marginal cells uniform with the rest. Lamina (fertile) thin but firm, 40 cm long,

20 cm wide, consisting of an apical portion and four pairs of upcurved pinnae, the basal

ones not greatly longer than those next above them; apical portion 15 cm long, ovate-

deltoid in outline, base short-decurrent, lobed to 10 mm from its axis, basal lobes to 8 x

2 cm, lobulate less than halfway to their costae, upper lobes progressively shorter and

less deeply lobulate; first pair of pinnae adnate to rachis, 10 x 3 cm, lobed to 4 mm from

costa, lobes falcate, obtuse and entire; second pair of pinnae short-stalked, 12 cm long

with basal basiscopic lobe 3.2 x 1.2 cm shallowly lobulate, acroscopic lobe a little shorter,

other lobes subentire; third pair of pinnae 16 cm long with one pair of subequal free

pinnules 4.5 cm long; basal pinnae to 18 cm long (including stalk 1.5 cm), free basis-

copic pinnule 8 cm long, acroscopic one 5.5 cm long, both deeply lobed with crenulate

lobes, rest of pinnae like frond apex but somewhat asymmetric; areoles along midribs

of pinnae rather irregular with frequent short included veinlets, the larger non-costular

areoles also often with included veinlets which in a few cases are forked; ower surface

of pinna midribs and costules of lobes bearing slender pale hairs to 0.3 mm long, short

and more sparse hairs on smaller veins; upper surface more densely hairy on midribs

of pinnae, no hairs present between veins except very few near sinuses. Sori mostly on

142 Gard. Bull. Sing. 34:1 (1981)

short free veins in areoles, in one row each side of costules of smaller pinna lobes, more

irregular on larger lobes; indusia rather large, rather thin, slightly reniform, with many

very slender hairs 0.1 — 0.2 mm long on the upper surface.

In general aspects this is rather intermediate between.T. amplifolia and T. coadunata.

It lacks the hairs on the upper surface which are distinctive of the latter and has more

free veinlets in areoles, but a much less ample reticulation than the former; scales at

the bases of stipes are not narrow and dark as in 7. amplifolia. As no sterile fronds

were found, one cannot be sure whether they would be very different from fertile ones,

but the fertile fronds are much less contracted than those of 7. brachiata and have more

amply anastomosing veins, differing also in the rather copious hairs on the lower surface

of midribs and in the character of stipe scales. The sori are rather young; the few ripe

sporangia have well-formed spores, so that the plant is probably not a hybrid.

15. Tectaria siifolia (Willd.) Copel., Philip. J. Sci. 2C (1907) 414; Backer & Posth.,

Varenfl. Java (1939) 75; Copel., Fern Fl. Philip. (1960) 314. — Polypodium siifolium

Willd., Sp. Pl. 5 (1810) 196. — Aspidium siifolium (Willd.) Mett., Ann. Mus. Bot. Lugd.

— Bat. 1 (1864) 237; v.A.v.R., Handb. (1908) 251. — Type: Ventenat, Java (B, Herb.

Willd. no. 19689).

Aspidium haenkei Presl, Rel. Haenk. (1825) 30; Holttum, Novit. Bot. Inst. Bot.

Univ. Carol. Prag. 1968 (1969) 9. — Tectaria haenkei (Presl) Copel., Bishop Mus. Bull.

59 (1929) 50, nomen tantum, — Type: Haenke, Mariana Islands (PRC).

Drynaria menisciicarpa J. Sm. in Hook., J. Bot. 3 (1841) 421, nom. nud. — Dryome-

nis phymatodes Fée, Gen. Fil. (1852) 225, t. 18A, fig. 1. — Dryomenis menisciicarpa

Moore, Ind. Fil. (1857) 47. — Aspidium menisciicarpum Mett., Farngatt. [V (1858) 121,

non B1. 1828. — Type: Cuming 4, Luzon (Fée’s specimen not seen; isotype at K).

Aspidium teysmannianum Hook., Spec. Fil. 4 (1862) 41, t. 236. — A. siifolium

var. teysmannianum Christ, Ann. Jard. Bot. Btzg. 15 (1898) 138. — Type: Teysmann

s.n., Pulu Pisang, W. Sumatra (K).

Aspidium ternifolium v.A.v.R., Bull. Jard. Bot. Btzg. II, 11 (1913) 3; Handb. Suppl.

(1917) 194. — Tectaria ternifolia (v.A.v.R.) C. Chr., Ind. Fil. Suppl. HI (1934) 185;

Holttum 1955: 516, fig. 303. — Type: Matthew 510, Gopeng, foot of Bukit Mensa,

on limestone (BO, K).

Aspidium papyraceum v.A.V.R., Bull. Jard. Bot. Btzg. III, 2 (1924) 131. — Tec-

taria papyracea (v.A.v.R.) C. Chr., Ind. Fil. Suppl. III (1934) 183. — Type: Brooks

377/S, Benkoelen, Sumatra (BO, BM).

Comparsion of type specimens cited above, and other specimens from many locali-

ties from Sumatra to the Philippines and New Guinea, convinces me that they all re-

present one species. In several cases there are two pairs of pinnae, and in the type of

A. teysmannianum four pairs. The fronds are always strongly dimorphous; all lateral

pinnae are broadly rounded on the basiscopic base (sometimes the base is quite cordate);

basal pinnae in smaller fronds are simple, but in larger ones have a large basal basiscopic

lobe; on old sterile fronds there are always buds (sometimes quite large) at the bases

of pinnae; the sori are on connected veins, in two slightly irregular rows between the

main veins, sometimes asymmetric or confluent; the indusia are small and thin appa-

rently soon caducous so that many specimens appear to lack indusia.

Holttum: Tectaria in Malaya 143

16. Tectaria keckii (Luerss.) C. Chr., Ind. Fil. Suppl. III (1934) 181. — Aspidium kec-

kii Luerss., Bot. Centralbl. 11 (1882) 76; v.A.v.R., Handb. (1908) 248. — Type: Keh-

ding 2817, “Klang bei den Gua Batu” = Batu Caves, Selangor (not seen).

Aspidium amplifolium v.Av.R., Bull. Jard. Bot. Btzg. II, 11 (1913) 2; Handb. Suppl.

(1917) 197. — Tectaria amplifolia (v.A.v.R.) C. Chr., Ind. Fil. Suppl. III (1934) 176;

Holttum 1955: 515. — Type: Matthew 509, Perak, G. Mensa, Gopeng, on limestone (BO,

K).

Distribution: Malaya, peninsular Thailand; on limestone only.

Kehding collected plants in N.E. Sumatra and in southern Malaya. Luerssen, when

describing A. keckii, stated that he did not know whether the locality was in Malaya

or Sumatra, and subsequent authors wrote Sumatra only. The limestone habitat and the

description indicate clearly that Kehding’s specimen and the type of A. amplifolium are

conspecific.

As noted in my book, Henderson’s specimen SFN 22325 from G. Senyum in Pahang

(of which a duplicate is at Kew) has many short hairs on the lower surface of the pinna

midribs and main veins, whereas these are quite glabrous on most other specimens;

the G. Senyum specimens are also very large with pinna lobes strongly lobulate. Mrs

AG. Piggott has recently collected small fertile plants on Bukit Cheras in Pahang which

also have many hairs on the lower surface and are more dissected than those of yound

plants of typical 7. keckii. Local study is needed to decide whether these hairy plants

represent a distinct species.

17. Tectaria semibipinnata (Wall. ex Hook.) Copel., Sarawak Mus. J. 2 (1917) 371;

Holttum 1955: 515. — Aspidium semibipinnatum Wall. ex Hook., Spec. Fil. 4 (1862)

59,t.231.— Type: Wallich 388, Penang (K).

Nephrodium nudum Bak., Journ. Bot. 17 (1879) 41. — Type: Burbidge s.n., Sara-

wak, Lawas River (K).

I suggest that this species would be an interesting subject for experimental study.

No one knows how the gametophytes become established on muddy river banks in

the tidal zone; how can they withstand regular flooding? Can they grow in a submerged

condition, like those of Ceratopteris?

Tectaria modesta C. Chr. ( Dansk Bot. Ark. 9:3 (1937) 72, t.6, fig. 1) was describ-

ed from a plant which had narrow simple fertile fronds, much like a single pinna of

T. semibipinnata, and a slender creeping rhizome (Sarawak Museum Collector 218,

Kuching). It is possible that this represents a young stage of the present species which

is only known in Malaya and Borneo.

18. Tectaria melanocaula (B1.) Copel., Philip. J. Sci. 2C (1907) 416; Backer & Posth.,

Varenfl. Java (1939) 74; Copel., Fern Fl. Philip. (1960) 306; Holttum 1968: 636; Mor-

ton, Amer. Fern Journ. 61 (1971) 148. — Aspidium melanocaulon B1., Enum. P1. Jav.

(1828) 161; v.A.v.R., Handb. (1908) 245. — Sagenia melanocaulis (B1.) Moore, Ind.

Fil. (1858) 96; Racib., Fl. Btzg. 1 (1898) 195. — Type: Blume, Java (L).

Aspidium microsorum Presl, Epim. Bot. (1851) 61; Holttum, Novit. Bot. Inst. Bot.

Univ. Carol. Prag. 1968 (1969) 28. — Type: Cuming 57, Luzon (PRC, K).

144 Gard. Bull. Sing. 34:1 (1981)

This species, distributed almost throughout Malesia, has been only once collected

in Malaya, by Betty Molesworth Allen (no. 3985) in the forest on steeply sloping ground

near the road to the Cameron Highlands at 3000 ft altitude.

Caudex short, erect or suberect. Stipe 30—60 cm long, black, glossy except near

the base which bears many short hairs; basal scales firm; to 12 mm long, to 2 mm wide at

the base, marginal cells conform with the rest. Lamina to 60 cm long, pale green when

living with black rachis and pinna midribs, consisting of a deltoid deeply lobed terminal

portion and 1—3 pairs of pinnae; lobes of apical lamina more or less deeply lobulate;

middle pinnae short-stalked, commonly to 18 cm long, lobed halfway towards the

midrib, lobes obliquely falcate with rounded to acute tips; basal pinnae of large fronds

bearing one pair of free (sometimes stalked) pinnules which are lobed like the upper

pinnae, the basal basiscopic pinnule or lobe always longer than the acroscopic; lower

surfaces of midrib and main veins sparsely and minutely hairy, upper surface densely

covered with thicker hairs. Sori small, scattered irregularly, mostly on short free vein-

lets in areoles; indusia small, shrivelling and mostly persistent, not hairy.

Blume wrote the specific epithet melanocaulon, meaning black stem; the latter

part of the name is derived from the Greek kaulos, which Blume altered to make it

conform with a neuter generic name. When transferring the species to Sagenia, Moore

changed the specific epithet to melanocaulis, and in this was followed by Copeland

when transferring again to Tectaria, but-caulis is not correct in Greek. Morton (1971)

Stated that melanocaulon should be treated as invariable, but Blume had already varied

it. Backer and Posthumus wrote melanocaula, treating the word as a feminine Latin

adjective, and I think this is the best solution of the problem.

19. Tectaria simonsii (Bedd.) Ching, Sinensia 2 (1931) 32. — Aspidium simonsii Bedd.,

Ferns Br. India Suppl. (1876) 15, t. 367. — Type: Simons 301, Nuku & Naga Hills,

Assam (K).

Sagenia longicruris Christ, Bull. Acad. Geogr. Bot. Mans. 16 (1906) 250. — Aspidium

longicrure Christ, ibid. 20 (1909) Mem. 169. — Type: Cavalerie 268, Kweichow (not

seen, but a duplicate of Cavalerie 7294, cited in 1909, is at Kew).

Aspidium subtriphyllum var. ebenosum Nakai, Bot. Mag. Tokyo 47 (1933) 157. —

Tectaria subtriphylla var. ebenosa DeVol, F1. Taiwan 1 (1975) 345. — Syntypes: three

cited, from Ryukyu Islands, not seen.

Stipe nearly black, to 60 cm long, minutely hairy at base, the rest glabrous; basal

scales very firm and dark. Lamina to 40 cm long, “dark bluish green” when living (Mole-

sworth Allen), consisting of a deeply trilobed apex and 1-3 pairs of pinnae; middle

pinnae sessile with subcordate base which is a little dilated, margins subentire; basal

pinnae of large fronds stalked, with one pair of free pinnules (basiscopic pinnule longer

than acroscopic), the distal part trilobed with subentire lobes; veins anastomosing to

form areoles with many included free veinlets, concolorous but distinct on both sur-

faces; lower surface of midribs of pinnae and main veins bearing scattered minute hairs,

longer dense antrorse hairs present on the upper surface. Sori small and uneven in shape,

sometimes a little elongate, scattered irregularly, usually not on free veins; no indusia

seen.

Distribution: Assam to southern China, Taiwan and Ryukyu Islands, Vietanam,

Thailand. The only specimen collected in Malaya is Molesworth Allen 4947 from the

forest near the 23rd mile, Cameron Highlands Road, at an altitude of 2000 ft.

Holttum: Tectaria in Malaya 145

20. Tectaria angulata (Willd.) Copel., Sarawak Mus. J. 2 (1917) 370; Holttum 1955:

511, fig.301, excl. syn. Nephrodium nebulosum Bak. — Polypodium angulatum Willd.,

Sp. Pl. 5 (1810) 185. — Aspidium angulatum (Willd.) Mett., Ann. Mus. Bot. Lugd. —

Bat. 1 (1864) 239. — Type: Ventenat s.n., Java (B, Herb. Willd. 19641).

Aspidium pentaphyllum v.Av.R., Bull. Dep. Agr. Ind. Neerl. 18 (1908) 16, non

Willd. 1810. — Dictyopteris pentaphylla v.Av.R., Handb. (1908) 519, nom. nov. —

Aspidium quinquefoliatum C. Chr., Ind. Fil. Suppl. 1(1913) 9, nom. nov. superfl. —

Type: Treub s.n., New Guinea (BO, K).

Aspidium trifolium v.AN.R., Bull. Jard. Bot. Btzg. II, 7 (1912) 4. — Tectaria folium

(v.A.v.R.) C. Chr., Bot. Jahrb. 66 (1933) 49; Copel., Fern Fl. Philip. (1960) 307. - Type:

Elmer 8239, Luzon (BO, K).

Aspidium terminale Rosenst., Meded. Rijksherb. 31 (1917) 4; C. Chr., Gard. Bull.

S.S. 4 (1929) 393. — Tectaria terminalis (Rosenst.) C. Chr., Ind. Fil. Suppl. III (1934)

185. — Type: Hallier 1821, Borneo (L, BO).

Distribution: Malesia, Solomon Islands.

21. Tectaria herpetocaulos Holttum, Dansk Bot. Ark. 23: 2 (1965) 241; Hottum 1968:

636. — Type: B. Molesworth Allen 4454, 22nd mile, Cameron Highlands Road, “in

steep rocky forest”, 2000 ft (K).

Caudex long-creeping, 6-10 mm diameter when dried, bearing stipes about 1.5

cm apart; scales 5 x 1 mm, base cordate, dark glossy with pale fragile edges. Stipe 30—80

cm long, lightly flushed with red, glabrescent, scaly near base, scales as those of caudex

but to 10 mm long. Fronds of young plants trifoliate, those of mature plants consisting

of an apical lamina and 4 pairs of pinnae; apical lamina to 26 x 11 cm (smaller on larger

fronds), entire, base cuneate, apex abruptly short acuminate; upper pinnae sessile, to

21 x 6 cm, with asymmetric base rounded on the basiscopic side; basal pinnae stalked,

to 30 cm long and 10 cm wide, asymmetric with a single basiscopic lobe to 20 x 6 cm

(largest fronds may have 2 pairs of pinnae with basal lobes); pinnae glabrous apart

from rather sparse short hairs on lower surface of midribs and main veins and dense

short hairs on their upper surface; main veins pale on lower surface, 8—10 mm apart,

at a wide angle and upcurved; smaller veins all slightly prominent, forming irregular

rather small areoles with included free veinlets. Sori mostly not on free veins, rather

small, in a single row on each side of the main veins but with additional sori between

the rows irregularly present; indusia thin, shrivelling but persistent; perispore consisting

of many narrow short projections.

In Malaya still only known from the original collection and no. 9948 from the

same locality, “spreading over humus-covered rocks’’. The species also occurs in Thailand,

Burma, and Bangladesh (see list of specimens in Holttum 1965). Several specimens

are recorded as growing on limestone.

22. Tectaria oligophylla (Rosenst.) C. Chr., Ind. Fil: Suppl. II] (1934) 183. — Aspidium

oligophyllum Rosenst., Fedde Repert. 5 (1908) 13; v.A.v.R., Handb. (1908) 237. —

Type: Burchard s.n., 1907, Sumatra, Indragiri (L; not seen).

Tectaria polymorpha vat. cuneifolia Bonap., Notes Pterid. 14 (1923) 50; Holttum.

1955: 518, fig. 304. — T. cuneifolia Love & Love, Taxon 26 (1977) 326. — Type: Holt-

tum 9566, G. Tampin (P, SING). Leptochilus rumicifolius Ridl.,J. Mal. Br. R. Asiat. Soc.

146 Gard. Bull. Sing. 34:1 (1981)

4 (1926) 116. — Tectaria rumicifolia (Bonap.) C. Chr., Ind. Fil. Suppl. HI (1934) 184;

Holttum 1955: 519 & Dansk Bot. Ark. 23: 2 (1965) 241. — Type: Ridley s.n. 1921,

Klang Gates, Selangor (K).

Distribution: Malaya, Sumatra, Sarawak; in lowland forests.

When writing my book of 1955 I overlooked the description of Aspidium oligo-

phyllum in van Alderwerelt’s Handbook. Rosenstock’s type is presumably at Leiden;

I did not find a duplicate at Bogor. His description is good and I cannot doubt that

it applies to the Malayan plants which I described as 7. polymorpha var. cuneifolia.

He described the indusia as peltate, stating that they were like those of T. singaporeana,

but I found that the indusia of the latter species are variable and usually not fully peltate.

The true 7. polymorpha is a quite different species with broad-based pinnae.

Ridley described Leptochilus rumicifolius from young plants which have fronds

rather copiously hairy on both surfaces. Mrs Allen sent similar plants, from rocks at

Klang Gates, to Kew; these plants later developed almost hairless fertile fronds. Pro-

bably the hairiness of young plants which grow on rocks is due to the habitat. Ridley

thought his plant was sterile, but it has a few sori on one of its simple fronds.

23. Tectaria fauriei Tagawa, J. Jap. Bot. 14 (1938) 102; Acta Phytotax. Geobot. 10

(1941) 203; J. Jap. Bot. 36 (1961) 208; DeVol & Kuo, F1. Taiwan 1 (1975) 342; E.H.

Walker, Fl. Okinawa (1976) 86. — Type: Faurie 67, Feb. 1914, Taiwan, Prov. Takao

(KYO, not seen).

Tectaria gymnosora Holttum, Dansk Bot. Ark. 23 (1966) 308. — Type: Hansen

et al. 11225, N. Thailand (C, K).

Tectaria vasta p.p. sensu C. Chr., Contr. U.S. Nat. Herb. 26 (1931) 289; sensu Hot-

tum 1955: 513, specimens from Thailand.

Caudex stout, erect. Stipe 40—60 cm long, narrowly winged in the upper part;

basal scales 10 mm or more long, to 2 mm wide at base, firm, with narrow margin of

thin-walled cells. Lamina firm, to 50 cm long, consisting of a trilobed apex which is

narrowly decurrent at the base almost to the attachment of the uppermost pair of pinnae,

and 2 or 3 pairs of pinnae, all with similarly narrow-decurrent bases; buds present at

the base of some pinnae on mature fronds; middle pinnae 20—25 cm long, entire or

nearly so, gradually narrowed towards their bases, apex narrowly acuminate, sterile pin-

nae often wider than fertile, to c. 6 cm wide; basal pinnae of small fronds narrowed

towards their bases on both sides, unlobed but the lamina on the basiscopic side of the

midrib wider than on the acroscopic side, of larger fronds with a narrowly winged stalk

and a rather long basal basiscopic lobe; pinna midribs and main veins reddish on the

lower surface when dried and minutely sparsely hairy, the upper surface glabrous; smaller

veins concolorous and distinctly prominent on both sides, forming many areoles which

enclose branched free veinlets. Sori small, round, in about 4 irregular rows between

the main veins, not on free veins; indusia rather small, thin, glabrous, persistent.

Distribution: Assam, northern Shan States and adjacent Yunnan, Taiwan, Ryukyu

Islands (Okinawa), N. Thailand, Pahang; in several cases reported as growing on rocky

stream banks.

The specimen from Pahang is Holttum 20071, Tahan River, below Kuala Teku,

on rocks and earth of river bank. I failed to notice this specimen when writing my book

of 1955 and referred to specimens from Thailand erroneously under T. vasta.

Holttum: Tectaria in Malaya 147

This species differs from 7. vasta in its narrow rachis wing which is decurrent from

each pair of pinnae but does not reach down to the next lower pair, in the much narrow-

ed bases of pinnae and their more narrowly acuminate apices, in the presence of buds

at the bases of some pinnae on old fronds, and in the indusia which are not fringed.

T. fauriei is nearer to T. oligophylla, agreeing rather closely in the shape of pinnae

and of sori and indusia, differing in the decurrent bases of pinnae and in the presence

of buds. 7. oligophylla also has rather long-stalked basal pinnae on old fronds. T. oli-

gophylla is always a fern of forest, not of river banks, in my experience. T. fauriei pro-

bably occurs on the banks of tributaries of the Kelantan River where natural forest re-

mains; it probably would not persist on expoed river banks from which forest had been

felled.

Though I have not seen the type specimen of TJ. fauriei, | have seen a specimen

from Thailand identified by Prof. Tagawa, author of the species, and also from his des-

cription I feel confident that my identification of the Tahan River plant is correct.

148

EXPERIMENTAL FINDINGS OF

THE TISSUE CULTURE OF ORCHID HYBRIDS

AT THE SINGAPORE BOTANIC GARDENS

Lim-Ho Chee Len

Botanic Gardens,

Singapore

ABSTRACT

A Tissue Culture Laboratory was established at the Singapore Botanic Gardens

in the early 1970’s. By 1975 several orchid hybrids had been successfully mericloned

and in 1978 the Laboratory started a tissue culture service to assist local orchid growers

and breeders. To date, the Laboratory has successfully mericloned six species and more

than one hundred orchid hybrids. Some effective media compositions have been establish-

ed and used in research and routine commercial production at the Laboratory. Tissue

culture techniques using leaves and lateral buds of flower stalks have also been employed.

Various tissue culture techniques and media used for local orchids is presented.

INTRODUCTION

Research work on orchids at the Botanic Gardens, Singapore started in 1928 and

early work was mainly on hybridisation. Following the success in tissue culture of orchids

by G.M. Morel (1960) in France, it was decided to venture into this new line of research

at the Botanic Gardens. A Tissue Culture Laboratory was established in the early 1970’s

and preliminary experiments were conducted with improvised equipment. In 1973 one

laminar flow and two shakers were acquired, however these shakers operated on a pulley

system which broke down frequently. In 1975 a Brunswick shaker was purchased. By

that time, several hybrids of Dendrobium (Singh, 1976), Aranda, Aranthera, and Burkillara,

had been successfully mericloned. After many attempts, Dendrobium attennatum

was also successfully mericloned in 1977.

In late 1978 the Tissue Culture Laboratory started a mericlone service to assist

local orchid growers and breeders. The Laboratory undertakes experimental tissue cul-

ture propagation of parent plants sent in by the public. When fully differentiated plant-

lets are obtained the experiment is considered successful and a contract is then agreed

upon between the owner of the parent plants and the Laboratory for the purchase and

supply of a certain number of plantlets. Plantlets supplied are at least 3 cm in length

with strong, healthy roots and in a condition ready for potting. Orchid hybrids which

have been mericloned at the Laboratory and supplied commercially on a large scale

include Dendrobium Tay Swee Keng, Dendrobium Yong Kok Wah, Dendrobium Jaquelyn

Hawaii, Aranda Chong Chee Yuen, Kagawara Teoline Fair, Mokara Sally Lim, and Aranda

Ng Chong Hang. Facing keen competition from Thailand and Malaysia on the inter-

national market for cut flowers, Singapore orchid growers and breeders have increasingly

sought the assistance of the Tissue Culture Laboratory for the mass propagation of

their choice hybrids. The Laboratory has, therefore, been able to contribute directly

to the rapid growth, both in terms of the total annual export and the level of technical

sophistication, of the local orchid industry by making readily available an efficient

mass propagation technique based on advanced biological knowledge.

Lim-Ho: Orchid Tissue Culture 149

Orchid Tissue Culture Techniques

In the poineering work of Morel (1960) shoot apices of Cymbidium were excised

to produce virus-free protocorms which later developed into plantlets. The techniques

used at the Tissue Culture Laboratory are basically variations of Morel’s methods. Young

leaves (Fu, 1979), axillary buds, lateral buds of flower stalks (Fu, 1978), and roots

are sometimes used instead of apical shoots. Various media have been tested and hor-

mones are occasionally added to the basic salt to induce protocorm formation, to acce-

lerate growth, and to promote protocorm differentiation.

Plant Tissues Used

Using the shoot apex of an orchid plant for tissue culture requires the sacrifice of

new growth or even, as in the case of monopodial orchids, the whole plant. Since success

cannot be guaranteed and most of the plants selected for tissue culture are valuable

(sometimes it is the only specimen of a new hybird of great commercial potential), the

cost and risk involved could be high. It is, therefore, desirable to develop techniques

using tissues which can be removed without doing much harm to the parent plants.

Experiments have been conducted using young leaves, axillary buds, lateral buds of

flower stalks, and roots as explants. Some of these experiments were successful and

the methods developed have been adopted in routine production at the Tissue Culture

Laboratory (see Tables 1 and 2). The parts taken from parent plants for the various

tissue culture techniques are summarised below:

Culture technique Part of plant used

Apical and axillary meristems

sympodial orchids young side shoot measuring 7—8 cm

monopodial orchids top 10 cm of the apical shoot

Leaf whole young leaf (3—4 x 1—2 cm)

from mature plant

Lateral bud (flower stalk) whole flower stalk about 20 cm |

in length

Root young aerial root about 5 cm in

length

Sterlization

Prior to excision, all plant parts must be sterilized. The methods for the various

plant parts are as follows:

1. Apical and axillary meristems: The leaves are removed to expose the apical or

axillary buds. The remaining shoot is then washed with antiseptic soap and soaked

in a chlorox solution (10—12%) for 10—15 minutes.

2. Leaves: Cut leaves are washed with antiseptic soap and soaked in a chlorox

solution (10—12%) for 15—20 minutes.

3. Lateral buds of flower stalks: Scale leaves are removed from the flower stalks

to expose the lateral buds. The remaining stalk is then washed with antiseptic soap

150 Gard. Bull. Sing. 34:1 (1981)

and soaked in a chlorox solution (12—15%) for 20 minutes.

4. Roots: Cut aerial roots are washed with antiseptic soap and soaked in a chlorox

solution (15%) for half an hour.

Excision

In meristem culture, 0.2 mm cubes are excised from sterilized plant parts and care is

taken to excise only the meristematic cells and not the surrounding tissues. In root

culture, young root tips 1—2 cm long are cut for use as explants. In leaf culture, whole

young leaves are used.

Inoculation

All excised plants are put into 100 ml conical flasks continaing 25 ml of liquid

media under sterilized conditions. These are then sealed and placed on shakers.

Shaking and environmental conditions

Shaking is essential during the early stages of protocorm formation (Wimber, 1963,

1965; Kunisaki et al, 1972). The basic purposes of shaking are to increase aeration, to

reduce plant polarity, to enhance cell multiplication, to dilute excretion of explants,

and to help distribute nutrients. From our experience, the mode of shaking is not signi-

ficant however, too violent shaking has been observed to cause browning-off of pro-

tocorms.

In the Tissue Culture Laboratory, explants in shakers are exposed to continuous

illumination of about 2000 lux and maintained at a temperature of 21—25°-C.

Media

The most commonly used media for tissue culture of orchids are the White (1943)

medium, the Murashige and Skoog (1962) medium, the Vacin and Went (1949) medium,

and the Knudson (1949) C medium. These and many modified forms of these media

have been tested at the Tissue Culture Laboratory. The most successful of the modi-

fied media are: KCBP medium — Knudson C basic salt supplemented with banana and

pineapple juice; and VWM medium — Vacin and Went basic salt supplemented with

banana, coconut milk, and tomato juice.

The media used depend on the stage of development of the protocorms, For pro-

tocorm initiaton, the Vacin and Went medium and the Knudson C medium have been

found to be far superior to the others for apical shoot tip and axillary bud cultures.

For most species and some hybrids it is necessary to add NAA or IAA to induce pro-

tocorm formation. In the case of tissue culture using leaves and lateral buds of flower

stalks, the Murashige and Skoog medium supplemented with 2—4—D and 6—amino—

benzyladenine has been found to be best for protocorm initiation.

For protocorm multiplication and differentiation, the Vacin and Went medium

supplemented with coconut milk and sugar is best for most hybrids, especially for Dend-

robium hybrids. Vanda hybrids, however, have been observed to multiply and differen-

tiate faster in the Vacin and Went medium supplemented with coconut milk only. For

most Aranthera hybrids, the best medium for multiplication and differentiation is the

Murashige and Skoog basal salt.

Lim-Ho: Orchid Tissue Culture 15]

The VWM medium has been found to be the best for the final stage of most hybrids.

For some slow-growing hybrids such as Aranda Majula ‘“Rimau’ and Aranda Christine

No. 5, root formation is much faster in the KCBP medium than in the VWM medium.

All Oncidium hybrids fail to form roots in the VWM medium, but satisfactory root

formation is obtained in the KCBP medium. Since species of Arachnis and Aranthera

hybrids excrete strong phenolic componds, it is necessary to change media every few

days during the first six months of their growth. The composition of the media used

in the Tissue Culture Laboratory is listed in Table 3. The appropriate media for some

local species or hybrids at different stages of development are given in Tables 1 and 2.

