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THE

GARDENS’ BULLETIN

STRAITS SETTLEMENTS

Volume IX
December, 1935-——-March, 1938

To be purchased at the Botanic Gardens, Singapore

Published by Authority

SINGAPORE :

PRINTED AT THE GOVERNMENT PRINTING Office, SINGAPORE.
sy W. T. Cuerry, GoveRNMENT PRINTER.

1938

|
.

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WWAjyps

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ak i . 2 ¥ » oe 7 a

CONTENTS

PAGE
Part 1: 20th December, 1935 (pp. 1-117).
Dedication of the Bulletin to Mr. H. N. RIDLEY
(with one plate) i 1

HENDERSON, M. R. & C. G. G. J. ae ae
Bibliography of the works of H. N. RIDLEY,

from 1872-1935 1 2
ANONYMOUS: A List of plants boned in CONGUE

of Mr. H. N. RIDLEY .. 29
SouTH, F. W.: Mr. RIDLEY’s work a tropical

agriculture 31
EATON, B. J.:, Mr. RILEY and Raikes Phy eae

ANONYMOUS: Mr. RIDLEY and Forestry in Malaya 42
SLOOTEN, D. F. VAN: Mr. H. N. RIDLEY and the

flora of the Netherlands Indies .. 44
MERRILL, E. D.: Some oo phytogeographi-

cal problems 49
DOCTERS VAN LEEUWEN, W. M.: The ae ae of

plants by fruit-eating bats ie 58

STEENIS, C. G. G. J. VAN: Open-air Hates i in
the tropics at 3100 metres altitude ey ith piste

1) 64
BACKER, C. ne t pani ciaaimeninn Ph as vatantar
preted genus of Balsaminaceze (with plate 2) 70

HOLTTUM, R. E.: The flowering of Tembusu trees
(Fagraea fragrans ROXB.) in sie aoe 1928—

1935 a 73
CORNER, E. J. H.: The: eee frultive of
Agarics in Malaya sah 19
SMITH, J. J.: A few orchids from ahs Malayan

Archipelago 89
HENDERSON, M. R.: The enpnaic flora of

Dipterocarpus oblongifolius Bl. (‘““Neram.’’) .. 93
LAM, H. J.: Phylogeny of single features

(Madhuca Ridleyi n. sp.) (with plates 3-7) . 98
FURTADO, C. X.: Liste Li ae ylla oH

allied species 113

Part 2: 4th May, 1937 me 119- oe,

HouLtTtum, R. E.: Notes on Malayan ferns, with
descriptions of five new species .. gar Rho

HOLTTUM, R. E.: Further notes on Stenochlaena,
Lomariopsis and Teratophyllum ..

CROIZAT, LEON: An early tertiary relict in Malsiver
Euphorbia Ridleyi

FURTADO, C. X.: Palme Malesicz VI (with ares

FURTADO, C. X.: Palme Malesice VII

STEENIS, C. G. G. J. VAN: On the application of
the terms Bien and Malaya in plant-
geography

HoLTTuUM, R. E.: The cn Lomagramna (with
plates 8-16)

Part 3: 9th October, 1937 (pp. 223-317).

FURTADO, C. X.: A Commentary on the laws of
Botanical Nomenclature (with Index)

FURTADO, C. X.: The Nomenclature of SS i
(with Index) iz

FURTADO, C. X.: Asterisks in Linea S Species
Plantarum
Part 4: 31st March, 1938 (pp. Headey

SYMINGTON, C. F.: Notes on Malayan Dipterocar-
pacee IV (with plates 17-27 and Index)

HOLTTUM, R. E.: A redefinition of the genus
Teratophyllum (with plates 28-30) be

( IARDE Ns’ BULLETIN

"STRAITS: SETTLEMENTS

“Vol. ° 20th December, 1935 2 Part

- CONTENTS»

eee

Peete eres ins ~' - i
vi Bibliography of the eeeba of Mr. ns Si N. Ridley. joe 1872 to 1935,
= compiled by M.:R, Henderson and.C'G. G. J. van Steenis- 2
) List of plants named in honour of Mr. H. N: Ridley sit - 29

SiMe, Ridley’s Work for Tropical Agriculture, sy F. W. penth : 31
ja Mr.-Ridley and Rubber, by B. J. Eaton - = - 89
« ‘Mr. Ridley and Forestry in Malaya - 42

|. Mr. Ridley and. the Flora of dic, Neiheranas Indies, bg D.. Fy; van.

cir): OOterE.) 8 S.2 44
Make Malaysian Phy ionebieniice! Helene by E D. Merrill - 49
/) The Dispersal of pias by: Peete. bats, . bx 1A M. Docters van
PA Beeuwen - 58
Open-air hothouses in’ the tropics at 8, 190 metres litude, by. C G.
. G.d. van Steenis Ga
- Semeiocardion Zoll., a ‘misinterpreted genus -of Palaaihacdu by Cc ;
A. Backer. = ey at
|The flowering. of Tembusu tees: (Fagroea frazrons Roxb: ) itt Singa;
pore, 19278-1985, by R: E. Holttum - 73
\ The seasonal ‘fruiting of Agarics in Malaya, by E. iz HAT, ghee - 79
Ny few Orchids from the Malayan Archipelago, by J J. Smith - - 89
The epiphytic flora of oh ile eet AG te Bl., ey M.
Wu Ffenderson .. =' ; 93
Ae he PR loneny of Single Features, es ETGARE: Fee - - 98

Oa ig ky as and shite’ species, Oy C X. Poresas = AIS

Te be purchased at the- Botanic Blons Singapore
Bras ) | Price $3.0 Og or att:

Di
Le

ARR Nah: ‘SINGAPORE:
Praxteb Ar THE: EEE REARS Printing Orrice, SINGAPORE,
eal i Lae gee pais ere PRINTER.

——

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Set

Ae

H. N. Raptey, C.M.G., F. B.S.

SHE

GARDENS’ BULLETIN

STRAITS SETTLEMENTS

Vol. IX 20th December, 1935.

Part 1

This Bulletin is dedicated to

HENRY NICHOLAS RIDLEY, C.M.G., M.A., F.R.S.

Director of Gardens, Straits Settlements, 1888-1911
on the occasion of his 80th birthday

Mr. Ridley’s work as a field botanist added more than
that of any other single man to our knowledge of the flora
of the Malay Peninsula; his Flora is the foundation upon
which all future work must rest; he actively pursued the
experimental cultivation of agricultural plants at a time
when agricultural interests in Malaya were at a low ebb;
in particular, his persistent advocacy of the plantation of
Para Rubber and his pioneer tapping experiments were
largely responsible for the foundation of that industry upon
which so large a share of the wealth of Malaya has rested ;
his reports on timbers, rattans, and other forest products
of the Peninsula were the first adequate works of their
kind and embodied much original information; he added
very considerably to the collection of cultivated plants, both
Malayan and exotic, in the Botanic Gardens, Singapore;
his wide interests, not only in botany but also in entomology,
geology and other branches of natural science, found
expression in a series of papers which are enumerated in
the bibliography which follows. Few men have accom-
plished so much in 23 years of tropical service, and few
have been able to complete their work after retirement as
Mr. Ridley has completed it.

The contributors to this Bulletin, and many more who
have followed in the paths where Mr. Ridley led, unite in
congratulating him, on the present happy occasion.

i

N

i
BIBLIOGRAPHY OF THE WORKS OF
MR. H. N. RIDLEY, FROM 1872 TO 1935,
compiled by
M. R. HENDERSON and C. G. G. J. VAN STEENIS

This bibliography has been made as complete as
possible, but as Mr. Ridley has published, over a long
period, a very large amount of material covering a very
wide field of pure and applied botany, the compilers realise
that a number of papers may have been overlooked.

During 63 years of activity Mr. Ridley has published,
according to this bibliography, more than 500 books, papers
and notes, amounting to nearly 10,000 printed pages.

Amongst the more important works may be mentioned
his various papers on Monocotyledons, on which he was a
well known authority; and his various articles on plant
dispersal, resulting in his monumental “Dispersal of. Plants
throughout the World’, published when he was 75, an age
at which most authors are content to enjoy a well earned
rest. His life-work, however, has been devoted to
Malaysian botany, to the knowledge of which he has
contributed largely, both as a writer and as a keen
collector.

1872

Mammals and Coleoptera of Haileybury (School Nat. Hist.
Report, 1872).

1875
Sphinx convolvuli_ at Cobham, Kent (Entomologist, 1875, p.
22a
1876
Hibernation of Hemerobius (Entomologist, 1876, p. 48).
1878

Coleoptera hunting in 1877 (Entomologist, 1878, p. 22).
Acronycta Alni in Hereford (Entomologist, 1878, p. 230).

1879
Doridicola anthez, n. sp. (Copepoda) (Ann. Mag. Nat. Hist. 4,
1879, Pp. 458).
1880

Spring flowering form of Colchicum autumnale (Journ. Bot. 18,
1880, p. 185).

Observations on some Larve of piaenne (Entomologist, 13,
£380, pit2T).

Some English Collembola (Entomologists’ Monthly Magazine,
17, , 1880) Spe I).

Gardens Bulletin, S.S.

8

1881

A new variety of Carex pilulifera L. (Journ. Bot. 19. 1881,
DP: 97-93) 152, .f/ 278),.

Notes on Radnorshire plants (Journ. Bot. 19, 1881, p. 170-174).

Degeeria Eatoni, n. sp. (Entomologists’ Monthly Magazine, 17,
TOS]; Pp. 270):

Thysanura of Canary Isles (Entomologists’ Monthly Magazine,
EO, LGOR Pe Ay, |

; 1882

(With Fawcertr) Additions to the flora of Dorset (Journ. Bot.
20, 1882, p. 246):

1883

Teratological notes on plants I. (Journ. Linn. Soc. London,
Bot. 20, 1883, p. 45-48, 3 figs., read 1882).

Descriptions and notes on new or rare monocotyledonous plants
from Madagascar, with one from Angola (Journ. Linn. Soc.
London, Bot. 20, 1883, p. 329-338).

On the generic names Dantia and Prouvenzalia (Journ. Bot. 21,
1883, p. 349).

1884

The Cyperaceae of the West Coast of Africa in the Welwitsch
Herbarium (Transact. Linn. Soc. London, Bot. 2nd ser.,
vol. 2, 1884, p. 121-172, pl. 22-23).

Cyperacez nove (Journ. Bot. 22, 1884, p. 15-17).

Kerry plants (Journ. Bot. 22, 1884, p. g1-92).

Shetland plants (Journ. Bot. 22, 1884, p. 301).

Bees and Erica cinerea (Journ. Bot. 22, 1884, p. 302).

A new Bornean Orchid (Journ. Bot. 22, 1884, p. 333).

On Didymoplexis sylvatica (Leucorchis-sylvatica Blume) (Journ.
Bot. 22, 1884, p. 345-346).

A new species of Albuca from Aden (Journ. Bot. 22, 1884, p. 370).

Additions to ‘“Topographical Botany’? (Journ. Bot. 22, 1884,
DP. 377-378).

1885

The Orchids of Madagascar (Journ. Linn. Svc. London, Bot. 21,
1885, p. 456-522, pl. 15).

Prodromus florze timorensis; compiled by the Botanical Depart-
ment of the British Museum, Monocotyledones (in H. O.
Forees, A Naturalist’s Wanderings, etc. 1885, p. 518-523).

On Juncus tenuis as a British plant (Journ. Bot. 23, 1885, p. 1-3,

eo 253)
A new Carex from Sumatra (Journ. Bot. 23, 1885, p. 35-36).

Vet, HX.) 61935).

4

A new Dendrobium from Siam (Journ. Bot. 23, 1885, p. 123).

A new Habenaria from Brazil (Journ. Bot. 23, 1885, p. 170).

Crocus Korolokawi in Afghanistan (Journ. Bot. 23, 1885, p. 185).

Peloria in Habenaria bifolia Br. (Journ Bot. 23, 1885, p. 218-2109).

Castanea sativa Mill. as a native of Britain (Journ. Bot. 23, 1885,
Pp. 253-254).

Two new British plants (Journ. Bot. 23, 1885, p. 289-291,
t. 261-262).

On a new species of Gussonea (Journ. Bot. 23, 1885, p. 310).
Notes on British Rubi (Journ. Bot. 23, 1885, p. 370).

Angroecum glomeratum, n. sp. (Gardeners Chronicle, 24, 1885,
p. 678).

1886

On the Monocotyledonous plants of New Guinea, collected by
H. O. Forbes (Journ. Bot. 24, 1886, p. 321-327, 353-360,
t. 270-271).

Notes on Orchids of tropical Africa (Journ. Bot. 25, 1886,
Pp. 291-2096).

A monograph of the genus Liparis (Journ. Linn. Soc. London,
Bot. 22, 1886, p. 244-297).

On Dr. Fox’s collection of orchids from Madagascar, along with
some obtained by the Rev. R. Baron, F.L.S. from the same
island (Journ. Linn Soc. London, Bot. 22, 1886, p. 116-127).

On the freshwater Hydrocharidee of Africa and its islands (Journ.
Linn. Soc. London, Bot. 22, 1886, p. 232-241, pl. 13-14).

Habenaria albida in Brecon (Journ. Bot. 24, 1886, p. 285). °

A new Amorphophallus from Gambia (Journ. Bot. 24, 1886,
Pp. 305-306).

Animal Life in High Latitudes (with S. O. RmpLeEy) (Zoologist,
1886).

1887

On anew genus of Orchideze from the island of St. Thomas, West
Africa (Journ. Linn. Soc. London, Bot. 24, 1887, p. 197-200,
pl. 6).

Orchideze, Scitamineze et Cyperaceze Africanze (Bot. da Soc. Bot.
Coimbra, 5, 1887).

Burmannia biocolor Mart. in Africa (Journ. Bot. 25, 1887, p. 85).

Angolan Scitamineze (Journ. Bot. 25, 1887, p. 129-135).

Botany of the Roraima Expedition, Orchidaceze and Cyperaceze
(Trans. Linn. Soc. London, Bot. 2, 1887, p. 281-288).

An unknown island (Globe, Nov. 1887).

Gardens Bulletin, S.S.

5
1888

Notes on self fertilisation and cleistogamy in Orchids (Journ.
Linn. Soc. London, Bot. 24, 1888, p. 389-395, pl. 16).

A revision of the genus Microstylis and Malaxis (Journ. Linn.
Soc. London, Bot. 24, 1888, p. 308-351).

Ponthieva grandiflora, n. sp. (Gardeners Chronicle, 3, 1885, p.
264).

Proliferous Orchids (Gardeners Chronicle, 4, 1888, p. 676).
A visit to Fernando Noronha (Zoologist, 12, 1888, p. 41-49).

1889

On the foliar organs of a new species of Utricularia from
St. Thomas, West Africa (Ann. Bot. 2, 1889, p. 305-307,
pl. 19).

Report on the destruction of coconut palms by beetles (Journ.
Str. Br. Roy. As. Soc. 20, 1889, p. 1-11).

1890

Notes on the Botany of Fernando de Noronha (Journ. Linn. Soc.
London, Bot. 26, 1890, p. 1-74, pl. 1-4)

New Malayan plants (Journ. Bot. 1890, p. 99).

On the method of fertilisation in Bulbophyllum macranthum and
allied Orchids (Ann. Bot. 4, 1890, p. 327-356, pl. 224).

On the habits of the Caringa (Formica gracilipes Gray) (Journ.
Str..Br. Roy. As. Soc. 22, 1890, p. 345-347).

Mosquito larvee in the pitchers of Nepenthes (Journ. Str. Br.
Rey, -As2o0c..22, 'I500,"p. 430).

Matonia pectinata in the Karimon Islands (Journ. Str. Br. Roy.
As. Soc. 22, 890, p. 430):

The Burmanniacez of the Malay Peninsula (Journ. Str. Br. Roy.
Fis, SOC. 22, 1890; p. 331-339).

On the so-called tiger’s milk ‘‘Susu Rimau’’ of the Malays (Journ.
Str. Br. Roy. As. Soc. 22, 1890, p. 341-344).

Coco-nut beetles (Journ. Str. Br. Roy. As. Soc. 22, 1890,
Pp. 429).

Notes on Zoology of Fernando Noronha (Journ. Linn. Soc.
London, Zool. 20, 1890, p. 473).

Thysanura and Collembola of Fernando Noronha (Journ. Linn.
Soc.: London, Zool. 20, 1890, p. 556).

Occurrence of a Crocodile on Cocos Island (Nature, 42, 1890,
D: 457)-

Calanthe rubens, n. sp. (Gardeners Chronicle, 7, 1800, p. 576).

Vol. IX) (1935).

6

1891
The genus Bromheadia (Journ. Linn. Soc. London, Bot. 28, 1891,
P. 331-339, pl. 42).
On two new genera of Orchids from the East Indies (Journ.
Linn. Soc. London, Bot. 28, 1891, p. 340-343, pl. 43-44).
Cypripedium insigne (Gard. Chron. 3rd series, vol. 10, 1891,
p. 94).

The grasses and sedges of the Malay Peninsula (Journ. Str. Br.
Roy. As. Soc. 23, 1891, p. I-33).

Plants collected at Bukit Etam, Selangor, by H. J. Kelsall (Journ.
Str. Br. Roy. As. Soc. 23, 1891, p. 72-75).

A day at Christmas Island, with list of animals and plants
recorded and bibliography (Journ. Str. Br. Roy. As. Soc.
23, I8QI, p. 123-140).

Discovery of stone implement in Singapore (Journ. Str. Br. Roy.
As. Soc. 23, 1831, p. I4I-142).

On the occurrence of a Rhododendron in Singapore (Journ. Str.
Br. Roy. As. Soc. 23, 1891, p. 144-146).

Pogonia punctata Bl. in Singapore (Journ. Str. Br. Roy. As.
Soc. 23, 1891, p 146-147).

The Keringga (Journ. Str. Br. Roy. As. Soc. 23, 1891, p. 147).

Frugivorous habits of the Tupaia (Journ. Str. Br. Roy. As.
S0C.323,. 1501) fe 149).

Diamonds in the Malay Peninsula (Journ. Str. Br. Roy. As. _
Soc. 24, 1891, p. 166-167).

On the occurrence of Petrosavia in Perak (Journ. Str. Br. Roy.
As. Soc. 24, 1891, p. 170-172).

Mr. Ridley’s expedition to Pahang (Straits Times Nov. 13, 1891,
cited in Journ. Bot. 30, 1892, p. 31-32).

The raised reefs of Fernando Noronha (Am. Journ. Sci. 41, 18or,

p. 406).
1892
Gambir (Agric. Bull. Mal. Peninsula no. 2 1892, p. 20-41).
1893

On the flora of the Eastern Coast of the Malay Peninsula (Trans.
Linn. Soc. Bot. ser. 2, 3, 1893, p. 267-408, pl. 61-66).

Dye Plants (Agric. Bull. Mal. Peninsula no. 3, 1893, p. 44-51).

Patchouli (Agric. Bull. Mal. Peninsula no. 3, 1893, p. 52-55).

Ngai Camphor (Agric. Bull. Mal. Peninsula no. 3, 1893,
p. 55-56).

Fibres (Agric. Bull. Mal. Peninsula no. 3, 1893, p. 56—60).

Expedition to the Tahan District, Pahang, Malay Peninsula
Proc. Roy. Geogr. Soc. 1892, p. 533-540).

Vanda Miss Joaquim (Gardeners’ Chronicle, 13, 1893, p. 740).

Gardens Bulletin, S.S.

7

1894

Note of Camphor Tree (Journ. Str. Br. Roy. As. Soc. 26,
1894, P. 35-39).

Stick insect destroying orchids (Journ. Str. Br. Roy. As. Soc.
26, 1894, p. 204).

On the Dispersal of seeds by mammals (Journ. Str. Br. Roy. As.
Soc. 25, 1894, p. Ii-—32).

Malay Materia Medica—lecture read to Straits Medical Society
1894, reprinted 1897. (Dates according to a note by Ridley
on the separate, which is otherwise without data).

Vegetation of Pahang (Journ. Str. Br. Roy. As. Soc. 25, 1894,
Pp. 49-56).

List of mammals recorded from Pahang (Journ. Str. Br. Roy.
As. Soc. 25, 1894, p. 57—60).

The Botanists of Penang (Journ. Str. Br. Roy. As. Soc. 25,
1894, p. 163-167).

Earthquake in the Malay Peninsula (Journ. Str. Roy. As.
Soc. 25, 1894, p. 169-171).

A large beetle caught in a pitcher of Nepenthes (Journ. Str. Br.
Roy. AS. Soc: 25, 1594; Pp. 172).

The bird-dropping spider (Ornithoscatoides) in Johore (Journ.
Str. Br. Roy. As. Soc. 25, 1894, p. 172-173).

List of plants collected by Lake and Kelsall during trip across
Johore (Journ. Str. Br. Roy. As. Soc. 26, 1894, p. 25-33).
Pothos flexuosa (Anadendron) (Gardeners Chronicle 15, 1894,

p. 526-527).

Orchideze of the Flora of Mt. Kinabalu’ (Trans. Linn. Soc.

London, Bot. 4, 1894, p. 233-240).

1895

The Mammals of the Malay Peninsula (Natural Science 6, 1895,
nos. 35, 36, 37, pp. 23-20, 89-96, 161-166).

Two new species of Thismia (Ann. Bot. 9, 1895, p. 323-326,
pl. 12).

On a new saprophytic monocotyledon (Ann. Bot. 9, 1895,
p. 56-57) (with P. Groom).

Utricularie in the Malay Peninsula (Journ. Bot. 33, 1895,
p. IO-ITI).

On two new species ef Clerodendron (Journ. Bot. 33, 1895,
p. 42-43).

Sago (Agric. Bull. Mal. Peninsula no. 4, 1895, p. 62-73).

Notes on soils and lallang grasses (Agric. Bull. Mal. Peninsula
no. 4, p. 73-86).

Vol. IX. (1935).

— *

8

Linospadix Micholitzii, n. sp. ‘(Gardeners Chronicle, 18, 1895,

p. 262).

A Strange Pet (Nature Notes, Mag. Selborne Soc. 6, 1895, p.
SQ).

Peperomia malaccensis, n. sp. (Kew Bull. 1895, p. 185).

1896

Cyrtandraceee malayenses (Journ. Linn. Soc. Bot. 32, 1806,
Pp. 497-528).

An enumeration of all Orchideze hitherto recorded from Borneo
(Journ. Linn. Soc. Bot. 31, 1896, p. 261-306, pl. 13-15, read
2I-Xi1-93).

The Dracaenas of the Malay Peninsula (Journ. Bot. 34, 1896,
p-. 162-168).

A new genus of Commelinaceae (Journ. Bot. 34, 1896, p. 329-330,
t. 360).

The Orchideze and Apostasiaceze of the Malay Peninsula (Journ.
Linn. Soc. Bot. 32, 1896, p. 213-416).

Report on the Diseases of the Nutmeg and Clove Trees in Penang
and Province Wellesley (Agric. Bull. Mal Peninsula no. 5,
1896, Pp. 91-95).

Dispersal of seeds by birds (Natural Science 8, 1896, p. 186-199).

Forests of Selangor (Selangor Journal, Sept. 1896, p. 444).

Habits of Macropisthodon rhodomelas (Zoologist, 1806).

1897

Facts about Ramie (reprinted from Straits Times, Singapore,
undated, probably Jan. 1897).

Malay Plant Names (Journ. Str. Br. Roy. As. Soc. 30, 1897,
p. 31-283).

Calanthe vestita, Lindl. in Selangor (Journ. Str. Br. Roy. As.
Soc. 30, 1897, p. 311-312).

Rubber cultivation (Agric. Bull. Mal. Peninsula no. 7, 1897,
p. 132-138).

Ramie (Agric. Bull. Mal. Peninsula no. 7, 1897, p. 138—141).

Notes on Sugar cultivation (Agric Bull. Mal. Peninsula no. 7,
1897, Pp. I4I—1I47).

Coffee Diseases (Agric. Bull. Mal. Peninsula no. 7, 1897,
p. 147-149).

Birds useful to Agriculturists (Agric. Bull. Mal. Peninsula no. 7,
1897, Pp. 149-150).

Spices (Agric. Bull. Mal. Peninsula no. 6, 1897, p. Hdadeat

Mr. H. Vaughan Steyens, Obituary (Journ. Str. Br. Roy. As.
Soc. 1897, p. X XI).

Jering Fruit (Pithecolobium lobatum). (Proc. As. Soc. Beng.,
1897, p. I).

Gardens Bulletin, S.S.

9

1898

Birds in the Botanic Gardens Singapore (Journ. Str. Br. Roy.
As. Soc. 31, 1898, p. 73-89).

The Peliosanthes of the Malay Peninsula (Journ. Str. Br. Roy.
As. Soc. 31, 1898, p. 91-98). |

The white snake of the Selangor caves (Journ. Str. Br. Roy. As.
Soc. 31, 1808, p. gg—ro0r).

Note on precocious coconuts (Journ. Str. Br. Roy. As. Soc. 31,
1898, Pp. 103-104).

The white-winged bat in Singapore (Journ. Str. Br. Roy. As.
Soc. 31, 1898, p. 104).

Hyblea puera Cram. (Journ. Str. Br. Roy. As. Soc. 31, 1800,
p. 104-105).

New Malayan Orchids (Journ. Bot. 36, 1898, p. 210-216).

New species of Entada from Singapore and Borneo (Journ. As.
Soc. Bengal 67, part 2, no. 2, 1898, p. 305-307).

Vegetables (Agric. Bull. Mal. Peninsula no. 8, 1808, p. 105—198).

The Poisonous plants of the Malay Peninsula (Agric. Bull. Mal.
Peninsula no. 8, 18908, p. 199-220).

Shade Trees (Agric. Bull. Mal. Peninsula no. 8, 1808, p. 222-230).

Para Rubber notes (Agric. Bull. Mal. Peninsula no. §, 1808,
p. 230-232).

Lemon Grass Oil (Agric Bull. Mal. Peninsula no. 8, 1808, p. 233).

Citronella Oil (Agric. Bull. Mal. Peninsula no. 8, 18608,
Pp. 233-234).

Report of Exploration of the Batu Caves (Brit. Assoc. Report,
1898, p. 2).

Ptychosperma Sanderiana, n. sp. (Gardeners Chronicle, 24,
1898, p. 330).

Livistona Woodfordii, n. sp. (Gardeners Chronicle, 23, 1808, p.
177).

1899

The habits of Malay Reptiles (with list) Journ. Str. Br. Roy.
Asiat, Soc. 32, 1899, p. 185-210).

Golden flowers (Journ. Str. Br. Roy. As. Soc. 32, 1809, p.
214-215).

An insectivorous squirrel (Journ. Str. Br. Roy. As. Soc 32,
1899, Pp. 217).

The Scitaminez of the Malay Peninsula (Journ. Str. Br. Roy.
As. Soc. 32, 1899, p. 85-184).

Note on Rhinoceros Hornbill (Journ. Str. Br. Roy. As. Soc.
32, 1899, p. 217).

Vol. IX. (1035).

Io

1900

Native rubbers of the Malay Peninsula (Agric. Bull. Mal.
Peninsula no. 9, 1900, p. 239-252).
Insect pests (Agric. Bull. Mal. Peninsula, no. 9, 1900, p. 253-264).

Kickxia africana (Agric. Bull. Mal. Peninsula no. 9, - 1900,
p. 286-287).

Injurious Fungi (Agric. Bull. Mal. Peninsula no. 9g, 1900, p. 287-

288).
The Flora of Singapore (Journ. Str. Br. Roy. As. Soc. 33, 1900,
p. 27-196).

The Orang Laut of Singapore (Journ. Str. Br. Roy. As. Soc.
33, IQ00, p. 247-250) (with W.W. Skeat).

Tiger’s Milk, further note (Journ. Str. Br. Roy. As. Soe. 34,
IQoO, p. IOI).

Some new Eastern Gingers (addendum to last paper) (Journ. Str.
Br. Roy. As. Soc. 34, 1900, p. 97-99).

Note on Malayan gingers (Journ. Str. Br. Roy. As. Soc. 34,
Ig00, Pp. QQ-IOI).

Plants collected at Penrissen, Sarawak, by R. S. Shelford (Journ.
Str. Br. Roy. As. Soc. 33, 1900, p. 22-24).

A botanical excursion to Gunong Jerai (Kedah Peak) (Journ. Str.
Br. Roy. As. Soc. 34, 1900, p. 23-30).

On the use of the Slow Loris in Malay medicine (Journ. Str. Br.
Roy. As. Soc. 34, 1900, p. 31-34).

Dammar and wood oil (Journ. Str. Br. Roy. As. Soc. 34, 1900,
p. 89-94).

New Malayan plants (Journ. Bot. 38, 1g00, p. 68-74).

Habenaria Columbe, n. sp. (Gardeners Chronicle, 28, 1900, p.
390).

1901

Supplementary notes on the Flora of Singapore (Journ. Str. Br.
Roy. As. Soc. 35, 1901, p. 84-90).
Rambong Beetle (Journ. Str. Br. Roy. As. Soc. 36, 1rgor,

p-. 138-139).

The flora of Mt. Ophir (Journ. Str. Br. Roy. As. Soc. 35, 1901,
p. I-28).

Garu and Chandu (Journ. Str. Br. Roy. As. Soc. 35, Igo1,
p- 73-82).

Calogramma festiva Walk. (Journ. Str. Br. Roy. As. Soc. 35,
I90I, p. 82-83).

Habits of the Drongo (Journ. Str. Br. Roy. As. Soc. 35, 1901,
p. 105).

Gardens Bulletin, S.S.

yt

The short-eared owl in Singapore (Journ. Str. Br. Roy. As. Soc.
35, IQOI, p. 105).

The Sumatran Rhinoceros (Journ. Str. Br. Roy. As. Soc. 35,
IQOI, p. IO5—I06).

In memoriam Dr. N. B. Dennys (Journ. Str. Br. Roy. As. Soc.
35, IQOI, p. I06—107).

1902

The timbérs of the Malay Peninsula (Agric. Bull. S.S. & F.M.S.
vol. I, nos. I, 2, 3, 4, 5, 6, 7 & 8, 1901-1902, p. I-13, 43-63,
Q5-II1, 135-145, 171-181, 209-220, 245-261, 289-292).

The dissemination of seeds by natural means (Agric. Bull. S.S.
& F.M.S,; I, 0. 4,.1902, p., 153-156).

Fruits of the Malay Peninsula (Agric. Bull. $.S. & F.M.S. 1,
10s. 10, Il), 12, '13;" 1902, "Pp. 371-382; 420-435, 4997503,
531-537). |

Sudu-sudu (Agric. Bull. S.S. & F.M.S. 1, no. 10, 1902, p. 389).

Volatile Oils (Agric. Bull. S.S. & F.M.S. 1, no. 9, 1902, p.
335- 342).

Ficus bracteata (Agric. Bull. S:S. & F.M.S. 1; no..10, 1902,
p. 387).

Some Malay Aroids (Journ. Bot. 40, 1902, p. 34-38).

A coffeebean pest (Agric. Bull. $.S. & F.M.S. 1, no. 10, 1902,
p. 387-388).

Fruit Cultivation (Agric. Bull. S.S. & F.M.S. 1, no. 8, 1902,
p. 298-300).

A large Indian Melon (Agric. Bull. S.S. & F.M.S. 1, no. 13,

us) ehs p.°548).

Funtumia elastica (Agric. Bull S$.S. & F.M.S. 1, no. 14, 1902,
Pp. 591-592).

Malay Tiger Beetles (Journ. Str. Br. Roy. As. Soc. 38, 1902,
Pp. 129-131).

Malay plant names (Latin-Malay) (with C. Curtis) (Journ. Str.
br. Roy. As. Soc. 38," 1902, p.- 30-122).

1903

Jelutong (Dyera costulata) (Agric. Bull. $.S. & F.M.S. 2, no. 3,
1903, DP. 95-97).
Sarcolobus globosus (Agric. Bull. S.S. & F.M.S. 2, n. 7, 1903,

Pp. 223-224).

Gunda Sikkima (Agric. Bull. S.S. & F.M.S. 2, no. 2, 1903,
p. 69-70).

The black Cobra (Agric. Bull. S.S. & F.M.S. 2, no. 7, 1903,
p. 225-226).

The Camphor Tree (Agric, Bull. $.S. & F.M.S. 2, no. 5, 1903,
p. 163-164).

Vor, 8X s\ 61935).

I2

Barringtonia seed as a Sakai Food (Agric. Bull. S.S. & F.M.S. 2,
NG. 5, 1903,.%. BOs):

Rattans (Agric. Bull. S.S. & F.M.S. 2, nos. 4 & 5, 1903, p. 129-
136, 157-159).

Turf and Fodder Grasses (Agric. Bull. S.S. & F.M.S. 2, no. 9,
1903, P. 273-279).

Datura Poisoning (Agric. Bull. S.S. & FMS. 2; no, ¥,- i603,
Pp. 224-225).

Boring Beetles in Para Rubber (Agric. Bull. $.S. & F.M.S. 2;
no. 7, 1903, Pp. 222—223).

A Funtumia Pest, (Agric. Bull. S.S. & F.M.S. 2, no. 11, 1903,
Pp. 355)-

De Maleische timmerhoutsoorten (Bull. 12, Kolon. Mus.
Haarlem, Amsterdam 1903, Translated from Agric. Bull.
VOl..1, G02)?

A Castilloa borer (Agric. Bull. $.S. & F.M.S. 2, no. 10, 1003,
p. 3a2):

List of plants collected by Mr. W. D. Barnes on Gunong Benom,
Pahang (Journ. Str. Br. Roy. As. Soc. 39, 1903, p. 10-18).

New Malay Orchids (Journ. Str. Br. Roy. As. Soc. 39, 1903,
p- 71-87).

Work on Sakais by Messrs. Skeat and Blagden (Journ. Str. Br.
Roy. As. Soc. 39, 1903, p. 204).

A new Balanophora from Tenimber Islands (Journ. Str. Br. Roy.
As’ Soc...30, 19085-2. 207).

1904

On the flowering of Barringtonia racemosa (Journ. Str. Br. Roy:
As.-Soe. 435) 1964p. F25—120).

Fertilisation of Webera stellulata (Journ. Str. Br. Roy. As.
Soc. 41, 2604, p."1520-127)".

The orang laut of Singapore (Journ. Str. Br. Roy. As. Soc. 41,
IQ04, Pp. 120-130).

The mosquito Plant (Agric. Bull. $.S. & F.M.S. 3, no. 1, 1904,
Bi. 2a):

Parasitic Fungi on Hevea brasiliensis (Agric. Bull. $.S. & F.M.S.
3, NO. 5, 1904, p17 3egs)- .

The Palm collections of the Botanic Gardens, Singapore (Agric.
Bull. S.S. & F.M.S. 3, no. 7, 1904, p. 249-266).

Pineapple cultivation (Agric. Bull. $.S. &. F.M.S. 3, no. 2, 1904,

P- 37-40):

Three new Bornean palms (Gard. Chron. 3rd series, vol. 35, 1904,
Pp. 50).

New Malayan plants (Journ. Str. Br. Roy. As. Soc. 41, 1904,
p. BEsSH)-

Insect Attractions in Flowers (New Phytologist, 3, 1904, p. 164).

Gardens Bulletin, S.S.

13

1905

The Gesneraceee of the Malay Peninsula (Journ. Str. Br. Roy.
As. Soc. 44, 1905, p. I-02).

The Aroids of Borneo (Journ. Str. Br. Roy. As. Soc. 44, 1905,
p. 169-188).

Nesting of the little grey Wood-pecker (Journ. Str. Br. Roy. As.
Soc. 44, 1905, p. 226-227).

Nesting of Draco fimbriatus (Journ. Str. Br. Roy. As. Soc. 44,
1905, Pp. 227).

A wasp attacking a leaf-mining caterpillar (Journ. Str. Br. Roy.
As. Soc. 44, 1905, p. 227-228).

On the fertilisation of Grammatophyllum (Journ. Str. Br. Roy.
As. Soc. 44, 1905, p. 228-229).

New and little known Malayan plants, series II (Journ. Str. Br.
Roy. As. Soc. 44, 1905, p. 189-211)

Fibres (contd.) (Agric: Bull:S:S. & F.M.S) 4) nos. 1, 2; 3, 5, 6,
1905, Pp. 6-8, 43-45, 64-67, 155-159, 213-218).

Glycine hispida (Agric. Bull. S.S. & F.M.S. 4, no. 2, 1905, p. 49)

Note on a peculiar flow of latex in a Hevea (Agric. Bull. S.S. &
FMS. 4) 602 t008) 'p.) 49):

Kumus (Shorea ciliata) Agric. Bull. S.S. & F.M.S. 4, no. 3, 1905,
p. 63-64). :

Tile pots for Casuarina seedlings (Agric. Bull. S.S. & F.M.S.
45,20. )9;) 1905 ,)'p.' 167-63).

Rubber pests (Agric. Bull. S.S. & F.M.S. 4, no. 3, 1905, p. 68-69).

Rubber experiments in the Botanic Gardens, Singapore (Agric.
Bull. $.S. & F.M S. 4, no. 4, 1905, p. 98-110).

Grammatophyllum speciosum (Agric. Bull. S.S. & F.M.S. 4,
no. 5, 1905, p, 141-143).

The pupoi (Connaropsis Griffithii) (Agric. Bull. S.S. & F.M.S.
A, HO 5, Pt 155).

Germinating Para rubber seeds (Agric. Bull. S.S. & F.M.S. 4,
no. 5, 1905, p. 159).

Note on Termes gestroi (Agric. Bull. S.S. & F.M.S. 4, no. 5,
$O05;-Pir THO). = | 7

Profits on a small rubber estate (Agric. Bull. S.S. & F.M.S. 4,
NWOs0;°1005,) Do 248):

Isoptera borneensis (Agric. Bull. S.S. & F.M.S. 4, no. 6, 1905,
Da Zip),

mie Hugh Low, obituary (Agric. Bull. $.S. & F.M.S. 4, no. 6,
1905, Pp. 239-241).

Notes on the collection of Aroids cultivated in the Botanic
Gardens, Singapore, with list (Agric. Bull. S.S. & F.M.S. 4,
HO. 7, 1905, p. 255-204).

V Oren X 3 (1035).

14

Cotton experiments in the Botanic Gardens, Singapore (Agric.
Bull. $.5. & F.M.S.. 4,.:no.: 7, .1905, .p.. 265-266).

Fibre of Melochia (Agric. Bull. S.S. & F.M.S 4, no. 7, 1905,
Di 203).

Fungus on Para rubber leaves (Agric. Bull. S.S. & F.M.S. 4,
1105175, ROGER, Du274)-

Red Coco-nut beetle (Agric. Bull. S.S. & F.M.S. no. 7, 1905,
Pp. -272-273).

The History and Development of Agriculture in the Malay Penin-
sula (Agric. Bull. S.S. & F.M.S. 4, no. 8, 1905, p. 292-317).

Rubber in Sarawak (Agric. Bull. S.S. & F.M.S. 4, no. 9, 1905,
Pp. 305-366).

The position of Rubber among cultivated plants (Agric. Bull.
S.S. & F.M.S. 4, no. 10, 1905, p. 382-384).

Caterpillar attacking tobacco plants (Agric. Bull. S.S. & F.M.S.
4, NO. 10, 1905, p. 391).

A bark fungus on Para rubber (Agric. Bull. $.S. & F.M.S. 4a,
no. II, 1905, p. 423).

Report on Experimental Tapping of Para rubber in the Botanic
Gardens, Singapore for 1904 (with R. Derry) (Agric. Bull.
S.S. & F.M.S. 4, no. 11, 1905, p. 424-443).

Rubber pests (Agric. Bull. S.S. & F.M.S. 4, no. 12, 1905, p. 457-

458).

Scitaminez philippinensis (Govt. Labor. Publ. (Philipp.) 35, 1905,
p. 83-87).

On the dispersal of seeds by wind (Ann. Bot. 19,° 1905,
D. 351-363).

1906

Exporting seed of Para rubber (Agric. Bull. $.S. & F.M.S. 5,
no. I, 1906, p. 1-2) (Reprinted in Kew Bull. 1906, p. 106).

The Engkala, a new fruit (Agric. Bull. S$.S. & F.M.S. 5, no. 1,
1906, p. 5-6).

New Palms and Aroids added to collection in Botanic Gardens,
Singapore (Agric. Bull. S.S. & F.M.S. 5, no. 1, 1906, p. 6-8).

East Indian Dragon’s Blood (Agric. Bull. S.S. & F.M.S. 5, no. 2,
1906, p. 28-34).

Price of Timber in 1867 (Agric. Bull. S.S. & F.M.S. 5, no. 2,
1906, p. 38-39).

Betis or Malay Bilian (Agric. Bull. S.S. & F.M.S. 5, no. 2, 1906,
Pp. 39-40).

The timbers and guttas of the Malay Peninsula (Agric. Bull. S.S.
& F.M.S. 5, no. 3, 1906, p. 62-64).

A fungus attacking roots of Para rubber (Agric. Bull. S.S. &
F.M.S. 5, no. 3, 1906, p. 64-65).

Gardens Bulletin, S.S.

15
Cassia fistula (Agric. Bull. S.S. & F.M.S. 5, no. 3, 1906, p. 68).
A parasitic fungus on Roses (Agric. Bull. S.S. & F.M.S. 5, no. 3,
s “1906, p. 6BY.

Crickets eating Rubber plants (Agric. Bull. S.S. & F.M.S. 5,
no. 3, 1906, p. 68).

Notes on Stebbing’s paper on Termes gestroi (Agric. Bull. S.S. &
F.M.S. 5, no. 4, 1906, p. rro—1IT).

Paspalum dilatatum (Agric. Bull. S.S. & F.M.S. 5, no. 4, 1906,
p. 113-114).

A warning to planters (Thefts of rubber seedlings) (Agric. Bull.
S:S. & F.M.S. 5, no. 4, 1906, p. 116).

The Water Hyacinth (Agric. Bull. S.S. & F.M.S. 5, no. 4, 1906,
DTI

Note on an old Rubber stump (Agric. Bull. S.S. & F.M.S. 5,
no. 4, 1906, p. 117-118).

Tapioca as a catch crop for Rubber (Agric. Bull. S.S. & F.M.S.
5, NO. 5, 1906, p. 133-135). ,

The Old Rubber tree at Penang (Agric. Bull. S.S. & F.M.S. 5,
no. 5, 1906, p. 138).

Malay Drugs (reprint in part of a lecture on Malay Materia
Medica read before the Medical Society in 1894, published
originally in Journal of that Society) (Agric. Bull. S.S. &
F.M.S. 5, no. 6, 1906, p. 193-206).

A new way of dealing with Termes gestroi (Agric. Bull. S.S. &
F.M.S. 5, no. 6, 1906, p. 207-208).

Malay Drugs (contd.) (Agric. Bull. S.S. & F.M.S. 5, no. 7, 1906,
Pp. 245-254).

Note on the method of preparing Dragon’s Blood (Agric. Bull.
S.S. & F.M.S..5, no. 8, 1906, p. 264-265).

Malay Drugs (contd.) (Agric. Bull. S.S. & F.M.S. 5, no. 8, 1906,

p. 269-282).

Synthetic Rubber (Agric. Bull. S.S. & F.M.S. 5, no. 10, 1906,
Pp. 369-373).

Creosoting Rubber (Agric. Bull. S.S. & F.M.S. 5, no. 11, 1906,
P. 395-396).

A Disease of Rubber seedlings (Agric. Bull. S.S. & F.M.S. 5,
no. II, 1906, p. 400-401).

Attacus atlas and Rubber (Agric. Bull. S.S. & F.M.S. 5, no. 11,
1906, p. 401).

Notes on the action of Copper Sulphate on Aquatic Plants (Agric.
Bull. S.S. & F.M.S. 5, no. 11, 1906, p. 403-408).

Second Annual Report on the Experimental Tapping of Para
Rubber trees in the Botanic Gardens, Singapore for 1905
(with R. Derry) (Agric. Bull. S.S. & F.M.S. 5, no. 12, 1906,
Pp. 440-464). :

Vol- EX. (1935).

16

Begonias of Borneo (Journ. Str. Br. Roy. As. Soc. 46, 1906,
Pp. 246-261).

Grasses and sedges of Borneo (Journ. Str. Br. Roy. As. Soc. 46,
1906, p. 215-228).

Sciatamineze of Borneo (Journ. Str. Br. Roy. As. Soc. 46, 1906,
Pp. 229-245).

Habits of the Tupaia (Journ. Str. Br. Roy. As. Soc. 45, 1906,
p. 279).

The Menagerie at the Botanic Gardens (Journ. Str. Br. Roy.
As. Soc. 46, p. 133-194).

Curious nesting place of Simotes octolineatus (Journ. Str. Br.
Roy. As. Soc. 46, 1906, p. 263).

Note on the foliar organs of Monophyllea (Ann. Bot. 20, 1906,
p. 213-214).

Fertilization of Barringtonia (Journ. Str. Br. Roy. As. Soc. 46,
1906, p.°2;'/263).

An expedition to Christmas Island (Journ. Roy. As. Soc. Str.
Br. 45, 1906, p. 137-271).

1907
The Oil Palm (Agric... Bull. S.S. & FM.S. 6, no) @ehree7,
P. 37-40).
Chinese Anti-opium Drugs (Agric. Bull. S.S. & F.M.S. 6, no. 2,
1907, Pp. 45-47).
Late flowering of Grammatophyllum (Agric. Bull. S.S. & F.M.S.
6, no. 2, 1907, p. 49).
Annual Report of the Botanic Gardens, Singapore for 1906
(Agric. Bull. S.S. & F.M.S. 6, no. 4, 1906, p. 92-1071).
African Rubber-vines (Agric. Bull. S.S. & F.M.S. 6, no. 5, 1907,
p- 125-126).

Funtumia elastica flowering in Johore (Agric. Bull. S.S. & F.M.S.
6, H0..5; 1007, Dp. 127).

Fruiting of the Frangipani (Agric. Bull. S.S. & F.M.S. 6, no. 5,
1907, p. 128).

Cyanogenesis in plants (Agric. Bull. S.S. & F.M.S. 6, no. 5,
1907, Pp. 131).

Abnormalities in the stem of Hevea (Agric. Bull. S.S. & F.M.S.
6, no. 6, 1907, p. 157-158).

Wind Twist (Agric. Bull. S.S. & F.M.S. 6, no. 6, 1907, D. 160).
Review of Notes on Rubber Cultivation by Wylie and Ferreira
(Agric. Bull. S.S. & F.M.S. 6, no. 6, 1907, p. 167-168).

Casuarinas (Agric. Bull. $.S. & F.M.S. 6, 1907, p. 169-170).
Some timber notes (Agric. Bull. S.S. & F.M.S. 6, no. 6, 1907,
p. I70-I71).

Gardens Bulletin, S.S.

17

A rubber factory in Singapore (Agric. Bull. S.S. & F.M.S. 6,
no,.6, 1007, pr 474).

Early fruiting of Para rubber (Agric. Bull. S.S. & F.M.S. 6, no. 6,
Dp. 176). |
Conifers cultivated in the Botanic Gardens, Singapore (Agric.

Bull. S.S. & F.M.S. 6, no. 7, 1907, p. 197-201).

Pandans (Agric. Bull. S.S. & F.M.S. 6, no. 8, 1907, p. 241-244).

The Herbarium (of the Botanic Gardens, Singapore) (Agric.
Biull. 8.5: & FES: 6, no. 10, 1907, p. 329-333).

Lallang as a paper material (Agric. Bull. S.S. & F.M.S. 6, no. 11,
1907, DP. 379-382).

Flora of Singapore Island (In Handbook to Singapore, Ed. 2, by
G. M. Reith, 1907, p. 117-123).

Malay Drugs (Agric. Bull. $.S. & F.M.S. 5, 1906, p. 193-206).
Transl. as ‘‘Maleische Geneesmiddelen’’ in Indische Mercuur
1907.

De inlandsche Geneesmiddelen der Maleiers (reprinted from ‘‘De
Indische Mercuur’’—Amsterdam, J. H. de Bussy, 1907).
New or rare Malayan plants, series III (Journ. Str. Br. Roy. As.

pie. 40, 1007.1." 11-52).

Christmas Island Flora (Journ. Str. Br. Roy. As. Soc. 48, 1907,

A Ba -1O7—108).

The Pagan races of the Malay Peninsula, by Skeat and Blagden.
A review (Journ. Str. Br. Roy. As. Soc. 49, 1907, p. I-5).
Branching in Palms (Ann. Bot. 21, 1907, p. 415-422, pl. 34-39).
Materials for a flora of the Malay Peninsula (Monocotyledons).
(Singapore 1907, pt. I, 1907, 22+233 pp.; pt. 2, 1907,

44+ 235 pp.; pt. 3, 1907, 75+197 pp.)

1908

Some notes on the acclimatisation of plants (Agric. Bull. S.S. &
F.M.S. 6, no. 12, 1907, p. 415-420, and 7, no. 1, 1908,
i: =O). |

Mr. A. B. Stephens, obituary (Agric. Bull. S.S. & F.M.S. 7
10; :2, 1908,: p: ‘66

Annual Report on the Botanical Gardens of the Colony for 1907
(Agric. Bull. S.S. & F.M.S. 7, no. 3, 1908, p. 104-100).

Heliconias (Agric. Bull. S.S. & F.M.S. 7, no. 4, 1908, p. 129-
%32);

Funtumia elastica fruiting in Perak (Agric. Bull. S.S. & F.M.S.
7, 10.4, 1908, p. 136).

A further note on Crinothrix ee Bull. S.S. si F.M.S.:7, no. 4,
1908, p. 136).

Vol. IX. (1935).
B

18

Curious root-development of Albizzia (Agric. Bull. S.S. & F. M.S.
7, NO. 4, 1908, p. 137).

Big rubber trees in the Botanic Gardens, Singapore (Agric. Bull.
5.8. & F.M.S. 7, no. 7, 1908, p. 253-255).

A disease of Cloves (Agric. Bull. S.S. & F.M.S. 7, no. 7, 1908,
p. 263-264).

Another coco-nut beetle (Agric. Bull. S.S. & F.M.S. 7, no. 7,
1908, p. 265-266).

Acalypha fruticosa. Forst. (Agric. Bull. $.S. & F.M.S. 7, no. 7,
1908, p. 268).

Fruiting of Sciadopitys verticillata (Agric. Bull. S.S. & F.M.S.
7, 2G.22) 1908, PDP. 200).

Germination of Para rubber seed (Agric. Bull. S.S. & F.M.S. 7,
no. 8, 1908, p. 333-334).

Size of Rubber seed (Agric. Bull. S.S. & F.M.S. 7, no. 8,1908,
P. 335-339).

Coca leaves (Agric. Bull. S.S. & F.M.S. 7, no. 8, 1908, p. 339).

Notes on some piscicides (Agric. Bull. $.S. & F.M.S. 7, no. 10,
1908, p. 443-445).

New or rare Malayan plants, series IV (Journ. Str. Br. Roy. As.
Soc. 50, 1908, p. III—I52).

On a collection of plants made by H. C. Robinson and L. Wray
from Gunong Tahan, Pahang (Journ. Linn. Soc. Bot. 38,
1908, p. 301-336). é

The Labiates of the Malay Peninsula (Journ. Str. Br. Roy. As.
Soc. 50, 1908, p. I05—107).

A list of the Ferns of the Malay Peninsula (Journ. Str. Br. Roy.
As. Soc. 50, 1908, p. I-59).

Bats in a bamboo (Journ. Str. Br. Roy. As. Soc. 50, 1908,
p. 103-104).

The Crackling Moth (Journ. Str. Br. Roy. As. Soc. 50, 1908,
p. I0Q—II0).

Gesneracece (with Gamble & Prain) in Materials for a flora of
the Malay Peninsula (Journ. As. Soc. Bengal 74, pt. 2,
Extra number 1908, p. 730-793).

Botany of British Malaya (in Twentieth Century Impressions of
British Malaya, by Wright and Cartwright, 1908, p. 331-336).

1909
New or rare Malayan plants series V (Journ. Str. Br. Roy. As.
Soc. 54, 1909, p. I-61).
Zingiberaceze from South Negros (Elmer, Leafl. Philipp. Bot. 2,
art. 31, 1909, p. 603-607).
The flora of the Telom and Batang Padang Valleys (Journ. Fed.
Mal. St. Mus. 4, 1909, p. I-98).

Gardens Bulletin, S.S.

ee)

The Scitamineze of the Philippine Islands (Philipp. Journ. Sci.
4, Bot. 1909, p. 155-199).
Musszenda erythrophylla (Agric. Bull. S.S. & F.M.S. 8, no. 3,
1909, p. 114).
Various notes (on rubber cultivation) (Agric. Bull. S.S. & F.M.S.
no. 4, 1909, Pp. 134-136).
Planting of trees (Agric. Bull. S.S. & F.M.S. 8, no. 5, 1909,
Pp. 239-240).
Variation in the flower of the Black Pepper (Agric. Bull. S.S. &
F.M.S. 8,.no.-5,: 1909, Pp.) 244).
Ferns and their cultivation, with list (with T. Wilson Main)
Agric. Bull. S.S. & F.M.S. 8, no. 6, 1909, p. 282-297).
A new Fungus pest on Para rubber (Agric. Bull. S.S. & F.M.S.
S. oo. 7, 1900, p. 310-312).
Improvements in the manufacture of Gambir (Agric. Bull. $.S. &
F.M.S. 8, no. 7, 1909, p. 312-314).
Castilloa elastica fruiting in Singapore (Agric. Bull. S.S. &
FMS! &, -tio.’-7,; 1909, p.' 420).
Effect of gale on a para rubber tree (Agric. Bull. S.S. & F.M.S.
8, no. 7, 1909, Pp. 322).
Azolla as an anti-mosquito plant (Agric. Bull. $.S. & F.M.S. 8,
no. 7, 1909, Pp. 322-323).
Remedies for snake-bite (Agric. Bull. S.S. & F.M.S. 8, no. 7,
1909, P. 323-324).
Rubber cultivation on so-called Peat Soil (Agric. Bull. S.S. &
F.M.S. 8, no. 10, 1909, p. 468-470).
Peat Soils (Contd.) (Agric. Bull. S.S. & F.M.S. 8, no. 11, 1909,
p. 516-518).
The Black Hevea fungus (Agric. Bull. S.S. & F.M.S. 8, no. 11,
1909, p. 521).
Brazil nuts (Agric. Bull. S.S. & F.M.S. 8, no. 12, 1909, p. 551—
553).
Gomphocarpus semilunatus (Agric. Bull. $.S. & F.M.S. 8, no. 12,
1909, p. 5600-561).
New Philippine Zingiberaceze (Hlmer Leafl. Philipp. Bot. 2, 1900,
Pp. 569-572).
Branching Palms (Ann. Bot. 23, 1909, p. 338-339).
Fruit of Burbidgea (Journ. Str. Br. Roy. As. Soc. 53, 1909, p.
175-176).
Plants of Prince of Wales Island coll. by Sir W. Hunter (Journ.
Str. Br. Roy. As. Soc. 53, 1909, p. 49-51).

Botanical Journey in the Malay Peninsula (Kew Bull. 1900,
p. 159).

Vol. IX. (1935).

20

1910
Symbiosis of ants and plants (Ann. Bot. 24, 1910, p. 457-483,
pl. 35-36).
A Peat-soil note (Agric. Bull. S.S. & F.M.S. 9, no. 1, 1910,
Pp! Is);

A nest of Termes malayanus with many queens (Agric. Bull.
5.9. & M.S: 9, no. 1910, p. 23)i

Tillage of Soil (Agric. Bull. S.S. & F.M.S. 9, no. 3, 1910, p. 80—
84).

The black Brazil cherry, Eugenia brasiliensis (Agric. Bull. S.S. &
F.M.S. 6, no. 3, roro,p. 85).

The abolition of the Botanic Gardens of Penang and History of
the Gardens of the Peninsula (Agric. Bull. S.S. & F.M.S. 9,
no. 3, IQIO, p. 97—I05).

Horticultural notes (Peliosanthes violacea var. Clarkei, Dendro-
bium spectabile, Hosea Lobbiana, Cleaning cement floors
(Agric. Bull. S.S. & F.M.S. 9, no. 3, 1910, p. 105-107).

Coco-nut Palm disease (Agric. Bull. $.S. & F.M.S. 9, no. 5, 1910,
p. 178-180).

Timber notes (including Shorea collina, n. sp., Shorea materialis,
n. sp.) (Agric. Bull. S.S. & F.M.S. 9, no. 5, 1910, p. 180-183).

Historical notes on the Rubber industry (Agric. Bull. $.S. &
F.M.S. 9, no. 6, 1910, p. 201-213). :

Coco-nuts in peat soil (Agric. Bull. S.S. & F.M.S. 9, no. 6, 1910,
p. iA).

Another Para rubber fungus (Agric. Bull. $.S. & F.M.S. 9, no. 6,
IQIO, p. 216-218).

Report on the Expenimental tapping of Para rubber trees in the
Botanic Gardens, Singapore for 1909 (with R. Derry) (Agric.
Bull. $.S. & F.M.S. 9, nos. 7 & 8, gto, p. 255-237, 289-297).

Further notes on Malay Camphor (Agric. Bull. S$.S. & F.M.S.
9, no. 8, I9IO, p. 299-300).

A. D. Machado and J. B. Carruthers, obituaries (Agric. Bull.
S.S5-& FOMES: 9; 110: '8) 1680} p. 320-396):

Rubber Fungi, a lecture delivered at the Agri-Horticultural Show
(Agric. Bull. S.S. & F.M.S. 9, no. 9, 1910, p. 380-384).
Chilis as a catch-crop (Agric. Bull. S.S. & F.M.S. 9, no. 11, 1910,

P. 450-452).

Disinfection of Rubber seeds (Agric. Bull. S.S. & F.M.S. 9,
no. II, I9I0, p. 453-456).

Coco-nut trees attacked by a Coccid (Agric. Bull. $.S. & F.M.S.
Q; 10. 11, TOTO; p. 465)s

Planting of trees (Agric. Bull. S.S. & F.M.S. 9, no. 12, 1910,
p. 481).

Gardens Bulletin, S.S.

21

Dr. Melchior Treub, Dr. Burck, obituaries (Agric. Bull. S.S. &
F.M.S. 9, no. 12, 1910, p. 495-496).

The Story of the Rubber industry (Malay States Development
Agency, undated, probably rgro).

Butomaceze, Hydrocharitaceze, Graminez, Cyperacee, Flagellari-
aceze, Pontederiacez, Liliaceze, Ameryllidaceze, Dioscoreacee,
Zingiberaceee and Marantacezee in WINKLER, Beitr. z.
Kenntnis der Flora und Pflanzengeographie Borneos (Engl.
Bot. Jahrb. 44, 1910, p.518—532).

A termite’s nest with 8 queens (Journ. Str. Br. Roy. As. Soc.
54, 1910, p. 157).

The abolition of the Botanic Gardens, Penang (Agric. Bull. S.S.
& F.M.S. 9, 1910, p. 100-105; reprinted in Kew Bull. rgro,
Pp. 153-158).

The Flora of the North-western States, Malay Peninsula (Kew
Bull. 1910, p. 202-204).

Monocotyledonze (with Hub. Winkler) (in Winkler, Beitrage
zur Flora und Pflanzengeographie Siidostborneos, Engl. Bot.
Jahrb. 44, 1910, p. 518-532).

New or rare Malayan plants, series V. Journ. Str. Br. Roy. As.
Soc. 54, I9I0, p. I-61).

Aristolochia minutiflora Ridley ms. (Kew Bull. rgro, p. 79).

Poisonous principles of Rhus (Pharmaceutical Journal, March 10,
IQIO, p. 360).

1911

Legislation against the dissemination of Pests (Agric. Bull. S.S.
& F.M.S. 10, no. 1, I9gII, p. 1-4).

Eutypa as a Parasite (Agric. Bull. S.S. & F.M.S. 10, no. 1, 1911,
p.°6).

Gum Benjamin (Agric. Bull. S.S. & F.M.S. 10, no. 1, rorr,
p. 12-16).

Rubber Smoking House (Agric. Bull. S.S. & F.M.S. 10, no. 3,
IQII, p. 64-66).

Science and Agriculture (Agric. Bull. S.S. & F.M.S. 10, no. 3,
IQII, p. 66-68).

Hevea disease in Ceylon (Agric. Bull. S.S. & F.M.S. 10, no. 3,
IQII, p. 70-71).

Mustard as a Catchcrop for Rubber (Agric. Bull. S.S. & F.M.S.
IO, NO. 3, III, p. 74).

Spangle-scale on Soursop (Agric. Bull. S.S. & F.M.S. 10, no. 3,
IQII, Pp. 74-75).

Chonemorpha rubber vines (orig. descript. of Ch. Rheedei and

Ch. penangensis) (Agric. Bull. S.S. & F.M.S. 10, no. 5, rorr,
p. 146-148).

Vol. 1X. (1935).

22

Fasciation in Eucharis (Agric. Bull. S.S. & F.M.S.. 10, no. 5,
IQII, p. 154).

The Drought spell (Agric. Bull. S.S. & F.M.S. 10, no. 6, rgrr,
p. 181-186).

Travelling of Rubber seed (Agric. Bull. S.S. & F.M.S. 10, no. 6,
IQII, p. IQI—Ig2).

Stumping on Estates (Agric. Bull. S.S. & F.M.S. 10, no. 6, rg11,

p. 195).

Birds and Crops (Agric. Bull. S.S. & F.M.S. 10, no. 8, 1911,
Pp. 232-235).

Cinnamomum Deschampsii (Agric. Bull. S.S. & F.M.S. ro, no. 8,
IQII, Pp. 235).

Abnormal Rubber tree (Agric. Bull. S.S. & F.M.S. 10, no. 11,
IQII, 351-352).

Root-shoots in Casuarina glauca (Agric. Bull. S.S. & F.M.S. 10,
no. II, IQII, p. 359).

A Bleeding Rubber tree (Agric. Bull. $.S. & F.M.S. ro, no. 11,
IQII, p. 361).

The Flora of Lower Siam (Journ. Str. Br. Roy. As. Soc. 59, 1911,
p. 15-26).

The Gymnosperms of the Malay Peninsula (Journ. Str. Br. Roy.
As. Soc. 60, I9II, p. 53-68).

An Account of a Botanical Excursion to Lower Siam (Journ. Str.
Br. Roy. As. Soc. 59, IQII, p. 27-234, map.).

The Rt. Rev. M. F. Hose, D.D. (Journ. Str. Br. Roy. As. Soc.
57, I9II, p. 3-4).

A scientific expedition to Temengoh, Upper Perak (Journ. Str.
Br. Roy. As. Soc. 57, I9II, p. 5-122).

Rengas Poisoning (Malayan Medical Journal, 9, 1911, p. 7-13).

A retrospect and some cautions (Grenier’s Rubber News, Kuala
Lumpur, ro1tr).

1912

A botanical excursion to Pulau Adang (Journ. Str. Br. Roy.
As. Soc. 61, 1912, 45-65).

New plants from Sarawak (Sarawak Mus. Journ. 12, rg12,
p. 30-38).
Spices (1912, Macmillan & Co., London, vili+ 449 pp.).

New or rare Malayan plants (series VI) (Journ. Str. Br. Roy.
As. Soc. 61, I912, p. I-43).

Some plants collected on Moulton’s Expedition to Batu Lawi
(Journ. Str. Br. Roy. As. Soc. 63, 1912, p. 59-62).

Decades Kewenses (Kew Bull. ror2, p. 380-380).

Gardens Bulletin, S.S.

23

1913
Contributions to a flora of Borneo (Sarawak Mus. Journ. 13, 1913,

| p- 67-98).

Some Bornean Aroidez (Journ. Bot. 51, 1913, p. 201-202, t. 527).

An expedition to Mount Ménuang Gasing, Selangor; with an
account of the journey by C. B. Kloss (Journ. Linn. Soc.
Bot. 41, 1913, p. 285-304).

A new Alpinia from Borneo (Journ. Bot. 51, 1913, p. 247).

Decades Kewenses (Kew Bull. 1913, p. 267-268).

African Zingiberaceze (in Talbot’s South Nigerian Plants, Brit.
Mus. Public., 1913, p. 110-112).

1914

On a collection of plants from G. Mengkuang Lebah, Selangor
(Journ. Fed. Mal. St. Mus. 5, 1914, p. 28-50).

Plante nove papuanz (Journ. Bot. 52, 1914, p. 289-296).

Decades Kewenses (Kew Bull. rg14, p. 188, 209-210, 211, 324,
327, 332, 377-378).

Moracez, Scitamineze, Heemodoracee, Iridaceze, Amaryllidacee,
Taccaceze, Dioscoreaceze, Liliaceze, Pontederiacezee, Xyrida-
ceee, Commelinaceze, Araceze, Eriocaulaceze, in L. S. Gress
Contribution Flora, etc. of Mt. Kinabalu (Journ. Linn. Soc.
London Bot. 42, 1914, p. 136-138, 162-167, 171-172).

Cypripedium Pereirz, n. sp. (Gardeners Chronicle, 55, 1914,

p. 236).
1915

On two new plants from Gunong Tampin, Negri Sembilan
(Journ. Fed. Mal. Stat. Mus. 6, pt. 1, 1915, p. 39).

A new Begonia from Borneo (Sarawak Mus. Journ. 26, 1915,
p. 177).

An expedition to Mt. Menuang Gasing, Selangor (Journ. Fed.

Mal. Stat. Mus. 6, 1915, p. I-21).

Plants from G. Kerbau, Perak (Journ. Fed. Mal. Stat. Mus. 6,
1915, Pp. 43-62).

New or rare Malayan plants series VII (Journ. Str. Br. Roy.
As. Soc. 68, 1915, p. 11-14).

Two new orchids from the Prov. of Bandon, S.W. Siam (Journ.
Fed. Mal. Stat. Mus. 5, 1915, p. 156-157).

The plants of Koh Samui and Koh Pennan (Journ. Fed. Mal.
Stat. Mus. 5, 1915, p. 158-168).