Data and observations on tissue culture of local orchids

To date the Tissue Culture Laboratory has successfully mericloned 6 species and

more than 110 hybrids of orchids. All the species are grown locally and out of the hy-

brids, 82 are local. Summaries of the basic data, part of plant used, media, growth rate,

and relevant remarks on the tissue culture of these local orchid species and hybrids

are given in Tables 1 and 2. These tables also show that many orchids have been success-

fully mericloned and it is possible that all orchid hybrids can be mericloned provided

the right media is used.

Coconut milk appears to be an important supplement to all the media used in every

stage of protocorm development. Coconut milk is believed to contain hormones (pro-

bably cytokinins) in a balanced composition (Letham, 1974) which are effective in

inducing protocorm formation and differentiation. Auxins such as NAA and IAA are

occasionally added to media to induce protocorm formation, however, they are found

to have an inhibitory effect on the differentiation of protocorms.

A wide range variation in growth rates has been observed. Fast-growing hybrids

produce marketable plantlets in about one year’s time while slow-growing hybrids may

take more than two years. Most Dendrobium hybrids have been found to be fast-growing

and except for some Aranda hybrids, monopodial ground orchids are generally slow-

growing. Experience seems to suggest, however, that the slow growth of many hybrids

may probably be due to the use of unsuitable media rather than to some basic phy-

siological characteristics of the hybrids. Considerable research effort in the Tissue Cul-

ture Laboratory has, therefore, been on the improvement of media in use and the search

for effective new media.

Acknowledgement

The Tissue Culture Laboratory was developed due to the efforts of many researchers,

especially Dr. Fu-Fan May Lay and Dr. Choo Yon Sen.

152

Gard. Bull. Sing. 34:1 (1981)

TABLE 1. LOCAL ORCHID SPECIES SUCCESSFULLY MERICLONED

Arachnis

hookerana

Dendrobium

antennatum

laciniosum

superbiens

‘Superba’

Oncidium

lanceanum

haematochilum

Aeridachnis

Bogor “Apple

Blossum’

Alexandra

Elizabeth

Howie

Arachnis

Capama’

‘merah’

Arachnostylis

Chorchalood

Aranda

Belzonica

. Bertha Braga

. Bintang

. Chong Chee

Yuen

. Chen Chin Mu

. Christine No. 1

. Christine No. 5

. Christine No. 9

. Christine No. 27

. Christine No. 80

Source of

meristem

explant Growth rate

apical & extremely

axillary slow

apical slow

apical & slow

axillary

apical fast

apical slow

apical fast

Parentage’

Arachnis hooke-

rana x Aerides

odorata

Aeridachnis

Bogor x Arach-

nis flosaeris

‘insignis’

Arachnis Ishbel x

Aerides lawren-

ceae

Arachnis Maggie

Oei x Arachnis

breviscapa’

Arachnis hooke-

rana x Rhyno-

chostylis gigantea

Arachnis Ishbel

x Vanda luzonica

Vanda tricolor x

Arachnis Maggie

Oei

Arachnis hooke-

rana x Vanda

Tatzeri

Aranda Blue

Star x Vanda

Rothschildiana

Aranda Bintang

Raffles x Vanda

Wangse

Arachnis hooke-

rana x Vanda

Hilo Blue

Arachnis hooke-

rana x Vanda

Hilo Blue

Arachnis hooke-

rana x Vanda

Hilo Blue

Arachnis hooke-

rana x Vanda

Hilo Blue

Arachnis hooke-

rana x Vanda .

Hilo Blue

Source of

meristem

explant

apical &

axillary

apical &

axillary

apical &

axillary

apical &

axillary

apical &

axillary

apical &

axillary

apical &

axillary

apical &

axillary

apical &

axillary

apical &

axillary

whole

leaf

apical &

axillary

Media for Media for multi- , Media for

initiation plication and plantlet

(liquid) differentiation formation Remarks

VWN + NAA VWN, VWS VWM Arachnis spp. often ,

excrete strong phenolic

compounds; need to

change medium weekly

VWS + NAA VWN, VWS VWM during the first six months

Vws VWN, VWS VWM

VWS + NAA VWN, VWS VWM

VWS VWS VWM

VWS VWN, VWS KCMP

Growth

slow

fast

slow

fast

slow

fast

TABLE 2. LOCAL ORCHID HYBRIDS SUCCESSFULLY MERICLONED

Media for Media for multi- Media for

initiation

(liquid)

VWN/KC

VWN

plication &

differentiation

VWN, VWS

VMN, VWS

VWN, VWS

VWN, VWS, MS

VWN, VWS

VWN, VWS, MS

VWN, VWS

MS+2-4-D+ VWN, VWS

VWN

VWN, VWS

plantlet

formation Remarks

VWM

VWM MS medium best

for differentiation

VWM

VWM/ MS medium best

KCBP for differentiation,

root formation faster

VWM in KCBP than in VWM

VWM

VWM/ root formation

KCBP faster in KCBP

than in VWM

VWN

VWM

Lim-Ho: Orchid Tissue Culture

TABLE 2. LOCAL ORCHID HYBRIDS SUCESSFULLY MERICLONED

Aranda

11. Christine No. 130

12.

20.

21.

22)

23.

24.

25:3

26,

28.

29.

30.

Christine No. 999

. Elizabeth

Douglas Home

. Eric Mekie

. Freckles

. Hatifah

. Hilda Galistan

. Imelda Romual-

dez Marcos

. Jessie

Lucy Laycock

Majula “Rimau’

Ng Chong Hang

Noorah Alsagoff

Peter Ewart

Queen of

Purples

Tourism

Singapura

Wee Huck Lay

Wendy Scott

‘Blue Bird’

Wendy Scott

‘Green Field’

Yvonne Tan

Aranthera

Anne Black

Beatrice Ng

“Yellow’

Dainty

Parentage’

Arachnis hooke-

rana x Vanda

Hilo Blue

Arachnis hooke-

rana x Vanda

Hilo Blue

Arachnis Maggie

Oei x Vanda

Radman

Aranda Lucy Lay-

cock x Vanda

luzonica

Arachnis Ishbel x

Vanda dearei

Arachnis hooke-

rana x Vanda

luzonica

Arachnis hooke-

rana x Vanda

tricolor suavis

Arachnis main-

gavi x Vanda

Harvest Time

Arachnis hooke-

rana luteola x

Vanda Ellen Noa

Arachnis hooke-

rana x Vanda

tricoler

Arachnis Maggie

Oei x Vanda

insignis

Aranda Bintang

Raffles x Vanda

Rose Davis

Arachnis hooke-

rana x Vanda

Dawn Nishimura

Arachnis hooke-

rana x Vanda

Kapoho

Arachnis hooke-

rana x Vanda

coerulea

Vanda Kapoho x

Arachnis brevis-

capa

Arachnis Ishbel x

Vanda lamellata

Arachnis hooke-

rana x Vanda

Rothschildiana

Arachnis hooke-

rana x Vanda

Rothschildiana

Aranda Queen

of Purples x

Vanda Roths-

childiana

Arachnis Maggie

Oei x Renanthera

coccinea

Renanthera

storiei x

Arachnis \shbel

Arachnis hooke-

rana var. luteola x

Renanthera

monachica

Source of

meristem

explant

apical &

axillary

apical &

axillary

apical &

axillary

apical &

axillary

leaf base

apical &

axillary

axillary &

leaf base

axillary

Media for Media for multi- Media for

Growth initiation plication &

rate

fast

slow

fast

slow

fast

slow

fast

slow

fast

slow

(liquid)

differentiation

piantlet

formation Remarks

MS+24-D VWN, VWS VWM

+BA

VWN VWN, VWS VWM

VWN/KC VWN, VWS VWM

VWN VWN, VWS VWM

VWN VWN, MS, VWS_ KCBP/

VWM

VWN VWN, VWS VWM/

KCBP

MS+2-4-D+ VWN, VWS VWM

VWN VWN, VWS VWM

VWN/KC VWN, VWS VWM

VWN VWN, VWS VWM

VMW VWN, VWS VWM

VWN VWN, VWS,MS_ KCBP/

(apical) VWM

MS+24-D+BA

VWN VWN, VWS VWM

VWN VWN, VWS,MS VWM

VWN+24-D VWN, VWS

+BA

VWN

VWN, VWS

VWM

MS medium is best

for differentiation,

root formation faster

in KCBP than in VWM

MS medium better

than VWS for

differentiation

MS medium better

than VWS for

differentiation

153

154

TABLE 2. LOCAL ORCHID HYBRIDS SUCESSFULLY MERICLONED

Aranthera

4. Francina Bruyns

5. Gracia Lewis

6. James Storie

Ascocenda

1. Heah Hock

Heng

2. Botanic Gardens

serial No. 4038

3. Botanic Gardens

serial No. 4766

Burkillara

1. Ong Thye Chiew

Cattleya

1. Meadii

2. Botanic Gardens

serial code C03

Dendrobium

1. Alice Spalding

2. Dendrobium

Barbara Moore

3 Caesar

4 Cheong Chee

Yon

5. Ellen Harris

6. Fiery Gold

Source of

meristem

Parentage’ explant

Aranthera Dainty apical &

x Renanthera axillary

storiei

Aranthera apical &

Mohamed Haniff axillary

x Renanthera

storiei

Arachnis hooke- apical &

rana x Renanthe- axillary

ra Storiei

Vanda Josephine whole

van Brero x leaf

Ascocenda Hilo

Rose

unregistered apical &

hybrid axillary

unregistered apical &

hybrid axillary

Aeridachnis apical &

Bogor x axillary

Vanda Dawn

Nishimura

Cattleya bow- apical

ringiana x

Cattleya

forbesii

unregistered leaf base

hybrid apical

Dendrobium apical &

tokai x Dendro- axillary

bium undulatum

Dendrobium affine axillary

x Dendrobium

gouldii ‘blue’

Dendrobium apical &

phalaenopsis x axillary

Dendrobium

sthatiotes

Dendrobium apical

Cheong Fook Sum

x Dendrobium

Queen Emma

Dendrobium apical &

phalaenopsis x axillary

Ursula

Dendrobium apical &

Ismail Treng- axillary

ganu x Dendro-

bium Guadalcanal

Growth initiation plication & plantlet

VWS/VWN VWN, VWS VWM VWS solid medium good

Media for

rate (liquid)

slow VWN

fast MS+2-4-D

+BA

slow VWN

slow VWN + NAA

slow VWN

fast MS+24D

+BA

fast

slow VWs

fast VWs

fast VWN

slow VWN

slow VWS

Gard. Bull. Sing. 34:1 (1981)

Media for multi- Media for

differentiation formation Remarks

VWN, VWS VWM

VWN,VWS,MS VWM MS medium better

than VWS for

differentiation

Aranthera spp. ex

crete strong phenolic

compounds, need to

change medium every

few days during the

first six months; when

comparing them to

Vanda and Aranda,

their growth is extremely

slow

VWN, VWS VWM

VWN,VWS,MS VWM MS medium better

than VWS medium

for differentiation

VWN, VWS VWM

VWN, VWS,MS VWM

VWN, VWS KCBP root formation

difficult, roots

form in KCBP

medium, but not

VWM, VWS VWM in VWM medium

for all Dendrobium spp.

for differentiation into

VWN, VWS VWM shoot; rate of differentia-

tion fast in this medium,

but not good for Vanda spp.

VWN, VWS VWM

Lim-Ho: Orchid Tissue Culture

Dendrobium

10.

Ng Eng Cheow

Sunny

. Tay Swee Keng

Yong Kok Wah

. Botanic Gardens

serial No. 4879

. Botanic Gardens

serial No. 8090

. Botanic Gardens

serial No. 8262

. Botanic Gardens

serial No. 8324

. Botanic Gardens

serial No. 8961

. Botanic Gardens

serial No. 9215

. Botanic Gardens

serial No. 12917

Doritaenopsis

Elizabeth

Waldheim

Botanic Gardens

serial PDI code 1

Holttumara

Cochineal

Kagawara

1. Teoline Fair

Mokara

Sally Lim

Parentage

Dendrobium

Alice Spalding

x Dendrobium

Jaquelyn Thomas

Dendrobium

Beach Girl x

Dendrobium

Jaquelyn Concert

x Dendrobium

Irene Cheong

Dendrobium

Mary Trowse x

Dendrobium

schulleri

unregistered

hybrid

unregistered

hybrid

unregistered

hybrid

unregistered

hybrid

unregistered

hybrid

unregistered

hybrid

unregistered

hybrid

Doritis pul-

cherrima x

Phalaenopsis

Lam Soon

unregistered

hybrid

Aranda Hilda

Galistan x

Renanthera

coccinea

Ascocenda

Meda Arnold x

Renanthera

storiei

Arachnis

hookerana x

Ascocenda

Meda Arnold

Source of

meristem

explant

apical &

axillary

lateral bud

of flower

stalk

lateral bud

of flower

stalk

apical

Growth

rate

fast

slow

fast

slow

fast

slow

intiation

(liquid)

VWS+

VWN

VWS

TABLE 2. LOCAL ORCHID HYBRIDS SUCCESSFULLY MERICLONED

Media for Media for multi- Media for

plication & plantlet

differentiation

VWN, VWS VWM

VWN, KCS, VWS VWM

VWN, VWS VWM

VWN, KCS, VWS VWM

VWN, VWS VWM

VWN, VWS Tch

VWN, MS VWM

VWN, VWS, VWM

VWN, VWS KCBP

formation Remarks

3 petals

Lae

Root formation

slow, successful

only in KCBP

medium

155

156 Gard. Bull. Sing. 34:1(1981)

TABLE 2. LOCAL ORCHID HYBRIDS SUCCESSFULLY MERICLONED

Source of Media for Media for multi- Media for

meristem Growth initiation plication & plantlet

Parentage explant rate (liquid) differentiation formation Remarks

Phalaenopsis

1. Botanic Gardens _ unregistered flower slow MS+24-D VWN,VWS Tch

serial No. 3596 hybrid stalk +BA

Renanopsis

1. Lena Rowold Renanthera apical & slow VWN VWN, MS VWM

storiei x axillary

Vandopsis

laissochiloides

Vanda

1. Chia Kay Heng Vanda Josephine apical & fast VWN VWN VWN VWN solid medium

van Brero x Vanda axillary good for protocorm

Dawn Nishimura multiplication and

2. Peopoe ‘Diana’ Vanda Cooperi differentiation for

x Vanda teres Vanda spp.

3. Norbert Vanda Alice Lay- apical & slow VWN VWN VWM

Alphonso cock x Vanda axillary

Cooperi

4. Patricia Low Vanda Josephine apical & slow VMW VMW VMW

van Bréro x axillary

Vanda Jennie

Hashimoto

5. Ruby Prince Vanda Ruby x apical & fast VWN VWN VWM

Vanda Cgoperi _ axillary

6. Sanada Kuma Vanda B.P.Mok apical & slow VWN VWN, MS VWM

x Vanda Bill axillary

Sutton

7. Botanic Gardens _ unregistered apical & slow VWN VWN VWM

serial No. 3102 hybrid axillary

Aranda Wendy Scott ‘Green Field’

Aranda Freckles

Lim-Ho: Orchid Tissue Culture 157

TABLE 3. NUTRIENT MEDIA COMPOSITIONS FOR TISSUE CULTURE OF LOCAL ORCHIDS

(ALL VALUES IN UNITS OF MILLIGRAMMES PER LITRE)

VWN VWN VWS- VWS-_ VWM KC KCS KCBP MS MS#+2-4-D+ Tch

(Liquid) (Solid) (Liquid) (Solid) (Solid) (Liquid) (Solid) (Solid) (Solid) | BA (Solid)

(Liquid)

Ca3 (POQ4) 5 200 200 200 200 200 200

KH, PO, 250 250 250 250 250 100 100 100 170 170 250

KNO3 525 525 525 525 525 1900 1900 525

MgSO, 250 250 250 250 250 100 100 100 370 370 250

NH, SO4 500 500 500 500 500 400 400 400 500

MnSO4 (ES) 75 Tes fies) Ths 0.1 0.1 0.1 16.9 16.9 1S

FeSO4.7H,0 27.84 27.84 27.84 27.84 27.84 27.84 27.84 27.84 27.84 27.84 27.84

Na, EDTA 37.24 37.24 37.24 37.24 37.24 37.24 37.24 37.24 37.24 37.24 37.24

Ca(N 03), 4H,0 400 400 400

NH,NO3 1650 1650

ZnSO4 1.0 1.0 1.0 8.6 8.6

CuSO4.5H,0 0.03 0.03 0.03 0.025 0.025

CaC1,.2H,0 332 332

KI 1 1 1 0.83 0.83

CoC1,.6H,0 0.025 0.025

H3 B, 03 1 1 1 6.2 6.2

Na,Mo0,4.2H,0 0.25 0.25

AIC13 0.03 0.03 0.03

Thiamine HC1 1 1

Myoinositol 100 100

Adenine sulfate 40 40

Nicotinic acid 5 5

IAA 0.1-0.5 0.3

NAA 0.2

6-amino-benzyladenine 0.5-2

2-4-D 1-2

Coconut milk 150ml 150ml 150ml 150ml 75ml 150ml 150ml 150 ml 150 ml 150 ml

Active charcoal 150

Tomato juice 75 ml

Banana 75000 75000

Pineapple juice 150

Sucrose -_ 1000-2000 1000-2000 2000 1000-2000 1000-2000 1000-2000 — 2000

Agar = 800-1000 = 800-1000 1000 - 800-1000 800-1000 800-1000 — 800-1000

Distilled water 850 ml 850 ml 850ml 850ml, 925ml 850ml 850ml 850 ml 850 ml 850 ml 850 ml

Vanda Sanada Kuma Vanda Chia Kay Heng

Gard. Bull. Sing. 34:1 (1981)

158

ORCHID SPECIES

winuvaoun] WNIpIIUC

oqdadng, suaiqaadns wniqoapuaq

159

Orchid Tissue Culture

Lim-Ho

ORCHID HYBRIDS

6L8p ‘ON [eleg suepiey o1uejOog WnIqoup

S176 ‘ON [Plas suopiey s1uejlOg wnigospuag

uaq

YM JOY BUOX wniqodpuagq

uOX V9YD suosyD wnigodpuaqg

160 Gard. Bull. Sing. 34:1 (1981)

REFERENCES

Fu-Fan, May Lay (1978). Studies on the tissue culture of orchids 1: Clonal propagation of Phalae-

nopsis by lateral buds from flower stems. Orchid Review 86, 308—310.

_ (1979). Clonal propagation of Aranda, Assocenda, Cattelya by leaf tissue culture.

Gard, Bull. Sing. 31:2, 132-138.

Knudson, L. (1946). A new nutrient solution for germination of orchid seeds. Am. Orch. Soc. Bull.

15, 214—217.

Kunisaki, T.J., K.K. Kim, and Y. Sagawa (1972). Shoot tip culture of Vanda. Am. Orchid Soc. Bull.

41, 435-439.

Letham, D.S. (1974). Regulation of cell division in plant tissues XX. The cytokinins of coconut

milk. Plant Physiol. 32, 66—70.

Morel, G.M. (1960). Producing virus-free Cymbidium. Am. Orch. Soc. Bull. 23, 473—478.

Murashige, T. and F. Skoog (1962). A revised medium for rapid growth and bioassays with tobacco

tissue culture. Plant Physiol. 15, 473—497.

Singh, H. (1976). Meristem tissue culture of Dendrobium Ng Eng Cheow. Gard. Bull. Sing. 28:2,

259-267.

Vacin, E. and F. Went (1949). Some pH changes in nutrient solution. Bot. Gaz. 110, 605-613.

White, P.R. (1943). A Handbook of Tissue Culture. Jacquens Catell Press; Lancaster, Pennsylvania,

USA.

Wimber, D.E. (1963). Clonal multiplication of Cymbidiums through tissue culture of the shoot meris-

tem. Am. Orch. Soc. Bull. 32, 105—107.

. (1965). Additional observations on clonal multiplication of Cymbidium through

culture of shoot meristem. Cymbidium Soc. News 20(4), 7—10.

161

A NEW KEY FOR BIOLOGICAL IDENTIFICATION

‘

K. SWARUPANANDAN, A. R. R. MENON. AND K. BALASUBRAMANYAN

Division of Ecology

Kerala Forest Research Institute, Peechi-680 653

Trichur, Kerala, India

ABSTRACT

Another biological identification key, the synoptical dial key, has been devised. A

description of this key, its operation, and improvements over other keys are discussed.

INTRODUCTION

Biologists are familiar with the various types of identification keys such as dicho-

tomous keys, polyclave punch cards and their various modifications; the commonest tools

of systematic biology (Bianchi, 1931; Ceska and Trumpour, 1979; Leenhouts, 1966,

1967; Mayr et al., 1953; Metcalf, 1954; Pankhurst, 1974; Saldanha and Kameswara Rao,

1975; Tomsovic, 1976; Voss, 1952, and others). All these keys are very difficult to

handle in the field; also whenever the number of species increases, identification becomes

all the more difficult. Aware of these difficulties, the authors have been attempting to

develop a key that will serve the purpose of the field biologists. These efforts resulted in

the invention of a simple synoptical dial key which is the subject of the present paper.

THE SYNOPTICAL DIAL KEY

The present device is also a modified version of the dichotomous key where all the

dichotomies are condensed into a single node. The system has been devised to combina-

tion reading between any of the characters in a series of complementary set of characters,

as in an n-coordinate device.

The functioning device (Plate I) comprises of a series of concentric discs of increasing

diameters made of thick paper. The largest one is pasted to a hardboard disc of equal

diameter and all the others are superimposed on it in the order of decreasing diameters.

Thus, only a peripheral ring of about one centimeter width of each of the discs and the

smallest one completely are exposed. All the discs can be rotated around a central

common axis. A narrow sector is marked in each disc and completely darkened for

distinction. These sectors are marked ¢ (zero, marked as ¢ to distinguish from the English

letter ‘O’). The discs are numbered I, II, III and so on (to n) in Roman numerals in

a centripetal sequence, just on the left of the ¢s. Thus, the zeros in the different discs

function as a ‘zero-set’.

Different characters of the taxa to be included in the key are studied and categorised

into different ‘major characters’, each comprising a set of complementary characters

162 Gard. Bull. Sing. 34:1 (1981)

(Chart I). Each disc is meant for recording the complementary set of characters within

a major character. Thus, the number of discs can be increased or reduced according to the

number of major characters selected. The exposed peripheral portion of the discs are

marked into several divisions as there are complementary characters in each major charac-

ter plus a few empty divisions. The empty divisions serve to.add some more characters if

found necessary. All the divisions are indexed by lower case English letters a, b, c, and

so on (to z) on the extreme upper right corner of the divisions. The respective com-

plementary set of characters is written in the individual divisions sequentially, in all

the different discs, so that each character is indexed by a letter, each letter being dis-

tinguished from the same letter of other discs by the disc number. Each division of

the discs is also furnished with small outline diagrams of the corresponding characters.

On the back of the disc device is a printed chart (Chart II). The chart includes as

many vertical columns as there are discs in the device, corresponding to each major

character, arranged in the sequence of the arrangement of discs. The major characters

are written on the top of the columns and indexed with corresponding Roman numerals

as in the discs. The characters of the different taxa included in the key are coded in terms

of alphabetic indices (as given in the discs) and written in the various columns. These

codes standing for the various taxa are further arranged in the alphabetic sequence;

zero (@) following the last letter in each disc. After the codes starting with a particular

letter, space for the inclusion of a few more codes is left blank. The name of the

taxa corresponding to the codes are written on the right of the columns. Some distinct

characters of the taxa that are not given in the discs are also included in the appended

chart, wherever possible, to facilitate confirmation.

The names of the taxa are serially indexed by Arabic numerals. These numerical

indices of all the taxa that share a particular character are also repeated in the discs as

in Leenhouts’ synoptical key. Such cross indexing is done for all the characters. Whenever

a taxon included in the key has some characters peculiar to it only, its numerical index

is given in block numerals in the discs. The alphabetic index for that character in the code

standing for the taxon is put in squares also.

PROCEDURE OF IDENTIFICATION

First zero-set the device and find out any one of the prominent characters of the

specimen in question of identity. Locate the character in the discs by determining the

major character to which it belongs. If the alphabetic index for that character bears a

bold number, the corresponding name can be obtained from the appended chart. If the

alphabetic index for the character does not carry a bold number, it is coincided with the

zeros (gs) of all the other discs. A second prominent character of the specimen is located

in another disc; if the alphabetic index for that also does not carry a bold number,

coincide it with the alphabetic index of the formerly located character. Go through the

number given at the bottom of these characters, and find the ones common to both.

This process of location of characters, coinciding, and finding out the common numbers

is continued with more characters till all except one are eliminated and the specimen is

identified as the taxon bearing the common number in the enumeration chart.

In this key the alphabetic codes given in the enumeration chart are scarcely used.

If necessary, the codes can be eliminated from the chart also; but the retention of the

codes is advantageous. By coinciding the discs according to the alphabetic code for a

given taxon, a skeletal description of the taxon can be obtained. Thus, if ‘Synoptical Dial

Keys’ are employed in Floras, descriptions can be minimised further.

Swarupanandan, Menon, Balasubramanyan: Biological Key 163

A SPECIMEN ILLUSTRATION

An example to illustrate the identification method is given below:

As described above, a ‘dial device’ was prepared (Plate I and Chart II) to identify the

seedlings of fifty Asian tree species, mostly using the descriptions in Burger’s (1972)

Seedling Flora. The different major characters and their complementary sets within,

employed in the key preparation, are given in Chart I. A drawing of the seedling of Vateria

indica Linn., is given (Plate II). The prominent characters of the seedling are the presence

of (1) simple, entire, stipulate leaves with pinnate nerves; (2) basal nodes with whorled

phyllotaxy, and (3) the two, thick, anisomorphic cotyledons with similar halves and areo-

late surfaces.

The procedure of identification begins with zero-setting the device (Plate III

Fig. 1). The first character pertains to leaf and is held in the division ‘b’ of disc I. Since

this character is shared by a large number of taxa (Plate III Fig. 2 — as there are more

than one number in the division) coincide the alphabetic index ‘b’ with zeros of all other

discs (Plate III, Fig. 2). The next character pertains to phyllotaxy, and is held in the

division ‘b’ of the VIIth disc. Coinside ‘b’ with ‘b’ of the first disc (Plate III, Fig. 3). Since

there is more than one cross over number (3 and 5) some more characters are to be

taken up for identification. The third character pertains to the cotyledons and is in the

division ‘¢’ of the IInd disc. Since ‘e’ bears a bold number ‘3’ the taxon is immediately

identified as Vateria indica from chart II, the corresponding name for the number. Also

by coinciding ‘e’ of the IInd disc with the formerly selected indices (Plate III, Fig. 4),

again we will get a single crossover number 3 and hence the taxon is identified.

ADVANTAGES OF THE PRESENT DEVICE

The present system shares many advantages of both punch cards and Leenhouts’

(1966 and 1967) synoptical key. Preparation of the key is simple with the procedure

being the same as for Leenhouts’ synoptical key. Characters or taxa can be added,

dropped, combined or split easily without any remake. Incompletely known taxa can be

included in the key without any difficulty. To a certain extent, imperfect specimens can

also be identified. If the discs are made of thin cellophane or plastic sheets, several

such devices can be included in Floras. Since identification gives a skeletal discription

of the taxon, use of synoptical dial keys in Floras will help to reduce descriptions further.

Unlike Leenhouts’ synoptical key, alignment of the characters is possible here without

the interruption of characters in-between. Finally, the most important advantage is that

the key, being in one unit, is very convenient to work in the field.

ACKNOWLEDGEMENTS

The authors are thankful to Sri. R. Vasudevan Nair, Government Victoria College,

Palghat; Prof. B. K. Nayar, University of Calicut; Dr. P. M. Ganapathy, Director, Kerala

Forest Research Institute, for their helpful criticisms and valuable suggestions. We also

wish to express our sincere thanks to Sri. P. M. Jacob and Sri. M. Muraleedharan Pillai

of the Kerala Engineering Research Institute, for their technical advice.

164 Gard. Bull. Sing. 34:1 (1981)

REFERENCES

Bianchi, A.T.J. (1931). Een Niewe Determinatie — Methode (A new method of indentification),

Tectona 24: 844-893.

Burger Hzn., D. (1972). Seedlings of some Tropical Trees and Shrubs mainly of South East Asia.

Centre for Agricultural Publishing and Documentation, Wageningen.

Ceska, A. and A.D. Trumpour (1979). Computer Editing of Serial and Indented Identification Keys.

Taxon 28: 329—335.

Leenhouts, P.W. (1966). Keys in Biology I and II. A survey and proposal of a new kind. Proc. Konik.

Nederl. Akad. Wentesch. (Amsterdam) 69 C: 571—596.

. (1967). A New Kind of Botanical Identification Key. Malay Forester 30: 104—108.

Mayr, E., E.G. Linsley, and R.L. Usinger (1953). Methods and Principles of Systematic Zoology.

McGraw-Hill, New York.

Metcalf, Z.P. (1954). The Construction of Keys. Systematic Zool., 3: 38—45.

Pankhurst, R.J. (1974). Automated Identification in Systematics, Taxon 23: 45—51.

Saldanha, C.J. and C. Kameswara Rao (1975). A Punched Card Key to the Dicot Families of South

India. Centre for Taxonomic Studies, St. Joseph’s College, Bangalore.

Tomsovic, P. (1976). Uncovaci Klice a Jejich Konstruce (Identification keys and their construction)

Preslia 48: 355—367.

Voss, E.G. (1952). The History of Keys and Phylogenetic Trees in Systematic Biology, J. Sci. Labor.

Denison Univ, 43: 1—25.

ed, note.

The following chart also has relevance to this article: Haflinger, E. and H. Eisenhurt (1975).

Circular Key for Identifying the Plant Families With Weeds of Worldwide Importance.

Supplement to Weed Tables, Ciba-Geigy; Basle, Switzerland; 1 page chart.

Swarupanandan, Menon, Balasubramanyan: Biological Key 165

Se;

a‘

6e "<1u9 Pessns. e1e2—s)

218 Bu 0404 09

7 ©181U0D s ©} eu 0812

MOdW! dwoo esu0 87 un

A synoptical dial key prepared for the identification of the seedlings

50 tree species.

Plate I.

[44

Perseu eseujed

166

Gard. Bull. Sing. 34:1 (1981)

CHART I

A list of major characters and their complementary sets

used in the preparation of the sample Dial-key

DISC I — FORM OF NRL. LEAVES

Nrl.

Nr.

Nrl.

Nil.

Nii.

Nrl.

Ls. simple, entire, parallel nerved

Ls. simple, entire, pinnate nerved

Ls. simple, crenate/serrate/dentate

Ls. simple, 2-lobed, palmate nerved

Ls. simple, 3—5-lobed, palmate nerved

Ls. simple, 3—5-lobed, parallel nerved

Ls. palmate comp.

Ls. once comp., imparip., entire

Ls. once comp., imparip., crenate/dentate/

serrate

Ls. once comp., parip., entire

Ls. twice comp., imparip., entire

Ls. twice comp., imparip., crenate/dentate/

serrate

Ls. twice comp., parip., entire

DISC Il — COTYLEDONS

Cots

Cot.

Cots

. absent

single, terete

. 4-5, terete

. 7-10, terete

. 2, thick*, anisomorphic, 2s similar

. 2, thick*, anisomorphic, %s dissimilar

. 2, thick*, isomorphic, “%s similar

. 2, thick*, isomorphic, %s dissimilar

DISC III - COTYLEDONARY SHAPE

Cots.

orbicular

lalongate, reniform/subreniform

lalongate, obreniform

lalongate, obtuse-trapeziform

lolongate, 3-lobed

lolongate, 5-lobed

lolongate, entire, falcate-curved

lolongate, entire, cordate

lolongate, entire, ovate/obovate

lolongate, entire, obtuse-tetragonal

DISC IV — COTYLEDONARY TIPS

Cot. tip acuminate

Cot. tip obtuse

Cot. tip round

Cot. tip emarginate

Cot. tip retuse

Cot. tip truncate

Cot. tip oblique-truncate

DISC V — COTYLEDONARY BASES

Cot. base acute-/round-sagitate

Cot. base cordate/round

Cot. base 1 half hastate

Cot. base 1 half with a denticle

Cot. base acute

DISC VI — PHYLLOTAXY OF NRL. LEAVES

Nrl. Ls. whorled

Nrl. Ls. opposite

Nil. Ls. opposite+whorled

DISC VII — BASAL LEAVES

Basal Ls. of scales

Basal Ls. foliaceous, whorled

*Thickness more than four millimeters.

Abbreviations: comp. — Compound;

Cot. — Cotyledon; Cots. — Cotyledons;

imparip. — imparipinnate; Ls. — Leaves;

NRL. — /Nrl. — Normal; parip. — paripinnate.

Swarupanandan, Menon, Balasubramanyan: Biological Key

CH sh So

The enumeration chart appended to the serree lialekey

(to be printed on the back of the ice)

= =

=.) > H :

4 > Columns correspond to the ciscs, 8

R arranged in centripetsel sequence

o €

y 2 +

® a a 4 3 SCeSe Cel SSSS SSS SSSSSSTSSSSSSSTHSSSESSHSSSSSSSSSESSSSSESSSS Pr

5 £&£-xu GH ae i

rw eae © 2 2 £

-kéé 66 é Pen ER Characters not given ;

Sess sg “ ee in the discs

no See a k

a ew ef & ev et c.)

E88882 a4 -

a Agathis GAMMATA ceocveccccecece St em with scale le. eece 1

Garcinia parvifolia eoeccece eeeceseseseesceseseeeee® 2

Vateria IVGICA. acast ceases coe Cots. areolate e@cesesese

Calophyllum inophyllum eevee @eeeeseeeeeeeeeeeee2ee2ee2e2e68 ¢

Lithocarpus kostermansii ... Hypocotyl Mil] wccccccece 5

Lurio zebethianus e.ecccccces Secor ercesccesessesesesecs 6

Quercus lineata cecceccccses Hypocotyl short ..cccccoe 7

Gluta renghas eeeseeoeoeaeoee eee @eeeeeeeseeeeeee2ee20202060080 8

Xylocarpus granatuM .cccceee BYBOCOLY I LED cs cccconece

Buchanania arborescens ecooe Te gee ee 0

Eucalyptus globulus eeocvecer eocccccccccccccccccccccs§ F1

Terminalia bellerica eeeeees @eeeeseveeeeeoeeeeeee eee eee 12

lTerminalia catapa eveveecsecce SCceecesessesooecesseosessese Ar

Cananga odorata cescccceccccs ecereecseseseseseseeseses 4

Alstonia angustiloba eeeeoeee @eeeoveeaeeoeceeeeeeeeee ee e@ 15

Phyllanthus emblica ceccccece le. bifarious ccccecccce 16

S BVessam assarsam fF] wy Ba aos Qeaoavaseussanmamep we

fe] Bee Sooo [oa] fh) @@ [8] 2020 eeoreeerercrrC00"0"" [a

kw FHSAA

Abbreviations: Cots.= Cotyledons;

en ro raf] erry mp H A HEA KH BUH OHHH) Oe He HD/]e eS »

EEG BerD

oecororo [p} Boo o cAA HofMooHHAAaGoOroBO BE Co

oa s aAcso™n

2am "m0"

Ba BH ABOPM fal oblo wn ao ea QR ST 2BBQO WEP GavrvBVausvVprpoeorsa Co

ow om oe

BS MOOS BOOWOBWAS BS B BO B oOo Boer BODO Sade so aT o

BSBassa.qa ~®wWB

~ Bo

A RQQVVAA BVVSVVasasB®e Bo BB VQ HH WAN BVNBVABVAWVWSSa! OP BP of oP

So BBO

Santalum album

Antidesma montanum ecceccecece

Flacourtia indica c.cccccces

Tectona grandi8 ceccoccceces

Edonymus javanicus eeecscece

Piliostigma malabaricum ....

Gmelina elliptica eeeccessee

Aleurites molluccana ececccee

Corypha utan eecesscccccvece

Bombax ceiba @eeeeveveeeeneeee8@

Lysoxylum densiflorum ....e-

Swietenia mahagoni ..ceeeees

Erythrina subumbrans ceccece

butea MONOSPECIMA eccesececee

Micromelum minutum o.....ce8

Pterocarpus indicus sece-cece

Lracontomelon puberulum ....

Dalbergia latifolia s.sccoees

Azaderachta indica ..ccrrcce

Aegle marmelos e@eeeoeeeveee ee

Protium javanicum ..cceccecs

Garuga floribunda ......-e0%

Bischofia javanica eeereeese

Vitex quinata seccreeeseeees

Cassia siamea eeeeeveeevee ee eee

Radermachera gigantea eesees

Melia azaceracn ceeeeseeseeee

Pithecellobium jiringo ...e.

Adenanthera pavonina eeseres

Acacia arabica «sesee-

FPeltophorum pterocarpa esses

Cupressus lusitimice sesseee

Pinus merkusii wescceseseees

Casuarina equisetifolia «ee.

Ls./lese= Lenves;

coilar swollén ..

Stipules spiny ..cccccce

Stem tetragonal .cccccce

Hypocotyl BShOrt secccece

Pinna: 3 06s 6 kind -<<0 ee

Pinnae 3 @cceceseseoeses

‘Pinnae 3, stem thorny .«.

Cots. palmate nerved ...

Pinnae 3 eeeeoeeeeeeeeee8

Cots. elliptic eeeeesees

Leaflets alternnte sees

hrl. ls. needle-like ..

seeeeeeeeeeeeeeeeeeeeeeees

hrl.e hormnl,.

168 Gard. Bull. Sing. 34:1 (1981)

Plate Il. A sketch of the seedling of Vateria indica Linn.

Swarupanandan, Menon, Balasubramanyan: Biological Key

Plate III.

3 Mple

Paraliel- nerve gilire

dentate,

19 20 21

Nel Ls gj

dentate » .

19 20 21

Cots 2 tin;

1/28 dissj

169

Figs. 1—4. Different stages in the procedure of identification of seedlings

of Vateria indica with the sample synoptical dial key.

170

CHANGES IN THE FERN FLORA OF GUNONG ULU KALI*

A. G. PIGGOTT

9A, S.P.B. Towers, Jalan Batai, Damansara Heights, Kuala Lumpur, Malaysia.

ABSTRACT

Changes in the fern flora of Gunong Ulu Kali, Malaysia, resulting from the develop-

ment there of a hill resort, are discussed. Fourteen fern species new to the mountain are

recorded from near the summit, 5,814 ft (1772 m), and a further thirteen from tall

montane forest at about 5,000 ft (1500 m).

INTRODUCTION

The realisation that large areas of the natural vegetation of Gunong Ulu Kali would

inevitably be destroyed during development of the Genting Highlands resort was a major

reason for recording the fern flora there. A list of species occurring above 5,000 ft

(1524 m) was compiled during the four years following the opening of the first hotels

(Piggott, 1977). The resort, which provides recreational facilities giving pleasure to

thousands of people, continues to expand. The mushroom farm was moved to a larger site

on the slopes below Location F and the original site converted into a flower nursery and

vegetable garden. Although the buildings remain, the new site has now been abandonned.

Colourful gardens now surround the resort buildings. Exotic ornamentals have been

introduced, and with them some weeds (Stone et al., 1977). Development of the area has

resulted in changes in habitat and, consequently, changes in the composition and distribu-

tion of the flora. Pioneer species are common in cleared areas and tangled thickets are

covering the exposed edges of the forest. Many of these species were previously present

only in small numbers and are now rapidly multiplying. Fourteen species new to G. Ulu

Kali have been recorded along the ridge between Locations B and I, and a further thirteen

along the 1500 m contour. Specimens have been deposited in the herbaria at Kew (K) and

Singapore (SING). Changes in the fern flora of the different habitats occurring on the

mountain are discussed in the following paragraphs.

ROADSIDE VERGES AND DRAINS

The road along the ridge had not long been constructed when the fern flora there was

first examined. Now the verges and drains have matured and weathered, and in places

fallen forest debris and sand have accumulated. Previously only infrequent plants of

Christella arida (Don) Holtt., Nephrolepis tuberosa (Bory) Presl, Pteris vittata L. and

Sphaerostephanos polycarpus (Bl.) Holtt. were seen. These have now become common.

Equisetum debile Roxb., first recorded as a single colony near Location C, is very

common on sandy patches and small cleared areas throughout the length of the ridge.

This fern ally has recently appeared at Fraser’s Hill and is spreading in similar habitats.

* q.v. “The Ferns of Gunong Ulu Kali”, Gard. Bull. Sing. 30 (1977) 31-43.

Piggot: Fern Flora 171

New record for G. Ulu Kali: Pseudophegopteris rectangulare (Zoll.) Holtt. at Loca-

tion E, collections 2627 (K) and 2628 (SING) of 26.10.80.

GRANITE WALLS

Walls were necessary at several places along the ridge. The new walls were not a

suitable habitat for plant growth and only occasional stunted plants of P. vittata and

S. polycarpus were recorded on them. Now they have weathered a little, and debris

has lodged between the granite rocks. More ferns are beginning to grow there. Three

species new to the summit, but recorded at lower altitudes on G. Ulu Kali, are:

Aglaomorpha heraclea (Kze) Copel. at Location H, collections 2357 (K) and 2358 (SING)

of 29.04.79.

Drynaria rigidula (Sw.) Bedd. at Locations B, F and H, collection 2359 (K) of 29.04.79.

Pyrrosia floccigera (Bl.) Ching at Location H, collections 2360 (K) and 2361 (SING)

of 29.04.79.

SMALL CLEARINGS AND GENTLE EARTH SLOPES

The acid peat surface soils and acidic subsoils (Piggott, 1978) were removed from

small clearings on the ridge at the time the road was constructed, leaving more neutral,

though often shallow, sandy weathered rock. These areas are being covered by carpets of

Crawfurdia trinervis, with purple berries, and thorny tangles of Rubus moluccanus, wild

raspberry. Here Pityrogramma calomelanos (L.) Link, Diplazium speciosum Bl. and Pteris

tripartita Sw. are frequent. Some plants of Plagiogyria tuberculata Copel. have also been

found in this rather exposed habitat. Pteridium aquilinum (L.) Kuhn var. wightianum

(Ag.) Tryon grows on the edges of the clearings. Gleichenia hirta Bl. var. amoena

(v.A.v.R.) Holtt. is now quite common on some of the gentle earth slopes. Five species

new to G. Ulu Kali were found in this habitat:

Diplazium esculentum (Retz.) Sw. at Location H, collections 2354 (K) and 2355 (SING)

of 29.04. 79.

Goniophlebium subauriculatum (Bl.) Presl at Location I, collections 2585 (K) and 2586

(SING) of 27.09.80.

Hypolepis brooksiae v.A.v.R. at Location H, collection 2695 (K and SING) of 01.01.81.

Microlepia speluncae (L.) Moore var. hancei (Prantl) C. Chr. at Location B, collection

2402 (K) of 14.10.79.

M. strigosa (Thbg.) Pr. at Location F, collection 2401 (K) of 14.10.79.

Plants matching the unidentified sterile Microlepia sp., collections 1175 (K) and

1176 (SING) of 09.11.74, which was under observation but lost during road improve-

ments, have been found in the same vicinity at Location G. A few sori were produced

and the species identified as M. todayensis Chr., collections 2352 (K) of 29.04.79, 2366

(SING) of 10.05.79 and 2644 (K) of 22.11.80.

172 Gard. Bull. Sing. 34:1 (1981)

LARGE CLEARING AT LOCATION H

Very little plant growth has occurred in this area, mainly due to its frequent use by

vehicles and as a camp site. P. a. var. wightianum, Hypolepis beddomei (Kunstler) Nair &

Ghosh (former name: H. punctata (Thbg.) Mett.) and some Gleicheniaceae are growing

along the outer edges.

ROAD CUTTINGS AND EARTH BANKS

Previously infrequent and often stunted, large plants of Blechnum orientale L. are

now not uncommon in this habitat on the ridge. Dicranopteris curranii Copel., D. linearis

(Burm.) Underwood var. linearis Holtt. and-D. I. var. montana Holtt. have covered some

exposed areas. Dipteris conjugata Reinw. has colonised others, often in association with

Sphenomeris chinensis (L.) Maxon var. divaricata (Chr.) Kramer. The deeper vertical road

cuttings are not yet much colonised and only a few small plants, some of them fallen

epiphytes, have appeared in cracks and erosion gulleys.

MOSSY BANKS ABOVE ROAD CUTTINGS

The mossy peat rootmat of the floor of the dwarf forest has been exposed along the

top of earth cuts. Where erosion has occurred it overhangs and is well-drained, supporting

colonies of Gleichenia vulcanica Bl., Coryphopteris gymnopoda (Bak.) Holtt. and C. g.

var. bintangensis Holtt. The two varieties of C. gymnopoda are present in about equal

numbers in this habitat, together with apparently intermediate forms. In shade the plants

produce rosettes of fronds, but on exposed rootmats this character is less evident. The

fronds appear evenly distributed and assume a drooping habit, although not wilted.

EDGE OF FOREST

Fern growth along the exposed edges of the forest is similar to that occurring in

upland areas throughout West Malaysia, and consists mainly of thicket-forming species.

Along the ridge these are G. longissima Bl.,G. truncata (Willd.) Spr. var. plumaeformis

(Presl) Holtt. and Histiopteris incisa (Thbg.) J. Sm. In several places G. vulcanica has also

formed delicate miniature thickets. Although trunk growth in Cyatheaceae is slow and

few tall tree ferns can yet be seen in this habitat, fronds of Cyathea contaminans (Wall. ex

Hook.) Copel. are now penetrating the thickets. At lower altitudes, along the road from

Genting Simpah to the hotels, tall tree ferns of this species are common on the edge of

the forest. Pyrrosia floccigera was recorded as an igen ba in an exposed position on the

edge of the forest at Location E.

DWARF FOREST

The less accessible areas of the ridge forest are still relatively undisturbed. Where

access is fairly easy, the forest has been penetrated by footpaths and camp sites have been

cleared. Culverts, constructed when the road was built, have improved the drainage of the

highest parts of the forest. The habitat has become drier and many of the small delicate

epiphytes are now less abundant. Whereas previously many trees were clothed with

Hymenophyllaceae, it is now necessary to hunt for them in the shelter of the remaining

wet mossy hollows. Phymatopteris laciniata (Presl) Pic.Ser. (former name: Crypsinus

laciniatus (Presl) Holtt.) now appears to be more abundant, often on the higher branches

of the trees. Matonia pectinata R. Br. has spread where the dwarf forest has been

ee and become more open. Four small fern epiphytes new to G. Ulu Kali were

collected:

Piggot: Fern Flora 173

Calymmodon gracilis (Fee) Copel. at Location E, collection 2630 (K) of 26.10.80.

Grammitis hirtella (Bl.) Tuyama var. coriacea at Location F, collection 2440 (K) of

218:

Scleroglossum debile (Kuhn) v.A.v.R. at Locations B and I, collections 2568 (K) of

22.03.80 and 2632 (SING) of 26.10.80.

Trichomanes digitatum Swartz at Location I, collection 2633 (K and SING) of 26.10.80.

ROAD TO WESTERN DAM, LOCATION L

During his visit to Malaysia in 1978, Dr. R. E. Holttum collected specimens of

Chingia perrigida (v.A.v.R.) Holtt., growing in fairly open scrub at 5,000 ft (1524 m).

Earlier specimens from the same place had been incorrectly named C. pseudoferox

Holtt. This is a new record for W. Malaysia.

STEEP SLOPE WITH BOULDERS

The sheltered area southeast of ‘Sri Genting’ had not previously been fully investi-

gated. Here, a little above the 5,000 ft contour, tall forest covers a steep slope. There are

piles of large granite boulders, and small, wet, deeply shaded valleys. Many of the tree

trunks are mossy. This area is relatively undisturbed except along the line of the cable-

car route. A form of P. laciniata, differing in appearance from the same species in other

locations on the mountain, was frequent on mossy trees and rocks. The pendent fronds,

thin but firm, were glaucous above and very glaucous below. Most fronds were simple

or trilobed, and many were fertile. Oleandra pistillaris (Sw.) C. Chr. on mossy boulders

exhibited a creeping habit rather than the usual shrubby growth. Filmy ferns were

common on mossy trunks and rocks. The following fern species, collected from this

location, are new to G. Ulu Kali:

Asplenium affine Sw., collection 2497 (K) of 13.01.80.

Diplazium malaccense Presl, collection 2495 (K) of 13.01.80.

D. subintegrum (Holtt.) Holtt., collections 2424 (K), 2425 (SING), 2427 (K) and 2428

(SING) of 04.11.79.

D. tomentosum (B1.) Milde, collections 2429 (K) and 2430 (SING) of 04.11.79.

Elaphoglossum callifolium (B1.) Moore, collection 2500 (K) of 13.01.80.

Hymenophyllum denticulatum Sw., collection 2453 (K) of 02.12.79.

H. polyanthus Swartz, collection 2457 (K) of 02.12.79.

Phymatopteris triloba (Houtt.) Pic. Ser. (former name: Crypsinus trilobus (Houtt.)

Copel.), collection 2464 (K) of 02.12.79.

Trichomanes idoneum Morton, collection 2455 (K) of 02.12.79.

T. maximum Bl., collection 2465 (K) of 02.12.79.

174 Gard. Bull. Sing. 34:1 (1981)

T. proliferum Bl., collection 2461 (K) of 02.12.79.

T. saxifragoides Presl, collection 2444 (K) of 02.12.79.

CONCLUSION

The genera Calymmodon, Grammitis, and Scleroglossum are mostly epiphytes of

mossy trees in mountain forest. Some species had already been recorded on G. Ulu Kali.

C. gracillis, G. hirtella var. coriacea, and S: debile have been little collected in W. Malaysia

but their occurrence in dwarf forest on the ridge could be expected. T. digitatum is not

common but has been found on mossy trees at lower altitudes in the mountains and also

on mangrove trees in the southern part of the peninsula.

Aglaomorpha heraclea and Pyrrosia floccigera are fairly common epiphytes of

mountain forest to about 4,000 ft (1,220 m) and have been recorded at that altitude

nearby. Drynaria rigidula is widely distributed, but not common, in the northern half of

the peninsula. It occurs at Fraser’s Hill, only about 20 miles (32 Km) to the north, and

also on the lower southern slopes of G. Ulu Kali. It is not, therefore, unlikely that spores

could be carried to these higher altitudes and lodge in positions suitable for successful

germination.

Diplazium esculentum is a fern of wet ground in open places in the lowlands. Al-

though the author has recorded this species at Fraser’s Hill and also near the southern end

of the Main Range, its occurrence on the ridge was quite unexpected. Microlepia spe-

luncae var. hancei is again predominantly a lowland fern, but has occasionally been

collected in the mountains. Neither M. strigosa nor M. todayensis is common in W. Malaysia,

although both occur in a few places in the mountains. Goniophlebium subauriculatum,

Hypolepis brooksiae, and Pseudophegopteris rectangulare are all species which become

common in mountain clearings, such as occur at Cameron Highlands and Fraser’s Hill.

Only a single large plant of G. subauriculatum and one small colony of P. rectangulare

were recorded, but both were sporing profusely and could be expected to spread.

The twelve species new to G. Ulu Kali collected from tall sheltered forest near ‘Sri

Genting’ are not uncommon in W. Malaysia, and have been collected from similar situations

elsewhere. But Chingia perrigida is only otherwise known from central and southern

Sumatra and Java at 5,000 — 8,500 ft (1500 — 2600 m) (Holttum, 1981).

Many of the fern species found on G. Ulu Kali are common in W. Malaysia, some at a

wide range of altitudes, others only in the mountains. Several are confined to high moun-

tains with dwarf forest frequently in cloud. A few species have not been recorded else-

where in W. Malaysia but probably occur on botanically unexplored mountains, and are

known to occur in other countries. The only species which might possibly be endangered

is Elaphoglossum robinsonii, known only from G. Ulu Kali and nearby G. Mengkuang;

but even this is not in danger of becoming extinct. It is now growing in the Fern House

of the Royal Botanic Gardens at Kew, and spores have also been distributed to pterido-

logists for propagation in other parts of the world.

ACKNOWLEDGEMENT

I must thank Dr. R. E. Holttum for his help in identifying the fern specimens and

also for accepting aninvitationto visit Genting Highlands. It was a most enjoyable and

instructive expedition.

Piggot: Fern Flora 175

REFERENCES

Holttum, R. E. (1981) The Fern Family Thelypteridaceae in Malaya. Gard. Bull. Sing. 33, 1—30.

Piggott, A. G. (1977) The Ferns of Gunong Ulu Kali. Gard. Bull. Sing. 30, 31—43.

1978. Notes on Soil Types on a Fern-rich Tropical Mountain Summit in Malaya.

Brit. Fern Gaz. 428 — 429.

Stone, B. C., H. J. M. Bowen, & J. F. Veldkamp, (1977) European Weeds Introduced to Gunung

Ulu Kali, Genting Highlands, Pahang. Mal. Nat. J. 30:1, 103 — 108.

Plate 2. Christella arida (Don) Holtt. growing in

sand and debris which has accumulated

in a roadside drain.

Plate 1. Pteridium aquilinum (L.) Kuhn var.

wightianum (Ag.) Tyron growing in

the crack of a granitic outcrop.

| Plate 3. Equisetum debile Roxb. on an eroding

roadside bank.

a eu yp 3

176 Gard. Bull. Sing. 34:1 (1981)

Plate 4. Plagiogyria tuberculata Copel., Cory-

photeris gymnopoda (Bak.) Holtt.,

Gleichenia vulcanica Bl., and Dip-

lazium speciosum Bl. growing on an

exposed earth slope.

Plate 5. Cyathea contaminans (Wall. ex Hook.)

Copel. penetrating thickets at the edge

of the forest, c. 1524 m.

Plate 6. Gleichenia longissima B1., Matonia pectinata R. Br., and Dipteris conjugata Reinw. (left

to right) in a “thinned” dwarf forest.

Ivy

A NEW SPECIES OF PANDANUS (PANDANACEAE) FROM BURMA

BENJAMIN C. STONE

Dept. of Botany, University of Malaya,

Kuala Lumpur, Malaysia

ABSTRACT

Pandanus burmanicus sp. nov. is described from the Upper Chindwin District, Burma.

It belongs in the Subgenus Rykia (DeVr.) Stone, the Section Rykia, Subsection Bidens

(St. John) B. C. Stone. To date it is known only from the type locality.

The species of Pandanus in Assam, Burma, and Thailand are still incompletely

known. The following new species was collected more than half a century ago and had

been determined as P. furcatus Roxb., to which it is indeed allied, but it is specifically

distinct in its several-celled phalanges, with the carpels arranged in adnate transverse

series. The forked styles with adaxial stigmatic surfaces clearly indicate that the species

belongs in Subg. Rykia (DeVr.) Stone, Sect. Rykia. Among the species of this group it

appears to find a place near P. piniformis Holtt. & St. John, which till now has con-

stituted the sole member of St. John’s Sect. Bidens, which I regard as a subsection of

Sect. Rykia. The character of this subsection is that the phalanges are several-celled.

Whether this is an artificial character of convenience remains to be seen.

Pandanus burmanicus B.C. Stone sp. nov. (Figures 1, 2).

Arbor habitu P. furcato. Folia ad 248 cm longae, 6.8 cm latae, lineari-ensiformia

apicem versus sensim attenuato-acuminata, marginibus antrorse denticulatis, costa dorso

distaliter antrorse denticulato; marginibus in basi dentibus 2—3 mm longis, 4—10 mm sese

separatis; in medio, dentibus acicularibus 1 mm longis valde antrorsis 7-12 mm sese

separatis; in apice dentibus vix 0.5 mm longis, 1.5—3 mm sese separatis; nervis longi-

tudinalibus c. 120 per foliam; nervis transversis inconspicuis; basis vaginantibus cupreo-

brunneis. Inflorescentia foeminea terminalis, solitaria. Fructus c. 20.5 cm longis, 15 cm

diametro (48 cm circumferentio), e phalangibus compositus. Carpidia in phalangiis

transverse-seriatim adnata; phalangibus 3—4-carpidiatis; cuneatis, compressis, c. 45 mm

longis, ad 30 mm latis, c. 10—11 mm crassis; pileo delapso intacto 3—4-stylophoro;

stylis antrorse deflectis c. 7 mm longis et 4—5 mm latis, brunneis, nitidis, oblique actuis

vel furcatis, stigmata infra stylos posita sursum spectantia. Mesocarpium apicalium

fibroso-medullosum; basalium fibrosum. Endocarpium 3—4-loculatum c. 24mm longum

ferrugineum osseum. Cetera ignota.

Type: BURMA: Upper Chindwin District, Kodan stream near Kyoktha Village,

600—1500 ft. alt., 30 November 1917, C. G. Rogers 1033 (Holotype in Dehra Dun

Herbarium). Vernacular name: “‘sathwa”’.

The collector notes that this is ‘“‘the common screwpine of the forests in the Kodan

chaung.”

178 Gard. Bull. Sing. 34:1 (1981)

NY ‘wlll

CE Rogers 1033

Fig. 1 — Pandanus burmanicus Stone. (From holotype). Leaf apex; short segment from near leaf apex;

segment from lower third of leaf. Phalange in side, top, and longisection views. Side view shows

distal adaxial face.

179

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180

ORTHORRHYNCHIUM ELEGANS (Hook. f. et Wils.) Reichdt.

(PHYLLOGONIACEAE), NEW RECORD FOR THE PHILLIPINE MOSS FLORA

BENITO C. TAN

Department of Botany, CAS University of the Philippines at

Los Banos, College, Laguna, Philippines

ABSTRACT

Orthorrhynchium elegans is reported for the first time from the Philippine archipe-

lago. Some noteworthy morphological features are also reported.

A recent study of many undetermined moss collections made by Prof. J. V. Pancho

of the University of the Philippines from 1960—70 and deposited at the University

herbarium (CAHP) yielded many new records for the local flora. Among which

Orthorrhynchium elegans (Hook. f. et Wils.) Reichdt., previously known only from

Australia and New Zealand, is of phytogeographical significance, and is, therefore,

reported here separately together with some new morphological information.

The collection (Pancho 3915B, July 23, 1965) came from the vicinity of Mt. Apo

on Mindanao Island, the second largest island of the country which is situated north of

Borneo. The packet contains abundant sterile plants that show the characteristic com-

planate and distichous arrangement of the conduplicate leaves as described by Sainsbury

(1955) and Wijk (1957) (Figs. 1 & 2). The leaf apices are blunt (Fig. 3). This feature can

be used to distinguish the taxon from the morphologically similar Horikawaea phyllo-

gonioides (Sull.) Nog. which has abruptly acute and slightly recurved leaf apices. The

latter was listed in Bartram (1939) as Orthorrhynchium phyllogonioides (Sull.) E. G.

Britton.

Wijk (1957) was the first one to point out the important difference between O.

phyllogonioides and O. elegans in terms of their phyllotaxy. He reported, in addition,

the identification confusion which existed among the many herbarium collections of the

two taxa which he had examined and concluded that O. phyllogonioides be best included

in the genus Neckera in the family Neckeraceae. Noguchi (1961), on the other hand,

contended that the taxon be placed in Horikawaea, a new genus which he had proposed

in 1937 for the family Phyllogoniaceae, after fully stressing the salient morphological

differences between O. phyllogonioides and species of Neckera.

I have been fortunate to have had the opportunity to study the isotypes of Neckera

phyllogonioides Sull. and Orthorrhynchium philippinense C.Muell., and also a Bartlett

collection (15935a) named O. phyllogonioides by E. B. Bartram. All collections men-

tioned here are from the Philippines and are part of Bartram’s Herbarium at the Farlow

Herbarium. I fully agree with Bartram (1939) in uniting the taxa in question under the

single species epithet, O. phyllogonioides. Nevertheless, this species, as suggested by

Noguchi (1961), is best placed in Horikawaea. The justification has already been con-

vincingly outlined by Noguchi (1961) and will not be repeated here.