The botany of G. Tahan (Journ. Fed. Mal Stat. Mus. 6, ro15,
p. 127-202).

Cytinaceze and Balanophoracez in Gamble, Materials for a flora
of the Malay Peninsula (Journ. As. Soc. Bengal 75, pt. 4,
_ IQI5, Pp. 393-399).

Vol. IX. (1935).

24

1916
Drimys elongata Ridl. (Hooker, Icon. Plant. 31, 1916, t. 3051).
Aglaia rubra Ridl. (Hook. Icon. Plant. 31, 1916, t. 3052).
Timonius vaccinioides Wernh. (Hook. Icon. Plant. 31, 1916,
t. 3054).
Vittadinia oan, S. Moore (Hook. Icon. Plant. 31, 1916,
t. 3055).

Mischopleura Ridleyana Wernh. and M. ovalifolia Wernh.
(Hook. Icon. Plant. 31, 1916, t. 3059).

Neowollastonia tabernemontanoides Wernh. (Hook. Icon.
Plant. 40,2010, t.. 20001,

Solanum peranomalum Wernh. (Hook Icon. Plant . 31, 1916,
t. 3062).

Eriolopha flagellaris Ridl. (Hook. Icon. Plant. 31, 1916, t. 3067).

New or rare Malayan plants series VIII (Journ. Str. Br. Roy. As.
50.0735. 1916, p. 139-140),

On endemism and the mutation theory (Ann. Bot. 30, 1916, p.
551-574).

Report on the Botany of the Wollaston Expedition to Dutch
New Guinea 1912-1913 (Trans.Linn. Soc. London Bot. ser.
2, 9, 1916, p. I-260).

The natural history of Kedah Peak V. Botany (Journ. Fed. Mal.
Stat. Mus. 7, 1916, p. 37-58).

The large Mias in the Zoological Gardens (Field, July, 1916).
Cultivation of Rubber in Malaya and Burma (Times Supplement,
1916).
1917

New or rare Malayan plants series IX (Journ. Str. Br. Roy As.
Soc. 75, 1917, p. 5-38).

The scientific exploration of the Malay Peninsula (Journ. Str.
Br. Roy. As. Soc. 75, 1917, p; VII—X]).

Botany of Kedah Peak (Journ. Fed Mal. Stat. Mus. 7, 1917, p.
12).

Lasianthus barbellatus, a new species from P. Tiuman (Journ. Str.
Br. Roy. As. Soc: 77, 1017, Pp. roy, 2 ee

Results of an expedition to Korinchi Peak, Sumatra (Journ. Fed.
Mal. Stat. Mus. 8, part 4, 1917, p. 9-145).

Moracee in L. S. Gress, Contribution to the Flora and Phyto-
graphy of the Arfak Mts., etc. 1917, p. 208-209.

Gardens Bulletin, S.S.

25

1918
New or rare Malayan plants series X (Journ. Str. Br. Roy. As.
Soc. 79, 1918, p. 63-100).
Hoseanthus Merr. nov. gen. (Journ. Str. Br. Roy. As. Soc.
79, 1918, Pp. 17).
Appendix II to Gibbs, Some peculiar Papuan customs (Journ.
Str. Br. Roy. As. Soc. 79, 1918, p. 16).

1919

The fern-allies and Characez of the Malay Peninsula (Journ. Str.
Br. Roy. As. Soc. 80, 1919, p. 139-164).

Cultivated Arundinas (Orchid Review, 27, 1919, p. 46).

1920
New Malayan plants (Journ. Bot. 58, 1920, p. 147-149, 195-196).

New or rare plants from the Malay Peninsula (Journ. Fed. Mal.
Stat. Mus. 102, 1920, p. 128-156).

New or rare species of Malayan plants (Journ. Str. Br. Roy. As.
Soc. 82, 1920, p. 167-204).

On a collection of plants from Peninsula Siam (Journ. Fed. Mal.
Stat. Mus. 10, 1920, p. 65-126).

Two new Siamese plants (Journ. Fed. Mal. Stat. Mus. ro, 1920,
p. 127).
Plantago Cynops L. in Kent (Journ. Bot. 58, 1920, p. 127-272).

1921

The Indo-Malayan species of Jussieua (Journ. Bot. 59, 1021,
Pp. 257-260).
Plant dermatitis (Journ. Bot. 1921, p. 310).

Plants from Annam, Scitaminez and Orchideze (Journ. Nat.
Hist. Siam, 4, 1921, p. 109).

1922

The Flora of the Malay Peninsula, Vol. I (Reeve & Co., London,
1922, p. XXxv+Q18).

Rigiolepis and other Vacciniaceze of Borneo (Kew Bull. 1922,
p. 106-108).

The Flora of Klang Gates, Selangor (Journ. Fed. Mal. Stat. Mus.
IO, 1922, p. 247-251).
New or rare Malayan plants series XII (Journ. Str. Br. Roy.
As. Soc. 86, 1922, p. 292-311).
Sandoricum Koetjape and Dendrobium caninum (Journ. Bot.
_ 60, 1922, p. 273-274).

Vol. IX. (1035).

26

Plant Dermatitis (Journ. Trop. Med. & Hyg. 25, no. 14, 1922,
Pp. 225-227).

Musacez, Zingiberacezee and Marantacee in C. T. Wuire, A
contribution to our knowledge of the Flora of Papua (British
New Guinea) (Proc. Roy. Soc. Queensl. XXXIV, no. 1,
1922, Pp. IQ—22).

Note on clearing ponds with Copper Sulphate (Field, May, 10922).

1923

New Euphorbiaceze from the Malay Peninsuta (Kew Bull. 1923,
p. 360-360).

The Flora of the Malay Peninsula Vol. 2 (Reeve & Co., London,
1923 p. vi+672, fig. 76-131).

A Botanical Excursion in Northern Sumatra (Journ. Mal. Br.
Roy. As. Soc. 87, 1923, p. 46-113).

The distribution of plants (Ann. Bot. 37. 37, 1923, p. I-29).

Koetjape and sentol (Journ. Bot. 61, 1923, p. 200-201).

Vanda punctata Ridley (Kew Bull. 1923, p. 118).

1924

The Flora of the Malay Peninsula vol. 3 ( Reeve & Co., London,
1924, p. vi+ 406, fig. 1-159).

New Malayan plants (Journ. Bot. 62, 1924, p. 294-301).

The Flora of the Malay Peninsula, Vol. 4 (Reeve & Co., London,
383 pp., fig. 169-209).

Lotus siliquosus L. in Berks (Journ. Bot. 62, 1924, p. 246).

Decades Kewenses (Kew Bull. 1924, p. 202, 204, 261-262, 263,
266).

Poisoning by Bamboo Hairs (Journ. Trop. Med. & Hyg. 27,
1924, p. 278).

Evodia quercifolia, n. sp. (Gardeners Chronicle, 76, 1924, p. 303).

1925

The Flora of the Malay Peninsula vol. 5 (Reeve & Co., London,
470 pp., fig. 210-229).

Plants of Bencoolen collected by Mr. C. J. Brooks (Kew Bull.
1925, Pp. 76-94).

New Diospyros from Sarawak (Journ. Bot. 63, 1925, p. 5I—52).

Endemic plants (Journ. Bot. 63, 1925, p. 182-183).

Dr. H. O. Forbes’ Malayan plants. Monocotyledones (Journ.
Bot. 63, 1925, Suppl. p. 115-127).

Globba Burbidgei Ridley n. sp. (Journ. Bot. 63, 1925, p. 204).

Decades Kewenses (Kew Bull. 1925, p. 280-281).

Use of Iodine in Beri-beri (Journ. Trop. Med. & Hyg. 28, 1925,
p. 102-103).

Gardens Bulletin, S.S.

27

1926
Spolia Mentawiensia (Kew Bull. 1926, p. 57-04).

The ferns of the Malay Peninsula (Journ. Mal. Br. Roy. As.
Soc. 4, 1926, p. I-12T).

Additions to the flora of Malaya (Kew Bull. 1926, p. 469-479).

Cause of hairlessness in Man (Journ. Trop. Hygiene, Sept. 15,
1926, p. 303).

_ White Blackbird in Kew Gardens (Country Life, Jan. 1926).

Habits of Pangolin (Country Life, Jan. 1926).

1927
The genus Argostemma (Journ. Bot. 65, 1927, p. 25-41).
A correction.—Sonerila ciliata (Journ. Bot. 65, 1927, p. 184).
Rev. W. A. Taylor, M.A. Obituary (Journ. Bot. 65, 1927, p. 317).
Voices of Apes (London Illustrated News, Sept. 17, 1927).
Letter on Rubber Industry (Journal of Heredity, 9, 1027, p. 485).

1928
Cotyledon Umbilicus as an epiphyte (Journ. Bot. 66, 1928, p.
273).
Charles Curtis. Obituary (Journ. Bot. 66, 1928, p. 332-333).
Decades Kewenses (Kew Bull. 1928, p. 72-78).

History of the evolution of the cultivated Rubber industry (Bull.
Rubber Growers’ Ass. 1928, p. 3-7).

1929

New species from the Maley: Peninsula and Borneo (Kew Bull.

1929, p.254—262).
Schefflera minimifiora Ridl. comb. nov. (Journ. Bot. 67, 1920,

p. 87).
H. C. Robinson. Obituary (Journ. Bot. 67, 1020, p. 26r).
Elephant Cemeteries (Times, London, August 5, 1929).
1930
Additions to the flora of Borneo and other Malay Islands (Kew
Bull. 1920, p. 364-373).

The Dispersal of Plants throughout the World. (Reeve & Co.,
Kent, 1930, xx+744 pp., 22 pl).

Myrtaceze malayenses (Journ. Bot. 68, 1930, p. 10-17, 33-39).
Asiatic species of Pleomele (Journ. Bot. 68, 1930, p. 177-181).
Additions to the flora of Borneo (Kew Bull. 1930, p. 74-87).

On the action of the poison of the Upas tree (Antiaris toxicaria
B1.) (Journ. Trop. Med. & Hyg. July, 1030, p.1-2).

Vol. IX. (1935).

28

1931
Additions to the flora of Borneo and other Malay Islands II (Kew
Bull. 1931, p. 33-39).
Additions to the flora of Borneo and other Malay Islands III
(Kew Bull. 1931, p. 493-499).
Decades Kewenses LX (Kew Bull. 1931, p. 448-449).
How Plants move about (The Listener, May 20, 1931).

1932
Taraktogenos calophylla Ridley (Hook. Icon. Plant. 32, 1932,
£. 3167).
Sonerila Ruttneri n. sp., Didissandra parviflora n. sp. (Arch. f.
Hydrobiol. Suppl. Bd. 11, 1932, p. 312-313, 327-328).
The genus Urophyllum in Malaya (Journ. Bot. 70. 70, 1932,
p. IQI-I97, 219-226).

Sir Everard Ferdinand im Thurm. Obituary (Journ. Bot. 70,
1932, DP. 341-342).

Poisonous Nature of Derris elliptica (Pharmaceutical Journal,
129, 1932, p. 387).

1933
G. Ramage. Obituary (Journ. Bot. 71, 1933, p. 75).
Rocks grooved by Ficus roots (Journ. Bot. 71, 1933, p. 75-76).

Additions to the flora of Borneo and other Malay Islands IV
(Kew Bull. 1933, p. 190-202).

Contributions towards a flora of British North Borneo III (Kew
Bull. 1933, p- 487-495).

Fruiting of Passion-flower (Passiflora) in England (Times,
London, March 10, 1933).

1934
Firmiana and Erythropsis (Kew Bull. 1934, p. 214-217).
Some dispersal notes (Journ. Bot. 72, 1934, p. 52-53).
J. D. Gimlette. Obituary (Journ. Bot. 72, 1934, p. 176).
New Malayan Rubiacee (Journ. Bot. 72, 1934, p. 251-257,
271-275).
Dorycnium gracile in Kent (Journ. Bot. 72, 1934, p. 349-350).

Review of Beccari’s Asiatic Palms, Coryphez (Nature, 133, 1934,
Dp. 122).

Notes on insects special protection (Proc. Roy. Soc. London, 7,
1934, p. 634).

1935
Malay violets (Journ. Bot. 73, 1935, p. 13-18).

Gardens Bulletin, S.S.

29

LIST OF PLANTS NAMED IN HONOUR OF
MR. H. N. RIDLEY.

GENERA.
RIDLEYELLA, Schlechter (Orchidaceae).
RIDLEYINDA, Kuntze (Dipterocarpaceae).

SPECIES.
Malay Peninsula:—

Acriopsis Ridleyi, Hook. f. (Orchidaceae).
Acroceras Ridleyi, Stapf (Gramineae).
Actinodaphne Ridleyi, Gamble (Lauraceae).
Actoplanes Ridleyi, Schum. (Zingiberaceae).
Alangium Ridleyi King (Cornaceae).
Alseodaphne Ridleyi, Gamble (Lauraceae).
Amoora Ridleyi, King (Meliaceae).
Ardisia Ridleyi, King & Gamble (Myrsinaceae).
Argostemma Ridleyi, King (Rubiaceae).
Bauhinia Ridleyi Prain (Leguminosae).
Bambusa Ridleyi, Gamble (Gramineae).
Calamus Ridleyanus, Beccari (Palmae).
Castanopsis Ridleyi, Gamble (Cupuliferae).
Cephaelis Ridleyi, King & Gamble (Rubiaceae).
Cinnamomum Ridleyi, Gamble (Lauraceae).
Clerodendron Ridleyi, King & Gamble (Verbenaceae).
Diospyros Ridleyi, Bakh. (Ebenaceae).
Embelia Ridleyi, King & Gamble (Myrsinaceae).
Endopogon Ridleyi, Clarke (Acanthaceae).
Epigynum Ridleyi, King & Gamble (Apocynaceae).
Eria Ridleyi, Rolfe (Orchidaceae).
Eugenia Ridleyi, King (Myrtaceae).
Eulalia Ridleyi, Stapf (Gramineae).
Fagraea Ridleyi, King & Gamble (Loganiaceae).
Filetia Ridleyi, Clarke (Acanthaceae).
Galearia Ridleyi, Gage (Euphorbiaceae).
Genianthus Ridleyi, King & Gamble (Asclepiadaceae).
Goniothalamus Ridleyi, King (Anonaceae).
Gnetum Ridleyi, Gamble (Gnetaceae).
Gymnostachyum Ridleyi, Clarke (Acanthaceae).
Hemigraphis Ridleyi, Clarke (Acanthaceae).
Henslowia Ridleyi, Gamble (Santalaceae).
Homalomena Ridleyana, Engl. (Araceae).
Horsfieldia Ridleyana, Warb. (Myristicaceae).
Hoya Ridleyi, King & Gamble (Asclepiadaceae).
Impatiens Ridleyi, Hook. f. (Balsaminaceae).
Kopsia Ridleyana, King & Gamble (Apocynaceae).
Kunstleria Ridleyi, Prain (Leguminosae).
Lasianthus Ridleyi, King & Gamble (Rubiaceae).
Lecanorchis Ridleyana, Schl. (Orchidaceae).
Lettsomia “Ridleyi, Prain (Convolvulaceae).
Litsea Ridleyi, Gamble (Lauraceae).
Lucinaea Ridleyi, King & Gamble (Rubiaceae).
Meliosma Ridleyi, King (Sabiaceae).
Morinda Ridleyi, King & Gamble (Rubiaceae).

Vol. IX. (1935).

30

Ochlandra Ridleyi, Gamble (Gramineae).

Oryza Ridleyi, Hook. f. (Gramineae).
Pachynocarpus Ridleyanus, Brandis (Dipterocarpaceae).
Palaquium Ridleyi, King & Gamble (Sapotaceae).
Pandanus Ridleyi, Martelli (Pandanaceae).
Pentaphragma Ridleyi, King & Gamble (Campanulaceae).
Piper Ridleyi, C. DC. (Piperaceae).

Piptospatha Ridleyi, Hook. f. (Araceae).
Polyosma Ridleyi, King (Saxifragaceae).

Premna Ridleyi, King & Gamble (Verbenaceae).
Psychotria Ridleyi, King & Gamble (Rubiaceae).
Saprosma Ridleyi, King & Gamble (Rubiaceae).
Schefflera Ridleyi, Viguier (Araliaceae).
Schismatoglottis Ridleyana, Engl. (Araceae).
Scleria Ridleyi, Clarke (Cyperaceae).

Scleropyrum Ridleyi, Gamble (Santalaceae).
Sclerostachyum Ridleyi, Stapf (Gramineae).

Shorea Ridleyana, King (Dipterocarpaceae).
Spatholobus Ridleyi, Prain (Leguminosae).
Stenothyrsus Ridleyi, Clarke (Acanthaceae).
Strychnos Ridleyi, King & Gamble (Loganiaceae).
Styrax Ridleyanum, Perkins (Styraceae).
Symplocos Ridleyi, King & Gamble (Symplocaceae).
Tarenna Ridleyi, Pearson (Rubiaceae).

Tylophora Ridleyi, Henders. (Asclepiadaceae).
Uvaria Ridleyi, King (Anonaceae).

Wikstoemia Ridleyi, Gamble (Thymeleaceae).
Xylopia Ridleyi, King (Anonaceae).

Xyris Ridleyi, Rendle (Xyridaceae).

Ferns:—

Alsophila Ridleyi, Baker.
Athyrium Ridleyi, Copeland.
Lastrea Ridleyi, Beddome.
Microlepia Ridleyi, Copeland.
Platycerium Ridleyi, Christ.
Trichomanes Ridleyi, Copeland.

Borneo:—

Coelogyne Ridleyana, Schltr. (Orchidaceae).
Dendrobium Ridleyi, Merr. (Orchidaceae).
Dioscorea Ridleyi, Prain & Burkill (Dioscoreaceae).
Panicum Ridleyi, Hack. (Gramineae).

Pollinia Ridleyi, Hack. (Gramineae).

Raphidophora Ridleyi, Merr. (Araceae).

Sarcanthus Ridleyi, J. J. Smith (Orchidaceae).
Saurauia Ridleyi, Merr. (Ternstroemiaceae).
Thoracostachyum Ridleyi, Clarke (Gramineae).

Siam:—

Dendrobium Ridleyanum, Kerr (Orchidaceae).
Hedyotis Ridleyana, Craib (Rubiaceae).
Ophiorrhiza Ridleyana, Craib (Rubiaceae).

.

Gardens Bulletin, S.S.

Mr RIDLEY and his collector ACHMAD BIN HASSAN in the Gardens Jungle, about 1909.

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MR. RIDLEY’S WORK FOR TROPICAL AGRICULTURE.

By F. W. SOUTH,
Chief Field Officer, Department of Agriculture S.S. & F.M.S.

Introduction.

Mr. H. N. RIDLEY, C.M.G., F.R.S., was Director of Gardens,
Straits Settlements from 1888 to 1911 and for the period 1888
to 1894 was also Director of Forests, Straits Settlements.
Probably but few people at the present time realise how great
a debt Malaya as a whole owes to Ridley for his services in
laying a sure foundation for that agricultural development
which subsequently brought so much prosperity to this country.

The work which Ridley did during his period of service in
Malaya was only rendered possible by his tireless energy and
enthusiasm and his wide knowledge of biology, including both
botany and zoology. He specialised more particularly in botany,
including in that term pure botany as a sience and botany as
applied to forestry and agriculture, but his knowledge of
zoology enabled him to render further services to agriculturists
by devising measures for the control of insect pests of various

- crops.

For about twelve years of his service Ridley with his staff
of two or three European Officers serving as Superintendents or
Assistant Superintendents of Gardens and Forests in Singapore,
Malacca and Penang, represented the only organisation in the
Colony or the Native States, as they were then known, to which
agriculturists could apply for technical advice and assistance.
The need for such assistance naturally increased steadily through-
out this period, since it covers the early extension of large scale
agriculture in the Malay States. When it is remembered that the
whole period of his service corresponds closely with that of the
growth of the rubber industry in this country from its earliest
infancy to the great boom of 1910-1911, it becomes possible to
realise how great were the calls made on Ridley and his staff.
The demands from the agricultural comtnunity for planting
material of Para rubber and for advice on every aspect of this new
and quite unknown crop occupied in themselves a large share
of the time of Ridley and his assistants.

In spite, however, of the demands of the growing rubber
industry and the claims of the more purely scientific aspect of -
botany as represented by the maintenance of the Botanic Gardens
in Singapore and the collection, identification and description of
local plants, Ridley was able to carry out work on a range of
plants which comprised most of the actual or potential tropical
crops known.

Vol. IX. (1935).

32

Ridley’s work on rubber is dealt with in a separate article in
this number, so that the remarks which follow deal only with
some of his work on other crops.

THE ECONOMIC GARDEN.

In 1888 when Ridley assumed duty there was in Singapore
under the Forest Department, an Experimental or Economic
Garden near the Botanic Gardens, in which experiments were
in progress with English vegetables and fruit trees of various
kinds, together with trials of patchouli, coca (the plant from
which cocaine is obtained) cubebs peper, ordinary black pepper,
cacao, tapioca and various trees yielding rubber. In 1889 Ridley
recommended that this Experimental Garden should be removed
from the Forest Department and be placed under the Gardens
vote. In making this suggestion he wrote :—

“I hope to increase the area under cultivation this year
and, wherever I can, to make the whole an Economic Garden
worthy of the Colony’’.

In the following year this transfer was effected.

In the years which followed Ridley gradually increased the
size of this Garden and added to the collection of economic
plants which it contained, until in his Annual Report for 1906
he was able with full justice to write of it :—

‘This part of the Gardens has steadily increased in
usefulness and importance, not only to the local community
but to all the tropical parts of the Empire, and indeed it
would be difficult to over estimate its utility. All tropical
plants are experimented with and stocks of all for which
there is or is likely to be an exceptional demand are
maintained in considerable quantities. It may give some
idea of the work carried out if it is mentioned that during
the year plants or seeds have been despatched on sale to
countries as remote as the West Indies, including Cuba,
Samoa, Honolulu, Queensland, New South Wales, West
Australia, Fiji Islands, Philippines, New Guinea, China,
Hongkong, Borneo, Sumatra, Java, India, Ceylon, Egypt,
Lagos, Nigeria 4nd Madiera, while the local demand from
the Colony and Federated Malay States has increased
enormously’’.

Two of the objects of this Garden are clearly stated in the
paragraph quoted, namely to experiment with all known tropical
crops, or tropical plants likely to prove of economic value, more
especially as regards their suitability for cultivation locally, and
to provide planting material of those for which there was a
demand. It also served a third purpose by providing demons-
trations to visiting planters of the alternative or new crops in
which they were interested.

Gardens Bulletin, S.S.

33

In addition to the Economic Garden in Singapore there were
in the earlier part of this period two other localities in the Colony
where work on economic crops was conducted.

One of these was the Bukit Sebukor Experimental Garden in
Malacca. This was under the Forest Department and was used
partly as a nursery for forest trees and partly for experiments with
agricultural crops such as Mauritius hemp, tobacco, castor oil,
pepper, Maltese oranges and lemons, cloves, nutmegs, Liberian
coffee, tea, cacao and Egyptian cotton. The cotton and cacao
were not successful. Liberian coffee was considered to require
manuring on poor soil. The tea, a Hybrid Assam type, did fairly
well, as did the nutmegs and cloves on the better soil in this
garden. In 1893, however, on the recommendation of the
Retrenchment Committee of that year this garden was abandoned,
partly because much of the soil on this site was very poor.

Another locality was the Hill Nursery and Government
Bungalow Garden on Penang Hill. The Nursery was used for
trials of fruit trees and it is mentioned in the Annual Report for
1888 that an apple tree from Australia bore 8 fruits of large size and
fair flavour, but olives produced no fruit. In 1891 an avocado
pear tree in this Nursery fruited for the first time and in 1894
three of the same trees bore a heavy crop. Lechee, loquat,
olives and peaches do not appear “to have produced fruit and
oranges from Malta gave a few fruits, but of moderate flavour and
not in economic quantities. The bungalow garden was used for
raising vegetables of which, according to the Report for 1894,
carrots, beet, leeks, lettuce, khol rabi, radish, peas and in the
dry season tomatoes were the most successful. In 1898 potatoes
gave a fair crop but several were diseased. The great handicap
on the Hill was the expense of farmyard manure when the
charges for transport, in the absence of a a road or railway, were
added, while another expensive item was the construction of a
terrace round each fruit tree to prevent soil erosion by heavy
rains. Little work was done in this fruit nursery after 1897
and an attempt was made to reafforest the site with useful trees
in 1900. The Bungalow Garden was, however, regularly upkept
and continued to give a supply of vegetables.

The Waterfall Gardens in Penang provided little room for
work on econmic crops, but well known tapping trials on one of
the rubber trees in that Garden were commenced in 1898 by the
Assistant Superintendent, Mr. Curtis, and were maintained
for at least eleven years. Other economic activities at this
garden were mainly confined to raising young plants for trial
on estates in the island or in Province Wellesley. Thus in 1896
Mr. CurTIs raised a fairly large number of Borneo sugar canes
from seed and in the following two years distributed them for
trial on estates. From 1895 to 1897 a stock of Ramie fibre plants
was built up for distribution and some interesting observations
on varieties of this plant were made. In 1903 and 1904,

Vol. IX. (1935).
c

34

Mr. Fox, then Superintendent, carried out experiments with
Egyptian and Sea Island cottons. These experiments in con-
junction with those carried out on estates in the Settlement led
him to conclude that in the cultivation of cotton the Straits
Settlements could not compete with other cotton growing
countries.

In the Malay States, Sir HuGH Low planted Hevea rubber
trees a few other economic trees behind the Residency at Kuala
Kangsar in 1878 and following years, but there was no Experi-
mental or Economic Garden of much size or importance until
Mr. ARDEN, Superintendent of Experimental Plantations,
Selangor, commenced the establishment of the Batu Tiga
Plantation towards the end of 1902.

In these circumstances the Economic Garden in Singapore
was the chief source of information on various crops and of
planting material for the whole Peninsula, until such time as the
Department of Agriculture, Federated Malay States, created in
1904, had developed sufficiently to take its share of this work.

Examination of the Annual Reports of the Botanic Gardens,
Singapore, shows how wide a range of plants was tried and
reveals several points of interest.

Throughout the period of Ridley’s tenure of office there was
a demand, varying somewhat from time to time, for planting
material of most of the well known local crops, such as pepper,
cloves, nutmegs, tapioca, gambier and local fruit trees. These
call for little comment beyond the fact that six good varieties of
tapioca from British Guiana were introduced in 1888.

Until the cultivation of Para rubber began to monopolise
attention in the first decade of this century, a number of plants
were tried in the Economic Garden with a view to ascertaining
those which would be most suitable for use in large scale agri-
cultural development in the Peninsula. Beverages included a
collection of varieties of cacao obtained from Ceylon in 1883,
a collection of different species of coffee assembled gradually,
and Assam, China and Paraguay tea planted in 1891. In 1888
the cacao fruited well and it was stated that there was no reason
for its failure if properly cultivated, and that it might be pro-
fitable in the Peninsula where the soil was better than in
Singapore. In later years there was some demand for planting
material for trial, but it never seems to have proved entirely
satisfactory. In the nineties there was a considerable demand
for liberian coffee for the estates in Selangor and Perak and some
demand towards the end of that decade for Stenophylla. It is
peculiar that so little interest seems to have been taken in the
possibilities of tea cultivation during these years, seeing how
promising it has shown itself to be in the last few years on both
the plains and hills of this country.

Gardens Bulletin, S.S.

one

35

The collection of fibre plants assembled in this Garden
included Ramie, Cottons of various kinds, Manila hemp (Musa
textilis), various species of Samsevieras (Bowstring hemp),
Mauritius and sisal hemp, Fourcroya gigantea imported from
Barbados and some Bromelias. Ramie cultivation attracted
considerable attention up to about 1904 and Ridley ‘had
hopes that the cultivation of this and some other fibres would
gradually come to occupy the definite place in the agriculture of
the country which he considered that they deserved, but the
overwhelming importance of rubber caused them to be neglected
in later years. In the years 1903 and 1904 shortage of the cotton
crops in America emphasised the importance of this cultivation
throughout the Empire. Numerous varieties of this plant were
imported but did not prove successful. Ridley states :—

““‘The climate and soil of Singapore seems, however,
quite unsuited for cotton culture, owing to the excess of
rain, and the red cotton bug, Dysdercus cingulatus, in spite
of the use of insecticides, destroyed nearly all the pods’’.

This conclusion was supported by that of the trials in Penang,
referred to above, and agrees with the results of trials conducted
in more recent years by the Department of Agriculture, S.S. &
F.M.S., so far at least as the western side of the Peninsula is
concerned. Mauritius hemp did well, but sisal was not a success.
Both these plants grew satisfactorily in trials during the last
ten years at the Central Experiment Station of the Department
of Agriculture in Selangor, but they gave little prospect of pro-
fitable cultivation except at high prices for the fibre. The
Sansevieras never became of much importance, but it is
still possible that Manila hemp may prove a useful crop on
certain types of soil in this country.

As early as 1888 the annual value of the preserved fruit
export from the Colony, chiefly preserved pineapples, was
$100,000, and in view of this Ridley imported in 1892, 1893, and
1895 better kinds of pineapples which included hot-house
varieties from Kew and Windsor Castle and West Indian strains
from Trinidad and Jamaica.

Various. other kinds of fruits were imported and tried. It
is impossible in the space available to enumerate these, but a few
records of interest may be mentioned. Figs fruited in 1888 and
the following year, but the fruits were of poor flavour. In the
first of these years the tree tomato was a failure in Singapore,
although it has in recent years grown and fruited well on the
hills of this country. The avocado pear fruited well in 1802
and more seeds from Kew and Trinidad were imported. In
view of its success both in Singapore and on Penang hill it is
remarkable that even now this fruit, so popular in many parts
of the tropics and sub-tropics, is still comparatively rare and of
little account here. Mangoes from Madagascar and grafted
kinds from Calcutta were imported in 1903, but the grafted kinds

Vol. IX. (1935).

30

of this fruit have so far never done well in this evenly moist
climate. In 1890 Maltese lemons fruited well, giving fruits of
good size and flavour, but even to the present day this fruit has
attracted but little attention.

Summarising the results obtained with English vegetables
in 1888, Ridley states that Tomatoes and Jerusalem artichokes
did well, while turnips, onions and carrots gave fair results,
though the flavour of the carrots was poor and they were apt to
be small and stringy. In 1889 he mentions the successful
cultivation of water-cress.

In 1907 when rubber was beginning to secure its place as the
main crop, Ridley received numerous enquiries for suitable catch
crops for planting in the young clearings. He was not able to
give this matter the attention which he felt that it deserved, but
in the following year he mentions that Ground-nut, Citronella
and Lemon grass were in demand for this purpose.

Although in the early days of rubber planting the fetish of clean
weeding held away, Ridley in 1909 planted up plots of crotalarias,
tephrosia, desmodiums cajanus, clitoria and other leguminous
plants which he considered suitable in various degrees for
covering and improving the soil of rubber and other estates.

In 1907 an article in the ‘‘Agricultural Bulletin’”’ called
attention to the value of the Oil Palm as a cultivated crop and
resulted in the distribution of 3,030 seeds. Substantial develop-
ments did not follow until some 10 more years had elapsed, but
this record may well mark the beginnings of what is now a well
established industry.

Considerations of space forbid reference to a number of other
plants of various kinds grown in the Economic Garden in
Singapore, but the following taken from the report for 1907
gives a picture of the extent of the planting material distributed :

Para Rubber seeds 405,600 and 13,100 plants, Ramie 3,170,
Sanseviera 10,000, Manila hemp 130, Mauritius hemp 150, fruits
of various kinds 1,700 plants and 12,000 seedlings, Tapioca 300
plants, Coffee 125, Citronella 160, Lemon grass 250, Camphor
750, Cocoa 160, Nutmeg 172, Oil Palm 3,030 seeds, Ground nuts
50 lbs. seeds. Most of these were sold to the Federated Malay
States and Johore, but some went to Lagos, British New Guinea,
Southern Nigeria and the Caroline Islands.

PESTS AND DISEASES OF CROPS.

Early in his period of office Ridley drew attention to the
damage caused to coconut palms by the black beetle (Oryctes
rhinoceros) and the red beetle, as it was then known, (Rhyncop-
horus sach). The outcome of this was the passing in July, 1890
of the Coconut Trees Preservation Ordinance. Asiatic Inspectors
were appointed in Singapore under the Director of Gardens and
in Penang Settlement under the Superintendent of Gardens to

Gardens Bulletin, S.S.

37

ensure that all suitable breeding places for these insects were
cleared up. Similar legislation was provided about the beginning
of this century in the Federated Malay States and until 1913 was.
administered by the Inspector of Coconut Trees and his staff.

In 1896 Ridley published a bulletin on the disease of nutmegs
in Penang due to a Scolytid beetle and another on diseases of
coffee. In 1899 he visited the districts in Selangor which were
affected by the plague of the coffee caterpillar (Cephonodes
Hylas) investigating its life history and trying experiments for
its control. He also investigated and gave advice on certain
disease of rubber and the white ant (Termes gestroi) attacking
rubber and coconuts.

Among other pests and diseases recorded by him were Sireh
disease on sugar cane in 1889 and in 1898 a borer caterpillar
(Chilo spp.) and red smut fungus on the same crop, while there
is also a record in the same year that the beetle Xylotrupes gideon
damaged canes in Province Wellesley by eating the roots. In
that year damage to coffee by a locust (Cyracanthus nigrovaria,
Walk.) is noted and also by a fungus believed to be a species of
Irpex. The Banana weevil (Sphenophorus sordidus) was
recorded in 1899 together with the rose sawfly and a species of the
fungus Rosellinia on the roots of a species of Ficus. In
1900 there is a note on the beetle, Batocera octomaculata, boring
in the branches of Rambong (Ficus elastica); the life history of
the caterpillar, Calogramma festiva, a Noctuid moth was worked
out; and a caterpillar pest proved very destructive to young
plants of Getta Percha.

In this connexion it is interesting to note that in 1909 Fox
records that coconut palms on Malakoff Estate in Province
Wellesley were badly attacked by the larve of a small moth
and that for a few months the trees presented the appearance of
having been burnt. Judging from this description it would
seem that this constitutes an early record of an outbreak of the
pest Artona catoxantha.

AGRICULTURAL BULLETIN.

It was necessary for Ridley to find some means, additional
to ordinary correspondence and personal visits, for conveying to
the planting community information resulting from his various
investigations and experiments. For this purpose he issued
between 1895 and 1900 Agricultural Bulletins on various:
subjects, which were published as information became available.
In October, 1901 he commenced to issue monthly ‘“The Agri-
cultural Bulletin of the Straits and Federated Malay States,’’
which he continued to edit until his retirement in 1911. This
was supported by numerous contributors and correspondents
including officers of the Forests and Agricultural Departments
established in the Federated Malay States during those years, as

Vol. IX. (1935).

38

well as several leading planters, but Ridley himself was always
a regular contributor in addition to carrying out the considerable
task of editor. This publication contains a mass of interesting
and valuable information on a number of crops besides mibber,
and shows, as do his other activities, that Ridley was no believer
in placing all the agricultural eggs in one basket.

HorRTICULTURAL AND AGRICULTURAL SHOWS.

Another well recognised method of propaganda utilised by
Ridley and his staff was the holding of Horticultural and Agri-
cultural Shows. A Flower Show, often including frit and
vegetables, was held in Singapore in 188q and the following
two years, but was then discontinued. A successful Agri-
Horticultural Show was held in Penang in 1893; another at
Butterworth in 1899; and again in Penang in 1901. Im 1904 a
Show was held in Kuala Lumpur, but what was then described
as probably the largest Exhibition of its kind ever held in the
East took place in Singapore in 1906 and was attended by
visitors and delegates from Java, India, Ceylon and Burmah.
This was followed by Annual Shows at different centres such as
Kuala Kangsar and Kuala Lumpur until 1910 when the last of
the continuous series was held in Singapore.

It is hoped that the above account will give some conception
of Ridley’s multifarious activities on behalf of Agriculture in
Malaya.

Since the days with which this article is concerned, the
development of agriculture in the Federated Malay States,
which resulted in the creation in 1904 of what is now the Depart-
ment of Agriculture, Straits Settlements and Federated Malay
States, and the establishment in 1926 of the Rubber Research
Institute, has caused the transfer of agricultural investigation
work from Singapore to Kuala Lumpur and as a result the
Economic Garden in Singapore has been abandoned. Ridley
himself lives far away, enjoying a well earned retirement in
England. But in spite of these changes the agricultural structure
which has arisen on the foundations which he so securly laid
remains as his permanent memorial in Malaya.

Gardens Bulletin, S.S.

39

# MR. RIDLEY AND RUBBER IN MALAYA.

By B. J. Eaton,
Director, Rubber Research Institute of Malaya.

As the name of the late Mr. (later Sir) HENRY WICKHAM is
synonymous with the introduction of the rubber tree to the eastern
tropics, so the name of Mr. RIDLEY is synonymous with the
establishment of the Para Rubber Tree (Hevea brasiliensis) as a
commercial plantation product in Malaya.

An anonymous article (probably by Ridley) published in
the Agricultural Bulletin of the Straits and Federated Malay
States (1903) gives the history of the introduction of the Para
rubber tree into the Malay Peninsula. A few extracts from this
article will therefore be appropriate in an appreciation of the
work of Ridley in connection with rubber cultivation in Malaya.

On August 11th 1876 fifty plants derived from seeds received
from Wickham on June 14th 1876 and grown in the Kew
Gardens were sent to the Botanic Gardens Singapore. Owing
to delay in payment of freight these plants all perished.

On June 11th 1877, twenty two plants were sent to the
Singapore Gardens. In October of the same year Mr. MurRTON,
Superintendent of the Gardens, planted nine of them in the
Residency Gardens, Kuala Kangsar, F.M.S.

In a report dated February 3rd 18790 Mr. HuGH Low (then
British Resident of Perak) writes ‘‘The Heveas are now 12-14
feet high. They take to the country immensely’’.

In a letter dated December 11th 1896 Sir HuGH Low writes
to the Kew Gardens ‘‘As I am writing I should like to mention
that the Hevea brasiliensis which having received from Kew
through Singapore I planted at Kuala Kangsar in Perak grew
magnificiently and fruited I believe 2-3 years before those of
Ceylon. I distributed the seeds to various places in the neigh-
bourhood and they are now to be found in Mr. Hm’s Coffee
Gardens in various parts of the peninsula and in several places
in Perak.

“When Mr. SWETTENHAM was at home in the summer I
enquired of him as to their condition and found they were not
thought to be of any value as some Dyaks had tapped some of
the largest trees and found that scarcely any juice exuded from
them (Note:—The effect of ‘‘wound response’’ on yield had not
then been discovered). This unfortunate statement seems to
have deterred Perak planters from paying any attention to Para
rubber for some time’’. In 1882 other seeds obtained by
Sir HucH Low were planted by Mr. Wray at Kuala Kangsar.

Vol. IX. (1035).

40

In 1881 Mr. CantLEy of the Botanic Gardens Singapore
writes ‘‘the tallest Hevea (in the Gardens) is now 25 feet tall and
14 inches round the base. These trees commenced to fruit in
isp2.”°

Later seeds were received in large quantities from Ceylon.

It was not until 1897 that the high price of rubber and low
price of coffee stimulated the interest of planters (Note:—Had
interest been aroused earlier and extensive planting taken place,
the extraordinary boom of r910 due to the rapid development
of the motor industry would probably not have occurred).

Mr. RrpLEy served as Director of Gardens Singapore from
1888 until 1911. During most of this period—from 1888 until
1905—he was largely responsible for advising on agriculture
throughout the peninsula, although during the later part of this
period in the Federated Malay States, two Superintendents of
Plantations in Perak and Selangor and a Federal Inspector of
coconuts were employed by the Government before the inaugura-
tion of the Department of Agriculture F.M.S. in 1905.

Ridley thus remained in Malaya sufficiently long to see the
fruition of his labour in regard to the commercial exploitation
of the rubber tree (Hevea brasiliensis) in the extraordinary boom
in rubber prices which reached its culminating point in 1910 and
which was responsible for the subsequent extensive development
not only in Malaya but also in the Netherlands Indies.

The writer of this article need make only a passing reference
to the many-sided nature of his work in connection with other
economic crops and in relation to his systematic work on the
flora of Malaya, since this is being dealt with by others.
Details of his work on numerous economic crops are contained
in Annual Reports of the Botanic Gardens and Forests Depart-
ment, S.S., of which he was also Director for some years and in
the Agricultural Bulletin, S.S. which he initiated and edited for
many years.

During the period prior to 1905 he was responsible for the
distribution of an enormous amount of rubber planting material,
seedlings and seeds from the experimental area planted in the
Botanic Gardens Singapore, which formed the beginning of the
oldest plantations in the peninsula. ‘This activity is particularly
noticeable from the years 1897 onwards. During this period
also various systems of tapping and methods of coagulation were
experimented with under his direction.

He also realised the possibility of utilising the seeds of the
rubber tree as a source of oil, although for various reasons the
marketing of seeds for this purpose has not developed on a
commercial scale.

In 1903 he carried out certain manurial experiments with
rubber plants in beds and showed the value of cattle manure in

Gardens Bulletin, S.S.

AT

improving growth. Experiments under his direction also
indicated that early morning and evening tapping gave the best
results. .

In 1904 rubber seeds from the Botanic Gardens were sent
to Burma and in the following year to Jamaica, Lagos and
Nigeria. It is perhaps fortunate that these did not develop
successfully in various countries to which they were despatched.

One interesting fact emerges from Mr. RipiLEy’s Annual
Report for 1909 which is probably not known generally, viz. that
the Government grant for the upkeep of the gardens was only
sufficient to keep them in a respectable condition and was
insufficient for tools, stores and improvements, so that the
revenue derived from the sale of rubber and rubber seeds was
used for this purpose. He had indeed to bemoan the fact that by
1909 plantation seed had become available—thanks to his
distribution of seed in previous years—and was able to compete
with his own sales. On this account he was seriously considering
concentrating on running his rubber area as a plantation in order
to make sufficient revenue from sales of rubber to enable him to
continue improvements in other directions. It is obvious there-
fore that “‘retrenchment’’ is by no means new in the history of
Malaya.

In this year also the records show that one tree in the
Penang Gardens tapped every year (not continuously as at
present) since 1885 had yielded approximately 52% lbs. over
the period.

It is also reported that ‘‘the success of this marvellous
product reached a height little dreamt of a few years ago, prices
well over 9 (shillings) a pound being realized’’. In spite of this
however it is amusing to note in the report of the late Mr. Fox
who served under Ridley in the Penang Gardens, that a small
parcel of rubber sent home in October was withheld as the sales
were unfavourable ! !

The honour of a C.M.G was conferred on Mr. RrpiEy on his
retirement from Malaya and he was also elected a Fellow of the
Royal Society for his botanical investigations.

In addition to continuing his publications on the flora of
Malaya, he is still a member of the Technical Sub Committee
of the London Advisory Committee for Rubber Research
(Ceylon & Malaya) and thus continues at the age of 80 to
maintain his interest in an industry of which he may be said to
have witnessed the birth. Those of us who knew his personally
as a fellow scientist and friend join in wishing him Many Happy
Returns of the birthday which he celebrates this month. We
in Malaya, not to mention other rubber growing countries in the
East owe him a considerable debt of gratitude for his pioneer
work in connection with the plantation rubber industry and his
optimism and faith in its future.

Vol. TX. (1035).

42

MR. RIDLEY AND FORESTRY IN MALAYA.

It is never an easy matter to assess the value of the services
of a biologist in any particular field, and when the boundaries of
the fields are as obscure as those that surround forestry and
agriculture, differentiation becomes impossible. Nearly every
issue of the official organ of the Indian Forest Service the
Indian Forestey————contains articles on crops that nobody in
this country would connect with forestry, but which are regarded
as coming within the purview of the Forest Department in India.
Coming nearer home there is the case of rubber. Rubber started
as a forest crop, but general responsibility for government acti-
vities and experimental work was, for purposes of convenience,
subsequently vested in the agricultural authorities. Are Ridley’s
attainments in this field to be labelled forestry or agriculture?
It may be supposed that Ridley was not particularly concerned at
the time with the labels on the pigeon-holes in which posterity
intended to docket the records of his activities, and we do not
think that he will worry a great deal now if some of them have
got a bit mixed.

That he was alive to the importance of maintaining supplies
of timber goes without saying, and it is to Ridley that we owe the
constitution of reserves in Malacca, whose forests would have
been in danger of total dissolution during the rubber boom but
for his foresight. The same applies to Penang, for aesthetically
spoiled though that delightful settlement may be from the clearing
of hill forest, it would have been completely ruined had reserva-
tion not put a stop to the (mostly unremunerative) efforts
of cultivators on the hill sides. In Singapore, the reserves are
of less importance and have been virtually abandoned. When
Ridley arrived the damage had already been done, and the areas
available for reservation were, for the most part, already past
praying for as forests, though possibly worth considering for
reafforestation, an aspect of forestry that loomed very much larger
in the Straits when comparative ignorance prevailed with regard
to the resources of the interior and communications were chiefly
remarkable by their absence. Such plantations as were formed
were, for the most part , confined to denuded tracts of country,
and except for a patch of quite fair mahogany and some unhappy
line plantings of high forest forms that continue to receive a
certain amount of attention at Bukit Beruang in Malacca, little
trace remains of these early efforts.

In 1895 Ridley paid a visit to Selangor and reported on the
forests of that state which, at that time, were in nominal charge
of the Land Officers, and it was largely due to his report that the
first Forest Officer, Mr. B. H. F. Barnard was appointed in that
State. He subsequently accompanied Mr. H. C. Hill, who was
seconded from the Indian Forest Service to report and advise on

Gardens Bulletin, S.S.

43

the forests of the F.M.S., a step which led to the appointment of
Mr. A. M. Burn-Murdoch and the establishment of the Forest
Department as a combined Federal and S.S. organization in 1goo0.

So much for his direct activities. Valuable though they
undoubtedly were they were as nothing compared with his work
in the field of systematic botany and plant economy. ‘Thanks
chiefly to Ridley, the Malayan forest organization was able to start
off with immeasurably more knowledge of the composition of
the forests than is the common lot of tropical foresters generally.
Without his Flora of the Malaya Peninsula and Malay Plant
Names most of our Forest Records would have been very incom-
plete or might never have been written. Though essentially a
systematist he never allowed himself to be tied to the
herbarium, and his contributions to the Agricultural Bulletin
included invaluable field notes on timbers and on the uses and
sources of all kinds of minor forest products. That there were
inaccuracies goes without saying, but it was just that quality of
recklessness in Ridley’s work that enabled him to turn out such
vast quantities of assimilable knowledge when it was most needed.
A lesser man would have feared for his professional reputation,
but Ridley was above such petty considerations. The work
needed doing, and with the means at his disposal he did it
magnificently.

Vol. IX. (1935).

44

y
Mr. H. N. RIDLEY AND THE FLORA OF THE
NETHERLANDS INDIES.

By D. F. VAN SLOOTEN,
.

Acting Director, Botanic Gardens, Buitenzorg.

The series of island-groups, which extend from further India
to Australia, and which are situated between 95° and 141° east
longitude and between 6° north and 11° south of the equator,
belong to the overseas territory of the Kingdom of the Nether-
lands, with the exception of the Philippines, Sarawak and British
North Borneo, Portuguese Timor, Christmas Island and the
Territories of British New Guinea. If one,.compares the extent
and area of the Netherlands East Indies with those of Europe
or the United States of America, one finds that the east to west
extent of the N.E.I. is approximately equal to the distance from
the West Coast of Ireland to the East Coast of the Caspian and
from 7° to the West of San Francisco to 7° to the East of New
York, the maximum length from West to East of the Netherlands
East Indies (i.e. from Pulau Weh to Humboldt Bay) being about
5,000 km. The maximum distance from North to South is
2,000 km., which is equal to the distance between Oslo and
Rome and as far as from the North coast of Michigan Lake to
New Orleans (Louisiana).

The total area covered by all the islands of the Netherlands
Indian Archipelago measures 1,900,152 km.?, which equals about
half the size of Europe, excluding Russia. In order to give
some idea of the size of the islands we mention the following
details: Java, including Madoera, has an area of 131,508 km.?
(i.e. about the same size as the State of New York), Sumatra
with the surrounding islands an area of 420,384 km.” (exceeding
the area of Great Britain), Borneo, as far as the Dutch territory
is concerned, covers an area of 553,341 km.? (which is as large as
France), Celebes and the small islands in the immediate
neighbourhood an area of 185,914 km.? (being about the same
size as New Zealand and Ceylon taken together) and the Dutch
section of New Guinea one of 397,204 km.? (viz. the same area
as Japan proper), while the rest of the Archipelago measures
211,801 km.?

The botanists, who have to study the vegetation and the
immensely rich flora of an area covering approximately 1,900,152
square kilometers, have to perform—as everybody will under-
stand—a very difficult and lengthy task. Though the climatic
conditions (uniformity of temperature and changes in rainfall,
both intensified by the high mountain ranges which intersect the

Gardens Bulletin, S.S.

45

islands) are very similar throughout this vast tropical area, so
that the number of vegetation types is very limited, yet from the
differences in the local soil conditions (water, edaphic factors,
voleanic material, etc.) has arisen such a complex flora, that
the very same vegetation-type shows in various places a large
diversity as to the constituent flora-elements.

The total flora of flowering plants may confidently be
estimated to comprise 30,000 species, belonging to about 2,500
genera. It will be many years yet before all islands of the
Archipelago are intensively explored botanically and before such
a wonderfully rich flora is thoroughly known, the more so as
the flora of each island presents a small or high percentage of
endemic species. This flora could not form a subject of systematic
investigation until the foundation of the Botanic Gardens at
Buitenzorg in 1817. Its study was taken up energetically by
the second Director of the Gardens, C. L. BLuMe, and has
been continued up till the present.

This study is progressing but slowly, on account of the
vastness of the territory, and also of the fact that during many
years this task was entrusted to a few botanists only. Moreover,
in Bu.tenzorg the standpoint is taken, that our knowledge of the
flora ought to be based on a thorough monographic study of the
families; this has found expression in contributions to the
Bulletin du Jardin Botanique de Buitenzorg. ‘To further this
object, contact has long been developed with botanists from
abroad for co-operation in the study of the Malaysian flora. The
researches of those botanists who concern themselves with the
study of a closely related flora, are therefore of direct importance.
Many individuals are cited in literature as having studied more or
less botanical material from this Archipelago. Among the most
prominent of these is Mr. H. N. RIpiey.

» Mr. Rip ey’s course of life once more proves how profitable
it is for men of outstanding ability to be employed in the tropics
for a considerable period of years; in this way important work
can be accomplished without interruption and along the same
lines. Well-known examples of this are the celebrated RUMPHIUS,
‘TEYSMANN and TREvp, the last two of whom by their very long
service in the tropics were able to found and consolidate the
Botanic Gardens at Buitenzorg.

It has been given to Mr. RImpLeEy also to spend a long period
in the tropics, as a result of which he became one of the founders
of our knowledge of the flora of the Malay Peninsula. Being
evidently very zealous in fulfilling his botanical duties, he based
the study of this flora not only on herbarium specimens, but also
himself collected large numbers of plants during inspection trips.

Vol. IX. (1935).

46

His large collections, of which he deposited duplicate sets in
many herbaria, including Buitenzorg, form, with other older
ones, the essential basis of our knowledge of the exceedingly rich
flora of the Malay Peninsula. He did not confine his trips to
this Peninsula, but he also made expeditions to the Malay
Archipelago, viz. to Christmas Island (1890 and 10904), Java
(1915), Sumatra (1897 and 1921) and Borneo, as a result of which
he was in a position to compare the floras of these regions. So
far as the Netherlands Indies are concerned, it is only from
New Guinea that he has studied an important collection of plants
without having visited the country of origin. The number of
species he named as new to science amounts to a few thousands;
the number described by him from the Netherlands Indies
(including British North Borneo, Sarawak and British New
Guinea) so far as I have been able to trace—is 1,053, divided
over the different islands as follows: British New Guinea 391,
Sumatra and surrounding islands 329, Sarawak British North
Borneo and Dutch Borneo 322, Java 6, Celebes 2, Ambon, Key
and Aru Islands each r.

Now it may be true that the majority of these plants
described as new by Mr. RIDLEy originated from outside the
Netherlands Indies; but phytogeographically the flora of the
Malay Archipelago appears to be so closely related to that of the
Malay Peninsula, that the systematic work of RIDLEY implies an
important progress for our knowledge of the flora of the Malay
Archipelago also.

Already before he became Director of the Botanic Gardens
at Singapore, viz. in 1884, he published in the Journal of Botany
some articles about the flora of the Malaysian Archipelago; the
description of a new Bornean orchid (a new diagnosis of Didy-
moplexis sylvatica (Bl.) RrpLEy) belonging to the Javan collection
of HorsFIELD, formed the first contact. In 1885 he described
in the same journal a new Carex (viz. C. tartarea RIDLEY),
collected in Sumatra by Dr. H. O. Fores, while one year later
he published a study on the Monocotyledonous plants of New
Guinea, collected by the same (followed in 1925 by a study of
ForBEs’s Javan and Sumatran Monocotyledones). Later, various
contributions about the flora of the Malay Archipelago have been
issued, most of them concerning Borneo and Sumatra.

RIDLEY made his first visit to Sumatra in 1897, to the Siak
district in the South. He did not have a second opportunity for
a visit to this island until 24 years later; he jumped at this °
occasion as he ‘‘was anxious to visit other parts of Sumatra
before (his) Eastern travels came to an end’’. Im spite of the
great amount of work which has since been done, the beginning
of RrpLEY’s account of this botanical excursion to Northern

Gardens Bulletin, S.S.

47

Sumatra, viz. to Brastagi and the Sibayak Volcano is still true :
‘’'The flora of Sumatra is at present still very little known, so
that the account of even a short excursion especially in the quite
unexplored northern part of the island is a contribution to the
history of its flora’. To know this history of the Sumatran -
flora as completely as possible is the prime requisite before
cultivation, which is rapidly extending, makes it impossible for
ever. Owing to the marked similarities that exist between the
flora of Sumatra and that of the Malay Peninsula, a thorough
knowledge of the flora of the latter territory is of the greatest
importance in a study of that of Sumatra, though the Sumatran
flora has certain peculiar characteristics, so that MERRILL’S.
opinion may be correct: ‘‘So far as I can judge at the present
time, somewhere between 40 and 50 per cent. of the known
Sumatran species are endemic, so that no published flora of any
neighbouring region can be expected to be more than an aid
to the determination of Sumatran material’’.

As to Borneo the words of RrpLEy in 1913: ‘‘very much
remains to be done even in the best-known district’’, agree also
with the present situation of 20 years later: on account of its
enormous densely forested areas, sparse population, few roads
and difficult transportation, Borneo has been only slightly
explored botanically. How exceedingly rich its flora is, is
illustrated by the extensive and very interesting Kinabalu
collections made a few years ago by Mrs. Mary and Chaplain J.
CLEMENS: these collections, among the largest ever made in
Borneo, are very important to our knowledge of the Bornean
flora, though that lofty mountain had already received attention
from several botanists. Ripi_ey himself, who visited Sarawak in
September 1905, has written many essays about the flora of
Borneo, working through the important collections made by Dr.
HAVILAND and Bishop Hoss in Sarawak.

From the foregoing it is evident that Mr. R1ipLEy’s work has
in large measure been devoted to systematic studies: the name
of Ripley will always be associated with work on the flora of the
Malay Peninsula. He has further been much concerned with the
geographical distribution of plants and with the development of
species; his interest in these subjects finds expression in several
of his published papers. This also may have underlain his great
work entitled ‘‘The dispersal of plants throughout the world’’,
which begins as follows: ‘‘The study of the Dispersal of Plants
BN is one of very great importance in its bearing on the
distribution of plants throughout the world, the causes of their
presence or absence in different localities, the changes of floras
at different periods and all that goes to make a local flora. It
has also been an important factor in the evolution of species,
genera, and, in some cases, even of orders’’. For long years

Vol X: (1035).

48

considerable attention was given by him to this subject: already
in 1905 in his description of the botany of Christmas Island, he
devoted a detailed chapter to the dissemination of the plants of
this island, which presented ‘such an excellent occasion for his
purpose, being a true oceanic island, as it ‘‘has never at any time
been connected with the mainland of Java or with any other
land’’, and as “‘it is an extinct volcano covered with coral reefs
of various ages from the eocene period to the present day’’.

Moreover by his studies of economic plants, RIDLEY proved
that there exists an admirable opportunity for co-operative
productive work. Some of his publications are of such great
importance for the Netherlands Indies, that they have been
translated into Dutch. In ‘‘The Timbers of the Malay Peninsula’’
(De Maleische timmerhoutsoorten) timbers are concerned, which
are distributed throughout the Malayan region. However,
RIDLEY’s excellent study was a valuable addition to publications
already existing, in which mainly Javan timbers were described,
the descriptions of RrpLEy referring for the first time to timbers
from Sumatra.

Ripiey’s Malay drugs (De inlandsche geneesmiddelen der
Maleiers) originated in 1894 from a lecture about the Malayan
‘‘Materia medica’’. The medicines, used in the Malay Peninsula,
agree so well in nature and even in their names with those from
Java and Sumatra, that in this case too the importance of a
translation is quite conspicuous.

This very year MERRILL, who has also done so much for the
knowledge of the Malaysian flora, has once more broken a lance
for the value of international co-operation among scientists in the
introduction of part 1 of the ‘‘Chronica Botanica’’ (1935). Let
us finish with his words, for which he could have cited RIDLEy’s
work in affirmation: “...... science is necessarily world-wide in
its scope. An outstanding scientific development in one country
is followed with interest and appreciation in all other countries,
to a degree scarcely reached in any other group of thinking
people. Particularly among botanists has international co-.
operation been developed, ever since the need of international
co-operation was indicated through the development of its basic
branch, Taxonomy. Plants are not subject to _ political
boundaries, and the problems of plant science cover the world
in all fields. The continued harmony among botanists in these
days of stress, international discord, and internal difficulties, as
widely separated as they are in the fields in which they specialize
and the institutions of which they are a part, may well influence
international co-operation among other groups’’.

Gardens Bulletin, S.S.

49

a SOME MALAYSIAN PHYTOGEOGRAPHICAL
PROBLEMS.

By E. D. MERRILL,
Professor of Botany, Harvard University.

Perhaps no part of the world is more intriguing from the
standpoint of phytogeography than is the great equatorial archi-
pelago lying between southern Asia and Australia. Malaysia is
by far the largest island group in the world, lies wholly within
the humid tropics, has great diversity of altitudes up to nearly
five thousand metres, and enjoys uniformly high low altitude
temperatures, and, except in limited regions, an abundant rainfall.
Almost continuous precipitation occurs over large sections,
accompanied by relatively high humidity; other large areas are
characterized by alternating wet and dry seasons. These factors,
combined with the characters of the soils, the topography, and
the position of mountain masses in relation to prevailing winds
provide optimum conditions for plant growth, and the net result
is a flora of tremendous richness and exuberance. The differentia-
tion of species has perhaps been in part favored by the geologic
development of the region, and its more or less insular character
over long periods of time. Under present conditions contiguous
parts of the same island may present rather strikingly different
floras, while certain islands separated from each other by only
relatively narrow arms of the sea may have very strikingly
different vegetative and floristic aspects.

Some years ago on the basis of a study of endemism of those
parts of Malaysia blessed with published floras or compiled
enumerations, such as the Malay Peninsula, Java, Borneo, and
the Philippines, I concluded that when the flora was approxi-
mately known, that in the Pteridophytes and the Spermatophytes
combined its total would approximate to 45,000 species. Thus
for the Malay Peninsula, Borneo, and the Philippines, considering
all known species, it was found that at least fifty per cent. of
those known from each major unit were endemic. Considering
how relatively little was then, and still is, known regarding the
floras of the large islands, including Sumatra, Borneo, Celebes,
and New Guinea, I am inclined to consider that this estimate of
45,000 species is perhaps too conservative.

With a land area in excess of one million square miles, this
region (including the Malay Peninsula, which is almost an island,
and which is essentially a part of the Archipelago) extends from
northern Sumatra to Java, New Guinea, and the Philippines, a
region of magnificent distances, extending from north-east to
south-east, a distance of over forty-five hundred miles. Among
its many thousand islands are found the largest ones in the
world: New Guinea, Sumatra, and Borneo, and such large

Vol. IX. (1935).
D

50

secondary ones as Java, Luzon, Mindanao, Celebes, Gilolo, and
Timor. The magnitude of some of these major islands is well
suggested by Wallace’s statement that within Borneo ‘“‘the whole
of the British Isles might be set down, and would be surrounded
by a sea of forests.’’

This is one of those parts of the world regarding which we
have only an approximate knowledge of its flora. Certain areas,
such as parts of the Malay Peninsula, Java, and parts of the
Philippines, may be considered to be well-known botanically,
_ but in fact only certain very limited parts are thoroughly known.
Vast regions have been only superficially explored, and great
areas have never been visited by any collector or botanist. A
distinctly high percentage of the known species is known only
from the single collections on which the original descriptions are
based. Wecannot, therefore, make any reasonably accurate com-
parison between any particular part of this vast region and
comparative areas in what may consider to be thoroughly explored
parts of Europe and North America. In any consideration of the
numerous phytogeographical problems of Malaysia we are handi-
capped by our own limited knowledge of its exuberant flora, and
must realize that future intensive exploration will yield data

which may conceivably alter some of our at present accepted
axioms.

Although the geologic history of Malaysia is reasonably well
understood in its salient outlines much detailed knowledge is
lacking. The magnificent work of the Dutch geologists and
hydrographers on the geological history of Malaysia forms a basis
that must be considered by those biologists interested in the
present-day distribution of plants and animals, for the key to
present-day distribution to a large degree is found in geologic
history. Whatever the causes may have been, it seems to be
evident that the great islands of Sumatra, Java, Borneo, and the
Palawan-Calamian group in the Philippines were in the Pleiocene-
Pleistocene at times connected with the Asiatic continent, and
New Guinea had the same relationship with Australia. There is
much evidence to support the idea that most of the Philippine
group, Celebes, Lesser Sunda Islands, and the Moluccas, at this
time in geologic history, as well as in the Tertiary, were con-
stantly archipelagic, with no direct land connections between
eastern and western Malaysia, and hence no direct land connec-
tions between Asia and Australia since the early Tertiary.

Botanists, probably because confronted by such myriads of
species, have not been as prone as zoologists to delimit life zones
or to map provinces. ‘The attempts of the zoologists to establish
lines of demarcation separating the Asiatic and Australian faunas
are interesting. Wallace’s line extending northward from
between Bali and Lombok through the Macassar Strait between
Borneo and Celebes is, at least popularly, the best known of
these. Its most noteworthy substitute is Weber’s Line, farther

Gardens Bulletin, S.S.

51

to the east. If these lines are plotted on a hydrographic map the
former approximates the eastern boundary of the Asiatic contin-
ental shelf and the latter the western boundary of the Australian-
Papuan shelf. It is reasonably good evidence that these two
shelves mark the approximate limits of previously existing
continental areas. These two more or less stable areas have been
constantly separated by a triangular unstable insular area, its
base the Lesser Sunda Islands, its apex Luzon.

There is no sharp line of demarcation between the Australian
and the Asiatic faunas and floras, but rather this unstable insular
area forms a great transition zone that has been in part populated
by infiltrations from the south and east and from the north and
west. No proposed line is absolute for any single group, much
less when all groups are considered.

It is a well-known fact that as we progress southward and
eastward from Asia, the continental types of animals and plants
become fewer and fewer, and as we progress northward and
eastward from Australia, the Australian types likewise decrease
even more markedly. This is exactly what we would expect if
our present concept of the geologic history of the region is
approximately correct.

One should not make the mistake of drawing final conclu-
sions on the basis of the present-day distribution of any one group
of organisms, whether it be the molluscs, insects, reptiles, birds,
freshwater fishes, mammals, or plants, and much less on any
single family or genus within these major groups; and yet there
are conspicuous cases of broad generalizations that have been
founded on the study of a single family, perhaps even a single
genus. While some such generalizations may ultimately prove
to be correct, it is always safer to consider what is known regard-
ing the geographic distribution of other and perhaps totally
unrelated groups and to attempt to correlate such data with what
is known of the geologic history of the entire region. It is a safe
assumption that the different groups have been developed and
have attained their present-day distribution at different periods
in geologic history; and that while some groups were rapidly
differentiating, others were perhaps static or were retrograding.
That the Philippine flora, as to the Myrtaceze, shows an un-
mistakably close relationship with that of north-eastern Australia
is no reason for considering that the entire Philippine flora and
fauna is allied to that of Australia. If one studies only the
Dipterocarpacee of the Philippines one would conclude that its
flora is overwhelmingly allied to that of Borneo, Sumatra, and the
Malay Peninsula. Clearly here, as with other groups, the periods
of development and expansion have occurred at different times
in geologic history, periods that permitted the distribution of
one group perhaps restricting that of another.

Vol. IX. (1935).

52

Botanical collectors who have explored Malaysia compared
with those who have worked in Europe and in North America,
north of Mexico, have been few, and the number of botanists who
have familiarized themselves with what has been published is
fewer still. For all practical purposes those botanists who have
concentrated on a study of the floras of the Malay Peninsula,
Sumatra, Java, Borneo, the Philippines, and New Guinea, in the
past two hundred years are merely the pioneers, and their work
should be considered as that of pioneers, to be correlated, polished,
and evaluated by future generations of botanists who will have
infinitely more to work with than did those who first exploited
the fertile fields of Malaysian botany. It will be many years
before the essential data are assembled on which we may base
final conclusions. Even such work as has been done by various
taxonomists on the Malaysian flora working in scattered centres
in Europe, England, and America, in Manila, Singapore, Buiten-
zorg, and Calcutta, has not been well correlated. Much needs to
be done to prove the validity or non-validity of some thousands
of proposed species. Many points can be settled only when
competent taxonomists, having access to comprehensive collec-
tions, shall have had the time and opportunity to prepare and
publish authoritative revisions of numerous natural groups. As
this is done, and as type collections and other authentic material
are critically studied and the species carefully compared with
each other, nomenclatural changes and many reductions of pro-
posed species will be the order of the day. In accordance with
the current aims and methods of botanical work such changes
will be inevitable. In the more thoroughly explored parts of
the world, and in those countries where their floras have been
long and intensively studied, we approach a condition of finality
but in such vast regions as Malaysia, where great areas still
remain to be explored and even existing collections have not
been intensively and comparatively studied, our taxonomic con-
clusions cannot be other than approximate.