A more recent collection (Alvarez 0-80553) of O. elegans at CAHP, which originated

from the same general locality of Mt. Apo, contains some fertile specimens. Three mature

capsules with pilose, mitriform calyptra were available for detailed examination. The

Tan: Orthorrhynchium elegans 181

peristome teeth, lacking the endostome, are only rarely perforated along the median

line. The dorsal (or outer) surface of the exostome clearly shows the diplolepideous

nature (Fig. 13). The ventral (or inner) surface develops irregular longitudinal (vertical

columns) and horizontal thickenings (trabeculae) (Fig. 12) resulting in the formation of

a series of thin-walled areas which may look like perforations under the light microscope.

The apices of the teeth are slender and of uniform thickness and are often broken. It

will be interesting to reexamine the peristome of O. elegans under the scanning electron

microscope in order to determine the cell patterns as seen in the light of the recent

report of Edwards (1979) on haplolepidous peristomes.

This peculiar pattern of cell wall thickening of the primary peristomial layer (cf.

Edwards, 1979) in O. elegans is not known from any Philippine Neckeraceous taxa.

Instead, a similar condition can be observed in the peristome of a few species of Oedicla-

dium Mitt., Palisadula Toyama, and other genera of the family Pterobryaceae (cf. Iwat-

suki, 1979).

The stem of O. elegans has been observed to consist of more or less uniform cells

without a differentiated strand (Fig. 6). Pseudoparaphyllia appear late in the development

of buds. When present, both filamentous and foliose types are seen. Interestingly, the

Philippine plants bear abundant filamentous gemmae on the stem and leaf apices, a

feature not mentioned in Sainsbury (1955) for the New Zealand population. These

asexual propagules are septate and strongly papillose (Figs. 15 & 16),. resembling the

thizoids which are always larger in size.

Dwarf male plants (Fig. 9) are discovered for the first time in O. elegans, Plants

consisting of profusely branched protonema with a few leaves were found growing inside

the folds of the conduplicate leaves of the female plants. Like the perichaetial leaves,

perigonial leaves are not conduplicate and their leaf margins are irregularly laciniate to

serrulate in the upper half (Figs. 9 & 10).

The spores (Fig. 14) are papillose and are of two sizes: the larger ones ranging

between 32.5 — 40 w (with a few becoming elongate and bean-shaped), and the smaller

ones less than 20 win diameter. Whether or not there is a correlation between the spore

sizes (anisospory) and sexual dimorphism (cf. Ramsay, 1979) is not known at present.

The distribution of O. elegans was mapped by Noguchi (1961) to be confined to

Australia and New Zealand. Its discovery in the Philippines is a significant extension of

its range while its presence in New Guinea is highly possible. I have seen several collect-

ions of Orthorrhynchium from the Christmas Islands, Papua New Guinea, and elsewhere

in the Malesian region deposited at the University of Michigan (MICH), Singapore

Botanic Gardens (SING), and other herbaria. All of them are Horikawaea phyllogonioides.

O. elegans, together with Dawsonia superba Grev., are two noteworthy Austro-New

Guinean elements in the local moss flora.

ACKNOWLEDGEMENTS

I am much indebted to the curators of the following herbaria for the loan of

specimens: MICH, SING, FH, and MONASH. Also, to Dr. H. Bishler (PC) and Dr. W. B.

Schofield for the procurement of rare literature. Lastly, to the University of the Philip-

pines at Los Bafios for the financial aid in my research. The illustrations were prepared

by Mr. Ver Aguila.

182 Gard. Bull. Sing. 34:1 (1981)

REFERENCES

Bartram, E.B. (1939). Mosses of the Philippines. Phil. J. Sci. 68, 1—423.

Edwards, S.R. (1979). Taxonomic implications of cell patterns in haplolepidous moss peristomes,

pp. 317-346. Jn: Clarke & Duckett (eds.). Bryophyte Systematics. Academic Press.

Iwatsuki, Z. (1979). Re-examination of Myurium and its related genera from Japan and its adjacent

areas. J. Hattori Bot. Lab. 46, 257-283.

Noguchi, A. (1937). Studies on the Japanese mosses of the Orders Isobryales and Hookeriales. J. Sci.

Hiroshima Univ., B—2, 3, 37—S6.

_ (1961). Notes on the genus Horikawaea Nog. (Musci). Bull. Bot. Soc. Univ. Saugar 13,

27-30.

Ramsay, H.P. (1979). Anisospory and sexual dimorphism in the Musci, pp. 281—316. Jn: Clarke

& Duckett (eds.). Bryophyte Systematics. Academic Press.

Sainsbury, G. O. K. (1955). A handbook of the New Zealand mosses. Royal Soc. New Zealand Bull.

5, 1-490.

Wijk, R. van der. (1957). Distichous and pseudodistichous mosses. Acta Bot. Neerlandia 6, 386—391.

Figures 1—16. Orthorrynchium elegans (Alvarez 0—80553).

1. habit sketch,

De frond or secondary branch (arrow pointing at perichaetial bud),

Be leaf apex,

4. laminal cells,

5; leaf base,

6. cross-section of stem,

7—8. leaves,

9, male plant,

HO, perichaetium,

en exothecial cells,

| ome peristome tooth, ventral view,

Lids peristome tooth, dorsal view,

14. spores,

15—16. gemmae.

183

Tan: Orthorrhynchium elegans

—_

184

BOTANICAL NOTE

J. F. MAXWELL

Botanic Gardens, Singapore

How to Cite the Name of Miss Joaquim’s Orchid

The national flower of Singapore, Vanda Miss Joaquim, was officially announced

as such by the Minister of Culture on 16 April 1981. This hybrid, the first cultivated

orchid hybrid recorded for Singapore, was selected from many other contenders because

of its historical significance for Singapore plus the fact that it is easily grown, and

well-known. This hybrid also stimulated the development of the presently thriving

orchid industry in SE Asia and has been one of the favourites of orchid enthusiasts

for decades.

Along with the development of the orchid industry and the development of many

new hybrids, there has been an increasing amount of confusion concerning the technical

details on how to properly cite these new creations.

Plant Names

In addition to common names, which are often variable and inconsistent, there are

two more official and internationally recognized kinds of plant names viz. scientific or

botanical names and the names of cultivated plants. It is generally understood that the

scientific name of a plant consists of two Latin or Latinized words viz. a generic name

and a specific epithet, in addition to the author’s name (or names). For example, the

scientific name of the para rubber tree is Hevea brasiliensis (Willd. ex A. Juss.) M.-A.

Hevea (derived from the Brazilian name for these trees) is the generic name, brasiliensis

(from Brazil), the specific epithet, and M.-A. is an abbreviation of the name of the botanist

(author) who placed this species under Hevea (Jean Mueller of Aargau, a 19th century

botanist). In addition to this, varieties and other subspecific taxa are also frequently used

to delimit variation within a species, for example Paphiopedilum stonei (Hk. f.) Pfitz. var.

platytaenium Rchb. f.

The names of cultivated plants are slightly different since many of them have arti-

ficial origins or cannot technically be considered as true species or varieties. Miss

Joaquim’s orchid can be used here as an example on nomenclatural propriety for cul-

tivated plants. It has been suggested that this orchid is probably a natural hybrid (Yeoh,

1963) first occurring in Miss Agnes Joaquim’s garden in Singapore in the latter years of

the 19th century. In her garden two Vanda species, Vanda hookerana Rchb. f. (from

Kinta Valley in the northern part of W. Malaysia) and Vanda teres Lindl. (from Burma)

were planted and to this day it is not clear which one was the pod (female) parent. Bo-

tanically this orchid is known as an interspecific hybrid as there are two different parent

species involved. The term grex (plural: grexes) is also used for such hybrids. In order to

indicate the parentage of such an interspecific hybrid or grex it is customary to cite both

parents’ names, but place the pod plant’s name first and to connect these two names with

an “x”, e.g. Arachnis flos-aeris x A. hookerana. Generally there are two ways to name

such a hybrid plant. The first is to give a new specific epithet in Latin preceeded by an

“x”, e.g. Tilia x vulgaris, a hybrid of two linden tree species viz. Tilia cordata Mill. and

T. platyphylla Scop. The second, by far the more conventional way, is to give a common

name, not a Latin one, to hybrid plants, e.g. Arachnis Maggie Oei, Oncidium Goldiana,

etc;

Maxwell: Vanda Miss Joaquim 185

H. N. Ridley (1893), then Director of the Singapore Botanic Gardens, reported

and described Miss Joaquim’s orchid in the Gardeners’ Chronicle and named it as Vanda

Miss Joaquim. To conform with the examples given by Brickell (1980) and Withner

(1980) this hybrid should be written as Vanda (italicised or underlined when written

or typed) Miss Joaquim (normal lettering).

Interspecific hybrids (grexes), such as Vanda Miss Joaquim, should technically be

written as Vanda (Miss Joaquim g.), however the extra details of the parenthesis and

“s”’ (grex) are usually omitted by horticulturists for simplicity. The author’s name

of these hybrids, in this case Mr. Ridley’s, is always omitted.

The names of variant plants derived from a single individual, i.e. clones (cultivars)

are written inside a pair of simple quotation marks, e.g. Cattleya Bow Bells ‘Singapura’

or by placing cv. (cultivar) before the name of the cultivar, e.g. Phalaenopsis Samba

cv. Mamamia. The cultivar name, as with the name of the hybrid, is not underlined or

printed in italics.

The following examples, in addition to those noted above, illustrate the proper

manner in which various orchids should be written:

1. Vanda hookerana Rchb. f. (a species with author citation),

Phalaenopsis amboinensis J. J. S. ‘Simanis’ (cultivar of a species, the author

citation of the species is often omitted by horticulturalists), and

3. Aranda How Yee Peng ‘Ada’ (a cultivar of an intergeneric hybrid under a col-

lective generic name, technically written as X Aranda).

Legitimate Names

Species and varieties of orchids and other plants must be published with a Latin

description in order to be botanically acceptable (legitimate). Hybrids and cultivars

are also affected by the rules of botanical nomenclature since each hybrid or cultivar

can have only one correct (legitimate) name (Gilmour, Greatwood, and Hunt; 1976

and Brickell etal, 1980). Orchids, however, have received special attention from

botanists and horticulturalists, thus the rules concerning these plants differ somewhat

from those regarding others, e.g. other ornamentals, crop plants, fruit trees, etc. (Gil-

mour, Greatwood, and Hunt; 1976). Orchid hybrids and cultivars, for botanical and

legal purposes, must have either published descriptions or approved registrations so

that the names are legitimate.

The naming of hybrids and cultivars is sometimes also a legal process since these

names are essentially trademarks and are often commercially valuable. The owners

or registrants of these names must, therefore, properly register or publish descriptions

of these hybrids or cultivars to attain botanical as well as legal recognition of these

plants. The names of orchid hybrids and cultivars are, with a few exceptions, based

on publication or registration priority; that is, the first valid publication or registra-

tion of a legitimate name has priority over all other names of the same hybrid or cul-

tivar. Hybrid names proposed prior to 1 January 1967 are based on the earliest date

of publication or registration, while after this date they are based on registration only.

Cultivar names are based on publication priority only, that is, registration does not

provide for botanical or legal legitimacy. Unfortunately, due to the enormous num-

bers of orchid cultivars produced, registration of all cultivars is impossible. Several

schemes have been proposed to deal with this problem, however no satisfactory solution

has been formulated and agreed on by the World Orchid Commission. It must be noted that

186 Gard. Bull. Sing. 34:1 (1981)

exceptions to these rules of priority can only be considered and resolved by the Royal

Horticultural Society, London which is the International Registration Authority for

Orchid Hybrids. The details of publication and registration are given by Gilmor, Great-

wood, and Hunt (1976) and Brickell et al. (1980).

Acknowledgements

I would like to thank Prof. Hsuan Keng of the National University of Singapore

for providing me with an initial memorandum on this topic and to Mr. Hugh Tan for

his technical assistance.

REFERENCES

Brickell, C.D. etal. (eds.) (1980). International Code of Nomenclature for Cultivated Plants. Bohn,

Scheltema, & Holkema; Utrecht. 31 pp.

Gilmour, J.S.L., J. Greatwood, and P. F. Hunt (1976). Handbook On Orchid Nomenclature and

Registration, 2nd ed. International Orchid Commission, London, 149 pp.

Ridley, H. N. (1893). Vanda Miss Joaquim. Gardeners’ Chronicle 24 June, p. 740.

Withner, C. L. (1980). How to Write, Type, or Print an Orchid Name Properly. American Orchid

Society Bulletin 49:4, 384-387.

Yeoh, Bok Choon (1963). Miss Joaquim’s Orchid. Malayan Orchid Review 7, 36—39.

Vanda Miss Joaquim, national flower of Singapore Photo: David Lim

Maxwell: Book Review 187

BOOK REVIEW

J.F. MAXWELL

Botanic Gardens, Singapore

P. R. Wycherley and Ahmad Azli bin Mohd. Yusof, Grasses in Malayan Plantations.

Rubber Research Institute of Malaysia, Kuala Lumpur, West Malaysia, 1974; 176 pp.

Price: Malaysian $15.00.

This handy book, intended for rubber tree planters, satisfies the need for an identifi-

cation manual of grass weeds in rubber estates of West Malaysia. Since many modern

herbicides are selective preparations intended to eradicate certain species of grasses, it is

imperative that these grasses be correctly identified so that the appropriate chemicals be

used for the most effective grass control treatment.

Wycherley and Ahmad Azli include 64 species of grasses which, throughout the

years, have been found to be noxious and otherwise detrimental to the growth of rubber

trees. Far from just being a picture guide, the authors include brief and succinct accounts

on the habit and morphology, and ecology of grasses as well as notes on the detrimental

or damaging effects of specific species of grasses in rubber plantations. Rubber planters

will find these notes as important and interesting as the detailed line drawings of each

species. The status of grasses as cover plants and grasses as intercrops are also discussed

with complete literature citations for those who care to investigate these sections more

thoroughly.

The key to grasses is, by virtue of the inherent complexities of grass morphology

and taxonomy, somewhat technical; however all the botanical terms used are illustrated

in the beginning of the book and defined in a glossary at the end. Perhaps in an effort to

avoid various complications in the use of the key, most leads in the key have more than

the usual two couplets. Fortunately, the key refers to more vegetative and structural

details of the inflorescences than to the minute and difficult morphological aspects of the

spikelets; however a thorough knowledge of all these features must be realized before the

key can be used. Since this book is intended mainly for agriculturists, most of whom have

limited botanical training, the inclusion of the common names of these species should

have been included.

Aside from being the best manual of its kind for rubber planters, this book will also

be useful for other agriculturists in oil palm estates, fruit orchards, livestock ranches,

farms, etc. as well as students and naturalists. This book is, therefore, highly recommend-

ed for all libraries dealing with natural science, agriculture, and for anyone interested in

gaining a basic understanding of some of the more ubiquitous grass species found from

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Lawson, R. H. (1970). Flower Necrosis in Cattleya Orchids. Am. Orchid Soc. Bull.

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Singh, H. (1967). Sclerids in Fagraea. Gard. Bull. Sing. 22:2, 193-212.

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BK KE” DE Ee” KE ROR RBEe” Fe\ DR ®7B@ Pen” De” IEEE DE OE

‘

THE GARDENS’ BULLETIN

- SINGAPORE ”

VOL. XXXIV (Part II) lst December 1981

eed en TS

WONG Yew Kwan: \

Horticultural Notes on the Angsana (Pterocarpus indicus) ............... 189-201 4

F LIM-HO Chee Len: (

a, n

[ Tissue Culture of Curculigo latifolia (Hypoxidaceae) .................. 203—208 (

g B.C. STONE: (

% Contributions to the Flora of the Solomon Islands ................... 209—222 5

a Nomenclature of Joinvillea (Joinvilleaceae)... 2... 2 ee eee eee 223-225 NY

§ eee emai nt WetNECe Gale Manel 2 nek wos sls cs ee ee we ee eee 227 é

wi) Subash CHANDRA: ‘

> a Some Aspects of Interrelationships among Drynarioid Ferns .............. 229-238 )

te X

- Jennifer NG: ‘)

_ In-Vitro Pollen Germination of Cassia fistula. ..........-000000200ee 239-242 j

)

(

Published by the Botanic Gardens

Parks and Recreation Department

Ministry of National Development

Singapore 1025

Printed by Amsterdam Type Printers, Singapore.

ona Wo OK

EO OYE ORE ERROR?

GARDENS’ BULLETIN

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Chairman: Y.K. Wong, B.Sc. Hons. (Singapore), M.A. (Oxon), M.S.I. Biol.

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Gardens’ Bulletin

Botanic Gardens

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Singapore 1025

THE GARDENS’ BULLETIN

SINGAPORE

VOL. XXXIV (Part II) Ist December 1981

ae ON, a oe B.S

PAGES

WONG Yew Kwan:

Horticultural Notes on the Angsana (Pterocarpus indicus) ........-..++.+.+.-- 189-201

LIM-HO Chee Len:

Tissue Culture of Curculigo latifolia (Hypoxidaceae) ...........-..2-++--. 203—208

B.C. STONE:

Contributions to the Flora of the Solomon Islands ..............+.+.+-+-. 209—222

Nomenelaturc of. Jomvillea (loinvilleaceac)..... . .. . 2s 2 se es eee eee ee 223—225

a memondam: Ie Monte (cor Manel. 5. 55 lo ee i ee ae we me SE ee Ee 227

Subash CHANDRA:

Some Aspects of Interrelationships among Drynarioid Ferns .............-.- 229-238

Jennifer NG:

In-Vitro Pollen,Germination.of Cassia fistula ..........-2-+2+++++++- 239-242

Published by the Botanic Gardens

Parks and Recreation Department

Ministry of National Development

Singapore 1025

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HORTICULTURAL NOTES ON THE ANGSANA

(PTEROCARPUS INDICUS Willd.)

WONG YEW KWAN*

Parks & Recreation Department, Singapore

ABSTRACT

Pterocarpus indicus, a species of Leguminosae and highly adaptable in a

variety of habitats, is extensively planted as a shade and ornamental tree

in Singapore. Propagation is commonly by woody cuttings and trees of

considerable size can be transplanted successfully. Growth rates under mesic

habitat conditions are fast. Transplanted stumps of 3.7 m height and 60 cm

girth breast height can achieve a height increment of 1.66 m and a girth

increment of 14 cm per annum.

General requirements, pest and disease association, phenology, horti-

cultural use of the species, and transplanting technique for large rooted stumps

are described.

INTRODUCTION

Pterocarpus is a large genus of trees of the family Leguminosae with distribution

throughout the tropics. The Angsana is indigenous to Peninsular Malaysia*Its natural

habitat is said to be on the banks of coastal creeks and rocky shores, but it is not certain

whether it occurred in Singapore (Corner, 1952). Pterocarpus indicus is extensively

cultivated in the Indo—Pacific region as an ornamental tree and for its shade. It has

massive limbs, both ascending and spreading, which produce slender, drooping branches.

Crowns of trees of various sizes which have not been subjected to pruning or pollarding

for some time assume a rounded appearance. Full-sized trees in Singapore measure

25 to 30 m tall with a crown-spread of almost equal dimension and a girth at breast

height of 5 — 5.5 m (Fig. 1). There are, however, larger specimens growing in Penang

with girths exceeding 6 m. In Singapore there are some old trees along the highways

and adorning institutional grounds, but in the past 14 years a vast number of the species

has been planted in a great variety of settings, so much so that many young trees are

now dominating many local scenes (Figs. 2—5). Its popularity lies in its ease of propaga-

tion, fast growth rate, ease of management and, above all, it lends itself to transplanting

as a fairly large tree with a high degree of success. Such transplants have been referred

to as “instant” trees because branches develop soon after transplanting and within a

matter of a few months fair-sized trees obtained.

PROPAGATION

Angsanas can be propagated from seed, however, in view of the ease of propagation by

woody cuttings in Singapore, propagation by seed is seldom, if at all, practised. Cuttings

of various sizes can be used. The Parks & Recreation Department has planted cuttings

of 3 m length, with a lower-end diameter measuring some 10 cm, directly into planting

Sites with some degree of success. More commonly smaller cuttings are used for rooting

* At the time of printing, the author had resigned.

* Angsana or Sena, as this species is also known in Malay, is also naturally found in India, SE. Asia,

Philippines, Sumatra, and Java.

189

190 Gard. Bull. Sing. 34:2 (1982)

in the nursery before field planting. A suitable cutting is one with a 1.5 m length and 4 to

5 cm diameter at the lower end. The cutting is inserted about 20 cm deep in a soil

mixture contained in a polybag 30 x 30 cm. The cutting can be in the form of a straight

stake or be treated so that the top end has a few branches. Treatment with a fungicide

will ensure greater succes — which may be as high as 90%. Such a cutting will flush within

2—4 weeks and be well rooted within 3 months. Cuttings of the size described can also be

planted unrooted directly into the field, but the maintenance of such plants (e.g. watering

and pest control) spread over a large area is of course more problematic and the

percentage of success will be lower.

For the production of “instant” trees, many of the rooted cuttings are planted out -

in road verges and roadside open spaces. In such areas planting is done at a higher

density than required by mature trees so that after a period of growth (about 2 to 3

years) the grove is progressively thinned for transplanting to places where the landscape

requires an instant effect for visual impact or for shade.

FIELD PLANTING AND TRANSPLANTING

Small Rooted Cuttings

If propagation is through woody cuttings planted in polybags in the nursery, the rooted

cuttings can be transplanted into the field when they are about 3 to 6 months old. A

cutting of this age will have produced a bunch of short slender branches at the top

end measuring about a metre long. Care should be taken not to break these branches

while the plants are being handled for transplanting. For cuttings of the size described it

is normal to plant them out in machine-dug holes of 1 m*. The planting technique is

similar to planting out a sapling grown in polybags. In Singapore it is essential to provide

a good loamy top soil for the planting holes because not only is the in situ soil clayey, but

in many a planting situation cut and fill operations could have either reduced the planting

site to the parent materials or given rise to filled areas consisting of compacted sub-soil

or stiff clay.

“Instant” Trees

Angsanas lend themselves to transplanting as large trees fairly easily and the survival

rate is usually as high as 90% provided the trees to be transplanted are not too large.

In the earlier days of the Garden City Campaign, trees with girths as large as 100 cm

(measured 1 m above ground level) were transplanted. It was, however, observed that

such large transplants would not grow well. It is now, therefore, the practice to transplant

trees with girths of not more than 60 cm.

The transplanting of such trees needs a small crane mounted ona truck, simply because

of their weight and size. A tree of 60cm girth is,in fact, already 7m tall. The branches

are first lopped off so that the resultant stump with the various snags is about 4.5 m high.

The stump should have 3 to 4 main snags for proper crown development. All slender

twigs and branches below the pollarded ends or the main branches are removed. The cut

surfaces of the pollarded branches are painted with a wound dressing so as to prevent

dessication and extensive rotting. This is of particular importance for the Angsana

because it has the habit of producing new branches almost flush with the cut surface of

the snag or just below it. If rotting occurs, proper callousing may not be achieved and

a developing branch will snap easily at the point of union during a storm.

After the tree has been prepared as described above, the next step is to excavate around

Z Y Zs

Fig 1. Full grown Angsanas near Anderson Bridge. The age of these trees is likely to be about 100

years. They are the remnants of a row destroyed by a disease in 1926. Their present height

is approximately 27m. with a girth at breast height of 4.57m.

Fig. 2. Year old transplants of Angsana at Ang Mo Kio Town Centre. Note the feathery appearance

of young, mostly ascending, branches at this stage of growth.

Fig 3. Angsanas lining a part of Orchard Mall. These transplants were 4 years old when the photo

was taken. Note the fully developed and rounded crowns with drooping branches.

Fig4. The Margin of the swimming Lagoon at East Coast Park lined with transplants 5 years old.

On Angsanas (Pterocarpus indicus) 193

Fig5. Six-year old Angsana transplants providing shade to the cycle and jogging tracks at East

Coast Park.

the tree base to create a proper ball of earth for eventual lifting. A trench of about 60 cm

depth is first dug around the tree base at a distance of about 50 cm from the base of the

stump. More earth is removed by undermining towards the trunk so that the ball of

earth is shaped like aninverteddome with the original soil surface on top. The ball of

earth is wrapped with gunny sacks or some other suitable material so as to prevent the

earth from breakirig up when the tree is lifted. A chain and choke arrangement attached

to a winch with power take-off from the lorry is used to sever the ball of earth from the

ground before the tree is lifted by the crane onto the lorry for transportation to the

planting site (Fig. 6).

Trenching can be done mechanically, although the shaping of the ball of earth and

wrapping it have to be done manually.

What is described has been the practice in Singapore up to now and this careful treat-

ment has ensured a high degree of success in transplanting. In the face of labour shortage

194 Gard. Bull. Sing. 34:2 (1982)

we have tried out some of the transplanting machines available today. The preliminary

trials showed that such machines do not give an intact ball of earth because the grabbing

and uplifting actions tend to loosen the earth too much, especially when the soil is loamy,

thus leading to root disturbance and greater transplanting shock, particularly in our hot

tropical climate in which evapo-transpiration is high. The Parks & Recreation Department is

using more clayey soil for growing future transplants and for species such as the Angsana

which has a high regrowth potential, there is a chance that transplanting can be done

mechanically.

Theoretically in the transplanting of “‘instant’’ trees one should ensure as large a ball of

earth as possible so as to preserve a large number of undisturbed roots and rootlets.

This will lessen transplanting shock. On the other hand, this will also increase handling

costs especially when trenching is done manually. Angsana has been found to be a very

rugged species which can withstand rough handling. In recent months transplants of

50 cm girths which were lifted with a ball of earth as small as 30 cm radius and 50 cm

depth were found to survive well. Further, it has been found that when the ball of earth

is very clayey no wrapping is needed and so long as the ball is not disintegrated the stump

can still survive.

The hole earmarked to receive the transplant can be 1.5 m square by a metre deep.

Enough top soil should be filled in so that it forms a flat-topped mound about 20 cm or

so above the original soil surface. It has been found that in transplanting “‘instant’’ trees

it is best to avoid putting sludge in the planting hole for this has been found to cause

higher mortality. One reason may be that the organic matter encourages greater moisture

retention leading to fungal infestation of the injured root system of the transplant.

The ball of earth of the transplant is placed so that the original surface is flush with the

flat-topped mound. Allowing for settling, the surface of the original ball of earth will

be flush with the surrounding land surface in due course. In places where there is a

tendency to water-logging a slightly higher mound can be used. One should avoid deep

planting which will result in poor plant growth even if the transplant were to survive.

For such large transplants there is no necessity to stake, but in areas such as car parks

there is a need to provide tree guards to prevent the transplants from being knocked

down by cars.

Sometimes during preparation for transplanting, the bark of the stem or snags might

be damaged. Should this happen one should use a wood preservative to paint over the

injury to prevent rotting. Unless damage is extensive callous formation will soon grow

over the wound.

Spacing ©

If final spacing is considered then the Angsana should be given 25 m between trees.

However, a planting scheme along roadsides with such spacing will give rise to a rather

Sparse appearance and will not have a green impact for a long time. It is normal,

therefore, for us to plant trees about 12 m apart so that if need be such trees can later be

progressively thinned for use elsewhere.

On Angsanas (Pterocarpus indicus ) 195

GROWTH

Crown Development

For transplants as “‘instant”’ trees, sprouting from near the pollarded tips of the snags

takes place within 2 weeks, although this can be as iate as 1 month from the date of

transplanting. It has also been observed that the early sprouts may suddenly die back,

especially during drought, only to be followed by the emergence of fresh shoots which

will eventually develop into the main branches of the tree.

The young branches tend to cluster around the pollarded tip. A few, however, may deve-

lop a little further down. Unlike the Yellow Flame (Peltophorum ptercocarpum (DC.)

Back.) epicormic shoots are uncommon under normal conditions of growth after trans-

planting. Within 3 months there can be as many as 10 young branches per snag (pollarded

Fig. 6. Transplanting a large rooted stump. The pollarded stump with its ball of earth is positioned

by the crane to go into the planting hole. Since the ball of earth has clayey soil, it has not

been wrapped.

196 Gard. Bull. Sing. 34:2 (1982)

branch) which measure about 1.5 m length and assume a feathery appearance. After a year,

the lower of these branches spread out while the upper ones become the ascending limbs

which increase rapidly in girth and length. Each of these ‘primary’ branches gives rise to

many subsidiary branches, many of which are slender and drooping. After two years a

well-formed crown develops. After 4 years, if no subsequent pruning is done, a rounded

crown is achieved. Crown density, however, varies from tree to tree, depending on the

original number of snags of the transplanted stump and the subsequent shoots which

develop therefrom.

Growth of large woody cuttings or rooted cuttings planted out in the field assumes

a similar pattern, but because of the smaller initial stature of such plants, growth in

size and crown proliferation will understandably be slower.

Bole Development

It has been observed that the original stump does not grow in height after transplanting.

Height is contributed by the numerous new branches which develop from the snags of

the original stump. The bole, however, increases in girth. At the time of transplanting

the bole of a stump of the usual girth size (60 cm) is rounded in cross section but as

the bole increases in size, it becomes fluted. The fluting is increasingly pronounced with

greater girth size, with the ridges developing into short buttresses at ground level.

The girth measurements presented below have been taken over the fluted boles.

Height & Girth Increments

Quantitative data are not available on height and girth increments of the Angsana be-

cause there was no systematic inventory to study growth. However, some figures of growth

based on estimates and measurements of a few trees planted in the past 10 years are

given here as an indication.