It has long been known that within Malaysia we find what
we consider to be Asiatic and Australian types. To a much
lesser degree we find scattered African and Mascarene types from
the west, Polynesian ones from the east, and New Zealand and
New Caledonian ones from the southeast. Both the Asiatic and
Australian types include high and low altitude forms. In refer-
ence to the high altitude and some of the low altitude Asiatic types
we are reasonably confident that they actually are continental in
origin, because in such genera as Taxus, Pinus, Ulmus, Salix,
Acer, Primula, Gleditsia, Lilium, Anemone, Sedum, Rosa, Buxus,
Potentilla, Lysimachia, Aster, Solidago, and many others, the
numerous species are characteristic of the north temperate zone
of both hemispheres. The occurrence of scattered species in
Malaysia is unquestionably due to the fact that somehow at some

Gardens Bulletin, S.S.

53

time in geologic history these representatives of northern genera
extended their ranges southward into Malaysia and have persisted
there.

As to the Australian types in Malaysia such as Eucalyptus,
Centrolepis, Patersonia, Drimys, Clianthus, Stackhousia, Pimelea,
Tristania, Leptospermum, Melaleuca, Xanthostemon, Halorhagis,
Didiscus, Stylidium, and others, we can also be reasonably certain
that they are true Australian types because of the great develop-
ment of these genera in Australia and, to a certain degree, in
New Caledonia; and that the few representations in Malaysia
migrated northward into the archipelago even as the Asiatic types
migrated southward into Malaysia.

There are, however, a very large number of genera common
to north-eastern Australia and Malaysia and southern Asia and
Malaysia, some covering the entire range from Asia to Australia,
whose origins it is practically impossible to determine and to

- assign to Asia rather than to Malaysia or vice versa. Doubtless

many of these were originally Malaysian and extended their ranges
southeastward to Australia and northward to Asia, but many
others probably Asiatic. Their universal distribution within the
monsoon region is unquestionably in part a reflex of the adapta-
bility of these plants to a certain type of climatic conditions.

There are relatively few African and Mascarene types in
Malaysia, such as Angrecum, Combretodendron (Petersia,
Petersianthus), Ervthrophleum, and Baphia, but definitely they
are African and Mascarene because the principal development of
species in these genera (at least Erythrophlaum, Angrecum and
Baphia) is found in those regions. When we come to the
so-called Polynesian types in Malaysia, such as Javea, Joinvillea,
Dolicholobum, Tetraplasandra, and Couthovia, the present
indications are that they are not really Polynesian types in
Malaysia but are actually Malaysian types in Polynesia, the
generic types merely having been first described from the latter
region. Thus Vavea is now known to occur from Java, Borneo,
the Philippines, and New Guinea; and New Guinea in definitely
the center of origin and dispersal of Couthonia and of Dolicholo-
bium. Jotnvillea and Tetraplasandra, both small genera, have
Malaysian representatives.

The numerous definitely Asiatic types in Malaysia apparently
followed certain distinct routes of migration into Malaysia from
the continent. In the west unquestionably the most important
route was through the Malay Peninsula to Sumatra, Borneo,
Java, and even further to the south and east. Considerable
evidence leads us to believe in the existence of a secondary
western route from Asia through the Andaman and Nicobar
Islands to Sumatra. In the east, considering the fact that certain
Asiatic types occur in the Philippines and in Celebes, but are
absent from other parts of Malaysia, there was apparently another
route from southeastern Asia to Formosa and the Philippines and

Vol. IX. (1935).

54

southward. We find in Formosa and on the higher mountains of
northern Luzon certain Himalayan types, in many cases these
representing the most eastern and southern extensions of the
Himalayan flora.

Increasing knowledge of the biology of Malaysia is tending
more and more to confirm the belief that there have been two
great centres of origin and dispersal of tropical types, one in the
west, the Sunda Islands, Java, Borneo, Sumatra, and the Malay
Peninsula, and one in the east, New Guinea. From the latter
plants and animals migrated southward into eastern Australia,
eastward into Polynesia, and northward into the Philippines, few
extending westward to the Sunda Islands From the Sunda
Islands the striking migrations were northward into tropical Asia,
northeastward into the Philippines, and to a more limited degree
eastward and southward in the direction of New Guinea. There
were, of course, intermigrations within each area.

Comparative studies of the climates and plants in tropical
Asia and Malaysia with those of America, emphasizes one striking
difference. In America the humid tropics are to a large degree
separated from the temperate regions by extensive arid areas. In
the Old World the intervening arid regions are lacking. There
has been a continuous humid north and south route connecting
Malaysia with subtemperate and temperate parts of Asia—the
Malay Peninsula northward through Burma, Siam and Indo-
China, to India and southern and eastern China. The limiting
factor here has been largely temperature, while in America it has
been aridity and temperature combined. We have little evidence
on which to base conclusions as to what the humidity and aridity
of the various parts of Malaysia may have been in past geologic
times, but the available paleobotanical evidence seems to indicate
continuously humid tropics in the past, in general similar to the
conditions existing to-day.

Outside of the taxonomic field, but nevertheless intimately
associated with and dependent upon it, there are an infinite
number of problems to be settled regarding the Malaysian fauna
and flora. In part these depend on an intensive study of existing
collections of reference material, in part on long continued inten-
sive field work. What was the original flora of Malaysia? What
are to be considered the relic species of earlier floras? What
factors permitted or encouraged the inter-migration of Australian-
Malaysian and Asiatic-Malaysian types and what were the
limiting factors? Were these geological or climatic in nature,
or both? Does the theory of continental shift throw any light
on the problem? What are the relations between the present
distribution of plants and the prevailing winds? What bearing
do typhoons have on the problem? What is the relationship
of existing ocean currents to the spread of plants? What does
the annual north and south flight of certain types of migratory
birds signify for those plants characterized by the production

Gardens Bulletin, S.S.

55

of myriads of minute seeds and adapted to muddy or wet
habitats? What are the effects of constantly moist and alter-
natingly wet and dry conditions? What is the effect of dominant
mountains and mountain ranges on prevailing winds and vice
versa, and how do these factors affect vegetation? What is the
bearing of geologic history and of hydrography and topography
on the problem ?

From long residence in the tropics and from numerous
observations made on the vegetation, some years ago I reached
the general conclusion that in modern times man is the most
important single factor in the dissemination of plants. Manifestly
much of Malaysia was originally covered by a forest vegetation
of one type or another. To-day vast areas are under intensive
cultivation, almost equally large areas are in primitive cultiva-
tion, enormous tracts are covered with coarse grasses, bamboo
thickets, and second growth forests. Man’s agricultural activities
have been the greatest single factor in the destruction of the
original forests over vast areas, but lumbering, and in some
regions mining, has played a part. It is axiomatic that in
Malaysia a primary forest once destroyed is never re-established
directly; its place is taken by coarse grasses, bamboo thickets, or
dense stands of rapidly growing small trees and shrubs. As the
primary vegetation has been destroyed the habitats have been
provided for plants from other regions. These, accidentally or
intentionally introduced by man, or through the agency of nature,
once man has paved the way for them, have become dominant
over great areas. Many of these were of American origin,
introduced since the sixteenth century, including representation
of such woody genera as Leucena, Psidium, Lantana, Pithecello-
bium and Prosopis, and a very large number of weedy herbaceous
plants in such genera as Drymaria, Malachra, Waltheria,
Stachytarpheta, Cassia, Hyptis, Ageratum, Cosmos, Elephan-
topus, Erechtites, and Synedrella, as well as numerous grasses
and sedges. Antedating these numerous American introductions
were many Asiatic types, and this introduction of Old World
forms is still progressing. There is much that needs to be done
in reference to the history of these introduced plants. Where
did they come from? When were they introduced, and how?
What are the reasons for their present dominancy? What is the
percentage of introduced species in primary forest areas as com-
pared with the percentage in those regions where man is directly
or indirectly responsible for the present vegetation ?

Endemism presents numerous fascinating problems.
Throughout Malaysia one finds in most of the large genera a few
strongly characterized, widely distributed species. At the same
time in these genera one notes that in such groups as Ficus,
Antidesma, Eugenia, Medinilla, Dendrobium, Quercus, Eria,
Bulbophyllum, Calamus, Aglaia, Begonia, Polyalthia, and
Canarium, to mention only a few, that there is a tremendously

Vol. IX. (1935).

56

developed local endemism in many of the larger islands and even
in some of the smaller ones. Invariably the percentage of
endemics is very low in the settled areas, open grass lands, and
secondary forests, and correspondingly is very high in those
groups that require primary forest conditions. One concludes
that the primary forests represent the original vegetation, and
that the low percentage of endemics in the settled areas, open
grass lands, and secondary forests is a reflection of the activities
of man on the vegetation as a whole; that many of the species
now dominant in these latter areas were introduced as the
primary forests were destroyed. We are reasonably certain that
this was the case in reference to the many species of American
origin, which predominate in the settled areas, for these did not
commence to come in until some time during the sixteenth
century. We are less certain in reference to those species of
Asiatic origin, largely groups characteristic of the second growth
forests and open grass lands.

The cultivated plants represent a series of problems in
themselves. Most of those in Malaysia are not natives of the
archipelago but were introduced by man from other regions.
The earliest invading peoples probably brought no plants for they
had no agriculture. Later waves of more advanced peoples cer-
tainly brought cultivated plants with them. Many basic
Malaysian food plants were introduced in prehistoric times, others
in modern times; and the introduction of weeds parallels the
introduction of economic species. Within the period of western
expansion the Malaysian flora has been greatly enriched through
the introduction of numerous economic plants and many weeds
of American origin.

Not to be disassociated from the cultivated plants are their
aboriginal names, which in many cases point definitely to the
place of origin and the approximate time of introduction of
individual species. In these fields of ethnobotany and compara-
tive philology of plant names much needs to be done, and
Malaysia with its very numerous dialects and its great racial
mixtures offers a marvellously fertile field.

Again in the field of plant biology there are numerous
problems. The relationships between insects and plants, birds
and plants, and even bats and plants in reference to cross
pollination. The biological relationships between ants and such
domiciliary plants as Hoya, Dischidia, Mvyrmeconauclea,
Neonauclea, Macaranga, Lecanopteris, Polypodium, Demonorops,
Myrmecodia, Hydnophytum, Myrmedoma, Saurauia, and Piper,
and representatives of other genera where piant organs such as
branches, stems, leaves, and sheaths have been modified to form
domiciles for colonies of ants. But this and numerous other
subjects is outside the scope of this paper.

Gardens Bulletin, S.S.

37

In this brief survey, hurriedly written because of the
exigencies of my present situation, and entirely undocumented, I
have attempted merely to enumerate some of the numerous
intriguing problems of Malaysian phytogeography. I have done
so in the hope that some who read it may be tempted to assist in
solving some of the problems by undertaking field work in this
too little explored part of the world or to co-operate in the study
of the rich reference collections already available. In Malaysia
one is constantly on the edge of the botanical unknown. ‘The
field is vast, existing problems create further ones, and the need
of more investigators is great. ‘The serious-minded interested
student who elects to assist in the study of the plants and plant
problems of Malaysia will be amply repaid by the productive
results secured.

Vor EX. (.1935.).

58

THE DISPERSAL OF PLANTS BY FRUIT-EATING BATS.

By W. M. DOCTERS VAN LEEUWEN,
Leersum, Holland.

Many data are at our disposal about the dispersal of plants
by birds, both endozoically and exozoically. Very little, however,
is known about the part played in plant dispersal by fruit-eating
bats. They are probably as important as the birds, but exact
observations are mostly lacking. In H. N. RipL&y’s well-known
handbook on the ‘‘Dispersal of Plants throughout the World”’
three pages are filled with what is known of plant dispersal by
bats. These data are based partly on his own observations made
during his long and fruitful stay in the tropics, and partly they
have been taken from literature. In the same work upwards of
130 pages are given to seed dispersal by birds.

Fruit-eating bats occur in the tropics of the old and of the
new world, but they are most numerous in the tropical territories
of Asia and Australia. Little is known about their way of living,
though they can be observed everywhere. In the twilight these
bats come forth from their sleeping places, just like the insect-
eating species, and look for their food during the night, to turn
back again to their sleeping places at day-break. A few species,
viz. of the genus Pteropus, the Fox-bats, hang down from the
branches of large trees, in great numbers together, during the
daytime. Others hide themselves, resting in holes, or under
folded leaves of palm-trees, or in dark parts of the forests, as I
observed several times in Krakatau. In the lake-plains of
northern New Guinea I saw large troops, consisting of thousands
of individuals, flying low in great circles above the vast
Saccharum-marshes, during the hottest hours of the day. The
animals circled in a quiet, regular flight above the grass covering,
and it seemed as if they were asleep. When they were startled
by a gun-shot, they flew restlessly in all directions, finally to
unite again into the rotary flight. Above these marshes this
phenomenon seems to be observed regularly; in 1926 I myself
saw it four times, and also during former expeditions it seems
to have been perceived, for this place, which is a large island in
the river Mamberano, has received the name of Kalong Island,
i.e. island of flying foxes.

Sopy' mentioned a few particulars about the modes of life
of fruit-eating bats in captivity. These animals seem rarely to
be kept in captivity, but Sopy succeeded in keeping a female of

1. SoDY. Jets over vruchtenetende vleermuizen in gevangenschap.
De Tropische Natuur. Vol. XVII. 1928, p. 170.

Gardens Bulletin, S.S.

59

Cynopterus sphinx tithecheilus, and her young alive for a
considerable time. According to him both specimens were
hostile to approach. At Bandoeng, I myself kept a specimen of
Pteropus species in my aviary for a long time; it was very affec-
tionate, crept up my coat, and nestled its head against my neck.
According to Sopy they seek food late in the evening, take it
along to the higher parts of the cage, and eat it while hanging
down. Sometimes they hang down by one leg only, using the
other to hold the fruit in front of them. In the field he observed
that this species broke off the fruits of Piper aduncum.
Unfortunately nothing is said about the food eaten by the
animals in captivity, only that they got soft fruits. Nor are any
data given about the digestion of fruits, and the excretion of
seeds.

The fruit eating bats mainly live on fruits, but other parts
of plants are eaten also. The fruits eaten by these animals are
either soft or hard. Soft fruits which are taken may be berries,
soft drupes, figs, and gourds. But they eat also fruits with a
hard, or rather hard, fruit-shell. These are mainly the larger
drupes. These are torn off with the beak, and brought back to
the resting places, to which the animals return again and again.
They usually choose a spot near the tree from which they gather
the fruits, using a branch, roof-gutter, or prominent parts under
the roofs or galleries of houses, to hang from. The soft or hard
wall is partly or completely gnawed off from the kernel, and the
latter is simply dropped. Often they seem to take the fruits along
over greater stretches on returning at dawn to their sleeping
places, for below them fruit-kernels and seedlings are often
found. At Buitenzorg, near my laboratory, there stood a few
old specimens of Corypha Utan, in which roosted a colony of
Cynopterus sphinx tithecheilus. The animals slept in the old,
more or less folded and drooping leaves. Under these trees the
grass was partly killed by the excreta, but a number of plants
developed from the fruits taken home by the bats, and eaten
there. They sometimes drop the fruits undamaged during the
flight. During the time the large Canarium commune bore
fruits in the Botanic Gardens, at Buitenzorg, we could often hear
the heavy fruits banging down on a zinc pent-house.

These animals probably cannot cover great distances with
these heavy fruits. Observations, however, are lacking, and it
would indeed be very difficult to make any. But yet many
fruits and plants which I found under the above-mentioned
sleeping places at Buitenzorg, must have been carried for a
distance of hundreds of metres, because in the immediate
surroundings of the Corypha there were no fruit-bearing trees
from which the bats could have taken their food. As the animals

Vol. IX. (1935).

60

are very voracious, and often eat only a part of the fruit-flesh,
these large colonies must destroy enormous quantities of fruits.
For the dispersal of fruits over moderate distances they must
undoubtedly be of the greatest importance.

Van Der Pit’ tells in an article of recent date that the
fruit-flesh is not eaten straight away, but chewed, sucked, and
ejected. These animals have a very narrow gullet, measuring
in case of Cynopterus 2 to 2%4 mm. And almost without excep-
tion, the intestinal canals of these animals are filled with a
colourless jelly. The larger species of Pteropus too are said
not to contain any seeds in the intestinal canals. Mc. CANN? also
found only a mucous fluid in the intestines of Pteropus giganteus,
although the animals had eaten fruits of various species of
Ficus only a short time before. VAN DER PyL, however,
mentions that seeds of Ficus fistulosa have been found in the
newly evacuated excrement of a Cynopterus sphinx, and that
the intestines of Cynopterus brachyotis, which had eaten the
catkins of Piper aduncum, contained seeds of this plant.

According to VAN DER PIL, fruits eaten by bats have certain
characteristics, viz. inconspicuous colour, a smell which is
neither fresh nor agreeable, large size, large kernels, hard flesh,
while the part of the plants from which the fruits are taken is
near open spaces. Such fruits are actually found in the food
of bats, but many others, besides which are juicy, fragrant,
small, and soft. At any rate, these animals are important
because they eat many fruits which are despised by most fruit-
eating birds.

Most remarkable is, according to VAN DER PIL, the cauli-
flory of many fruits eaten by bats. He observed the following
fruits eaten by bats: Achras Zapota, Anona muricata, Averrhoa
Carambola, Bouea macrophylla, Canarium spec. div., Carica
Papaja, Ceiba pentandra, Cynometra cauliflora, Eleocarpus
Ganitrus, Eugenia javanica, malaccensis, and polyantha, Ficus
fistulosa, and glomerata, Garcinia Mangostana, Lansium
domesticum, Mangifera ce@sia, fatida, and indica, Musa
paradisiaca, Melia Azedarach, Nephelium lappaceum, Persea
americana, Psidium Guajava, Phyllanthus acidus, and Emblica,
Sechium edule, Spondias, and Zalacca edulis.

In case of species of large fruits with large kernels only the
wall is eaten away, but the soft fruits with small seeds-are eaten
either whole, or part of them. A great number of seeds are

1. L. VAN DER PIJL. Zaadverspreiding door vleermuizen. De
Trepische Natuur, Voi. XXIV. 1935, p. 131.

2. Mc. CANN. Notes on the flying fox (Pteropus giganteus).
Journ. Bomb. Nat. Hist. Soc. Vol. XXXVII.
(Quoted from VAN DER PIJL.).

Gardens Bulletin, S.S.

61

eaten together with the flesh in which they are imbedded. In the
case of figs they are really small fruits covering the inner wall of
the receptaculum, but I will call them ‘‘seeds’’. The juicy
fruits are digested in the intestinal canal, and the seeds come
forth undamaged together with the excreta. These excreta are
dropped, not only in the resting places, but also during the
flight, so that the seeds can come down in all kinds of places.
They are hardly ever found on the ground, because here they are
hidden between plants, but often on leaves, stones, and walls.
The excreta are of a very fluid consistency, often jelly-like. I!
often saw the seeds of Piper aduncum on leaves in the
surroundings of Buitenzorg. I saw leaves with excreta
containing seeds of Ficus in Krakatau, where there is a very great
number of two species of fruit-eating bats, Cynopterus Horsfieldii
and C. sphinx tithecheilus; in the higher parts of the same
island I found excreta principally containing seeds of Cyrtandra
sulcata. The excreta also often come down on buildings, and
young plants of Ficus Benjamina and of F. retusa can often be
seen growing in the gutters. When the seeds come down on old,
badly kept buildings, or on old temples, then the roots of these
large species of Ficus can slowly break the stones asunder, and
do much damage.

The animals need not rest while eating these juicy fruits,
and thus the seeds can be carried over greater distances than in
the case of big fruits with a hard kernel. It is not known how
long a time the seeds need to pass through the intestinal canal of
bats in the digestive process. It would be important to study this.

In the islands north of Batavia there are large colonies of
Pteropus, and the animals fly over to the shore of Java, late in
the afternoon, or early in the evening, in order to look for
food. When at sea one can often see large groups of these
animals crossing wide sea-arms. Undoubtedly these animals have
played a part in the dispersal of seeds to the islands of the
Krakatau-group. It is not known when these animals have
arrived there. JACOBSON’ who examined the fauna of these
islands in 1908 fails to mention them, but in rg19 they were quite
common, and DAMMERMAN? mentions them in 1922.

The first to write about plant dispersal by bats was
RUMPHIUS, in his well-known Herbarium Amboinense. He
mentions Nauclea elegans, Artocarpus integra and communis,
besides Anona mucosa, and two species of Mangifera, the seeds
of which are spread by bats. In addition many data can be found
in the above mentioned work of RIDLEY.

1. E. R. JACOBSON. De nieuwe fauna van Krakatau. Jaarverslag
van den Topographschen Dienst in Nederlandsch
Indié over 1908. Batavia, 1909, p. 1092.

2. K. W. DAMMERMAN. The fauna of Krakatau, Verlaten Island
and Sebesy. Treubia, Vol. III. 1922, p. 65.

Vor. EX.) (1035).

62

An examination of the contents of the intestinal canals of
these animals might produce important results. Up till now only
a few facts are known. One might also collect the fruits and
seeds from the excreta. Another way of carrying out the
investigations with a greater chance of success, is the observation
of plants which regularly come up under the sleeping and resting
places of these animals. During the many years which I stayed
in Java, I noted down, here and there, which species of plants
occurred in such places. In a few cases it was not possible to
recognize the species of plant from the young, non-flowering
materials. It appeared from these observations that numerous
species of plants are spread by these bats; the appended list,
undoubtedly, does not contain all the species which are actually
eaten and dispersed. The list, moreover, shows that it is mainly
species of trees from which the bats take their food. Yet one
sees among them also smaller plants, such as Clibadium
surinamense, Passiflora fotida, a Zingiberacea, and an Aroid.

Some plants owe their quick dispersal mainly to these animals.
Piper aduncum is a small tree which has been imported to the
Botanic Gardens at Buitenzorg from tropical America. ‘The
trees, which are 2 to 6 m high, often bear large quantities of
fruit-catkins, which are principally eaten by the smaller species
of fruit-eating bats. Owing to this the plant has spread very
quickly in the surroundings of Buitenzorg, and farther in West
Java. Large quantities occur in grass jungles, in valleys, and in
kampong wildernesses, where often it may crowd out the original
flora.

Below is given a list of the species of plants which I have
found. Most observations have been made in Buitenzorg, but
also in Batavia, Bandoeng, Samarang, Cheribon, and other
smaller places, wherever I had the opportunity. As I was not
able to state the species of bats, I give the list without indicating
them. Among the plants found in Buitenzorg there are several
species of trees which are cultivated in the Botanic Gardens.

Young plants found under the resting and sleeping places
of fruit-eating bats :—
MORACEZ= URTICACEZ,

1. Artocarpus communis FORST.| 16. Pipturus incanus WEDD.
elastica REINW.

N

e et BL PIPERACE.
3. mm integra MERR. 17. Piper aduncum L.
4. Ficus ampelas BURM. 18. ,, -retrofractum VAHL.
Buys ee L. EUPHORBIACEZ.
ane mea Arana 19. Antidesma Bunius SPR.
Ms * - Fctulosa. REINW 20. Baccaurea racemosa M. A.
9. ,, fulva REINW. ANONACEE.
EOLA) oy) hispida TL. 21. Anona muricata L,
Hes Hd infectoria ROXB. 22. Species ignota.
iz. +; "pulosa (REIN W..
13. 4» - iretusa B) LAURACEZ.
14. ,, septica BURM. 23. Litsea polyantha Juss.
15. 4, | ¥arieeata Bx. 7 ie species.

Gardens Bulletin, S.S.

25.

26.

ie meee Fapaja:L.

. Calophyllum Inophyllum L.

3°
. Ochrocarpus

. Cynometra cauliflora L.
. Maniltoa gemmipara SCHEFF.

. Eugenia aquea BuRM. f.

. Terminalia Catappa L.

. Durio Zibethinus L.
. Eleocarpus species.
. Evodia species.

. Canarium commune L.

. Aglaja species.

63

-_FLACOURTIACEZ.
Flacourtia Rukam Z. et M.

PASSIFLORACEZ,
Passiflora feetida L.

CARICACEZ.

GUTTIFERZ.

“species.
congregatus
BOERL.

LEGUMINOSZ.

MYRTACEZ,

. malaccensis LAM.
i polyantha WIGHT.
x species. |

COMBRETACE.

aa species.

MELASTOMATACE:.
Melastoma malabathricum L.

MALVACEZ.
ELAZOCARPACE.

RUTACE.

BURSERACE.

be species.
MELIACE.

Dysoxylum species.

3 Lansium domesticum CORR.
. Melia Azedarach (sa
. Sandoricum Keetjape MERR. |

Vol. IX. (1935).

58.

50-

60.

OL.

. Actinophlaas

ANACARDIACEZE.
Gluta Renghas L.

. Mangifera indica L.,

species.

. Spondias dulcis FORST.

SAPINDACE.

. Nephelium lappaceum L.
. Sapindus Rarak DC.

EBENACE.

. Diospyros species.

- species.

LOGANIACE&,
Fagrea species.

RUBIACEZ.
Morinda citrifolia L.

CUCURBITACE.
Trichosanthes species.

COMPOSIT.

Clibadium surinamense L.
var. asperum BAKER.

ZINGIBERACEZ.,

. Amomum species.

MUSACE.

. Musa species.

PALM.

angustifolius
BECCc.

. Arenga pinnata MERR.

. Bactris speciosa KARST.

. Caryota Rumphiana Mart.

. Corypha Utan LAMK,

. Livistona rotundifolia MART.
. Oncosperma filamentosa BL.

ARACE.

71. Colocasia species.

64

OPEN AIR HOTHOUSES IN THE TROPICS AT 3100
METRES ALTITUDE.

By C. G. G. J. VAN STEENIS,
Botanist ai the Herbarium, Buitenzorg, Java.

In this article I discuss the ecological and morphological
reponse of some plant species to the action of fumaroles in the
highlands of Java and Bali. I also discuss briefly the dispersal
of some of these plants. |

In June of this year I climbed Mt. Ardjoeno’ in East Java.
This lofty volcanic mountain complex, attaining more than 10,000
feet above sea level, is built upon a fault running north and south.?
It consists of 5 separate peaks, the northernmost of which is
called Mt. Welirang. Welirang is the Javanese name for
sulphur, and this peak is so called because of the occurrence of
several enormous solfatara, which have been commercialised for
centuries, probably even in Hindu times. Numerous Hindu
monuments are found on Mt. Ardjoeno at nearly 10,000 feet
altitude in the form of remains of well built rooms. Even today
on several Javan mountain peaks priests live a poor life and
offerings are regularly made by the people.

The southernmost peak of the mountain complex is Mt.
Ardjoeno itself in a restricted sense. This peak, the highest of
all, is practically extinct. It does not possess such huge craters
and solfatara as Welirang. Between Ardjoeno and Welirang
there are three other peaks of lesser altitude. The southernmost
and lowest is Mt. Bakal, a rough and rocky, extinct lava-stop.
The two others are both called Kembar; we shall distinguish them
as Kembar S. and Kembar N. Both are characterised by the
occurrence of numerous fumaroles near their summits. They
are reached easily along the road from Trétés to Lalidjiwo; in the
latter place there is a resthouse at about 2500 m. altitude, situated
within the pure Casuarina Junghuhniana forests, which cover the
entire northern side of the mountain. From Lalidjiwo one
reaches the pass between the Kembars, 2900 m. altitude, in about
one hour and a half. Near the pass I observed the first small
and hitherto unrecorded fumaroles in the northern of the two
blowholes. The soil, consisting of weathered volcanic rock, is
hot, and steam—which condenses perms at this altitude—
is continuously produced.

1 Oe in Dutch is pronounced as oz in would
2 Cf. N. J. M. Taverne, Vulkaanstudien op Java (Vulkan. Med.
Ned. Ind. 7, 1926, p. 109, seq.)

Gardens Bulletin, S.S.

65

It is a well known fact that the local distribution of
mountain plants is often very curious. So for instance
Lycopodium clavatum, L. and Gaultheria fragrantissima, Wall.
have only been found on the Kembar peaks, but never on the
much loftier Welirang or on Ardjoeno itself.

From the pass I climbed Kembar S., which is 3100 m. high,
in about half an hour, over the gently sloping northern ridge,
which is covered by a very short vegetation of small herbs such
as Stvphelia pungens, Koord., grasses such as Danthonia pilosa,
R. Br., Festuca nubigena, Jungh., Tripogon exiguus, Buese,
and especially Imperata (lalang), and several orchids such as
Thelymitra javanica, Bl., and Microtis unifolia, Rchb. Remains
of charcoal from burned shrubs confirm our idea, derived from
the occurrence of lalang, that the absence of forest is due to fire
and to the grazing of deer. Deer are very common here, as
shown by their numerous droppings, especially towards the
summit. I had very bad weather, and made my way upwards
first in a cold drizzle, then under heavy rain. Whereas all the
slopes, except the ridge towards the pass, are covered by fairly
tall tjemara forest, the top region proper is devoid of forest. It
consists of a horseshoe-shaped ridge in the centre of which a
so-called sawahan is situated, that is a cup-shaped basin
covered entirely with the tuft grass Festuca nubigena Jungh.
Everywhere on the summit steam escapes from cracks in the
hot rock; the peak may be said to be more or less crowned by
numerous steam clouds. It was a very agreeable sensation to
sit on the hot soil within reach of the steam, which escapes
under pressure with a dull droning sound. Less agreeable is the
smell which is halfway between that of a rotten egg and that of
a phalloid. However, no SO: was observed. The natives told
me that deer came here at night to sleep.

A very peculiar vegetation is developed round these
fumaroles, and it shows a distinct zonation. Some metres from
the centre, where the steam loses its heat, Lycopodiums
(L. cernuum. L. and L. clavatum, L.) thrive in a rather dense
thicket along with Gleichenia linearis, Clarke and some poor
tjemara, only one metre tall, densely clad with Usnea. Still
nearer to the steam-holes the vegetation is much lower and
consists of a carpet of very small herbs. Here Lycopodium
cernuum, L. shows a very peculiar modification, becoming a
flat plaited mat of hardly 1 dm. thickness, covering several
square metres. It has just the appearance of a moss (Pl. 1).
Other weeds are also able to approach the hot steam, namely a
common lowland Rubiacea, Oldenlandia herbacea, L., which I
Was very astonished to find in such a peculiar locality, forming

Vol. IX. (1935).
E

66

a very low dense mat. Still other weeds show the stunted
dwarf habit. These are Fimbristylis capillaris, A. Gray, and
also, though less typical, Lindernia crustacea, F. v. M., Senecio
sonchifolius, Moench, and a Blumea. All of them, however,
do not form mats. Moreover, Lindernia and Fimbristylis are
found elsewhere in Java in dwarf forms, notably on poor soil,
e.g. on gravel round houses. It must be noted that Lycopodium
cernuum and Oldenlandia are found thriving more luxuriantly
some metres away from the steam spouts. Hence the fumarole
modification is effected by the tolerance of these plants. All
the plants mentioned live in continuously humid and warm
surroundings, which I call an open-air hothouse in a temperate
mountain zone. The wind often alters its direction and sways
the steam clouds all over the summit ridge and on account of
this several of the plants mentioned occur scattered amongst -
the lalang. In some places swollen semiglobular white cushions
of the moss Racomitrium lie loosely on the hot wet rocks.

After I had made these observations I unexpectedly received
confirmation of the phenomena described when my friend
C. N. A. de Voogd, chief forester of Bali, etc., sent me a collec-
tion of plants from the Lesser Soenda Islands and gave me
permission to publish concerning them. He collected in 1934
on Mt. Agoeng, the highest peak of Bali island, 3100 m. altitude.
It is thus of the same height as Mt. Kembar. Agoeng also has
fumaroles on the top ridge and there De Voogd collected some
other plants which show the same steam habit as the Kembar
plants. He too collected the stunted form of Oldenlandia
herbacea, L. (Pl. 1) and also found some lowland plants in the
Bali open air hothouse, viz. Mitrasacme nudicaulis, R. Br.,
Bidens pilosus, L. and Senecio sonchifolius, Moench. His best
finds were, however, the steam-modifications of two species
which I did not get on Kembar—of:Dicrocephala chrysanihemi-
folia, DC. and of Hyptis brevipes, Poit., the former a mountain
plant, the latter a typical lowland plant (Pl. 1). Mr. de Voogd
informed me that the fumaroles of Mt. Agoeng also produce
steam without any smell of SO2. He camped on the summit
ridge and his coolies pitched a tent above one of the fumaroles
to keep warm. The condensation of the steam during the night,
however, caused ‘‘rainfall’’ in the tent, so that this idea turned
out to ®: less practical than it was ingenious.

It is definitely shown that steam, at least that of fumaroles
at high altitudes in the tropics, has a very uniform effect on
plants which are not in the least allied to one another. It is
of interest to note, too, that the steam does not diminish fertility,
as both Oldenlandia and Fimbristylis were in flower.

Gardens Bulletin, S.S.

67

The sketches given above lead to another general considera-
tion. The question arises how these lowlan¢ plants were
dispersed to this high altitude. Dicrocephala chrysanthemifolia
and Lycopodium cernuum are of general occurrence in the Javan
high mountain zones, but what about Oldenlandia and the
others? Of these common weeds (only Mitrasacme nudicaulis
being decidedly rare) we have abundant herbarium material at
Buitenzorg, and on the labels of most specimens the altitude is
noted. From this material, collected throughout the Malaysian
region, I made the following table of the greatest altitudes :—

In Malaysia from sealevel !

) .

i Malaysia : Kembar | Agoeng
Bidens pilosus, L. c. 2100 m. — 3100 m
Hyptis brevipes, Poit. ie.. E2003, — 3100 m
Lindernia crustacea, F. v. M. c. 1450 m. 3100 m —
Mitrasacme nudicaulis, R. Br. |c. 1700 m. — 3100 m
Oldenlandia herbacea, L. c. 1450m. 3100m.| 3100 m
Senecio sonchifolia, Moench. c. 2000 m. 3100m. | 3100 m

This table shows distinctly that there is a distinct gap
between the common occurrence in the lowlands and the hills and
that near the fumaroles on the mountain tops. This cannot be
attributed to the absence of waste ground, where these pantropic
weeds generally grow, in the mountains. On the contrary,
waste ground is very common on the mountain slopes, where
natives burn the vegetation, cut paths, build houses and try to
cultivate crops such as maize and potatos. I am of the opinion
that the plants concerned cannot stand the cold of the higher
mountain zones above 2000 m. This would fit in with their
occurrence in local hotter spots, such as the fumaroles.

The question of vertical dispersal still remains. In the case
of the only Composite, Senecio sonchifolius, Moench., the
pappus is no doubt responsible for dispersal by ascending winds,
which, as I observed frequently, may be very strong. But the
other plants do not possess any special means for wind dispersal.
Neither their seeds or their fruits are fleshy and endozoic
dispersal seems out of the question. Exozoic dispersal must be
held responsible. Their seeds or fruits are all small and
inconspicuous, as are the plants themselves. The only notes on
their methods of dispersal that I could find in Ridley’s well-
known work ‘‘Dispersal of Plants throughout the World’’ are
that Lindernia may possibly be dispersed by mud on birds’ feet,

Vol. IX. (1935).

68

Oldenlandia washed away by rain, and that it might further be
possible that the adhesive calyces of Hyptis and the pappus of
Senecio sonchifolius and of Bidens could become entangled in
the hair and feathers of animals.

I prefer to put forward the following hypothesis. It is
common knowledge that deer migrate through the forests not
only in a horizontal but also a vertical direction. Mr. W. H. de
Kramer’ wrote on migration of tiger, python, bear and pig
during the fruiting season of the durian (Durio) in South
Sumatra. The then prevailing western winds bring the smell
of durian towards the interior, attracting the pig, and with the
latter comes the whole fauna living on them. Migration of
elephant and of rhinoceros from sea level to high up the
mountains is well known. In January 1935, it was observed that
on the north slope of Mt. Slamat in Central Java a continuous
migration downwards of the fauna, including deer, was taking
place. The natives attributed this to a future eruption, which
the animals knew by instinct was coming.? That their instinct
proved to be at fault does not matter here. It might be that
the grazing on higher levels was poor at that time. It is a fact
that several animals when old migrate to mountain tops to die in
solitude, the reason for which is unknown. Several notes on
this phenomenon have been published in our semi-popular
naturalists’ journal ‘‘De Tropische Natuur’’.* I may mention
here of the animals found on high mountain tops—often dead—
the flying squirrel (at 3675 m. on the summit of Mt. Smeru and
at 3800 m. on the summit of Mt. Kerintji), the kambing utan
(the Sumatran mountain goat) and a rat on the top of Mt.
Kerintji, 3800 m., several birds, black monkeys (Semnopithecus
maurus) on Mt. Smeru and so on. It seems to me that exozoic
dispersal by migrating animals, either ascending or descending
for food supplies, for dying or for other reasons is the most
reasonable explanation of the occurrence of lowland plants near
fumaroles. The seeds may adhere with mud to hair or feathers,
or, aS seems more probable to me, to the hooves and feet of
animals, and be occasionally dispersed in that way. They may
adhere in no less degree to the master of all animals, man, who
frequently visits mountain tops and may unintentionally carry
with him seeds from lower down. I refer especially to the
statement made above of priests living and people making
offerings on mountain summits, a practice which dates back to

1 De Tropische Natuur 23, 1934, p. 119-120.

Verslag Ned. Ind. Ver. tot Natuurbescherming 1933-1934, D- 33:
1035-

3 Cf. volumes 19, 1920, p. 197, and 20, 193I, DP. 57.

wv

Gardens Bulletin, S.S.

rn) lf h,
Bae

Gard. Bull. S.S. Vol. IX. Plate 1.

Fimbristylis capillaris Oldenlandia herbacea j Lycopodium cernuum

a i
PK

a.

| ) Dichrocephala. ~~
Hyptis brevipes chrysanthemifolia®

69

Hindu times. De Voogd informed me that the Balinese regularly
make offerings on the summit of Mt. Agoeng, and that many
monkeys live there, feeding partly on these offerings. ‘These
grey monkeys seem to be more or less accustomed to the
presence of man, as they took small pieces of bread without
hesitation. De Voogd also found a skeleton of a monkey on the
summit of Mt. Agoeng.

The gap in altitudinal distribution I believe to be due to the
fact that these plants are not able to thrive, or at least to settle,
between the highest border of distribution and the abnormal
high altitude of the fumaroles, of which the hot soil and atmos-
phere offer lowland conditions. It would be of great interest
to make field experiments on this matter. It would be
interesting, too, to make experiments in the laboratory to discover
the physiological effect of steam on plants. I do not know of
any such experiments. Can the tufted fumarole habit of plants
be brought about artificially in the laboratory by the continuous
action of pure steam? And in all plants? I much regret that
I cannot give any exact data regarding the temperature of the
fumarole steam. All I can say is that even near its point of
escape it was quite bearable on the human skin.

Plate 1. Modifications of plants by fumaroles, the left figures of each
species with the normal, the right with the abnormal habit.
Fimbristylis capillaris, A. Gray, a, from Mt. Tengger,
b. from Kembar S., x 1; Oldenlandia herbacea, L., a. and
b. from Kembar S., c. from Agoeng, x 1; Lycopodium
cernuum, L., a. and 6. from Kembar S., x 4; Hyftis
brevipes, Poit., a. from medium altitude, b. from Agoeng,
x 4; Dicrocephala chrysanthemifolia, DC., a. and 6 from
Agoeng, x 4.

Vol. IX... (10935).

70°

SEMEIOCARDIUM ZOLL., A MISINTERPRETED
GENUS OF BALSAMINACEZ.

By C. A. BACKER,

Heemstede, Holland.

In the Natuurkundig Tijdschrift voor Nederlandsch Indié
XVII (1858-59), pp. 243 seq. the celebrated ichthyologist
P. Bleeker published a letter, dated 8 August, 1858, written to
him by the Swiss botanist-collector Heinrich Zollinger, then
residing in the Dutch Indies, on the Natural History of Madura,
a small island north of the eastern part of Java. In this letter
Zollinger describes a trip made that year by him over the island
in the company of N. A. Th. Arriéns, then Resident (i.e.
Commissioner) of Madura. On page 245 of his description
Zollinger says: ‘‘A new genus of the family of Balsaminacece
‘ig perhaps the most important discovery. The plant grows on
“‘naked calcareous rocks between 600 and 800 feet above sea
‘level and has entirely the structure of the common Balsams.
“Instead of a spur one of the petals bears a short sack ending in
“two obtuse points. ‘The same petal’’ (this is an error; see des-
cription below) ‘‘is provided at its apex with a beautiful cordate
“standard. On account of this character I have called the plant
“‘Semeiocardium arriensii. ‘To my regret I found only two ripe
seeds which, however, did not germinate.’’

Here matters rested until J. K. Hasskarl in 1863 (Annales
Musei Botanici Lugduno-Batavi, I, pp. 142 seq.) gave a mono-
graph of Indian Polygalacee. He examined i.a. the type-speci-
men of Semeiocardium arriensii collected by Zollinger (No. 3956)
bearing on its ticket the note: ‘‘corolla 4-color; vexillum cordatum,
“‘cyaneum, albido-marginatum; saccus bifidus, luteus, purpureo-
‘“‘maculatus’’. The flowers of Zollinger’s specimen being clearly
in a bad state of preservation Hasskarl not only could not find the
sack or spur but moreover was led into the error of mistaking
the plant for a Polygalacea. The genus Semeiocardium, of which
he gave no diagnosis, was retained by him and he brought to it,
besides S. arriensii Zoll., three species from British India. Of
S. arriensii he gave a diagnosis, but his description of the inflore-
scences, flowers, fruits and seeds, apparently taken from another
specimen, is entirely erroneous.

In Hooker’s Flora of British India 1, 201 (1872) Alfred
W. Bennett reduced the three Indian species of Semeiocardium
to Polygala triphylla Ham., a plant which does not occur in
Java, nor in Madura. He did not mention Semeiocardium
arriensit.

R. Chodat in his Monographia Polygalacearum, 2me partie
(Mémoires de la Société de Physique et d’Histoire Naturelle de
Genéve XXXI, Seconde Partie, 1892/93, p. 41), apparently not

Gardens Bulletin, S.S.

ya!

having seen good specimens of Semeiocardium arriensti considered
Semeiocardium to be a section of Polygala. To this section he
reckoned, besides Polygala triphylla Ham., only the insufficiently
known P. cardiocarpa Kurz. Semeiocardium arriensii Zoll. was
reduced by him to Polygala triphylla.

I myself, having seen only very bad materials, committed
the same mistake in my Schoolflora voor Java (1911), p. 77.

But in 1915 I had, thanks to Dr. J. C. Koningsberger, then
Director of the Buitenzorg Botanical Gardens and an ardent pro-
motor of the floristic investigation of Java and surrounding
islands, the occasion of making a botanical trip to Madura. On
this trip I had the good fortune to re-discover the plant evidently
meant by Zollinger, who was, after all, much too able a botanist
to mistake a Balsaminacea for a Polygalacea. The plant, which
showed the characters mentioned by Zollinger in his short des-
cription, and which grew indeed on calcareous rocks, proved to
be a true Balsaminacea, nearly related to Impatiens but differing
from this genus (see below) by the 2 lateral and the 2 posticous
petals being all of them connate into a rather large, bifid
standard, and by the 4-celled (not 5-celled) ovary. In rIg19 a
slightly differing form of the same species was discovered by me
in the Kangean Archipelago (east of Madura). I succeeded in
bringing over living plants and seeds to the Buitenzorg Botanical
Gardens. The seeds germinated poorly, but seeds thrown out
by cultivated plants germinated now and then, always on rocks.
In 1930 plants, offspring of those raised from seeds collected in
Kangean, were still living in the Buitenzorg Gardens.

I give below a description of the genus and the species, based
on living plants.

SEMEIOCARDIUM Zoll. (descriptio emendata).

Cymee axillares paucifloree. Flores more generis Impatientis
resupinati, zygomorphi, bisexuales, majusculi. Sepala 3; duo
lateralia alte connata, tenuiter membranacea; tertium posticum
(antrorsum versum) majus, petaloideum, cymbiformi-
infundibuliforme, breviter calcaratum; calcare disticte bilobo,
antrorsum spectante. Petalum anticum (retrorsum vergens)
liberum; duo lateralia quam cetera minora, fere ad apicem usque
connata cum unguibus posticorum (specie anticorum); laminze
petalorum posticorum connate in vexillum antrorsum vel
deorsum spectans, incisurd angusta fere ad medium usque
bifidum; hisce 4 petalis connatis floris labium inferius biungui-
culatum fingentibus. Stamina 5; filamentis liberis, brevibus,
latis, ovarium celantibus; antheris crassis, in tubum brevem
connatis; loculis apice poro magno dehiscentibus. Ovarium
oblongum, 4-loculare; loculis pauciovulatis. Stylus 1, brevis,
subulatus; capsula lineari-oblonga, 1-5-sperma; valvulis 4 elastice
dissilientibus, semina ejaculantibus. Folia alterna, longiter
petiolata, ovato-lanceolata, leviter serrata. Herba annua, erecta,
patule ramosa; caule valde succulento; radicibus brevibus.

Pon WX (r9.35).

72

Genus Impatienti proximum sed bene diversum petalis 4
superioribus (specie inferioribus) in laminam bifidam, biungui-
culatam connatis, necnon ovario 4-loculari.

Species adhuc unica:

Semeiocardium arriensii Zoll. (see the plate).

Cymeze breviter pedunculatz, 1-3-florze; pedicellis 10-16 mm.
longis. Sepala lateralia oblique deorsum spectantia, mucronu-
lata, pallide viridia, 10-18 mm. longa, 7-11 mm. lata. Sepalum
posticum a lateralibus partim tectum, mucronulatum, album vel
pallide flavum, intus pulchre rubro-striatum et rubro-guttatum.
Calcar inter sepala lateralia occultum, 3-7 mm. longum, latum,
basi et apice angustatum, petalum anticum ovali-ellipticum,
7-11 mim. longum, 6-9 mm. latum, viride, dorso carinatum; carina
acumine recurvo terminata. Petala lateralia quam postica multo
breviora; ungues petalorum posticorum curvati; labium (a petalis
lateralibus et posticis formatum) late ovale, 15-20 mm. longum,
in speciminibus madurensibus pallide violaceum, infra incisuram
cum macula intense violacea, basi seepe notatum punctis rubris
quasi continuationem efformantibus punctorum striarumque sepali
postici (unde error zollingerianum), in speciminibus kangeanensi-
bus candidum, basi luteum. Tubus antherarum recurvum
(specie procurvum). Ovarium glabrum. Capsula_ glabra,
+ 20 mm. longa, 5-6 mm. lata. Caulis crassus, apice pubescens,
deorsum glaber. Petioli 30-70 mm. longi; foliis ovato-
oblongis, basi obtuso-rotundatis, apice acute acuminatis, leviter
serratis, junioribus subpubescentibus, adultis glabris. Herba
erecta, annua, 40-80 cm. alta.

Crescit preecipue in rupibus calcareis.
Distributio:—Madura (Backer 20268, 20911).

Archipelago Kangeanensis (Kangean et Paliat)
(Backer 26962, 27376, 27409, 27863, 27950,
29565, 29966, 30004).

Gardens Bulletin, S.S.

Gard. Bull. S.S. Vol. IX. Plate 2.

Semeiocardium arriensii ZOLL. — Photo made in 1917 after a speci-
men cultivated in the Buitenzorg Botanic Gardens, raised from seeds
collected in Madura.

73

THE FLOWERING OF TEMBUSU TREES (FAGRAA
FRAGRANS Roxb.) IN SINGAPORE, 1928—1935.

By R. E. Hortrum.

The gregarious flowering of the Tembusu trees (Fagrea
fragrans Roxb., or Cyrtophyllum fragrans DC) which are so
abundant, both wild and planted, in the neighbourhood of the
town, is one of the most striking seasonal phenomena to be
observed in Singapore. Beautiful at all times, with their finely
moulded masses of foliage, at the flowering season the trees
are transformed by their host of small creamy-white fragrant
blossoms. For a fortnight or more, some of the suburbs of the
town are filled with their heavy fragrance, both day and night.
This impressive spectacle occurs nearly always in April or May
(usually May), and is followed in August by the almost equally
impressive general fruiting of the trees, accompanied by nightly
visits of hordes of giant fruit-bats. The whole phenomenon
is one of the most regular features of the somewhat irregular
general behaviour of plant life in Singapore. It is fairly certain
that such a general flowering, as in other similar cases, is a
response to some definite climatic change, and the question
naturally arises what such a change could be.

In the Gardens Bulletin S.S. Vol. V, pp. 173-206, I gave
an account of my observations on periodic leaf-change and
flowering in Singapore, chiefly as regards deciduous trees, during
the years 1927-1930. In the introductory part of that paper is
a short account of the salient features of the climate of
Singapore, and to this the reader is referred for further details.
The distribution of rainfall throughout the year, though on the
average very even, is so irregular in detail, and the incidence of
dry weather so uncertain, that many species of plants which are
native of seasonal climates behave quite abnormally. As regards
the native evergreen trees, it appears that any pronounced dry
spell will cause most of them to flower, provided they have had
an appropriate period in which to recover after the previous
flowering and fruiting season. Some trees respond very quickly
to the dry weather, and flower within two or three weeks; others
take much longer to develop their flowers. It is fairly clear
however that most of them are dependent on pronounced dry
spells, and that their flowering is much less regular than that of
the Tembusu.

We have therefore to look for a seasonal phenomenon,
perhaps relatively slight and not always effective on most trees,
which is as regular as the flowering of the Tembusu. ‘The one
almost regular feature of our climate is the rather sudden break in
the wet weather which usually comes some time in January. ‘The
Wet season, which is most intense in November and December,
is characterised by cloudy skies, prolonged slight drizzling

Vol 1 .\\(19 35).

74

rains, and a very high general atmospheric humidity, with little
bright sunshine and no drying winds. he actual rainfall may
not be very heavy, but it has a far greater effect than at other
times of the year, when heavy showers, followed by bright
sunshine, are the rule. Then rather suddenly, usually in
January, the cloudy weather ceases, and there is a shorter or
longer period of drying winds. This drier weather may not
last very long, but its onset, after the rather depressing wet
season, is very noticeable.

The thought therefore occurred to me that perhaps the
Tembusu trees are very sensitive to this change from prolonged
wet to drier conditions, responding even though the dry weather
is not sufficiently intense to affect some other trees. I have
examined the rainfall records made at the Botanic Gardens
during the period of my observation on the trees, and believe
that they indicate a definite relationship between the change of
season and the onset of flowering of the Tembusu. The
irregularities of one are followed by irregularities of the other,
and the last wet season, by stopping at an unusually early date,
gave a most instructive example. Unfortunately the figures
showing change of rainfall do not give a complete account of the
climatic changes; the actual change, as above remarked, is
accentuated by the presence of clearer skies and drying winds.
The figures given are however perhaps enough to demonstrate
a connection between the phenomena concerned.

The Tembusu tree is slow in all its activities. Its growth
is slow, but seems to be an almost continuous process, unlike
that of most evergreen trees, which produce new flushes of leaves
at intervals of a few months (or sometimes many months).
The development of flower buds is slow; though the flowers are
quite small, the buds take something like two months from the
time they are first visible to the opening of the flowers. The
inflorescences are borne in the axils of quite young leaves; at
flowering, the subtending leaves are usually the second pair
from the apex of the shoot. Then the fruits, though small, take
more than three months to ripen (it is usually about three and a
half months from the height of the flowering to the full ripening
of the fruit crop). These slow processes are perhaps connected
with the soil conditions favoured by the trees, which grow best
in the very hard compacted soil formed as a result of exposure
to rain and sun. Under such conditions very few other trees
will thrive. The soil is very slightly pervious to water and air.
I have seen tembusu trees in the neighbourhood of Palembang
(Sumatra) growing in swampy ground, and they also grow under
similar conditions in Pahang; possibly the lack of aeration is
the chief character in common between the swamp conditions
and the compacted clay of Singapore. It is notable that the
palms Oncosperma and Cyrtostachys, both native of wet ground,
also flourish in hard clay soils suitable to the Tembusu. -

Gardens Bulletin, S.S.

a yn

ee

75

In addition to the heavy flowering which is usual in May,
the trees often have a subsidiary flowering towards the end of the
year, in October or November, and occasionally a few flowers
may be seen at other times. I have noted particularly the
behaviour of three individual Tembusu trees from the latter part
of 1927 up to the present (the observations are not quite complete,
especially as regards the subsidiary flowerings, which are
often not easy to observe). I have also noted the seasons of
principal flowering of the trees in Singapore generally, and any
pronounced subsidiary seasons. The main flowerings are
therefore all noted, but not all subsidiary flowerings. Of the
three individual trees, one stands almost alone, and has its lower
branches intact and widely spreading over the ground all around
it; the others are near other trees, one on a hill and the other in
a more sheltered position near the bottom of a slope. They are
referred to as E, X and P respectively.

The exact date of the beginning of flowering is not easy to
fix; I have therefore indicated a date at which a general flowering
was well in progress, perhaps a week after the first flowers open.
As regards subsidiary flowerings, they are so incompletely
recorded that I only quote here a few cases when the flowering
was fairly general, or when one of the trees had rather numerous
flowers at an unusual time.

I have indicated below the dates at which wet weather
ceased, giving rainfall for the previous ten days, and for the
week or more of dry weather following. It will be seen that
general flowering is in progress about four months after the
onset of the dry weather. To test whether the flowering dates
could be connected with the length of the dry period, or with
the time at which wet weather followed the dry, I examined the
subsequent rainfall figures, but the incidence of subsequent
rainfall and its amount are so variable that it appears most
unlikely that it can have any effect on the time of flowering. As
regards sudden dry weather following wet at other times of the
year, this is rarely so pronounced as in January, as heavy rainfall
at other seasons is usually of the heavy shower type; figures of
relative humidity and perhaps of duration of sunshine would also
have to be added to give a full account of the climatic
conditions. However, I have traced some of the more
pronounced subsidiary flowerings to dry weather following
heavy rain, and these are indicated.

As the time from the stimulus to flower-development to the
ripening of fruit is about seven months and a half, and as it is
unlikely that further stimuli during that period could cause
anything in the nature of a second general flowering, it is most
unlikely that two full flowerings could occur in any calendar
year; a second flowering would ‘alw ays be a subsidiary one, and
probably affects those trees or branches which have for some
reason not been affected by the main flowering. Flowering on

Vol. IX. (1935).

76

separate branches is a not uncommon phenomenon on trees of
various kinds in Singapore; also leaf-change on one branch at a
time.

In connection with the following figures, it may be noted
that the average rainfall of the wettest month in the year
(December) is about 12 inches, while May and June, which on
the average are the driest months, have about 6 inches each;
but the rainfall in any one calendar month varies considerably
from year to year.

PRINCIPAL FLOWERINGS.

1928
Rainfall January 5-14 7.99 inches
I5—21 14
Flowering: May 12
1929
Rainfall December 28 (1928)—January 6 5.62 inches
January 7-13 15
Flowering: May 5.
1930
Rainfall January 3-16 4.01 inches
17-26 .07
Flowering: May 20
1931
Rainfall January 15-22 5.22 inches (Jan. 12-14, 4.28 ins.)
January 25—February 2 .24 inches

Flowering: middle of May.

1932
Rainfall December 21-30 (1931)
December 31—January 5 .07
Flowering: May 1.

1933
Rainfall January 16-25 4.19 inches
January 26—February 1 .O4
Flowering: May 27.
1934
Rainfall December 17-26 (1933) 3.81 inches
27—-January 2 .0O
Rainfall January 12-21 Sy.
22-Feb. 3 .0O

Flowering: April 25—June ro.°

Note:—The duration of flowering was recorded by
Mr. Corner, and he did not note whether the main flowering
began on April 25th. As a heavy flowering usually lasts for at
least three weeks, two flowerings, one caused by dry weather

Gardens Bulletin, S.S.

77

beginning December 27, and the other January 22, would produce
the effect of a continuous flowering. The dry period December
27-January 2 was followed by more heavy rain, and then a more
intense dry period, which evidently caused a second flowering.

1935
Rainfall November 28—-December 7 (1934) 3.44 inches
December 8—27 .07
Flowering: April 7.

SUBSIDIARY FLOWERINGS.
1928

Rainfall September 30—October 9 (1927) 6.33 inches
October 10-21 sE2
Small flowering on trees E & X, 19 February 1928.
Rainfall July 3-12 4.89 inches
13-30 ‘12
Small flowering on trees EK & P, also general small flowering
about November Ir.

1929
Rainfall June 19-28 6.24 inches
26-July 12 .02
Small general flowering October 27.

1930
Rainfall November 26—December 5 (1929) 3.68 inches
December 6-15 .20

Tree X with many flowers on low branch, and tree P with
some flowers, April 20 and 26.

Rainfall June 13-22 8.62 inches
23—-July 5 .00

October flowering not noted, but tree E had many fruits
ripening early in February 1931.

1931
Rainfall June 2-11 4.34 inches
12-20 a1
Tree X fairly full flowering in October.
1933
Rainfall November 30—December 9 (1932) 3.29 inches
December 10-19 sb7

Small general flowering April 16, 1933.

Vol. £X.\\(.19035,).

78

1934

Rainfall July 16-27 6.95 inches
28—-August 16 38

Many trees have small flowering in early December.

1935

Rainfall December 30 (1934)-January 8 — 7.56 inches
January 9-13 .00

General small flowering, May 16.

The year 1935 was particularly interesting for two reasons.
In the first place, the general flowering was a full month earlier
than usual, following upon a most unexpected break in the wet
season in the second week of December. In the second place,
there was a quite well-marked second small flowering considerably
more than a month after the first, and this corresponded to a
second and more severe onset of dry weather, following much
rain in the latter part of December and early January.

The above information needs to be supplemented by more
detailed observations of the vegetative growth of Tembusu trees.
What is required is regular measurement of the growth of all
parts of a small branch, and also of the development of the
inflorescence. It would be interesting also to make careful record
of subsidiary flowering on a few individual branches, to find the
minimum interval between two flowerings on the same twig.

It is quite likely that some other trees respond to the change
of season in a similar manner to the Tembusu; but the only
records I possess are of a much less regular nature. Fagrea
auriculata, for example, sometimes flowers with the Tembusu and
sometimes a full month later. The common scorpion orchid,
Arachnis flos-eris, though flowering very regularly about “May
and November, does not appear to follow the fluctuations of the
Tembusu. Saraca taipingensis and Cinnamomum iners, both of
which flower gregariously not long after the change of season, are
evidently more rapid in their response, and probably do not react
to quite the same stimulus as the Tembusu, for I can find no
relation between their flowering dates. Possibly some of the
common species of Eugenia and Eleocarpus may have a similar
response, but insufficient records of these are available.

SUMMARY.

In Singapore there is an almost regular general heavy
flowering of trees of Fagrea fragrans Roxb. in May. This
flowering is shown to follow about four months after the break
in the wet season which usually occurs some time in January.
Observations of rainfall for ten days before the break in the
weather, and for a week or more afterwards, are given; the
fluctuations of date of this break are followed closely by
fluctuations in the date of the general flowering of Cyrtophyllum
trees.

Gardens Bulletin, S.S.

79

THE SEASONAL FRUITING OF AGARICS IN MALAYA.
By E. J. H. Corner,

Singapore Botanic Gardens.

There is no day in Malaya when the fruit-body of some
agaric cannot be found in the high forest, but twice each year,
in the south at least, there is a season of three months when a
“run of fungus’? develops like the autumn crop in temperate
countries. Fruit-bodies, never seen at other times, then appear
in great numbers with a succession of early, mid- and late
seasonal forms. ‘The forest may be so full of toadstools of all
sizes, shapes and colours, in troops on the ground and on fallen
wood and in myriads on dead leaves and sticks, that one cannot
step without treading on them, and each kind has its season of
but a few days or a week when it is in full fruit.

I learnt of these seasons first in the Gardens Jungle, over the
few acres of which I searched almost daily in the year 1929, and
then from repeated collecting in the forests of Singapore and
south Johore and from occasional visits to more distant parts of
the country. In the Gardens Jungle, also, I set myself to dis-
cover the exact times of inception of fruit-bodies and their periods
of development and maturity so that I could determine the life of
different kinds of fruit-body and the climatic stimulus to which
their mycelia responded. I have collected much information on
nearly a hundred species but only a few general facts can be
mentioned in this place.

THE SEASONS.

In Singapore the first toadstool-season of the year extends
from March to May and the second, less regularly, from some
time in August or September, rarely October, to October or the
beginning of December. From 1929 till the beginning of this
year Boleti, for instance, fruited chiefly in March and August or
September; Hygrophorus firmus B. and Br. fruited in April and
October or November; Amanita virginea Mass. in May and
November or the beginning of December. ‘The lowlands of south
Johore appear to have the same seasons though the dates may
differ by two or three weeks through local variations in the
weather. At Tembeling, in the centre of Pahang, a run of
fungus was in mid-course at the beginning of November 1930,
and one had just ended at the beginning of June 1931. On
Fraser’s Hill, at 4000’ in Pahang, a run had just ended in the
middle of November 1930 and 1932. About 20 miles inland from
Chukai, Kemaman, there was no run of fungus at the end of
June 1932, though the forest was very wet. In Penang a run was
starting at the beginning of March 1929. ‘These dates agree with
the seasons in Singapore. But in Perlis, in the north, a run was
starting at the end of November 1929: at Slim River, Perak, and

Vol. IX. (1935).

80

on Fraser’s Hill, Pahang, a run was starting at the beginning
of May 1930: round Kuala Pilah and on the lower slopes of
Gunong Angsi, in Negri Sembilan, a run was starting in the last
week of June 1930: at Cameron Highlands, at 5000’ in Pahang,
a run was finishing at the beginning of August 1934. ‘These
dates disagree, yet one may be certain that, while there is a
seasonal fruiting in all parts of Malaya following changes in the
weather, the date of the season in each place depends on local
conditions, for the times of the weather-changes vary considerably
not only in the lowlands but from mountain to mountain.

THE SUCCESSION.

Without a systematic account of malayan agarics, only an
outline of the seasonal succession can be given. Briefly it has
this order :—

1st Crop :—epiphyllous species of Marasmius, Andro-
saceus, Mycena, Collybia, etc. with small fruit-bodies : some
horse-hair Marasmius: Coprinus, Psathyrella and

Leucocoprinus.

2nd Crop :—developing 4-8 days after the first: most
kinds with large fruit-bodies but especially Amanita,

Amanitopsis, Lepiota, Collybia radicata, Laccaria, Lactarius,

Russula, Volvaria, Pluteus, Clitopilus, Leptonia, Inocybe,

Astrosporina, Cortinarius, Hypholoma janus (B. and Br.)

Petch, Psathyra_ trechispora Petch, Psaliota, Boletus,

Paxillus, Lentinus giganteus B.

3rd Crop :—about two weeks after the second: Hygro-
phorus, Cantharellus, Pleurotus, Entoloma, Clitopilus
flavidus Mass. (really an Entoloma), Lentinus connatus,

L. blepharodes, L. hookerianus, L. similis, and other horse-

hair Marasmius.

4th Crop :—about two weeks after the third: Hygre-
phorus firmus B. and Br. and several miscellaneous species.

5th Crop :—about 2-4 weeks after the fourth: Amanita
virginea Mass. and many species of Hygrophorus and

Marasmius.

The crops are not actually distinct but peaks, rather, on a
summation curve in which the number of species fruiting in a
locality and the number of fruit-bodies would be plotted against
time, and with fewer, less conspicuous species fruiting in the
intervals. ‘he succession is so constant under normal conditions
that a few easily recognisable species serve as indicators :—

Crop. Indicator.
Ist .... Epiphyllous Androsaceus and _ Filoboletus
(= Poromycena)
and =... Amanita, Russula, Boletus in great variety
3rd ... Clitopilus flavidus Mass.
4th ... Hygrophorus firmus B. and Br.
5th ... Amanita virginea Mass.

Gardens Bulletin, S.S.

81

Amanita virginea, like Collybia velutipes in England,
signifies the end of the season.

THE CAUSE OF THE SEASONS.

In most of Malaya rain falls so frequently that the forest is
nearly always damp. A week without rain scarcely affects the
humus. But twice each year, after the wet weather of November
and December and April to June or July, there comes a spell of
several weeks with sunny, windy days and only occasional
showers. The forest then dries; the humus becomes friable and
brittle; streams shrink in their beds; and the leeches disappear.
Mycelial growth in the soil gradually stops and the hyphe remain
dormant for a week or more. At the next rains the hyphz revive
and fruit. Thus spring the first two crops of toadstools which
begin the succession.

It may be thought, now, especially from the evidence of
fungi grown in culture, that a mycelium would fruit so soon
as it had gathered enough food, but I am convinced direct fruiting
is exceptional among forest fungi and pertains more to species
of transient or confined habitats, like coprophytes and carboni-
colous forms. This succession in the rain-forest shows that most
forest-species need an external stimulus to fruit, which is
primarily the check to vegetative growth given by mild drought.
For, two or three days after the forest floor is soaked again, fine
rhizomorphs begin to grow in the humus and there is the mouldy
smell of active mycelium. A few days later the primordia of
Russules, Boleti and Amanitas appear and these, I find, take only
8-1o days to mature. There is, therefore, no time between the
onset of the rain and the fruiting for vegetative growth. Indeed,
the first mycelial growths of these fungi after the soaking of the
soil are slender rhizomorphs—somatic strands, not diffuse feeding
hyphe.

The relation between the dry weather, the onset of the rain
and the crops of Boleti in the Gardens Jungle is shown in the
following table. The period of the dry weather is taken from the
daily rainfall record in the Botanic Gardens and measured from
the end of continually wet weather to the heavy rain which caused
the resumption of mycelial growth in the soil. Actually it is
an interrupted period of dry rather than wet weather, and single
storms and light rain on a few successive days I have reckoned
in the dry weather because I found, by carefully digging in the
fhumus where toadstools always grew, that such rain never
penetrated or lasted long enough to evoke the development of
primordia. By digging in the humus after each likely storm
toward the end of the dry weather, I determined to a few days
When mycelial growth was resumed.

[Vol. IX. (1935).

F

: Pee é Mature
Year Dry Weather Rain begins Boletus-crop
1929 6. 7th-«sJan.—8th March oth March 24th March
29th June—23rd Aug. 24th Aug. roth Sept.
1930 20th Jan.—6th March 7th March 22nd March
23rd June—6th Sept. 6-14th Sept. 24th Sept.
15—27th Sept. 28th Sept. 16th Oct.
I93I 26th Jan.—28th Feb. Ist March 18th March
roth June—18th July roth July 7th Aug.
1932 «Ss ¥st' —1Sth Jan: 19-26th Jan. 8th Feb.
(small fruiting)
27th Jan.—16th Feb. 17th Feb. 6th March
(heavy fruiting)
roth July—2nd Sept. 3rd Sept. 20th Sept.
1933 26th Jan.—23rd Feb. 24th Feb. 14th March
No record in second half
of year
1934 20th Jan.—27th Feb. 28th Feb. 20th March
4th Sept.—8th Oct. oth Oct. 27th Oct.
1935 6—25th Dec. 1934 26th Dec. 1934 20th Jan.
—21st Jan.
22nd Jan.—12th Feb. 13th Feb. 6th March
oth July—oth Aug. roth Aug. 31st Aug.

Two crops of toadstools immediately follow the rain because
the fruit-bodies of the epiphyllous species of the first crop are
small and develop more quickly than the larger ones of the second
and, moreover, growing on the surface of the humus, their
mycelia revive sooner than the more deeply situated and, perhaps,
mycorhizal mycelia of the second crop.

The toadstools of the third crop, which mature about 4-5
weeks after the wet weather has returned, may need to vegetate
before fruiting or they may be delayed merely because their
mycelia recover more slowly or take longer to develop rhizo-
morphs than those of the early crops: in general, their primordia
develop at the same rate as those of Boleti. The toadstools of
the fourth and fifth crops must have a preliminary vegetative
stage yet they do not fruit indiscriminately as shown by their
constant sequence.

Gardens Bulletin, S.S.

83

Heavy rain must follow the dry weather to revive the
mycelia but rain in itself is no stimulus to fruit. In June and
December, when the forest is wettest and mycelia have been
growing during several months of rainy weather, fruit-bodies
are scarce. ‘Toadstools of the second crop, which fruit about
August, do not fruit in December though they have been
vegetating for four months.

Many epiphytic orchids, it is well known, flower after a
heavy storm not because of the wetting but because of the
sudden drop in temperature during the storm. There is no
evidence, however, that agarics respond to this stimulus. The
drop in temperature must be appreciably less, both in degree and
severity, in the humus than on the boughs of trees and, were
the stimulus effective, runs of toadstools would be as frequent
as the flowering of pigeon-orchids (Dendrobium crumenatum).
I have, moreover, grown lignicolous agarics in a cool frame,
where the temperature varied very gradually from 73-78° F.
daily, and found that by drying the logs for two weeks and then
soaking them fruit-bodies would develop without any marked
change in temperature. And it seems that, except the highest
mountain tops, nowhere in Malaya is the temperature low
enough to limit the fruiting or the distribution of toadstools.
All the common lowland species occur in the mountains at
4000-5000, where the soil-temperature in the forest is, on the
average 60-70° F. and 10-15° below that in the plains.

Some of these agarics, especially the Russules, Boleti and
Amanitas, must be mycorhizal. In forests which I have
repeatedly visited they occur season after season in the same
places under the same trees, among which dipterocarps, oaks
and Castanopsis are prevalent, and not elsewhere. ‘They should
fruit, one might imagine, according to some general activity of
their host yet, as they do so at the same time or in sequence with
saprophytic species, their mycelia must react like the
saprophytic. Most forest trees respond to one or both dry
spells of the year by flowering or leafing so that a run of fungus
coincides with a flowering season, but without exact knowledge
of the habits of the trees or of their mycorhizal fungi the ways
of the two cannot be related. A remarkable Russule, for
instance, I found in fruit in the mid-weeks of April and October
In 1930, 1931 and 1932 only in one small area, about 30 yards
across, on the lower slopes of Gunong Panti in Johore; it is
clearly a seasonal, mycorhizal species, but its host cannot be
determined by inspection for some twenty kinds of tall tree grow
in proximity.

THE SHORTNESS OF EACH SEASON.

During a normal year in Singapore and, probably, in any
one locality in Malaya, Boleti will fruit for two weeks, the first
in March, the second about September, and a botanist visiting
the forest at other times could have no inkling of their existence

Vol. IX. (1935).

84

apart from a few species which fruit sporadically. (I have
collected about 60 species without particular search). So, too,
with other forest agarics: their seasons are remarkably short.
Unless one can visit the forest at least once a week during a run
of fungi many species, if not an entire crop, may be missed not
merely because each has its place in the succession but because
the fruit-bodies develop rapidly, especially through the later
stages of expansion when they emerge from the humus, and
they are short-lived. Those of Boleti, Paxillus, Flammula,
Inocybe and Hygrophorus last about 4—5 days at maturity; those
of Entoloma, Cortinarius and Lentinus may last a week or ten
days; those of Amanita, Lepiota, Collybia, Russula, Lactarius,
Volvaria, Psaliota and Psathyra last only 2-3 days: though there
are a few exceptions such as two Boleti which I find last nearly
three weeks.

When Patouillard came to Singapore on August 21st 1917
(Journ. Straits Branch R.A. Soc. no. 78, p. 67), there was a
Boletus season in the Gardens Jungle and he described sixteen
new species, though only seven had not already been collected
by Ridley and Burkill in the same place and described by
Massee (Kew Bull. 1901, 1909, 1914). But it could never have
befallen Westerdijk that he went into the forest during a toad-
stool-season, when he wrote of the malayan region ‘‘In the virgin
WOOGS). cle. soe even the flora of mushrooms on the ground
Latin ce te EE is absent. Everything seems to point to the con-
clusion that conditions are unfavourable to fungus growth’’
(Ann. Miss. Bot. Gard. 1915, 2, 308).

IRREGULARITIES IN THE SEASONS.

A run of toadstools in the forest in a sure sign of a climatic
change. ‘he weather must have been dry enough to have
affected the humus and the ensuing rain heavy and prolonged
enough to have soaked it. A dry week might stimulate trees to
flower or change their leaves and a storm might cause the
pigeon-orchid to flower but the effects of such would not
penetrate the dense canopy of the forest. Nevertheless there
are minor irregularites in the toadstool-crops, which appear to be
caused more by topographical features than climatic. A small,
isolated and deteriorated patch of forest, such as the Gardens
Jungle or the Bukit Timah Forest Reserve in Singapore, 1s ex-
posed and a short dry spell or single storm might affect the humus;
the wind could blow through such forest and dry the soil in weather
which would not affect the continuous virgin forest. Thus I
have noticed that there may be two crops of Boleti at the
beginning of a season. In the Gardens Jungle in 1930, after a
dry July and August, a great many Boleti began to develop
about the 16th Sept., following a rainstorm on the 6th with
showery weather in the ensuing week. But from the 15-27th
the weather turned dry and windy and the floor of the jungle
soon dried out; most primordia shrivelled and only a very few

Gardens Bulletin, S.S.

85

came to maturity. Rainy weather followed from the 27th, with
a heavy storm on the 28th, and the normal run of fungi began
in October with a second, rather feeble, crop of Boleti about 16th
October. It is unlikely that the rain from the 6—14th September
would have soaked the humus in virgin forest or, if sufficient,
that the following dry spell would have checked the new growth
of mycelia, and there would thus have been the one normal
succession. For the same reason there were two crops of Boleti
at the beginning of 1932 and 1935; and the first crop of 1935 was
exceptionally early because the wet weather broke a month
before usual. In each of these instances only the first two crops
of the succession were repeated; presumably the dry weather
which returned after the first heavy rain prevented the mycelia
of species of the subsequent crops from reviving, and one might
prefer therefore a late-seasonal species as Hygrophorus firmus
to indicate the seasonal run. Similarly, the crop of epiphyllous
agarics may occur out of order: in the Gardens Jungle
there may be a small crop in June because the surface of the
humus dries in a few windy days and a single heavy storm may
penetrate the rather light canopy and soak it. But how often
or at what short intervals of dormancy or vegetative growth the
toadstools of the first two crops can be made to fruit under such
circumstances I cannot state. Nor is it known how mycelia
remain dormant in dry soil; few species have sclerotia.

Observations on crops of Boleti, as the most easily
recognised indicators of a succession, would in the virgin
mountain forest give a valuable clue to the climate. For
instance, in the ridge-forest on Gunong Panti in S.E. Johore I
have never seen a run of fungus though I have visited it in most
months of the year and when there has been a run on the lower
slopes: it is a long, narrow and disconnected mountain about
1600’ high and its climate, which must be intermediate between
that of the East and West coasts, is generally wet.

SPECIES IN SECONDARY HABITATS

The environmental factors controlling the growth of fungi
are much more equable in the high forest than in the open or in
secondary jungle, and the fruiting of agarics in such places may
be irregular. All such places, in fact, as lawns, pastures,
orchards, estates and other clearings, must be considered
secondary habitats because the land was originally covered with
high forest and malayan fungi are primarily forest dwellers.
Agarics of secondary habitats fruit also in crops in response to
the same stimulus as the forest species but, because of the
openness of their habitat, their mycelial growth is checked more
readily by short dry spells and promoted again by lighter and

Vol. IX. (1635).

86

less prolonged rain. Of this kind are several species of Hygro-
phorus related to H. puniceus and H. conicus, Lepiota,
Marasmius, Clitopilus, Leptonia, Galera, Naucoria, Coprinus and
Psaliota on lawns or pastures, and of Pleurotus and Pluteus
together with Schizophyllum commune, Collybia apalosarca,
Flammula dilepis, Lentinus sajor-caju, L. subnudus, and
L. Lecomtei on logs in the open. ‘They do not fruit continuously
in wet weather nor. will they fruit without an interval for vegeta-
tive growth after each crop. As already mentioned, I have
proved this by growing several of the lignicolous kinds under
equable conditions: no fruit-bodies grew when the logs were
kept continuously damp during 3—6 months but, when they were
dried for 10 days and then sprayed, an abundant crop was
obtained. Indeed the best way to study lignicolous agarics is
by collecting logs from the forest in the wettest weather, drying
them gradually in a shady place for 10 days and then spraying
them frequently: the variety of basidiomycetes which develop
is often incredible.

The only agarics which I have found fruiting continuously
under wet conditions are coprophilous ones, which grow and
fruit till their habitat is exhausted, and some lignicolus species of
Mycena, Collybia, Marasmius and Pleurotus with small fruit-
bodies, yet even they respond to the stimulus of drying and
wetting.

THe TERMITE-AGARIC, COLLYBIA ALBUMINOSA (BERK.) PETCH.

So irregular is the fruiting of this species that I can form no
opinion about it. As Petch noted (A. R. B. Gdn. Perad. 19713,
5,323) it fruits from inhabited nests, but it seems not to fruit
more than once from the same nest, at least in Singapore where,
however, most nests are rather small, and so one cannot watch
the same place repeatedly. It fruits at any time of the year,
even in the dryest weather on lawns which are browning, and
though there are many nests in the Gardens Jungle from which
it could develop it gives no special crop in a seasonal run of
toadstools.

THE FRUITING OF OTHER BASIDIOMYCETES.

It appears that all kinds of Basidiomycetes, fruit in Malaya
according to their season exactly as the agarics, but I have few
detailed observations on others. The rain which starts a run of
agarics staits also the fruiting of the woody Thelephoraceze, Hyd-
naceze and Polyporacez, most of which become noticeable rather
late in the succession because their fruit-bodies grow slowly, though
a few, like Favolus and some Hydnums and Polypores, grow fast
and come with the second crop of agarics. The dry weather

Gardens Bulletin, S.S.

87

checks also the growth of the perennial fruit-bodies of Fomes,
Stereum amd Hymenochezte and they form a new layer of tubes
or hymenium in the wet weather: hence, in a normal year,
these fruit-bodies develop two layers of tubes or hymenium,
unless they have grown in secondary habitats when they may
develop more numerous and narrower layers. Even the
resupinate fruit-bodies of Poria, Corticium and Odontia become
stratified in this way.

Most Clavarias fruit with the second or third crop of agarics,
but the coriaceous fruit-bodies of Lachnocladium generally
develop later with the fourth crop. There are Clavarias,
however, which fruit late in the season.

Lycoperdon, Calvatia, Scleroderma, Mitremyces and the
subterranean forms, as Hymenogaster, Hydnangium, Leucogaster
and Dendrogaster, fruit with the second or third crop of agarics:
the fruit-bodies of some develop as rapidly as those of Amanita.
The Nidulariaceae, Geasters and Tylostoma exasperatum fruit
later with the fourth or fifth crops, but I have not been able to
watch them. In the forest at Tembeling, in Pahang, during the
second and third weeks of November 1930, there was a remarkable
fruiting of some ten species of Geaster : the fruit-bodies developed
mostly on the deep humus between the buttresses of tall trees and
at the base of some the humus was almost covered: yet most of
these I have not seen elsewhere in Malaya or, indeed, another
such crop at any time. The season of the phalloids I do not
know. Dictyophora, in the Botanic Gardens, fruits irregularly
With the toadstools of the lawns: other kinds I have found too
seldom to generalise.

Most Auriculariaceae, Tremellaceae and Caloceraceae fruit
early in the succession, with the second and third crops of
agarics, but there are some which fruit late.

COMPARISON WITH TEMPERATE COUNTRIES.

Of the genera of agarics common to tropical Asia and the
north temperate region, those are best represented in the tropics
Whose species fruit chiefly in the late summer or early autumn
in the temperate region. Dry weather in May or June in England,
followed by rain in June or July, will the cause the fruiting in
July or August of many species of the following genera :—
Amanita, Amanitopsis, Lepiota, Hygrophorus, Laccaria, Russula,
Lactarius, Marasmius, Cantharellus, Clitopilus, Leptonia, Ento-
loma, Pluteus, Inocybe, Astrosporina, Psaliota, Hypholoma
(sect. H. appendiculatum), Psathyra. Boletus and Paxillus.
And these genera are well represented in Malaya, there being,
for instance, several species related to Amanita caesarea of
southern Europe, and malayan toadstools in general resemble

Vol. IX. (1935).

88

those of the southern United States. In contrast, Tricholoma,
Clitocybe, Collybia, Pholiota, Cortinarius, Hebeloma, Naucoria,
Tubaria, Flammula, Stropharia, Hypholoma (sect. H. fasciculare),
Gomphidius, Psilocybe and Panaeolus, whose species fruit mostly
in late autumn and early winter in Europe, are poorly re-
presented. I have not seen a single specimen of Tubaria or
Gomphidius in Malaya (G. roseus Mass. non (Fr.) Quel.,
described from Singapore, is probably Paxillus lateritius Petch
and, certainy, no Gomphidius) and only one Phlegmacium,
one Hebeloma and one Psilocybe. Gomphidius, however, is
typically associated with forests of Abietineze and these are absent
from Malaya.

It seems that the agarics of temperate countries can be
divided roughly into two groups :—1. those which are related
to tropical species and which fruit, like them, at relatively high
temperatures in response to summer drought followed by late
summer rain; 2. those which are truly temperate and fruit at
relatively low temperatures in late autumn, winter or spring and
perhaps only after night-frosts or prolonged cold rain.

SUMMARY.

Each normal year agarics have two general fruiting seasons
in south Malaya, perhaps over the whole country. A succession
of species then develops for some three months. Each has its
own short season of about a week in full fruit and most,
especially Amanita, Russula and Boletus, fruit at the beginning
of the succession.

The first season beings in March, the second about August
or September. Both follow immediately the change from dry
to wet weather. The stimulus to fruit is primarily the dry
weather which checks mycelial growth, and ensuing rain causes
the development of fruit-bodies. Seasonal species do not fruit
in prolonged wet weather.

Observations on crops of Boleti in the forest would give
valuable evidence of the local climate.

Agarics in open places fruit more often because their habitat
is more exposed to alternate drying and wetting.

Most other basidiomycetes fruit in the same _ seasons.
Perennial fruit-bodies normally develop two new layers each
year.

Malayan agarics in their kind and manner of fruiting
resemble the summer and early autumn agarics of temperate
lands.

Gardens Bulletin, S.S.

-_

89

A FEW ORCHIDS FROM THE MALAYAN
ARCHIPELAGO.
By J. J. SMITH,
| Oegstgeest, Holland.
Oberonia (sect. Hymenobractea, incl. Platystreptus) Latifil

WW ).-S:; ap.

Caules approximati, elongati, compressi, serpentini, cum
alis vaginarum recti, 12-21 cm. longi, cum vaginis c. 1.2 cm. lati,
10-21-folii. Folia inarticulata, erecto-patentia, plerumque plus
minusve falcatula, basin versus decrescentia, summa etiam
minora, breviter acuta vel brevissime acute acuminata, parte
patente elongato-triangula, basi haud vel vix fissa, 2.1-3.3 cm.
longa, 0.6-0.8 cm. lata, cum vagina 4.2-4.6 cm. longa, vaginis
sensim in folia vergentia, 0.75-0.8 cm. latis antice fissis, basi
tantum tubulosis. Inflorescentia elongata laxe multiflora, c.
20-22 cm. longa, angulata, papillosa. Bracteze oblong, versus
apicem angustate, irregulariter eroso-fimbriato denticulate,
concave, c. 0.225 cm. longe, fere o.1 cm. late. Flores in
pseudoverticillis 5-6-meris, in inflorescentize apice basique irre-
gulariter dispositi, patentissimi vel subpatentissimi, parvi,
c. 0.2 cm. diam., sepalis petalisque patentissimis, apice leviter
recurvulis. Sepalum dorsale ovatum, obtusum, convexum, basi
concavum, bene o.1 cm. longum, 0.06 cm. latum. Sepala later-
alia ovata, acutiuscula, convexa, bene o.1 cm. longa, 0.07 cm.
lata. Petala divaricata, leviter recurvula, oblonga, obtusa, basi
paululum angustata, convexa, margine papillosa, fere 0.1 cm.
longa, 0.04 cm. lata. Labellum basi auriculis oblique rotundatis
gynostemium semiamplectens, concavitate antice rotundata ad
basin, 3-nervium, antice in lobum medium cum auriculis angulos
rotundato-obtusos facientem ambitu quadrangulum apice leviter
angustatum angulato-retusumque margine papillosum con-
tractum, expansum o.r cm longum, ad auriculas fere 0.1 cm.
latum, lobo intermedio 0.06-0.07 cm. longo, 0.05 cm. lato.
Gynostemium breve, rectum, a dorso compressum, dorso con-
vexum, cum anthera 0.04 cm. longum. Anthera cucullata,
transverse ovalis. Pollinia 4, in corpuscula 2 ovalia plus minusve
obovata unita. Stigma transversum, semilunatum. Ovarium
densius patentissime papillosum, 0.06 cm. longum; pedicellus
minus dense papillosum, 0.16 cm. longum.

Sumatra :—West Coast, environs of Kajoe Tanam, 150 m.
(S. M. Latif n. 13, November, 1934).

This belongs to the group of species on which Schlechter
based his sections Hymenobractea and Platystreptus. In habit it
approaches Hymenobractea, in the shape of the flowers Platy-
streptus, but it is questionable whether these groups, even when
united, may be retained as a section.

O. Latifii is a near ally of O. monstruosa Lndl. but the leaves
are much more numerous and denser and the lip much less
contracted.

Vol. IX. (1935).

go

Description from herbarium specimen, for which I am
indebted to Mr. Latif.

Dendrobium (sect. Calcarifera) Paathii J. J. S. in ‘‘De Orchidee’’
IV (1635) 163;

Caules elongati, deinde ramosi. Inflorescentize e nodis
caulium defoliatorum, ut videtur semper uniflore, pedunculo
tenui c. 1.3 cm. longo. Bractea adpressa, triangula, c. 0.26 cm.
longa. Flores conspicui c. 4.4 cm. longi, 2.75 cm. lati. Sepalum
dorsale erectum, recurvulum, convexum, oblongo-ovatum, apice
angustatum, obtusum, I-nervium cum nervis minoribus inter-
mixtis, c. 1.85 cm. longum, I cm. latum. Sepala lateralia lacinia
conspicua oblique lineari-oblonga concava ad pedem gynostemii
decurrentia, marginibus anticis versus pedis apicem contigua
longitudine c. 0.45 cm. connata, mentum magnum calcariforme
cum ovario angulum acutum vel subrectum faciente fere rectum
ultra medium leviter recurvulum apice abrupte incurvum
truncato-obtusum antice canaliculatum apice antice bilobulum
2.4 cm. longum apice 0.525 cm.latum formantia, parte libera
recurva, oblique ovato-triangula, obtusa, marginibus leviter
curvatis, c. 8-nervia, c. 1.95 cm. longa, margine antico 3.65 cm.,
basi 2.5 cm. metientia. Petala apice recurva, laxe undulata,
oblique rhombea, obtusa 3-nervia, nervis exterioribus bene ramosis,
bene 2 cm. longa, 1.15 cm., basi 0.33 cm. lata. Labellum basi
longitudine c. 0.6 cm. pedi gynostemii adnatum, deinde arcuato-
recurvum et incurvum, subparallelum pedi gynostemii erectum
unguiculatum, cuneato-spathulatum, 3-lobum, lobis lateralibus
erectis concavum, leviter undulatum, gynostemium superans, sub-
tus tota longitudine sulcatum, ungue ad basin partis liberze incras-
satione transversa carnosa margines unguis connectente postice
in crura 2 basin versus producto antice fimbriato instructo, costa
brevi longitudinali (glandula) inter crura, intus ante incrassa-
tionem ad basin laminz verruculosum, 3-costatum, costis approxi-
matis angustis convexis simplicibus, costa media basin lobi inter-
medii usque producta, lateralibus humilioribus brevioribusque inex-
planatum 3.85 cm. longum, explanatum fere 4 cm. longum, ad
lobos laterales 2 cm. latum, ungue toto c. 1.2 cm. parte connata
0.8 cm. longo; lobi laterales expansi sinibus subrectangulis
plicaque introrsa a lobo intermedio separati, oblique triangulli,
obtusi, margine antico leviter rotundati, costula inconspicua intus
marginem lateralem mesochylium cuneatum antice truncatum
infra apicem satis distincte contractum 2.1 cm. longum apice 2
cm. infra apicem 1.6 cm. slatum formantes lobus intermedius
erectus, leviter sigmoideus angulato-convexus, anguste
triangulo-oblongus, obtusus, basi margine minute transverse
rugulosus, carnosulus c. 1.1 cm. longus, basi 0.6 cm., medio
0.73 cm. latus. Gynostemium breve, oblique conicum, dorso
convexum, absque anthera 0.5 cm. longum, clinandrio excavato,
transverse quadrangulo, filamento satis longo, vix curvulo lineari,
auriculis porrectis, verticalibus, filamento cquilongis, ovato-
triangulis, breviter acuminatis, acutis vel obtusis. Anthera

Gardens Bulletin, S.S.

gI

cucullata, oblique cubica, dorso transverse gibbosa, postice
canaliculata, apice producta truncata papilloso-hirtella, c. 0.4 cm.
longa, 0.3 cm. lata. Pollinia 4, lateraliter compressa anguste
obovato-pyriformia, 0.3 cm. longa. Rostellum appendice semor
biculari margine incrassato (viscido?) basin stigmatis obtegente
instructum. Stigma conspicuum, basin gynostemii _ fere
attingens oblongo-quinquangulare circiter dimidio inferiore
contractum. Pes gynostemii cum ovario angulum acutum vel
subrectum faciens elongatus leviter sigmoideus, apice bene
incurvus et ungui labelli adnatus, linearis canaliculatus apice
incurvo magis excavatus et glandula costiformi longitudinali
donatus 2.45 cm. longus. Ovarium teres 6-sulcatum, 0.9 cm.
longum, cum pedicello tereti leviter sigmoideum.

Borneo:—Westerafdeeling, Goenoeng Raja near Singkawang.

(J. J. Paath, April, 1935; ‘‘sepals and petals white, lip yellow,
streaked orange in the throat, midlobe white’’).

A very distinct member of the section Calcarifera of which
I do not know a very near ally.
Description from 2 flowers preserved in alcohol.

Dendrobium (sect. Distichophyllum) singkawangense J. J. S. n.
sp.
Caules approximati, c. 20 cm. longi, flexuosi internodus c.
1.7 cm. longis. Folia patentissima, apice recurvula, oblonga,
apice oblique rotundato-bilobula, ut videtur initio nigro-pilosa,
costa media supra sulcata subtus prominente, c. 4.3 cm. longa,
2.3-3.3 cm. lata; vaginis internodia zequantibus.

Inflorescentiz vaginam dorso ad basin perforantes, brevis-
simz, 3-4-floree, rachide nonnullis pilis adpressis partim ramosis
nigris inspersa c. 1-2 cm. longa. Bractee oblonge acutiuscule,
concave, dorso et etiam intus ad apicem marginemque pilis 1-3-
ramulosis grossiusculis nigris vestite, c. 8-nervize nervis 3 median-
si usque ad apicem productis membranacee 1.175 cm. longz, 0.475
em. late. Flores carnosuli, c. 2.9 cm. lati, 2.8 cm. longi. Sepalum
dorsale cum ovario angulum obtusum ffaciens, levissime
sigmoideum porrectum, secundum costam mediam canaliculatum
subovatum, apice triangulo-contracto-acuminatum, satis acutum,
margine superne subirregulare basi latum 7-nervium, c. 1.6 cm.
longum, fere o.8 cm. latum. Sepala lateralia cum pede
gynostemii mentum conicum rectum cum ovario angulum acutum
faciens dorso longitudinaliter bicanaliculatum basi utrinque
gibbiforme parte superiore calcariformi-contractum cum margini-
bus anticis fere omnino liberis contiguisque obtusum I cm.
longum formantia divergentia angulum acutum facientia,
secundum costam leviter canaliculata obtusangule triangula,
acuta breviter apiculata, apice subirregulariter marginata, dorso
obtuse carinata, 7-nervia c. 1.75 cm. longa, margine antico
2.5 cm. basi 1.35 cm. metientia. Petala divergentia, apice
recurvula oblique ovata, 2/5 partibus superioribus triangulo-
contracto-acuminata acuta, basi utrinque rotundata medio tantum

Vol. IX. (1935).

Q2

affxa, 3-nervia, nervis exterioribus ramosis, textura sepalis
tenuiora, 1.75 cm. longa, 1.05 cm. lata. Labellum ungue parti
superiori pedis gynostemii calcariformi-connatum, 3-lobum,
concavum, lobis lateralibus gynostemium zquans, subtus inter
lobos laterales leviter longitudinaliter canaliculata, intus inter
lobos laterales incrassatione longitudinali leviter 3-costulata in
lobo intermedio in costas 3 valde verruculosas apicem labellifere
attingentes exeunte, 5-nervium, nervis lateralibus valde ramosis
nervis ramisque omnibus in lobo intermedio valde verruculosis
ceterum serie unica verrucularum instructis, inter nervos in lobo
intermedio humilius verruculosum, in basi laminz fascia longi-
tudinali glabra excepta papillosum, expansum totum ambitu,
rhombeum, fere 2.5 cm. longum, 1.5 cm. latum, ungue oblongo,
longitudine 0.6 cm. pedi gynostemii adnatum; lobi laterales erecti,
rotundato-trapezoidei, antice vix producti margine antico leviter
irregulares, mesochylium zquilateraliter trapeziiforme c. I.I cm.
longum formantes; lobus intermedius cum lateralibus angulos
acutos faciens, porrectus, recurvus, undulatus, irregulariter mar-
ginatus, expansus quinquangularis, apice angustatus, obtusius-
culus, 0.9 cm. longues, 0.75 cm., basi 0.575 cm. latus. Gynoste-
mium rectum subconicum dorso convexum, subtus planum, cum
anthera 0.525 cm. longum, filamento longo, filiformi-subulato, auri-
culas paululum superante, auriculis triangulis, acutiusculis margine
postico leviter serrulatis concavis. Anthera cucullata antice visa
quinquangularis, dorso obtusa retusaque, apice in appendicem
transversam subtruncatam vix retusam dense hirtellam producta,
omnino minute cellulosa, 0.25 cm. lata. Pollinia 4 in corpuscula
2 unita lateraliter compressa, oblique oblonga vix curvula, 0.175
cm. longa. Stigma basin gynostemii attingens, oblongo-ovale,
vix urceolatum, basi rotundatum pice truncatum. Pes
gynostemii cum ovario angulum acutum faciens, rectus, canali-
culatus, fere dimidio superiore labelli ungui adnatus et angustior,
parte lata dorso comspicue bicanaliculatus, 1 cm. longus.
Ovarium oblique obconicum, 6-suleatum, cum pedicello angulum
obtusum faciens, 0.65 cm. longum; pedicellus teres c. 2.5 cm.
longus.

Borneo :—Wester Afdeeling, Singawang, alang-field with
scattered trees, epiphytical (F. J. Paath, 20 October, 1934;
‘flowers creamy white, midlobe of the lip inside bright orange’’).

A distinct species of the group Nigrohirsuto of the section
Distichophyllum. The lip is very warty inside, as in D. ovi-
postoriferum J. J. S., D. Hallieri J. J. S. etc., but it is very
different from any of the species known to me in the rather short
mentum, in the side lobes, which are very large as compared with
the midlobe, and in other characters. In the shape of the ovate
acute petals it somewhat resembles D. Hallievi.

By the good offices of the editor of ‘‘De Orchidee’’ I received
flowers preserved in spirit and phctographs from which the
description has been made.

Gardens Bulletin, S.S.

* 93

THE EPIPHYTIC FLORA OF DIPTEROCARPUS
OBLONGIFOLIUS, BI. (‘‘Neram’’).

By M. R. HENDERSON,

Botanic Gardens, Singapore.

Dipterocarpus oblongifolius, Bl. is a common tree on the
banks of rivers and streams, especially east of the Main Range
of the Malay Peninsula, and is abundant on some of the
tributaries of the Pahang river. It appears to be confined to
riverbanks, and is a massive tree, though not usually tall as
compared with other Dipterocarps.

Its ability to grow on steep banks and its resistance to floods
are remarkable. ‘This is probably due in part to its root system.
Judging from specimens seen which had their roots exposed by
floods, the roots do not penetrate deeply into the soil but spread
over a large area and are so tenacious that the whole top of a
tree may be wrenched off by pressure of floodwater and drift-
wood, leaving the roots and lower part of the trunk undisturbed.

Where the trees grow on steep banks, the base of the trunk
is almost at right angles to the bank. The more nearly
perpendicular the bank the more nearly horizontal the base of
the trunk, which then curves gradually upwards but seldom
attains a perpendicular position. The same method of growth
may be observed in such plants as Pandanus growing on very
steep limestone cliffs. The upper surface of the basal portion of
the trunk of the neram is extended into a flange, so that a cross
section of the trunk at this point would be roughly a triangle with
a rounded base, or somewhat pearshaped. This extension of the
trunk no doubt acts as a stay, and the tree is actually not
buttressed in any way, as there is no buttress development on the
lower surface of the trunk, and less development of roots in the
direction of the water than in other directions.

In some localities the neram trees carry a large and varied
epiphytic flora, which is more easily examined than that on tall
forest trees, as the former are usually not difficult to climb and
can be examined from the streambed with comparative ease. A
census of this epiphytic flora was attempted during an expedition
to one of the small tributaries of the Tembeling river (Sungai Sat)
in July, 1929 and to Sungai Ketil, Kelantan in the following
month. It is noticeable that on the banks of the larger rivers,
such as the Tembeling in Pahang and the Lebir in Kelantan,
where the original vegetation has been cleared, the neram trees,
where they have been allowed to remain, carry few epiphytes,
but on the smaller rivers, where the original vegetation has not
been destroyed, and where the trees meet overhead, epiphytes
are much more numerous. The luxuriance of the epiphytic flora,

Vol. IX. (1935).

04 :

however, varies in a curious way. For instance, the trees along
the Sungai Terang, a small tributary of the Lebir, carried few
epiphytes compared with those on the Sungai Sat, although one
would have said that the conditions were more favourable on the
former river. The same applies to the epiphytic flora of Saraca.
In some places these trees are loaded with small orchids, while
in other apparently similar localities they bear none.

It was found that the orchids were the most numerous and
for this reason they were given more attention than other groups.
For the identification of these I am indebted to Mr. C. E. Carr,
whose detailed knowledge of the local orchid flora enabled him
to identify most of the specimens although they were not
flowering.

The method adopted was to send one or two men up the tree
with instructions to strip off and throw down every epiphyte they
could see. Branches were cut off and examined for such small
plants as Taeniophyllum, and field glasses were used to ensure
that nothing was missed. As far as possible identifications were
made on the spot, and anything which could not be dealt with
in this manner was preserved for future study. Probably about
I0o—20 per cent. of the total area of each tree was examined, the
search being naturally restricted to the trunk and lower, stouter
branches. Without felling the tree the canopy could not be
examined. Many Taeniophylla are often confined to one or two
branches and do not spread over the whole tree, and some species
such as Thrixspermum grow amongst the leaves towards the
tips of the branches in more or less full sunlight. However, the
proportion of the epiphytic flora enumerated here is considerably
more than 20 per cent. of the total, as the bulk of the epiphytes
grow on the trunk and lower branches.

It was found impossible to make a count of individual plants.
Some orchids and such ferns as Drynaria quercifolia were
exceedingly abundant on some trees.

The following are the flowering plants (excluding orchids}
and ferns :—

Vitis macrostachya, Miq.; Vitis sp.; Pachycentria
tuberculata, Korth.; Medinilla Hasseltii, Bl.; Medinilla sp-.;
Schefflera Ridleyi, Vig.; Myrmecodia tuberosa, Jack; Hydno-
phytum formicarium, Jack; Vaccinium acuminatissimum,
Mig.; Rhododendron Teysmanni, Miq.; Hoya latifolia, Don;
Hoya sp.; Fagraea auriculata, Jack; Aeschynanthus radicans,
Jack; Helixanthera coccinea, Dans.; Ficus consociata, B1.;
Ficus diversifolia, Bl. var. Kunstleri; Ficus diversifolia, Bl.
var. ovoidea; Ficus parietalis, Bl.; Pothos sp.; ?Anadendrum
sp.; Asplenium pellucidum, Lam.; Asplenium spathulinum,
J. Sm.; Cyclophorus varius, Gaud.; Drynaria quercifolia, L.;
Humata angustata, J. Sm.; Humata heterophylla, Desv.;
Platycerium Ridleyi, Christ.; Polypodium lomarioides, Kze;

Gardens Bulletin, SS.

95

Polypodium mettenianum, Ces.; Polypodium phymatodes,
L.; Polypodium revolutum, C. Chr.; Polypodium sinuosum,
Wall.; Trichomanes minutum, Bl.; Trichomanes pallidum,
Bl.; Vittaria elongata, Sw.; Vittaria ?ensiformis, Sw.;
Lycopodium laxum, Spr.; Lycopodium nummularifolium,
Lycopodium phlegmaria, L.; Lycopodium phyllanthum,
= & A...

Six trees were examined particularly for orchids and the
results are given in tabulated form below. ‘The first three trees
were in a group together, numbers four and five were together
some distance upstream, and number six was on another small
stream about a mile distant from the others.

)
— @ 2)3)/4/5]6
, " D. oie
Oberonia anceps, Lindl. ~ Ps
Hippeophyllum Scortechinii, ‘Schitr. ... x
Platyclinis longifolia, Hemsl. ve x x
Dendrobium acerosum, Lindl. ae x x
on atropurpureum, Mig. ote x
A: bifarium, Lindl. fe x
i eulophotum Lindl. ie x | x
* Farmeri, Paxt. Mag. eh Bina) x
af fugax, Schltr. iS ee x
" grande, Hook. f. aun
A- Hosei, Ridl. ... r18 Bbc
indragiriense, Schltr. ... x
m leonis, Rchb. ... = x x
Ms lonchophyllum, Hook. f. | x
Af pensile, Ridl. ... Lee x te one Se .
oi pumilum, Roxb. PF x > ie ie
* serra, Lindl. ... Sl | x x
teres, Lindl. ... iz x
Bulbophyllum odoratum, Lindl. oe | =.
- lepidum, i ie die x x x x
2 medusae, Rchb. f. zt Ye om :
be purpurascens, Teysm, ... x x
oF restrepia, Ridl. a x
vaginatum, Rchb. f. ... x x x
Eria floribunda, Lindl. x ¥ oe
» neglecta, Ridl. a ap | x x
En suaveolens, Ridl. bin Novi x x ee
,, Xanthocheila, Ridl. Fa x x
Trichotosia Teysmanni, Kranzl. 5a x *
velutina, Kranzl. ae x |
Tylostylis rigida, Bi. at / x |
Coelogyne Rochusseni, De Vr. ah x i On oe
ml testacea, Lindl. — x
Cymbidium pubescens, Lindl. ree x | > x
Grammatophyllum speciosum, Bl. acer | x
Saccolabium Hendersonii, Carr x
~ undulatum, Ridl.

Vol. IX. (1935).

96

yea | 1 | 2 | 4|5 | 6
Renanthera elongata, Lindl. tt x |
Sarcanthus Scortechinii, Hook. f. aa x, / x
Pomatocalpa Kunstleri, J.J.S. x /
Sarcochilus pusillus, Rchb. f. ¥
virescens, Ridl. x
Taeniophyllum obtusum, Bl. x x
Poaephyllum pauciflorum, Ridl. sie a x
Appendicula cornuta, Bl. ... ie. x 4
Podochilus tenuis, Lindl. ... ay D4
Agrostophyllum bicuspidatum, J.J.S. ... x Re Sed a ee ee
Hasseltit, J.J.s; x x | 24 eee
Oxyanthera decurva, Hook. f. sai x
Thrixspermum pardalis, Carr ee x |
Number of species on each tree ... 16 17 | 19 | 14 | 18 | 20

There were also two Eria spp. of the section Hymeneria,
and a Bulbophyllum of the section Aphanobulbon, unidentified.
Fifty known species and three unidentified species were therefore
found on these six trees. The two species occurring on all six
trees were Agrostophyllum bicuspidatum and A. Hasseltii. The
only species common to the three trees close together were
Agrostophyllum tnmathscas A. Hasseltit and Dendrobium
penstle.

The following orchids were found on other neram trees :—

Liparis lacerata, Ridl.

Dendrochilum sp.

Dendrobium aggregatum, Roxb.
crumenatum, Sw.
a Haniffii, Ridl.
iS luxurians,, ‘Carr

se salaccense, Lindl.

ae flavidulum, Hook. f.
Bulbophyllum abbrevilabium, Carr

A clandestinum, Lindl.

oe tortuosum, Lindl.

Eria pendula, Ridl.

49. Danes, ods:

», leiophylla, Lindl.

,, stellata, Lindl:

5 biflora, Gris:
Trichotosia vestita, Kranzl.
Chelistonele sp.

Gardens Bulletin, S.S.

97

Coelogyne Kingii, Hook. f.

re stenophylla, Ridl.
Cymbidium aloifolium, Sw.
Acriopsis javanica, Reinw.

feelin ve
Agrostophyllum majus, Hook. f.
Bromheadia alticola, Ridl.

a rupestris, Ridl.
Dilochia Wallichii, Lindl.
Thrixspermum arachnites, Rchb. f.

a rubrocallosum, Carr
Taeniophyllum pahangense, Carr
Sarcochilus biserratus, Rid].
Ceratostylis pendula, Hook. f.
Sarcanthus halophilus, Ridl.

The total number of species of orchids found on neram
within a limited area is thus eighty-seven.

As a contrast to the neram flora, two trees of Saraca bijuga
were examined for orchids and their flora was found to consist
mainly of the following :—

Dendrobium subflavidum, Ridl.
Bulbophyllum nigropurpureum, Carr
Chamaeanthus laciniatus, Carr
Saccolabium Hendersonii, Carr

mi undulatum, Ridl.
Sarcochilus caligaris, Ridl.

és eihiatus;, Jul S.

a incurvicalcar, J.J .S.

. pusillus, Rchb. f.

: a siamensis, Carr
Pomatocalpa Kunstleri, J.J.S.
Thrixspermum arachnites, Rchb. f.
Taeniophyllum obtusum, Bl.

Ys rugulosum, Carr
Appendicula pendula, Bl.

It will be noticed that none of the commonest orchids on
neram occur on Saraca, e.g. Dendrobium pensile, D. pumilum,
Bulbophyllum lepidum, Eria floribunda, E. suaveolens, Agrosto-
phyllum Hasseltii, A. bicuspidatum, are absent. The orchid:
flora of Saraca consists mainly of small plants such as
Saccolabium, Sarcochilus, Thrixspermum, Taeniophyllum, etc.,
and large plants, such as Grammatophyllum, Coelogyne,
Cymbidium, etc. are conspicuous by their absence. _

Vol. IX. (1935).
G

98

PHYLOGENY OF SINGLE FEATURES,

as illustrated by a remarkable new Sapotaceous tree
from British Malaya (Madhuca Ridleyi, n. sp.).

By H. J. Lam,

Professor of Botany, and Director National Herbarium, Leiden
(Holland ).

In older works concerning the field of phylogenetic taxonomy,
families and genera were considered as more or less static entities.
It was accepted that they wandered both through time and
through space, and also that, as they developed, or in general
changed by evolutional processes during that journey they
helped to form that much-discussed, but ever-mysterious struc-
ture, the genealogical tree. Such expressions as: the genus A
has probably given rise to the genus B, the genus C is to be
derived from the genus D, the genera EK and F have common
ancestors with the genus G, etc., are characteristic of those days.
Although dynamic in the sense of evolution, these taxonomic
units were, in fact, static entities, as they were considered to
travel and to change as units.

However, taxonomic ideas are not the same nowadays.
Genetics arose and developed and, being in some way the branch
of science, which covers the phylogeny of living species, greatly
influenced the ideas of taxonomical and phytogenetical investi-
gators. Instead of a static treelike structure as the old extreme,
we got, as the extreme opposite concept, a sort of dynamic net-
work, comparable with the creeping plasmodium of a slime-mould.
No fixed units any more, only a series of ever-changing
combinations of a constant number of genetical factors
(Havyata). Although few will agree with an extreme dynamic
system like his, the remarkable ideas of this author are certainly
both characteristic of, and, to some degree, also responsible for
the change of our thoughts. At any rate, the former families,
genera, etc., are considerably less solid, not only in a taxonomic,
but especially in a phytogenetic sense. Feeling that the old tree
is as unsatisfactory as the slime-mould plasmodium to serve as a
symbol for our ideas concerning phylogenetical processes, we are
nowadays trying to find some “‘golden mean’’. We do that,
for instance, according to an old and approved recipe, by analysis
of the material available; in other words, we are no longer examin-
ing the tree and the slime-mould as a whole, but we try to look
for what parts they are made of : we study single features. The
features, whether determined by a single genetical factor or by a
group of such factors, certainly have more right to be called the
true taxonomical-genetical units than the individuals. During
my investigations on Malaysian Burseraceze (5, 6) I became more
and more convinced of the value of this phylogeny of features

Gardens Bulletin, S.S.

99

(“Merkmalsphylogenie’’; ZIMMERMANN); next to this my atten-
tion was drawn to the idea that features could, alone or in
combination with others (coupling of factors), travel, fairly
independently, not only through time but also through space for
long periods (Paleogeography of features as e.g. practised by
LYMAN B. SmiTH, 7). It seems possible that a middle course
between the tree and the slime-mould may be found by accepting
that all combinations of factors (features) and all grades of
strength or duration of a coupling may occur. Evolution would
mean in that case: a certain, yet undiscovered, trend in the way
in which features cling together or in which certain combinations
succeed each other. Both hybridization (Lotsy) and mutations
(HUGO DE VrRiEs) find their place in this concept.

As in all phylogenetic studies, paleobotanic data are of
principal importance, but as these are too often wanting we have
to look for what taxonomy (morphology) and geography (distri-
bution) may teach (VON WETYSTEIN). A careful and critical
study of these data may give us an idea of the possible phylogeny
of what we call a taxonomic unit. We shall then see, I expect,
that generalisation is not possible in the way we were used to;
that every genus, etc., has to be investigated quite separately,
and that conclusions of a general nature may only be relative to
the fundamental laws of evolution. These considerations leave
the way open to a process of the origin of genera and families,
which I think has been taken into account only too little up to
now, viz. through polyphyletic evolution. On account of our
present knowledge I prefer a phylogenetic symbol in the form of
a network-structure, extremely minute and dynamic at every
point representing the present, and extremely coarse and static,
as viewed from these points, in the farthest past; we have to bear
in mind, however, that this structure is no reality but just a
human attempt to grasp a complicated problem.

At the time that I studied Malaysian Sapotaceze (1, 2, 3, 4),
these ideas had not yet evolved in my mind. However, having
taken up these studies? again with the aid of some collaborators,
I have come upon some facts which perfectly fit into the above
considerations, and I am glad to mention here an example of the
probable phylogeny of a single feature, illustrated by a new
species of Madhuca, which I am very much pleased to dedicate
to the nestor of Malayan and Malaysian; botanists, Mr. H. N.
RIDLEY, on the occasion of his 80th birthday. I have named it
Madhuca Ridleyi; a diagnosis and a picture of the species may
be found at the end of the present paper.

1 I hope to report on a striking example concerning this before
ong.

2 Publication of a revision may be expected next year.

3 Malaya=Malacca=the Malay Peninsula (British Malaya),
which in this paper is considered as forming phytogeographically a
part of Malaysia: Malaysia=the Malay Archipelago; the word has,
if I am not mistaken, been introduced by E. D, MERRILL.

Vol. IX. (1935).

LOO

As appears from may earlier papers (1, p. 11, 110-112),
Madhuca is one of four genera (the others are Payena, Ganua and
Burckella,), together forming the group of the Palaquiine-
Madhucee-Eumadhucee, well characterized by the possession of
two whorls of two sepals each.

Let me begin with some words as to what may be suggested
regarding the phylogeny of these four genera. As we know
nothing about fossil Madhucee, we have to look for what data
recent representatives have to yield. First of all we have to state,
that all of the four genera are very closely related, which is
illustrated by the great trouble it took to find a means of sub-
division, as all features are very irregularly distributed over the
genera. Apparently the Eumadhucee form a group which is in
a state of initial differentiation into four branches; a differentia-
tion, nevertheless, distinct enough to claim different names for
the branches.

Valuable data may now be expected in the present distribu-
tion of species and features.

Distribution of species. As appears from the accompanying
table (Pl. 3) and the map (PI. 4), the group is restricted to S.
Eastern Asia (Western limit: Deccan Peninsula and Ceylon) and
Polynesia (as far east as Fiji, Samoa and Tonga) with a very
distinct centre of differentiation in Western Malaysia (British
Malaya inclusive).

According to orthodox ideas of historical phytogeography,
our table (Pl. 3) confirms some well-known facts, obvious also in
many other plant families, e.g.

1. that the phytogeographic relations between British
Malaya and the greater Sunda Islands (especially
Sumatra and Borneo) are extraordinarily strong
(cf. 2, p. 384-385, table; and 3);

2. that next to a line of dispersion from Malaya to
Sumatra, a very important line is that from Malaya
to Borneo and the Philippines (Palawan !);

3. the hypothesis that this region which in a former
period is supposed to have formed one big land
mass that split up in postglacial times, has been an
important source and centre of dispersion of plant

| groups, the areas of which are now disjunct.

On account of these considerations I would suggest that the
Eumadhucee have originated in that former land mass, the so-
called Sunda-land. From this centre, the group has spread
geographically, during its morphological differentiation, both in
western direction (continent of Asia) and towards the East, and
the isolation of the various islands then gave rise to endemic
species (the Philippines—Madhuca-—and also New Guinea—
Burckella—have apparently acted as secondary regions of differ-
entiation), while other species remained unchanged everywhere
or perhaps evolved in the same way in different places.

Gardens Bulletin, S.S.

IOI

Of the three genera Madhuca, Payena and Ganua (cf. Pl. 3,
4 and 5), Madhuca covers the largest area and contains the
greatest number of species. The differentiation of these genera
mutually is both geologically and morphologically very young;
Ganua lias apparently its centre in Borneo (3 endemic species)
with offshoots to British Malaya, Sumatra and Palawan (non-
endemics) and, curiously enough, one isolated species in the
Moluccas;' Payena, with greater area than Ganua, and also with
stronger differentiation, is geographically intermediate between
Madhuca and Ganua. Morphologically Ganua is developing in
the direction of a peculiar venation of the leaves, of imperfect
septa in the ovary and of dry fruits. Payena is distinguished
by a well-characterized venation and a tendency to form a volu-
minous endosperm and Madhuca, with which Payena is connected
by many transitional species, shows, besides a type of venation
of its own, a tendency to reduce the endosperm. Burckella is
the only genus that is more distinct morphologically, but especi-
ally geographically. In contradistinction to the other genera it
does not follow the age-and-area theory, as the number of its
species is small and its area large. Yet I see no reason to accept
a reduction of the area and to consider the species as relic-
endemics, but a biphyletic origin seems to be possible. In its
venation it reminds one of eastern Ganua-species as well as of
a certain type of mostiy western Madhuca-species (Sect. Dasy-
aulus). It must certainly be remarked that its venation much
resembles the type of the two geographically nearest species :
Ganua Berlageana and Madhuca Burckiana. It shows, further-
more, a tendency to form very big fruits.

These general indications led me to the assumption that
Madhuca is the basal group in which the other genera are rooting.
Further evidence for this supposition may be found in some
features, intentionally left unmentioned thus far.

Distribution of features. The characteristics purposely not
yet mentioned are the number of parts in the corolla, the andrce-
ceum and the gyneceum. According to what is generally
accepted, high numbers may also in this group be considered as
representing the more primitive state. In the Eumadhucee
there are two tendencies to be noticed : one to fix the said numbers
at P. 8, A. 16, G. 8, and the second (which is, by the way, an
extremely general one) to reduce the G. number beyond 8. Now
we.see that Madhuca is the only genus in which all whorls show
a great variation viz. P. 18-6 (-5?), A. 4o-12, G. 21-6. In the
other genera P. and A. are, to a much higher degree, fixed to 8
and 16 respectively; G., however, shows a further reduction as
far down as 3. It is certainly a striking fact that the highest
numbers in the last-named whorl occur in or near the geographical

I aw may be a case of convergence (polyphyletic origin of the
genus).

Vol. IX. (1935).

I02

centre of the group and that the lowest ones occur in the most
remote parts of the area (Burckella in Polynesia and Madhuca,
although not exclusively, in India and Ceylon).

Our new species, Madhuca Ridleyi is, in this respect, to be
considered as a representative of a most primitive state, the
numbers of parts in its P. and G. whorls being by far the highest
known in the whole group, viz. P. 18, A. 37—36, G. 21-18!

The following facts may elucidate this somewhat more in
detail : —
Madhuca. P. 18-6 (-5?) A. 40-12 G. 21-6.

Between these extremes there is a very rich variation. Most
species show a variability in all whorls, and often a considerable
one, also those which cannot be said to be polymorphous in other
respects. Some examples are:

P.. A. G.
M. lancifolia ... I4-I0 28-22 I4—I0
Kingiana <1? OKs 36-32 10-8
betis a 10-8 20-16 8
sericea Pn? ees) 24-18 10-8
Endertii Pe 6) I3-I2 7-6
macrophylla sag, EO 28-20 18-II

etc.

M. Ridleyi has the highest numbers for G. As to P. it is
only equalled by M. macrophylla, as to A. it is only surpassed by
M. Lecomteana (nom. nov. for Bassia Thoreliana Pierre), in which
P.= 40-35. Only 3 other species, all from the Malay Peninsula
show similar P. numbers, except a Burckella species, discussed
below.

Payena. P. 8-6 A. 20-12 G. 8-6.

These extremes are exceptions. About 12 species out of 16
of which all whorls are completely known, show the 8—16-8
arrangement, many others show little variations and only a few,
some of which are polymorphous throughout, are more various,
viz. P. lucida (8-6, 16-13, 8-6), P. dasyphylla (8, 20-16, 8),
P. Griffithii (8-6, 16-12, 8) and P. truncata n. sp. (8-7, 16-4, 8?).

Ganua. P. 8-6 A. 24-16 G. (12?-) 9-6.

In this small genus the variability is somewhat ‘more pro-
nounced than in Payena. Yet, most of the species are fixed to
the 8—16—8 type or almost, and, as in Payena, the most striking
variability occurs in species that are also polymorphous in other
respects, such as G. Motleyana (10-8, 20-16, 8-6) and
G. monticola (8, 24-16, 12 ?—9-8).

Burckella. P. 8-6 A. {40?-) 18-9 G. 6-3.

Gardens Bulletin, S.S.

103

There is hardly any variability in the corolla; only one species
is said to possess 6 petals. In the andrceceum three species have
18-16 stamens, one has 16-9 and one (B. Thurstonii [HEMSL. |
H. J. Lam, from Fiji) is said to have as many as 40 (p. 8, G. 4).

This fact gives me an opportunity to say a few words on the
phylogenetic value of the three whorls mentioned. As may
appear also from the following statement, the corolla and the
androeceum seem, in general, not to yield valuable and trustworthy
data for phylogenetic conclusions. ‘There is only a slight indi-
cation as to a reduction of the P and A. numbers in three Ceylon
species of Madhuca (6—-12-6). For the rest, it may even be that
these whorls are not always subject to a reduction but may, on
the contrary, in some cases, attain higher numbers by progression.
This not uncommon phenomenon is e.g. illustrated by the figure
of the andrceceum of Burckella (Bassia) Thurstonii in HOOKER’S
Icones Plantarum t. 2569 (X XVI, 1899), in which a petaloid
stamen is figured and also by the fact that stamens with halfway
split anthers are not rare in the group of the Palaquiine. This
made me refrain from laying any particular stress upon the
phylogenetic value of the characters of these two whorls.

The gyneceum, however, being an organ of much less
phylogenetic plasticity as a rule, is more reliable in this respect,
and shows a distinct reduction in the more remote parts of the
area. It does so in all of the four genera; in Payena this reduction
is hardly conspicuous (down to 6 in 3 out of the 19 species in
which this feature is known); in Ganua, being also in other
features more isolated (ovary, fruit, venation) it become more
apparent (down to 6 in 3, to 7 in another of 9 species); in Madhuca
the genus with the highest G. numbers this reduction is occa-
sionally found (viz. down to 6 in 2, 7 in 1 W. Malaysian, down
to 5 in 1, to 6 in 7 and to 7 in 1 Continental species). It is also a
striking fact that in M. mindanensis the Philippine specimens
possess 8, the North Borneo ones (living in a place which is pro-
bably colonized from the main area in the Philippines) 6 cells in
their ovaries. None of these genera, however, have a G. number
below 6. This is only reached in Burckella where there are 6,
5, 4 or 3 cells, however, wihout more detailed geographic
aia with the accepted general migration from West to

ast.

Poin X (1935).

104

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Gardens Bulletin, S.S.

105

It appears, I think, especially from the percentage data, that
there is a tendency to be traced to reduce the number of carpels
as the distance from the centre of dispersion is greater, both to
the West (India, Ceylon) and the East (Polynesia). That this
reduction is more distinct in the Hast may readily be ascribed
to the preponderant insular character of those parts. In the
other whorls, there is no correlation, possibly partly due to an
eventual tendency to raise the number of petals and stamens e:g.
by what has sometimes been called a_ reduplication
(‘“dédoublement’’).

The above facts have been combined with my general ideas
on the dispersion of the whole group in a tentative ‘‘genealogical.
branch’’ (Pl. 6) which may speak for itself, but concerning
which I must state that, of course, complicated processes as the:
phylogeny of a group of plants can never be symbolized in a
thoroughly satisfactory way on a sheet of paper; furthermore I
have to emphasize that the ‘‘branches’’ are not meant to be:
““solid’’, but should be imagined as sponge-like tissues of.
extremely complicated structure. For the rest, I am quite aware
of the very problematic value of such considerations and ideas.

A description and analysis of the new species may follow

here¥

Madhuca Ridleyi (Sect. Dasyaulus), nova species, Pl. 7—
Arbor mediocris, ramuli crassi; internodiis brevibus bulboso-
inflatis, 1.5-2 cm. longis et latis rugosis; folia ad eorum apices
dense conferta; stipule deciduze 0.3-1.2 cm. longz pubescentes;
folia coriacea glabra, obovata vel oblongo-obovata, 14-25 cm.
longa, 5—10.5 cm. lata, basi in petiolum glabrum 1.5-3.5 cm.
longum, 0.2—0.3 cm. crassum attenuata, apice acuta vel subrotun-
data; costa media 1.s. supra canaliculata, subtus valde prominens;
nervi secundarii subtus prominentes, 18-25, angulo 60°—70 de
costa adscendentes, recti, prope marginem curvati, diminuti, haud
conjuncti, 0.7-1 cm. distantes; nervi tertiarii transversi, angulo
circ. 120° de costa descendentes, subtus conspicui, reticulatione
perminuta, interdum nervo brevi secondariis parallelo a costa
media usque ad %4—-% adscendente; inflorescentie ex foliorum
delapsorum axillis orte, nonnulle ad ramulorum longorum
rectorum apices dense conferte, 15—25-—floree, pedicelli minute
tomentosi, graciles, 2-2.5 cm. longi; alabastra acute ovoidea;
calyx 1I-1.3 cm. longus, sepala extus dense minute fulvo-
ferrugineo-tomentosa, intus glabra, angula apicale circ. 90°,
Eeeriora 1.25-cm. longa, x cm. lata, imteriora 1.31.1. cm.;
corolla in alabastris vix exserta (flores novellos tantum vidi) 1.1—
1.2 cm. longa, extus glabra vel subglabra, intus glabra; petala 18
uniserialia dextrorsum tegentia o.95-1 cm. longa, 0.4 cm. lata,
margine plus minusve irregulariter minute denticulata; stamina
glabra 36-37, plerumque uniserialia, interdum nonnulli serie
imperfecta addita, 0.5-0.6 cm. longa, nova filamenta usque ad
0.2 cm. longa, antherz 0.2-0.35 cm. longe, 0.15 cm. latze, apice

Pore? xX... (1935).

106

acumine acuto 0.075—0.15—-0.25 cm. longo ornate; ovarium gla-
brum depressi-globosum extus paulo sulcatum, 18—21- loculatum;
loculi 1-ovulati; stylus solidus in alabastro 0.9 cm. longo; infrutes-
centie ramulorum apicibus subumbellatim conferte, pedicelli
2.5 cm. longi, 0.25-0.3 cm. crassi, apice vix dilatati; calyx
persistens, sepala 1-1.8 cm. longa, extus tomentosa; fructus
globosi, apice szepe stylo 3 cm. longo persistente ornatus, dense
ferrugineo-pubescentes, pericarpio crasso subduro; 4-6 semina
oblonga, testa tenuis, extus nitida; cetera ignota.

Malay Peninsula: Pahang, Bukit Serdam, Raub, 1300 ft.,
on dry limestone rocks (HENDERSON 25055, flowers, on 6 Oct.
1931, type specimen)—Upper Perak, Gunong Runto, Lenggong,
500 ft., on limestone (HENDERSON 23834, fruits on 11 June, 1930).

A small but rather stout tree, both specimens known growing
on limestone; bark pale, horizontally and longitudinally fissured;
fruits rusty brown.

I have to thank Mr. K. GRIFFIOEN for his assistance in pre-
paratory work for this paper, and my colleague, Dr. L. G. M.
BAAS BECKING for kindly looking over the English.

Summary.

After some theoretical considerations on modern phylogene-
tical Taxonomy as a discipline, dealing with morphological as
well as geographical evolution and newly influenced by Genetics,
it is stated that next to Paleobotany two categories of data may
be considered important for our ideas on Phylogeny, viz. the
present distribution of species and that of features. This is
illustrated by examples taken from the Eumadhucee, a group of
Hast-Asiatic to Polynesian Sapotaceee. It is shown that the
former Sunda-land is the most probable place of origin of this

group and that the species have migrated both westward (India, |

Ceylon, Indochina) and eastward (Polynesia). Evidence for this
suggestion is found in the distribution and evolution of certain
features, especially in the reduction of the number of carpels in
both directions mentioned. The paper was stimulated by the dis-
covery of a new Madhuca-species, M. Ridleyi, from British
Malaya, the gynzeceum of which shows the highest number of
carpels known in the whole group. A description of this species
is added.

Literature.
1. Lam, H.J., The Sapotaceae, etc. of the Dutch East Indies, etc.—
Bull. Jard. bot. Buitenz. 3e Sér. 7, 1-289,: 1925.

2. ——————-, Further studies on Malayan Sapotaceae, I—Bull. Jard.
bot. Buitenz. 3e Sér. 8, .381-493, 10927.

3. —————» Some remarks on the genetic phytogeography of the
Malay Archipelago—Ann. Jard. bot. Buitenz. 37,
33-48, 1927.

4. —————-» Enumeration of the Sapotaceae thus far known from
New Guinea—Nova Guinea 14, 1, 349-570, 1932.

Gardens Bulletin, S.S.

107

5. Lam, H.J., The Burseraceae of the Malay Archipelago and
Peninsula—Bull. Jard. bot. Buitenz. 3e Sér. 12,
281-561, 1932.

5. ———_—_——., Beitrage zur Morphologie der Burseraceae II—Ann.
Jard. bot. Buitenz. 42, 35-164, 1932.

7, Smith, Lyman B., Geographical evidence on the lines of evolution
in the Bromeliaceae—Engl. Bot. Jahrb. 66, 4,
446-468, 10934.

Explanation of Plates 3-7.
Plate 3 |
Present Distribution of the Eumadhucee.

Well known species only; numbers between brackets under
genera indicate total number of species, in the other columns they
indicate the number of endemic species in the region concerned;
however, it must be borne in mind that amongst these are a
number of new and inore recent species.

Plate 4

Areas of Madhuca (M), Payena (P), Ganua (G) and Burckella
(B) and total number of species (Eumadhucee) in the separate
regions.
Plate 5

Probable lines of migration.

Plate 6
Probable phylogeny of Eumadhucee.

Plate 7

MADHUCA RIDLEYI, n. sp.—a. branch with leaves; b.
shoot with inflorescences; c. id. with fruits; (d) flowerbuds; e.
outer sepal, in- and outside; f. id. inner sepal; g. corolla inside;
h. petals; k. stamens; m. pistillum; n. ovary, cross-section—after
HENDERSON 25055, except c. which is taken from HENDERSON

23834.

alo tx: (1035).

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Gardens Bulletin, S.S.

Plate 3

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Gardens Bulletin, S.S.

Plate 5

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Plate 7 Gardens Bulletin, S.S.

II3

CHONEMORPHA MACROPHYLLA AND ALLIED
SPECIES.

C. X. FurTApbo,
Botanic Gardens, Singapore.

Introduction.

Although Chonemorpha macrophylla G. Don is the type of
the genus as currently interpreted, it is a species whose limits
are not yet clearly defined, with the result that more than -one
species are included under that specific name, and this holds
good even after the exclusion of the Philippine and Javanese
material that was previously referred here. It is true that
Ridley, acting on Stapf’s suggestions on this matter, had (in
Agric. Bull. Straits & Fed. Maiay States X, 1911, pp. 146-148)
created two new specific epithets, C. penangensis and C. Rheedet,
hoping to straighten out the matter. But, as he did not bring
out fully the chief characters that distinguish the three species,
his paper did not succeed in making matters much clearer, and
moreover ‘his paper with Stapf’s suggestions being published in a
periodical of purely agricultural and local interest, failed to
attract the attention of horticulturists and systematists even to
the fact that C. macrophylla as then interpreted was being
regarded as a mixed species. It is not surprising therefore that
the two Supplements to Index Kewensis for t1g1t1I-15 and
1916-20, published respectively in 1921 and 1926, were issued
without including Ridley’s two new species, their inclusion not
appearing until 1929 when the Supplement for 1921-25 was
issued and when presumably the compilers’ attention was drawn
to Ridley’s paper through a reference made to it in Ridley’s
Flora IT, 1923.

In the hope, therefore, that the circumscription of C. macro-
bhylla G. Don may become clear, and that the separation of the
species usually confused with it may be easier, I have prepared
the Key and the Enumeration given below. This paper not
being a complete revision of the genus, I limit myself to giving
only those synonyms and references which I consider may be
useful to interpret the species correctly. I have also described
a new species C. assamensis, based on a specimen from Assam.

The species are so variable as regards the hairiness and size
of the leaves that it is not easy to distinguish them on that
character. The general outline of the leaves, the comparative
size of the corolla and the hairiness, size and shape of the calyx
are more stable characters and so advantage is taken of them in
framing the key. The hairiness of the pistil is apparently a
useful character in separating these species, and may form an
important basis in subdividing such polymorphous species as

Vol. IX. (1935).

II4

C. macrophylla and C. mollis; but the material at my disposal
was so meagre in flowers that I did not feel myself justified to
detach flowers from the authentic specimens for the purpose of
dissection.

From the enumeration given below it will be evident that
the success of my inquiry into the various species commonly
referred to C. macrophylla is due principally to the rich material
preserved in the herbarium of the Royal Botanic Gardens,
Calcutta. My thanks are therefore due to the Director of that
institution for the loan of the specimens.

Key to the Species.