Three trees along Maxwell Road immediately next to the Maxwell Market planted in

1970 have an average girth of 2.15 m, measured 1.5 m above ground level, and a total

height of about 17 m. At the time of planting the average girth was about 60 cm and

the stump height was about 3.7 m. This means the trees have put on over 11 years an

average height increase of 13.3 m and a girth increment of 1.55 m or an equivalent of

1.2 m height, and 14 cm girth growth per annum. These trees now have a crown spread

of 12 to 18m.

Another group of 6 trees growing along Maxwell Road in front of the Tamil School and

planted in 1973 have an average girth of 1.63 m, measured 1.5 m above ground and

an average height also of 17 m. The other parameters are the same as for the trees

mentioned in the preceding paragraph. Calculating in the same way a girth increment

of 12.9 cm and a height increment of 1.66 m per annum.

It will be noted that the trees in front of the Tamil School have faster height growth,

but slightly smaller girth increments. This could be due to the fact that the spacing of

the 2 groups of trees are different since those by the side of the market are spaced 18 m

apart while those in front of the school are spaced 12 m apart. The closer spacing in the

latter group has caused crown competition to push the trees skywards, while the girth

increment is retarded.

On An*sanas (Pterocarpus indicus) 197

Given the right root environment the Angsana grows well and fast, but is sensitive

to root-bound conditions. This is shown by the trees growing along the centre divider

of Paya Lebar Road. Angsanas stumps of similar sizes were planted there in 1974. Some-

how at the time of planting several layers of the old road surfaces were not broken

through. In effect, the trees were standing on a concrete surface 1 m below the present

road surface and as a result all the trees stagnated. Although they are now 7 years old

their average girth is only 0.9 m (0.6 m at the time of planting) or the girth increment

was only 4.3 cm per annum compared with the more rapid growth of 14 cm and 12.9 cm

girth per annum of the trees mentioned earlier.

Another example of stunted growth is seen in the concourse of the National Develop-

ment Building where bed rock is near the soil surface and where lateral root spread is further

restricted by the concrete surface. Two Angsanas planted there in 1970 have hardly

grown, but nevertheless are still alive.

GROWTH REQUIREMENTS

Habitat

The habitats of the Angsana along the East Coast of Peninsular Malaysia are said to be

riparian along coastal tidal creeks (Corner, 1952). In Singapore it is one of the most

adaptable tree species and is able to grow well in a wide range of habitats and soil condi-

tions. It grows on flat sandy areas near the coast as well as in hilly areas with stiff clay.

It is also able to withstand a considerable degree of water-logging as evidenced by its

fair performance at the East Coast Park which has reclaimed land with extremely

compacted soil. The compaction and the flat topography of the Park cause poor percola-

tion and surface run-off. As a result the planting holes into which loamy top soil has

been filled became sumps for rain water. Many other species have failed to establish

under such conditions, but the Angsana is able to grow. It is, however, expected that

growth will not be as good as that achievable by the individuals under more mesic

conditions.

The Angsana is a light demander and is, therefore, not suited for underplanting in an

existing grove of trees. Left to compete amongst themselves, Angsana trees can be planted

close together or in groves to form good screens or other ornamental features.

Nutrients

Little is known about the exact nutrient requirements of ornamental trees.Our manurial

programme is, therefore, a tentative one until more reliable data can be obtained at

a later date. One approach is to study the annual litter fall of trees of different ages

to gauge the amount of nutrient loss through this factor. In an urban environment it

can be assumed that all litter is swept away and the manurial programme should aim

at replacing this amount of loss. Further refinement can be achieved through a more

quantitative estimate of the stem and branch growth.

The Department has just begun to establish leaf traps in different places to measure

annual litter fall for Angsana. Perhaps after a year or so some information can be obtained.

Meanwhile, the practice is to apply 4% kg, 1 kg, 1.5 kg of a compound fertilizer to trans-

plants respectively in the 1st, 2nd and 3rd year of growth. Thereafter, application is based

on the visual conditions of the trees and the manurial rates can vary between 3-10 kg

per annum depending on the size of the trees.

198 Gard. Bull. Sing. 34:2 (1982)

In parks and gardens the fertilizers are broadcast evenly over the soil surface under the

tree crown but in roadside situations with concrete sidewalks, the manure is applied

in the grooves between concrete slabs or in the holes in the slabs. In such concrete

areas it is usually difficult to apply the full rate for a tree of a particular size because

of the limited unsealed area of the tree base. The annual rate is, therefore, given in two

or more applications.

PESTS & DISEASES

In urban sidewalks, small urban parks, and open spaces the Angsana is particularly free

from pests and diseases. When planted in large parks or in areas bordering natural vege-

tation, foliar damage by pests, mostly grasshoppers (Valanga) and night flying beetles

(Apogonia, Adoretus, and Anomala), is not uncommon. Occasionally the young leaves

can also be damaged by leaf miners.

As the trees grow fast and new flushes are produced continually it is seldom the practice

to control pest damage. If, however, young trees produced by short cuttings are badly

damaged, an organophosphate such as “‘Dipterex’’ can be used for a weekly spray. More

persistent pests can be tackled with “‘Basudin’’.

The Angsanas planted along Mandai Road and those growing in the Zoological Gardens,

both localities being surrounded by forests, suffer from a kind of leaf blight— the fungus

of which has been identified as a species of Gleosporium. Phenologically the Angsana

does shed its leaves occasionally but seldom with the crown completely bare. The trees

planted along Mandai Road, however, undergo a cyclical rhythm with the crown

completely defoliated by the leaf blight followed by luxuriant new flushes. As the leaves

get older, they are afflicted by the same malady and the cycle is repeated. The association

of the disease with proximity to the forest is due perhaps to the higher relative humidity

associated with such a habitat which favours fungal growth.

Angsana trees in Singapore and Peninsular Malaysia suffered extensive mortality from

1875 to 1925. Trees 10 to 40 years old died progressively and sometimes whole rows

were wiped out. It appears the present trees standing near Anderson Bridge along

Connaught Drive are remnants of a whole row growing along that road. The disease

was never identified. Furtado (1935) noted that the malady could have been spread

by a Jassid (leaf hopper) while Corner (1952) stated that the disease was akin to the

Dutch Elm Disease of the Temperate Zone.

PHENOLOGY

In the northern part of Peninsular Malaysia where the climate tends to be drier and the

dry spells more pronounced, gregarious flowering is said to take place frequently. In

Singapore mature trees do not flower often and flowering is normally solitary. One

exception to this rule was the general flowering which took place in June/July 1976 when

old trees littered the roads with carpets of their fragrant yellow flowers, so much so

that special efforts were made to sweep them off daily. Apparently this general flowering

was in response to the prolonged and pronounced dry spell which stretched from January

to March 1976. For the first time many other flowering trees planted in Singapore also

flowered gregariously in response to that stimulus—notably amongst them being Gliricidia

sepium (Jacq.) Kunth ex Walp., another Leguminous tree.

When a tree flowers, normally a large part of the crown is covered with the flowers

On Angsanas (Pterocarpus indicus) 199

and the whole affair is over within 4 days to a week (Fig. 7). According to Corner (1952)

the disc-like fruits take 4 months to mature.

In Singapore, young trees or new transplants assume an evergreen habit when grown

under mesic conditions and given adequate manuring. The crown puts on new flushes

continually throughout the year. Old trees or younger transplants which are grown under

exacting conditions such as in rocky areas with shallow soil, tend to be deciduous at the

top of the crown and such trees assume a stag-headed appearance. Strangely enough

such trees also tend to flower and fruit slightly more often.

SPECIAL HORTICULTURAL CHARACTERISTICS

Adaptability

As indicated before, Angsana is a tree which lends itself to propagation by large woody

cuttings and large transplants. Apart from this, Angsanas are highly adaptable. In Singa-

pore they are now planted in a great variety of habitats ranging from well-drained hill

slopes to flat, water-logged reclaimed land.

Aggressive Roots

Perhaps one reason why the Angsana is able to grow well under a wide range of habitats

is its powerful root system. It has been observed in trenching operations by Services

Departments that the roots are found to be able to spread under road foundations, well

beyond the crown limit of the tree, although the soil is very clayey. Figure 8 shows

part of the root system of an Angsana tree which proliferated under a service road.

A peculiar feature is the presence of anastomosis in the system.

Fig. 7. Inflorescences of the Angsana associated with leafy twigs. The petals are golden-yellow

in colour.

200 Gard. Bull. Sing. 34:2 (1982)

In many instances the roots have been found to grow underneath large concrete

monsoon drains and then emerge in the surface of lawns located some distance from the

base of the tree. Such rugged root systems mean that it is able to exploit nutrient and

moisture resources over a wide area, hence the vigorous growth.

Response to Pruning, Pollarding, or Crown Damage

The Angsana is able to recover from pruning or pollarding very quickly. This has advan-

tages in horticulture, but at the same time poses a problem to the management of roadside

trees. In areas where the trees need pruning it is desirable aesthetically that the crown

recovers its shape rapidly; on the other hand, this means higher frequency of pruning or

pollarding to ensure the branches will not get into the way of traffic (especially buses and

containers).

In May 1976, during a severe thunderstorm, the crown of a large tree was severely

damaged. After the usual tree surgery to harmonise the branching and to prevent rotting,

the tree put on new flushes within a couple of weeks and 7 months later had reformed a

well-shaped crown.

The Production of Adventitious Roots in Response to Injury

It has been observed that the Angsana has another very interesting characteristic in

that, for mature or old trees extensive injury to any part of the trunk or stem causes the

callousing bark to produce adventitious fibrous roots which can be extensive and matted.

Likewise, fibrous roots are produced in the arch of a cracked bifurcation. The roots

grow in size and number rapidly within the crack, presumably because of the collection

of water and nutrients therein, thus causing the two arms of the bifurcation to be forced

apart. If this happens to be between the main ascending limbs it is most dangerous for it is

only a matter of time before one of them collapses. It is, therefore , better to have such a

tree felled or proper bracing must be carried out. If the phenomenon occurs in a less

critical part of the tree, then that portion should be pruned off.

GENERAL HORTICULTURAL USE

The Angsana is certainly a graceful tree with considerable ornamental value and because

of its large size it also provides good shade. It is a tree for open spaces and wide sidewalks

or centre dividers where shade is important and needed quickly. It is also a tree to be

chosen where the edaphic conditions are too exacting for establishing other trees, i.e.

if space permits the planting of such a tree. It is not recommended for narrow sidewalks

or in small home gardens, not only because of its big size, but also because of the aggres-

sive roots. For any particular landscape which needs an “‘instant’’ effect, the Angsana

can be used judiciously, but because of the ease of transplant and fast growth, the Angsana

has been chosen for many localities where it should not have been planted.

Angsana root systems are wind-firm, but the young branches of transplants tend to

snap in stormy weather at the point of union with the pollarded branches. Smaller

rooted stumps with smaller pollarded snags tend to give better union and hence more

wind resistance. |

BRAVE

LOW

Fig. 8. Vigorous roots of a 25-year old Angsana under a service road. Note the unusual phenomenon

of anastomosis of the roots. The tree was grown from a seedling.

202 Gard. Bull. Sing. 34:2 (1982)

ADDENDUM TO PTEROCARPUS INDICUS

With reference to the Introduction on page 189, subsequent information received

shows that there is an exceptionally large tree with a girth size measuring 7.5 m in the

grounds of the Fairy Point chalets at Changi.

REFERENCES

Corner, E.J.H. (1952). Wayside Trees of Malaya I. Government Printing Office, Singapore. 375—376

Furtado, C.X. (1935). A Disease of the Angsana Tree. J. Mal. Br. Roy. As. Soc. 13:2, 163-192.

(

”

;

TISSUE CULTURE OF CURCULIGO LATIFOLIA DRY. ex W.T. AIT.

(HYPOXIDACEAE)

LIM-HO CHEE LEN

Botanic Gardens, Singapore

ABSTRACT

This paper reports the success in clonal propagation of Curculigo latifolia

at the Tissue Culture Laboratory, Singapore Botanic Gardens. Experiments

using leaf blades, petioles, apical shoots, and rhizomes were carried out and

the best results were obtained from rhizome cultures. The medium used

contained the Murashige and Skoog inorganic salts (half strength), sucrose

(30 g per litre), thiamine (0.4 mg per litre), coconut milk (150 ml per litre),

kinetin (5 mg per litre), and IAA (2.5 mg per litre). Mass propagation of

this species is in progress.

INTRODUCTION

Singapore, often called the Garden City, is famous for its parks, gardens and, most of

all, the refreshing sight of the green canopy of big trees and luxuriant ornamental plants

that grace the city especially along the roadsides. Rapid urban development has created

many new environments, such as open spaces, under flyovers, or reclaimed land, where

many plants find it difficult to grow and thrive. There has been considerable effort in

trying to cultivate more species of native plants that can grow in these various new

growth conditions. Curculigo latifolia is one species which has been found suitable for

landscaping needs under flyovers which are shaded and sometimes rather dry.

Curculigo latifolia is an elegant native herb which grows under fully shaded forest condi-

tions. It is a stemless herb with long leaves and short, thick rhizomes. The inflorescences,

which are produced throughout the year, are compact, head-like, close to the ground,

and have numerous flowers with bright yellow tepals (Fig. 1). Although this species

offers good prospects for ornamental purposes in Singapore, the problem of obtaining

large numbers of this species has yet to be solved. Curculigo latifolia is not abundant

in Singapore’s forests, thus collection of many living plants would have an undesirable

effect on the floristics and ecology of their native habitats. In an effort to find alternative

means to supply abundant material of this species for roadside planting, tissue culture

techniques were investigated. This paper reports the success in mass propagation of

Curculigo latifolia using tissue culture techniques at the Tissue Culture Laboratory,

Singapore Botanic Gardens.

METHODS AND MATERIALS

Young shoots of Curculigo latifolia about 8—10 cm in height were used for tissue

culture. The plants were thoroughly washed to remove dirt on the leaves and soil on the

rhizomes. The outer leaves and roots were then removed, leaving the apical tissue and the

rhizomes.

The apical shoots, rhizomes, and leaves were then sterilized by soaking sequentially in:

(1) 10% detergent for 10 minutes,

(2) 75% ethyl alcohol for 30 minutes,

(3) 15% chlorox for 30 minutes, and

(4) three changes of distilled water for five minutes each time.

203

204 Gard. Bull. Sing. 34:2 (1982)

The following three different plant parts were excised for tissue culture:

(1) Shoot apices — The sterilized apical tissue was further trimmed to a small

cube of about 1 cubic mm in size containing the apical meristematic tissue

of the shoot.

(2) Leaves — The whole leaf, blade and petiole, was cut into 5mm segments

and placed proximal side down on agar medium.

(3) Rhizomes — The outer layer of the rhizomes was removed to reduce risk

of contamination. The remaining part was then cut into sections and trans-

ferred onto an agar medium.

The agar medium used contained the inorganic salts of Murashige and Skoog (1962)

at half strength with the addition of sucrose (30 g per litre), thiamine (0.4 mg per litre),

coconut milk (150 ml per litre), and various hormones.

All cultures were incubated at a temperature of 20—28°C and exposed to Grolux light

for 8 hours per day.

RESULTS

A total of 11 plants were used in the experiment. The number of cultures prepared using

the various plant parts and their state of growth at the end of a 3 month period are listed

in Table 1.

Table 1: Results of the tissue culture experiment of Curculigo latifolia

Explant no. of only callus callus, callus no. conta—

cultures callus and shoots, and growth minated

prepared formed shoots &roots roots

formed formed formed

Leaf blade 18 3 0’ 0 0 7 8

Petiole 20 3 6 0 0 8 3

Rhizome 20 3 l fi 2 3 4

Apical i l 6 0 0 3 ]

shoot

The most unsuccessful results were from those using leaves. Except for a few cases

where callus formed (Fig. 2), the leaf cultures either showed no sign of growth or were

contaminated. The difficulty is probably in the sterilization technique. To kill all the fungus

spores stuck on the leaves, it was found necessary to use a sterilization duration which

was probably too long in that the leaf tissue was also damaged.

The petiole and apical shoot cultures formed callus and shoots quite readily (Fig. 3),

but were slow to form roots (root formation was observed 2—3 months after the end of

the experiment).

Tissue culture, Curculigo latifolia 205

The best results obtained were from rhizome cultures. Satisfactory growth of callus,

shoots, and roots were observed in 7 out of the 20 cultures prepared (Fig. 4).

In order to encourage root formation, it was decided that hormones such as kinetin and

IAA should be added to the agar medium used for incubation. A simple factorial experi-

ment was conducted with three levels of kinetin (1, 2.5, 5 mg per litre) and five levels

of IAA (0, 1, 2.5, 5, 10 mg per litre). Five replicates were prepared. In each test a single

rootless shoot was introduced and incubated for three months. The results are listed in

Table 2.

Table 2 shows that the best root formation and shoot multiplication were obtained at

5 mg per litre kinetin concentration. At 5 mg per litre kinetin concentration, although

root formation improved with increasing concentration of IAA, shoot multiplication

reached a clear peak at 2.5 mg per litre [AA concentration. Increasing the IAA concentra-

tion further appeared to strongly favour the formation of callus at the expense of shoot

multiplication. Media discolouration also appear to become worse with increasing concen-

trations of kinetin or IAA.

Based on the above results, it was decided to use a medium containing the Murashige

and Skoog (1962) basic salt (half strength), sucrose (30 g per litre), thiamine (0.4 mg per

litre), coconut milk (150 ml per litre), kinetin (S mg per litre), and IAA (2.5 mg per

litre) for mass propagation work.

MASS PROPAGATION

Starting from the tissue taken from 11 plants, a stock of about 70 flasks of cultures

were obtained after about 10 months. A production programme was then planned and

put into operation. A culture flask normally contains 2—8 shoots at different stages of

development. At 3 month intervals the shoots were taken out and separated into indivi-

Table 2 : Results of the factorial experiment on hormone-induced root formation of

Curculigo latifolia shoots.

IAA kinetin medium callus root shoot multi-

(mg/L) (mg/L) discolou- formation* formation* plication

ration* (average no.)

0 1 none + none 1 (1)

1 1 none + none 1-2 (1.1)

25 1 none ~ none 1-3 (1.8)

5 1 2 + none 1-2 (1.1)

10 1 ee oa + 2-3 (1.8)

0 fe, - + + 1-2 (1.1)

1 2 + a tee 1-2 (1.1)

2.5 25 ++ ++ ++ 0 (0)

2 2.5 ++ ++ ++ 2 (2)

10 24 ++ +4 ++ 2-5 (2.6)

0 5 ++ + ++ 1-2 (1.1)

1 5 ++ + +t 3-5 (3.6)

pe 5 +4 ++ +++ 2-8 (5)

5 5 tat +++ T+t 1-5 (3)

10 5 +++ +++ +++ 1-5 (3)

*+ = slight ++ = moderate +++ = high

206 Gard. Bull. Sing. 34:2 (1982)

dual plants. Those taller than 8 cm with strong roots were rinsed in distilled water and

potted out. The smaller plants were transferred individually onto agar medium for multi-

plication.

The plants potted out were protected under a plastic cover for one week to help them

adjust to nursery conditions. About 90% of the plants potted out have survived. After

another 5-6 months these plantlets are normally ready to be planted in field. Fig. 5 shows

some plants in the nursery.

About 90% of the plants produced so far have remained identical to the parent plants.

The rest seem to produce more leaves and show some variation in leaf colour.

Based on the observed growth rate, it has been estimated that about 5 000 plants from

tissue culture can be produced within the first year of mass propagation, sufficient to

meet the current landscaping needs of the Parks and Recreation Department.

ACKNOWLEDGEMENT

The Curculigo latifolia tissue culture project was initiated on the direction of Mr. Wong

Yew Kwan, recently Commissioner, Parks and Recreation Department. The author wishes

to thank Miss Lee Swee Meng, Miss Lein Lee Jiuan, and Mrs. Teo-Lee Guek Choon for their

competent assistance in the experiment.

REFERENCE

Murashige, T. and F. Skoog 1962. A revised medium for rapid growth and bioassays with tobacco

culture. Plant Physiol. 15, 473-497.

Fig 1. Curculigo latifolia, mature plant.

Fig. 3. Callus and shoots formed from apical shoot.

and roots

shoots

i tisfactory growth of callus,

wing sa

Fig 4. Rhizome cultures sho

+ ee

SGA

the

SKNCRK x

Qg QBW ww

MN 0R#—-D?3?)

tissue culture technique

latifolia produced by

ligo

Fig 5. Curcu

CONTRIBUTIONS TO THE FLORA OF THE SOLOMON ISLANDS

BENJAMIN C. STONE

Department of Botany, University of Malaya, Kuala Lumpur

Malaysia

From 14 September to 10 December 1957, 1 collected plants in the Solomon Islands

on an expedition made possible by a National Science Foundation grant (G1834) to

Professor H. St. John of the University of Hawaii. Although the main purpose of the

expedition was to investigate the genus Pandanus, of which 86 numbers were collected

in the Solomon Islands, 25 in the New Hebrides, and 39 in the Bismarck Archipelago,

there was also opportunity to collect other plants of which 232 numbers were collected

in the Solomon Islands, 40 in the New Hebrides, and 14 in the Bismarck Archipelago.

The very imperfect state of our botanical knowledge of the flora of these three island

groups stimulated a desire to study all the available material, and has led so far to eleven

papers (Stone & Lane, 1960; Stone 1961, 1962, 1963, 1965, 1966, 1969, 1970, 1972,

1973; Stone & Whitmore, 1971) dealing respectively with a few ferns, Sararanga

(Pandanaceae) the genus Boerlagiodendron (Araliaceae), new Pandanus and Freycinetia

(Pandanaceae), a new species of Polyscias (Araliaceae), an account of Pandanus in the

Solomon Islands, and a study, with descriptions of two new species, of Tapeinosperma

(Myrsinaceae). The paper of Sararanga was monographic in scope, including the Philippine

species. Material on Pandanus is accumulating and some is awaiting publication. A com-

plete account of the Solomons Freycinetia is now in preparation. The Myrsinaceae

are being worked up in collaboration with Dr. T.C. Whitmore, formerly of the Forest

Dept., B.S.1-P.

The present account is an attempt to deal with all but the most intractable collections

from the 1957 expedition, and is set out, in the traditional Englerian arrangement of

families, purely for convenience. The order of genera and species within each family is,

however, alphabetical. Each collection is documented with appropriate notes, and where

needed, descriptions and illustrations.

The first set of the specimens is deposited in the herbarium of the Bishop Museum,

Honolulu (BISH); a second set, not so complete, is at the U.S. National Herbarium,

Washington DC. (US); a further set is for the time being kept in the author’s possession;

and a very few fourth replicates are in the Arnold Arboretum, Cambridge , Massachusetts

(A).

I should like to acknowledge the thanks I owe to several colleagues; the late Mr. James

Sinclair (Singapore), Dr. T.C. Whitmore (Kepong, W. Malaysia), Dr. T. Koyama (New

York), and Prof. A.C. Smith (Honolulu).

Literature Cited

Stone, B.C. 1961. The Genus Sararanga (Pandanaceae). Brittonia 13: 2; 212-224.

1962. Boerlagiodendron (Araliaceae) in Eastern Melanesia. Proc. Biol. Soc. Wash. DC.

75, 25-32.

1963. The genus Freycinetia in the Solomon Islands. Proc. Biol. Soc. Wash. DC. 76, 1-8.

1965. Melanesian plant studies I, II. Suppl. Micron. 14. Kuala Lumpur.

210 Gard. Bull. Sing. 34:2 (1982)

Stone (cont.)

1966. Polyscias verticillata (Araliaceae), a new species from the Solomon Islands.

Jour. Arnold Arb. 47: 3, 279-272.

1969. Pandanus decus-montium, a new species from the Solomon Islands. Proc. Biol.

Soc. Wash. DC. 82, 439-442.

1970. Materials for a Monograph of Freycinetia, 1X. The genus Freycinetia in the Solo-

mon Islands. Kew Bull. 24: 2, 355-375.

1972. The genus Pandanus in the Solomon Islands, with notes on adjacent regions.

Malaysian J. Sci. IA: 93-132, figs. 1-18, Pls. 1, 2.

1973. The Genus Pandanus in the Solomon Islands, with notes on adjacent regions

(Concluded). Malaysian J. Sci. 2A: 59-80.

and I.E. Lane, 1960. On a small collection of ferns from the New Hebrides and the

Solomon Islands. Bot. Notis. 112, 371-376.

and T.C. Whitmore, 1971. Notes on the systematy of Solomon Islands plants and some

of their‘New Guinea relatives. XI. Reinwardtia 8, 3—11.

Walker, F.S. 1948 and 1962. The Forests of the British Solomon Islands Protectorate. Honiara,

B.S.LP. (Reprint of the London, 1948, edition).

Whitmore, T.C. 1966. Guide to the Forests of the British Solomon Islands. B.S.I.P. Oxford Univ.

Press, London. (Forest Record No. 2, Forestry Dept. B.S.I.P.).

ULMACEAE

Celtis L.

Celtis kajewskii Merril & Perry, J. Arn. Arb. 22 (1941), 254.

SANTA YSABEL: Tatamba Lagoon, Horara Islet; alt. 15 m. tree with triplinerved ovate leaf

blades, 24 Oct. 1957, Stone 2509 (BISH).

MORACEAE

Ficus L.

Ficus verticillaris Corner, Gard. Bull. Sing. 18: (1960) 27.

MALAITA: Kwara-ae District, near Tantalau Village, c. 325 m. alt., tree about 13 m. tall,

leaves clustered near ends of slender branches; figs globose, 4 mm diam., n.v. “fa ‘adi’ila”, 23 Sept.

1957, Stone 2347 (BISH).

Ficus indigofera Rechinger, Fedde Rep. Sp. Nov. 11 (1912) 179.

MALAITA: Kwara-ae District, Tantalau, 325 m, tree 15 m tall, leaf blades with tiny basal

auricles, n.v. “aitia”; 23 Sept. 1957, Stone 2340 (BISH).

Ficus dissipata Corner, Phil. Trans. Roy. Soc. Lond. 253: 783 (1967) 149.

MALAITA: Fiu River 4 miles N. of Aulei near the coast, 7 m, tree, cauliflorous, leaf blades

reddish-hispid; 19 Sept. 1957, Stone 2298 (BISH).

Ficus erinobotrya Corner var. solomonensis Corner, Phil. Trans. Roy. Soc. Lond, 253: 783

(1967) 108, f. 34.

Flora of Solomon Islands zit

MALAITA: Kwara-ae District Kwalo and vicinity, alt. 650m, tree 10 m tall, in a clearing;

figs orange, hairy, maroon-brown when ripe; leaf blades scabrous, asymmetric, harshly scabrid. 24

Sept. 1957, Stone 2394 (BISH). — Tantalau Village, 325 m alt., tree in adjacent forest, n.v. “‘samota’’;

23 Sept. 1957, Stone 2323 (K).

Streblus Lour.

Streblus glaber (Merr.) Corner, Gard. Bull. Sing. 19 (1962) 221.

MALAITA: Kwara-ae District, Kwalo and vicinity, c. 650 m. alt., tree with milky sap, n.v.

“tou”; 29 Sept. 1957, Stone 2425 (BISH).

Streblus solomonensis Corner, Gard. Bull. Sing. 19 (1962) 224.

MALAITA: Kwara-ae District, Kwalo and vicinity, c. 650 m. alt., tree, flowers in short spikes

resembling Gnetum, n.v. ‘“‘aingaro’’; 29 Sept. 1957, Stone 2437 (BISH).

PROTEACEAE

Finschia Warburg

Finschia chloroxantha Diels, Engl. Bot. Jahrb. 54 (1916) 204; White, Pac. Sci. 3:3 (1949),

190-194, f. 3.

MALAITA: Tantalau Village, nearby forest, tree with yellow fruits; 23 Sept. 1957, Stone 2322

(BISH). TULAGI: Ridge above docks, tree with yellowish— or orange—brown fruits. 10 Nov. 1957,

Stone 2538 (BISH).

Both specimens are in complete accord with the description and figure in C.T. White’s mono-

graph of 1949. The species was already known from the Solomon Islands from Kolombangara Is.

in the New Georgia Group and is known to occur in New Guinea, the Caroline Islands, and the New

Hebrides where it has passed under several specific and generic names (e.g. Finschia densiflora C.T.

White, Grevillea elacocarpifolia Guillaumin. and Helicia micronesica Kanehira).

LORANTHACEAE

Amyema V. Tieghem

Amyema rigidiflora Danser, Bull, Jard. Bot. Buitenz. 3: 10 (1929) 298; ibid. 11 (1931) 344.

MALAITA: Kwara-ae District, near Tantalau Village, c. 325 m alt., epiphytic hemiparasite,

leaf blades narrowly deltoid—ovate, flowers in open, few-flowered, axillary umbels; leaves subopposite

or alternate; 23 Sept. 1957, Stone 2334 (BISH).

I am not entirely convinced of the correctness of the determination, though it is the closest

I have been able to come. The following slight differences may be pointed out: the above collection

has somewhat smaller flowers; acuminate rather than rounded leaf apices; glabrous inflorescence

axes and calyx, except for the marginal cilia; and slightly longer petioles (to 22 mm, not 12 mm).

There being little if any real distinction in the essential floral characters and the inadequate material,

this is provisionally considered as a slightly abberrant specimen of Danser’s specics. Known from

Talaud Is. and N.E. New Guinea.

Amyema aff. sogerensis (S. LeM. Moroe) Danser, ibid. 10 (1929) 299; ibid. 11 (1931) 246.

MALAITA: Kwara-ae District; Tantalau-Launguata trail via Kwai’ afa River, c. 325 m, alt.,

hemiparasite on tree (host = Phyllanthus sp.) shrubby, leaves mostly strictly opposite, green. corolla

red; 22 Sept. 1957, Stone 2308 (BISH, US).

212 Gard. Bull. Sing. 34:2 (1982)

This plant differs from Danser’s A. sogerensis in the absence of corolla scales. There are supposed

to be six scales ‘“‘directed upwards inside (the corolla) at 1 mm above the base” which cannot be

located in the flowers of the cited specimen. This is perhaps due to the undeveloped state of these

flowers, which appear only about half-grown.