A Leaves of the flowering shoots elliptic to obovate, usually more
narrowed in the lower half than in the upper; base rounded,
obtuse or slightly cordate. Calyx glabrescent or distinctly
hairy, urceolulate or companulate or crateriform; lobes
triangular.

a: Calyx distinctly hairy or velvety, about 10-12 mm. long,
slightly contracted at apex; lobes triangular, about 4
the total length of the calyx. Corolla tube 3-3.5 cm.
long; limb 5-6.5 cm. across.

1. C. penangensts.
aa: Calyx glabrous or glabrescent, not contracted at apex;

b: Calyx glabrescent, 9-12 mm. long, campanulate ;
lobes about 4} the length of the calyx.
Corolla tube 3-4.5 cm. long, contraction
towards the base begins at the mouth of the
calyx; limb 7-9 cm. across.

2. C. macrothylla.

bb: Calyx 4-5 mm. long, crateriform, glabrous ;
lobes about 4 the length of the calyx.
Corolla tube 3-4.5 cm. long, contraction
towards the base begins much above the
mouth of the calyx; limb 7-8 cm. across.

3. C. assamensis.

AA Leaves of the flowering shoots ovate, oval elliptic-oblong, or
almost orbicular, usually more narrowed in the upper half
than in the lower; base rounded, cordate. Calyx glabrescent,
campanulate; lobes triangular or ovate.

a: Calyx 10-13 mm. long; lobes } the length of calyx,
triangular. Corolla tube 2-2.5 cm. long; limb 6-7
cm. across.

4. CP piollis,

aa: Calyx 7-10 mm. long; lobes ovate, graduaily and
arcuately narrowed towards the apex, overlapping at
basé, 3-3 the length of calyx. Corolla tube 2-2.5
cm. long ; limb. 7—o cm? across:

5. C. fragrans.

Gardens Bulletin, S.S.

715

Enumeration.
1. C. assamensis Furtado spec. nov.

A C. macrophylla G. Don, cui affinis, calycibus glaberrimis
brevioribus, cupuliformibus; corollarum tubo supra (haud ad)
calycum orificium dilatato; heec species facile distinguenda.

A C.. fragrante Alston calycum lobis_trianglaribus,
ramulorum foliis floriferorum obovatis distat.

INDIA: Assam, loc. incert (collected by a native collector
under the supervision of G. Mann, in May 1893. Holotype in
the Calcutta Herbarium).

2. Chonemorpha fragrans (Moon) Alston in Ann. Roy. Bot.
Gard. Perad. XI (1929) 203 excl. syn., et in Trimen FI.
Ceyl. VI (1931) 192 p.p.

Echites fragrans Moon Cat. (1821) 20. ‘Type.

C. Rheedei Ridl. in Agric. Bull. S. & F.M.S. X (r1o11z) 146.
Syn. nov.

C. macrophylla G. Don sensu Thwaites, Enum. (1864) 194;
Hook. f., Fl. Brit. Ind. III (1882) 661 p.p. sake, F). Ceyl.
III (1895) 138 quoad specimina zeylanica; Cooke, Fl. Pres.
Bombay II (1904) 139 quoad descriptionem et specimina citata;
Gamble, Fl. Pres. Madras V (1923) 818 quoad specimina Indize
australis. Syn. nov.

Belutta-Kaka-Kodi Rheede, Hort. Malab. IX (1680) p. 7, tt.
5 & 6. (Type of E. fragrans Moon).

CEYLON : loc. incert. (Thwaites, C.P. 2467 in Calcutta herb).

CULTIVATED SPECIMENS: Singapore: Botanic Gardens
(Ridley, in 1911, holotype of C. Rheedet).

3. Chonemorpha macrophylla [Roxb.] G. Don, Gen. Syst.
Wien. Ft. iV (14636) 76; DC., Prodr. VIII (1344) 430, p.p.;
Miq., Fl. Ind. Bot. II (1856 444 p.p; Thwaites, Enum.
(1864) 194 p.p.; Hook., f., Fl. Brit. Ind. III (1882) 661 p.p.;
Tom,., Fl. 'Ceéeyl.’ LIT ‘(1895)’ 138 ip.p. ‘Hook.f. _Bot. Mag.
(1896) t. 7492. Gamble in King, Mat. Fl. Mal. Pen. IV
(1907) 693 p.p.; Ridley in Journ. Roy. Asiat. Soc. Str. Br.
BOL EOLO) OS...
C. macrophylla var. grandis A.DC., Prodr. VIII (1844) 450.

Syn. nov.

C. penangensis Ridl. in Agric. Bull. S. & F.M.S. X (1911)

147 quoad specimina perakensia, et Fl. Mal. Pen. II (1923) 360

my. SVR. 2Ov:.

Echites macrophylla Roxb. sp. nov., Hort. Beng. (1814) 20,
goa. mud., et Fl. Ind. Il (1824) 13 excl. syn. Type. [non

E. macrophylla H.B.K., Nov. Gen. et. Spec. III (1819) 219].

E. grandis Wall. Cat 1658, nom. nud.
E. latifolia Wall. Cat. no. 1657 E, nom. nud.

Vel FX. (1035).

116

InpiA: N. W. Himalaya, Dehra Dun (Mackinnon in Aug.
1897). W. Duars, Buxa (Gamble, -7608). Prov. ~— Ieee
‘Teendaria (Gamble, 3223 B & C); loc. incert. (Falconer).
Assam and Sikkim: Sillet (Wall. Cat. 16574); Kom (Wall. Cat.
1658); Kumaon (leg.?, in 1871); Goyalpara (Wall. Cat. 16578);
Slik (Rebu, 873).

BuRMA Shan States, loc. incert. (Allen, 2); Lashio alt
2,000 14; (lace; 5533):

ANDAMANS: (Prain’s coll., 57; Parkinson, 619).

MarAy PENINSULA: Perak, Temengo (Ridley, 14271); Larut,
alt. 300-500 ft. (Kunstler, 3636); Gopeng, alt. 300-500 ft.
(Kunstler, 6,000).

CULTIVATED SPECIMENS: INDIA, Botanic Gardens, Calcutta
(Wallich, 16578; Lane, 11,270). MALAY PENINSULA: Singapore,
Botanic Gardens (Furtado, Nov. 15, 1935). Selangor, Kuala
Lumpur in Public Gardens (Nov. 27, 1932).

With the exception of those from the Malay Peninsula, all
specimens cited above are from the Calcutta Herbarium. The
species is variable as regards the breadth and glabrescence of the ©
calyx.

4, Chonemorpha mollis Mig., Fl. Ind. Bot. II (1856) 444.

Stat. Mov,

C. macrophylla G. Don sensu Vidal, Cat. Pl. Prov. Manila
(1880) 36; Hook. f., Fl. Brit. Ind. III (1882). 661 p.p; Koorders,
Exkursionsfl. Jav. III (1912) 77; Heyne, Nutt. Plant. Nederl.
Ind. IV (z917) 85, et and Ed. IT (7927) 120%. Son.cwog:

C. Blancoi Merr., Spec. Blancoanze (1918) 312, et Enum.
Philipp, Pl. Ill (1923) 335. Syni-see-

C. elliptica Merr. & Rolfe in Philpp. Journ. Sci. III (1908)
E21, SVR. 2OVn

Tabernemontana elliptica Blanco sp. nov. Fl. Filip. (1837)
115 (non T. elliptica Thunb., Fl. Jap. 1784, p. 111). Syn mow:

Java. (Horsfield, duplicate of the Type, in Calcutta herb.).

CELEBES: Bonto Parang (Biinnemeijer, 10572).

PHILIPPINES : Luzon, Borhoso (Ahern, 1141; Ramos, 1065).
Palawan, Taytay (Merrill, 92409).

This species, like C. macrophylla, is very variable as regards
the pubescence and the size of leaves and calyces. A further
inquiry into the hairiness of the pistil and the corolla tube may
be desirable to understand the exact significance of the several
variations.

5. Chonemerpha penangensis Ridley in Agric. Bull. S. & F.M.S.
X (1911) 147 excl. specimina perakensia, et Fl. Malay Pen.

Il (1023) 360.p.p.

Gardens Bulletin, S.S.

117

C. macrophylla G. Don sensu Hook. f., Fl. Brit. Ind. II]
(1882) 661 quoad specimina malaccensia; Gamble in King,
Materials IV (1907) 693 p.p.

MaLAy PENINSULA: Penang, Balek Pulau (Ridley, 9441);
Wang Seraya, alt. 500 feet (Curtis, 832); Ayer Etam (Haniff,
in April 1916, vern. name: Gérip Merah), Malacca, Brisu (Derry,
543, vern. name: Akar Gérip-Gérip Merah.; loc. incert (Maingay
1074=Kew n. 1837, in Calcutta herb.). Negri Sembilan,
Tampin Hill (Burkill, 556). Selangor, Kepong (Symington,
20148; Pawanche, 14673, vern. name Akar Gérit Tembaga.).

Vol. IX. (1935).

22525—400—12/35

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"STRAITS "SETTLEMENTS

4th May, 1937 | Part 2

CONTENTS

7~_—_-_o_oo

N eh on cc Makyen Fen with Descriptions of 5 new
- Species, by R. B. Holttum = E - 119

‘urt r notes on Stenochlaena, Lomariopsis and Terato-
| pion. by. R. E. Holtturn - 139

4 bp Ren Crolaay: - 145
: Malesicae VI, by C. X. eee os se 152
e Malesicne VIL, by C. X. Furtado : : 182

in Plant Gecganhy. by C.G.G. J. van Steenis - 187

a purchased ime ‘ie Botanic Gardens, Singapore
oe titan ae Price $3. 00 or 7s.

ee He: - SINGAPORE:

NA at tHe GOVERNMENT? PRINTING Orpice, SINGAPORE,
ee i “By V. CG, G, GaTRELt, BoS Sie cra PRINTER.

; :

rey “1987 | ws

SUX 8 108% <<

THE

GARDENS’ BULLETIN

Vol. IX 4th May, 1937 Part 2

NOTES ON MALAYAN FERNS, WITH DESCRIPTIONS
OF FIVE NEW SPECIES.

By R. E. Hovrrum.

At various times during the past few years I have collected
notes on ferns of the Malay Peninsula, and have gathered them
together here, with descriptions of five new species, Davallia
dimorpha, Diplazium insigne, D. latisquamatum, D. subintegrum,
and D. velutinum. The larger Diplaziums, inhabiting the
mountain valleys from about 2,000 ft. upwards, still require
much field study, and also careful comparison with the species
of neighbouring countries. There are certainly several further
additions to be made to the Peninsula species of this genus.

The notes here published are of various kinds. Most are
additions to the recorded fern flora of the Peninsula, chiefly
based on recent collections, but in some cases on redetermination
of old collections. Some are critical notes on nomenclature,
based either on my own study of type specimens, or copied from
other sources, which are duly acknowledged. ‘There are also
field notes on many species, based on my own observations.

Acrostichum speciosum Willd. Spec. Plant. 4 Aufl. 5: 117. 1810.
Troll, Flora N. S. 28: 301 — 327. 1933.

As pointed out by Dr. Troll, this species is quite distinct
from A. aureum, with which it has been united by most authors.
Watson, in Malayan Forest Records No. 6, pp. 152-157, gives
an account of the two forms of Acrostichum occurring in Malayan
mangrove; they are so distinct that local Malays have given them
different names. The larger A. aureum, which may reach 4
metres in height, is called Piai raya; the smaller A. speciosum
is called Piai lasa. Both species are abundant. A. speciosum
occurs especially ‘‘where there are plenty of prawn heaps on
which it can grow above the reach of the medium high tides.”’

Aplenium glaucophyllum v.A.v.R.; Bull. jard. bot. Buitenz. II
ser. VII: 6. 1912. A. polystichoides Bl. Mett. Asplenium
204. 1859 (nomen).

This species, described from Borneo (Nieuwenhuis no. 864),
is not uncommon in the lowlands of the Malay Peninsula. ‘The

TIQ

I20

specimens have hitherto been referred with doubt to A. nitidum
Sw., to which species Mettenius referred as a variety the specimen
of Korthals named A. polystichoides in Blume’s herbarium. The
species has been adequately described by van Alderwerelt van
Rosenburgh, but some additional notes may be of interest.
These are based on the material in the Singapore herbarium, and
on a plant cultivated in the Botanic Gardens, Singapore. The
cultivated plant occurred spontaneously on the base of an old
Asplenium nidus on an African Oil Palm (El@is guineensts),
and was subsequently removed to a hanging basket.

The characteristic features of the species are the very long
relatively narrow fronds (lamina commonly about 85 by 30 cm.),
and the large pinnules, which are not pinnate but are commonly
rather deeply incised once on the upper margin towards the base.
Occasionally a specimen has the pinnules quite entire, except
for the rather distant teeth (e.g. Burkill 17265 from Pekan,
Pahang), while on very large fronds the largest pinnules may
have several lobes, the lowest almost separated as a distinct
leaflet. An extreme example of a lobed form is a specimen of
Burkill’s from Pulau Tioman (s.n., June 1915), but a quite
comparable development is seen on the largest frond of the
cultivated plant. The smallest fronds on the cultivated plant
are quite small, but are fully fertile. The pinnules are almost
all rounded at the apex, whereas on larger fronds the pinnules
are gradually narrowed to an acute apex.

The stipes of young fronds are green, blackening with age;
on old fronds the rachises are all black. ‘The stipes and rachises
are dull, not polished, and when young are clothed with very
small narrow dark scales. The densely crowded scales on the
apex of the rhizome are thin, dull purplish-brown, to at least
2 cm. long and 1 mm. wide. The colour of dried fronds varies
from a rather pale olive to brown, and the colour of stipe and
rachis from greenish-grey to black. Specimens in the Singapore
herbarium are as follows. ‘The Perak specimens are all small,
and were labelled A. nitidum var. obtusatum; they match very
closely the smallest fronds of the cultivated plant above
mentioned.

SINGAPORE: Bukit Timah, Ridley s.n. 1906; do. 1907;
Woodlands, Matthew s.n.; Mt. Austin, Ridley s.n. 1906.
JoHORE: G. Pulai, Hullett s.n. Aug. 1880; Haji Senawi, Tem-
payan River, Ridley 10965; Tempayan River, Ridley 13286;
Gunong Lambak, Holttum 9375; Mengkibol Station, Gossett
8314. SELANGOR: Batu Caves Woods, Ridley 8144. Matacca:
Harvey s.n. PAHANG: Pulau Tioman, Ridley s.n. 1889, Burkill
s.o. June 1915; between Pekan and Ayer Tawar, Burkill 17265;
Kuala Lipis, Burkill 17078, Tahan River, Ridley s.n. 27-9-1893.

Gardens Bulletin, S.S.

£27

KELANTAN: Sungei Keteh, Md. Nur 12037. PERAK: Gopeng
district, King’s Collector 8180; Upper Perak, L. Wray Jr. 3657.

Asplenium subnormale Copel. Perk. Frag. p. 183, pl. 4, £.B. 1905.

This species has not hitherto been recorded from the Malay
Peninsula. It has been collected by Burkill on Pulau Tioman,
S. Cape, close to the sea, no. 1036. I have seen specimens from
Sarawak and British North Borneo.

Asplenium subscalare v.A.v.R. Bull. Jard. bot. Buitenz. II
ser XA: 6 Fors.

This species was described from a specimen originating from
“‘Malacca’’, cultivated in the Botanic Gardens, Buitenzorg. The
specimen was at first named A. spathulatum Bak. by v.A.v.R.,
whence the statement in his Handbook that A. spathulatum
occurs in the Peninsula. A specimen of Scortechini’s from
Taiping, no. 129, labelled in his hand A. abruptum (a nomen
nudum), is apparently the same as this species. Scortechini 243,
from Maxwell’s Hill, 3,000 ft., is perhaps also to be referred
here; the plants are young.

Athyrium macrocarpum (BI.) Bedd. F.S.I. t. 152. 1863.
Aspidium Bl. Enum. 152. 1828.

This species has not hitherto been recorded as occurring in
the Malay Peninsula. It has been found by me (no. 23462) on
wet rocks at Cameron Highlands, beside the stream, with
Lindsaya cultrata.

Athyrium nigripes (Bl.) Moore, Index XLIX. 1857. Aspidium
Bl. Enum. 162.

This species was collected by Ridley on G. Berumban, no.
14001, and the specimen was named Asplenium tenutfolium by
Beddome (Kew Bull. 1909: 423). I collected it later (no. 23275)
it Cameron Highlands, where it was abundant in the forest
near the stream at about 4,800 ft. altitude. This is the first
published record of its occurrence in the Peninsula.

Bolbitis diversifolia (Bl.) Schott, Gen. ad t. 14. 1834.

Bolbitis subsimplex (Fée) Ching in C. Chr. Ind. Suppl. HI:
50. 1934. with synonyms.

At the herbarium at Leiden I examined three specimens of
Blume’s, referred to this species, and I can see no difference
from the common fern in the Peninsula which is clearly Campium
subsimplex as described by Copeland (Phil. Journ. Sci. 37: 356.
1928). One of Blume’s specimens has simple fronds, the others
pinnate. The species is common on rocks, usually by streams,
in lowland forest throughout the Peninsula. Young plants
always have simple fronds, and occasionally simple fertile fronds
are produced.

Vol. IX. (1937).

122

Bolbitis Quoyana (Gaud.) Ching in C. Chr. Ind. Suppl. III:

49. 1934.

This species has not hitherto been reported as occurring in
the Malay Peninsula. The only specimens which can now be
so referred are from Bukit Timah, Singapore: Ridley 12560,
Holttum 19799, Hullett s.n. March 1882. ‘These specimens are
not typical of the species, having pinnze only very slightly lobed,
but I do not know any other species to which they could be
referred.

Cheilanthes varians (Wall.) Hk. Spec. Fil 2: 89, t. 103A. 1852.

This species occurs in the north of the Peninsula, but has
not hitherto been reported. Collections in the herbarium at
Singapore are: KEDAH, Kampong Naka, Holttum 19814; Lang-
kawi, Curtis s.n. I found the fern not uncommon, under the
shade of rather open secondary forest.

Cyclopeltis crenata (Fée) C. Chr. Ind. Suppl. III: 64. 1934.
Hemicardion Fée Gen. 283, t. 22A, f. I. 1852.

Specimens of Cyclopeltis from the Malay Peninsula have
previously been referred to C. semicordata, the type of which
is from tropical America. Fée’s description and figure agree
well with Peninsular plants which are quite distinct from the
Philippine C. Presliana.

Cystopteris tenuisecta (Bl.) Mett. Ann. Lugd. Bat. I: 241. 1864.

This interesting species has lately been collected at Cameron
Highlands (S.F.N. 3120907, Holttum). Plants were found
growing abundantly on the banks of a path cut in rather moist
but lightly shaded forest at 5,000 ft. altitude. The same locality
was visited before the path was cut, and no plants of the species
noticed; they evidently found the moist newly cut earth a suitable
habitat. No other plants were noticed in the neighbourhood,
where the species is certainly not common, and no previous
collections have been made in the Peninsula.

There are notes on C. tenuisecta by Posthumus in Bull.
Jard. bot. Buitenzorg, Ser. III, vol. 13, p. 91. In the Malayan
region it has been rarely collected, being known from N. Sumatra,
a few localities in Java, and Luzon. It evidently requires a
specialised habitat.

Davallia dimorpha Holttum sp. nov.

D. Griffithii affinis, differt: frondibus dimorphis,.in sicco
rufobrunneis, segmentis ultimis laminz fertilis perangustatis.

Scales of the rhizome red-brown, shining, to 1 cm. long and
3 mm. wide above the peltate base, narrowed gradually to
acuminate apex, edges subentire. Stipes of large fronds 30—40
cm. long, red-brown to dark purple-brown. Lamina of sterile
frond deltoid, to about 45 cm. long and 30 cm. wide, pinne

Gardens Bulletin, S.S.

i

123

subcontiguous, the segments of the pinnules contiguous or sub-
contiguous; pinnules separated by not more than half their width,
more or less elliptical in shape, the edges bluntly serrate, apices
rounded; texture coriaceous; colour of dried fronds rather light
red-brown (often darker on older fronds); veins distinct, elevated;
no spurious veins. Lamina of fertile frond to about 45 cm. long
and 30 cm. wide, the pinnez and pinnules of similar shape, size
and spacing to those of the sterile frond, but the lamina of the
pinnule-segments cut down to a narrow wing on either side of
the veins, each segment terminating in a semicircular widened
portion to accomodate the sorus; occasionally but not commonly
the lamina prolonged into a tooth or horn beyond the sorus on
one side; sori broader than long, often twice a broad, the
indusium almost crescent-shaped, occupying almost the whole
of the widened ends of the segments.

Type.—Pahang: Brinchang, Cameron Highlands, alt.
5,000 ft., S.F.N. 31289 (Holttum), in Herb. Singapore.

This remarkable species has only been found at Cameron
Highlands, on the Main Range of the Peninsula. It is evidently
allied to D. Griffithii, with which it agrees in the very broad sori,
but differs very strikingly in the much-narrowed fertile pinnule-
segments, which carry at their ends the almost crescent-shaped
sori. The rather bright red-brown colour of dried fronds which
are not too old when collected is also notable.

Diplazium insigne Holttum sp. nov.

Caudex brevis erectus. Stipites 1 m. longi, basin versus
aculeati; aculei tenues, ad 2 mm. longi, primo paleas sordides
angustas, ad 1.5 cm. longas, margine nigro-dentatas, ferentes,
demum nudi. Lamia 1.5 m. longa, bipinnata. Raches
minute paleacece; palee irregulares, majores dentate. Pinneze
infimee 28 cm. longe, 6.5 cm. late, basi angustateze, medio ad
costam lobatze, lobi leviter obliqui, rotundati vel acuti, margine
minute dentati. Pinnee maxime ad 60 cm. longe, 16 cm. late,
pinnate. Pinnule leviter oblique, omnes ad rachin adnate;
pinnularum infimarum basis adnata 5 mm. lata, superiorum
latior. Pinnule maximz 9 cm. longe, 2.3 cm. late, basi angulo
45. cuneate, e basi sensim angustatee, apice 1.5-2 cm. multo
angustatee, margine leviter serrate, textura firmiter herbacee,
colore subtus pallide, supra atrovirides. Venule more
D.accedentis anastomosantes, et soros ferentes.

Tyvpr.—Pahang, Fraser Hill, in Tras valley, 3,800 ft.,
Singapore Field Number 21635 (Holttum), in Herb. Singapore.

This species is in every way closely allied to D. accedens
Bl., but is bipinnate (with slightly adnate pinnules) instead of
simply pinnate. D. stenochlamys C. Chr. appears to be an
allied but much smaller species. The type of D. insigne was
a very fine large plant, inhabiting moist shady valley forest.

Votrrides.. (1037):

124

The mountain valleys of Malaya at 2,o00—4,000 ft. altitude have
been much less explored than the ridges, and, especially in ferns,
contain a very rich flora. It is possible that D. insigne may be
fairly widely distributed at such altitudes on the main range of
the Peninsula. I found a young sterile plant, probably of this
species, at about 2,000 ft. altitude near Ginting Simpah, some
25 miles south of Fraser’s Hill, on the other side of the range.

Diplazium latisquamatum Holttum sp. nov.

Caudex brevis erectus. Frondes juveniles circinnatae
squamis nigris rotundatis vel ovatis peltatis integris imbricatis
vestitae. Stipites ad 90 cm. longi, basin versus vel omnino
scabridi vel papillati, cum rachibus minute paleacei. Lamina
ad 1.5 m. longa et zim. lata, tripinnatifida. Costae supra
bialata, subtus squamis parvis subrotundatis paucis munitae,
famina cetera glabra. Pinnule sessiles basi late cuneate vel
subtruncatae, apicem versus leviter angustatae, ad alam 2-3 mm.
latam lobate, apice acuminate serrate, textura firmiter
herbaceae; lobi leviter obliquae, plerumque 4 mm. lati, oblingi,
apice truncati, subintegri vel serrati. Venulae in _ lobis
5-6-jugatee, simplices vel superiores furcatae. Sori dimidium
inferum venularum solum occupantes; indusia persistentia.

Type. Pahang, Sungei Terla (Ulu Telom), S.F.N. 31311
(Holttum), in Herb. Singapore.

Other specimens.—PAHANG: Cameron Highlands $.F.N.
23207, 23360, 31353, 31361. .B. N. Borneo: Mt. Kamabala,
S.F.N. 25429; Clemens 27122, 32952, 32551, 33082, 32454, 325106.

The characteristic features of this species are the broad
almost black imbricating scales on the young circinnate fronds,
the tripinnatifid habit, and the sori confined to the basal half of
the veins. In other characters there is much variation, due in
part to the fact that plants immature in size may be fertile. The
scales on the stipes, as the frond expands, are mounted on soft
papillz, which remain after the scales have fallen; but the degree
_ to which these papillee are developed seems to be very variable.
S.F.N. 23297 has abnormally broad lobes to the pinnules, the
ends rounded, and sometimes lobed again. In other characters it
agrees with other specimens, and in view of the variation of
cutting which clearly exists in the species, I can only regard it as
a local variation. ‘The Kinabalu specimens agree well, the only
notable differences being that the scales tend to be ovate-acute,
and the stipes on the whole more smooth.

This species is characteristic of moist shady valley forest in
the neighbourhood of Cameron Highlands, at 4,000—5,o000 ft.
The plants often grow in very wet ground.

The Kinabalu specimens were at first identified with
D. atrosquamosum (Copel.) C. Chr. ‘This is clearly an error, in

Gardens Bulletin, S.S.

125

view of the very broad scales, the scales of D. atrosquamosum
being narrow.

A recent collection (S.F.N. 31193, Holttum) from an
altitude of 2,500 ft. at Ginting Simpah (on the Main Range, south
of Cameron Highlands) appears to belong to this species. The
pinnules are broader than in any other collection, but the plants
agree in other respects.

Diplazium subintegrum Holttum sp. nov.

Caudex brevis erectus. Stipites ad 80 cm. longi, basin
versus squamis brunneis nitidis integris ad 1.5 cm. longis 2 mm.
latis acuminatis vestiti, cetera nudi, straminei. Lamina pinnata,
ad 70 cm. longa et 4o cm. lata, pinne ad 16-jugatze; apex
laminze basin versus lobatus, cetera more pinnarum serratus.
Rachis supra sulcata, in sulco minute papillata, subtus nuda.
Pinnee sub-basales maxime (infime leviter breviores, non
deflexee), petiolutatee 8 mm., ad 24 cm. longe et 2.5 cm. latae,
basi cuneatze vel rotundatee, margine serrate, apice acuminate;
textura subcoriacez; colore supra atrovirides, subtus pallide.
Costee subtus squamis linearibus brunneis paucis munite, cetera
nudze; venze prope costam furcatee, ramus acroscopicus simplex,
ramus basiscopicus I- vel 2-furcatus. Sori semper in venula
acroscopica, e costa % — 34 ad marginem, sedentes; rami venulz
basiscopicee interdum soros breves ferentes. Indusia simplicia vel
duplicia, lata, persistentia.

Tyvpr. Pahang, Sungei Burong, Cameron Highlands, 5000
ft. S.F.N. 31350 (Holttum), in Herb. Singapore. Frequent in
forest.

Other specimens.—PAHANG: Cameron Highlands, 4800 ft.
S.F.N. 23338 (Holttum); G. Berembun, Ridley 13969; Fraser’s
Hill, gooo ft. S.F.N. 8818 (Burkill and MHolttum), 21539
(Holttum), G. Hose 15. Perak: ‘The Cottage, Taiping,
H. D. Hervey s.n. 1889; G. Batu Puteh, summit, Wray 314.
SELANGOR: Bukit Kutu, 3,000 ft. Ridley 7844.

This species is allied to D. Prescottianum (Wall.) Moore,”
but differs constantly in the pinnee never being in the least degree
lobed nor auriculate at the upper base. It is frequent in the
mountain forests on the Main Range, at least at Fraser’s Hill
and Cameron Highlands and is so constant that I cannot but
regard it as a good species, though intermediates between it and
D. Prescottianum appear to exist at lower altitudes.
D. Prescottianum, founded on a specimen from Singapore, needs
more study in the field. There are some specimens from Ginting
Simpah (alt. 1500 ft.) which seem to connect it with
D. subintegrum; but as D. Prescottianum has been rarely collected
in true lowland forest we still have no proper understanding of its
range of variation.

Vol. IX. (1937).

126

Diplazium velutinum Holttum sp. nov.

Caudex brevis erectus. Stipites plerumque 20 cm. longi,
fusci, omnino pilis mollibus pluricellulis brunneis, basin versus
squamis angustis nigris integris vestiti. Lamina anguste
deltoidea, ad 25 cm. longa et 15 cm. lata, bipinnata. Raches
pilis brunneis omnino dense velutinze, squamis angustis nigris
paucis intermixtis. Pinnze liberze 12-15-jugate. Pinnze sub-
basales maximee, ceterze sensim reductee, superiores in apicem
brevem lobatum laminze confluentes. Pinnee maximz ad 8 cm.
longee et 3 cm. late, plerumque 6 cm. longe et 2 cm. late, sub-
sessiles, e basi lata ad apicem sensim angustatee; basin versus
pinnate, apicem versus ad alam angustam lobate. Pinnule
liberee pauce, cetere ad rachin plus minusve adnate.
Pinnule vel lobi maximz oblongze, leviter oblique, margine
serratee vel lobate, apice rotundate, textura chartacee.
Venule in pinnulis ad 7-r10-jugate, simplices vel furcate.
Sori e costa % — % ad marginem in venulis sedentes : in venulis
furcatis ramum acroscopicum solum occupantes. Indusia
sororum infimorum duplicia, cetera simplicia.

Type.—Pahang, Cameron Highlands (47th mile, Telom Rd)
4800 ft. S.F.N. 31221 (Holttum), in Herb. Singapore. Frequent
in shady forest by rocky stream. Also Cameron Highlands, 4800
ft. “S/F IN. 23428 "\(Holttim):

This species is closely related to D. tomentosum Bl., and
there are a few specimens which are intermediate between the
two species, but it differs very strikingly from _ typical
D. tomentosum. in the following characters: the pubescence on
stipe and rachis is more abundant, and the lower pinnze are always
pinnate at the base, even in fronds of moderate size. The most
extreme form, represented by one plant, has the pinnules lobed
half way to the costa. In most cases the pinnules are serrate, the
upper base tending to be auriculate.

D. tomentosum is a common fern of lowland forest in the
Malay Peninsula, and there are a large number of collections in
the Singapore herbarium. ‘The range of variation is considerable,
and in some cases the lower pinne are dissected to the costa.
This extreme form is connected with D. velutinum by a few
specimens from altitudes of about 3000 ft. In view of the con-
stancy of D. velutinum in the neighbourhood of Cameron
Highlands, I regard it as a mountain derivative of
D. tomentosum; the intermediate specimens are possibly hybrids.

D. velutinum grows in the wettest and shadiest mountain
forest, near streams.

I have to thank Dr. Carl Christensen for examining
specimens, and for suggesting the name velutinum.

Gardens Bulletin, S.S.

127 ,

Dryopteris brunnea (Wall.) C. Chr. var. glabrata (Clarke).

Ching places this species in the genus Thelypteris (Bull. Fan.
Mem. Inst. 6: 272. 1936), in which he is doubtless right, but
I am not yet in a position to segregate the Malayan species of this
genus. The object of this note is to record the collection of the
var. glabrata at Cameron Highlands, alt. 5000 ft. (Holttum s.n.,
May 1936). ‘The specimens are small, but clearly referable here.
The solitary plant seen was growing among rocks beside a stream
in a recently made clearing. D. oppositipinna (see below) is
clearly allied, but is much smaller with most of the pinnze broadly
- sessile.

Dryopteris crenata (Forsk.) O. Ktze.

This widely distributed species has recently been collected
for the first time in the Peninsula, on limestone at Gua Teja in
Kelantan (S.F.N. 29682, Henderson).

Dryopteris dissecta (Forst). O. Ktze.

The first recorded collection of this species in the Malay
Peninsula is Holttum 31292 from Cameron Highlands, alt.
5000 ft. The plants were found growing on the banks of a
stream in shady forest. They are large, and agree with some
Philippines specimens. They have the general aspect of
Tectaria. For a note on this species see C. Christensen in this
Bulletin, Vol. 7, p. 253.

Dryopteris glandulosa (Bl.) O. Ktze Rev. Gen. Pl. 2: 812. 1891.

Aspidium glandulosum Bl. Enum. 144. 1828.

Dryopteris ividescems v.A.v.R. Bull. Jard. bot. Buitenz. IL
peel Y Il. 1Q13.

Dryopteris malayensis C. Chr. Vid. Selsk. Skr. VII. 16:
iat, nota. 1913.

It appears to me that by the rules of nomenclature
Christensen was wrong in transferring Polypodium glandulosum
Desv. to Dryopteris (see Index Suppl. 33. 1913), when the
name D. glandulosa was already occupied by Blume’s species;
the new name D. malayensis was therefore unnecessary. I
have examined the type of D. glandulosa (Bl.) O. Ktze at
Leiden, and I am of opinion that certain Peninsular ferns,
which I had noted as being identical with D. iridescens v.A.v.R.
are referable to this species. From description, it appears to me
likely that D. excrescens Copel, and possibly also D. Bartlettii
Copel. fall also within the range of variation of the species.

Blume’s specimen at Leiden has the lower surfaces very
uniformly glandular, with very short hairs on the veins, the
upper surfaces uniformly appressed-hairy. Among Peninsular
specimnens the pubescence of the upper surface is very variable,
even on the same plant; fertile fronds may be hairy when the

Pol. PX: (1937):

128

sterile are almost glabrous. Characters of pubescence can
therefore not be relied upon in this species. A character which
seems to be uniform and distinctive is the presence of spherical
yellow glands on the sporangia; these glands are similar to those
on the lamina. In the subgenus Abacopteris, the sporangia are
nearly always setose or glandular in some characteristic way.
Occasionally neither setze nor glands are present; I have noted
that this is the case in D. lineata (Bl.) C. Chr.

The Peninsula ferns of D. glandulosa are all of recent
collection, and with one exception are from Pahang; there is
also a specimen from Patani in Lower Siam. ‘The duplicates of
the specimens have been distributed variously as D. lineata,
D. malayensis and D. iridescens. Older collections named and
distributed as D. glandulosa are mostly D. mirabilis Copel. (see
C. Chr. in Gard. Bull. S.S. 4: 391. 1929). The specimens are
as follows:

MALAY PENINSULA.—NEGRI SEMBILAN: Ulu Bendul,
Holttum 9886. PAHANG: Base of G. Senyum, Henderson
22289; K. Tembeling, Holttum 20533; Ulu Sungei Merapoh, Md
Nur 11938; Tembeling, abundant near stream, Holttum 24702.

SUMATRA.—MENTAWI ISLANDS: Siberut, Boden Kloss 13087.
S1aM.—PATANI: Banang Sta, Eryl Smith 1858.

Dryopteris lineata (Bl.) C. Chr. Ind. Fil.
Aspidium lineatum Bl. Enum. 144. 1828.

As this species has been confused with D. glandulosa, the
specimens regarded as true D. lineata are enumerated below.
D. lineata is clearly distinct in the absence of glands on the
lower surface of the pinnee, the absence of hairs on the upper
surface, the somewhat pustular nature of the surfaces, the upper
base of the pinne strongly auricled, the pinnz almost entire.
In the Peninsula the species has only been found in the
neighbourhood of Cameron Highlands (on the Main Range,
near the Perak-Pahang boundary). All specimens of which the
altitude is given were collected in the mountains; but a specimen
from Patani in Lower Siam was collected at a low altitude. The
plants grow on rocks by forest streams. The specimens are:

PERAK.—Gopeng, King’s Collector 497; no _ locality,
Scortechini. PAHANG: Telom, Ridley 13997; Cameron High-

lands, 4800 ft., Holttum 23362 and s.n. PATANI: Banang Sta, .

70 m., in earth on rock in large stream, Eryl Smith 2026; near
Betong, on log in stream, 220-270 m., Eryl Smith 2027.

Dryopteris multisora C. Chr. Gard. Bull. S.S. 7: 241. 1934.

This species, described from Mt. Kinabalu, B.N. Borneo,
has been collected also in the Malay Peninsula, at Cameron
Highlands, Holttum 23345, 23437 It belongs to a group of
species which are only found in mountain forests.

Gardens Bulletin, S.S.

Eee

129

Dryopteris oppositipinna v.A.v.R. MS in Herb. Bog.
Phegopteris oppositipinna v.A.v.R. Bull. Jard. bot. Buitenz.
mrser. XVI:»24 Tord.

The type of this species was collected by Matthew on Mt.
Singgalang, Sumatra (no. 518). I found it for the first time in
the Malay Peninsula at Cameron Highlands (Holttum 23261),
on wet rocks by a small stream in a rather open place at 4000
to 4,500 ft. altitude. The peculiar almost opposite broadly
sessile pinnze are very distinctive; the lower surfaces of rachis,
coste and lamina bear numerous short stiff pale erect hairs;
similar hairs are also sometimes borne by the sporangia. There
is no trace of indusium.

Dryopteris Parishii (Hk.) O. Ktze Rev. Gen. Pl. 2: 813. 1891.
Nephrodium Parishti Hook. Spec. Fil. 4: 131, t. 260. 1862.

Ridley does not record this species as occurring in the
Malay Peninsula, but it was collected by Curtis at Langkawi
(Nov. 1889), in crevices of limestone rocks; it has since been
found on the mainland, in Perlis, at Bukit Lagi (Corner and
Henderson 22876, 23135). Plants were brought to Singapore,
and cultivated in pots under glass. The fronds died down
periodically; the plants were then dried off and rested until new
growth began. ‘The resting habit appears to be obligatory in
this species. The same is true of the allied Dryopteris
chupengensis (Ridl.) C. Chr. Plants from the type locality were
brought to Singapore by Messrs. Corner and Henderson in 1929,
and grown in pots alongside D. Parishii. The two species are
so closely similar in form of frond that I thought at first that
D. chupengensis might be only a young stage. The two
remained distinct however, and grew side by side for about
three years. The largest dimensions of lamina attained by
D. chupengensis under cultivation were 10 by 9 cm.

Dryopteris polita Rosenst. Fedde Report. 13: 218. ro14.

A record of the occurrence of this species in the Peninsula
was nade in this Bulletin, Vol. 7, p. 252. It has been found
only at Cameron Highlands (Holttum 23451), and was
distributed as D. sparsa. I have seen the Leiden duplicate of
the original collection of D. polita (Batakerland, F. Winkler;
Rosenst. Fil. sumatr. exsic. 178); it is clearly the same as the
Peninsula collection above cited.

Dryopteris pseudocalcarata C. Chr. Ind. Suppl. III: 95. 1934.

Aspidium ciliatum Wall. 1828 (nomen); Nephrodium Clarke
Trans. Linn. Soc. Bot. 1: 514. 1880 (non Desv. 1827).

Lastrea calcarata var. sericea Ridl. Journ. Mal. Br.
Peasiatic Soc. 4: 65. 1926.

Roms PX. (7937).

130

This species is very distinct from D. calcarata. It is
common on mossy rocks beside mountain streams in the
Peninsula, but does not occur in the lowlands; it has not been
found in the south of the Peninsula.

Dryopteris sparsa (Ham.) O. Ktze Rev. Gen. Pl. 2: 813. 1801.

The only specimen of this species I have seen from the
Malay Peninsula is Holttum 25303 from Sungei Sengam, Perak,
alt. 3500 ft., on rocks and damp ground near stream, in forest.
This specimen is near var. nitidula, but the segments are more
rounded than in Beddome’s illustration. The Sungei Sengam
is a small stream on the Perak side of the main range, just
below the road up to Cameron Highlands. Ridley quotes a
specimen of Matthew’s (at Kew) from Maxwell’s Hill.

Dryopteris stipellata (Bl.) GO. Ktze Rev. Gen. Pl. 2: 813. 1801.
Aspidium stipellatum Bl. Enum. 152. 1828.

I have examined Blume’s specimen at Leiden, and find that
some specimens from the Peninsula agree with it. The species
is rather abundant by streams in shady valleys at Cameron
Highlands; the young unexpanded fronds are covered with
mucilage, through which the prominent pale aerophores appear.
The surfaces beneath are glabrous, not glandular. A charac-
teristic feature is the sudden narrowing of the apices of the
pinne. The specimens at Singapore are:

PERAK: King’s Collector 6345, 7174 (distributed as
D. heterocarpa). PAHANG: Cameron Highlands 4800 ft. Holttum
23352. JOHORE: Gunong Muntahak, 600 ft., Holttum 19923.
Hymenophyllum exsertum Wall. Hook. Spec. Fil. 1: 109, t. 38A.

1844.

‘This species has not hitherto been recorded as occurring in
the Peninsula. "The following specimens appear to belong to it;
the indusia tend to be narrowed to the apex and entire, but
there is much variation, and the sori are often aggregated in the
apical portion of the frond. PAHANG: Gunong Tahan, Holttum
20738; Fraser’s Hill, Holttum 21560.

Hypolepis bivalvis v.A.v.R. Bull. Jard. bot. Buitenz. IJ ser.

mm VISTO OMA:

Christensen identified a specimen from G. Tahan as this
species; I find that other specimens are also referable to it. All
have been distributed as H. tenuifolia. ‘The type of H. bivalvis
is from G. Sago, Sumatra. Peninsula specimens are charac-
terised by the presence throughout of viscid glandular hairs;
in herbarium specimens these hairs. are dried and their viscid
character lost. The species has only been found in open places
on mountain ridges.

PAHANG: G. Tahan, 6000 ft. Holttum 20763; Fraser’s Hill,
4000 ft. Burkill and Holttum 8498, Holttum s.n. 1923, Eryl
Smith 817, G. Hose 7.

Gardens Bulletin, S.S.

&

EE

Leptochilus decurrens Bl. Enum. 206. 1828.

This species is placed in Campium by Copeland (Phil.
Journ. Sci. 37; 351).. In his key to the genera concerned in
the paper quoted, Copeland gives as a distinction between
Campium and Leptochilus that the former has a creeping
terrestrial rhizome, the latter being scandent or epiphytic. It
may be interesting to record that I collected at Cameron
Highlands a number of plants of L. decurrens (Holttum 23546)
the rhizomes of which were scandent to the extent of at least
one metre from the ground, on small trees by a stream in shady
forest.

Leuccstegia immersa (Wall.) Pr. Tent. 95, t. 4, f. 11. 1836.

This species has not been recorded as occurring in the
Peninsula. There is a specimen in the Singapore herbarium,
from Gunong Tahan (Ridley 16088).

Lindsaya javanensis Bl. Enum. 219. 1828.

L. flabellulata var. gigantea Hk. Spec. Fil. 1: 211, t. 63¢.
1846.

Ee tenera var. giganiea Holttum Gard. Bull..S.S. 5: 65.
1930.

Dr. O. Posthumus has called my attention to this species,
and thireugh the-courtesy of Prof..Dr. H. J.. Lam I have
received a photograph of the type. I believe that it is identical
with Hooker’s variety above cited, of which I have seen the
type at Kew. In my paper on the species of Lindsaya in the
Malay Peninsula, I maintained this fern doubtfully as a variety
of L. tenera. I believe that it is quite distinct; it should there-
fore bear Blume’s name.

Lindsaya lucida Bl. Enum. 216. 1828.

L: Lobbiana Hk. Spec. Fil. 1: 205, t. 62c. 1846. Holttum
Gard. Bull..S.S. 5: 61. 1930.

I have examined the type of this species at Leiden, and
I can see no distinction from L. Lobbiana Hk., the type of
which also came from Java. ‘The Peninsula plants hitherto
called L. Lobbiana are now therefore referred to L. lucida; there
is however some possibility that they are distinct from the Java
species.

Lindsaya nitens Bl. Enum. 217. 1828.

L. recurvaia Wall. Hk. Spec. Fil. 1: 222, t. 7oa. 1846.
Holttum Gard. Bull. S.S. 5: 66. 1930.

I have seen the type of L. nitens at Leiden, and find that
it is indistinguishable from L. vrecurvata; Blume’s name
should therefore be adopted for the species. The Leiden
specimen has simply pinnate fronds, the lower pinnze somewhat
recurved and their upper edges incised.

Vol. 1X .1(1637):

132

Lindsaya nitida Copel. Phil. Journ. Sci. 6c: 138, t. 21. Ig1t.
L. integra Holttum Gard. Bull. $.S. 5: 67. 1930.

I have seen the type of Copeland’s species in Brooks’s
herbarium at the British Museum. It is bipinnate, but in other
respects agrees with my L. integra. Brooks’s specimens was
gathered at 3000 ft. on Mt. Penrissen, and it is possible at that
this altitude the species grows to a larger size than by the
lowland rivers of the Peninsula. Many species of Lindsaya have
fully fertile fronds before they are large enough to bear fronds
of fully mature size and branching. Christensen in Gard. Bull.
8: 237 suggests that this species may be identical with L. nitens
Bl. In my opinion the latter species is quite different, being
identical with L. recurvata Wall.

Lygodium borneense v.A.v.R. Bull. Jard. bot. Buitenz. II ser.

XX: 29. 1915. Holttum, Journ. Mal. Br. R. Asiat. Soc. 6:
16.. 1928:

This remarkable fern, first discovered in Borneo, was
collected by Boden Kloss on the Mentawi Islands, off the west
coast of Sumatra. The gap in distribution has now to a small
extent been filled by the discovery of plants in Johore, by
Mr. E. J. H. Corner (Sungei Dohol, Ulu Sedili, Corner 26053).
The plants were found in fresh-water swamp forest. ‘This
peculiar habitat may be part of the reason that the species has -
been so little collected. Fresh-water swamp forests are difficult
botanical hunting grounds, and have been inadequately explored,
a fact which is well shown by several recent discoveries by
Mr. Corner.

Microlepia pilosula (Wall.) Pr.

A specimen recently collected at Cameron Highlands
(S.F.N. 31293, Holttum) is closely similar to a specimen from
Yunnan determined as M. pilosula by Mr. R. C. Ching. I have
sent a specimen of the Cameron Highlands plant to Mr. Ching,
who confirms the identification. The plant was found growing
in the bed of a forest stream, under very moist conditions, at
about 5000 ft. altitude. Its characteristic feature is the rather
stout stiff pale glistening hairs on the rachises and veins beneath.
A similar specimen from northern Siam (Doi Sutep, 1250-
1400 m:, Winit 1212), in the Singapore herbarium, was also
found on moist rocks in a stream bed in evergreen forest. ‘This
is a great contrast to the dry open habitat favoured pe the
common M. spelunce.

Nephrolepis barbata Copel. Perk. Frag. 178. 1905.

Christensen has seen the type of this species, and has
identified with it specimens from the Peninsula; I am thus able
to enumerate the specimens in the Singapore herbarium, which
have hitherto been named WN. exaltata. The species is
characterised by relatively narrow fronds (to about 9 cm. wide),
often of great length, the pinnz close, strongly falcate, usually

Gardens Bulletin, S.S.

133

drying rather pale green. The plants nearly always grow on
rocks, or sometimes on trees, their fronds being pendulous,
sometimes well over two metres in length.

PENANG: Waterfall, Ridley 7038. PERAK: Bujong Malacca,
Ridley 9607; Relau Tujor, L. Wray 2207; Larut, King’s
Collector 5229, 5220. NEGRI SEMBILAN: Ulu Bendul, Holttum
9820. SELANGOR: Gua Batu Woods, Ridley s.n. Dec. 1806.
PaHANG: Tahan River, near K. Teku, Holttum 20807; Gua
Tipus, Chigar Perah, crevices in limestone, Henderson 19405.

There are also specimens from Siam.

Ophioglossum simplex Ridl. ex Bower, Nature 64: 617. 1901.

This curious species, originally reported from Sumatra, has
been found on the summit of Gunong Panti, Johore. It was
growing on the ground in the shade of a thicket which had
grown up on the summit after clearing. ‘This is the first record
for the Peninsula.

Osmunda javanica Bl. Enum. 252. 1828.

The first collection of this species in the Peninsula was made
by me in the valley of the Sungei Sengam, Perak (Holttum
25302), at about 3500 ft. above sea. The plants were quite
abundant in shady forest, on the valley-sides. "They are in
appearance very like Cyathea brunonis, for which reason they
may have been overlooked in the past, but when fertile they are
naturally quite unmistakable. The very young unexpanded
fronds are covered with mucilage, in the manner of Plagiogryria,
but there are no aerophores. ‘The pinnez are articulate, another
resemblance to C. brunonis. Plants are growing quite well in
cultivation in Singapore and have produced spores. It has since
been discovered that Osmunda javanica is very abundant beside
the Telom river at about 3900 ft. alt., growing in large clumps
on rocks, and also in the forest near by.

Plagiogyria adnata (Bl.) Bedd. F.B.I. t. 51. 1865.

No record has been made of the occurrence of this species
in the Peninsula. There is however a collection of Mr. Ridley’s,
from Gunong Tahan (Ridley 16002); this was distributed as
Polybotrya appendiculata var. subintegra.

Polypodium albidosquamatum Bl. Enum. 132. 1828.

This species has never been recorded as occurring in the
Malay Peninsula. It was collected by Kunstler in Perak, and
was distributed as Polypodium leiorhizon Wall. ‘The specimen
in the Singapore Herbarium is labelled ‘‘King’s Collector, no.
10703’’, and bears no exact locality. The species is of wide
distribution, and it is rather surprising that it has only once
been found in the Peninsula. ‘The Singapore specimen has only
fertile pinnee, which are rather unusually long.

Vol. IX. (1937).

134

Polypodium grandidentatum (Cesati) Bak. Ann. Bot. 5: 470.
18o1I.

P. dilatatum var. grandidentatum Ces. Att. Ac. Nap. 7, 8:
272070:

This species, allied to P. Hancockii (see below), appears to
be quite distinct. In the Peninsula it is a lowland plant,
whereas P. Hancockti has only been found at about 5000 ft.
P. grandidentatum when mature has much larger fronds than
P. Hancockii, and also differs in having the lower lobes gradually
reduced in size, whereas in P. Hancockii the lowest lobes are
the largest. P. grandidentatum was described from a plant from
Matang, Sarawak, which I have seen. ‘The species usually grows
on rocks, but in my experience has not the stiffly erect habit of
P. Hancockii.

Polypodium Hancockii Bak. Journ. Bot. 1885 106.
P. euryphyllum C. Chr. Index.

In the Peninsula this species has only been collected at
Cameron Highlands (Holttum 23501); the specimens formerly
distributed as P. dilatatum Wall. or P. euryphyllum are P. grandi-
dentatum. P. Hancockii is found on mossy rocks beside small
mountain streams at about 5000 ft. altitude, in shady places.
The rhizome clings to the rocks and the fronds are erect.

Polypodium insigne Bl. Enum. 127. 1828.

This species has been collected in the Peninsula, but not
reported. It is apparently very variable in the width of the
pinne; in this character Peninsula specimens agree well with
the type. ‘The species appears normally to grow on rocks beside
forest streams, but two specimens collected at Fraser’s Hill were
climbing trees to a few feet from the ground. Ridley’s specimens
were distributed as Aspidium decurrens.

PAHANG: Telom River, Ridley 13978; Fraser’s Hill, G. Hose
9, Holttum 8789. PERAK: Sungei Sengam, 3500 ft., Holttum
Stl. LOZ F.

Polypodium normale Don, Prodr. Fl. Nepal. 1. 1825.

This species has not yet been recorded as occurring in the
Malay Peninsula. I collected it at Cameron Highlands, at
4800 ft., on a tree on the edge of the forest (Holttum 23363).
The specimen appears to me quite identical with P. normale from
Assam, and also with P. subnormale v.A.v.R. from Sumatra.

Polypodium malaicum v.A.v.R. Handb. 577. 1909.

Polypodium sessilifolium Hk. Spec. Fil. 4: 168, pl. 268.
1863.

The question of the type of this species is a rather difficult
one. Hooker in his original description quoted three specimens,
but did not indicate a type. The specimens are: Luzon, Cuming

Gardens Bulletin, S.S.

135

382; Penang, Sir W. Norris; Java, Blume. An inspection of
the sheets at Kew indicates that Hooker’s figure was drawn
from another specimen not cited by him, namely Cuming 222
from Luzon. ‘This figured specimen is on the same sheet at
Kew as Cuming 382 and Norris’s plant from Penang; both of
the latter are very much smaller than the figure, which agrees
closely with Cuming 222. Examination of the lists of Cuming’s
numbers show that Hooker made a mistake in regarding no. 382

~~ as from Luzon; it was actually collected in Malacca, and it is

certainly identical with P. swhevenosum Bak., the type of which
is from Penang.

Hooker’s description indicates that he included Cuming 222
in his species (he gives the length of the fronds as 3—8—9 inches);
in fact, the species is a composite one, including P. subevenosum
from the Peninsula and a distinct species, represented by Cuming
222, from Luzon. ‘There are clearly two alternatives: either we
may regard Cuming 382 as the type of P. sessilifolium Hk.,
in which case that species becomes a synonym of P. subevenosum
Bak. and a new name is required for the Luzon plant; or we may
select Cuming 222 as the type of P. sessilifolium Hk., in which
case the species is quite distinct, and should bear the name
P. malaicum. I prefer the latter alternative, more especially as
it agrees with the treatment which has usually been given to
the species. Christensen refers to the species in this sense in
this Bulletin, vol. 7, p. 292, and remarks that a specimen from
Kinabalu is identical with Copeland’s Pterid. Philip. exsic. 141,
distributed as P. sessilifolium, and notes the presence on both
- of setose sporangia. Cuming 222 has not been examined to find
out whether it has this character.

My proposal is therefore to regard Cuming 222 as the type
of this species, excluding the Peninsula specimens quoted by
Hooker (Cuming 382 and Penang, Norris); it is much larger
than P. subevenosum and most of the veins are forked, with a
probable distinguishing character in the setose sporangia.

Polystichum prolificans v.A.v.R. Bull. Jard. bot. Buitenz. III
ser. 2: 170. 1920.

A species of Polystichum is quite common on the large
granite boulders in shady gullies on Penang Hill; the same
species has been collected in Perak (King’s Collector 6258).
These plants have been named P. biaristatum Bl. (Ridley,
Journ. Mal. Br. R. Asiat. Soc. 4: 61. 1926) I have examined the
type of P. prolificans (from Sumatra) and find that the Perak
specimen is identical; the Penang specimens in the Singapore
herbarium are all smaller than the type, but are otherwise similar.

Pteris platysora Bak. Journ. Bot. 1880: 211.

_ A specimen from Gunong Kerbau (Haniff 14743) in the
Singapore herbarium appears to be referable to this species, the

Vol. IX. (1937).

B

136

type of which is from Sumatra. The specimen represents a
large and handsome plant, quite unlike any other Pteris so far
known in the Peninsula. This species is apparently very near
P. insignis Mett.

Pteris scabripes Wall. Hook. Spec. Fil. 2: 165. 1858.

The type of this species is Wallich no. 94 from Penang.
As stated by Hooker, it is very near the widely distributed P.
cretica L., and I am not in a position to judge how good its
claims may be to rank as a district species. The sterile pinnze
are very close together, so that the frond is often almost flabellate;
the edges of the pinne are finely serrulate throughout, by which
this species is clearly distinguished from the following. The

species is rather local in its occurrence, though sometimes fairly —

abundant. Specimens from the Peninsula are:

SELANGOR: 15th mile Pahang Track Ridley s.n. July 1897.
PENANG: Jerantut, Holttum 24791; Tembeling, Holttum 24690.
PERAK: Upper Perak, growing on rocks, 300 ft., L. Wray Jr.
3699; Telok Pinang, Ipoh, Ridley s.n. Oct. 1808. KeEpDAH:
Burau Bay, Langkawi, Ridley 15789; Langkawi, Curtis s.n. Sept.
1890; Foot of G. Raya, Langkawi, Curtis 3381.

Pteris venulosa Bl. Enum. 209. 1828.

It appears probable that the specimens from the Peninsula
hitherto referred to P. pellucida Pr. are P. venulosa Bl. I could
not find the type of the latter species at Leiden, but saw several
other specimens from Java, with which Peninsula specimens
agree, with the exception that most, of the latter have fewer
pairs of pinne, and some have the pinne so close as to be almost
digitate. In habit, Peninsula plants agree rather closely with
P. scabripes, with which they have been confused. They differ

very strikingly however, in having the margins of the sterile ©

pinnze entire and slightly wavy or crisped, except towards the
apex, where they are broadly and rather bluntly toothed; this
contrasts with the pinnz of P. scabripes, which are finely serrate
throughout. Peninsula specimens are as follows:

JoHORE: G. Pulai, Hullett s.n. Aug. 1880; G. Pulai, rocks
by the stream, Ridley s.n. Dec. 1905; G. Berhidong, on wet rocks
1,800 ft. by stream, Holttum 10995; G. Pulai, G. A. Best 7721.
NEGRI SEMBILAN: G. Angsi, 2000 ft. Md. Nur 11582.

SELANGOR: C. Curtis s.n. May 1902. PERAK: Larut, 2800- ~

3000 ft., King’s Collector 3811; Maxwell’s Hill 3000 ft. Scorte-

chini 405: 444. PENANG: Government Hill 2000 ft. Haniff ©

7590; Hullett s.n. December 1881.-

Quercifilix zeyianica (Houtt.) Copel. Phil. Journ. Sci. 37: 409.
1928.

This species, though of wide distribution, has never been

collected on the mainland of the Malay Peninsula. It has been
found on the island of Pulau Tioman, where it was collected

Gardens Bulletin, S.S.

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137

at the South Cape, nestling in rocks, by Burkill in r9r5 (Burkill
1034).
Rumohra aristata (Forst.) Ching, Sinensia 5: 50. 1934.

Under Polystichum aristatum, Ridley (1926, p. 61) quotes his
no. 15966 from Gunong Jahan; this is however R. puncticulata
(see below). The only collections of Rumohra aristata known
to me from the Peninsula are as follows. In one case plants
were found growing on rocks beside the Tahan river at about
500 ft. altitude (Holttum 20805); these were small, but quite
similar to specimens from southern China. The other collection
(Holttum 24999) was from a more shady position, from rocks
beside a small forest stream near the Sungei Sengam, Perak, at
about 3000 ft. (just below Renglet); the fronds of this were
larger and thinner in texture.

Rumohra puncticulata (v.A.v.R.) Holttum comb. nov.

Polystichum puncticulatum v.A.v.R. Bull. Jard. bot.
Buitenz. III ser. 2: 171. 1920.

I have seen the type of this species, from Sumatra, and I
believe that specimens from the Peninsula (also Mount Kinabalu)
are also referable to it. It is allied to R. speciosa (Don) Ching.
The Peninsula specimens are from G. Tahan and G. Kerbau,
our two highest mountains. They are: G. Tahan, Ridley 15066,
Holttum 20779; G. Kerbau, Haniff 4033.

Stenosemia pinnata Copel. Phil. Journ. Sci. Suppl. II: 146. 1906.

All Peninsula specimens of Stenosemia appear to agree with
this species rather than with S. aurita, to which they have been
previously referred. Typical specimens of S. aurita from Java
have triangular fronds, the lowest pinnze largest, and the lowest
basiscopic segments longest. In most Peninsula plants the
lowest pinne are not much larger than the next, and the lowest
basiscopic segments of the lower pinne are small, the largest
ones being at about one-third towards the apex. There is some
variation, but the Java form is never found. Stenosemia appears »
to be confined to limestone rocks in the Peninsula. It is well
established in shady rockeries at the Singapore Botanic Gardens,

_.where it grows on old coral and reproduces freely. The type

of S. pinnata is from Mindanao, and a study of the geographic
distribution of the forms of Stenosemia would be interesting.

Tectaria decurrens (Pr.) Copel.

This species has not yet been collected in the Malay
Peninsula. The specimens so named by Ridley (1926, p. 79)
are T. crenata Cav. and Polypodium insigne Bl.

Tectaria Keckii (Luerss.) C. Chr. Index Suppl. III: 181. 1934.

Aspidium Luerss: Bot. Centralbl. 11: 76. 1882.

The type of this species is from ‘‘Klang, bei den Gua Batu’’.
I have not seen it, but it seems to me highly probable that it is

Vol. IX. (1937).

138

the same as Aspidium amplifolium v.A.v.R., a common fern of
limestone rocks in the Peninsula, formerly referred to A. subtri-
pbhyllum. It is certainly allied to A. subtriphyllum, but is a
much large fern. Kehding collected both in the Peninsula and
Sumatra, and the locality Gua Batu is probably the same as
Batu Caves, near Kuala Lumpur.

Tectaria Maingayi (Bak.) C. Chr. Index Suppl. III: 182. 1934.

For sysnonymy of this species, see Holttum in this Bulletin,
Vals. p. 307.

Tectaria ternifolia (v.A.v.R.) C. Chr. Index Suppl. III: 185.

1934.

Aspidium v.A.v.R. Bull. Jard. bot. Buitenz. II ser. XI: 3.
TOES.

The type of this species is a specimen collected by Matthew
at Gopeng. The species has since been collected in Pahang, as
a locally common terrestrial fern in forest on ‘‘Raub Series’’
rocks. The fertile fronds are produced seasonally, on much
longer stipes than the sterile. Specimens seen are: Tembeling,
Holttum 24693; Henderson 21682; G. Senyum, Henderson s.n.
Ridley quotes a specimen from Pekan as Aspidium ternatum;
this species has not otherwise been found in the Peninsula, and
the specimen is perhaps T. ternifolia.

Trichomanes exiguum (Bedd.) Baker, Syn. Fil. 464. 1874.
Hymenophyllum Bedd. F.B.I. pl. 275. 1868.

A specimen (s.n.) gathered by me at Ginting Simpah
appears to be this species, which has not hitherto been recorded
as occurring in the Peninsula. The plants were growing on a
large boulder beside a stream in forest near 2000 feet above sea
level.

Trichomanes radicans Sw.

No specimen of this species, even widely interpreted as in
Copeland’s recent work (Phil. Journ. Sci. 51, p. 213), has
hitherto been found in the Malay Peninsula, the nearest being
a specimen from Siam. A recent collection from Cameron
Highlands (S.F.N. 23466, Holttum) fills this gap. The plants
are small, with fronds to 7 cm. long.

Vittaria malayensis Holttum Gard. Bull. S.S. 4: 409. Jan. 1929.

Apparently V. stenophylla Copel. from Luzon (name
published in October 1929) is the same as V. malayensis; there
is a specimen of Copeland’s (Pterid. Philip. exsicc. 122) in the
Singapore herbarium and I have compared it with Malayan
specimens. V. malayensis is not uncommon as an epiphyte on
the main range of the Peninsula.

Gardens Bulletin, S.S.

139

FURTHER NOTES ON STENOCHLZAENA,
LOMARIOPSIS AND TERATOPHYLLUM.

By Ry. EB. .HOLTTom.

In this Bulletin, Vol. 5, nos. 9-11 (1932), I gave an account
of the above genera, based on all the material then available to
me. It subsequently appeared that the bulk of the specimens of
“‘Stenochleena’’ from the Herbarium of the Bureau of Science,
Manila, had previously been sent to the U.S. National Herbarium
on loan and were not with the material I had for study from
Manila in 1932. Some of the Philippine collections were
represented by duplicates at Buitenzorg, and in the Singapore
herbarium, but a great many were not. I was therefore very glad
of the opportunity of examining the full Manila collections, which
has been possible through the courtesy of Dr. W. R. Maxon of
the U.S. National Herbarium and Dr. Quisumbing of the
Philippine National Herbarium. At the same time, Dr. Maxon
sent me on loan the specimens from the U.S. National Herbarium,
which in some cases supplied further information.

The result of the examination of these additional specimens
is that I am able to add a few notes about the Philippine species
of the three genera, and I am able also to correct a misapplication
of the name Lomariopsis leptocarpa. My thanks are due to
Dr. Maxon and Dr. Quisumbing, and also to the authorities of
the herbaria at Kew and the British Museum, where I discovered
some further specimens during my leave to Europe in 1934.

This paper contains the new combination Teratophyllum
leptocarpum (Fée) Holttum.

STENOCHLAENA.
S. fraxinifolia Presl, Epimel. 164. 1849.

The type of this is Cuming 347 from Negros. There is a
specimen (including fertile frond) in the British Museum

a herbarium and another in the U.S. National Herbarium; I can

see no distinction from S. palustris. The characters mentioned
by Presl seem to me of little diagnostic value.
S. juglandifolia Presl, Epimel. 164. 1840.
Lomaria? juglandifolia Presl, Rel. Heenk. 1: 52.
Stenochlena Milnei Und. Bull. Torr. Bot. Cl. 33: 38. 1906.

The type of this species is a specimen of Heenke’s from
Luzon; I have not seen it. In Epimel. bot. Presl quotes also
Cuming 133 partim. Cuming 133 is also quoted under
Stenochlena scandens (the species now called S. palustris) both
by Presl and John Smith (Journ. Bot. 3: gor).

Vol. IX. (1937).

140

All the specimens of Cuming 133 which I have seen (in the
herbaria of Kew, the British Museum, and Dr. E. B. Copeland)
are identical, and are intermediate in character between
S. palustris and S. laurifolia. ‘There are some remarks on the
subject in this Bulletin Vol. 5, p. 258. I now find that these
specimens agree with S. Milnei, a species overlooked when pre-
paring my former paper, the type of which I have examined at
Kew. The question therefore arises as to whether S. juglandifolia
and S. Milnei are synonymous.

The principal characters distinguishing S. Muilnei from
S. palustris are the broadly rounded bases of the sterile pinne,
and the width (5 mm.) of the fertile pinne. Presl gives the first
of these as distinctive of S. juglandifolia; as regards the fertile
pinne he states that they have a broad margin on either side ot
the narrow costal fertile zone, but he does not give any
measurements. Presl may perhaps have had a young fertile
frond. These facts, together with Presl’s citation of Cuming 133
seem to me sufficient evidence for accepting the identity of the
two species in question. The only doubtful point in Presl’s
citation of Cuming 133 under S. scandens; it is possible that this
was copied from John Smith.

S. juglandifolia is intermediate in character between
S. palustris and S. laurifolia. When preparing my former paper
I was in some doubt how to treat intermediate specimens, and
finally referred them to S. palustris, as they were clearly not
S. laurifolia, which is a very well-marked species as compared
with the variable S. palustris. I think however that
S. juglandifolia is sufficiently distinctive to warrant specific
separation. Its intermediate character is seen in the following
points :

. S. palustris S. juglandifolia S. laurifolia
Base of sterile cuneate broadly rounded cordate
pinne
articulation of complete partial not articulate
pinnee
width of fertile 2-3 mim. 5 mm. 7-10 mm.
pinnze
vascular bundles to about 40 intermediate over 100
1n stipe
spores tuberculate irregular parallel
broken ridges ridges

The statement about the spores of S. juglandifolia is based on an
examination of the spores of Milne 518 (the type of S. Milnei);
this specimen was referred by Hooker (Spec. Fil. 5: 251) to
S. laurifolia.

Gardens Bulletin, S.S.

I4I

The known distribution of S. juglandifolia is as follows: the
Philippines, New Guinea and the Solomon Islands. It therefore
corresponds approximately with that of S. laurifolia and is within
the range of S. palustris. It is possible that S. juglandifolia
represents a natural hybrid between the other two species. The
Philippine specimens referred to S. juglandifolia are as follows:
LUZON. Cuming No. 133 (K, B.M., W, Copel.), Irosin (Mt.
Bulusan), Prov. Sorsogon, Elmer 16253 (W, M). Los Banos,
Elmer 18242 (Copel., M, W). MINDANAO. Camp Keithley,
Lake Lanao, M. S. Clemens 944 (M). Butuan Subprov. Fenix
15907 (W).

S. laurifolia Pres], Epimel. 164. 1849.

I have examined the sheets of Cuming 226 (the type
collection) at Kew, and find that glands are present at the bases
of the pinne as in S. palustris. Hooker (Spec. Fil. 5: 251)
remarks on the absence of glands as a character distinguishing
this species from S. palustris; apparently he failed to observe
them. Owing to the broad cordate bases of the pinne of this
species, the glands are not so obvious as in S. palustris.
Additional specimens of S. laurifolia are:

Luzon. Guintacutan, Tayabas, Foxworthy and Ramos
13170 (M).

MINDORO. Bulalacas, J. Bermejos 1545 (M).

LOMARIOPSIS.

L. spectabilis (Kze) Mett. Fil. Lips. 22. 1856.
Lomaria Kze Bot. Zeit. 1848: 144.

Lomariopsis leptocarpa Holttum, Gard. Bull. S.S. 5: 270,
not of Fée.

In my paper above quoted I followed Underwood in
regarding Lomariopsis spectabilis as synonymous — with
L. leptocarpa. Fée’s figure of the latter agrees very well in
shape of pinnze with specimens of the former from Java. When
_however I examined the type of L. leptocarpa (Cuming 132), I
found it to be a Teratophyllum; it is in fact T. Williamsii Und.
A close inspection of Fée’s figure reveals the articulation of the
terminal pinna; inspection of the specimens reveals also the
great difference in texture and colour. This being the case,
another name is needed for the species of Lomariopsis, and
fortunately such a name exists in L. spectabilis.

The type of L. spectabilis is from Java. I have not seen
it, but I have seen a number of other specimens from Java, and
I think there can be no doubt of the identity with this
species of the specimens quoted in my former paper. I have
seen one more Philippine specimen (Mt. Mariveles, Bataan Prov.,
Luzon, Topping 423). This is fully pinnate, but small.

Vol. IX. (1937).

142

There are no specimens from the Philippines which equal
in size those from Java, and there is an almost complete lack of
specimens from the intervening area. The Philippine specimens
however are not sufficiently distinct to warrant the founding of
a new species. The whole question of the distinction between
species in the genus Lomariopsis is a very difficult one, and
must wait for much more evidence in the shape of collections
and field observations.

L. Brackenridgei Carr. in Seeman, Flora Vitiensis, 373. 1873.

As a synonym to this species should be added Gymnogramme
subtrifoliata Hook., formerly included by me _ under
L. oleandrifolia. I have seen the type of Hooker’s species at
Kew, and in texture and colour it clearly resembles
L. Brackenridgei rather than L. oleandrifolia, which has very
coriaceous fronds of a peculiar brown colour. The type in
question is a trifoliate frond of a young plant; its reference to
L. Brackenridgei is therefore not quite certain, but there is no
other recorded species from Fiji to which it could belong. The
name Lomariopsis subtrifoliata is already occupied for Copeland’s
Philippine species, so that there is no question of this name
replacing L. Brackenridgei, which is fortunate, as Hooker’s type
is from a very young plant, and also the validity of his original
description, with two question marks, is perhaps open to doubt.

TERATOPHYLLUM.

T. Clemensiz Holttum, Gard. Bull. S.S. 7: 262. 1934.

This species is closely allied to T. rotundifoliatum (Bonap.)
but differs in the bathyphylls, which are all simply pinnate, and
in the narrower fertile pinne (about 5 mm. wide). The
original specimens were obtained on Mt. Kinabalu, British North
Borneo. A further collection from Palawan has now to he
reported, consisting of the characteristic bathyphylls only. It is
Merrill’s No. 862 from San Antionio Bay, Feb. 1903 (Philippine
National Herbarium).

Teratophyllum luzonicum Holttum Gard. Bull. S.S. 5: 297. 1932.

This species was based on a single specimen from Luzon;
the material did not include bathyphylls. A relationship to
T. rotundifoliatum seemed probable, but was not proved owing
to the incomplete specimen. Among the further Philippine
collections from the herbaria at Manila and Washington which
I have since examined are some from the island of Palawan
which seem to be referable to this species. They consist of
bathyphylls from the youngest stage onwards to sterile
acrophylls, all associated together but without fertile fronds; the
feature in common with the Luzon plant is therefore sterile
acrophylls. It is a little uncertain in this genus to base a

Gardens Bulletin, S.S.

143

deterinination of an incompletely known species on one stage
alone, and collections of bathyphylls from the type locality of
T. luzonicum, definitely associated with typical acrophylls, are
necessary to give a final proof of identity. I think however that
the identity is extremely probable; if it fails, then the Palawan
plant must be a new species.

The youngest stages (simple fronds) are shown by Merrill’s
No. 770. The fronds are closely similar to those of
T. rotundifoliatum and T. Clemensic (see this Bulletin Vol. 5,
moon. ig. 37,-and Vol. 7, p. 263, fig. 1). The basal-lobe is
perhaps more pronounced than in the other species. No. 771
shows transition (on the Washington sheet) from the youngest
stage to small bipinnate bathyphylls; the latter are up to about
4 cm. long and 1.6 cm. wide, the pinnules obovate, to almost 2
mim. long. No. 773 has both sterile acrophylls and bathyhylls,
and in the Kew specimen both are attached to the same rhizome.
The bathyphyll on the Kew specimen is large, tripinnatifid
(almost tripinnate), the segments as in the small fronds. Fox-
worthy’s specimen has rather large bipinnate bathyphylls. The
distinction from the bathyphylls of T. aculeatum is the obovate
shape of the segments, which are rarely bifurcate.

The sterile acrophylls associated with these bathyphylls
agree well with those of the type callection of T. luzonicum, the
difference being that some of the pinne are a little larger. The
specimens are:

PALAWAN (Paragua). E-wi-ig- River, 1700 ft. Merrill 770
(W, M). zor (K, W, M), 773 (K, W, M). Rio Balsajan, 700 m.
Foxworthy 577 (M).

T. leptocarpum (Fée) Holttum comb. nov.
Lomariopsis leptocarpa Fée, Acrost. 69, t. 29. 1845.

Teratophyllum Williamsti Holtt. Gard. Bull. S.S. 5: 292,
with synonyms. 1932.

As mentioned above, I have discovered that Lomariopsis
leptocarpa Fée, represented by the type Cuming 132, is a
Teratophyllum, and is identical with the species later named
~ Stenochlena Williamsii by Underwood. Fée’s name must
therefore be used, as the oldest. The specimen at the British
Museum consists of an adult sterile frond, agreeing in all
respects with T. Williamsii. No bathyphylls were represented
in the collection, for which reason Fée included the species under
the heading ‘‘frondibus homomorphis’’. As the shape of the
pinnze is very like that of the pinnae of Lomariopsis spectabilis
the discrimination of the two species from figures is not easy.

Among the Philippine specimens which I have recently
examined are a few which show the early stages and normal
bathyphylls more completely. The youngest stage is still not

ep. AX «(1637 ).

144

represented, but there are bathyphylls of only 2.8 cm. long;
these show no signs of a dimidiate condition. The small
bathyphylls show the same character of narrowly linear ultimate
segments as the larger bathyphylls; only the very youngest
stages have rather broader segments, still with parallel sides:
Nearly all specimens, bathyphylls included, show the pronounced
red colour on drying. Additional specimens are:

LUZON. Prov. Bataan, Curran 19152 (M), C.B. Robinson
s.n. Aug. 1908 (M), P. T. Barnes 85 (M, W). Apayao subprov.
Fenix 28276 (M).

T. aculeatum (B1.) Mett.

There are a number of further collections from the
Philippines, chiefly of bathyphylls, which indicate that this
species is typically represented in the Philippines, though there
are no very young or fully dimidiate stages.

One collection (Rizal Prov., Ahern’s Collector, Forestry
Bureau No. 2695) has very remarkable large normal bathyphylls,
to 16 cm. long and 8 cm. wide. The general character of the
pinnee is similar to T. aculeatum, but there are no other speci-
mens at all comparable in size. It is possible that they
represent another species.

Gardens Bulletin, S.S.

145

AN EARLY TERTIARY RELICT IN MALAYA,
EUPHORBIA RIDLEYI nom. nov., descr.
emend. (E. SYNADENIUM RIDL.) AND
ITS POSITION IN THE
LINNEAN GENUS.

' By LEON CROIZAT,

New York.

A singular Euphorbia was presented by Ridley (Journ. Roy.
As. Soc., Str. Br., No. 61: 36, 1912) as follows :—

“Euphorbia Synadenium n. sp. A shrub about 2 feet
tall, stem smooth brownish rather succulent. Leaves alter-
nate oblanceolate spatulate subacute or blunt narrowed to
the petiole fleshy, green, nerves invisible, 4 inches long 1%

' inch wide, petiole % inch long. Inflorescence solitary or in
pairs 1 male and 1 female axillary. Male, peduncle % inch
long thick with two short ovate bracts purple pink.
Involucre as long as pink lobes 5 thick semiorbicular light
green punctate. Glands separate truncate oblong narrowed
at the base, apex laciniate pink ten. Flowers numerous.
Stamens solitary, filament thickened at the base above filiform
surrounded by a fimbriate calyx of narrow white linear lobes
about 8, anther cells globose separate. Female involucre
smaller subangled with 2 dark pink lobes much shorter not
expanded greenish yellow, glands flat incurved laciniate pale
green. Pistil solitary surrounded with pale green laciniate
calyx, overy subglobose shortly stalked. Styles three
subulate not plumed twisted together.

Penang Hill: near the Waterfall: Selangor: Langat.

In rocky woods. I have this curious plant in cultivation
in the Botanic Gardens in Singapore.

Baillon described a species of Synadenium as Euphorbia
Synadenia but this has been referred tc its genus Synadenium,
so that the above name will stand.’’

The description, no doubt, suffers on account of careless
typesetting. A carefully edited rearrangement of it (Fl. Mal.
Pen., 3: 180-181, 1924), however, is barely more intelligible to
the average taxonomist; the revised description is as follows :—

“FE. synadenium Shrub about 2 ft. tall. Branches rather
succulent, brown. Leaves alternate oblanceolate spathulate
sub-acute or blunt, fleshy dark green: nerves invisible; 4 in.
long, 1.5 in wide; petioles 5 in long. Involucres solitary
unisexual. Male on a peduncle .12 in. long with 2 ovate
purple-pink bracts; involucre pink, lobes thick semiorbicular,
green. Glands ro truncate oblong laciniate pink. Stamens

Vol. LX. (1937).

146

solitary surrounded by a calyx of about 8 white linear lobes.

Female involucre smaller with 2 pink lobes, glands flat

incurved green. Pistil surrounded by a laciniate green calyx.

Ovary subglobose stipitate. Styles 3 subulate. Hab. rocky

woods, local, Selangor, Langat (Ridley). Penang Hill near

the waterfall (Curtis, Ridley) Native name: Poko Sesudu

Bukit. I have only seen this plant about 2 ft. tall.

Mohammed Haniff reports it as growing to 6 to 8 ft.”’

In both accounts the terminology is Ridley’s own and highly
unconventional. So much, of course, does not in itself call for
comment or censure, but any one familiar with the genus is
puzzled by the mention of ro glands and of 8 lobes that occurs
in the later version. Not only in the species, as described, are the
glands twice as many as usual: the apparent quinary symmetry
of the cyathium itself is denied. The letter of the document
reveals E. Synadenium either as not less extraordinary a form
than E. pedilanthoides, or a downright pathologic case.

Before investigating the species, it is necessary to establish
its correct name according to the present Rules of botanical
nomenclature. In his original publication, Ridley practically
admitted that his name E. Synadenium (which clearly referred
to the genus Synadenium) was homonymous with E. Synadenia
Baillon. At that time, it was still customary among many
European botanists to regard such a homonym as valid, if the
earlier name, as in this case, was transferred to another genus.
By the recent change in the Rules of botanical nomenclature,
however, all later homonyms are invalid. Ridley’s name can
therefore stand only if we regard it as distinct from E. Synadenia.
This raises the nice question of what differences may be admitted
as constituting distinct names, and conversely. The Rules are
decidedly vague on this matter, but in view of Ridley’s tacit
admission, and of the fact that the two names can legitimately
be regarded as of different gender only, it appears right to regard
them as identical. A new name is therefore needed, and is
given below. It may also be mentioned that Boissier mistran-
scribed Baillon’s name as E. Synadenium in the synonymy of
Synadenium arborescens (D.C. XV: 1269, 1866); but by Art. 40
of the Rules such a synonym has no validity as a published name.
The loss of the name Synadenium for the present species is not
to be regretted, as the ‘‘glands’’ are far from connate.

The writer stands under deep obligation to Mr. Holttum who
kindly consented to send on loan the material preserved in the
herbarium of the Botanic Gardens of Singapore (No. I1I99,
VII. 13. 1916, leg. Mohammed Haniff; s.n., III. 6. 1926, leg.
E. Seimund) and arranged for the transmission of live rooted
cuttings and detached cyathia in fluid. The description to follow,
unfortunately, is still short of the data from the 2 cyathium,
although a nearly ripe capsule is recorded.

Gardens Bulletin, S.S.

147

E. Ridleyi nom. nov., descr. emend.—Leaves alternate,
crowded at the tip of carnose slender twigs, glabrous, up to 21
em. long, 7 cm. broad; blade oblanceolate to subelliptic, very
entire, occasionally sublobate (1 lobule seen on 1 leaf), more or
less deeply notched at the rounded apex, when dried thickened at
the margin and smooth or obscurely and irregularly veined
beneath, mostly tapering from the upper third or fourth to a
distinct roundish subwinged petiole up to 3% cm. long; midrib
conspicuous beneath canaliculate above ending in a short stiff
deciduous mucro; leaf-stipules 2, minute carnose, irregularly
conic, grooved, persisting or not at the slightly upraised leaf-
scars. Inflorescence strictly axillary of 1-2 (and more ?) variously
arranged cymes, mostly bistipulate at base, up to 2.5 cm. long;
cyathia unisexual, clustering, on monocephalous short peduncles
about 2 mm. long, or single on a peduncle 6 mm. long extended
beyond a pair of minute bracts into a pedicel 8 mm. long, or in
pairs on a common peduncle 5 mm. long and individual pedicels
up to 7 mm. long. Cyme-bracts opposite, free or subconnate at
base, carnosule, subquadrangular to broad ovate, mostly apiculate,
to 3 mm. long. Cyathium (only o& seen in fluid and dissected)
up to II mm. wide with a very shallow involucre and manifestly
deflected (at ca. 45°) nectaria [glands], these about as broad as
the involucre itself is deep; cyathium-bracts 2-3 spirally ascending
or subopposite, clasping, occasionally exceeding the cyathium,
carnose, broad ovate to round triangular, apiculate or entire at
apex, keeled or rounded on back, up to 4 mm. long and broad;
nectaria [glands] 4—5, contiguous, quadrangular to elliptic,
rugulose pitted, shallowly and irregularly crenate at margin,
5-6 mm. long, 3-3% mm. wide; lobes 4-5 elliptic, denticulate
3 mm. long, 2 mm. wide; co flowers [stamens] numerous in (4-)
5 fascicles; staminodes very numerous, carnosule, spatulate, laci-
niate cleft, to 4 mm. long. Capsule glabrous, smooth or rugulose,
I.3 cm. wide, 8 mm. long. exserted on a thin stipe 8 mm. long;
cocci 3, often unequally developed, discrete, the rounded keel
becoming more or less thin-acute towards the recurved end;
calyculus irregularly 3-5 lobulate, scarcely 3 mm. broad; styles
3, cleft to base, subulate, entire, spreading. Seed not seen.(')

(1) &. Rzdleyz nom. nov., descr. emend. (£. Synadenium Ridl.,
nec Boiss. (sphalm. Baill.)) Frutex diffusus 2-8 ped. altus, ramulis
succulentioribus ultimis crassitie calami scriptorii. Foliis alternis
subverticillatis carnosis glabris, ad 21 cm. longis, 7 cm. latis, apice
rotundatis, oblanceolatis subellipticisve, rarissime sublobatis, emargi-
natis; petiolo ad 33 cm. longo; stipulis petiolaribus binis subconicis
rimosis minutis deciduis_ vel ad _ folii cicatricem subelatiorem
persistentibus. Cymis axillaribus ad 2.5 cm. altis; cyathiis mono-
sexualibus pedunculatis pedicellatisve ad 11 mm. latis; nectariis 4~5
contiguis foveolatis, ad 45° deflexis, involucri ambitum aequantibus ;
lobis 4-5 ellipticis, denticulatis, 3 mm. longis, 2 mm. latis:
staminibus multis (4-) 5-fasciculatis; staminodiis plurimis; capsula
exserta stipitata ad 1.3 cm. lata; coccis 3, discretis recurvis; calyculo
aoe epulato; stylis 3, partitis, subulatis inmtergris. Semina
desiderantur.

Vol. IX. (1937).

148

Loc.: Penang Hill, near the Waterfall: Selangor, Langat
(Ridley). Pulau Rumbia (specimen). An indication that the range
might tend to become more circumscribed is apparently given in
a note from Mr. Holttum (in litt. Aug. 1935) mentioning that
the plant was not found at an easy accessible former habitat at
Penang, although it was still being reported from the south of
Penang Island.

The habit is described as follows by Mr. Holttum (in litt.,
Febr. 1936) ‘‘I recently saw an old plant of the species in Penang;
it was scrambling over a large rock in half shade and had reached
a considerable length, though the stem was still quite slender’’.
This peculiar habit is observed in several Mexican and Tropical
American species.

Plainly interpreted E. Ridleyi is a form less remarkable in
itself than Ridley’s diagnosis suggests. Truly remarkable,
however, is this species in floristic and in phytogeography.

The understanding of Euphorbia at this hour is still funda-
mentally what it was in the days of Linneus. In the opinion
of the writer the “‘genus’’ is a threadbare net of diverse evolutive
phyla artificially connected by a common type of inflorescence
(cyathium) which the Linnaean understanding consciously or
unconsciously treats as a flower. ‘The sections which attempt to
present the countless species of Euphorbia as a rational whole
use in the majority of the cases imperfect concepts, and uncriti-
cally accepted are an hindrance rather than a help to study. The
fact that N. E. Brown who enjoyed unsurpassed facilities for
studying the Linnzean genus disregards sections altogether in his
classic works may be invoked to absolve the writer from the duty
of proving his statements at this hour. It is plainly not
advisable to abound in citations of modern works which, e.g.,
introduce sect. Tenelle in the Gangetic peninsula in the wake
of E. dracuncoloides, or very nearly propose a new section on the
strength of the digitate nectaria of E. clavidigitata from Burma.
For the purpose at hand the sections of Hooker (FI. Brit. Ind. V:
245-246, 1890) are as good as any. 4

Hooker accepts (o.c., l.c.):

Sect. I.—Anisophyllum with subss. Elegantes,
Hypericifolia, Chamesycee.

Sect. II.—Euphorbium with subss. Tirucalli, Dican-
thium, Rhizanthium.

Sect. III.—Tithymalus with subss. Galarheus, Esula.
The principles of a sound classification of the Linnzean genus
were perfectly understood by Roeper (En. Euph., 1824), who

emphasized the faculty peculiarly inherent to Euphorbia to
suppress and to shorten axes (0.c., 28 et passim). It may be that

Gardens Bulletin, S.S.

149

the Enumeratio Euphorbiarum is a juvenile work weak in
taxonomic understanding: the fact stands that Roeper had a
sounder grasp of the nature of Euphorbia than Boissier and other
well-known botanists, and one may regret that Roeper failed to
present the monograph of the genus on which he was known to
work (Ch. des Moulins in notes to E. Milii, Bull. Hist. Nat. Soc.
Linn. Bordeaux 1: 27, /!1826).

Judged as natural groups in the light of Roeper’s basic.
premises Anisophyllum and Tithymalus have a value that beyond
comparison exceeds that of Euphorbium. In Anisophyllum the
main axis aborts immediately above the first pair of leaves, and,
therefore, may be technically described as indefinite. The species
of the section ate all harmonious in the sense of evolution: their
development can take place only from buds in the aborted main
axis, which simple fact accounts for their axial arrangement,
habit and interpetiolar stipules. In Tithymalus the main axis
develops and is definite, to say ends with a terminal cyathium.
The sections lacks stipules and has fundamentally alternate
leaves at the axes that are not immediately flower-bearing. Any
one who has worked with Euphorbia recognizes Anisophyllum
and Tithymalus at sight, for, it has been noticed, they are
natural groups. The subsections need not be discussed at present.

Euphorbium covers Tirucalli, Diacanthium, Rhizantium.
The last may be disregarded here. It probably includes regres-
Sive and depauperate forms, or highly specialized species from
different evolutive phyla and, appearances to the contrary, has
never been fully explored. Tirucalli has axes that are definite
precisely as are those of Tithymalus. ‘The main stem evolves
freely and the floriferous axes end with a cyathium surrounded
by a pseudo-umbel of more or less long-pedunculate and long-
pedicelled cyathia. The fact that some species of the Tirucalli
of common understanding have stipulate leaves may not be
discussed even briefly at this hour: suffice it to establish that
Tithymalus and Tirucalli, naturally interpreted, stand on the
same phylogenetic and systematic line. Diacanthium abhors
Tirucalli. Its axes are indefinite, not carrying true terminal
cymes (pseudo-umbels). Its prickles are spinescent aborted axes
prescisely matching those observed in Rosacee, Rhamnacee,
Rutacee, etc., and not relicts from a ternate foliar verticil
or from lobes and auricles at the base of the blade. In
brief: Tirucalli and Diacanthium are ‘‘classified’’ under one
section for no better reason than their being succulent. (')

Concluding: FE. Ridleyi can not be a Tithymalus, a
Diacanthium or a Tirucalli, and still less an Euphorbium which
sectio ineptissima is wholly an horticultural figment. EF. Ridleyi

(1) The issues that here must be briefly outlined in order to define

the position of £. Ridleyi will be treated in detail in a coming
publication.

Vor. IX. \(1937,)..

150

naturally falls in with certain species of sect. Goniostema, a
section which so far has not been known to exist in Asia.

The Goniostema species with which E. Ridleyi has affinity
are those that Boissier describes (D. C. Prodr. XV 2: 77, 1862)
as having stipulas indivisas (typ. E. pyrifolia). The species
which Boissier lists stipulis verticalibus cristeformibus (o.c., 78;
typ. E. leuconeura) have nothing to do with the true Goniostema,
Their affinities wholly le in the direction of the group represented
by the so-called E. splendens(?). Denis correctly shifts them
(Euph. II. Austr. Afr., 72, 1921) there, unfortunately, -however,
offsetting this pertinent change with the inclusion of
E. orthoclada under Goniostema. E. orthoclada has definite axes
and pseudoumbels, and belongs with Tithymalus.

Compared with the species of Euphorbia of the Malaysian,
Indochinese and Australasian domain E. Ridleyi is found
naturally to agree only with E. epiphylloides, an endemic of the
Amdaman Islands (v.v.) which has no place with the Tirucalli
species among which it has so far been listed. The peculiar
flattened stems of E. epiphylloides do not alter the basic
characters of inflorescence, leaf-stipule and habit, sensu lato, that
make of it an outspoken Goniostema. E. plumerioides with range
from the Philippines to Java belongs to Tithymalus. Two species
respectively from the Straits of Torres and from Queensland and
New Zealand, E. corynoclada and E. glauca abhor E. Ridleyt.
The former, as Manson Bailey appears to claim correctly (Queensl.
Fl., V: 1400, 1902) is close to E. aphylla which is a Canarian
endemic of the Tirucalli description. The latter is a strand-
species which is similar to the Euphorbia species from the
Macaronesian domain included by Boissier in Tithymalus subs.
Pachyclade.

There is a bare possibility that E. Ridleyi is allied with E.
Cleopatra (n.v.), from New Caledonia. This island, and its
archipelago, are a domain of extraordinary interest for the student
of Euphorbiacee. Regardless, however, of a possible connection
with E. Cleopatra the affinities of E. Ridleyi are in the direction
of species from Madagascar and the groups of islands in the
Indian Ocean. Thus E. Ridleyi and E. epiphylloides stand at
the eastern, E. pyrifolia at the western pillar of a collapsed
bridge : they brood over a continent that has been engulfed by the
Indian Ocean. In the truest sense of the word they are relicts,
which is it proper to designate as Gondwanic relicts.

Gondwanaland up to the beginning of Tertiary times

connected Africa with Malaysia and Australia, and sound
evidence has been unearthed to the effect that at the same time or

(1) Bojer’s binomial, ex Hooker, owes its success to the circulation
of pl. 2902, Bot. Mag. 1829, but loses precedence to E. Mila Ch. d.
Moul., Bull. Hist. Nat. Soc. Linn. Bordeaux, I: 27—30, cum ic. 1 e.
1826. Des Moulins’ binomial is far from being the nomen nudum
which it is generally supposed to be.

Gardens Bulletin, S.S.

I5I

shortly before Africa was connected with America (*). Under
the standpoint of palaeogeography a relationship between the life-
forms of Madagascar and Malaysia, at least as far east as the
Wallace’s Line, is not only posible but probable. Botanically
the evidence in favour of such connection is quite strong. Only
to mention one family, Euphorbiacew, have identic genera in
Tropical America and in Africa (cfr. Lanjouw, Euph. Surin.,
79-84, 1931). “The connection established between the Malagasy
and the Malaysian domain by the species of sect. Goniostema has
just been recorded. Merrill pointedly remarks (Ess. Geob., 247,
1936) that the flora of Polynesia is essentially Malaysian. The
same phyla have ‘‘tapered’”’ to closely allied genera in distant
continents (cfr. Richeria and Richeriella; Baccaurea and
Aporosella). The phylum of Euphorbiacee is not holartic: it
harks back, it seems, to forms the beginning of which must be
sought at the Permo-Carboniferous glaciation, and its axis of
migration is roughly parallel to the equator of to-day in a west-to-
east direction. The migration of the holartic phyla is roughly
perpendicular to the equator, and confusion need not be made
between the two main streams of diffusion of plant-life.

The persistency of isolated species at extreme points of a
geologic plant-domain is not surprising. Van Steenis remarks
(Bull. Jard. Bot. Buitenz. XIII: 393, 1935) that Malaysia from
the Cretaceous on has practically always been wholly within the
tropical zone. In contrast to the widespread assumption life does
not naturally exhibit an aptitude to change. It reacts against
the forces in the ambient, and ultimately dies or migrates or is
transformed when the pressure of the ambient becomes too strong.
It would seem, lastly, that the more a plant-group has been
capable of changing in the past the more it is apt to change in the
future. E. Ridleyi is an archetype lingering in a favoured area,
which is matched by E. epiphylloides and, if the writer be correct,
by certain genera of Malaysian ferns. It is curious to notice that
an analogy appears to exist between the rhythm of plant-life
at one hand and the laws of motion at the other.

(1) The bibliography on this subject vastly exceeds the limit of
-space allowed to these cursory notes. The reader may consult Th.
ARLDT, Handbuch der Paldogeographie, 2 vols., 1919-1922; J. W.
GREGORY, The Rift Valleys and the Geology of South Africa, 1921;
A. P. COLEMAN, Ice Ages Recent and Ancient, 1926; A. DE TOIT,
Geology of South Africa, 1926; (T. H. GOODSPEED, edit.) Essays in
Geobotany in honor of W. A. SETCHELL, 1936. A masterful study
on polar shifts is D. BLACK’s, Palaeogeography and Polar Shift, Bull.
Geol. Soc. China, X, 10931.

Vol. IX. (1937).

Cc

152

#
~ PALM MALESICZ.

VI.—Notes on Some Dzmonorops of the Section
CYMBOSPATH~.

By C. X. FurtTapo,

Botanic Gardens, Singapore.

Summary and Introduction.

Systematic Difficulties Inherent in the Species of the Section.
Good and Ample Collections are Needed.

Specimens should be Adequately Numbered.

Data to be Noted on the Field-labels.

Distribution of Sex in the Individual Plant and Spadix.
Sex of the Clump.

Is Dzemonorops Parthenogenetic ?

Systematic Notes.

eee ae

1.—Summary and Introduction.

This paper embodies some results obtained in the course of
my study of rattans belonging to the genus Demonorops of the
section CYMBOSPATH. My inquiry is based mainly on the
herbarium material, but I have also had opportunities of study-
ing some few species in the field and the results of these also
are embodied herein. Both these investigations have streng-
thened my previous conviction that much of the vagueness
hitherto observable in the specific circumscription of many
species is due to the inadequacy of the herbarium material
coupled with its faulty numbering. ‘These two aspects of the
subject I have already touched on in previous papers (cf. Gard.
Bull. S.S. VIII, 1035, pp. 241-261, and 339-367). But so
important is this matter to the systematist that I make no
apology for reverting to it here. If the systematist is to clear —
up the status of the doubtful species and to devise a simpler ©
means of classifying rattans, a large series of herbarium —
specimens, prepared according to the needs of the systematist,
becomes an indespensible preliminary, and part of this pre-
liminary is the issuing of detailed instructions to plant collectors.

The opinion that normally the genus Demonorops is —
moncecious with unisexual spadices, is herein shown to be ~
fallacious. All the specimens so far seen go to support Beccari’s
contention that the genus is dicecious, the male-like acolyte of —
the female being a neuter flower. In some species, however, —
of the section CYMBOSPATH the male plant seems to play ©
no part in the development of the fruit, the female plant itself ©
being apparently parthenogenetic. Further observations are ~
needed to verify this apparent parthogenesis in Demonorops. —

Gardens Bulletin, S.S. -

153

My inquiry into the systematics of the Malayan species of
Demonorops of the section herein treated was much simplified
by obtaining on loan authentic specimens of Griffith’s species
from the herbarium of the Royal Botanic Gardens, Calcutta, and
some critical specimens from the herbarium of the Botanical
Gardens, Buitenzorg. It is my pleasant duty here to record
my indebtedness to the respective Directors, of the two institu-
tions named, for their kindness in allowing the loan. The
results obtained may be summarised as follows :—

(1) New Species: (a) D. angustispathus and D. Curtisii
from the Malay Peninsula; and (b) D. javanicus from Java.

(2) New Records: (a) for the Malay Peninsula:
D. melanochetes; (b) for Borneo: D. hygrophilus (doubtful)
and (c) for Sumatra: D. hygrophilus, D. Lewisianus and
D. pseudosepal.

(3) New Interpretations: D. intermedius (Griff.) Mart.,
D. Lewisianus (Griff.) Mart., D. malaccensis Mart., and
D. monticolus (Griff.) Mart., all of which are indigenous
and endemic in the Malay Peninsula. ‘These interpretations
have been based on the authentic specimens, (when available
in the Calcutta herbarium), as well as on the description
and the drawings published by Griffith.

(4) New Synonyms (Species newly reduced to the
synonymy): D. aciculatus Ridl., D. bakauensis Becc.,
D. carcharodon Ridl., D. congestus Ridl., D. imbellis Becc.,
D. intermedius var. nudipes Becc., D. malaccensis Mart.,
D. microthamnus Becc., D. monticolus var. pinangianus
Becc., D. petiolaris Mart, and D. Schmidtianus Becc. The
possibility of D. fissus var. cinnamomeus Becc. being
identical with D. hygrophilus is indicated.

2.—Systematic Difficulties Inherent in the Species of the Section.

Anyone who has attempted to study the rattans of the genus
Demonorops, belonging to the section CYMBOSPATH, will

__ at once realise that, although the species can be classified readily
into definite groups, to make a clear cut delimitation between

the different members constituting each group, is a different,
and rather a difficult, matter. Beccari, in spite of his prolonged
studies, was unable to overcome this difficulty, and therefore he
ventured a suggestion, according to which each one of the
groups would be regarded as a collective species (synspecies), the
different individual members of the groups being treated as so
many sub-species. (Calc. Ann. XII, 1011, p. 28, footnote).

No doubt a chief reason why the separation of the species
is a difficult task is the very close relationship existing between
them; in many cases no conspicuous and easily describable
diagnostic characteristics distinguish one group from another.

Polk, LX. (1937).

154

But perhaps a reason of no less importance is our ignorance of
the full amount of variation in each species: for the species of
the section CYMBOSPATH vary greatly in regard to the size
of the leaflets, spathes, and spadices; and also to a certain extent
in regard to the nature and arrangement of bristles or spines on
the spathes and leaf-sheaths in different parts of the same plant,
or else at different ages, or under different ecologic conditions.
Moreover, the material available in the several herbaria of the
world is often inadequate as a source from which to draw reliable
distinctions between different but closely related species. While
such distinctions are, or at least seem, quite obvious in the field,
they cannot be verified in the herbarium on account of the

poorness of the material there available. :

3.—Good and Ample Collections are Needed.

In order to obviate in their studies complications arising
from difficulties in regard to species, systematists have been
content hitherto with the expedient of classifying species
geographically, and so, of dividing the entire Demonorops area
into several geographic sub-units. Under this expedient they
describe as new every find that does not correspond with any
species previously described in relation to its respective
geographical sub-region or sub-unit. Only when it is quite
obvious that a specimen belongs unmistakably to a quite distinct
species already defined as inhabiting another sub-region or
sub-unit will it be admitted as an old species into the region
where it is newly found. While this prodecure had the advant-
age of throwing some light on the rattan floras of the regions
studied, its legitimacy as a form of classification can scarcely be
defended, consequently it is high time that a pause should be
agreed upon to enable the conception of species to be
de-geographized. But to do this thoroughly would entail the
critical examination of a large series of material of the
‘‘synspecies’’ from the different regions.

No doubt the first disclosure would be the paucity and
insufficiency of the material represented in the herbaria of the
world. Next, it would necessitate collectors being asked
to contribute towards discovering the true species by striving
to show in their collections the extreme variations that occur
in the adult forms of the rattans. They should be encouraged
to give one number to each individual plant and to include in
the principal sets two complete specimens showing the extremes
of variability of the individual. If the collectors could be
induced to make complete specimens, as recommended in my
previous papers (Gard. Bull. S.S. VIII, 1935, p. 242, and 342),
much of the present confusion would vanish like a guilty thing
or like the furrow cut by a ship’s keel in the sea.

Gardens Bulletin, S.S.

155

4.—Specimens should be Adequately Numbered.

In this connection one cannot emphasize too strongly the
importance of numbering rattan specimens carefully in the field.
Ridley tried to study the Demonorops species from the Malay
Peninsula without numbering the specimens in the field, and
obtained but very poor results. Ridley alludes to his studies
in the field in the following remarks made in his Flora V, 1925,
Pp. 34:—

‘These plants vary much in the size of leaflets, spathe
and spadix, bristleness and arrangement of spines on the
sheaths in different parts of the same plant. Many species
have been based on totally inadequate herbarium material.
I have seen many of the species wild, often in abundance,
and cultivated many of them. Beccari’s volume Ann. Bot.
Gard. Calc. XII, with large photographs of the herbarium
specimens is invaluable, though I am by no means agreed
as to some of his determinations. Specimens are so bulky
and spiny that collectors are apt to bring home mere scraps
of the plant which are utterly unreliable. They can only
be properly studied in the forests’’.

A close examination of Ridley’s determination of the species |
of Demonorops of the Malay Peninsula (cf. my paper in Gard.
Bull. S.S. VIII, 1935, pp. 339, 367, and vide infra) reveals the
fact that the specimens, owing to Ridley’s not having numbered
them in the field, were often so mixed that Ridley himself could
not disentangle them correctly. Naturally, therefore, when
coordinating his field observations in the herbarium, Ridley
came to several erroneous or misleading conclusions; and but
for the specimens collected and numbered according to the
system introduced into Malaya by Ridley’s successor, Mr. I. H.
Burkill, as the Director of Botanic Gardens, Singapore, Ridley’s
errors in classification and determinations of the Demonorops
species would have remained beyond correction, or anyhow
extremely difficult to correct.

This miscarriage of Ridley’s long and painstaking studies
in the field, coupled with his systematic investigations in the
herbarium, should prove to botanical institutions how imperative
it is to impress upon the departmental plant-collectors, or upon
the collectors of the expeditions supported by them, the real
necessity not only for making their rattan collections as complete
as possible as directed in my previous notes (Gard. Bull. VIII,
1935 pp. 242 and 342), but also for numbering them adequately ©
in such a manner as to prevent the component parts of one from
getting irretrievably mixed with the component parts of
another. I am firmly convinced that good specimens, if
properly labelled, would also render the systematist less
dependent on field notes for their accurate determination.

Vol, UX..\(.1037).

156

5.—Data to be Noted in the Field-Labels.

It is evident, from a reading of the several floristic works,
that very little information is available about the actual habit
of many rattans. ‘The data recorded in the field-notes is likewise
so meagre that it often takes one not a step forward. After a
thorough study of all the information presently available in the
published works, and particularly those carefully noted on the
herbarium labels, our knowledge of the habit of certain rattans
is still obscure; thus it is difficult to say whether they are solitary
or tufted, whether they are long climbing, like Calamus
scipionum, attaining a length of over 150 feet, or whether they
are short, like Demonorops grandis, or erect or subscandent,
like D. Lewisianus. From the description available, for
instance, one would not imagine that the stems of D. calicarpus
are very short, being not more than 1-3 feet high and scarcely
visible to a casual observer, hidden as they are by long radical
leaves, which grow all round the bush and attain a length often
many times greater than the actual stem and its leaves together.
But if the specimen sent in by the collector, besides being
collected according to the necessities of the systematics explained
herein, were accompanied by field notes or photographs indicat-
ing the habit of the species chosen, then the systematist could
get a better conception of the species and might eventually
succeed in devising a classification that would render less
dificult the circumscription of many a puzzling species.

In the case of tufted rattans, specimens of the radical leaves
are not absolutely necessary, the adult leaves are the most
important. But if the radical leaves are taken and numbered
separately, so as to prevent them being confused with the adult
ones, but at the same time to show their relation to the latter,
they might help to bring out the characteristics of the species
more clearly.

6.—Distribution of Sex in the Individual Plant and Spadix.

There still exists considerable confusion regarding the
distribution of sex in the species. In the Materials II (1907}
pp. 173 and 174, Ridley definitely states that the spadices of
D. angustifolius are bisexual, a statement he repeats later in
Flora V (1925) >. 36. In the latter work Ridley notes that he
has not seen any specimens of D. Scortechini and D. sepal;
yet in his Key to the species (p. 35), he lists these two species
with D. angustifolius as having bisexual spadices. This is sur-
prising, seeing that Beccari, the author of the two namings, has
definitely stated the opposite regarding the sexes of the spadices
in the genus Demonorops; nor has any one else described the
species at length; it must therefore be regarded as gratuitous
that Ridley should re-define the sex without giving any support-

Gardens Bulletin, S.S.

157

ing observation. Beccari’s remarks on the question seem worth
quoting here:

‘As in the Calami, the spadices of Demonorops are
always dioecious, though one may sometimes happen to meet
with a male spadix with a hermaphrodite flower here and
there. It is possible that the female spadices of some
CYMBOSPATHAE are cleistogamous—at least, I suppose
they may be having observed that certain still closed
spadices of D. fissus contained ovaries in process of develop-
ment, which had every appearance of having been
fecundated. The closure may however have been more
apparent than real, and thus an insect may have been able
to penetrate into the interior through the half closed
Spaunes.2. (Calc, Anim. TOLI,'.p. 2).

Regarding Ridley’s views on the subject, Beccari comments
as follows :-—

“Mr. H. N. Ridley (Mat. Fl. Malay. Pen. II, 1907
p. 171) says that the spadices of the Demonorops are
usually unisexual, and that male and female inflorescences
are on the same stem——a fact which I have never observed.
The same author (1. c. p. 172) adds that both sexes occur
on the same plant: furthermore, he describes the spadix
of D. angustifolius as having a ‘bisexual spadix, male and
female flowers on the same spadix in pairs.’ Apparently
Mr. Ridley has considered as a fertile male flower the neutral
or sterile flower which accompanies every female flower on
the female spadix in all the species of Deemonorops I have
examined.’’

After examining several Demonorops of the section
CYMBOSPATHAE cultivated in the Singapore and Buitenzorg
Gardens, I can testify to the authenticity of Beccari’s observations
on the distribution of sexes in the individual spadices and plants
of the genus Demonorops, even of D. angustifolius, in which
species Ridley appears to be quite positive of the bisexuality of
the spadices. No doubt Ridley’s collections in the Singapore
Herbarium often show male and female spadices on the same
sheet, or on different sheets, but bearing the same number, as
if the specimens were collected from the same plant or clump.
But still there are strong objections to basing conclusions upon
such an evidence. Ridley numbered his specimens, not in the
field, but in the herbarium and after study; and there are many
instances to show that he did not hesitate to give one and the
same number to specimens that, although obviously not derived
from the same clump, appeared to him to represent the same
species (cf. my remarks on D. angustifolius and D. grandis).
As to his observations in the field, besides mistaking a neutral
flower for a male in D. angustifolius, it has to be noted that he
has sometimes taken a young female spadix for a male (vide

Vel. 12211037):

158

my remarks on D. setigerus in Gard. Bull. $.S. VIII, 1935
p. 363); and from such erroneous observations, he has concluded
that the genus Demonorops is normally monoecious, male and
female flowers being found in different spadices (Materials, II
1907, pp. 171 and 172), and in his Flora (V, 1925, p. 34) Ridley
has persisted in his opinion despite Beccari’s adverse criticism.

It is true that, as pointed out by Beccari (op. cit. XI, 1908,
p. 18; XII, rors, p. 2), a rattan female spadix might occasionally
bear, at its apex, a few spikelets composed entirely of male
flowers (e.g., Calamus perakensis, not observed in Demonorops),
or that the hermaphrodite flowers might be found here and there
on a male spadix. But this is not the usual phenomenon either
in Demonorops or in Calamus; therefore one cannot rightly
regard it as establishing that either of these genera is normally
monoecious with unisexual spadices.

7.—Sex of the Clump.

Though observations so far made show clearly that
Demonorops plants bear male and female spadices in different
individuals, it is not yet very clear whether the individual stems
in a clump are all alike in sex. I have seen several live clumps
of Demonorops, and, as far as it could be ascertained, the clumps
were always of one sex; that is, if one individual was found to
be male, then all the other individuals in the same clump were
noticed to bear male spadices; the same was true for the female
sex. But in one single clump of D. fissus, growing in the
Singapore Botanic Gardens, I found male and female plants (male
in the majority) in the same clump. Now the question arises
whether this clump with male and female individuals, is a result
of one or two seedlings being planted in the place. Our records
do not throw any light on the matter; but further observations
in the forests might enable one to say definitely whether a male
clump will occasionally produce female individuals, or vice-versa.

8.—Is Dzmonorops Parthenogenetic?

Though Demonorops species are normally dioecious, it does
not follow that they are dependent invariably on the male plants
for the production of the fruit. In some species of Demonorops
of the section CYMBOSPATHZ4 the female flowers attain their
full maturity and develop into fruits when the spadix is still
enclosed in the spathes. Hardly any pollinating agent can have
access to the flowers shut within the spathes, and excepting the
ants, which seem to have a special predilection to all kinds of
rattans, and which possibly are the only visitors that might gain
admission to the flowers through the enclosing spathes, one
rarely, if ever, sees any insect or bird visiting the female spadices.
For carrying pollen for long distances, however, the ant is an

Gardens Bulletin, S.S.

159

almost negligible agent. Unless we assume, then, that the
fruits develop parthogenetically—a fact that would mean the
male plants are useless,—it is difficult to explain the development
of good fruits in such species as D. angustifolius, D. grandis,
and D. fissus (species observed by me) when there exists no male
plant of any Demonorops species within a radius of about 300
feet, or, may be, considerably more.

In this connection it is to be observed that the spadices
in some cases show two kinds of female flowers, one with ovaries
fully expanded before the anthesis of the inflorescence, and the
other not so expanded. Normally only the expanded flowers
develop into fruit. But may it not be that the development of
the other flowers are dependent upon their receiving pollen from
male plants? This explanation, which might account for the
existence of the male plants in Demonorops as well as for the
two kinds of female flowers on a spadix, is suggested here with
a view to focussing attention on this problem and to encourage
field observations.

On the other hand it is possible that, during the early stages
of the development of the spadix, the smaller flowers that later
fail to develop, are merely those that lag behind in the compett-
tion between the flowers for food, and for space in the spadix.

It may be observed that in the Singapore Botanical Gardens
there is a female clump of D. Kurzianus Becc. ex descriptione
which always produces female individuals, and, though fruits
develop to a large adult size, seeds within the scale-covering
remain always abortive. ‘The plant is a native of the Andamans,
and, as this group of islands have a somewhat different climate
than that of the Singapore Garden, it is possible that this lack
of development of good-seeded fruits is not attributable to the
absence of male plants of the species in the Singapore Gardens :
A species that in its native habitat is parthenogenetic, may
refuse to produce, in a different climate, any seeds even when
pollinated.

. All this will show how sparse is our knowledge about certain

biological problems concerning the sex in Demonorops and will
indicate the necessity of utilizing every available opportunity
of making field observations on the subject.

9.—Systematic Notes.

IP Bes on angustifolius (Griff.) Mart., Hist. Nat. Palm.
hl -( (1850) 229; Beec.im Hook. f., Fl. Brits Ind. VI (1393)
464, in Rec. Bot. Surv. Ind. II (1902) 219, et in Calc. Ann.
XIT (1911) 69 t. 18; Ridl. in Journ. Roy. Asiat. Soc. Straits
r33 (1900) 175 et 57 (1911) 120, Mat. Fl. Malay Pen. II
(1907) 174, et Fl. Mal. Pen. V (1925) 36, omnino pro parte.

Mot. PX 901937 ).

160

Calamus angustifolius Griff. in Cale. Journ. Nat. Hist. (1844)
89, et Palms Brit. Ind. (1850) 95 tt. 213 A and B (Type locality =
Malacca).

D. carcharodon Ridl., Mat. Fl. Mal. Pen. II (1907) 178 et.
Fl. Mal. Pen. V (1925) 4o.

fie hygrophilus Mart. sensu Ridl. in Journ. Roy. Asiat. Soc.
Str. Br. 33 (1900) 175, Mat. Fl. Mal. Pen. IT (1907) 178, et F1.
Mal. Pen. V (1925) 40, pro parte. syn. nov.

MaLay PENINSULA: Pahang, Sungei Bera near Tasek Bera
(Henderson, 24,620). Perak, Temango (Ridley, 14,719, female).
Malacca, loc. incert. (Griffith, [vern. name Rotan Ghitah] in
Cale. Herb. Holotype). Selangor, Batang, Berjuntai (Ridley,
7880); Batu Tiga (Ridley 11,983*). Johore, Mawai Jemaluang
Road (Corner, 29,749, vern. name Rotan Manau); Sungei
Semangat in Muar (Fox, 11,299); Danan, along Sungei Sedili
(Corner, on 27—III-32, veru. name Rotan Tawar). Singapore,
Bukit Panjang (Ridley in 1907; and n. 13,302); Choa Chu Kang
(Ridley, in 1896*); Bukit Timah (Ridley in 1900); Bukit Mandai
(Ridley in 1900); Selitar (Ridley, 6279*); Chan Chu Kang
(Ridley, 4622, vern. name: Rotan Sepat).

Griffth’s type specimens in the Calcutta herbarium consist
of the terminal portion of a leaf and the portion of a male spadix,
figured in Griffith’s plate 213-A, as well as of the petiole with
its sheath figured in the plate 213-8. The unopened inflore-
scence and the detached spathes figured in the second plate are
not represented in the specimen; judging by the description
Martius gives of the species they are to be found perhaps in
the Munich herbarium. Certainly they are not in Kew. A
comparison of the authentic material in the Calcutta herbarium,
and of Griffith’s plates, with the Peninsular material from the
Singapore herbarium leads me to concur fully with Beccari’s
interpretation of D. angustifolius. ‘The prickles on the leaf-
sheath are rather short in the authentic specimen, but this is no
doubt due to the young age of the specimen.

Ridley was much confused over this species. Apparently
he took as equivalent all the specimens bearing his field number
5123, though he had given this number to specimens collected
from more than one clump, growing in the Singapore Botanical
Gardens, and belonging to more than one species (e.g.
D. angustifolius, and D. grandis). Further, from some in-
sufficient data he came to the erroneous conclusion that spadices
in D. angustifolius are always bisexual. It is not surprising
therefore, that, with such a mixed herbarium material to go by
and with a mistaken conception concerning the sex of the
spadices of the species, Ridley not only erected the new species
D. carcharodon to include the specimens that did not fit in with

* Syntypes of D. carcharodon Ridl.

Gardens Bulletin, S.S.

161

his acceptation of D. angustifolius (Materials II, 1907, p. 178),
but later (Flora V, 1925, p. 40) dismissed as erroneous Beccari’s
placing of the new species among the synonyms of D. angusti-
folius (Calc. Ann. XII, ro11, p. 69).

Ridley’s specimen no. 5123 that Beccari quoted as having
been collected from a wild plant at ‘‘Lawas’’ in Singapore, is
derived from a plant cultivated in the Botanic Gardens, Singapore,
the word ‘‘Lawas’”’ being a result of misreading of the word
Lawns, the garden being divided into a number of sections called
lawns. Ridley’s no. 11,983 from Selangor—a syntype of
D. carcharodon—is mounted on two sheets and consists of a
mixture of three species, as follows: leaflets of Calamus
scipionum ; fruits and fruiting spadix of D. grandis; and a leaf-
sheath, a portion of leaf-lamina and an unopened, male spadix
of D. angustifolius.

The Peninsular material varies a good deal regarding the
armature of the petiole, leaf-sheath, the leaf-rachis, and second
spathe, so that it might be possible to reduce D. trichrous Miq. as
a synonym of this.

II. Dzmonorops angustispathus Furtado spec. nov.

D. grandis Mart. sensu Ridl., Mat. Fl. Mal. Pen. II (1907)
177 et Fl. Mal. Pen. V (1925) 38 quoad specimen Ridleyanum
infra citatum. Syn. nov.

D. Lewisianus Mart., Hist. Nat. Palm. III (1850) 327;
Becc. in Cale. Ann. XII (1911) 87 t. 30: Omnino quoad specimen
Gaudichaudianum in herb. Parisiensi. Syn. nov.

A D. grandi, cui affinis est, inflorescentiis spathisque
angustioribus, ante anthesin lineario-lanceolatis; spathee
extimee rostro multo longiore quam corpore; frondium
segmentis angustioribus hzec species facile distingui potest.

Caudex, vagina. petiolusque ignoti. Lamina frondium,
secundum partem visam, magna, rhacheos dorso inermis, vel
aculeis solitariis tenuibus brevibus distantibus armata. Cirrhum
non visum. Segmenta plura, subequidistantia, 4-5 cm inter se
remota, lineario-lanceolata vel ensiformia, maxima latitudine ad
medium vel infra sita, 45 cm longa, 15-18 mm lata, apicem
versus valde angustata, acuminatissima; supra secus tres costas
setulosa, vel interdum in costa media inermia vel apicem ejusdem
versus tantum armata; subtus ad costam mediam tantum setulis
approximatis predita. Inflorescentia faminea sessilis, cum
tostro lineario-lanceolato 25 cm. superante 45 cm. longa. Spathe
persistentes: extime corpus 18-20 cm longum, bicarinatum,
(tractu intercarinali ad 1 cm. usque lato), deciduo fusco-
furfuraceum, aculeis paucis, distantibus, solitariis, interdum
confluentibus, laminaribus vel subcriniformibus, nigritellis, ad
2.5 cm. longis armatum; rostrum clausum permanens,

Vol. IX. (1937).

162

eodemmodo armatum, 25 cm. superans, 5 mm. latum: secunda
(spatha) et tertia paulo latiores aculeis minoribus, solitariis,
secus carinas obsoletas apicem versus tantum parcissime armate :
interne inermes. Rami spadicum primarii 4, porrecti, ad 6 cm.
longi, alternati, 5-6 cm. inter se dissiti; secundarii ad 2.5 cm.
-usque longi, axi rigido, crasso, flexuoso, floribus utrinsecus
2-4. Involucrophorum crassum, paulo’ exsertum, apice
crateroideum. Involucrum cupuliforme vel _ pateriforme,
I-—2 mm. longum; floris aureola neutrius conspicua.
Perianthium fructiferum explanatum, tepalis interioribus sub-
duplo longioribus quam exterioribus. Fructus immaturi tantum
visi, ovato-elliptici, cum perianthio rostroque 18 mm. longi,
I0O-I2 mm. in diam. Squame brunneo-graminez, in 15 ortho-
stiches imbricatee, in linea infra marginali atriores.

_Maray PeninsurA: Province Wellesley, Bukit Juru
(Ridley, 7002. Holotype in Singapore). Penang, loc. incert.
(Gaudichaud, in the Paris Herbarium).

I have not seen the actual specimen collected by Gaudichaud
but only its photographic plate given by Beccari (plate 30 in
the Calc. Ann. XII, 1911). Apparently Beccari mistook the
outermost spathe for one of the internal and so he remarked
that the specimen of Gaudichaud in the Paris Herbarium as well
as its duplicates in the Geneva and the Florence herbaria bore
male spadices without their outermost spathes; and yet the outer-
most spathe is represented in Beccari’s plate of Gaudichaud’s
specimen. The body of this spathe measures only 18 cm. in
length, while the beak, which is placed vertically on the extreme
left of the plate; is more than 35 cm. long.

Were it not for the narrow leaflets, the long beaked inflore-
scence and the narrow spathes, D. angustispathus would have
been extremely difficult to distinguish from D. grandis with
which the species was confused by Ridley. lLeaf-sheaths are
wanted to bring out further differences.

IiI. Dzemonorops calicarpus (Griff.) Mart., Hist. Nat. Palm.
ILL: (2850)-\.326 't. 275)fpowi., Bece: mehiooke Tir agen
Ind. VI (1893) 466; Ridl., Mat. Fl. Mal. Pen. II (1907)
174 pro! parte; Bece..,imiCale: » Annz eR (aor) posi
Ridl., Fl. Malay Pen. V (1925) 37 pro parte (ex altera
parte=D. intermedius, D. Lewisianus, D. sepal, and
D. tabacinus).

Calamus calicarpus Griff. in Cale. Journ. Nat. Hist. V
(1844) 92, et Palms Brit. Ind’ (7850) Oo tt, 215-45, ana
et t. 216-A f. V and VI. (Type locality = Malacca).

MALAY PENINSULA: Kemaman, Bukit Kajang, near Ulu
Bendong, alt. 500 ft. (Corner, 30,077; 30,389). Perak, Bujong
Malacca (Curtis, 3160); Tapak (Ridley, 14,115). Dindings,

Gardens Bulletin, S.S.

163

Lumut (Ridley, 7899; Curtis 3444, female). Malacca, Chinana
Puteh (Alvins, 979, vern. name Rotan Chucho Miniak); Bukit
Sadanan (Pun Bebas=Derry’s collector, 112, vern. name Rotan
Sabut). Selangor, Kuala Lumpur (Ridley in 1899); Dusun Tua
(Ridley, 7878).

IV. Dzemonorops congestus Ridl., Mat. Fl. Malay Pen. II
(1907) 178 et Fl. Mal. Pen. V (1925) 4o.

The type of this species is a specimen collected by Ridley
in Singapore in a place that was called the Stagmount by the
English, and Bukit Panjang by the Malays. The specimen does
not bear any date or number. After examining a duplicate of
the type Beccari reduced the species to D. leptopus, making at
the same time the following remarks on its status: ‘‘I have
reduced D. congestus of Ridley to D. leptopus after a very
careful analysis of a specimen kindly sent to me by its author.
It is true that this specimen (which is very incomplete, and has
very immature fruits) has spikelets with the flowers more
approximate than usual; but this specimen does not appear to
ime quite normal’ (cf. Calc. Ann. XII, 1911, pp. 128, 131).

As Ridley in his later work did not agree with this reduction
made by Beccari, and moreover he insisted in placing the species
in section CYMBOSPATH (D. leptopus belongs to the section
PIPTOSPATH®) I re-examined the type of D. congestus,
which is in the Singapore herbarium, and as a result I fully
concur with Beccari’s remarks on the status of the species.
Apparently Ridley relied for his information of the specific
description more on his fieldnotes than on the actual type
specimen, and so in his Flora V, 1925 p. 40, he repeated the
erroneous statement originally made in the Materials II, 1008,
p. 179 even as to the length of the spadix. A fragment of the
peduncle of the spadix in the type specimen measures more than
4 inches. This fact and the length of the spikelets or of the
portions of partial inflorescences (primary branches) show that
the spadix is very long, belonging to the section PIPTOS-
PATH. Ridley’s original statement that the spadix is only
2 inches long was perhaps due to a lapsus calami in his field-
notes for 2 feet long—a mistake that would have been easily
detected had Ridley, instead of relying on his notes, re-examined
the type specimen before preparing the specific description for
his Flora.

Two more specimens collected by Ridley on the Stagmount
are likewise labelled D. congestus by Ridley himself. But,
though they both belong to CYMBOSPATH, it is obvious
that they did not form the basis of D. congestus, for they not
only do not agree with Ridley’s description of the species, but
they were collected only in 1908, a year after the original

Mor. 1X. (1637).

164

publication of the species. These latter specimens, (numbered
13,302), are D. angustifolius as interpreted herein.

V. ‘Dzmonorops Curtisii Furtado spec. nov.

A D. Lewisianus, cui valde affinis, vagina frondium
inerme, haud geniculata; spathis inermibus hzec species bene
distincta.

Caudex gracilis, cum vagina 8 mm. in diam., erectus vel semi-
scandens. Frondium vagina inermis, haud gibbosa, pro maxima
parte atro-ferrugineo leprosa, apice oblique truncata. Ochrea
inermis, ad 5 mm. longa. Petiolus circ. 15 cm. longus, dorso
convexus, supra concavusculo-applanatus, inermis vel secus
margines dorsumque spinis perpaucis, solitariis, brevissimis vel
obscuris remote armatus. Lamina 25-30 cm. longa, secus
rhacheos dorsum unguibus indivisis, reduncis, brevibus preedita,
in cirrhum ad ro cm. longum unguiculatum transiens. Segmenta
circ. 12 cm. longa, r cm. lata, ensiformia, apicem quam basin
versus magis attenuata; supra secus costam medianam et
interdum duas costas subprimarias, subtus secus costam mediam
tantum, setulosa; ad margines eodemmodo ciliolata. Spadix ante
anthesin elongato-fusiformis, cum rostro circ. 12-14 cm. longus;
spatha extima inermis, dorso superficialiter bicarinata, in rostrum
circiter corpore ejusdem equilongum contracta. Perianthium
fructiferum explanatum. Fructus immaturi tantum visi, ovati,
apicem versus angustati, circ. 12-13 mm. longi, 8-10 mm. in
diam. Squame in orthostichos 15 disposite, in medio sulcate,
basi stramineze, apicem versus rebescentes, linea marginali
fuscente.

Matay PENINSULA: Province Wellesley, Tasek Glugor
(Curtis in April 1902. Type in Singapore).

Obviously an ally of D. Lewisianus from which it is easily
distinguished by unarmed leaf-sheath, non-gibbous petiole, and
unarmed spathes. The two last mentioned characters are some-
times found in the spathes of D. Lewisianus produced in the
‘‘senile’’ stages, but even then one can always detect some
spines or bristles on the leaf-sheaths and very often on the
outermost spathes. Curtis’s specimen is evidently taken from
the terminal portions of an old stem; because owing to the
disarrangement in the rate of growth of the internodes and leaf-
sheaths, the spadices appear as if they have originated in the
axil of the leaf immediately below it. The basal portion of
the spadices being thus caught up in pseudo-axils of the leaves,
the outermost spathes do not fall off even after they are detached
from the inflorescence axis, thereby making the spathes appear
as persistent. Such a condition is sometimes obtained in many
rattans in their ‘‘senile’’ stages, that is, stages immediately
preceding their death. :

VI. ‘Dzmonorops grandis (Griff.) Mart., Hist. Nat. Palm. III

(1850) 327 t. 175 f. 9 et t. Z-xii f. 11; Becc. in Hook: f., F.

Brit. Ind. VI (1893) 463, in Rec. Bot. Surv. Ind. II (1902)

Gardens Bulletin, S.S.

=

165

219 etin, (Caic.sAnn.: XE (r91r)''58t. 11; Ridl;;: Mat. F1.
Mal. Pen. II (1907) 177 et Fl. Mal. Pen. V (1925) 38.
pro parte.

Calamus grandis Griff. in Calc. Journ. Nat. Hist. V (1844)
eperst aime Orit. Ind/(1850) or tt. 2t0 A,B, and C, et t.
aio tf, 3.

D. angustifolius Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 174 et Fl. Mal. Pen. V (1925) 36 pro parte. Syn. nov.

D. hygrophilus Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907)178 et Fl. Mal. Pen V (1925) 40 p.p. Syn. nov.

D. intermedius Mart. sensu Becc. in Hook. f., Fl. Brit. Ind.
VI (1893) 464 et in Calc. Ann. XII (1911) 73 t. 20 pro. maj.
parte and excl. spec. Griffithianum. Syn. nov.

if D. intermedius var. nudinervis Becc. in Rec. Bot. Surv. Ind.
i. (1902) 220 nomen,nud., et in Calc, Ann. XII (1911) 75--tt.
21 et 22. Syn. nov.

D. Kirtong Griff., Palms Brit. Ind. (1850) 102 t. 216 B.
Syn. nov.

D. malaccensis Mart. tom. cit (1850) 327; Bec. in Rec. Bot.
Surv. Ind. IF (1902) 222 et in Cale. Ann. XII (1911) 61 quoad t.
216 B. syn. nov.

D. malaccensis Mart. tom. cit (1850) 327, Bec. in Rec.
Bot. Surv. Ind. II (1902) 222 et in Cale. Ann. XII (1911)
61 quoad t. 216 B. Syn. nov.

¥.Calamus acanthopis Griff., Palms Brit. Ind. (1850) t. 216 B.
Syn. nov.

Martay PENINSULA: Kedah, Kedah Peak=Gunong Jerai
(Ridley in 1893), Province Wellesley, Tasek Glugor (Burkill,
6536). Pahang, T'asek Bera (Henderson, 19484, vern. name Rotan
Relang. and Sakai name=Chongkak). Negri Sembilan, Bukit
Sutu in Sungei Ujong district (Alvins, 1878, vern. name Rotang
Sunang); Senaling Inas (Holttum 9778, vern. name Rotang
Kerotong); Tampin (Burkill; 1372; 2551). Selangor, Pataling
(Ridley on 27-vi-1889) the spathes only. Singapore, Bukit
Arang (Goodenough, 1660); Bukit Mandai (Ridley in 1890);
Changi (Mat in March 1894); Bidadari (Ridley 8957); Ang Mo
Kio (Ridley 6672); Bukit Timah (Goodenough in 1892, vern.
name Rotang Chinchin; Ridley in Feb. 1900; in 1890; 1892;
and in 1898); North Selitar (Ridley, 3506 and 3499); Selitar
(Goodenough in 1890); Ridley, 3498); Gardens’ Jungle (Ridley
5122; 5123* and 5124). Malacca: Chinana Puteh (Alvins, 890,
vern. name Rotang Sunang); Bukit Sadanen (Pun Bebas=
Derry’s collector, 110, vern. n. Rotan Chichik Landak; ibid.,
108, vern. n. Rotang Cheranang).

* This number is also given to specimens of D. angustifolius.

Mor. 1M. { 1937):

166

Calamus acanthopis and Demonorops malaccensis are exact
synonyms of D. kertong, all the three names being based on
the same material referred to as Rotang Kertong by Griffith in
a foot-note under C. petiolaris var. B. in Calc. Journ. Nat. Hist.
V (1844) 94. As Griffith had considered it to represent a new
species but had supplied no botanical name, Martius supplied
the name in 1850 by calling it D. malaccensis, Griffitth’s names
Calamus acanthopis and Demonorops kertong being published
in a posthumous work in the same year. The type material
consisted of two spadices |though a portion of radical leaf is
also represented in the figure given in Griffith’s posthumous
work, Palms Brit. Ind. (1850)]. As neither of these spadices
nor the leaf portion is in the Calcutta herbarium, and as Beccari,
who had examined Griffith’s specimens preserved in the Kew
herbarium, could not see the type specimens, and as Martius
noted that he had seen the spadices, it is to be presumed that
the spadices are in Martius’s herbarium in Munich. From the
drawings given by Griffith, it is evident that the two spadices
represent D. grandis, each having lost its outermost spathe.
The vernacular name Rotang Kirtong with its variants
is still being used by the Peninsula Malays to denote D. grandis,
though the name is sometimes used for Calamus scipionum or
for an allied species or for a species of Plectocomiopsis, which,
owing perhaps to its tufted growth and broad and distant
leaflets, are classed by the Malays with the former. The
specimens that Beccari identified as D. malaccensis do not
belong here (cf. my remarks on D. melanochetes).