A. sogerensis is reported from S.E. New Guinea and New Ireland. If, as I suppose, the above

specimen proves to be of the same taxon it is a logical range extension. One specimen cited by Danser

(Forbes 682) was collected at 510 m alt., the other from New Guinea (Forbes 715, Type BM) at 1500

m. The New Ireland specimen must have been from a low altitude (Port Carteret, Barclay 3517).

AMARANTHACEAE

Deeringia R. Br.

Deeringia arborescens (R. Br.) Druce, Rep. Bot. Exch. Club Brit. Is. 1916 (1917) 619. Lesti-

budesia arborescens R. Br., Prod. (1810) 414.

MALAITA: Kwara-ae District, Kwalo, c. 650 m. alt., climber in secondary growth of clearing,

leaves simple, alternate; blades elliptic. petals 5, stamens 5; fruit 4-lobed, baccate, magenta-pink,

slightly fleshy; seeds 4-7, glossy black, subcordate; 24 Sept. 1957, Stone 2368 (BISH).

A new record for the Solomons flora. Reported so far from Malaysia, New Guinea, and Queens-

land.

D. amaranthoides (Lamk.) Merr. was first reported from Bougainville by Merrill & Perry in

J. Arn. Arb. 23: (1942) 385.

WINTERACEAE

Belliolum Tieghem

Belliolum haplopus (Burtt) A. C. Smith, J. Arn. Arb. 23 (1942) 438.

MALAITA’: Kwara-ae District, Kwalo and vicinity, alt. 650 m ; tree with simple, oblanceolate,

spiralled leaves; blades grey—green beneath; 28 Sept. 1957, Stone 2429 (BISH).

ANNONACEAE

Phaeanthus Hk. f. & Thoms.

Phaeanthus macropodus (Miq.) Diels var. mollifolius Diels, Bot. Jahrb. 49 (1912) 161.

MALAITA: Kwara-ae District, Kwalo and vicinity, alt. 650 m; in forests; small tree with red or

yellowish fruits on gynophores from a torus; 28-29 Sept. 1957, Stone 2434 (BISH, US).

A new record. Determined by the late Mr. James Sinclair.

MY RISTICACEAE

Myristica L.

Myristica fatua Houtt. var. papuana Markgraf ex Diels, J. Arn. Arb. 10 (1929) 77.

MALAITA: Kwara-ae District, near Tantalau Village, in forests, c. 325 m alt., tree to 30 m.

eraser eo - =

Flora of Solomon Islands 213

high, leaves alt., blades fulvous beneath; flowers and fruits along twigs, n.v. ‘“Kakala‘a’’, 23 Sept.

1957, Stone 2346 (A, BISH, SING, US).

NEW HEBRIDES: Neguna Island, just N. of Ile Vate (Efate) Is. near summit of Mt. Mawasi,

c. 325 m. alt., tree with 2—locular fruits, red aril, n.v. “‘na pakurya niuta’”’ (transl. “the Calophyllum

of the hills’); 4 Sept. 1957, Stone 2211 (BISH, US).

All specimens were determined by the late James Sinclair. The New Hebridean specimen

has been included here for convenience and phytogeographic interest.

LAURACEAE

Litsea Lam.

Litsea perglabra C.K. Allen, J. Arn. Arb. 23 (1942) 126.

MALAITA: Kwara-ae District, Kwalo, 650 m; rain forest tree, leaf blades broadly ovate, n.v.

“arasibolu’’, 29 Sept. 1957, Stone 2421 (BISH).

Litsea solomonensis C.K. Allen, J. Arn. Arb. 23 (1942) 127.

MALAITA: Kwara-ae District, Kwai-afa River, between Tantalau and Launguata, 325 m.,

22 m tree; flowers white in axillary clusters, stamens 8 or 9, n.v. “‘sasasu’’; 22 Sept. 1957, Stone 2313

(BISH)

SANTA YSABEL: Ridge between Koloteve village and Sesedo village, southern tip of island,

175 m., tree with white flowers, n.v. “‘giagila’’ (Bugotlang.); 14 Oct. 1957, Stone 2492 (BISH).

The herbarium annotation by A. Kostermanns (SING) reads “‘Litsea timoriana Span.”’

This is a very common Litsea in the Solomons.

RUTACEAE

Micromelum B1.

Micromelum minutum Wight & Arnott, Prod. (1834) 448.

MALAITA: Kwara-ae District, Tantalau-Kwalo trail at c. 500 m. alt., rain forest tree to 15 m.

high, leaflets paler below; petals 5, creamy-white; stamens, 10; fruits with odor of lime (Citrus auranti-

folia); 24 Sept. 1957, Stone 2365 (BISH, US).

Zanthoxylum L.

Zanthoxylum megistophyllum (Burtt) Hartley, J. Arn. Arb. 47 (1966) 193. Fagara megisto-

phylla Burtt, Kew Bull. (1935) 301.

MALAITA: Kwara-ae District, Kwalo and vicinity, c. 650 m. alt., tree in forests, n.v. “‘bulungali”

(though this name is usually applied to species of Canarium); 23 Sept. 1957, Stone 2420 (BISH).

According to Hartley, material with flowers is a desideratum for this very distinct species.

The cited specimens, however, regrettably are all in ripe fruit.

214 Gard. Bull. Sing. 34:2 (1982)

MELIACEAE

Chisocheton B1.

Chisocheton sp.

MALAITA: Kwara-ae District, near Tantalau Village, 325 m. alt., young tree 7 m. tall; fruits

ramuline, subglobose, 3—4—celled, brown-fuzzy, 3 cm diam.; seeds 3 or 4, dark brown with red hilum

and white funicle; n.v. “‘latareko”; 23 Sept. 1957, Stone 2352 (BISH). — Kwalo and vicinity, 650 m.

alt., young tree, n.v. “latareko”’; 29 Sept. 1957, Stone 2432 (BISH).

Dysoxylum B1.

Dysoxylum caulostachyum Miq., Ann. Mus. Bot. Lugd.—Bat. 4 (1868) 12.

MALAITA: Kwara-ae District, Kwalo and vicinity, 650 m. alt., forest tree, fruits ramuline

or cauline, brown-fuzzy, subglobose, 5-seeded, leaves large, pinnate, with opposite leaflets; n.v.

“airande”’, 24 Sept. 1957, Stone 2382 (BISH).

EUPHORBIACEAE

Pimeleodendron Hassk.

Pimeleodendron amboinicum Hasskarl in Versl., Med. Kon. Akad. Wet. 4 (1855) 140.

MALAITA: Kwara-ae District, near Tantalau, alt. c 325 m, tree 20 m tall, in forest, leaves

simple, alternate; blades crenate; petiole basally swollen, curved; fruit elliptic, magenta-brown, red

when ripe; seed 1, brown, in elastic, milky-sapped pulp, n.v. ‘“‘aisumbu’’; 23 Sept. 1957, Stone 2349

(BISH, US).

Macaranga Thouars

Macaranga aleuritoides F'.v. Mueller, Papuan P1. (1875) 21.

GUADALCANAL: Tenaru River lowlands, swamp forest, 135-180m alt., along the river bank,

trees with 3—lobed leaves; lobes long-apiculate; 15 Sept. 1957, Stone 2282 (BISH).

Macaranga polyadenia Pax & Hoffm., Pflanzenr. Heft 63, Euph. — Mercurial. (1914) 25.

”

MALAITA: Kwalo and vicinity, alt. 650m., Kwaro-ae District, rainforest tree, n.v. ““Kweto

or ‘“‘abe-Kweto”; 29 Sept. 1957, Stone 2419 (BISH).

Aporosa B1.

Aporosa papuana Pax and Hoffm., Pflanzenr. Heft 81, Euph. — Phyllanth. (1922) 83.

MALAITA: Kwalo and vicinity, alt. 650 m., Kwara-ae District, 14m tall tree; fruits along the

twigs, small, grey, globose, fuzzy; stipules large; 24 Sept. 1957, Stone 2386 (BISH).

ANACARDIACEAE

Rhus L.

Rhus taitensis Guill., Ann. Sci. Nat. ser. 2; 7 (1837) 361.

Flora of Solomon Islands 215

MALAITA: Tantalau, forest near village, tree to 17m tall, n.v. “areko” (Kwara-ae); 23 Sept.

1957, Stone 2325 (BISH, US); same locality, tree 12m tall, leaves with 5—7 pairs of pinnae, fruiting,

n.v. ‘“‘akwasi”’; same date, Stone 2339 (BISH).

Of the vernacular names “‘akwasi’’ is preferred as “‘areko”’ is usually given to Garuga floribunda.

COR YNOCARPACEAE

Corynocarpus J.R. & G. Forst.

Corynocarpus cribbeanus (Bail.) L.S. Smith, Proc. Roy. Soc. Queensland 67 (1956) 3h:

MALAITA: Kwara-ae District, Kwalo and vicinity, alt. 600 m, tall tree in an old secondary

forest; leaves opposite, blades elliptic; fruit light green, ellipsoid, 7.5 x 4.8 cm., 1-seeded; n.v. “tbo”;

24 Sept. 1957, Stone 2367 (BISH).

ICACINACEAE

Stemonurus B1.

Stemonurus sp.

MALAITA: Fiu River, 4 mi. N. of Auki, on the W. coast, 45m alt., secondary forest, 8m tree in

a swamp area; leaves alternate, up to 15 x 6.5 cm; flowers in axillary, cymose panicles, or on the twigs;

19 Sept. 1957, Stone 2296 (BISH).

LEEACEAE

Leea Royen ex L.

Leea tetramera Burtt, Kew Bull. (1935) 304. Leea suaveolens Merrill & Perry, J. Arn. Arb.

22 (1941) 381.

GUADALCANAL: Tenaru River, lowland swamp forest, alt. 135 m, small tree with bipinnate

leaves; flattened, purplish fruits, n.v. “‘borabora’’; 15 Sept. 1957, Stone 2280 (BISH, US).

RHAMNACEAE

Alphitonia Reissek

Alphitonia incana (Roxb.) Teysm. & Binn. ex Kurz, J. Bot. (Lond.) 11 (1873) 208. Rhamnus

incana Roxb., Hort. Beng. (1814) 17; F1. Ind. 1(1820) 603. Alphitonia moluccana Teysm, & Binn.,

Cat. Hort. Hog. (1866) 221, nomen. A. zizyphoides (Spt.) A. Gray, Bot. U.S. Expl. Exped. 1 (1854)

278, t. 20. Rhamnus zizyphoides Spr., F1. Hal. Mant. (1807) 37; Syst. 1 (1825) 768.

MALAITA: Kwara-ae District, Kwalo and vivinity, 650m. alt., tree with leaf blades almost

white beneath except on the nerves, wood sweet-fragrant, n.v. “kwanasia’’, 28-29 Sept. 1957, Stone

2424 (BISH, US).

A common tree, frequent in secondary forests.

216 Gard. Bull. Sing. 34:2 (1982)

ELAEOCARPACEAE

Aceratium DC.

Aceratium insulare A.C. Smith, J. Arn. Arb. 25 (1944) 118.

MALAITA: Kwara-ae District, near Tantalau village, in forests, 325 m alt.; 20 m tree; leaf

blades finely toothed, elliptic, glossy green and slightly pubescent above, flat green and pubescent

beneath, base slightly emarginate, tip acuminate-apiculate; flowers axillary; fruit short-cylindric,

3 cm long, 2 cm diameter, greenish at base, ripening to pink and reddish, attached by a long, slender

peduncle; n.v. “‘sura’u’u’’; 23 Sept. 1957, Stone 2341 (BISH, US).

TILIACEAE

Trichospermum B1.

Trichospermum psilocladum Merrill & Perry, J. Arn. Arb. 20 (1939) 341. Fig. 1.

SANTA YSABEL: Horara Islet, Tatamba Lagoon, near sea-level, in dry forests, tree 15 m. tall,

with Calophyllum inophyllum, Dillenia crenata, Parinarium glaberrimum, Casuarina nodiflora, Gnetum

gnemon, Evodia sp., etc.; 24 Oct. 1957, Stone 2514 (BISH).

This is very similar in general appearance to T. ikutai Kanehira (1932), which is probably the

earliest valid name.

Trichospermum kajewskii Merril & Perry, J. Arn. Arb. 20 (1939) 340.

MALAITA: Kwalo and vicinity, alt. 650 m, Kwara-ae District, rain forest; tree 15 m tall, n.v.

“sulu’’,; 29 Sept. 1957, Stone 2430 (BISH). (Vegetative).

Determined by T.C. Whitmore.

DILLENIACEAE

Dillenia L.

Dillenia crenata (A.C. Smith) Hoogland, Blumea 7 (1952) 42. Wormia crenata A. C. Smith,

J. Arn. Arb. 22 (1941) 498.

SANTA Ysabel: Horara Islet, Tatamba Lagoon 15 m alt., common big tree in dry forests;

petals white, connate by their tips, falling as a unit; stipules adnate to the petioles, but deciduous;

31 Oct. 1957, Stone 2520 (BISH).

This specimen is an excellent match for the description of this clear-cut species.

GUTTIFERAE

Calophyllum L.

Calophyllum paludosum C.T. White, J. Arn. Arb. 31 (1950) 98.

MALAITA: Kwara-ae District, Kwalo, 650 m alt., in rain forest; large tree; n.v. “‘koumanu”’

(compare Hawaiian “kamani’”’); 29 Sept. 1957, Stone 2431 (BISH, US).

+ise So”

rROGKXS

OX Sats vor

Neuse

Fig. 1. Trichosperma psilocladum Merr. & Perr. (Tiliaceae). From Stone 2514. a— habit of fertile

branch. b— portion of inflorescence with flower buds. c— floral details: longitudinal section

of flower, with apical view of compressed ovary; petal, stamen, and (below) bracteole. d— fruit,

side view. e— fruit, top view. f— fruit, transvers section, g— fruit, longitudinal section. h— side

and top views of seed with hair-tufts. i— detail of leaf margin showing trichome positions.

j— two representative stellate trichomes.

218 Gard. Bull. Sing. 34:2 (1982)

LEGUMINOSAE

Desmodium Desv.

Desmodium umbellatum (L.) DC., Prodr. 2 (1825) 325.

MALAITA: Kwara-ae District, Tantalau area, forest, alt. 325 m, tree 12 m. tall, petals white,

n.v. “‘aida-ahi’’; 23 Sept. 1957, Stone 2329 (A, BISH, US).

MYRTACEAE

Decaspermum J. R. & G. Forster

Decaspermum fruticosum J. R. & G. Forster, Char. Gen. P1. (1776) 74, pl. 37.

SANTA YSABEL: Horara Islet, 15 m. alt., small tree, flowers with greenish-white petals; 31

Oct. 1957, Stone 2519 (A, BISH, US).

MALAITA: Kwara-ae District, vicinity of Tantalau village, alt. 325 m, small tree 10 m. high;

small flowers with white petals and pink stamens, n.v. “‘oridi’’, 23 Sept. 1957, Stone 2338 (A, BISH,

US).

This is an attractive little tree and occurs in abundance in the cited localities.

MELASTOMATACEAE

Melastoma L.

Melastoma malabathricum L., Sp. P1. (1753) 390.

MALAITA: Tantalau village, in secondary growth; shrub, petals pink; stamens pink, yellow,

white; n.v. “‘maragwana’’; 23 Sept. 1957, Stone 2324 (BISH, US).

Walker (1948, 79) states that the same vernacular name is also applied to a species of Maesa

(Myrsinaceae).

ARALIACEAE

Plerandra A. Gray

Plerandra solomonensis W.R. Philipson, Bot. Bull. Brit. Mus. (N.H.) 1:1 (1951) 10. Fig. 2.

MALAITA: Kwara-ae District; Kwalo and vicinity, alt., c. 650 m, tree with palmate leaves,

infructescence umbellate, fruits black, n.v. ‘‘singoria’’; 28-29 Sept. 1957, Stone 2438 (BISH).

There are some points of discrepancy between this specimen and the description provided by

Philipson which is very short and somewhat incomplete. The above collection shows the following

features: mature flower bud 6 mm long or slightly longer; calyx undulate-discoid, the 5 lobes obscure,

spreading; petals 5, nearly 1 mm thick; stamens (in staminate flowers) approximately 160; ovary

(in female flowers) almost always 11-celled; fruit (seemingly ripe) only 18 x 17 mm (rather than 28 x

15 mm, Philipson’s dimensions); leaflets 5 or 7, their petiolules 4-5 cm long (not 2.5 cm), with

evident lateral nerves; pedicels about 5 per umbellule (not 20).

TMI

\ | )

8Eve SOG

Fig. 2. Plerandra solomonensis W.R. Phil, (Araliaceae). From Stone 2438. a— branch and leaf.

b— small portion of inflorescence. c— umbellule. d— floral details: longitudinal section of flower;

separate stamens, e— flower. f— fruit. g— fruit, enlarged, side and top views.

220 Gard. Bull. Sing. 34:2 (1982)

Gastonia Comm. ex Lamk.

(Peekeliopanax Harms)

Gastonia spectabilis (Harms) Philipson, Blumea 18 (1970) 494.

Peekeliopanax spectabilis Harms, Notizbl. Bot. Gard. Berlin 9 (1926) 478.

MALAITA: Kwara-ae District, Kwalo and vicinity, c. 650 m. alt.; large, single-boled tree, about

30-35 m. tall, distally few and short branched, dense leafy crown of very long, 1—pinnate leaves (to

2 m long) with c. 14 pairs of leaflets, rachis swollen at nodes; not flowering; n.v. “‘simalau”’; 28 Sept.

1957, Stone 2409 (BISH).

This is certainly one of the most splendid plants in this interesting family. The height, massive-

ness of the leaves, strength of the wood, and the relative rarity of this species are all notable charac-

teristics that impress the botanical traveller in the Solomons. It was described from material collected

in the Bismarck Archipelago (New Britain) by Peekel. Only two trees of this species were seen during

my entire stay in the Solomons and Bismarcks.

STYRACACEAE

Styrax L.

Styrax agreste (Lour.) G. Don, Gen. Syst. 4 (1837) 5.

MALAITA: Kwara-ae District, Kwalo, 650 m. alt., 20 m. tree, ovary superior, corolla 4-5-lobed,

white; stamens 8 or 10, epipetalous; ovules few, pendulous; leaves simple, alternate; stipules none;

n.v. “aigasi’”’; 29 Sept. 1957, Stone 2407 (A, BISH, US).

APOCYNACEAE

Carruthersia Seemann

Carruthersia mollis Markgraf,Gard. Bull. Sing. 22 (1967) 25.

MALAITA: Tantalau-Fiu River area, Kwara-ae District; vine, corolla white, lobes with yellowish

or pinkish spot at base (where they overlap); leaves subcordate; sap milky; alt. 260 m; 25 Sept. 1957,

Stone 2399 (BISH, US).

This specimen differs from the original description, which was based solely on the holotype

collection, Whitmore 2749, from Bogotu, Santa Ysabel, in having white corollas with only a spot

of pink (or yellow) at the base of each corolla lobe, rather than red corolla tubes and pink lobes.

No important differences were observed in other features so it would appear that the species is rather

variable in corolla color.

- ASCLEPIADACEAE

Anodendron A. DC.

Anodendron paniculatum (Roxb.) A. DC., Prodr. 8 (1844) 444.

MALAITA: Kwalo — Tamba’a, Kwara-ae District, alt. 650m., a vine, climbing trees, with terete,

brown stems; opposite leaves, nodes annular; fruit of 2 twinned carpels, terete-conic, 12 cm long,

nearly 2 cm diam. at the base; seeds plumose, body of seed oblong-elliptic, flat, thin, 17 x 5 mm.,

attenuate at the apex to a short column 6 x % mm, bearing a white plume 6 cm long of numerous,

straight, silky hairs; 28 Sept. 1957, Stone 2447 (BISH).

Flora of Solomon Islands 271

LOGANIACEAE

Fagraea Thunb.

Fagraea racemosa Jack in Roxb., F1. Ind. ed. Carey, 2 (1824) 35.

MALAITA: Kwara-ae District, near Tantalau Village, c. 325 m alt., 7 m. tree with drooping

branches, large, dull green leaf blades; white, zygomorphic flowers in terminal racemes, n.v. “‘ngara’’;

23 Sept. 1957, Stone 2328 (BISH).

Geniostoma J.R. & G. Forster

Geniostoma brassii Merr. & Perry, J. Arn. Arb. 23 (1942) 409.

MALAITA: Kwara-ae District, near Tantalau; 12 m tree, fruits 1.4 cm long, n.v. ‘“‘mahus-husi’”’

(one of several spp. so named); 23 Sept. 1957, Stone 2356 (BISH). - Near Kwalo, c. 650 m. alt., 7m

tree, or shrubby, leaves opposite, stipulate; blades elliptic, acuminate, paler beneath, rather thin and

fragile; petiole 2 cm long; fruits axillary, short-pedicellate, ovoid-ellipsoid, greenish-white, 2-celled;

seeds numerous, yellow, in juicy pulp, placenta axile; calyx 5-lobed; 24 Sept. 1957, Stone 2392

(BISH).

SAPOTACEAE

Planchonella Pierre

Planchonella obovoidea Lam, Bull. Jard. Bot. Buitenz. III: 7 (1925) 207.

MALAITA: Kwara-ae District, Kwalo and vicinity, Tamba’a path, rain forest at c. 650 m.

altitude, small tree with alternate leaves with close, minute, rusty coloured, scaly indumentum, n.v.

“mumu’’, 29 Sept. 1957, Stone 2423 (BISH).

Vegetative. The holotype is from Kei Is.

COMPOSITAE

Blumea DC.

Blumea riparia (B1.) DC., Prod. 5(1836) 444.

MALAITA: Kwara-ae District, Tantalau village, 325m alt.; scandent shrub in clearings; 23 Sept.

1957, Stone 2332 (BISH).

Determined at the Rijksherbarium, Leiden, in 1970.

FLAGELLARIACEAE

Flagellaria L.

Flagellaria indica L., Sp. P1. (1753) 333.

MALAITA: Fiu River, at 200 m. alt., in secondary forests with Macaranga, Hibiscus, and

Ficus; large vine with leaf tips as coiled tendrils; inflorescence large, terminal, branched. Perianth

white; 25 Sept. 1957, Stone 2395 (BISH).

222 Gard. Bull. Sing. 34:2 (1982)

COMMELINACEAE

Pollia Thunb.

Pollia thyrsiflora Endl. ex Hassk. in Miq., P1. Jungh. (1852) 150.

GUADALCANAL: Tenaru River, in lowland forest, 135-175 m alt.; 15 Sept. 1957, Stone 2286

(BISH).

New record for the Solomons; not listed in Whitmore (1966).

PALMAE

Heterospathe Scheffer

Heterospathe woodfordiana Becc., Webbia 4:2 (1914) 281.

MALAITA: Kwalo and vicinity, K wara-ae District, alt. 650 m, small, erect, pinnate-leaved palm;

fruits red, n.v. “araramei’’, 28 Sept. 1957, Stone 2408 (BISH, 3 sheets).

CYPERACEAE

Scleria Berg.

Scleria lithosperma (L.) Sw., Prod. Veg. Ind. Occ. (1788) 18.

SANTA YSABEL: Above Tatamba, on ridge between Koloteve and Sesedo, c. 275 m. alt.,

in forests; 15 Oct. 1957, Stone 2494 (BISH).

Scirpus L.

Scirpus fuirena T. Koyama, J. Fac. Sci. Univ. Tokyo III, 7:6 (1958) 361.

Fuirena umbellata Rottb., Descr. Icon. P1. (1773) 70, t. 19, f. 3.

MALAITA: Auki swamps, sedge with pentagonal stems, associated with Pandanus; 2 Oct.

1957, Stone 2461 (BISH).

NOMENCLATURE OF JOINVILLEA (JOINVILLEACEAE)

BENJAMIN C. STONE

Department of Botany, University of Malaya, Kuala Lumpur

Malaysia

ABSTRACT

The nomenclatural status of the name forthe genus Joinvillea and for its

two species is reviewed. It is concluded that the correct (first validly publ-

lished) generic name is Joinvillea Gaudich. ex Brongn. & Gris, 1861; Gau-

dichaud’s original publication of 1841 contravenes Articles 42 and 44 of the

International Code of Botanical Nomenclature. The two species are correctly

J. plicata (Hk. f.) Newell. & Stone, and J. ascendens Br. & Gr.

Joinvillea, a small genus previously attributed to the Flagellariaceae, hasnow been shown

on good evidence to constitute a small independent family (Tomlinson & Smith, 1970;

Lee, Yap & Liew, 1975). A recent taxonomic revision has been published by T. K.

Newell (1969). In this work, evidence is given to show that the generic name, coined by

C. Gaudichaud, dates as a legitimate botanical name only from a publication by A.

Brongniart & Gris (1861). Therefore, the correct names of all species of the genus must

date either from this same publication or from later publications, earlier appearances of

the name in print being considered illegitimate. Newell regards the genus as being com-

prised of two species, both of them being further divided into subspecies. For these

species, Newell concluded that the correct names are (1) J. ascendens Brongn. & Gris

(1861), with four subspecies; and (2) J. plicata (Hk.f.) Newell & Stone (1967), based

on Flagellaria plicata Hk. f. (1855), with two subspecies.

Lately this view has been challenged by St. John (1978), who maintains Newell’s

taxonomic conclusions but, on the basis of a different interpretation of the validity of

publication of the generic name, regards J. plicata as a synonym of J. elegans Gaudich.

In this note, evidence is reviewed and it is concluded that the nomenclatural discussion

of St. John is unconvincing, and that therefore, the nomenclature adopted by Newell &

Stone (1967) and by Newell (1969) is correct.

The crux of the nomenclatural problem is the applicability of Article 44 of the ICBN

(International Code of Botanical Nomenclature). This article reads as follows: “The

name of a species or of an infraspecific taxon published before 1 Jan. 1908 is validly

published if it is accompanied only by an illustration with analysis showing essential

characters (see Art. 32, Note 2). Note: Single figures of microscopic plants showing

the details necessary for identification are considered as illustrations with analysis

showing essential characters.”

This article refers back to Article 32 (to Note 2, but in fact this is an error, and the

reference should be to Note 3), which defines valid publication. Note 3 reads: In certain

circumstances an illustration with analysis is accepted as equivalent to a description

(see Arts. 42 and 44).

We are thus led also to Article 42, which reads as follows: ‘‘The publication of the name

of a monotypic new genus based on a new species is validated either by (1) the provision

of a combined generic and specific description (description generico-specifica) or diag-

223

924 Gard. Bull. Sing. 34:2 (1982)

nosis, or (2), for generic names published before 1 Jan. 1908, by the provision of an

illustration with analysis showing essential characters (see Art. 32, Note 3).”’

Gaudichaud conceived the genus Joinvillea. Instead of publishing it as a new name with

a diagnosis or description, he merely coined the name of the genus and of two species

which he believed the genus to be composed of. The two binomials were published as

titles to illustrations (Plates) with ‘“‘analyses’’ in his ‘Botanique’ Atlas, embodying results

of the world tour of the French exploring expedition of 1836-37 in the corvette “La

Bonite’ commanded by Capt. Vaillant (Gaudichaud, 1841). Two such Plates were pub-

lished, Pl. 39 and Pl. 40, the former showing “‘Joinvillea ascendens’’ (figs. 1—6, showing

short sections of a stem, a sterile shoot, and short cross sections of part of a leaf), the

remaining figures (7-26) showing ‘‘Joinvillea elegans’’ (figs. 7—9 show stem, leaf, and

inflorescence, figs. 10-26 show analytical details of the flowers and fruit and sections

thereof). Details of both species can be found in both Plates.

St. John’s argument in briefis this: Joinvillea ascendens is not a validly published name,

since no flowers or fruits or other “‘essential’’ parts were shown in Gaudichaud’s illus-

trations. Therefore Joinvillea as a genus is monotypic. The illustration of Joinvillea

elegans shows the “‘essential’’ parts, i.e. flowers and fruits; it is therefore validly published,

and can be taken as a joint or generico-specific publication of the genus and species.

The genus, therefore, dates from 1841, as does the species J. elegans, which therefore has

priority over the synonymous Flagellaria plicata Hk. f. of 1855.

This argument is however unconvincing, indeed specious, for the following reasons.

First, it is obvious that Gaudichaud intended the genus to consist of two species; it was

not monotypic in his original conception. Secondly, the insistence by St. John that the

analyses provided for J. ascendens are unacceptable because they do not show flowers or

fruits or other essential characters, is misleading. The ICBN does not specify what is meant

by ‘‘analyses’’ or by the term “essential characters.’ Inherently, both can and often do

include non-floral and non-fruit characters. If Gaudichaud meant to support his species

concept by the use of features of the stem or leaf anatomy, this cannot be rejected on

legalistic grounds (whether the anatomical features are significant is another matter).

In other words, the analyses provided for J. ascendens, of vegetative characters, are just

as satisfactory, from the legalistic point of view, in conforming to the stipulations of

Art. 42 and Art. 44, as those of floral and fruit details provided for J. elegans. With

our retroactive Code, we are thus forced to acknowledge that the generic name Joinvillea,

as it appears in the original publication of 1841 by Gaudichaud, is illegitimate, as it

clearly is intended to designate a genus of two species. Article 42 is very clear that only

monotypic new genera can be validated by the publication of a plate with analyses.

Joinvillea fails to meet this stipulation in 1841.

Newell & Stone (1967) and Newell (1969) accept the name Joinvillea plicata (Hk. f.)

Newell & Stone for J. elegans Gaudich. nom. illegit. (1841). The genus itself was validly

established and attributed to Gaudichaud by Brongniart and Gris (1861); they also

established the species J. ascendens in the same publication. As shown by Newell, Brong-

niart and Gris also published a synonymous name for this species (J. gaudichaudiana),

which previously has been used for Samoan plants. Three other species have been described,

all now reduced to the status of subspecies. A short synopsis of the nomenclature follows.

JOINVILLEA Gaudich. ex Brongn. & Gris, Bull. Soc. Bot. France 8 (1861) 268.