Ridley gave the number 5123 to specimens taken from more
than one clump growing in the Singapore Botanic Gardens.
Since a specimen bearing this number was determined hy
Beccari as D. angustifolius (later figured in the Plate 18 of the
Calcutta Annals XII, 1911), Ridley concluded that all his
specimens bearing the same number were D. angustifolius,
though a greater part of them are D. grandis. I think this
faulty numbering has been one of the main causes why Ridley
was much confused over D. grandis and D. angustifolius (see
my remarks on the latter species).

Ridley has quoted the Pataling specimen (Selangor) and
Pun Bebas (=Derry’s Collector) n. 110 (from Malacca) under
D. hygrophilus. But with the exception of the leaf on the
former, which by reason of its being from a radical leaf, is not
identifiable, both the specimens consist of unopened spadices
belonging to D. grandis. On Pun Bebas specimen n. 108,
Rotan Cheranang, is recorded as its vernacular name, but
obviously through a mistake for the name Cheranang or Jernang
(or Jeranang), is invariably applied by Malays to a Demonorops
species yielding dragon-blood.

Gardens Bulletin, S.S.

—————<—<<_<_ rr.

167

In the Materials II, 1907, p. 178, Ridley has left the
following remarks on D. grandis: ‘‘I am quite unable to
distinguish D. intermedius Mart. (Calamus intermedius Griff.),
though Beccari keeps it separate and describes [ ?] a variety
D. intermedius var nudinervis. Griffith, the original describer
of it, was dubious [ ?] as to its being distinct, and the two very
slightly different forms pass readily into each other. Both grow
side by side in the gardens and garden jungle, and as usual in
this group there is a great variation in the size of spathe, dimen-
sions of spines, and bristleness of the leaves, according to the
age of the plant and its habitat’’.

These remarks of Ridley are certainly applicable to
Beccari’s interpretation of Calamus intermedius Griff., but not
to this speeies in its original sense. Griffith’s species is allied
to D. Lewisianus, having much narrower and closer leaflets than
in D. grandis. (cf. my remarks on D. intermedius). Another
important character that distinguishes D. intermedius from
D. grandis, and which was also noted by Griffith, lies ‘‘in the
armature of the spathes, the thorns of which are very slender,
more deflexed, and much more numerous’. Beccari’s
D. intermedius var. nudinervis was included in Griffith’s
D. grandis as the leaflets of the latter were described as being
‘‘without setee on either surface or along the margins, or with
the midvein setigerous on both surfaces, as well as on the
margins.’’

Griffith’s plate 211-A of D. intermedius is also very clear
and cannot be easily mistaken for D. grandis. It approaches
some forms of D. angustifolius with rather distant leaflets and
narrower thorns on the’ spathes. (cf. my remarks on
D. intermedius).

Ridley gives Rotan Sumang as a vernacular for the species,
but ae spells the name as Rotang Sunang.

VII. Dzmonorops hygrophilus Griff. ex Mart., Hist. Nat.
a Falm II) (x850), 204 et 328, t: 177 f. II; Becc. in Hook. f.,
Fl. Brit. Ind. VI (1893) 464; Ridl., Mat. Fl. Mad. Pen. fI
(1907) 178 quoad specimen Griffthianum tantum; Becc. in
Calc. Ann. XII (1911) 62 t. 13; Ridl., Fl: Mal. Pen. V (1925)
40 pro parte.
| Calamus hygrophilus Griff., Palms Brit. Ind. (1850) 96 t.
213-C. (Basinym) Malacca.
. angustifolius Mart. sensu Ridl. in Journ. Roy. Asiat.
Soc. Str. Br. 57. (1910) 110 pro parte. Syn. nov.
2D. fissus var. cinnamomeus Becc. in Cale. Ann. XII (rortr)
a7. t.-16.
MarAy PENINSULA: Perak, R. Temango (Ridley, 14,7109,
male only; female=D. angustifolius). Kemaman, Ulu Kajang,

Mal, 1X2 -(1037)}.

168

alt. 500 ft. (Corner, 30,447, vern. name Rotan Tawar). Selangor,
Telok Forest Reserve (Burkill, 6520). Malacca, loc. incert.
(Fernandez, vern. name Rotan Ayer. ‘Type; not seen). Johore,
Kota Tinggi, by Sungei Johol (Corner, 29,750). Singapore,
Bukit Panjang, at Stagmount (Ridley, in 1907).

SuMATRA: Palembang, Lematang Ulu, alt. circ. 500
(Grashoff, 196=Herb. Bog. n. 404, vern. name Uwi Getah);
between Banjuasin and Kubus, alt. 60 feet (Grashoff, 639, vern.
name Rotan Kelemunting) (both specimens from herb. bogor.).

BORNEO; Pontianak (Heyne, 2548= Herb. Borgor. n. 222, in
herb. Bogor.). Sarawak, Bau (Ridley 11,826).

The species, which had up till now been regarded as endemic
in the Peninsula, was not clearly defined, and its*status had
therefore been rather doubtful. I have not seen Scortechini’s
specimen cited by Beccari, but from its plate, Beccari’s identi-
fication appears to be correct. The type is not in the Calcutta
herbarium, but might be found in the Munich herbarium, where
Griffith had sent the material for Martius’s study. But the
two plates by Martius, and by Griffith, bring out clearly the
characters in the petiole that distinguish the species from
D. angustifolius. viz., Geniculum unarmed, or armed only
along the ridge with a few, small, rigid prickles; and the petiole
armed along the margins, and occasionally with a few solitary
prickles along the dorsum. In addition the spathes and sheath
are usually blackish with crustaceous indumentum. In
D. angustifolius, the geniculum and both the surfaces (occasio-
nally one) of the petioles are armed with scattered rigid prickles.
The spines on the spathes and also on the leafsheaths of
D. hygrophilus are short and narrow, and scattered; not
broad and long, often laciniate and very close, as in
D. angustifolius.

The Singapore and the Johore material is fruiting and shows
that the scales are arranged in 16-18 vertical series. "The
mature fruits (in the Singapore specimen) are ovoid-globose,
depressed at base, shortly beaked at apex, 14-15 mm. in length
(excluding the beak), 15-17 mm. in diam. ‘They resemble very
much the fruits of D. angustifolius but are lighter coloured.

The Sumatran material cited above, which was obtained
from the Buitenzorg herbarium on loan, has been named by
Beccari as D. trichrous Miq. The Bornean ‘material, which
appear to be D. fissus var. cinnamomeus may belong here.

VIII. Dzmonorops intermedius (Griff.) Mart., Hist. Nat.
Palm. III (1850) 327 t. 175 f. 8; Bece. in Hook: £., Fl. Brae

Ind. VI (1893) 464 et in Cale. Ann. XII (1911) 173 quoad ~

specimen Griffthianum (altera=D. grandis).

Gardens Bulletin, S.S.

169

Calamus intermedius Griff. in Calc. Journ. Nat. Hist. V
(1844) 86 et Palms Brit. Ind. (1850) 93 t. 211 A and B (Type
“saa ai = Malacca).

D. calicarpus Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 174 et Fl. Mal. Pen. V (1925) 37 pro parte. Syn. nov.

D. grandis Mart. sensu Ridl., Mat. Fl. Mal. Pen. II (1907)
177 et Fl. Mal. Pen. V (1925) 38 quoad syn. D. intermedius.
Syn. nov.

MaLAy PENINSULA: Malacca, loc. incert. (Fernandez, vern.
name Rotan Crysa. Type in Calc. Herb.). Johore, Bukit Soga
(Ridley, 11,205, female specimen only; male=D. melanochetes).

The type specimen in the Calcutta herbarium is in a
fragmentary condition and without external spathes, which,
however, seem to have been present when Beccari examined it.
This specimen has given rise to a considerable confusion, because
the absence of the spathes, in addition to the incompleteness of
the extant parts, makes extremely difficult the identification of
the species with any certainty. Anyhow, that it cannot be the
species accepted as such by Beccari is evident, for that has very
broad leaflets, whereas Griffith’s description and the specimen
itself indicate that D. intermedius is a narrow-leaved species like
D. angustifolius and D. Lewisianus.

Beccari’s interpretation of D. intermedius was based on
Scortechini’s collection that seems to have been so mixed with
parts of another species that Beccari must have been misled to
regard all the specimens in the collection as having been derived
from the same plant or clump; and presumably this is the reason
why he explained away the differences he detected principally
in the armature of the outermost spathes by considering the
species as very variable or polymorphous. From Beccari’s plate
20, I would regard the detached spathe (on the right hand side
of the plate) to be D. angustifolius; but obviously the major
part of D. intermedius sec. Beccari is D. grandis.

In making the identification of D. intermedius in the
Griffthian sense, I have used the type specimen together with
the description and the plates publised by Griffith. The impor-
tant clue to the identification of Griffith’s species with Ridley’s
specimen quoted above is the armature of the second spathe,
which is noted by Griffith as thickly armed. There are only two
species known to occur in the Malay Peninsula (in Malacca or in
its neighbourhood), that would answer this description: the
one is represented by the specimen I refer here to D. intermedius;
and the other is D. melanochetes. 'The latter, which also pro-
duces spathes very like the example depicted by Griffith, bears
bristles down to the very base of the inflorescence and even on
the peduncular part that is adnate to the sheath, the bristles
in the latter part being as a rule very closely set. There are

For, FX. (1937).

170

also noticeable differences in the armature on the basal portion
of the petiole and on the apex of the sheath in these two species,
D. melanochetes being more thickly and conspicuously armed
in these parts than D. intermedius. In Ridley’s specimen
quoted above the spadix is practically unarmed at the base; and
Martius, who had seen a specimen from Griffith, described the
peduncle as ‘“‘subinermis’’. But D. intermedius as interpreted
here appears to be a very close relative of D. melanochetes, and
observations in the field and better herbarium specimens are
necessary to determine whether the former species represents
merely an ecologic or varietal form of the latter. To achieve
this purpose, it would be useful if the collectors could procure
specimens showing the full range of variability of sheaths and
spathes in the two _ species. See also my _ remarks on
D. melanochetes, D. grandis, and D. monticolus.

IX. “Dezmonorops javanicus Furtado spec. nov.

D. stenophyllo et D. singalano propinqua est, sed ab illo
spatharum spinis subacicularibus valde gracilioribus ac pluribus;
et ab hoc spadicibus sessilibus vel penduculis valde brevioribus,
erectis, haud reflexis, hzec species facile distingui potest. A
grege, cui D. angustifolius pertinet, spatharum spinis valde
gracilioribus; fructibus immaturis sphzeroidioribus, haud elliptico-
ovatis; squamis (margine albescente excepta) fere concoloribus,
cum canaliculo profundiore differt.

CAUDEX scandens, cum vagina 2.5-3 cm. in diam.; frondibus
magnis cirrhiferis. Vagina frondium infra petiolum gibbosa,
apice suboblique truncata, aculeis laminaribus, elasticis schistosis,
ad 3.5 cm. longis, basi 3.4 mm. latis, per series obliquas
interruptasque confluentibus, apice ejusdem spinis longioribus,
porrectis. Petiolus 15 cm vel longior, subtus secus dorsum
aculeis dimensione variablilibus dispersis, in geniculo ad dorsum
tantum armatus, supra canaliculatus levigatus, inermis. Lamina
magna, in cirrhum terminans, secus rhacheos dorsum unguibus
3-7-fidis armata. Segmenta plura, ensiformia, 2.5-3.5 cm. inter
se remota: 30-38 cm. longa, 13-18 mm. lata, maxima latitudine
infra medium sita, apicem verstts sensim acuminata, basi abrupte
contracta, plicatula, nervis transversalibus conspicuis; supra ad
costas duas laterales et margines, ad costam mediam in tertia
parte terminale tantum, setulosa; subtus inermia. Inflorescentie
foemineze sessiles vel pedunculo 1— 1.5 cm. longo praedita, cum.
rostro 5-7 cm. longo, inermi vel fere, 22-25 cm. longe.
Spathe primarie: infima cymbiformis, bicarinata, aculeis
1-2.5 cm. longis, subaciculatibus, basi 1-2 cm. latis, solitariis
vel confluentibus armata; interiores inermes, conspicue nervose.
Spathe secundarie bracteiformes amplectentes. Spathelle con-
similes, in apicem triangularem involucronhorum zequantem vel
sunerantem producte. Involucrophorum obconicum obsoleto
angulatum, apice in cupulam asymmetricam oblique truncatam

Gardens Bulletin, S-S.

:
'y
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ee

E7E

expansum. Involucrum cupulare, truncatum, in involucrophoro
fere totum immersum. Perianthium fructiferum explanatum.
Fructus immaturi tantum visi, globosi, 10-12 mm. in diam (sine
apiculo 2-3 min. longo). Squame per series 15-18 imbricate,
concolores vel fere, fusco-graminez, secus marginem albescentes,
apice atrescentes, dorso conspicue-canaliculatee.

Java: Preanger, Gedeh (Heyne, 2,670=Herb. Bogor. num.
468. Holotype in Herb. Bogoriensi).

Besides the widely distributed, polymorphous D. melano-
chetes, no Demonorops of the section Cymbosthe had been
recorded from Java. D. javanicus records therefore another
species of the section for Java. "The nearest ally of this are the
mountain species of Sumatra and the Malay Peninsula, belong-
ing to the group of D. singalanus. ‘The leaflets are quite unarmed
in the lower surface, a condition that might not be constant in
the ‘species. In the gibbose part of the petiole there is a band
on either side of the ridge where the spines are absent, though
a little turther on in the sheath (in the portion that may be called
the wings of the puckered part of the petiole) the spines are both
conspicuous and large.

X. Mi erpnOrGya Lewisianus (Griff.) Mart., Hist. Nat. Palm.
III (1850) 327 t. 175-iv f. 1-7 excl. specimen “Gaudichau-
diame deec, 1m Hoek. this Brit: Ind. Vi (1893) 465;
Ridl., Mat. Fl. Mal. Pen. II (1907) 176 pro parte (ex altera
parte=D. melanochetes and D. grandis); Becc. in Calc.
Amn. XIT (1911) 87 t. 30 pro parte (ex altera parte=
D. angustipathus); Ridle., Fl. Mal. Pen. V (1925) 38 partim.
Calamus Lewisianus Griff. in Calc. Journ. Nat. Hist. V

(1844) 87, et Palms Brit. Ind. (1850) 94 t. 212. Type locality =

Penang.

D. angustifolius Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 174 et Fl. Mal. Pen. V (1925) 36 pro parte. Syn. nov.

D. calicarpus Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 174 et Fl. Mal. Pen. V (1925) 37 pro parte. Syn. nov.

D. microthamnus Becc. in Rec. Bot. Surv. Ind. II (1902)
221; Ridl., Mat. Fl. Mal. Pen. II (1907) 175; Becc. in Calc. Ann.
XIT (911) 92 t. 32; Ridl., Fl. Mal. Pen. V (1925) 37. Syn. nov.

D. monticolus var. B Becc. in Rec. Bot. Surv. Ind. II (1902)
221 nom. nud.

D. monticolus Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 175 et Fl. Mal. Pen. V (1925) 37 pro parte (quoad
specimina penangensia) Syn. nov.

D. monticolus var. pinangianus Becc. in Calc. Ann. XII
(1911) 85 t. 29 excl. tab. Griffithianam. Syn. nov.

Vols PX (1937).

E72

“D. petiolaris Mart., Hist. Nat. Palm. III (1850) 326 t.
Z-xvili f. 3; Becc. in Hook. f., Fl. Brit. Ind. VI (1893) 466 et
in Cale. Ann. XII (1911) 90 t. 31. Syn. nov. :

YD. petiolaris var. nudipes Becc. in Rec. Bot. Surv. Ind. II
(1902) 221. Syn. nov.

C. monticolus Griff. sensu Griffith, Palms, Brit. Ind. (1850)
97 quoad specimen penangense. Syn. nov.

ve: petiolaris Griff. (incl. vars. A and E) in Calc. Journ. Nat.
Hist. V (1844) 93, et Palms Erit. Ind. (1850) ror t. 216 f. vil.
Syn. nov.

MaLAy PENINSULA: Penang, loc. incert. (Griffith, in Calc.
Herb.; Lewis vern. name Kichun, in Calc. Herb. Type of
D. Lewisianus; Curtis, 2232); Waterfall (Burkill, 376); Penara
Bukit (Curtis, in July 1893); Penang Hill=Government Hill
(Abrams, in 1893; Ridley: 7034; 7895; 7896); Balik Pulau
(Ridley; 7900; 7902); Moniots Road (Curtis, 764). Province
Wellesley, Bukit Juru (Ridley; 7003; 7102). Dimdings, Pangkor
(Ridley, 7897, vern. name Rotan Kusom); Lumut (Ridley in
Feb. 1892; Curtis, 3444, male specimens only, female specimens =
D. calicarpus, vern. name Rotan Chuchok). Perak, Pulau
Rumbia (Seimund, on 15-iii-1925); Gunong Kledang (Curtis,
3351, male and female). Malacca: loc. incert. (Griffith, in Caic.
Herb).

SuMATRA: Lingga Archipelago: Pulau Selajar (Biinnemeijer,
7451 in Herb. Buitenzorg); Pasir Panjang (Biinnemeijer, 7016);

At first I thought D. petiolaris to be a good species, but
after comparing the material with that of D. Lewisianus, I find
difficulty in keeping it as a species quite distinct from the latter.
Ridley’s 9897 from Dindings, Ridley’s 7003 from the Province
Wellesley, Seimund’s specimen from the Rumbia island in Perak,
Curtis’s 3351 (male) from Perak, and Burkill’s 376 and Ridley’s
7895 from Penang form, so to say, the transitional stages
between the extreme forms, which may be called D. monticolus
var. pinangianus and D. petiolaris. Griffith’s specimen cited
here as from Penang bears no locality and is named Calamus
monticolus var. 6 (=D. monticolus); but in all probability it is
the Penang specimen Griffith cited under C. monticolus, in his
second monographic work on Palms (1850). Regarding the
armature of its spathes and the apical wings of the leaf-sheaths,
the specimen agrees well with Ridley’s n. 1895, though the
latter is much less leprose. The authentic specimen of
D. petiolaris (Griffith’s specimen from Malacca cited above),
which consists of a terminal portion of a radical leaf and an
entire upper leaf with its sheath, is too meagre to be of any
great help in the identification of the species. The upper leaf
measures about 12 inches and the petiole is not terete. Similar
leaves are produced by forms of D. calicarpus as interpreted here

Gardens Bulletin, S.S.

173

and so I think Beccari’s acceptation of this species is. correct,
though the species has to be considered as a synonym of
D. Lewisianus.

The type specimen of D. Lewisianus in the Calcutta her-
barium (the Penang specimen collected by Lewis) was derived
from rather thick rattan and does not correspond to Griffith’s
plate 212 which is a representation of a terminal portion of the
stem approaching a stage referred by Beccari to D. petiolaris. In
its shape, armature and indumentum this type specimen, which
has lost the terminal portion of its spadix, corresponds to Ridley’s
7902 referred by Beccari to D. monticolus var. pinangianus.
The species varies a great deal not only in the size and shape
of the spadices but also in the length and bristleness of the
peduncle; but no important differences can be found to keep
these various forms separate as varieties or species. The
indumentum on the leaf-sheaths, petioles, and external spathes
is also variable, being in some:cases very thick and persistent
and in others entirely or almost deciduous very early during
the course of the development of the organs. These differences
are perhaps due to the age and the habitat of both the individual
plants as well as of the clump, the suckers produced at a time
when the soil around it has been exhausted by preceding suckers
and when the competition for space and light has become very
acute, becoming very stunted and weak. Perhaps Curtis’s 3351
(female) from Gunong Kledang in Perak, which corresponds to
Beccari’s plate 32 of D. microthamnus, represents such a stunted
plant, while Lewis’s specimen in the Calcutta herbarium and
Ridley’s 7902 in the Singapore herbarium (forma D. monticolus
var. pinangianus) represent forms produced at a time when the
competition between the suckers of the clump for space, food,
etc. is not so keen and therefore does not affect the vigour and
luxuriance of the individual plants.

The specimen Gaudichaud collected in. Penang (preserved
in the Paris herbarium), and which Eeccari referred to
D. Lewisianus, is referrable to D. angustispathus.

Beccari also referred Ridley’s specimens no. 9466 from the
Kew and the Calcutta herbaria to D. Lewisianus. Beccari’s
description of these specimen is applicable to Ridley’s 9466 in
the Singapore herbarium, which I cite under D. melanochetes.

XI. Dzemonorops melanochztes Bl. in Roem. et Schult., Syst.
Veget. VII (1830) 1333 excl. syn.; Mart., Hist. Nat. Palm.
ITI, ed prima (1836 ?) 198 t. 117 f. II et ed. 2 (1850) 203 et
326 Kunth, Enum, Pl, III (1841) 202; Blume, Rumphia II
(1845 ?) tt. 134 et 137, et III (1849) 3; Mig., Fl. Ind. Bat.
III (1855) 68; Becc. in Rec. Bot. Surv. Ind. II (1902) 210,
et in Cale, Ann.) AI(19n1) 47. win

Vol. 1X....( 1937).

174

D. angustifolius Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 174, et Fl. Mal. Pen. V (1925) 36, pro parte. Syn. nov.

D. Lewisianus Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 176-p;p:; Bec.1n Cale. Ann. MID (ior) 87 yprpese eens
Fl. Mal. Pen. V (1925) 38 pp. Syn. nov.

D. malaccensis Mart. sensu Becc. in Cale. Ann. XII (1911)
61 t. 12 (excl. spec. Griffthianum) Syn. nov.

D. monticolus Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 175 quoad specimina borneensia; Becc. in Cale. Ann. XII
(1911) 84 quoad specimen Ridleyanum; Merr., Enum. Bornean
Pl. (1921) quoad loc. Borneo tantum; Ridl., Fl. Mal. Pen. V
(1925) 37 quoad Borneo. Syn. nov.

Calamus melanochetes (Bl.) Griff. in Cale. Journ. Nat.
Hist. V (1844) 86, et Palms Brit. Ind. (1850) 92 in nota; Migq.,
Palm.» Arch: cid) (1566)-128:

D: Schmidtianus Becc: in “Schmidt., Fl) Kol lame
IX (1909) 330; et Calc. Ann. XII (1911) 52 t. 7. Syn. nov.

? D. Hallierianus Becc. in Calc. Ann. XII (1911) 218 t. g9
quoad specimen Hewittianum. ? Syn. nov.

MALAY PENINSULA: Kemaman, Bukit Kajang alt. 500 ft.
(Corner, 30428). Pahang: Pulau Tioman: at Sedagong, alt.
1,000 ft. (Nur, 18891); Bukit Surin alt. 1,000—2,o00 ft. (Hender-
son, 21710); Joara Bay (Burkill: 983, vern. name Rotan Langah;
1016). Penang, Moniot Road (Ridley, 9466). Province
Wellesley, Bukit Juru (Ridley 7100). Negri Sembilan, Gunong
Tampin (Holttum 9561). Johore, Serom (Ridley, in Nov. 1900);
Bukit Soga (Ridley, 11205, male only; the female bearing this

no.=D. monticolus). Pulau Aor, alt. circ. 600 ft. (Henderson
18358).

BORNEO: Sarawak, Satubong (Hewitt, in 1906, vern name
Rotan Bahut); Matang (Ridley, in July 1903); Quop, (Hewitt,
23-1v-1907, n. R.A.).

This species was recorded by Beccari for Java, Borneo and
Sumatra, but not for the Malay Peninsula, though Martius had
recorded as occuring in the Malay Peninsula in the island of
Penang. Merrill (Univ. Calif. Publ. Bot. XV 1929, p..24) refers
to this species Elmer’s specimens n. 21102 from ‘Tawao in
British North Borneo, ‘but the duplicate material (consis-
ting of a few fruits and leaflets) in the Singapore herbarium
does not appear to represent this species.

The Peninsular material of the species has been much con-
fused by Beccari and others. Beccari has regarded Ridley’s n.
11205 (male) from Johore as D. malaccensis (Griff.) Mart., while
he determined a similar specimen collected by Ridley in Penang
sub. n. 9466 as D. Lewisianus (Griff) Mart. Previous to the

Gardens Bulletin, S.S.

175

issue of his monograph on Deemonorops (1911) Beccari conimuni-
cated his determinations to Ridley, who took it for granted that
Beccari’s acceptation of the two species of Griffith were correct;
and as both the specimens referred by Beccari to the above two
species were conspecific, Ridley made D. malaccensis a synonym
of D. Lewisianus, interpreting the species in the sense given to
them by Beccari. On a sheet belonging to D. melanochetes,
Ridley also noted that a plant conspecific with the specimen was
cultivated in the Kew Gardens under the name of D. Lewisisanus
If so, I suspect that this name was given to the plant after its
determination by Beccari.

In the first and the second editions of the Historia, and
also in its Appendix, Martius quoted under D. melanochetes Bl.
a specimen collected by Wallich in Penang. ‘This reference has
been noted by Griffith, Kunth, Miquel, etc. In the Calcutta
Annals XII, 1911, p. 47 Beccari has cited all the references under
D. melanochetes without making any exclusions, though he does
not credit the species to the Malay Peninsula. Neither he, nor
Ridley has tried to identify the Wallichian specimen from
Penang. Presumably the specimen is not at Kew; and a search
may have to be made in Martius’ herbarium in Munich*
Similarly the Philippine material referred by Martius under this
species has to be examined to see whether it had been rightly
identified; for neither Beccari nor Merrill (Enum. Philipp. Pl. I,
1922) credits the species to the Philippines.

The Peninsula material of D. melanochetes varies a great
deal regarding the spininess of the spathes, and the width and
the remoteness of its leaflets; and I am unable to say whether or
not this variability is dependent on the age and the habitat of
the plants. But according to Beccari, this species seems to vary
extremely even in Java itself, and so before separating the various
forms into distinct varieties, one will have to devote a great deal
of time to study the variations due to the age and the habitat
of the plants. Some of the Peninsular forms look (e.g.
Henderson n. 21710) exactly like Beccari’s plate of
D. Schmidtianus from Koh Chang Island off the coast of Siam,
while some others (e.g. Corner, 30428) like Blume’s plate 134,
with the exception of the armature of the basal part of petiole,
which appears to have been wrongly depicted as having numerous
acicular spines; while still others could be easily mistaken for

* A specimen from Wallich’s herbarium consisting of a spadix
without the outermost spathe is in Martius’s herbarium in Munich.
From the photograph kindly sent to me by Prof. M. BuRRET of the
Berlin Herbarium, it appears to be the subject of Martius’s plate 117
fig. I.; but I am unable to decide whether the specimen is really
D. melanochetes, and not D. grandis which also has its second
spathes somewhat similarly armed. The plate of Martius depicts the
thorns on the second spathes with very much swollen bases as in
D. grandis, whereas the second spathes in D. melanochetes and
DPD. intermedius have thorns less tumescent at base.

Vor, EX. (1937).

176

D. intermedius or D. monticolus; in fact so great is the resem-
blance of the armature and shape of some spadices in our material
to Griffith’s plate 211-B (D. intermedius) that I had at first
thought to have recognized the latter in D. melanochetes. But
Griffth’s plant is at once distinguished in having peduncled
spadices and in being quite bare on the peduncle as well on its
portion that is decurrent on the sheath. These characteristics
were described for D. intermedius by Griffith and can be noticed
both in Griffith’s plate and the type specimen preserved in the
Calcutta herbarium. On the other hand the spadices in
D. melanochetes are sessile and thickly armed with nigrescent
spines up to the base and often also on the decurrent peduncular
part.

The Bornean material cited above was referred by Ridley to
D. monticolus, a species Merrill credited for Borneo on Ridley’s
authority. Beccari suggested the possibility of the material
belonging to D. Hallierrianus; and indeed Beccari’s plate 99 fig.
3, which is of the specimen collected by Hewitt in Quop, seems
represent D. melanochetes.

Possibly Ridley’s n. 11461 from the Government Hill in
Penang also belongs here, but the petiole is not typical.

The beak of spadices may be longer or shorter than the body.
The fruit scales are arranged in 15-19 vertical series.

XII. Dzmonorops monticolus (Griff.) Mart., Hist. Nat. Palm,
IIT (1850) 328. t. 175'f.. v; Bece.in- Hook.f, Fl. Bee ie.
VI (1893) 465 et in Rec. Bot. Surv. Ind. II (1902) 221;
Ridle., Mat. Fl. Mal. Pen. II (1907) 176 pro parte et excl.
specimina borneensia; Becc. in Cale. Ann. XII (1911) 84
excl. specimen Ridleyanum; Ridl., Fl. Mal. Pen. V (1925)
37 pro parte.

Calamus monticolus Griff. in Cale. Journ. Nat. Hist. V
(1844) 90, Palms Brit. Ind. (1850) 97 tt. 214 A, B, & C (excl.
spec. pinangense) Type locality = Malacca).

D. angustifolius Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 176 et Fl. Mal. Pen. V (1925) 36 pro parte. Syn. nov.

D. monticolus var. pinangianus Becc. in Cale. Ann. XII
(1911) 85 quoad tabulam Griffithianum. Syn. nov.

MaLAy PENINSULA: Malacca, Bukit Dusun Paya (Alvins,
1217, vern. name Rotan Miniak). Negri Sembilan, Gunong
Tampin (Goodenough, 1857).

The two specimens cited above agree very well with
Griffith’s plate and specific description and represent a species
likely to be confused with D. Lewisianus from Penang (as
Griffith had done). The authentic specimen in the Calcutta
herbarium appears to be the Penang specimen that was referred
here by Griffith in his second work, and which I cite under D.

Gardens Bulletin, S.S.

177

Lewisianus. The Malacca species, D. monticolus, seems to be
closely related to the latter, but the absence of leaf-sheaths and
petioles in the two specimens at my disposal precludes a deeper
inquiry into the affinities existing between the two species.

Beccari interpreting D. monticolus attached undue
importance to the following words in Grifhth’s Latin diagnosis :
“spatha eatima secus carinas duas spinis gracilibus deflexis
armata’’; for this description, if taken to mean that the spines
are present only along the two carinz, corresponds neither to
Griffith’s description in English, nor to his plates 212-B and C.
The English rendering of the description of the outermost spathe
runs thus:—‘‘The outer one [spathe| 16-17 inches long,
bicarinate, armed between and along the carine with weak long,
subulate, deflexed, black spines’’:

In view of this I regard both Griffith’s plates as representing
the typical plant.

At first sight, D. monticolus might be mistaken for some
forms of D. melanochetes and for D. intermedius, and in fact
Griffith’s plates of the last and the first mentioned species show
many points of resemblance. But unlike that of D. intermedius
or of D. melanochetes the second spathe of D. monticolus is
unarmed, or armed only with a few bristles here and there to-
wards the apex and along the carine, a probable reason why
the species was confused by Griffith himself with D. Lewisianus.
The peduncular portion may be armed or unarmed.

Both the specimens quoted above were cited by Ridley under
D. angustifolius. See also my remarks under D. melanochetes,
D. grandis, and D. Lewisianus.

XII. Dzmonorops pseudo-sepal Becc. in Hook. f., Fl. Brit.
Ind. VI (1893) 465; Ridl., Mat. Fl. Pen. II (1907) 180; Becc.
in Cale. Ann. XII (1911) 79 t. 25; Ridl., Fl. Mal. Pen. V
(1925) 40 inter species dubias.

: _D. bakauensis Bece. in Cale. Ann. XII (1911) 221 t. 100
Syn. nov.

Matay Peninsuta: Johore, Pulau Tinggi (Burkill, 866).

SUMATRA: Banka Archipelago. Bakau Isl. (Teysmann,
herb bogor. n. 277, type of D. bakauensis, vern name Rotan
Rundang); Banka, on Bukit Tampang, near Sungei Liat
(Bunnemeijar, 1736, vern. n. Rotan rondang); Banka by Sungei
(Teijsmann, herb Bogor. n. 269, bern n. Rotan Rundang); Banka
on Gunong Menumbing, near Muntok (Biinnemeijer, 1465, vern
name Rotan Getah). (All the Sujatran specimens are from the
Buitenzorg herbarium). ee

The species appears to represent a form midway between
D. Lewisianus and D. grandis. In-fruits and in the armature

Pon LX..( 1937).

178

of its leaf sheath, the species resembles very much D. grandis,
but the leaflets are much narrower and the spadices shorter. The
species should be included among the Deémonorops having sessile
or subsessile spadices, and not among those which have stalked
spadices. Judging from the outermost spathe present in
Bunnemeijer’s 1736, the species appears to be closely related
to D. Binnendijki Becc.

Beccari separated D. bakauensis from D. pseudosepal because
of the broader leaflets and smaller, ovoid fruits in the former;
but the fruits in the type specimen are young. Biinnemeijer’s
specimen 1465 shows both kinds of leaflets and the fruits which
are better developed than in the type of D. bakauensis are
spherical as in D. pseudosepal. In view of this I do not see
any reason to maintain D. bakauensis as a good species.

XIV. Dzmonorops sepal Becc. in Hook. f., Fl. Brit. Ind. VI
(1893) 469 et in Cale. Ann. XII (2011) 77 tea; Ridie wee
Mal. Pen. V iro25) 30;

D. aciculatus Ridl., Mat. Fl. Mal. Pen. II (1907) 176 et F1.
Mal. Pen. V (1925) 38; Becc. in Cale. Ann. XII (1911) 231 inter
species dubias. Syn. nov.

D. angustifolius Mart. sensu Ridl., Mat. Fl. Mal. Pen. II
(1907) 174 quoad syn. D. sepal Becc.

D. calicarpus Matt. (sensu \Ridl.;° Mat. Fie oMal Peart
(1907) 174 et Fl. Mal. Pen. V (1925) 37 pro parte. Syn. nov.

D. hygrophilus Mart. sensu Ridl., Fl. Mal. Pen. V (1925)
40 pro parte. Syn. nov.

D. imbellis-Becc. in Rec. Bot. Surv. Ind. II (1902) 220;
Ridl., Mat. Fl. Mal: Pen. 11. (i907): 279; _Beéc. im Caleta:
XII (1911) 80 t. 26; Ridl.. FL Mal. > Pens WW, (ae@25)e aie eees
species dubias. Syn. nov.

MALAY PENINSULA: Perak, Gunong Hijau (Ridley in 1892,
female. Syntype of D. aciculatus); ibid., southern summit, alt.
4,300 ft. (Burkill and Haniff: 12763; 12944); Sungei Mengkarau
in Larut Hill (Anderson: 176; 166); Maxwell Hill (Ridley, in
June 1893, male. Syntype of D. aciculatus). Selangor,
Sempang Mines (Ridley, 15876). Pahang, Sungei Lemoi at Ulu
Anak (Dolman, 28157); Cameron Highlands by Sungei Ikan near
Ulu Telom, alt. 4,500 ft. (Dolman, 27631, vern. name Rotan
Baik); Fraser Hill alt. 4,o00-4,370 ft. (Burkill. and Holttum,
7785). |

The fruiting specimens may be arranged in three classes
according to the number of the series into which the scales are
arranged on fruits, vizg:— «|

(a) 15 series: BurkillSd#ind Haniff: 12944 and 12763.

(b) 15-17 series: Ridley on Gunong Hijau; Dolman:
28157 and 27631.

(c) 18 series: Anderson 176.

Gardens Bulletin, S.S.

179

The fruits in Burkill and Holttum, 7785 are injured and so
the number of the scale series cannot be counted. Anderson’s
176 bear some fruits which are globose as in Beccari’s plate of
D. Scortechinii, though other fruits are normal. This shows
the possibility of D. Scortechinii being but a form of D. sepal.
Ridley’s specimen from Gunong Kledang has a very diffuse
spadix in fruit resembling that of D. imbellis. Some spadices
I have seen are distinctly ventricose and nodding, some straight
and not so ventricose. I think this variability is due to the
vigour of the plants and of the spadices.

Ridlev based his D. aciculatus on two specimens he had
collected in Perak. He gave the length of the peduncle as
4 inches, misjudging as free the decurrent portion of the
peduncle that has become detached off the sheath on being
pulled in the living state. The actual length of the peduncle
in Ridley’s specimens is about 3-4 cm. only. But the species
itself is very variable in respect to the length of peduncles. In
the Maxwell Hill specimen quoted above the spadix bears a
neduncle which is somewhat longer than in all the others I
have so far examined, and apparently is nodding as in
D. imbellis; in some other specimens the peduncle is subsessile or
hardly measurable. This variability in length of peduncles and
in the number of the vertical series into which the fruit scales
are arranged, as well as a comparison of the plates 24 and 26
given by Beccari with the specimens in the Singapore herbarium
lead me to consider D. aciculatus, D. imbellis, and D. sepal as
ae aa

XV. Dzmonorops sepal var. pahangensis Furtado var. nov.

A forma typica spatha infima aculeis subulatis vel
setiformibus appressis vel arrectis atro-fuscis densissime armata
(spatharum D. melanochetes Bl. instar) heec varietas differt.
Fructuum squamz per series verticales 18 imbricate.

MaLAy PENINSULA: Pahang, Fraser Hill, alt. 4,000 ft. (Nur.
11208. Holotype in Singapore).

The petiolar and laminar portions, present in the specimen,
appear to be of the radical leaves. The petiole, which is covered
with brown indumentum, bears long and short spines all over
on the dorsal side, the longest spines being 3 cm. long. At the
mouth of the sheath the spines are still longer (up to about
6 cm.). .Leaflets about 4o cm. long. 2.5 cm. wide, unarmed
beneath or occasionally with a few prickles in the apical half
of the midrib beneath; in the unper surface the 4 lateral nerves,
and occasionally the mirdib also, are remotely setulose. The
spadix with its fruits is like the typical of D. sepal.

D. sebal is a long climbing rattan: but this variety has been
noted by the collector as short stemmed plant. The collector’s
notes run thus: ‘‘Stem short, heavily armed with straight spines.

Vol. LX2.(.1937).

180

Leaves 6-7 ft. long, armed with straight spines. No leaf-
flagellum. Fruits pale green edged brown.’’ Further field
observations are needed to see whether this variety represents
D. sepal in the early stages of its development, for I note in the
apical parts of the outermost spathes a tendency for the softer
bristles to fall off and become like that of D. sepal.

Index to Collectors’ Numbers.

Abrams s.n.-X. Alvins: 979-III; 980-VI; 1217-XII; 1878-VI.
Anderson: 166 & 176-XIV. Bunnemeijer: 1465 & 1736-XIII;
7016 & 7451-X. Burkill: 376-X; 866-XIII; 983 & 1016-X];
1372-VI; 2551-VI; 6520-VII; 6536-VI. Burkill. &
Haniff: 12763 & 12944-XIV. Burkill & Holttum:
7785-XIV. Corner: s.n.-I; 12749-I; 29750-VII; 30077 &
30389-III; 30428-XI; 30447-VII. Curtis: s.n.-V, X, 764-X;
2232-X; 3160-III; 3351-X 3444-IJ] & X. Derry: 108 &
r10-VI; 112-IJI. Dolman: 27631 & 28157-XIV. Elmer:
21102-XI obs. Fernandez: s.n.-VII & VIII. Fox: 11299-I.
Gaudichaud: s.n.-II. Goodenough: s.n.-VI; 1660-VI;
1857-XII. Grashoff: 196 & 639-VII. Griffith: s.n.-I & X.
Henderson: 18358-XI; 19484-VI; 21710-XI; 24620-I. Herb.
Bogor. nos. 222-VII; 269-XIII; 404-VII; 468-IX. Hewitt:
sn. R.A.-XI. Heyne: 2548-VII; 2670-IX. Holttum:
9561-XI; 9778-VI. Lewis: s.n.-X. Mat: s.n.-VI. Nur:
11208-XV;- 18891-XI. Ridley: s.n.-I, III, VI, VII, X,
XI, XIV; 3498 & 3499-VI; 3506-VI; 4622-1; 5122-VI; 5123-
I obs. & VI; 5124-VI; 6279-I; 6672-VI; 7002-II; 7003-X;
7034-3 7100-XI1; 7102-X; 7878-III; 7880-1; 7895, 7896 &
7897-X; 7899-III; 7900 & 7902-X; 8957-VI; 9466-X obs. &
XI; 11203-VIII & XI; 11461-XI obs. 11826-826-VII; 11983-
I; 13302-I & IV obs.; 14115-III; 14719-I & VII; 15876-XIV.
Seimund: s.n.-X. Teijsmann: 260 & 277-XIII.

Gardens Bulletin, S.S.

—_ en

INDEX TO BOTANICAL AND VERNACULAR NAMES

Calamus :
acanthopis VI
angustifolius I ...
Calicarpus III °..
grandis VI
hygrophilus VII
intermedius VIII
Lewisianus X
melanochaetes XI
monticolus X; XII
petiolaris X

Demonorops :
aciculatus XIV

angustifolius I, IV, VI ene :
Bek OS, - RT; x1: XIV

angustispathus sp. nov. II

bakauensis XIII

ealicarpus III; VIII: X; XIV

carcharodon L es

congestus IV

Curtisii sp. nov. V

fissus var cinnamomeus VII

erandis:Il; Vi, VIII ...

Peilierianus a) wa

hygrophilus I; VI; VII; XIV

imbellis XIV

intermedius VI;

intermedius var.
V1. acy

VIII
nudinervis

javanicus sp. nov. IX ...
Kirtong VI

Kurzianus Ne

Lewisianus II, VL xX, ‘XI
malaccensis VI; XI ... a
melanochetes VIII obs., XI
Miscrothamnus X :
Moticolus X, XI, XID

monticolus var.
ae Qae .

petiolaris X /
pseudosepal XIII

penangensis

Vol. IX. (1937).

181

Schmidtianus XI
sepal XIV

sepal var.

pahangensis
nov. XV ¥, be

Rattan=Rotang=Rotan ...

Rotan :
ayer VI
baik XIV
bahut XI
cheranang VI obs.
chichik landak VI
chrysa VIII
chuchok X
chongkak VI
chucho miniak III
chinchin VI
vetah: X EIT
ghetah I ...
kichum X
kikir I
kerotong VI
kertong VI obs.
kelemunting VII
kusom X
langah XI
manau I
miniak XII
rondang XIII
rundang XIII
relang VI
sabut III
sepat I
sumang VI obs.
sunang VI
tawar I; VII

Uwi bungkus I

Uwi getah VI

var.

182

4

“ PALM MALESICZ.
VII.—Two new CALAMI from the Buitenzorg Gardens.

By CX. FURTADO,
Botanic Gardens, Singapore.

Early this year (1936) I noticed cultivated in the Botanic
Gardens, Buitenzorg, Java, two Calami showing some characters
that are rather infrequent in the genus, and on further study
both proved to be new. The one, C. Steenisiti—named in honour
to Dr. C. G. G. J. van Steenis of the Buitenzorg Herbarium, who
had taken a great deal of trouble to facilitate my studies during
my visit to the Gardens—is recorded to have been introduced
from New Guinea, and possesses so many anomalous characters
that it cannot be made to fit in any of the sixteen groups into
which the genus Calamus has been divided by Beccari (Ann.
Roy. Bot. Gard., Calcutta, XI, 1908, pp. 64-69), though it shows,
at the same time, several marks of affinity to some of the New-
Guinean species belonging to Group V (vide infra). ‘The second,
described here as C. inopitatus, is of unknown origin. It is a
stemless palm and produces leaflets which are covered in the
under surface with minute pinkish tomentum, both of which
characters are not common in the genus. It is closely related
to C. Burckianus Becc. from Java, a member of Group II of
Beccari and possibly also stemless.

“Calamus Steenisii Furtado spec. nov. (Planta faminea tantum
visa). Caudex soboliferus, alte scandens, circ. I7 m.
longus, cum vagina 2.5 cm. in diametro. Vagina frondium
flagellifera, infra petiolum gibbosa, furfureo ferrugineo,
deciduo, dense leprosa, aculeis numerosissimis, tenuibus,
applanato-acicularibus, primum ferrugineis, mox levigatis,
stramineis per series transversales interruptas, 3-5 mm. inter
se remotas dispositis, basi connatis, in partibus liberis
fragilibus armata. Ochrea papyracea, exsuccosa, fibrosa,
auriculo-lanceolata, circ. 50 cm. longa, e basi 2.5 ‘cm. lata
in apicem acuminatissimam interdum fissam transiens, intus
leevigata, extus obscure bicarinata, deciduo fusco leprosa,
spinis, flaccidis, laminaribus, exsuccosis, solitariis, vel basi
connatis, ad 6 mm. longis przedita. Petiolus subtus
convexus, usque geniculi basin inermis, supra applanatus,
basin versus concavusculus, secus margines aculeis 3-4 mm.
longis, rigidis per tres lineas dispositis armatus, furfureo
fusco deciduo leprosus, 8-12 cm. longus, ad 1 cm. latus.
Lamina circ. I-1.20 longa, ecirrhifera vel interdum in

Gardens Bulletin, S.S.

183

cirrhum abortivum, filiformem, inconspicuum, szepe cum
segmento terminali adnatum, ad 7 cm. longum exeuns, in
rhachi eodemmodo leprosa, subtus secus rhacheos dorsum
marginesque aculeis dispersis, reduncis, basi tumescentibus,
apice atris, solitariis, interdum transverse connatis armata.
Segmenta plura, untrinsecus circ. 50, alternantia vel sub-
opposita, ad angulum 45° divergentia, lineario- ensiformia,
maxima latitudine in media sita, circ. 20-23 cm. longa, 12-15
mm. lata, acuminata, basin versus attenuata et in marginibus
crispata, infima subito oblique contracta, ad margines subtus
secus costam mediam tantum et supra ad costas tres setulosa.
Inflorescentie famine foliis longiores, circ. 2 m. longe,
flagelliformes, in axi aculeis reduncis, solitariis vel digitatis,
rigidis, ad 5 mm. longis armatee, deciduo furfuracez, in
ramos primarios 2-4 divisee, pedunculo ancepiti, secus
margines preesertim aculeis minutis preedito suffulte. Rami
primarii ad spatharum os orientes, statu juvenili in lobo
spatharum libero obtecti, circ. 30 cm. longi, spiculas
utinsecus 5—7 ferentes. Spathe: primarie basi tubulose,
cylindricee vel ancepites, aculeis dimensione variabilibus, ad
3 mm. longis rigidis, raro digitatis armatz, apice in lobum,
liberum, mox marcescentem, parce aculeolatum, in parte
summa fere inermem, interdum fissum, 1n modo auris asini
producte; secundarie perbreviores, circ. 2 cm. longe, papy-
racez, infundibuliformes, in lobum acuminatum unilateraliter
productee, parce aculeolate. Spicule ad spantharum os
secundarium orientes, axi flexuosz, 8-12 cm. longae, florum
glomerulos distichos, alternatos gerentes. Spathelle in-
fundibuliformes, oblique truncate, acuminate, aculeolis
perbrevissimis, rigidis, basi tumescentibus scabride.
Involucrophorum cupulare. Involucra profunde cupularia,
in singulo involucrophoro germinata, connata. Flores ad
singula spathella germinati, ambo fceminei vel in parte
spicularum apicali alter neuter (masculus?). Perianthium
fructiferum pedicelliforme vel subpedicelliforme. Fructus
immaturi tantum visi, ovoidei, circ. 12 mm. alti, 6-7 mm.
crassi, squamis per series verticales 18-22 disposities,
concoloribus, stramineis. Albumen homogeneum.

PatriA: Nova Guinea borealis.

The plant described here is cultivated in the Botanic Gardens,
Buitenzorg, under the gardens’ no. X.E. 2, and is said to have
been introduced from the northern parts of New Guinea. The
herbarium collection studied by me bears the Singapore Field
No. 30840. The holotype is preserved in the Singapore
herbarium, and the isotypes will be distributed to the herbaria
of Buitenzorg, Leiden, Kew and Berlin.

The edges of the leaflets are slightly turned upwards.
Judging from the marks of the fallen flowers and fruits, it

Vol. IX. (1937).

E

184

appears that the two female flowers are borne at each spathel
except in the distal ends of the spikelets where one neuter
(or male?) and a female occur at each spathel.

It is extremely difficult to make this species fit in any of
the sixteen groups into which Beccari has subdivided the genus
Calamus. If the presence of the long dilated, auriculiform, free
apical extension of the primary spathes is disregarded, the
species would certainly fall into the Group V. Further the
scabridity on the spathes and spathels would place C. Steenisii
into the Group V-A, of which sub-division no species is known
to occur in the part of the Indonesia lying to the east of Borneo.
No member of this group is known to possess very long ocrea at
the mouth of the leaf-sheaths, a character which, if one dis-
regards the abovementioned scabridity, would put the species
into the Group V-8, and precisely in the section which includes
the species that occur in New Guinea alone.

But if the dilated ends of the primary spathes are taken into
consideration, two Groups VI and XVI may partly lay claim to
this species, though again no species belonging to both these
groups are known to occur eastwards beyond Borneo. All the
species of Group VI have spathes which split or open longitudin-
ally almost down to the base into loriform laminze. The species
of Group XVI, though have the primary spathes similar those
in C. Steenisii, differ from the latter in that they all have very
much compressed, short spadices and ruminate seeds (C. Steenisii
has very long spadices and non-ruminate seeds).

All this suggests the necessity of redefining Groups V, VI
and XVI, or of creating a new one so as to keep several of the
New Guinea species together and to include such species which,
together with C. anomalus Burret, C. Lauterbachii Becc., C.
nannostachys Burret, and C. Steenisti, cannot be made to fit
definitely into any of the sixteen groups of Beccari, though all
of them show marked affinities to the species in the three groups
referred to above.

‘Calamus inopinatus Furtado spec nov. (Planta mascula tantum
visa). Caudex soboliferus, acaulis, interdum ad 35 cm.
longus, 2.5 cm. crassus, per totam altitudinem radices zerias
ad 8 mm. crassas solum attingentes gerens. Vagina
frondium 10-15 cm. longa, haud flagellifera et non gibbosa,
in latere ventrali ad basin usque.vel fere aperta, in petiolum
sensim transiens, aculeis laminaribus, elasticis, ad 5 cm.
longis, acuminatis, basi ad 4 mm. latis, digitatis vel interdum
per series obscuras dispositis armata. Ochrea in modo alee
secus margines vaginee enata, apice truncata, haud in lobos.
liberos producta, in alis utraque 6-10 mm. lata, rigida,
fragilis, aculeis rigidis, acicularibus, atris, ad 5 mm. longis,
basi, per series transversales interruptas connatis preedita,
basin versus sensim in vagina evanescens. Petiolus 20-30

Gardens Bulletin, S.S.

185

cm. longus, circ. 6 mm. crassus, teretiusculus, supra anguste
canaliculatus, sulcos aculeorum exhibens, aculeis acuminatis,
laminaribus, elasticis, stramineis ad 2.5 cm. longis, basi ad
4 mm. latis, apicem petioli versus solitariis, basin versus
seepe digitatis vel oblique subverticillatis. Lamina ad 1 mm.
longa, imparipinnata vel fere; secus rhacheos dorsum in
duabus partibus inferioribus tantum aculeis solitarlis
laminaribus, distantibus preedita, signis aculeorum conspicuis
impressa. Segmenta numerosissima, eequidistantia vel fere,
alternantia vel subopposita, 3-3.5 cm. inter se remota, ad
35 cm. longa, ad 2 cm. lata, lineario ensiformia, in apicem
filiformem, setulosam terminata, intermediaria longissima,
subtus roseo-tomentellosa, secus costas tres supra medium
vel paulo infra setulosa, supra inermia, levigata. Inflores-
centie mascule inermes, haud flagelliformes, frondibus
breviores, ad 60 cm. long, in ramis primariis 3-4 divise,
in rhachi spathisque deciduo fusco-furfuraceze, pedunculo in
vagina includente, nonnihil infra ochrez apicem attigente.
Spathe: primarie plus minusve striate, ad 6 cm. longe,
tubuloseze, apice oblique truncate, unilateraliter acuminatze
et in altero latere fisse; secundarie consimiles, cequilongze
vel minores; tertiarie minores, infundibuliformes, acumin-
ate, striate. Spathelle consimiles, apiculate, in altero
latere fissee. Rami primarii 10-15 cm. longi, interdum
ramus basalis ad 40 cm. longus, nonnihil infra os spatharum
inserti, in ramos secundarios, interdum tertiarios, eodemmodo
infra orificium spatharum orientes divisi. Spicule prope
basin spatharum orientes; primariz ad 9 cm. longe, basin
versus secundarias 2-3 cm. longas szpe ferentes. Flores
solitariil, alternati, confertissimi, ex spathellis usque duas
partes exserti, in involucro profundo cupulari insidentes,
utrinsecus in speculis primariis ad 15, ac in secundariis 9-12,
ad 4 mm. longi, petalis sesqui longioribus quam sepalis,
ambobus striatis, ovato lanceolatis.

PATRIA: ignota.

The plant is cultivated in the Botanic Gardens, Buitenzorg,
and bears the gardens’ no. X.E. 24. The specimens
described here were collected by me under the Singapore Field
number 30919. ‘The holotype is preserved in the Singapore
herbarium. Duplicates will be distributed to the herbaria of
Buitenzorg, Leiden, Kew and Berlin.

This peculiar species belongs evidently to the Group II of
Beccari and seems to be closely related to C. Burckianus Becc.
from Java, which is probably also a stemless species; but it is
described to have its petiole unarmed, leaf rachis unarmed or
armed with small claws beneath, and the primary (three) costze
of the leaflets setose above and sometimes also beneath, whereas
C. inopinatus has its petiole and rachis of the leaves armed with

Vol. IX. (1937).

186

rather long, laminar spines, its leaflets quite smooth above but
armed along the three ribs beneath, as well as covered with
minute, pinkish tomentum in the lower surface. "The only other
stemless Calamus is C. pyg@us Becc., a species belonging to the
Group V of Beccari and, therefore, differing in its spikelets and
spadix-branches being sessile, that is, arising at the mouth of
their respective spathes.

Gardens Bulletin, S.S.

a Se

187

ON THE APPLICATION OF THE TERMS MALAYSIA
AND MALAYA IN PLANT GEOGRAPHY.

By C. G. G. J. VAN STEENIS,
~Buitenzorg, Java.

There is rather a lot of confusion about these terms, as
pointed out by HAVILAND (1) so long ago as 1897.

The Malayan language is properly limited to Western
Malaysia (Sumatra, Banka, Billiton, Riouw, Lingga, the Malay
Peninsula and Anambas-Natoena Islands). It is in common
use over the whole of the Netherlands Indies, originally on
account of the seafaring habit of the Malays, who introduced
their language along all coasts. Hence, the extremely wide
distribution of the common Malay language. As soon as one
meets inland people, one finds local dialects and languages.

Already H. ZOLLINGER (2) had a good idea of the use of
the word Malaysia (Malesia in Latin and German). He rightly
proposed to indicate the whole of the tropical flora between
Asia and Australia (including the Malay Peninsula) by the word
Malaysia.

The Rev. J. E. TeENrson-Woops published in 1889 an essay
“On the Vegetation of Malaysia’’ (3). This is principally con-
cerned with the Malay Peninsula, but it is clear from the text
that he used the term Malaysia in a wider sense, though he did
not define it exactly. He consistently used the term Malay
Peninsula when referring to that area only, and other resident
botanists have usually done the same. ‘They tend to avoid the
use of the word Malaya, but the term British Malaya is sometimes
used. Recently H. J. LAm (4) used the latter term, but in the
same article used also the term Malay Peninsula.

H. N. RIDLeEy (5) used in general the term Malay Peninsula
or Malaya for the Peninsula south of 7°, and Malay Islands for
Netherlands India. Most other authors however follow Zol-
linger’s proposal, made 80 years ago, and repeatedly advocated
by E. D. MERRILL (6), H. BANcRoFT (7), myself (8), H.J. Lam
(4), and others.

F. N. CHASEN (9) uses the term Malaysia to comprise only
the Sunda shelf (Malay Peninsula, Borneo, Sumatra, Java, with
the intervening small islands); BoDEN Koss also used the
term in that sense.

This is in slight contrast with the terms used by
ZOLLINGER -'(2); Merrit. (6), and,-l..,.H..) BURKILL, (ro), who
call the Sunda shelf region Western Malaysia, in contrast to the
Sahul shelf which they indicate as Eastern Malaysia. It must be

Mol LX. (1937,).

188

said, that ZOLLINGER did not include the whole of New
Guinea in his Malesia, but only its westernmost part. This is

doubtless to be ascribed to the inadequate knowledge of the

Papuan flora at the middle of the 19th century.

Several botanists, e.g. BECCARI in his ‘‘Malesia’’ exclude
New Guinea from Malaysia, but include Assam, Burma and
the Andaman Islands. The same view is held by ENGLER
(rr), WARBURG (12) and others who include in Malaysia the
Indo-Chinese Peninsula, in contrast with ZOLLINGER (2),
BURKILL (10) and RIDLEY, who take the NW. boundary of
the true Malaysian stock at about 7° N. across the Siamese
isthmus, where a rather abrupt change in flora is indicated.

Though nowhere in the world can absolute, pure and abrupt
floral regions be distinguished, and gradual transition is the
general rule, there is good reason to speak of an Indo-Malayan
flora, comprising S.E. Asia and the Indo-Malayan Archipelago,
which falls into two subregions; the Indian flora and the
Malaysian flora, the latter having a common stock distributed
from the Malay Peninsula as far as New Guinea. According to
HOOKER, BENTHAM and others the Australian flora is also
a modified part of the Indo-Malaysian stock but this dates back
to the far past and Australia is to be distinguished nowadays as
a separate centre.

Uniformity in the use of the terms, both geographically,
historically and phytogeographically, is much to be desired.
There are ample reasons for the following definitions :

MALAY PENINSULA, comprising the Malay States and Straits
Settlements, is a topographical term. In phytogeographical
sense its N. boundary is situated at ca. 7° N.

MALAYSIA includes the whole of the Netherlands Indies,
Philippines, the Malay Peninsula, Sarawak, Br. N. Borneo,
Brunei, Portuguese Timor, Christmas Island, and the whole of
the island of New Guinea (Papua). Itis a biogeographical term.

MALAYA is according to Mr. FurTApDo and myself a word
which is ambiguous and likely to cause confusion, especially to
non-British botanists. [There have been made several attempts
to restrict it to mean ‘‘Malay Peninsula’’, but even today writers
of local history and folklore have to define the word to show
what they mean. Only a few years ago the term ‘‘Malaya’’
included Sumatra, Borneo, Rhio Archipelago, etc., so that the
use of the term ‘“‘British Malaya’’ was not uncommon and was
used even on the survey maps of the Peninsula, and there are
still books in the Malay Peninsula like ‘‘Official Guide to British
Malaya’. For phytogeographical purposes it would be better
to use the word Malay Peninsula and to avoid on the whole the
single term ‘‘Malaya’’ which can only give rise to confusion.

Gardens Bulletin, S.S.

189

Literature mentioned :—(1) J. Linn. Soc. Lond. Bot. 33, 1897, 2.
(2) Natuurk. Tijdschr. Ned. Ind. 13, 1857, 293-322. (3) Proc. Linn.
Soc. N.S.W. ser. II, 4, 1889, 9-108. (4) Gard. Bull. S.S. 9, 1936, go.
(5) Journ. Bot. 1935, 13; 1034, 251; 1932, 101; 1935, 342; Kew Bull.
79033, 190, etc.,! (6). Philipp.’ J: Sc. #2)11923;, ¥;° Enum. Philip.
pk 4)-D. ii, mote tr (7) Geol. For.-Steckh. Forh. 55, 1933, 80,
note 3. (8) Bull. Jard. Bot. Btzg. 13, 1933, 136, note 1. (g) Bull.
Raffl. Mus. 11, 1935, p. 1. (10) Gard. Bull. Str Settlem. 3, 1923, 21.
(11) Versuch einer Entwicklungsgeschichte, etc. vol. 2, 1882.
{12) Verh. Ges. Deutsch. Naturf. & Aerzte 1890, Allg. Teil.

Vol. IX. (1937).

190

THE GENUS LOMAGRAMMA
By R. E. Hoirrum

Historical.

ae
.

Biology and Morphology.

Essential characters of the genus.
Distribution, and status of species.
Excluded species.

Keys.

Description of species (alphabetical order).
Species dubia.

Se Sho a ee

Doubtful specimens.

i.) HS LORICAL

Among the curious groups of climbing ferns of the Eastern
tropics, Lomagramma seems to have attracted little attention,
and there is no general account of the genus, chiefly perhaps
owing to the inadequacy of collections. The various species,
as indicated below, have been described by various authors at
different times, and in many cases descriptions are very incom-
plete, so that a comparison based on published descriptions is
quite impossible.

My interest in this genus was aroused when I undertook the
study of Stenochlena and the genera confused with it, all of them
climbing ferns of similar habit to Lomagramma. I studied these
ferns in the field in many places in the Malay Peninsula, and
became also acquainted with all the stages of the life history
of the two Peninsula species of Lomagramma. I then realised
that these climbing ferns (except Stenochleena proper) have two
distinct leaf-forms, produced in response to the two different
environments occupied by the plants: the moist shady lower
levels of the forest, and the lighter and more airy upper regions
reached by the climbing stems. This distinction has not been
fully appreciated by most authors, and the low-level fronds have
usually been ignored, or regarded as ‘‘abnomal’’. ‘These fronds
clearly play an important part in the life of the plants; they have
distinctive characters and are important diagnostically. Having
discovered these facts by observation of the local species, I feel
that I am perhaps in a better position than others to deal with
this genus, and have accordingly attempted to piece together an
account of it, based upon the available specimens.

The genus Lomagramma was founded by John Smith in
1841 (Journ. Bot. 3: 402). He cited the species L. pteroides,

Gardens Bulletin, S.S.

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|

IQI

which he based upon Cuming’s no. 223, collected in the island of
Luzon, and his description is as follows :—

The single species upon which this genus is founded has the
habit of Stenochlzna, but differs in the venation being reticulate.
It is distinct in habit from the following, and in the sporangia
forming a broad marginal line or sorus; and therefore in that
respect partaking of the character of Lomaria, but differing in
having reticulate veins, and being without a special indusium.

It is curious that the author should have happened to take as the
type of his genus a species which is unique (so far as present
knowledge goes) in having a marginal sorus instead of fully
acrostichoid fertile pinnee, and should have based the name of
the new genus on that very character.

In his next paper (l.c. 4: 152), John Smith gave a further
diagnosis of the genus, and added ‘‘presumably this fern may be
the Leptochilus lomarioides of Blume, but his description is too
brief to enable me to determine with certainty.’’ The remark
about Blume’s description is correct. An examination of the
types of both species, and of a number of other specimens, leads
me to believe however that they are quite distinct; they differ
constantly in various characters at all stages of their life history.

John Smith overlooked a yet earlier species, Aspidium
sorbifolium Willd. ‘This species, based on a young plant, was
ignored by most later authors. Christensen in his Index Filicum
doubtfully identified it with Lomagramma polyphylla Brack.
Ching, examining the type in the Berlin herbarium, considered
it to be identical with Leptochilus lomarioides Bl., but I give
below reasons for believing him to be mistaken. In my opinion,
Aspidium sorbifolium is one of the species L. sumatrana,
L. grosseserrata and L. Mathewii, but I do not think it is at
piesent possible to say which. Therefore I leave it as dubious
species.

L. pteroides and L. lomarioides were lumped together by
Hooker in Species Filicum under the name Acrostichum
Blumeanum (Chilolepton Blumeanum Fée, based on specimens
from Java). Hooker also included a specimen from Samoa, which
1 now describe as a new species L. cordipinna.

The next well founded spccies was L. polyphylla Brack.
(1854), a very remarkable outlier in Fiji, having bipinnate fronds.
Twenty years later Baker described another peculiar species
from the same region, L. cultrata (Solomon Islands), having
deeply lobed pinnze, both sterile and fertile; this still remains
known only from the original collection.

Beddome came next with Lomagramma perakensis (1892),
a very distinct species from the mountains of the Malay
Peninsula. Copeland distinguished the Bornean L. Brooksii in
1908; but he appears to have misunderstood the Philippine species
in making two new varieties of L. pteroides the same year. Van
Alderwerelt van Rosenburg, recognising for the first time that

Pals iks (tesz7)s

192

there were two species of the genus in Java, described
L. abscondita in 1922; unfortunately he omitted to examine the
type of L. lomarioides, and the type of L. abscondita is identical
with L. lomariodes, so that the second Java species still lacked a
name. Van Alderwerelt described also L. suwmatrana in the
same year; this species occurs also in the Malay Peninsula.

In 1920 Brause described L. novoguineensis, a very distinct
species from New Guinea. Two years later, Christensen
described L. sinuata, from a specimen from Celebes; this species
is in my opinion represented also in Borneo, East Java, and New
Guinea, and our knowledge of it is increased by a recent collection
from Java, including bathyphylls, kindly sent to me by Dr.
Posthumus. L. melanolepis was described by van Alderwerelt
from Ternate in 1922; it is probably closely allied to L. sinuata.

In 1930 Ching described Campium Matthewii from south
China. ‘The specimens were clearly bathyphylls of a Loma-
gramma. Ching subsequently recognised the genus, and
regarded the species as identical with Aspidium sorbifolium
Willd. This however appears doubtful, and I am still in doubt
about assigning acrophylls to L. Mathewii.

Among the specimens I have examined, there are in my
opinion four additional species which I here describe as new:
L. Copelandii, a common Philippine species which has hitherto
not been clearly distinguished from L. pteroides; L. Merrillii,
an interesting small species from Mindanao; L. cordipinna from
Samoa; and L. grosseserrata from Siam. ‘The lowland species
from Java remains unnamed, as it may prove to be identical with
one of the species already described. It is likely that there are
other distinct bipinnate species in Fiji and neighbouring island
groups.