Flagellaria subg. Chortodes Hook. f. in Hook., J. Bot. Kew Gard. Misc. 7(1855) 200,

tab. VI. Type species: J. elegans Gaudich. ex Brongn. & Gris,1. c. which = J. plicata

(Hk. f.) New. & Stone subsp. plicata.

Nomenclature of Joinvillea 22>

(1) Joinvillea ascendens Brongn. & Gris, 1.c. 1861.

(1a) subsp. ascendens — Hawaiian Islands, endemic.

(1b) subsp. samoensis Newell, Journ. Arn. Arb. 50 (1969) 548. J. gaudichaudiana

sensu Christophersen, Bishop Mus. Bull. 128 (1935) 46, pp. J. gaudichaudiana

var. samoensis (Newell) Deg. & Deg., Fl. Haw. fam. 53a: 3.31.1973.

Western Samoa, endemic.

(1c) subsp. borneensis (Becc.) Newell, 1. c. (1969) 549. J. borneensis Becc., Nelle

For. Borneo (1902) 198. J. malayana Ridl.,J. Str. Br. Roy. Asiat. Soc. 44

(1905) 199 and F1. Mal. Pen. 4 (1924) 368. Malesia (Malaya, Sumatra,

Borneo, Philippines (Palawan, Jolo), and Caroline Islands (Ponape).

(1d) subsp. glabra Newell, 1. c. (1969) 550.

New Caledonia, endemic.

(2) Joinvillea plicata (Hk. f.) Newell & Stone, Taxon 16(1967) 193. Newell, J. Arn.

Arb. 50 (1969) 550. Flagellaria plicata Hk. f J. Bot. Kew Gard. Misc. 7 (1855) 200.

Pl. VII.

(2a) subsp. plicata. — East Melanesia (Solomon Islands, New Hebrides, Fiji, and

New Caledonia, including Isle of Pines). J. elegans Gaudich., Atl. Voy. Bonite

(1841) Pl. 39. 40, figs. 7-26, nom. nud. illegit. J. elegans Brongn. & Gris,

1. c. (1861) 268. Flagellaria elegans Seem., F1. Vit. (1868) 315.

(2b) subsp. bryanii (E. Christophersen) Newell, 1. c. (1969) 552. J. bryanii E.

Christophersen, Bishop Mus. Bull. 128 (1935) 44. J. elegans subsp. bryanii

(E. Christophersen) St. John, Phytologia 40 (1978) 372.

Western Samoa, endemic.

References

Brongniart, A. & A. Gris. 1861. Note sur le genre Joinvillea de Gaudichaud et sur la famille des Flagel-

lariees. Bull. Soc. Bot. France 8, 264-269.

Gaudichaud, C. 1841. Atlas .Botanique. Voyage autour du monde execute pendant les annees 1836 et

1837 sur la corvette la Bonite, commandee par M. Vaillant. A. Bertrand,. Paris. (no pagin.).

Cfr. Plates 39, 40.

Hooker, J. D. 1855. On Chortodes, a subgenus of Flagellaria, from the Isle of Pines (New Caledonia).

Hook, Jour. Bot. Kew Gard. Misc. 7, 198-200.

International Code of Botanical Nomenclature. Adopted by the International Botanical Congress.

Regnum Vegetabile, Utrecht. 1972, 1977, et seq.

Lee, David W., K.P. Yap, and F.Y. Liew. 1975. Serological evidence on the distinctness of the mono-

cotyledonous families Flagellariaceae, Hanguanaceae, and Joinvilleaceae. Bot. J. Linn. Soc. 70,

77-81,

Newell, T.K. and B.C. Stone. 1967. Flagellaria (Chortodes) plicata Hooker fil. is a Joinvillea, Taxon

16, 192-194.

Newell, T.K. 1969. A study of the genus Joinvillea (Flagellariaceae). J. Arnold Arboretum 50,527-55S5.

St. John, H. 1978. Revision of Joinvillea (Joinvilleaceae). Pacific Plant Studies 37. Phytologia 40,

369-374.

Tomlinson, P.B. and A.C. Smith. 1970. Joinvilleaceae, a new family of Monocotyledons. Taxon 19,

887-889.

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IN MEMORIAM

Dr. Monte Gregg Manuel, bryologist, 1947 - 1981.

Friends, bryologists, and Malesian botanists will be saddened to learn of the death

of Monte G. Manuel, who for three years past was staff bryologist and bryological curator

in the Department of Botany, University of Malaya, Kuala Lumpur. Monte had just

departed from Malaysia after his contract and was taking up a new post at the University

of Nairobi, Kenya, but suffered a sudden stroke and died in Nairobi in April 1981.

Monte G. Manuel was born in Missoula, Montana, U.S.A., on 29th May 1947, and

received his early education in Lakes High School in Tacoma, Washington. After

obtaining an Associate in Arts degree from Lower Columbia College in Longview,

Washington, in 1968, he entered Central Washington State College in Ellensburg and

majored in botany, with a minor in geology, receiving his B.A. in 1970. He then continued

at the University of Southwestern Louisiana in Lafayette, studying under William D.

Reese, and gained the M. Sc. degree there. He completed his academic work at the Univer-

sity Park laboratory of Dr. Ronald A. Pursell (The Pennsylvania State University) and

received his Ph. D. in 1976. He spent a postdoctoral year at the Farlow Library and

Herbarium of Harvard University, Cambridge, Massachusetts, where he worked partly

with Dr. Norton G. Miller.

Monte had worked for the U.S. Forest Service in Oregon and had a taste for nature

and the outdoors. During his graduate work he taught in the biology laboratories and

did duties in the herbaria. His Research Fellowship at Harvard was a very valuable expe-

rience and he then took a teaching post at Fort Lewis College in Durango, Colorado,

for a year (1977-8) where he was very successful. But the tropical floras enthralled him

and he won a post at the University of Malaya, where he worked from May 1978 to

March 1981. Here he accomplished much in both teaching and research, publishing

several papers, and instituting advanced courses in bryology. A devoted botanist, Monte

had high research ideals. Before he left Malaysia, he had prepared a generic moss Flora

for the area and luckily this was published, albeit posthumously*. This work will be a

permanent memento of his application, ability, and enthusiasm, and will certainly become

a standard text in the region at universities.

Monte, a bachelor, had many friends in Malaysia and Singapore, in many fields

apart from the academic life. His untimely demise, at the age of 34, has deprived tropical

bryology of an exemplary taxonomist, and many of us of a good and gifted friend.

B.C. STONE

Department of Botany

University of Malaya

Kuala Lumpur,

Malaysia

***\ Generic Moss Flora of Peninsular Malaysia and Singapore" by M. G. Manuel. Federation Museums’

Journal (new series) vol. 26, part 2. Pp. 1—163.1981.

227

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SOME ASPECTS OF INTERRELATIONSHIPS AMONG THE

DRYNARIOID FERNS*

SUBHASH CHANDRA

Pteridology Laboratory

National Botanical Research Institute

Lucknow-226001, India.

ABSTRACT

Morphological characters of 15 species belonging to seven genera of dryna-

rioid ferns are presented and their significance relative to their origin, inter-

relationships, and trend of specialization discussed. Based on morpho-

anatomical evidence accumulated, two morphological groups of drynarioid

ferns are proposed: Drynaria and Aglaomorpha. The attributes considered are:

palea, rhizome; vegetative frond including epidermal cells, stomata, hypo-

dermis, mesophyll, venation; fertile frond including sorus, sporangium, and

spores; and gametophyte.

Existing knowledge of the formal taxonomy and phylogenetic trends

among the drynarioid ferns are compared with this study and the different lines

of descent are traced. It is suggested that the drynarioid ferns are probably

diphyletic in origin and the trend of evolutionary progression has been

towards reduction.

INTRODUCTION

The drynarioid group of ferns is an assemblage of large epiphytes of the Polypodiaceae

regarded by most pteridologists as intimately related to each other. As currently under-

stood, they consist of 8 genera (Copeland,1947) and 37 species (Christensen: 1905—1906,

1917, 1934) throughout their range of distribution. Drynaria includes 20 species

(Copeland, 1947; Holttum, 1954; Tindale, 1961) while the other genera are each repre-

sented by one or two species in the Indo-Malayan region. Several authors have expressed

different opinions on the systematic position of the drynarioid ferns (Christensen, 1938;

Ching, 1940; Holttum 1947, 1954; Copeland, 1947, 1960; Alston 1956; Pichi-Sermolli,

1958, 1959; Mehra, 1961; Nayar, 1970, 1974; Bierhorst, 1971; Wagner, 1973). Recently

Crabbe et al. (1975) proposed a different classification in which drynarioid ferns were

treated as a subfamily of Polypodiaceae without any further subdivision, while Ching

(1978) gave the drynarioid ferns a formal status as a very natural and distinct family-

Drynariaceae.

This group is quite different from most other polypodiaceous ferns and is considered as

one of the most controversial with regard to their relationships and evolution. The views

of Goebel (1928), Copeland (1929, 1947), Christensen (1938), Ching (1940), Holttum

(1947, 1954), Nayar (1954, 1955, 1959, 1965), Nayar and Kachroo (1953), Zamora

(1975), and Zamora and Vargas (1973 a, b) are varied and are highly susceptible to

revision because they are based only on a limited number of species or on selected

morphological characters. In view of the controversy regarding the relationship among

the drynarioid ferns, the present investigation aims to explain the possible relationship

among them and to discuss whether the splitting of the drynarioid ferns into two distinct

morphological groups on the basis of morphological attributes is justifiable.

*N.B.R.I., Research Publication No. 121 (N.S.)

een

230 Gard. Bull. Sing. 34:2 (1982)

MATERIALS AND METHODS

The present study is based on 15 species of drynarioid ferns (Table 1). Most of the

materials used were collected from the fernery of the Natural Science Research Centre,

University of the Philippines, Diliman, Quezon City except D. fortunei, D. laurentii,

D. propinqua, Pseudodrynaria, and Thayeria, which were studied from herbarium

specimens (Table 1).

Since the relationship and classification based on similarities and differences in one

organ are unreliable unless supported by the totality of evidence from other parts of the

plants, all possible morphological characters have been taken into account in this study.

For the convenience of description, all the species are arranged in two morphological

groups viz. Drynaria and Aglaomorpha. The Aglaomorpha group includes Drynariopsis,

Aglaomorpha, Merinthosorus, Pseudodrynaria, and Thayeria, while the Drynaria group

has Drynaria and Photinopteris (Chandra 1979 b, 1980).

TABLE 1. SPECIES STUDIED

Taxa

Drynaria descensa Copel.

D. fortunei J. Sm.

D. laurentii (Christ) Hieron

D. propinqua (J. Sm.) Wall.

D. quercifolia (L.) J. Sm.

D. rigidula (Sw.) Bedd.

D. sparsisora (Desv.) Moore

Photinopteris speciosa (Willd.) Mort.

Drynariopsis heraclea (Kunze) Ching

Pseudodrynaria coronans (Wall.) Ching

Aglaomorpha splendens (J. Sm.) Copel.

A. meyeniana Schott

A. pilosa (J. Sm.) Copel.

Merinthosorus drynarioides (Hook.) Copel.

Thayeria cornucopia Copel.

Source of Material

U.P., NSRC, Fernery; P.M. Zamora,

73-0408 (PUH)

S. Suzuki, PNH 63594; 63605

Jean Louis, PNH 29441

Cavalerie, PNH 26053; Petelote, PNH 63596;

B.K. Nayar, LWG, 74339

U.P. NSRC, Fernery; S. Chandra, 62(PUH)

U.P., NSRC Fernery; N.S. Vargas, 72-028 (PUH).

U.P., NSRC, Fernery; S. Chandra, 16 (PUH)

U.P., NSRC, Fernery; N.S. Vargas, 72-0186,

(PUH).

U.P., NSRC, Fernery, N.S. Vargas,

73-0449 (PUH)

S. Chandra, LWG, 94125; Poilana 1382 MICH;

Y.W. Taam; 1861 MICH: Thakur Rup Chand;

3988 MICH.

U.P., NSRC, Fernery;S. Chandra,48, 105 (PUH).

U.P., NSRC, Fernery; S. Chandra, 109 (PUH).

U.P., NSRC, Fernery; S. Chandra, 86 (PUH).

D.F. Grether, PNH 3680; Grether and Wagner,

PNH 3973.

Edano, PNH 8770; Jacobs, PNH 10420.

On drynarioid ferns 231

OBSERVATIONS

Among the Drynaria group there appear to be two independent lines of evolution

from the ancestor of the Drynaria group of species where one line gave rise to Drynaria

and the other to Photinopteris. Contrary to the observations of Copeland (1947) and

Nayar (1965), evolution appears to be in two directions from the Drynaria line (Fig. 2).

One line gave rise to D. rigidula and the other to D. quercifolia, independently of one

another.

D. rigidula is unique and is quite different from all other species of Drynaria in having:

(1) rhizomes with undissected dorsal median vascular strands (as in Photinopteris), (2)

stipitate pinnate fronds, (3) small foliar epidermal cells with shallow sinuous walls,

(4) small stomata, (5) fewer number of areoles between main veins, (6) prominently

impressed sori, (7) stellate paraphyses, and (8) small spores and sporangia. This possibly

indicates that D. rigidula represents a line of evolution separate from that of other

species of Drynaria but from the same common ancestor. This species, however, shows

some similarities with Photinopteris in the following features: (1) undissected dorsal

median vascular strand of the rhizome, (2) stipitate pinnate fronds, (3) two-ranked

leaf arrangement, (4) small stomata, and (5) smaller sporangia and spores.

All other species of Drynaria have presumably evolved from D. quercifolia in two direc-

tions. One line gave rise to D. sparisora and the other to all other Drynaria species.

D. sparsisora exhibits relatively advanced features, viz. (1) thick coriaceous texture of

the lamina with cartilaginous margins, (2) vestigial stipe articulation, (3) absence of

xylem parenchyma between the tracheids in the meristeles of the rhizome, (4) slightly

thickened upper foliar epidermis, (5) laminar hypodermis, (6) small stomata, (7) small

upper foliar epidermal cells with shallow undulations, (8) small spores with spinulose

exine ornamentation, (9) smaller sporangia (10) slightly reduced fertile fronds with

minute sori, and (11) few elongated soral receptacles. This propbably indicates that

D. sparsisora is comparatively more advanced than D. quercifolia and has possibly evolved

from it in a separate line. In contrast to Copeland (whose opinions of 1929 and 1947

are diagrammatically interpreted in fig. 1) this idea confirms the view of Holttum (1954)

who considered D. quercifolia to be closer to Drynariopsis than to D. sparsisora. Com-

parative morphology of the sporophyte suggests that all other Drynarias are presumably

evolved from D. quercifolia.

Among all Drynaria studied, D. descensa appears to be a reduced derivative of D.

quercifolia which it resembles most (Copeland, 1960). D. descensa exhibits relatively

advanced features viz. (1) small stature, (2) small nest leaves, (3) reduced xylem tissue

(1-2 layers of tracheids) in the rhizome, (4) absence of xylem parenchyma between the

tracheids in the meristeles of the rhizome, (5) paleae with shield-like bases and abruptly

narrowed apices, (6) paleal margin with minute dentations, and (7) localized geographical

distribution. This supports the contention that D. descensa is a reduced dervative of D.

quercifolia and is therefore comparatively more advanced.

Photinopteris is similar to Aglaomorpha in the following features: (1) paleal morpho-

logy (A. splendens), (2) reduced fertile fronds, (3) development of coenosori, (4) verrucate

areolate spores (A. splendens and A. meyeniana), and (5) small spores (A. pilosa).

Photinopteris is similar to Drynariopsis in the following features: (1 ) indistinctly clath-

rate paleae with (2) dentate margins, these (3) with glandular hairs, and (4) presence of

foliar hypodermis and hydathodes. Though similar to Aglaomorpha and Drynariopsis in

232 Gard. Bull. Sing. 34:2 (1982)

some respects, Photinopteris differs from Aglaomorpha and Drynariopsis in such features

as: (1) two-ranked leaf arrangement, (2) stipitate pinnate fronds (as in D. rigidula), (3)

undissected dorsal median vascular strand of the rhizome (as in D. rigidula), (4) paleae

with hairy, dentate margins, (5) club-shaped glandular hairs with waxy cap-like secretions,

(6) large epidermal cells with deeply sinuous walls, and (7) low stomatal frequency.

Photinopteris is similar to the Drynaria group in the following features: (1) paleae with

hairy, dentate margins and glandular marginal hairs with waxy cap-like secretions, (2)

two alternate rows of fronds on the rhizome, (3) pinnate fronds (as in D. rigidula), (4)

undissected dorsal median vascular strands of the rhizome (as in D. rigidula), (5) large

foliar epidermal cells with deeply sinuous walls, and (6) low stomatal frequency. In view

of the above, it is here suggested that Photinopteris be included in the Drynaria group.

Photinopteris, however, is comparatively more advanced than Drynaria in having: (1)

long, wide-creeping rhizomes, (2) small stature with thick coriaceous fronds, (3) dimorp-

hic fronds with reduced fertile laminae, (4) laminae with a hypodermis, hydathodes,

and differentiated mesophyll, and (5) small spores with verrucate exine ornamentation.

Photinopteris has, therefore, evolved separately from the ancestors of Drynaria and in a

different line of evolution.

This combination of characters possibly indicates that Photinopteris is probably evolved

independently of Drynaria but from the same common ancestor and that its similarities

with Drynaria (especially with D. rigidula) are probably due to paralled evolution. D.

rigidula probably branched off from the Drynaria line independently of the other

Drynaria species. In contrast to Copeland (1947) and Nayar (1965), this view appears

to support the earlier contention of Holttum (1954) who considered the origin of

Photinopteris as being more uncertain and pointed out that its origin might have been

through Drynaria, but in any case not immediately from Drynariopsis (Fig. 2)

In view of the two-ranked leaf arrangement characteristic of the Drynaria group, undis-

sected dorsal median vascular strand of the rhizomes (as in D. rigidula), and stipitate

pinnate fronds (as in D. rigidula), Photinopteris is included in the Drynaria group.

D. rigidula is quite different from all other species of Drynaria in having:(1) rhizomes

with undissected dorsal median vascular strands, (2) sclerenchyma strands in the rhizomes,

(3) stipitate pinnate fronds, (4) small stomata, (5) stellate foliar hairs, (6) prominently

impressed sori, and (7) small sporangia and spores. This possibly indicates that D. rigidula,

though similar to D. quercifolia in spore exine ornamentation, is probably nearer to it

and branched off from the Drynaria line independently of other Drynaria species (Chandra

& Zamora, 1979). Thus, D. rigidula represents a separate line of evolution from the same

common ancestor. This combination of unique features in D. rigidula is possibly sugges-

tive of the validity of its separation from Drynaria (it constitutes the section Poronema

J.Sm.).

Among the Aglaomorpha group, Drynariopsis is considered as the most primitive

genus. Thus, from D. heraclea, there have possibly evolved three genera of drynarioid ferns

naturally similar in many respects, but strikingly different in other aspects. Pseudo-

drynaria, though similar to Drynaria in unreduced fertile lamina and spinulose spores,

differs from the Drynaria group in such features as: (1) one-ranked leaf arrangement,

(2) broad humus-collecting leaf bases, (3) invaginated vascular cylinder of the rhizomes,

(4) basally attached paleae with non-glandular apices and dentate margins, (5) presence

of foliar hydathodes and.a hypodermis, (6) elongated irregular coenosori, and (7)

gametophytic trichomes. On the other hand, Pseudodrynaria is similar to the Aglaomorpha

< ic 4 4

% e % %

X oe t Y,

v 4

So Tai Y % ce L

YHd SOWOV IDV “Wovrn, % r

RN °

cow %

Fig. 1. A schematic representation of the inter-relationship of the drynarioid ferns (E. B. Copeland;

1929, 1947)

Fig. 2. A proposed schematic representation of the probable inter-relationship among the drynarioid

ferns.

234 Gard. Bull. Sing. 34:2 (1982)

group in having: (1) uniform fronds with broad humus-collecting bases, (2) one-ranked

leaf arrangement, (3) paleae with dentate margins, (4) foliar hydathodes and a hypo-

dermis, (5) similar size of sporangia and spores, and (6) hairy gametophytes. Pseudodry-

naria however, shows some advanced features over Drynariopsis viz. (1) tendency to form

coenosori, (2) high stomatal frequency, (3) non-glandular apices of the paleae, (4) non-

glandular paleal margins, and (5) spinulose spores. Thus, contrary to the observations

of Nayar (1965), it is here suggested that probably Pseudodrynaria is more allied to the

Aglaomorpha group and is posssibly derived from Drynariopsis in a separate line of

advancement (Fig. 2).

Another species presumed to have been derived from Drynariopsis is Merinthosorus

drynarioides, which is regarded as more specialized than Drynaria because of the differen-

tiation of its fronds into fertile and sterile portions. Based on gametophyte development,

Nayar (1965) pointed out that Merinthosorus is not very close to any other drynarioid

ferns, but its mature gametophyte is very much like that of Drynaria, Pseudodrynaria,

and Microsorium. Merinthosurs however, differs from Drynaria and Microsorium in

having: (1) one row of fronds on the rhizome, (2) basally attached paleae, (3) foliar

hypodermis and hydathodes, (4) smaller stomata, (5) dimorphic fronds, (6) verrucate

spores, and (7) small sporangia. Merinthosorus shows some similarities with Aglaomorpha

(A. meyeniana) in having: (1) ribbon-shaped, basally attached paleae with similar

dentations and without marginal glandular hairs, and (2) similar fertile fronds, spores, and

gametophyte trichomes. These similarities appear to support the contention of Copeland

(1947) that Merinthosorus is closely related to Aglaomorpha through. A. meyeniana (Fig.

1). The similarity of gametophyte development of Merinthosorus (not found in any other

drynarioid ferns) with that of Drynariopsis indicates that it is probably more intimately

related to Drynariopsis than to any other drynarioid ferns and is possibly derived directly

from Drynariopsis (Chandra, 1979a). Its similiarities with Aglaomorpha are probably due

to parallel evolution. The study of gametophyte development of Drynariopsis (Chandra

1979a) considerably strengthened the earlier view of Holttum (1954) who considers

Merinthosorus to have been derived from Drynariopsis (Fig. 2).

Still another genus thought to have sprung from an ancestor like Drynariopsis is

Aglaomorpha. Of the three species of Aglaomorpha, Copeland (1929, 1947) considered

A. meyeniana as the source of A. splendens which in turn gave rise to A. pilosa (Fig. 1).

A. splendens possesses comparatively primitive features such as: (1) indistinctly clathrate

paleae with glandular apices, (2) large foliar epidermal cells with deeply sinuous walls,

(3) large stomata, (4) low stomatal frequency, and (5) large sporangia and spores. A.

pilosa is, however, comparatively more advanced than A. splendens in having: (1) long,

wide-creeping rhizomes where each frond is associated with two branch buds, (2) high

stomatal frequency, (3) small stomata, sporangia, and spores, (4) further reduced fertile

fronds, and (5) localized distribution. Similarly, A. meyeniana also possesses some com-

paratively advanced features over A. splendens (i.e. absence of xylem parenchyma

between the tracheids of the meristeles in the rhizome, further reduction of the fertile

laminae, differentiated palisade, presence of upper and lower hypodermises, verrucate

spores, and restricted-~distribution). This suggests the probability that A. splendens

is the source of both A. meyeniana and A. pilosa and that A. meyeniana and A. pilosa

evolved independently of each other in a different line of evolution from A splendens

(Fig. 2). Thus, the earlier view of Copeland (1929, 1947) that A. meyeniana is the source

of A. splendens is refuted.

The genus Thayeria has presumably evolved from A. meyeniana (Figs. 1&2). Above the

cornucopia base, the frond and fertile apex of Thayeria are like those of A. meyeniana

to which it is most likely related (Copeland, 1929, 1947). Available information strongly

On drynarioid ferns

10.

12.

‘3.

14.

15.

235

TABLE 2. CHARACTERISTIC FEATURES OF THE TWO GROUPS OF

Attributes

Paleae structure

Frond arrangement

Nest leaves

Foliage leaves

Foliar hypodermis

Foliar epidermal cell

size

Stomatal size

Stomatal frequency

. Venation Pattern

Hydathodes

Nectary types

Leaf shape

Sori

Sporangial size

Spore size

DRYNARIOID FERNS

Drynaria Group

peltate, lancecolate or shield-shaped,

marginal glandular hairs with cap-

like waxy secretion

fronds in two alternating rows

on the dorsal surface of the

rhizome

present except in Photinopteris

stipitate

absent except in D. sparsisora

and Photinopteris

large

varies from 29 Ux 38 Lto 42

Uxs4u

varies from 38 to 79 per mm

finely reticulate, free included

veinlets found only in a few

areoles except in Photinopteris

absent on the lamina surface

except in Photinopteris

epicostular or hypocostular

fertile and sterile alike except

in Photinopteris

usually punctiform throughout

the frond except in Photinopteris

varies from 200U-400M x 2100p

-380

varies from 26-43 Ux 40-70

Aglaomorpha Group

peltate or basally attached,

lanceolate, shield-shaped,

or ribbon-like; Marginal

glandular hairs without

cap-like waxy secretions

fronds in one row on the

dorsal surface of the rhizome

absent as a rule

sessile except in Aglaomorpha pilosa

always present

smaller

varies from 27 Ux 32 Uto 34

x42u

varies from 59 to 140 per mm

broadly reticulate, free included

veinlets present in all the areoles

present as a rule

always hypocostular

fertile fronds more contracted

than sterile ones except in

Drynariopsis and Pseudodrynaria

coenosoral or acrostichoid, only at

at the upper portion of frond

except in Drynariopsis and

Pseudodrynaria

varies from 200-240 Mx 240-390 UW

varies from 26-36 Mx 42-56

236 Gard. Bull. Sing. 34:2 (1982)

indicates that Thayeria is the most advanced among the drynarioid ferns. Thus, dorsiven-

tral creeping, scandent habit, and supressed leaves on the dorsal surface of the rhizome

appear to support the above contention and have finally strengthened earlier conclusions

(Dickason, 1946; Holttum, 1964; Chandra, 1976) that dorsiventral creeping and a scandent

habit is biologically specialized and is more probably a derived one. Other morphological

attributes such as: (1) woody rhizomes, (2) paleae with closely spaced, broad, dentate

margins, (3) lamina with adaxial and abaxial hypodermises, (4) smallest paleae with non-

glandular apices, (5) smallest stomata, (6) highest stomatal frequency, (7) highly specilised

humus-forming habit, and (8) reduced fertile portion, also support the above contention.

CONCLUSIONS

During the course of this study I have come across evidence indicating reduction in

different parts of several species, i.e. stature, paleae, fronds, epidermal cells, epidermal

cell wall undulations, stomata, sporangia, and spores. Thus, it is suggested that one evolu-

tionary trend has been towards reduction. It appears that the Aglaomorpha group

represents the advanced group among the drynarioid ferns, in particular A. meyeniana,

Merinthosorus, and Thayeria (all of which possess thick adaxial foliar hypodermis and

nearly similar fertile laminae) represent the most advanced species among the Aglaomorpha

group (Chandra, 1979b).

From the foregoing facts it seems that these morphological attributes are of potential

value and may be useful for the assessment of relative specialization and relationships

among the drynarioid ferns.

On the basis of the accumulated evidence from comparative studies of the sporophytes

and gametophytes, it is concluded that the contrasting combination of distinct characters

among the drynarioid ferns supports the establishment of two natural, readily recognizable ,

and definable morphological groups namely Aglaomorpha and Drynaria (Table 2). It also

indicates that the drynarioid ferns are probably diphyletic in origin, i.e. one line represen-

ting the Drynaria group and the other line representing the Aglaomorpha group.

ACKNOWLEDGEMENT

I wish to thank Professor Dr. Prescillano M. Zamora of the Botany Department,

University of the Philippines, Diliman, Quezon City, Philippines, for providing constant

encouragement and helpful suggestions during the course of this study.

REFERENCES

Alston, A.H.G. 1956. The Subdivisions of the Polypodiaceae. Taxon 5, 23-25.

Bierhorst, D.W. 1971. The Morphology of Vascular Plants. Macmillan Co., New York.

Chandra, S. 1978. Morphology and Phylogeny of Drynarioid Ferns. Ph.D. thesis, University of the

Philippines, Diliman, Quezon City, Philippines (Unpublished).

1979a. Gametophyte Morphology of the Fern Genus Drynariopsis (Polypodiaceae).

Amer. Fern J. 69, 111-118.

1979b. Taxonomic use of Foliar Epidermis and Hypodermis in Drynarioid Ferns.

Kalikasan: Philipp. J. Biol. 8,211-226.

1980. A New Concept of Drynarioid Ferns. Ferns Gaz. (in press).

On drynarioid ferns 237

and P.M. Zamora, 1979. Spore Morphology in Relation to Taxonomy of Drynarioid

Ferns. Phytomorphology 29,252-255.

Ching, R.C. 1940. On the Natural Classification of the Polypodiaceae. Synyatsenia 5, 201-268.

1978. The Chinese Fern Families and Genera: Systematic Arrangement and Historial

Origin. Acta Phytotax. Sin. 16, 1-37.

Christensen, C. 1905-1906. Index Filicum and Suppls. II & III. (1917, 1934) H. Hagerup, Hafniae.

1938. Filicineae, In F. Verdoorn (ed.) Manual of Pteridology, Martinus Nijhoff, The

Hague. 512-550.

Copeland, E.B. 1929. The Oriental Genera of Polypodiaceae. Univ. Calif. Publ. Bot. 16, 45-128.

1947. Genera Filicum. Chronica Botanica; Waltham, Mass., U.S.A.

1960. Fern Flora of the Philippines 3, 377-555.

Crabbe, J.A., A.C. Jermy, and J.T. Mickel, 1975. A New Generic Sequence for the Pteridophyte

Herbarium. Fern Gaz. 11, 141-162.

Goebel, K. 1928. Morphologische and Biologische Studien XIII: Weiters Untersuchungen uber die

Gruppe der Drynariaceen. Ann. Jard. Bot. Buitenz. 39, 117-169.

Holttum, R.E. 1947. A Revised Classification of Leptosporangiate Ferns. J. Linn. Soc. Bot. 53,

123-158.

1954. Ferns of Malaya. Government Printing Office, Singapore.

Mehra, P.N. 1961. Cytological Evolution in Ferns with Particular Reference to Himalayan Ferns.

Proc. 48th Indian Sci. Congr. II Section Botany: 130-153.

Nayar, B.K. 1954. Studies in Polypodiaceae II: Contributions to the Morphology of Pseudodrynaria

coronans (Wall.) C. Chr. Phytomorphology 4, 379-390.

1955. The Gymnogrammoid and the Pleopeltioid Ferns: A summary. J. Univ. Gauhati

6, 47-62.

1959. Studies in Polypodiaceae VI. Further Observations on the Morphology of Drynaria

Bory. J. Univ. Gauhati 9, 95-103.

1965. The Gametophyte and Juvenile Leaves of the Drynarioid Ferns. Bot. Gaz. 126,

46-52.

1970. A Phylogenetic Classification of Homosporous Ferns. Taxon 19, 229-236.

1974. Classification of Homosporous Ferns II. In B.K. Nayar and S. Kaur (eds.) Com-

panion to R.H. Beddome’s Handbook of the Ferns of British India. Chronica Botanica, New

Delhi. 111-201.

and P. Kachroo, 1953; Studies in Polypodiaceae I: Contributions to the Morphology

of Drynaria Bory: D. quercifolia (L.) J. Sm. and D. propinqua (Wall.) J. Sm. Phytomorphology

3, 411423.

Pichi Sermolli, R.E.G. 1958. The Higher Taxa of the Ptcridophyta and their Classification. In O.

Hedberg (ed.), Systematics of Today Uppsala Univ., Arsskrift. 70-90.

1959. Pteridophyta. In W.B. Turrill (ed.), Vistas in Botany London; Pergamon Press.

241-493.

Tindale, M.D. 1961. Pteridophyta of South Eastern Australia: Polypodiaceac. Contr. Natl. Herb.

NSW Flora Ser. 210, 2446.

238 Gard. Bull. Sing. 34:2 (1982)

Wagner, W.H. Jr. 1973. Some Future Challenges of Fern Systematics and Phylogency. Bot. Jour.

Linn. Soc. 67 Suppl. 1, 245-256.

Zamora, P.M. 1975. Additional Notes on the Nectary-Costule Associations in Drynarioid Ferns. U.S.T.

J. Grad. Res., Philippines 4, 31-33.

and N.S. Vargas. 1973 a. Notable Variations in Leaf Forms of Drynaria. Philip. Argic.

57, 55-71.

1973 b. NectaryCostule Association in Philippine Drynarioid Ferns. Philip. Agric.

57, 72-88.

IN-VITRO POLLEN GERMINATION OF CASSIA FISTULA L.

JENNIFER NG

Botanic Gardens, Singapore

ABSTRACT

The germination of Cassia fistula L. pollen, ranging from fresh samples to

those stored for various periods, is reported. At the time of writing, the oldest

samples studied were stored for 4 weeks. It is intended to extend the observa-

tions for longer periods.

INTRODUCTION

Under the research programme at the Botanic Gardens, the author is involved in the

crossbreeding of ornamental plants. Among the various projects is an attempt to produce

new hybrids of Cassia. One of the species used is Cassia fistula, the Indian Laburnum, a

leguminous tree which has spectacular inflorescences. In connection with this research, it

is necessary to study the viability of the pollen, especially when it may have to be stored

for varying lengths of time to await the anthesis of flowers of other species.

MATERIALS AND METHOD

1. PREPARATION OF CULTURE MEDIUM

A sucrose-agar medium was made by dissolving agar strips (5 g), sucrose (100 g)

and boric acid (0.1 g) in 1 litre of distilled boiling water. The solution was poured into

petri dishes which were then packed in aluminium foil, autoclaved, and allowed to

set at ambient temperature.

2. COLLECTION AND STORAGE OF POLLEN

Cassia fistula has been observed to flower at least once a year, usually after a dry

spell. The whole crown is often covered with flowers at the beginning of the flowering

season, with sporadic flowering lasting up to 3 months.

Flowers were collected at anthesis, between 8 am and 10 am from 10-12 Nov 1981.

On each day , eight flowers were collected from the same tree. Pollen was taken from the

three anterior fertile stamens, distinguished by their long and curved filaments (Venkatesh,

1956). This decision was made after a series of viability tests of pollen from each of

the ten stamens showed that those taken from the three anterior stamens were viable,

whereas the other stamens yielded pollen which proved to be nonviable. Pollen to be

tested immediately was sown on agar for germination, the rest were stored in tiny pill

capsules. Silica gel was used to keep the vials containing the capsules moisture free.

The vials were then stored in a refrigerator at 4°C. Pollen was taken out subsequently

for testing: one week, two weeks, and four wecks after storage.

3. ASSESSMENT OF GERMINATION

Germination percentage was used to assess the viability of the pollen. Pollen grains

were dusted and smeared evenly on the agar using a fine brush and left to germinate

239

240 Gard. Bull. Sing. 34:2 (1982)

at room temperature (about 24°C). Although some pollen grains were observed to

germinate after half an hour, the assessment was not done until after 24 hours to ensure

maximum germination.

The germination percentage was derived in the following manner: For each of the

three pollen samples pertaining to a particular storage period, a viability test was done

on only one petri dish. Counting was done under a microscope fitted with a grid in the

eye-piece. A field was selected in which the pollen grains were not clustered together

to facilitate counting. An even distribution in the microscopic field selected for counting

generally produces lower variation between samples (Stanley, 1974). When such a field

was obtained, the whole field was systematically counted using a tally counter. It was

found that each field had between 100-150 pollen grains. A second count was made

on only the pollen grains with germinated pollen tubes. The total number of pollen grains

observed was then compared to the number with germinated pollen tubes.

RESULTS AND DISCUSSION

Fresh pollen showed the best viability. The average germination was 91.2%. Storing

the pollen for one week reduced the average germination by 2.4% (Table 1). The decline

was progressive after one week and by the fourth week the average germination was

66.9% (Fig 1).

100

PERCENTAGE GERMINATION

20 i STANDARD DEVIATION

10 2

1 3 3 4

STORAGE PERIODS (WEEKS)

FIG 1 PERCENTAGE GERMINATION OF CASSIA FISTULA POLLEN

STORED FOR VARIOUS PERIODS

i. Be

ad ee ait ith et ele I

Pollen germination, Cassia fistula 241

Stored pollen

Fresh pollen

So SE ee

Sample Date of collection

Number

germinated/ | % germ % germ

Total :

% germ

10 Nov 81 112/118 949 118/120 983 87/125 69.6 72/103

11 Nov81 102/127 803 93/109 85.3 88/125 704 72/107

12 Nov 81 130/132 985 120/145 82.7 95/118 805 75/118

MEAN

STANDARD DEVIATION (on—1)

912 88.8 73.5 66.9

96 83 6.1 32

Table 1. In-vitro germination percentage of fresh and stored pollen of Cassia fistula

It was observed that germination was very rapid. After 3 hours most of the pollen

grains had germinated. After 24 hours the pollen tubes had grown to an average length of

0.1 mm (Fig 2). The rapidity and high percentage of fresh pollen germination indicated

that the sucrose-agar medium used was suitable for in-vitro assays of Cassia fistula pollen.

The reasonable germination percentage obtained after four weeks of storage indicated

the capacity of the pollen to tolerate the storage conditions provided. The optimum

storage factors, like temperature, relative humidity and atmosphere surrounding the

pollen, would have to be determined for longer storage periods and better germination

potentials.

re ~~ , : ; >

< rie, ee as > Tt

S «Ae. « 3.4 a >

are”. gt Bt si

Fig. 2. In-vitro pollen germination of Cassia fistula on sucrose-agar (x 109). Note the long pollen tubes.

242 Gard. Bull. Sing. 34:2 (1982)

CONCLUSION

The storage conditions employed in this experiment were adequate in keeping the

pollen reasonably viable after 4 weeks of storage.

These findings indicate that even if the other species of Cassia were to be out of

phase in flowering by as much as | month and possibly longer, stored pollen of Cassia

fistula could still be used for crossbreeding work.

ACKNOWLEDGEMENTS

I wish to thank Miss Chang Kim Yam for her assistance, Mr Douglas Teow and Mr

Ong Tang Kwee for the photograph, and Mr Wong Yew Kwan for his comments.

REFERENCES

BULLOCK, R. M. and J. C. SNYDER (1946). Proc Wash. State Hort. Ass.

CORNER, E J H (1951). Wayside Trees of Malaya, Govt. Printing Office. Singapore, p. 386-388

STANLEY, R.G. and H.F., LINSKENS, H.F. (1974). Pollen: biology, biochemistry and management,

Berlin, Springer. P. 75-76

VENKATESH, C.S. (1956) Phytomorph 6. p. 168-175

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Vol. XXXIV

Basionyms and synonyms appear in italics. Page numbers in italics indicate substantial

treatment of the taxa or the presence of illustrations. Names on chart 2 on page 167 have

not been indexed.

Aceratum insulare 216

Acronychia 93

Acrostichum semipinnatum 138, 139

x Aeridachnis Alexandra 152

Bogor ‘Apple Blossum’ 152

Elizabeth Howie 152

Aglaomorpha 230, 231, 232, 234

heraclea 171, 174

meyeniana 231, 234, 236

pilosa 231, 234

splendens 231, 234

Alphitonia incana 215

moluccana 215

zizyphoides 215

Ampacus daniellii 105

meliaefolia 109

rutiecarpa 116

trichotoma 114

Amyema rigidiflora 211

aff. sogerensis 211, 212

Anodendron paniculatum 220

Aporosa papuana 214

Apteuxis 2,4

trinervis 2

x Arachnis Capama ‘Merah’ 152

hookerana 152, 158

species 151

x Arachnostylis Chorchalood 152

x Aranda 150, 151

Belzonica 152

Bertha Braga 152

Bintang 152

Chen Chin Mu 152

Chong Chee Yuen 148, 152

Christine no. 1: 151

nos S:/451.'152

nos 9, 27, 80: 152

nos 130, 999: 153

Elizabeth Douglas Home 153

Eric Mekie 153

Freckles 153, 156

Hatifah 153

Hilda Galistan 153

Imelda Romualdez Marcos 153

Jessie 153

Lucy Laycock 153

Majula ‘Rimau’ 151, 153

Ng Chong Hang 148, 153

Noorah Alsagoff 153

Peter Ewart 153

Queen of Purples 153

Tourism Singapura 153

Wee Huck Lay 153

Wendy Scott ‘Blue Bird’ 153

‘Green Field’ 153, 156

Yvonne Tan 153

x Aranthera Anne Black 153

Beatrice Ng ‘Yellow’ 153

Dainty 153

Francina Bruyns 154

Gracia Lewis 154

James Storie 154

x Ascocenda Heath Hock Heng 154

Bot. Gard. ser. nos. 4038, 4766: 154

Aspidium alatum Ridl. 140

alatum Wall. ex Hook. & Grev. 140, 141

amplifolium 143

angulatum 145

brachiatum 137

cardieri 136

coadunatum 137

decurrens 139

devexum 136

giganteum var. minor 136

griffithii 137

hankei 142

emersum 137

intermedium 136

keckii 143

kuhnii 138

kunstleri 138

longicrure 144

maingayi 139

malayense 138

melanocaulum 143

membranaceum 136

meniscticarpum 142

microsorum 143

multicaudatum 138

oligophyllum 145, 146

pachyphyllum 141

papyraceum 142

pentaphyllum 145

pinfaense 137

oligocarpum 138

pteropus 139

quinquefoliatum 145

repandum 141

ridleyanum 140

sagenioides 136

semibipinnatum 143

siifolium 142

var. teysmannianum 142

simonsii 144

singaporeanum 138

subdecurrens 139

243

744 Gard. Bull. Sing. 34

(Aspidium cont.)

subtriphyllum var. ebenosum 144 ; x Jaquelyn 148

terminale 145 laciniosum 152

ternifolium 142 x Ng Eng Cheow 155

teysmannianum 142 x Sunny 155

tricuspe 140 x superbiens ‘Superba’ 152, 158

trifolium 145 x Tay Swee Keng 148, 155

variolosum 137 x Yong Kok Wah 148, 155, 159

vastum 140 x Bot. Gard. ser. nos 4879, 9215: 155, 159

zollingerianum 137 x Bot. Gard. ser. nos 8090, 8262, 8324,

Asplenium affinis 173 8961, 129172 435

Astromia 8 Desmodium umbellatum 218

Belliolum haplopus 212 Dicranopteris curranii 172

Blechnum orientale 172 linearis var. linearis 172

Blumea riparia 221 var. montana 172

Boerlagiodendron 209 Dictyopteris baberi 138

Boymia 91, 92, 94 heterosora 139

glabrifolia 109, 113 pentaphylla 145

martinicensis 122 polycarpa 138

ruticarpa 94,97, 116, 118, 119 Dillenia crenata 216

Brucea 93 Diplazium esculentum 171, 174

javanica 122 malaccense 173

trichotoma 114 speciosum 171, 176

x Burkillaria Ong Thye Chiew 154 subintegrum 173

Calophyllum inophyllum 216 tomentosum 173

paludosum 216 Diospyros 79

Calymmodon 174 Dipteris conjugata 172, 176

gracilis 173 x Doritaenopsis Elizabeth Waldheim 155

hirtella var. coriacea 174 Bot. Gard. ser. PDI code 1: 155

Campylogramma trollii 139 Drynaria 229 et seq.

Cardiochlaena alata 139 descensa 231

Carruthersia mollis 220 fortunei 230

Cassia fistula 239 et seq. laurentii 230

Casuarina nodiflora 216 menisclicarpa 142 :

Cattleya x Meadii 154 propinqua 230

x Bot. Gard. ser. code C03: 154 quercifolia 230, 231, 232

Celtis kajewski 210 rigidula 171, 174, 230, 232 :

Chingia perrigida 173, 174 sparsisora 231

pseudoferox 173 Drynariopsis 230, 231, 232, 234

Cinnamomum 5 heraclea 232

Chisocheton sp. 214 Dryomenis menisciicarpa 142

Christella arida 170, 175 phymatodes 142

Corynocarpus cribbeanus 215 Dryopteris raciborskii 136

Coryphopteris gymnopoda 172, 1/76 Dysoxylum caulostachyum 214

var. bintangensis 172 Elaphoglossum callifolium 173

Crawfurdia trinervis 171 robinsonii 174

Crypsinus laciniatus 172 Equisetum debile 170, 175

trilobus 173 Eukibessia 3

Ctenitopsis matthewii 136 Euodia 91, 92

setulosa 135 ailantifolia 109, 113,122

Curculigo latifolia 172, 175, 203 et seq. austrosinensis 120

Cyathea contaminans 172, 176 tee balansae 113

Cymbidium 149 bodinieri 120

Dawsonia superba 181 calcicola 109

Decaspermum fruticosum 218 compacta 120

Deeringia amaranthoides 212 var. meionocarpa 120

arborescens 212 colorata 116

Dendrobium daniellii 98, 106, 108

x Alice Spalding 154 fargesii 110

antennatum 148, 152, 758 fraxinifolia 98

x Barbara Moore 154 glauca 97,110

x Caesar 154 hainanensis 116

x Cheong Chee Yon 154, 159 hirsutifolia 120

x Ellen Harris 154 hupehensis 98

x Fiery Gold 154 impellucida 104

Index

(Euodia cont.)

var. macrococca 104

labordei 108

lenticellata 116

meliifolia 97,100, 113, 114

officinalis 97,98, 120

var. officinalis 117

sambucifolia 101

sambucina 101

subtrigonosperma 95, 102

sutchuenensis 105, 108

245

poilanei 104 tonkinensis 122

robusta 104 trichotoma 94, 114

rugosa 120 var. pubescens 114

ruticarpa 97, 98, 118, 120 velutina 105

subtrigonosperma 97, 104 vestita 105

sutchuenensis 106 viridans 114

trichotoma 97,116 yunnanensis 110, 114

velutina 98 Eywickia 2, 4

vestita 106 capitellata 26

viridans 116 cordata 23,24

yunnanensis 114 cyanea 2, 26, 29, 30, 31

Eurycoma dubia 109, 113 echinata 42

Euscaphis japonica 122 galeata 2,49

Evodia jackiana 26

Sect. Evodioceras 95, 97

Sect. Oxyactis 95

Sect. Subtrigonospermum 95, 97

Sect. Tetradium 94

ailantifolia 109, 122

austrosinensis 120

baberi 105,117

balansae 109

bodinieri 116

calcicola 108

chaffanjoni 122

colorata 114

compacta 117

forma meionocarpa 117

daniellii 95, 97, 105

var. delavayi 105

var. henryi 106

var. hupehensis 106

var. labordei 106

var. villicarpa 106

delavayi 105

fargesii 109, 113

fraxinifolia 102

glauca 109

hainanensis 114

henryi 105, 108

var. villicarpa 105, 108

hirsutifolia 117

hupehensis 105, 108

impellucida 102

vat. macrococca 102

labordei 105

lenticellata 114

meliaefolia 109

var. celebica 122

mollicoma 122

odorata 122

officinalis 117

parviflora 122

poilanei 102

robusta 102

rugosa 117

korthalsiana 24, 31, 73, 74

latifolia 26, 29

var. membranacea 26, 29, 31

medinilliformis 26

paniculata 26, 30, 79

tuberculata 73

Fagara 98

martinicensis 122

megistophylla 213

Fagraea racemosa 221

Ficus 221

dissipata 210

erinobotrya var. solomonensis 210

indigofera 210

verticillaris 210

Finschia cloroxantha 211

densiflora 211

Flagellaria subg. Chortodes 225

elegans 225

indica 221

plicata 223, 224, 225

Freycinetia 209

Fuirena umbellata 222

Gastonia spectabilis 220

Geniostoma brassii 221

Gleichenia hirta var. amoena 171

longissima 172, 176

truncata var. plumaeformis 172

vulcanica 172, 176

Gnetum gnemon 216

Goniophlebium subauriculatum 171, 174

Gonus amarissimus 122

Grammitis 174

hirtella var. coriacea 173

Grevillea elaeocarpifolia 211

Gymnogramma maingayi 139

Helica micronesica 211

Hemionetis maingayi 139

Heterospathe woodfordiana 222

Hibiscus 221

Histiopteris incisa 172

x Holttumaria Cochineal 155

rutaecarpa 116 Horikawaea phyllogonoides 180, 181

var. bodinieri 117 Hymenophyllum denticulatum 173

forma meionocarpa 117 polyaxnthus 173

246

Hypolepis beddomei 172

brooksiae 171, 174

punctulata 172

Joinvillea ascendens 223, 224, 225

ssp. ascendens 225

spp. borneensis 225

spp. glabra 225

ssp. samoensis 225

borneensis 225

bryanii 225

elegans Gaud. 223, 224, 225

elegans Gaud. ex Brongn. & Gris 225

ssp. bryanii 225

gaudichaudiana sensu Christ. 225

var. samoensis 225

malayana 225

plicata 223, 224, 225

ssp. bryani 225

ssp. plicata 224, 225

x Kagawara Teoline Fair 148, 155

Kibessia 1,4, 17

acuminata 2,42, 44, 70

acuminatissima 15, 17

affinis 62, 64, 65

angustifolia 9, 13, 30, 43, 45, 80

azuria 15,44

var. subalata 15,17

var. typica 15

celebica 80

cordata 18, 19, 23, 24

coriacea 37

cupularis 43, 45

echinata 42

elmeri 23, 52, 54

galeata 49

var. latifolia 80

var. pluriflora 49, 50

gracilis 54

hirtella 58

korthalsiana 3, 23, 24, 54

ledermannia 80

pachyphyllum 69, 70

pubescens 2, 42

rostrata 62

simplex Korth. 15, 16, 17, 19, 44, 69, 79

var. oblonga 15, 17, 30, 79

simplex auct. non Korth. 2, 43

subalata 15

tessellata 67

tetraptera 18, 23

teysmanniana 69, 70

var. grandifolia 63, 64, 65

tuberculata 2, 30, 73

verrucosa 62, 64, 65

Leea suaveolens 215

tetramera 215

Leptochilus rimicifolius 145, 146

Lestibudesia arborescens 212

Litsea perglabra 213

solomonensis 213

“timoriana” 213

Macaranga 221

aleuritoides 214

polyadenia 214

Gard. Bull. Sing. 34

Macroplacis 2, 3, 4

Sect. Macroplacis 3, 54

cordata 2, 23

Maesa 79

Matonia pectinata 172, 176

Medinilla 3, 80

muricata 80

myrtiformis 80

Megabotrya 91, 92, 94

meliaefolia 94, 109, 113

Melastoma azureum 2, 15

malabathricum 218

Melastomataceae

Tribe Dissochaeteae 7

Tribe Oxysporeae 7

Meliocope 91, 92, 93

ternata 92

Memecylon 5, 8

oligoneuron vat. maluense 26, 31

Merinthosorus 230, 234, 236

drynarioides 234

Microlepia 171

spp. prev. unidentif. 171

speluncae 171

var. hancei 171, 174

strigosa 171, 174

todayensis 171, 174

Micromelum minutum 213

x Mokara Sally Lim 148, 155

Myristica fatua var. papuana 212

Myrsinaceae 209

Neckera 180

Nephrodium griffithii 137

ingens 135

multicaudatum 138

nebulosum 145

nudum 143

setulosum 135

Zollingerianum 137

Nephrolepis tuberosa 170

Oedicladium 181

Oleandra pistillaris 173

Oncidium

x Goldiana 155

haematochilum 152

hybrids of 151

x Josephine 155

lanceanum 152, 158

Orthorrynchium 181

elegans 180, 181, 183

philippinense 180

phyllogonioides 180

Palisadula 181

Pandanus 209

Sect. Bidens 177

Sect. Rykia & Subsect. Rykia 177

burmanicus sp. nov. 177,118, 119

piniformis 177

Parinarium glaberrimum 216

Peekeliopanax spectabilis 220

Phaeanthus macropodus var. mollifolius 212

Phalaenopsis, Bot. Gard. ser. no. 3596: 156

Phegopteris baberi 138

Index 247

(Phegopteris cont.)

polycarpa 138 var. capitellata 26, 28, 29, 30

subdecurrens 139 var. cyanea 26, 29, 31, 74

Phellodendron 93, 94, 98 var. jackiana 26, 28

burkillii 110, 112, 114 var. membranacea 31

Philagonia 91, 92, 94 var. paniculata 26, 28, 30

fraxinifolia 102, 104 cogniauxii 3, 5-8, 23, 25, 50

procera 100 cordata 18

sambucifolia 101 coriacea 5-8, 37, 38, 39, 74, 80

sambucina 94, 100, 101 crassicalyx sp. nov. 5-8, 40, 41

Phlebigonium impressum 137 discolor 80 =

Photinopteris 230, 231, 232 echinata 2, 5-9, 13, 16, 17, 30, 42, 47, 48, 69,

speciosa 230 70, 76, 80

Phyllanthus sp. 211 var. bracteata 42, 45

Phymatopteris lacinata 172 var. pubescens 8, 42, 45

triloba 173 elmeri 52, 54

Pimeleodendron ambonicum 172, 214 forbesti 49, 50

Pityrogramma calomelanos 171 galeata var. galeata 5-8, 24, 27, 49, 51, 64, 70,

Plagiogyria tuberculata 171, 176 76, 81

Planchonella obovoidea 221 var. elmeri stat. nov.5-8, 24, 52, 53

Pleocnemia membranacea 136 gracilis 6-8, 54, 56, 57

Pleopeltis grandidentata 139 grandifolia 63

Plerandra solomonensis 218, 219 griffithii 3, 24, 31, 65, 69, 73, 74, 78, 79

Pogonanthera pulverulenta 80 var. cordata 73,74

Pollia thyrsiflora 222 hirtella, ‘birtella’ 3, 5-8, 19, 42, 58, 59

Polypodium jackiana 26

angulatum 145 Korthalsiana 23, 24, 73

baberi 138 latifolia 26

dilatatum vat. grandidentatum 139 ledermannii 3, 80, 81

grandidentatum 139 moluccana 3, 80

heterosorum 139 multiflora 6-8, 26, 27, 60, 61

polycarpum 138 pachyphylla 69

semipinnatum 139 paniculata Benth. ex Cl. 26, 31, 74

stifolium 142 paniculata Benth. ex Ridl. 73

Polyscias 209 paniculata (Miq.) Triana 26, 31, 79

Poronemia 232 parasitica 80

Pseudodrynaria 230, 232; 234 rostrata 3, 6-8, iy, at; 42, 62, 66, 69, 73;

coronans 230 76, 79-81

Pseudophegopteris rectangulare 171, 174 simplex 15-17, 19, 80

Pteridium aquilinum var. wightianum 171, 172, 175 tessellata Ohwi 67

Pteris tripartita 171 tessellata (Stapf) Nayar 6-8, 17, 19, 23, 67, 68, 80

vittata 170, 171 teysmanniana 6-8, 37,69, 71, 72

Pternandra 1, 4, 17, 76, 79 tuberculata (Korth.) Maxw. 73

Tribe Dissochateae 7 tuberculata (Korth.) Nayar 2, 5, 7, 8, 23, 24, 30, 31,

Tribe Oxysporeae 7 37, 42, 44, 55, 63, 65, 69, 73, 75, 77, 78, 79, 81

geographical distribution 12 verrucosa 62, 64

key to species 10 viridula 54

relative abundance 8 Pterocarpus indicus 171 et seq.

acuminata 43 Pyrrosia floccigera 23, 171, 172, 174

acuminatissima 15 Rectromitra Sect. 3

affinis 62, 64 Rectomitra 2, 3, 4

angustifolia sp. nov. 6-8, 12, 14 cordata 23

azuria var. azuria 3, 5-8, 15, 16-20, 44, 60, galeata 49

76, 79, 80 tuberculata 73

var. cordata stat. nov. 6-8, 16, 18, 21, 22 x Renanopsis 156

23, 24, 60, 69 Rhamnus incana 215

var. jackiana 26 zizyphoides 215

var. membranacea 26 Rhodamnia 5

var. paniculata 26 dumetorum 80

var. subalata 15 Rhus 104

bornaeansis 15, 17 fraxinifolium 104, 107

capitellata 2, 26, 31,74 taitensis 214

cardiophylla 23 Rubus moluccanus 171

coerulescens 26, 32, 36, 44, 45, 50, 54, 62, Rutaceae 91

248

(Sagenia cont.)

decurrens 139

gigantea var. minor 136

griffithii 137

longicruris 144

macrodonta 137

melanocaulis 143

pachyphylla 141

pteropus 139

Sararanga 209

Scirpus fuirena 222

Scleria lithosperma 222

Scleroglossum 174

debile 173, 174

Selliguea maingayi 139

Sphaerostephanos polycarpus 170, 171

Sphenomeris chinensis var. divaricata 172

Stemonurus sp. 215

Streblus glaber 211

solomonensis 211

Strychnos 5

Styrax agreste 220

Tapeinosperma 209

Tectaria

key to Malayan species 133

amplifolia 133, 142, 143

angulata 145

barberi 133, 138

brachiata 137, 142

cardieri 136

celemensiae 140

cherasica sp. nov. 132, 141

christii 137

coadunata 137, 141, 142

crenata /4]

cuneifolia 145

decurrens 132, 139

devexa 133, 136

dimorpha 141, 146

fauriei 132, 146, 147

folium 145

fuscipes 146

grandidentata 139

griffithii 137

gymnosora 146

haenkei 142

herpetocaulos 132, 145

ingens 135.

keckii 143

macrodonta 137

maingayi 139

malayensis 138

matthewii 136

melanocaula 132, 1/43

modesta 143

multicaudata 138

oligophylla 145, 147

pandurifolia 138

paradoxa 136

polymorpha 145, 146,

var. cuneifolia 145, 146

ridleyana 140

rumicifolia 145, 146

Gard. Bull. Sing. 34

semibipinnata ]43

semipinnata 138

siifolia 142

simonsii 132, 144

singaporeana 1/38

subtriphylla var. ebenosa 144

terminalis 145

ternifolia 142

tricuspis 132, 140

variolosa 137

vasta (Bl.) Copeland 140, 146, 147

vasta non (Bl.) Copeland 146, 147

Tetradium 91-97

index to collections 124

keys to species 98-100

amarissimum 122

austrosinense comb. nov. 95, 97, 120, 122

calcicolum comb. nov. 96, 97, 108

daniellii comb. nov. 96, 97, 98, 105, 108, 117

fraxinofolium comb.nov. 95-98, 102, 104, 122

glabrifolium comb. nov. 96, 97, 98, 109, 113, 114,

ruticarpum comb. nov. 96, 97, 98, 105, 108, 116, 1

sambucinum comb. nov. 96,97 100

sumatranum sp. nov. 91,92, 95,97, 121, 122

trichotomum 91, 94, 96, 97, 114, 116

Thayeria 230, 234, 236

cornucopia 230

Toddalia 94, 98

Trichomanes digitatum 173, 174

idoneum 173

maximum 173

proliferum 174

saxifragoides 174

Trichospermum ikutai 216

kajewski 216

psilocladum 216, 217

Turpinia parviflora 122

Vanda 150

x Chia Kay Heng 156, 157

hookerana 184

hybrids of: 150

x Miss Joaquim 184-186

x Norbert Alphonso 156

x Patricia Low 156

x Poepoe ‘Diana’ 156

x Ruby Prince 156

x Sanada Kuma 156

teres 184

x Bot. Gard. ser. no. 3102: 156

Vateria indica 163, 168

Wormia crenata 216

Xanthoxylum

subgen. Oxyactis 94

daniellii 94, 105

Zanthoxylum 93, 94

bretschneideri 105, 108

daniellii 108

martinicense 122

megistophyllum 213

molle 122

odoratum 122

thetsoides 122