Among the specimens also are several which cannot
definitely be assigned to any of the known species, but which
are not adequate to serve as types for new species. It is possible
that some of them represent bathyphylls of known species, but
there are no definite connecting links. A list of these specimens
is given at the end of this paper; they are from Celebes, Luzon,
New Guinea, and Tahiti.

Ching has included in the the genus a species from South
Ameiica, Leptochilus guianensis (Aubl.) C. Chr. I give else-
where in this paper my reasons for excluding this species from
Lomagramma, which thus appears as a genus extending from
Assam and southern China through the Malayan region to the
Pacific. Within this region occur also other species which have
by some authois been regarded as belonging to Lomagramma
but are here excluded, namely Polvbotrya Wilkesiana Brack.
and its allies; these I regard as allied to Teratophyllum.

Gardens Bulletin, S.S.

0

193

2. BIOLOGY AND MORPHOLOGY

The species of Lomagramma are very interesting biologically.
They are climbing plants, starting life (in all recorded cases) on
rocks beside streams in shady forest, and later climbing high
up large trees. As with some other climbing plants, the fronds
they hear in their low creeping condition are different from those
borne by the high-climbing stems. In an account of Terato-
phyllum, I proposed the name bathyphylls for the fronds
produced in the low levels of the forest. A term is also needed
for the high-level fronds; these will be called acrophylls, a term
suggested to me by Dr. Christensen. ‘There is sometimes more
or less of a transition between the two, and there is a further
difficulty that the bathyphylls are not all similar, those borne
by the very young plants naturally being smaller than those
produced later. There is however a real distinction between the
two. A plant may go on creeping on the rocks for a considerable
period with a quite slender rhizome and fronds which do not
increase beyond a certain size. When it starts to climb a tree,
the rhizome increases very quickly to a much greater thickness,
and bears the much larger acrophylls. ‘The production of fertile
fronds appears in most species to be rather infrequent, or
possibly seasonal; at least in the Malay Peninsula, where we
have relatively slight seasonal change, fertile fronds have rarely
been found. Probably, as in Stenochlena and other ferns, the
fertile fronds are produced during dry seasons. I once found
fertile fronds of L. sumatrana on a plant which had been suddenly
exposed by the fall of a neighbouring tree; this may have had a
similar effect to that of a dry season. ‘The spores are very thin-
walled, without perispore. They appear unlikely to be able
to withstand dessication, and the character may be connected
with the streamside habitat of the plants.

The acrophylls of all species except L. polyphylla, and
L. cultrata are simply pinnate with almost entire pinne. ‘They
are thus all very similar, differing in rather small characters, and
some not very easily distinguishable. On the other hand, the
bathyphylls are usually much more distinct, and from my field
observations of the Peninsula species, and herbarium study of
the large number of bathyphylls of the unnamed Java species
which I have received from Buitenzorg and Pasoercean, I am
convinced that the bathyphyll characters are good and constant.
It is of course necessary to compare bathyphylls of similar size.

This state of affairs is almost exactly paralleled in Terato-
phyllum, and as in Teratophyllum, the earliest stages of growth,
bearing immature fronds, are often quite distinctive. Full-size
bathyphylls have usually all the pinne articulated; but in some
species the earlier fronds, of smaller size, have the apex not
articulated, but consisting of a narrow lobed lamina continuous
with the rachis and different from the lateral pinnze. I believe
this character to be a good one; it certainly is so in the Peninsula

Foot. BX «'01937)).

194

species, which are the only ones of which I have examined all
stages in the field. The larger bathyphylls have pinne which
are characteristic in number and spacing, size and shape, characters
of base, margins, apex, texture and venation. The marginal
teeth are more pronounced than in the acrophylls. I have not
seen bathyphylls of all species; and in some cases, where bathy-
phylls have been collected separately, I am uncertain to which
species they should be ascribed. I have not described them
definitely unless I am sure of their species.

3. “ESSHNITAL CHARACTERS “Or “THE GENGe

The essential characters which are common to all species of
the genus are as follows:

Rhizome dorsiventral, when creeping slender, when
climbing much stouter; the vascular system in section a single
ring with several leaf-gaps, the ventral vascular strand (bearing
the roots) double; fronds rather distant, especially on the
creeping rhizome; surface with several longitudinal ridges, with
grooves between them. The ridges are pale in colour in the
living rhizome. ‘They ate decurrent continuations of the bands
of aerating tissue on either side of the stipe; beneath them are
gaps in the coruical sclerenchyma.

Scales abundant on younger parts of rhizome, and on young
stipes, rachis and lamina, all similar in structure but differing in
size according to their position; the base of the scales (especially
the smaller ones) more or less bullate, the apex narrowed and
sometimes acuminate, the cells usually translucent, with thin
dark lateral walls; occasionally some of the larger rhizome
scales not translucent (L. sinuata and L. melanolepis).

Stipes with an open ring of vascular strands, to at least 18
in number.

Rachis with anatomy as stipe; more or less winged towards
the apex, the wing sometimes almost absent in the largest fronds,
and often present throughout the rachis in the smallest.

Sterile pinne all articulated to the rachis (except the apex
of small fronds of some species); the base rather unequal; the
margins more or less toothed; venation without any main lateral
veins, consisting of several series of areoles, the costal ones
usually largest, without free veins (except sometimes at the
margin). .

Fertile pinne articulated, usually much narrower than the
sterile; the lower surface, except for the midrib, entirely
covered with sporangia (except in L. pteroides where the
sporangia are confined to marginal bands towards the base of
the pinnze), the sporangia when young accompanied by peltate
paraphyses of the same structure as the scales; the venation as
in the sterile pinnee but the areoles smaller, usually clearly
visible on the upper surface.

Gardens Bulletin, S.S.

195

Spores thin-walled, without perispore or tubercles, usually
quite transparent.

The most distinctive of these characters are the scales, the
venation, and the spores.

a DISTRIBULION AND “STATUS OF SPECLES

So far as I can understand, no one species of Lomagramma
is very widely distributed geographically as compared with the
others. Our knowledge of the geographical distribution may
be summarised as follows:

ASSAM, TONKIN & S. CHINA. LL. Matthewii.

SIAM. L,. grosseserrata.

MALAY PENINSULA & SUMATRA. L. perakensis,

L. sumatrana (L,. lomarioides in S. Sumatra).

JAVA. JL. lomarioides, lL. sp., L. sinuata.

TERNATE. IL. melanolepis.

BORNEO: <1); Brooks; 4. sinuata

CELEBES: Ly sinwata:

NEW GUINEA. L. sinuata, L. novoguineensis.

PHILIPPINES. L. pteroides, L. Copelandii, L. Merrillii.

SOLOMON: ISLANDS. .: 1. cultrata:

NEW HEBRIDES. L. polyphylla?

FIJI. L. polyphylla, L. cordipinna.

TONGA. IL. polyphylla?

SAMOA. Ly. cordipinna.

TAHITI. Species unidentified, new?

It is evident that much still remains to be discovered
regarding these species, especially as regards their distribution.
Collections are on the whole meagre, no doubt due to collectors
being discouraged from collecting sterile materia’. Sterile
fronds of Lomagramma (bathyphylls and acrophylls) would
usually enable a species to be determined, and would be
invaluable as indications of the distribution of the species,
and of the constancy or otherwise of characters of the
bathyphylls. They would also give definite correlation between
the bathyphylls and acrophylls, which is lacking or doubtful in
some species.

The species of Lomagramma are certainly closely allied
together but I believe that they are distinct, and that sufficient
mate1ial is available for an account such as the present paper
to be of value. It is clear however that to characterise the species
fully a description of all stages of their development is necessary.
Some species are most easily distinguished in their young stages

Vol. IX. (1937).

196

ot bathyphylls, some in their sterile acrophylls, and some in the
fertile fronds.. Therefore we shall not have a full understanding
of the genus until we know all species in all stages, and we are
still very far from this. For the same reason I find it very
difficult to construct a key to the species.

Here is one of the many cases in which field observation is
essential to supplement herbarium study, and if this paper should
reach any persons who may be in a position to undertake such
field study, or to instruct collectors, I would appeal to them to
bear in mind the case of Lomagramma.

5. EXCLUDED SPECIES

L. guianensis (Aubl.) Ching, Amer. Fern Journ. 22: 17.
1932.
Leptochilus (Aublet) C. Chr. Index, cum syn.

I examined a specimen of this species at the British
Museum. ‘The spores have a large perispore, which I have
never found in any species of Lomagramma. Further, the
apical p nna is not articulated, but is continuous with the rachis.
The venation is connected to typical Bolbitis venation by
Bolbitis serrata, which has the pinnz not articulated. In the
British Museum Herbarium is also a specimen from Mexico
(C. A. Purpus no. 7246); labelled Leptochilus Turckheiimii
C. Chr.; this has main lateral veins as in Bolbitis, but pinnze
articulate. It is evident therefore that the American Bolbitis
species have produced a number of variants from the typical
condition both in venation and articulation of pinne, and I
suggest that Lomagramma guianensis (Aubl.) Ching is only a
rather unusually aberrant species of Bolbitis which simulates the
condition found normally in Lomagramma.

Lomagramma Wilkesiana (Brack.) Copel. Phil. Journ. Sci.
aC 7 Ae. TOOS.

L. articulata (J. Sm.) Copel. lc. L. bitpinnata Copel.
Phil. Journ. Sci. 11¢: 114. 1916. "These three species, and perhaps
also Lomariopsis Balanse Fourn. (Ann. Sci. Nat. V, 18: 271.
1873), evidently all belong to the same genus, but I do not think
that they are members of the genus Lomagramma. ‘They have
spores with perispore, and their venation is that of Teratophyllum,
not Lomagramma. I have not yet been able to examine
sufficient material of the species to understand them well enough
to found a new genus for them, but I believe that this should be
done when the material is forthcoming. The species are in my
opinion closely allied to Teratophyllum and could possibly be
included in that genus; to do this would necessitate an emenda-
tion in the definition of the genus which I gave in this Bulletin,
Vol. 5, p. 277, a new section being constituted for the species
under consideration.

Gardens Bulletin, S.S.

197

6; EKEVS

Owing to the fact that the species of Lomagramma in some
cases differ most strikingly from each other in one only of the
three leaf-forms, and also to the fact that in a few cases
bathyphylls are still unknown, it is hardly possible to make a
satisfactory key. It appears to me more useful to summarise
briefly the distinctive characters of the various species under the
headings bathyphylls, sterile acrophylls, and fertile fronds;
whichever frond-type is then available, this summary should at
least locate it within a small group of species, the detailed
descriptions of which will provide further information.

BATHYPHYLLS

Pinne of largest fronds 25—30—jugate;
apex of frond continuous with rachis,
except in some of the largest fronds.

pinnze entire L. lomarioides
pinne toothed
pinnz to about 2.5 cm. long, drying
reddish A ... L. pteroides
pinnee to 5 cm. or more long, not
reddish when dry L. grosseserrata
L. Matthewit
L. sumatrana
Pinne of largest fronds 10—18—jugate;
apex of frond always an articulated
pinna, except in young plants of some
species
pinnee entire... A ... L. perakensis
pinnee toothed
margins shallowly lobed, lobes
about 5 mm. wide, each with
a—4, teeth L. Copelandiu
L. sb. (Java)
margins not lobed
upper base suddenly subtrun-
cate from a narrow beginning;
lower base very marrow; apex
blunt bad ve ifes ee Siwata
base unequally cuneate; edges
sharply serrate; apex acute ... L. Merrillii

upper base broad subtruncate;
lower narrow; edges toothed
towards apex; apex in small
fronds somewhat rounded ww! BL. Brooksa

Bathyphylls unknown in L. cordipinna,
L. cultrata, L. melanolepis, L. novo-
guineensis; incompletely known in L.

bolyphylla.

moe £ xX. { TH37).

198

STERILE ACROPHVLLS
Bipinnate
Pinnz lobed to costa

Pinne at most shallowly ished
Pinne, at least of old fronds,
coriaceous, veins not distinct
Stipes 4 cm. long a
Stipes at least 15 cm. long
Pinne to 26 cm. long
Pinnz not usually more than
72\5°em. long
Pinnze ee
Pinne closely
towards apex
Pinne thin in texture, the veins ee
and distinct

Pinnz with a winged stalk to r cm.
long

Pinnz hardly eles
Base of pinnz cordate, to 2.9 cm.
wide
Base of pinnze at best subcordate,
narrower
Edges deeply crenate-serrate
Edges not more than slightly
toothed
Pinnzeto.aae by “173 ce
Pinnze larger, usually to 2
cm. wide
Pinnz to 18 by 3 cm.
edges not toothed
Pinnee smaller, edges
slightly toothed

sub-

toothed

FERTILE FRONDS

Fronds bipinnate
Pinnze lobed to costa

Pinne entire

Base of pinnz always dilated, freile at

the edges only
Base of pinne not dilated, whole lower

surface soriferous
Widest pinne 8 mm. or more wide
Pinnze about 4.5 cm. long, apex
rounded

Gardens Bulletin, S.S.

BS
. cultrata

polyphylla

L. Brooksii

L. perakensis

. lomarioides

L. Merrillii

Pe tae

. sinuata

. cordipinna

. grosseserrata

novoguineensis

. Copelandit

. Matthew
. melanolepis
. sumatrana

sp. (Java)

polyp hylla
. cultrata

. pteroides

L novoguineensis

“

oy

» New Te
pp liz te
Sn at
¢

Gard. Bull. S.S. Vol. 1X. Plate 8.

Lomagramma borneensis. Above: apex of bathyphyll from young
plant (Penrissen, Shelford). Below: apex of full-sized bathyphyll,
showing articulated pinna (Holttum 25251). Both x 1.5.

199.

Pinne usually not less than ro

cm. long, apex acute L. cordipinna
L. grosseserrata
L. Matthewii
L. sinuata
Widest pinnze to 5 mm. wide
Pinnze with stalks at least 3 mm.
long
Pinnze about 9 cm. long... L. Merrillii
Pinne at least 15 cm. long L. perakensis
Pinnze almost sessile
Pinnee I-2 mm. wide when
dry L. lomarioides
Pinnee mostly Saar L. Brooksii
L. Copelandii
L. melanolepis
L. sumatrana

7. DE SCRIF LION. OF “THE . SPECIES

In the following pages I have usually given the original
description of the species, with an amplified description of my
own, based on all material which I have examined. Critical
remarks and notes on the diagnostic characters are also included.

For permission to examine specimens, and for loan of
material, I am indebted to the heads of the following Herbaria :
Buitenzorg (B), Manila (M), U.S. National “Herbarium,
Washington (W), Kew (K), British Museum (B.M.), Calcutta
(C), Pascercean (Pas.). The herbaria are referred to by the
initials in brackets after the citation of specimens; Singapore
herbarium is indicated by the letter S. I am also indebted to
Prof. H. J. Lam for the loan of the type of Leptochilus
lomarioides Bl. and to Dr. L. Diels for a photograph of the type of
Aspidium sorbifolium Willd.

Lomagramma Brooksii Copel. Phil. Journ. Sci. 3c: 345. 1908.
Also l.c. Te: 60. 1912. Plate 8.

Rhizomate alte scandente, stipiteque 4 cm. longo paleaceis ; rhachi
deorsum paleacea, sursum fere glabra et anguste alata; pinnis
sterilibus ca. 8 cm. longis, 15 mm. latis, fere integris, coriaceis, nisi ad
costas glabris, basibus truncatis, apicibus falcatis breviter acuminatis ,
venulis immersis, areolis costalibus parvis, lamina viride, venulis
infra rubellis; pinnis fertilibus plus falcatis, ca. 3 mm. latis, basibus
dilatatis non auriculatis.

Sarawak, Bongo Mountain, leg. Hewitt and Brooks.

Near L. pteroides var. subcoriacea and L. perakensis Bedd. In
the specimen sent me the pinnz are sterile throughout the most of
the frond, the apical ones being fertile. The occurrence of sterile and
fertile pinne on the same frond is hitherto unknown in the genus.
It is possibly abnormal in this case, but I believe the species is
sufficiently distinct without this character. —

Vol. IX.. (1937).

F

200

In the latter publication above cited Copeland writes: Mr.
BROOKS has sent me a very complete specimen, collected at Bau.
It has the fertile and sterile fronds distinct, as in other species of the
genus. The frond of a juvenile plant ‘“‘which germinates on the
limestone and may grow in large masses before reaching a trunk,
and becoming scandent’’, has membranaceous, serrate, and very
oblique pinne. Fronds of adult plants are as described, except in
size. The sterile fronds are a meter or more long and 30 cm. wide.
The fertile frond is still wider, its pinne 2 to 4 mm. wide, and
straight or curved.

The species is nearest to Lomagramma perakensis Bedd, differing
conspicuously in the very short stipes and persistent scaliness.

I have not seen the type of this species, but I have seen a
considerable number of other specimens from Borneo which
clearly belong to one species and agree pretty well with
Copeland’s description, except that I have seen none so large as
indicated by Copeland’s note of 1922 and I would not describe the
pinnee of bathyphylls as very oblique. ‘The following description
is based on specimens examined by me.

Immature Bathyphylls. Stipes to about 10 cm, fronds up to
about 20 cm. long and 7 cm. wide with 12-15 pairs of pinnz;
apex continuous with the rachis; the lower pinnze at right angles
to the rachis, the upper oblique; pinnz close, largest usually
about 4.5 cm. long and 1.5 cm. wide; the upper base broad
subtruncate, sometimes slight!y auricled; lower base very
narrowly cuneate or narrowly rounded in lowest pinnee; midrib
fulcate towards apex; edges toothed towards apex only, apex
rounded in youngest plants, usually acute in older ones, width
greatest about one third from apex. ‘Texture thin; veins distinct
and slightly raised on both surfaces, in 2-3 series of areoles,
costal areoles not much larger than others.

Largest bathyphylls. Stipes to 10 cm. lamina to 4o cm.
long and 12 cm. wide, pinne to at least 18 pairs, uppermost
pinna articulate and like the others, largest pinnee about 6 cm.
long and 2 cm. broad, lower base rounded, much narrower than
subtruncate upper base, margins toothed towards falcate acute
apex.

Sterile acrophylls. - Stipes usually about 4cm. long. Fronds
(Copeland’s specimen to 1 1m. long and 30 cm. wide; largest speci-
men below cited 75 by 20cm. Largest pinne in middle third of
frond, upper and lower gradually reduced. Middle pinne at
right angles to rachis, to about 2 cm. wide, sessile, lower base a
little narrower than the upper, and coming closer to the rachis,
rounded to subcordate; upper base subtruncate; edges entire,
sometimes wavy especially towards apex, but not toothed; apex
acute, gradua'’ly narrowed from widest part of the pinna which
is usually one third from apex. ‘Texture thin but very firm; veins
in three series of areoles, slightly raised and usually quite
distinct on both surfaces. The lowest pinnz usually slightly
stalked, the base equal and subcordate.

Gardens Bulletin, S.S.

Gord: Bull, Ses. Vole tae Plate 9g.

Ses
&

SUP

&

ATA

ST ROOT

cs

¥

UG

|
|

Lomagramma Copelandii, Type, showing the sessile narrow fertile
pinnze, with base not dilated) 2/3 mat, ‘size,

201

Fertile fronds. In the specimen seen by Copeland these are
more than 30 cm. wide, the pinnze 2-4 mm. wide. The only
fertile fronds seen by me are those of Clemens from Penibukan,
Kinabalu; these have stipes to 10 cm. long, and lamina to about
30 cm. long and 10 cm. wide. The pinne are up to 7 cm. long
by 2 mim. wide, on winged stalks 2 mim. long; the veins usually
distinct, the costal areoles elongate, the others almost round.

BORNEO. Kinabalu: Tenompok, 5,000 ft., Clemens
23038 (B); near Dallas, 3,500 ft.,.S.F.N. 25251, Holttum (S);
Penibukan 4,000 ft., Clemens 30636 (M), 5,500 ft. Clemens
50263 (M). Exp. Nieuwenhuis, Jaheri 574 (B). N. Borneo
Grens Comm. Amjah 517 (B), 525 (B), 417 (B), 585 (B).
_ Penrissen, Sarawak, Shelford, s.n. 1899 (S, very young).

CELEBES. Menado, afd. Donggala, Kcelawi, 1000 m.
Posthumus 2335 (Pas).

This species has been found in various parts of Borneo, and
I believe also in N. Celebes; the Celebes specimen consists of
rather large bathyphylls, which I cannot assign to any othei
species. The occurrence of the species in north Celebes,
separated by a relatively narrow sea from B. N. Borneo, is not
unlikely.

L. Brooksii (as here interpreted) is certainly similar to
L. perakensis in its sterile acrophylls, differing in somewhat
smaller pinnz of rather thinner texture with more rounded base.
The bathyphylls of the two species are very different, those of
L. perakensis having quite entire pinne, the apical one articulate,
from a very early stage.

Lomagramma Copelandii Holttum sp. nov. Plates 9 & 10.
L. pteroides auctt. p.p. non J. Sm..

L. pteroides var. negrosensis Copel. Leafl. Philip. Bot. Il:
$032; 1908:

A lL. pteroidei differt: pinnee bathyphyllorum maximoruin
Cc. 15-jugate, ad 6.5 cm. longe, 1.8. cm. late, apicales articulate;
- pinnee acrophyllorum sterilium ad 18 cm. longe, 3 cm. late, basi |
superne late cuneate inferne rotundate, textura tenues; pinnz
fertiles ad 20 cm. longee, 4 mm. late, sessiles, basi non dilatate.

Bathyphylls. Stipes from 2 cm. long in fronds of 12 cm,
to 18 cm. long in fronds of 40 cm. In fronds to about 15 cm.
long apex a narrow lamina continuous with the rachis; in full-
sized bathyphylls, apical pinna like others and articulated, but
smallest. Bathyphyll pinne to about 15-jugate, largest seen 6.5
em. long, 1.8 cm. wide; base unequal, upper broadly cuneate,
lower narrower, rounded except in lowest (somewhat reduced
and reflexed) pinnee in which lower base is narrowly cuneate;
edges broadly crenate (lobes to 5 mm.), each lobe with 2-4 short
teeth, largest pinnze toothed only towards apex; apex not falcate,
acute in pinneze of larger fronds, blunt in pinnz of smaller

Pol oAX. (1937).

202°

fronds; texture thin; veins slightly raised on both surfaces, areoles
in two series, costal ones usually largest.

Sterile Acrophylls. Stipe 15-25 cm. long. Pinne to 18
cm. long, 3:cm. wide, sessile; upper base usually broadly cuneate

to subtruncate, more narrowly cuneate in the upper pinnz; lower

base usually rounded; edges slightly undulate, not toothed; apex
acuminate, hardly falcate.

Fertile pinne to about 20 cm. long, 4 mm. wide when dry,
sessile, the base not dilated, whole lower surface except midrib
soriferous, areoles long and narrow.

TYPE. Luzon: Mt. Binuang, Prov. Tayabas, Bur. Sci-

no. 28826, Ramos and Edano, in Philippine National Herbarium
(duplicate in Herb. Buitenzorg).

This species differs very strikingly from L. pteroides in the
various characters mentioned in the latin diagnosis. The
bathyphylls with their large broad pinnz are very distinct, being
nearest to those of the unnamed species from Java; the pinnz of
sterile acrophylls are very broad and thin in texture, never sub-
cordate as is often the case in L. pteroides; the fertile pinnze
are never dilated and partly sterile at the base, and are practically
sessile.

L.. pteroides var. negrosensis seems to me to belong here, and
probably consists of rather small plants of this species, perhaps
from a shady place. The specimens show fronds on a rather
slender rhizome, the pinne about intermediate in size between

those of the largest bathyphylls and of the typical acrophylls, ©

and somewhat toothed towards the apex.

L. Copelandii, like L. pteroides, is apparently distributed
throughout the Philippine Islands. The specimens examined
are as follows.

Luzon: Irosin (Mt. Bulusan), Prov. Sorzogon, Elmer 16919
(B, M.); Los Banos, Laguna, Copeland s.n. Mar. 16, 1906 (M,
bathyphylls); Infanta, Tayabas Prov. Bur. Sci. no. 9331, C. B.
Robinson (M, bathyphylls). Catanduanes: Bur. Sci. no. 30358,
Ramos (B, M). Samar: Catubig River, Bur. Sci. no. 24810,
Edano (M). Bohol: Bur. Sci. no. 42972, Ramos (M). Mindanao:
Surigao Prov. Bur. Sci. no. 34816, Ramos and Pascasio (M); Mt.
Apo, Cotabato Prov., Phil. Nat. Herb. no. 3244, Copeland (M);
Camp Keithley, Lake Lanao, M. S. Clemens s.n. Sept. 1907.
' Negros: Dumaguete (Cuernos Mts), Prov. of Negros Oriental,
Elmer 9678 (M, W, B; type of L. pteroides var negrosensis).

Lomagramma cordipinna Holttum sp. nov.

Bathyphylla ignota. Acrophylla sterilia ad 33 cm. lata;
pinnee mediz 3.5—5 cm. inter se remote, breviter petiolulate, ad
17.5 cm. longze et 2.9 cm. late, basi inzequaliter cordate vel

subcordatee, margine integre vel leviter undulate, apice |

Gardens Bulletin, S.S.

hee Moy

? .
eee

, ay ay -
" +
eS ee ee ee -—— <9 .

Gard: Bull. S:S2 Velo rx. Plate to.

Above
Below

: Lomagramma grosseserrata, pinne of bathyphyll.
: L, Copelandii, pinne of bathyphyll. Both x 1.5,

‘

203

acuminate; textura tenues; venulz prominentes, areolas 3-4
seriatas formantes; pinnee apicales minores, basi rotundatz apice
leviter serratee. Pinne fertiles petiolulate (petioluli ad 5 mm.
longi), ad 12 cm. longe et 1.1 cm. late, tota pagina sorifere.

SAMOA. Pango-pango Bay, W. E. Safford, March 1888,
U. S. National Herbarium (Type, 3 sheets). U.S. South Pacific
Exploring Expedition 1838-42, s.n. (W).

FIJI. Korumbamba, July 1932, A. Meebold 16856 (B.M.)

This remarkable species is well represented by the fine collec-
tions of Safford. "Though these include only the central and
upper portions of the fronds of acrophylls, the pinnz are so
distinctive that there is no doubt of this being a good species.
The cordate or subcordate base of the middle pinnze, combined
with the long-stalked and very broad fertile pinne (which are
completely soriferous) are the peculiar features of the species.

Of the Fiji specimen, I have seen only a single pinna which
I judge to be from the base of a frond; it is short and broad (8
em. by 3.3 cm.) with almost equally subcordate base and rounded
apex, the margins minutely irregularly crenulate towards the
apex.

Lomagramma cultrata (Baker) Holttum comb. nov.
Acrostichum cultratum Bak. Syn. Fil. 524. 1874.
Baker’s original description is as follows :

A. (Chrysodium) cultratum Baker; climbs on trees with fronds
6 ft. long ; barren pinne ligulate, 6-8 in long, 1-14 in broad, cut down

_to the rachis or a narrow wing into entire close blunt obovate lobes

3-4 in broad, cuneately narrowed to a broad base, or rarely produced

and oblong, often several of the lower and sometimes the central
lobes confluent; texture moderately firm; surfaces green, naked;
areole small, exappendiculate : fertile pinne and lobes like those of
the barren fronds, but the latter distant, 2-24 lin. broad.

Hab. San Christobal, Solomon Isles, H. Richards, in Herb.
Macleay.

The type sheet at Kew consists of pinnee only. Most of the
lobes are obovate, but some are almost evenly elliptical with a
narrow base, almost like separate pinnules but not stalked nor |
articulate; these elliptical lobes vary in length. The fertile pinnze
are 10-12 mm. broad, the lobes almost circular and rather distant.
The scales aie typical of the genus, and the spores also.

This species is referred to L. lomarioides in Christensen’s
Index Filicum. It seems to me a very distinct and interesting
species, a possible link between the pinnate and bipinnate species.
of the genus.

Lomagramma grosseserrata Holttum, sp. nov. Plates 10 & 11.

Species IL. sumatrana affinis, differt pinnis acrophyllorum
sterilum magnis, longe acuminatis, margine late crenato-serratis,
pinnis fertilibus ad 1 cm. latis.

iol. TX. (1937).

Fi

204

Bathyphylls. Stipes 8 cm. long (in the available specimen);
lamina 40 cm. long, Io cm. wide, lowest pinne slightly reduced,
reflexed and more separated, upper six pairs gradually reduced;
apex continuous. with the rachis. Pinnz 29 pairs, almost
touching (except the lowest), largest 5 cm. long and 1.3 cm. wide,

at right angles to the rachis, base only slightly unequal, upper -

base subtruncate, lower broadly rounded, apex slightly falcate
acute, edges shallowly crenate-serrate, the teeth 4-5 mm. apart;
texture thin translucent; venation in 2-3 series of areoles, the
costal ones largest, the veins distinctly raised; scales with bullate
base numerous.

Sterile acrophylls. Stipe to at least 20 cm. long. Pinne
about 3 cm. apart, to about 19 cm. long and 2.5 cm. wide, sessile,
the upper base broadly cuneate, the lower rather broadly rounded,
margins rather deeply crenate-serrate, especially towards the apex,
at intervals of 5-7 mm, the distal third gradually long-acuminate;
colour dark dreen when dry; texture thin; veins slender, raised,
brown on both surfaces, in about 3 series of areoles; scales
numerous.

Fertile fronds. Stipe 18 cm. long. Lower pinnz on stalks
to § mm. long, pinnze to 20 cm. long and I cm. wide, the upper
pinne gradually more shortly stalked and smaller (in the specimen
at hand the upper pinne are incompletely expanded); colour dark;
veins distinct on the upper surface, in about 3 series of areoles,
areoles more or less rounded except the costal ones; sporangia
covering the whole lower surface except the midrib.

TYPE. Siam: Pre, Me Sai, 620-880 m. Winit no. 1019
(27-11-22), in Herb. Singapore.

This species is clearly allied to L. sumatrana and L.
Matthewii, but it differs in the points mentioned in the diagnosis
so strikingly that I cannot but consider it distinct. "The bathy-
phylls are similar to those of the other species mentioned, but they
differ from those of L. sumatrana in their greater length, and from

those of L. Matthewiiin the greater number of pinne, very regular —

and closely placed. As regards the apex of the bathyphylls, the
specimens at hand has indications of articulation, but it is
a very large bathyphyll, and I believe that smaller ones would
have the apex clearly continuous with the rachis.

To this species should probably be referred the specimen cited
by Beddome on p. 106 of the Supplement to his Handbook
(Makum Forest, Lukhimpore, Assam). I have not seen this
specimen.

Lomagramma lomarioides (Bl.) J. Sm. Hist. Fil. 143. 1875.
Plates 12 & 13.

Leptochilus lomarioides Bl. Enum. 206. 1828.
Chilolepton Blumeanum Fée, Hist. Acrost. 89, t. 51. 1845.
Acrostichum Blumeanum Hk. Spec. Fil. 5: 268. 1864. p.p.

Gardens Bulletin, S.S.

Gard. Bull. S.S. Vol. 1X. Plate 12.

Lomagramma grosseserrata, Type; sterile acrophyll. 2/4. Mat. Size.

Gard. Bull. SiS. Velo TX. Plate 12.

Lomagramma lomarioides, specimen from Blume’s herbarium,
showing sterile and fertile acrophylls, 2/3 nat. size.

» 205

Lomagramma abscondita v.A.v.R. Bull. Jard. bot. Buitenz.
iisser: Dk: 26, 2613 sc1bids TW Iser..2: 159. i920.

This species was described by Blume as follows:

“‘L. frondibus pinnatis, pinnis sterilibus _lineari-lanceolatis,
fertilibus angusto-linearibus elongatis stipite rachique paleaceis’’.

By the courtesy of the Director of the Rijksherbarium at
Leiden, I have been able to examine Blume’s specimens of this
species. ‘There are four sheets, one bearing the name in Blume’s
hand, the other three all clearly from one collection (perhaps the
same collection as the type), and one of them labelled ‘‘Herb.
Dr. Bl.’’ Judging from the fact that Blume described the fertile
pinnee, which are not represented on the sheet actually bearing
his writing, it seems probable that all four sheets were actually
seen by him. In any case, they are quite certainly all of the
same species, and they agree closely with the type L. abscondita
v.A.v.R. and a number of other specimens from Java at Buiten-
zorg. I can therefore with confidence give a full description of
the species, except that there are only three specimens which could
be regarded as bathyphylls.

Bathyphylls. (description based on three collections from
Tjibodas: Fleischer 276, Burck s.n., Palmer & Bryant 1190).
Stipes to 15 cm. long, frond to 40 cm. long by 9 cm. wide; pinnze
to 25-jugate, the terminal ones articulate but smallest, with very
narrow base. Middle pinne to about 5 cm. long and 1.0 cm.
wide, the upper base broadly cuneate to subtruncate, the lower
base narrowly rounded, edges quite entire, often slightly sinuate
especially towards the apex, the apex falcate, acuminate. Texture
firm, veins hardly visible on the upper surface, slightly prominent
below, the areole rather small, usually in 2 rows. Palmer and
Bryant’s specimen from Salak represents a much younger stage,
stipes 22 cm., frond 21 cm. pinne 16 pairs.

Sterile acrophylls. Young stipes and fronds very scaly,
the scales thin and 1ather pale, mostly falling as the frond
developes. Stipes to 20 cm. frond to 1 m. long and 20 cm. wide,
the upper and lower pinnze gradually reduced, the lowest only
5 ecm. long, and more distant. Largest pinne to about 12.5
cm. long and 1.5 cm. wide, the upper base broadly cuneate
to truncate, the lower base narrowly rounded, occasionally the
upper base or both subauriculate. Pinnze usually widest about
the middle, apex gradually acuminate, falcate, the edges entire
or undulate, often inrolled when dry. Texture of old fronds
subcoriaceous, veins hardly visible on upper surface, usually
somewhat raised below but rather broad. Blume’s type has the
veins more clearly visible than most of the other specimens;
young fronds show the veins much more clearly than old ones?

Fertile fronds. Pinnze sessile, to at least 18 cm. long, 1-2
mm. wide.

Pol. 1X...°( 1937).

206

This species appears to be common on the mountains in the
west of Java, and there are also specimens from east Java which
I believe must also be referred to it, though they are not typical.
The characteristic features are the narrow rather coriaceous pinnze
of the sterile acrophylls, the very narrow pinne of the fertile
fronds, and the numerous narrow entire pinne of the bathy-
phylls. The sterile acrophylls are perhaps nearest to L. Brooksii,
but L. lomorioides differs markedly f:om that species in the
narrow fe1tile pinnze and the narrow pinnz of the bathyphylls.

TYPE: Java, Salak, Blume (Leiden).

Other specimens. Res. Preanger: G. Malang, Raciborski
s.n. (B, M); G. Rasamala, Raciborski s.n. (B), Hallier s.n. (M);
Tjibodas, Arsin 19624 (B), Burck s.n. (B), Bruggeman 438 (B),
Palmer & Bryant 1190 (W), 212 (W), M. Fleischer 276 (M);
Tjitibo, Tjidadap, Tjibeber, r000 m., Bakuizen 2402 (B); Tjadas
Malang, z.v. Tjibeber, 1000 m., Winckel 1699 (B); Mt. Salak
1ooo m., Palmer & Bryant 550 (W), 557 (W); 700 m., 700 (W).
Res. Besoeki: G. Idjen 1000 m., Posthumus 3774 (B); Weg v.
Lidjen halfweg Kawah Idjen, 1025 m., Posthumus 378 (Pas);
Mont. Tengger 1200 m., M. Buysman 108 (W).

Lomagramma Matthewii (Ching) Holttum comb. nov.

Campium Matthewii Ching, Bull. Fan. Mem. Inst. Biol.
1: 158, fig. 3, 1930.

This species was based on a collection of large bathyphylls
made by Dr. C. G. Matthew in Kwangtung. It was subsequently
identified by Mr. Ching (Lingnan Sci Journ. 12: 566. 1933) with
Aspidium sorbifolium Willd. ‘This identification however appears
very doubtful, as explained under Lomagramma sorbifolia at the
end of this paper.

I have seen Matthew’s specimen in the Hong Kong her-
barium, and judging from this and Mr. Ching’s description
I think it probable that certain specimens of bathyphylls from
Assam are to be referred to this species. Also from Assam, and
from Tonkin, are specimens of sterile acrophylls which are cer-
tainly not L. grosseserrata and perhaps belong to L. Matthewit.
The only fertile frond which might be associated here is on the
same sheet as one of the bathyphylls from Assam.

It is clear from the original collection of bathyphylls that
L. Matthewii is closely related to L. grosseserrata, and in the
absence of the quite distinct sterile acrophylls above mentioned,
I should be inclined to include it with that species; these acro-
phylls show however that another species is present, but whether
that species is L. Matthewii is another matter. The present
collections are in fact inadequate for more than a tentative
account of the species in the region between Assam and southern
China to be given. I here quote the original description of
Campium Matthewti, and add some notes about the other
specimens which I think may belong to the species. It is just

Gardens Bulletin, S.S.

Gord. Bull: S.S. Vol. EX Pigie ui.

Above: Lomagramma fpteroides, a bathyphyll.
Below: L. lomarioides, a bathyphyll. Both x 1.5.

- xe pas aber
Ae eo

a

; f LAr dy ;
a - an ik oe ee ae 4 y
FE oe co 4
eee Fa ee ‘

207

possible after all that the type of Campium Matthewti may
represent bathyphylls of L. grosseserrata.

Campium Matthewii. Rhizome wide-creeping, naked, fleshy,
light stramineous, terete, 5 mm. in diam.; stipes remote, pale green,
12-17 cm. long, terete below, deeply bisulcate throughout the rachis
above, moderately paleaceous throughout the rachis, palez small,
brown, membranaceous, lanceolate, acuminate; frond 35-50 cm. long,
10-13 cm. broad, linear oblong, slightly narrowed at base with the

_ apex of rachis often lengthening out into a stout naked prolongation

with proliferous tip, pinnate, pinnz close, 20-27 on each side under
the impari-terminal one, 7 cm. long, 1-1.5 cm. broad, gradually
decrescent upwards, the uppermost ones 4 cm. or less long, sessile
throughout, broadly linear-lanceolate, broadened downward, truncate
or cuneato-truncate, acute, margin shallowly crenate above the base,
texture thin papery, glabrous except the midrib which is sparsely
scaly; venation distinct, main veins wanting, veinlets pauci-anasto-
mosing in 2—3 rows of oblique irregular rhombic areolz, of which the
costal ones much the largest, extending to the half breadth of the
pinne, all without included veinlets, the marginal areole mostly
close, very rarely with excurrent free veinlets; fertile fronds not seen.

Distribution: Kwangtung: Lienchow, Dr. Matthew, No. 5108,
Dec. 1907.

There is one curious point in the above description, namely
‘‘apex of rachis often lengthening out into a stout naked prolong- -
ation with proliferous tip’’. This I have not seen in any
specimen of Lomagramma. ‘The apex of the specimen of the
type collection of L. Maithewii in the Hong Kong herebarium is
missing, so that I am unable to verify the matter.

Bathyphylls of other collections. ‘The following agree very
well with Matthew’s specimen, chiefly differing in the fact that
the costal areoles are not so very much larger than the others.

ASSAM. Wallich, s.n. (C, from T’. Moore’s Fern Her-
barium). Chima Forest, Garo Hills, 300 ft., Dec. 1890, Gustav
Mann (W).

Sterile acrophylls. I have seen one good collection from
Manipur (Meebold 6233, in Calcutta herbarium). ‘This has
pinne very like those of L. sumatrana, differing chiefly in the
very broadly cuneate bases of the middle pinne. The pinnz are
up to about 12 cm. long and 2 cm. wide, margins entire except
towards apex where they are rather sharply toothed.

There is another collection from Tonkin (Province de Hoa
Binh, vers 500 m., Avril 1926, U.S. Nat. Herb.) with rather
stout climbing rhizome, but the frond a rather intermediate one,
having apex continuous with the rachis as in typical bathyphylls.
It is in fact rather intermediate in character between the bathy-
phylls and acrophylls from Assam and Manipur above mentioned,
and forms the strongest link connecting these specimens together.

Fertile fronds. Mann’s specimen from Garo Hills, Assam,
above cited, has part of a fertile frond, attached to a typical
stout climbing rhizome, along with a large bathyphyll. ‘The

Vol. IX. (1937).

208

fertile frond has a stipe 28 cm. long, and pinne to 16 cm. long
and 8 mm. wide, on stalks 3 mm. long.

I am reasonably certain that the specimens above described
from Assam, Manipur and Tonkin are one species, but I do not
feel so sure that they are L. Matthewii. They are also rather
near L. sumatrana. ‘There the matter must be left until fresh
collections are made.

Lomagramma melanolepis v.A.v.R. Bull. Jard. bot. Buitenz.
ser. II], Ssi212., -¥922.

L. pteroidee J. Sm. affinis.—Squamz rhizomatis, stipitum,
rhachidum costarumque sparse, nigrz vel obscure castaneze. Frondes
steriles c. 60 cm. longe; rachis apicem versus anguste alata; ala ad
basin pinnarum superiorum brevius vel longius dentiformi-auriculata.
Pinne flaccide firmiter herbacez, subapproximatz, sessiles, patentes,
in sicco olivacez, supramedianz maxime, lineares, 12-13 cm. longe,
I1.75-2 cm. latz, integerrime vel apice acuminato, recto vel subfalcato
remote subserrulato-crenulatez, basi oblique et plus minusve inqe-
quilatere, rotundatz vel late rotundato-cuneate; venz distincte,
pallide (colore pinnarum). Frondes fertiles minores, usque as 40 cm.
longe, pinnis remotis, horizontalibus, contractis. Sori capsulis
juvenilibus paraphysis longe stipitatis, peltatis copiose intermixtis.

Fertile frond of the specimen on hand stili young, with the
pinnz 5 cm. long, 3 mm. broad.

TERNATE: Foramadjahi, in forest, 700 m. Beguin 1114 (B).

I have examined the type of this species. Like L. sinuata,
it has narrow black scales on the rhizome and on the very young
unexpanded fronds. Through an ordinary hand lens these scales
appear black with irregular narrow paler margins. ‘Through a
microscope, the central cells are seen to have very thick black
lateral walls, and the lumen is a brownish colour. The marginal
cells have thin walls and some of them are elongated to give an
irregularly fringed condition. The small scales on the frond
are few; they are rather dark, but of typical bullate form and the
cells normal.

The sterile pinne are hardly toothed, in which they agree
with L. sinuata; but they are much smaller than the pinne of
that species, and are sessile, lacking the peculiar basal characters.
The fertile pinne are up to 4 mm. wide (not 3 mm. as stated in
the description above quoted). They seem to be practically
mature and are almost sessile.

The distinctive characters therefore are the rhizome scales,
the medium sized almost entire sessile sterile pinneze and the
subsessile short and relatively broad fertile pinne.

At Ternate, this species falls within the range of distribution
of L. sinuata and is presumably a local derivative. Further
collections may possibly show it to have a range of variation
bridging the gap with L. sinuata.

Lomagramma Merrillii Holttum sp. nov. Plate 14.

Bathyphyllorum lamina 16 cm. longa, 4 cm. lata, pinne c.
10-jugatee; pinne laterales maxime, 3.5 cm. longe, 1.2 cm.

a

Gardens Bulletin, S.S.

Gard. Bull. S.S: Vol. TX. Plate 14.

Lomagramma Merrillit, Type. The fertile frond is incompletely
expanded, Twice nat. size,

Gard. Bull. SSo Vok TX, Plate 15.

Lomagramma novoguineensis (Docters van Leeuwen 9616), showing
the upper surface of the broad blunt fertile pinnze. Twice nat. size.

209

latee, basi supra late infra anguste cuneate, margine acute
serrate, apice acute; pinna apicalis 6 cm. longa, 1.4 cm. lata,
acuminata; textura tenuis, venule infra elevate, supra haud
elevate.

Acrophyllorum sterilium stipites ad 15 cm. longi, lamina ad
45 cm. longa et 15 cm. lata; pinnze superiores maxime, inferiores
sensim reducte, infimz c. 4 cm. longe, stipitatee 3 mm.; pinnae
maximze g cm. longe, 1.7 cm. late, basi superne late inferne
anguste cuneate, apice acute, falcatze, margine apicem versus
irregulariter dense szpe obtuse serrate, (margines pallide
cartilagineze), textura subcoriacez, venulis inconspicuis areolas
parvas formantes predite. Pinne fertiles stipitate 5 mm., ad
9 cm. longee, 3 mm. late.

TYPE: Mindanao, Dist. Zamboanga, Merrill 8282 (M),
‘‘very damp shaded ravine, on trees climbing, at 400 m.”’

Other specimens. Mindanao, Camp Keithley, Lake Lanao,
M. S. Clemens s.n. May 1907, Sept.—Oct. 1907 (3 sheets, M).

Merrill’s specimen is the only one showing bathyphylls, for
which reason I take it as the type. His sterile acrophyll is
thinner in texture and more toothed than those of Mrs. Clemens;
i.e. it is nearer to the bathyphyll condition. I believe however
that all represent the same species. It is a rather small species,
and is characterised by the combination of characters: small —
narrow bathyphylls with large articulated apical pinnz, subcoria-
ceous pinnee of sterile acrophylls, fertile pinnze with stalks 5 mm.,
the pinnee of moderate length. A character which seems to be
almost general is that the stipes and rachis are usually reddish
when dry; some are however stramineous.

The species has so far only been found in the island of
Mindanao. It is very different from the two other species
occurring in that island, L. pteroides and L. Copelandit.

Lomagramma novoguineensis (Brause) C. Chr. Index Suppl. III,
124, 10934. Plate 15.

Leptochilus novoguineensis Brause, Engl. Jahrb. 56: 117.
1920.

Rhizoma scandens, validum, 5-7 mm. crassum, radices numerosas
1.5-3 cm. longas, fusco-pilosas gerens paleisque fuscis, sublinearibus,
acuminatis margine subintegris, ca. 7 mm. longis 0.7 mm. latis,
dense instructum, folia dimorpha interstitiis ca. 20 cm. longis emittens.
Foliorum sterilium petioli 7-9 cm. longi, 4-5 mm. crassi, supra sulcati
infra teretes, untrinque paleis lis rhizomatis subzequalibus densissimis
muniti; lamina coriacea, ca 62 cm. longa, 22 cm. lata, supra glabra
infra paleolis bulbosis predita, ambitu oblonga, basi vix angustata,
ad apicem versus e media laminze parte decrescens pinnaque terminali
sublineari, ca 3.5 cm. longa 2-3 mm. lata, basi leviter incisa desinens ;
pinnis articulatis, subsessilibus, e basi paulo angustata, subrotundo-
truncata lineari-lanceolatis acuminatis, ad apicem versus levissime
serratis, cetera in parte subintegris, 1-2.4 cm. inter se (costis) distanti-
bus, alternis vel suboppositis, superioribus patentibus, intermediis
inferioribusque horizontalibus, infimis 1-2-jugis interdum _ paulo

Po. £X2'\ 61037):

210

abbreviatis et deflexis, maximis (intermediis) 11 cm. longis, 1.3 cm.
latis; rachibus petiolis similibus paleatis; costis utringque paleis
similibus sed minoribus supra sparsis infra densis armatis; nervis
inconpiscuis areolas obliquas formantibus, nervis apice liberis nullis.
Folia fertilia differunt petiolis paulo longioribus, ca. 12.5 cm. longis,
pinnis e basi truncata, non angustata linearibus abrupte in apicem
obtusiusculum desinentibus, margine subintegris, subpetiolulatis,
maximis 4.5 cm. longis, 0.8 cm. latis. Sori totam inferiorem pinnarum
partem occupantes.

TYPE: N. E. New Guinea, Kaiserin-Augusta (Sepik)
River, Etappenberg, 850 m., Ledermann 9524.

I have not seen the type, but have examined another speci-
men from New Guinea: Mamberamo bij Albatros Bivak, 50 m.,
Docters van Leeuwen 9616 (Expeditie Nieuw Guinea 1926), in
Herb. Buitenzorg. The striking feature of this specimen is the
fertile pinnee which are about 4.5 by 0.6 cm., with short stalks
and blunt apices, as described by Brause. ‘The sterile pinnze are
all somewhat damaged, and are rather smaller than described
(to 9.5 cm. long), but I have little doubt that this is Brause’s
species.

Lomagramma perakensis Bedd. Handb. Suppl. 107. 1892.
Plate 16.

Beddome’s description is as follows :—

Rhizome 40-50 feet long, 1 in. diameter, epiphytic on trees; stipes
6-8 inches apart, about 1 foot long, slightly scaly, but scales soon
deciduous, rachis not winged upwards; frond 3-4 feet long, very
similar to those of Blumeana, but coriaceo-herbacecus in texture,
pinne entire or obscurely crenated, quite glabrous or costa very
slightly scaly, veins sunk and scarcely visible, venation very similar

to Blumeana but closer and costal areoles smaller; fertile pinnz

1-14 lines broad.

Perak, 400 ft. alt. (Day). Referred by me to Blumeana in my
list of Day’s Perak ferns. Perak, 2,500~-3,000 feet alt. (Dr. King’s
collectors, No. 8345), the pinnze all fall off in drying, much more so
than in Blumeana.

In Beddome’s list of Day’s collections (J. Bot. 1888, p. 2},
the altitude is given as 4,000 ft.; the 400 ft. of the Handbook is
doubtless an error. I have not been able to find Day’s specimen,
and it seems rather doubtful whether the specimen exists. It
would therefore be best to take Kunstler’s collection as the type
of the species; specimens were distributed to several herbaria.

Th following description is based on the specimens in the
Singapore herbarium.

Bathyphylls. Largest fronds commonly about 30 cm. long
and 12-15 cm. wide, the stipe to about 25 cm.; pinnez about
I0O-15 pairs, or even less, the apical pinna articulate and similar
to the others; lower pinnze largest in larger fronds, apical pinne
largest in fronds of very young plants; rachis winged almost
throughout. Pinnz distinctly oblique or the lower ones some-
times horizontal, sessile; middle pinnee with upper base cuneate

Gardens Bulletin, S.S.

Gard. Bull. S.S. Vol. 1X. Plate 16.

Above,: apex of bathyphyll of ZL. sumatrana (the rachis continuous).
Below: apex of bathyphyll of L. perakensis, from a young plant
(apical pinna articulate): Both x 1.5.

"7

i

211

about 45 to the midrib, lower base very narrow; edges entire,
usually somewhat undulate, not toothed; apex subacuminate;
midrib often somewhat falcate; texture thin but firm; veins
slightly prominent above and below, often not distinct.

Plants with fronds only 4 cm. long have the apical pinna
(15 by 4 mm.) articulate, the veins free or forming an incomplete
series of areoles. Smaller fronds sometimes have an equally
forked not articulate lamina at the apex.

Sterile acrophylls. Stipes to 40 cm. long; fronds to about
1.25 m. long. Rhizome to at Ieast 1.5 cm. in diameter, densely
scaly towards apex; stipes to 1 cm. thick at base, roughened from
bases of scales. Pinnze oblique, gradually smaller towards apex.
Largest pinne 26 cm. long, 2.5 cm. wide, base rather narrowly
unequally cuneate (upper base as much as 45° to midrib, lower
always very narrow), edges slightly undulate, not toothed, more
or less reflexed when dry, apex acuminate; midrib often slightly
falcate towards apex. Texture of fully matured fronds subcori-
aceous; colour olive green, often rather pale; veins usually more
prominent on lower surface, but often almost immersed in the
substance of the frond, rarely very sharply defined on the surface,
usually in three series of areoles, the costal ones often smaller
than the next series.

Fertile pinne to 40 cm. long (commonly 15-20 cm.) and
2-5 mm. wide when dry, on stalks 3 to 7 mm. long; midrib with
two dorsal and three ventral vascular bundles (one in small
pinnee); venation in two or three rows of small areoles. The
Pulau Tioman specimen has pinnz only 12 cm. long; its sterile
pinnee are small also (13 cm. long, 2 cm. wide).

MALAY PENINSULA (all in H. S.). Sungei Ujong:
Hullett sn. Aug. 1880; Penang: Curtis 618. Pahang:
Cameron Highlands 4,500 ft., Holttum s.n. 31.3.30; Robinson’s
Falls, 4,600 ft., S.F.N. 24904,, 23259 (Holttum); Sungei Terla,
gyo0o0: ft.,; S.F .N:))313290 (Holttum);)Fraser’s Hill; 3,600 ft.,
Holttum s.n. 18-9-23, 4,000 ft., Holttum s.n. 21-3-29; Bukit
Kajang, Pulau Tioman, 2,000 ft., S.F.N. 18945 (Henderson).
Ferak: Taiping, 2,500-3,000 ft., King’s collector 8345.

SUMATRA. G. Keerintji, 1600 m., Bunnemeijer 8711
(B, two sheets, fertile only).

Bunnemeijer’s specimens have long pinne, agreeing with
L. perakensis, but in the absence of sterile fronds there is some
doubt of the identity. They cannot be either L. sumatrana nor
L. abscondita; if not L. perakensis, they represent a new species.

This very fine species appears to be quite distinctive in its
bathyphylls; in its acrophylls it is nearest to L. Brooksii and
L. lomarioides, which are mountain plants of Borneo and Java
respectively.

Vol. IX. (1937).

212

Lomagramma polyphylla Brack. Cryptogamia, Filices, in
Wilkes, U.S. Expl. Exped. XVI: 83, t. 12. 1854.

L. rhizomate scandente,; stipitibus sulcatis furfuraceo-squamosis ;
frondibus bipinnatis; pinnis petiolatis alternis; rhachi marginata
paleaceo-hirsuta; pinnulis glabris approximatis subalternis sessilibus
vblongo-lanceolatis crenatis; costis sparsim paleacets.

Hab. Ovolau, Feejee Islands: on trunks of trees.

Stipes thick and of a pale straw colour, about 8 inches long, with
a few brown furfuraceous scales, which extend to the rhachis. Sterile
fronds bipinnate, large, 2 to 3 feet long. Pzznules numerous, approxi-
mate, glabrous, subalternate, sessile, about 1 inch long and 3 to 4
lines broad, oblong- lanceolate, the margin irregularly crenate.
Rhachis of ‘the pinnz with two broad marginal lines in front (a
character not well represented in the figure), beset with slender
fugacious palez. Fertile fronds wanting.

In the absence of the fertile fronds we infer from the habit of
the sterile ones, with their articulated pinnze and uniform reticulated
venation, that our plant belongs to the present genus.

I have seen the type specimen in the U.S. National Her-
barium. It has 9 pairs of pinnze, oblique, all articulated; the
middle pinne largest, about 20 cm. long and 5 cm. wide; pinnules
all articulate about 20 pairs, slightly oblique, about 1 cm. apart,
the largest about 2.7 cm. long and 0.7 cm. wide, the base not
quite equally cuneate, the apex gradually tapering and slightly
falcate, colour rather pale, veins distinct, pale, in two series of
rather narrow oblique areoles, about 9 areoles in the costal series.

At the British Museum and Kew there are several other
collections of bipinnate fronds from Fiji and the New Hebrides
(Aneiteum); in the U.S. National Herbarium are also specimens
from Eua Island, Tonga. Both the Aneiteum and Tonga
specimens differ rather strikingly from the type of L. polyphylla
and possibly represent distinct bipinnate species. The other
specimens from Fiji however show in some cases intermediate
characters. I think.therefore that we must wait for further
information before attempting to describe any new bipinnate
species. Probably the matter will not be settled until someone
makes careful field observations on these plants in the various
island groups concerned.

The characteristic feature of the Aneiteum specimens is the
very short pinnules (to about 1.4 cm. long), with few rather large
marginal teeth and areoles to correspond. The specimen from
Eua Island, Tonga, has pinnules slightly larger than those of the
type of L. polyphylla, the margins broadly and slightly lobed,
the costal series of areoles about 6 in number, to correspond with
the lobes.

Bathyphylls are present in one of the Fiji specimens at the
British Museum (Harvey), in one collection from Aneiteum and
in the Tonga collection. In all cases they are simply pinnate,
the pinne larger than the pinnules of the acrophylls, with
shallowly lobed margins. One collection from Fiji (Meebold

Gardens Bulletin, S.S.

SOR at en

cee net (ate step at Ero Races ca cask en oe LN

Gatd> Bulle S3S.- VoL Plate. 73:

Above: Lomagramma pteroides, a bathyphyll.
Below: L. lomarioides, a bathyphyll. Both x 1.5.

213

16885, B.M.) is a small bipinnate frond with rather large pinnules
(to 4.8 cm. long and 1.5 cm. wide); this is perhaps a stage inter-
mediate between typical bathyphylls and acrophylls, but it might
possibly represent a distinct species.

Fertile fronds are present in one collection from Aneiteum
(McGillivray, Kew) and in the Tonga collection. Both are very
similar, the fronds bipinnate, of much the same size as the sterile
ones, the pinnules reduced in size, to about 10 mm. long by
3mm. wide. ‘The spores are characteristic of the genus.

The following list includes all the bipinnate specimens
I have seen.

FIJI. U.S. South Pacific Exploring Expedition 1838-42
(2 sheets, W, type collection). Seeman 713 (2 sheets, B.M.),
421 (B.M.). W.H. Harvey (2 sheets, B.M.). Nine miles above
Suva, Meebold 16885 (B.M.). Korumbamba Hill, near Suva,
Meebold 16831 (B.M.).

NEW HEBRIDES. Aneiteum, McGillivray 44, ‘‘woods,
climbing trees’? (Kew). Moore 1890, no. 17 (B.M.,).

TONGA IS. Eua Island, H. E. Parks 16311 (W, 3 sheets).
“‘Johannsen’s plantation, climbing tall trees, alt. 150 m. Fruiting
branches in top of tallest trees only. Vines many feet long’’.

Lomagramma pteroides J. Sm. Journ. Bot. 3: 402. 1841. ibid.
4: 152. 1841. Hook. Gen. Fil. t. 98. Plate 13.

Lomagramma subcoriacea Copel. Phil. Journ. Sci 40: 308.
1920.
_ L. pteroides var. subcoriacea Copel. ibid. 3C: 32. 1908.

The type of this species is Cuming 223, from Luzon. I have
examined the specimens of this collection at Kew and the British
Museum, and I can see no distinction from Copeland’s species,
of which I have seen the specimens of the type collection in the
herbaria of Manila and Buitenzorg and also several specimens
later distributed by Copeland. Copeland’s specimens have rather
narrower firmer pinnee, drying reddish; but the essential charac-
ters of the broad base of the sterile pinnze and the stalked fertile
pinnee with part of the lamina sterile towards the base are the
same in both. The reddening of the fronds on drying is rather
variable, and is perhaps to some extent connected with the
method of drying and perhaps with the position of the plant and
age of the fronds at the time of gathering.

The following description is based on all the specimens at
my disposal. ‘The description of the bathyphylls, which are
quite distinctive, is taken from Merrill 780 (Paragua) and Cope-
land 1736 (Mindanao). These are perhaps not of a fully mature
size.

Bathyphylls. Stipes short (longest seen 4 cm.). Largest
frond seen about 28 cm. long by 4.5 cm. wide. Pinnz to about

Pol IX ..{ 1037).

214

28-jugate, gradually reduced from the middle of the frond to
the base, the lowest very small (7 mm. by 3 mm.) but not much
more distant, the upper pinnz gradually reduced towards the
apex, Which is continuous with the rachis and narrow, unlike
the other pinne. Largest pinnze about 2.5 cm. long and 0.7 cm.
wide, the base unequal, the upper side broadly, the lower
narrowly cuneate, the edges crenulate (teeth 2-3 mm. apart),
the apex acute to somewhat blunt; bases and apices of lower
pinnze more rounded. Stipe, rachis and coste rather densely
scaly; scales rather dark. Pinnze reddish when dry; veins
strongly raised on both surfaces, in one or two series of areoles.

Sterile acrophylls. Rhizome with up to 5 leaf-gaps in trans-
verse section, very stout in well-grown plants. Stipes usually
20-30 cm. long. Largest pinnz seen 22 cm. long, 2.2 cm. wide;
pinneze commonly about 12-15 cm. long and 1.2 to 1.8 cm. wide,
the base slightly unequally subtruncate to subcordate, the lower
base narrower and more rounded to cordate, the sides subentire
parallel, the apex acuminate, somewhat falcate, irregularly
undulate or bluntly toothed; the uppermost pinne gradually
smaller, often less truncate at the base, the apex more clearly
toothed; lowest pinnze (except on very small fronds) not much
~reduced, the base often equally cordate and distinctly stalked.
Texture thin; colour often reddish; veins raised slightly on both
surfaces, usually more or less brown on lower surface, the areoles
rather small, in 2-3 series. Scales rather numerous, dark.

McGregor 235 from Negros shows an interesting transition
from bathyphyll to acrophyll. The Washington specimen has
the apex continuous with the rachis, the Manila specimen has
the apical pinna articulate; in each case a fertile frond is attached
to the same rhizome as the sterile.

Fertile fronds. Pinnze to 27 cm. long, commonly about
20 cm., base unequally cuneate, widening rather suddenly from
a winged stalk up to 1 cm. long; base of pinna 7-10 mm. wide,
only the edge fertile, the upper part gradually narrowed, the
distal half sometimes entirely covered with sporangia, but often
with a narrow sterile lamina on either side of the midrib to the
apex. |

ITVPE. LUZON; Crine 234.

Other specimens. LUZON: Lucban, Tayabas, Elmer 9068,
fert. (W, B). PARAGUA: E-wi-ig River, Merrill ‘780, bath.
(W, M). NEGROS: Dumaguete (Cuernos Mts.), Prov. Negros
Oriental, Elmer 10175, fert. (B, M, W). PANAY: Mt. Bulilao,
Capiz Prov., Bur. Sci. no. 35754 (Martelino & Edano, S, M).
MINDORO: Baco River, McGregor 235, fert. (W, M); Alag
River, Merrill 5654, fert. (M); Mt. Halcon, Bur. Sci. no. 40657,
fert. (Ramos & Edano, M). MINDANAO: Camiguin de Min-
danao, Bur. Sci. no. 14764, fert. (Ramos, M); Cotabato, Distr.

Gardens Bulletin, S.S.

SS ee

215

of Zamboanga, Copeland 1736, fert. & bath. (W,-M). SULU
ARCH: Jolo, Clemens 9371, fert. (M).

This species has only so far been found in the Philippines,
but within those islands it is widely distributed. It is very
clearly characterised by the fertile pinnez, which are the only
ones having regularly a partial sterile lamina that I have seen
in the genus. The bases of the sterile pinnz are also distinctive
but this character varies from base to apex of the frond and needs
careful observation. There is much variation in the width of the
pinnee of the sterile acrophylls; and in the development of the
reddish colour.

Lomagramma sinuata C. Chr. Svensk Bot. Tidskr. 16: 98, f. 5.
1922. ;

Rhizomate ad caudices arborum late repente, lignoso, longissimo,
I.5-2 cm. crasso, inferne radicibus brevibus dense radicante, superne
5-sulcis, rimis acutis separatis, profunde sulcato, squamis minutis
anguste lanceolatis adpressis atrofuscis sparse obtecto, denique
dunudato. Foliis difformibus ex schedula pendentibus, valde remotis.
Stipitibus foliorum sterilium cum rhizomate continuis ad 25 cm.
longis, basi 6 mm. crassis, cinnamomeis, glaberrimis, superne auguste
profundeque sulcatis. Lamina sterilis ovato-lanceolata, verisimiliter
1 m. vel ultra longa herbacea, siccitate brunnea, glaberrima; rachi
non alata. Pinnis 10-15-jugis, usque ad 6 cm. inter se remotis alternis,
erecto-patentibus (nec falcatis), petiolis 1 cm. longis stipitatis ad
rachin articulatis, facile deciduis, e basi postice cuneata actice rotun-
dato-truncata in petiolum decurrente oblongis, breviter acuminatis,
20 cm. longis 4 cm. latis, sinuato-repandis, margine ipso integerrimis,
pinna terminale conformi. Venis distinctis reticulatis inferne elevatis,
areolas 4-5 inter costam et marginem formantibus, areolis elongatis
plerumque 5-angulatis, costalibus valde inequalibus.—Folzis fertilibus
minoribus, lamina ad 35 cm. longa, pinnis configuratione sterilibus
similibus, 10 cm. longis, 8 mm. latis; stipite rachique paleis minutis
lanceolatis adpressis centro atris marginibus basique lutescentibus
onustis; sporangiis breviter pedicellatis paginam inferiorem excepto
spatio mediano 1 mm. lato sparse paleaceo (paleis adpressis e basi
peltata suborbicularibus in apicem longum acuminaftum contractis)
occupantibus; marginibus ipsis soris destitutis.

Bolaang-Mongondou, Modajag, North Celebes, 750 m., on stems
of living trees, leaves pendent, fertile ones. rare. W. Kaudern,
mo. 73° (B).

The characteristic features of full-sized acrophylls of this
species are the long, very broad, thin, almost entire sterile pinnze
with their long narrowly winged stalks and the broad fertile
pinnee, also stalked; the venation of the sterile pinnz is in several
series of areoles, those towards the margins being very small.
The scales on rhizomes and very young fronds resemble those of
L. melanolepis.

I have seen a pinna of the type. I have also seen an entire
fertile frond of another collection from Celebes, which agrees
with Christensen’s figure and description. These specimens have
no bathyphylls.

There are specimens (including a fertile frond) from British
North Borneo which appear to me to be referable to this species;

Vol. IX. (1937).

216

there are also specimens from East Java. From New Guinea
(Papua) are specimens of sterile and fertile acrophylls collected
by Carr; these agree exactly with Celebes specimens, except that
the sterile pinne are rather long-acuminate. There is another
specimen, from southern Dutch New Guinea (von Romer 216),
which has smaller (but still acuminate) sterile pinnee than the
Papuan specimen, and the pinne are unstalked; the specimen
consists of the apical part of frond, and it is possible that the
basal part might have had larger and stalked pinne.

Of all these collections, only that of Dr. Posthumus shows
bathyphylls, and it is fortunate that there can be no doubt of
their authenticity. The collection consists of several fronds,
both bathyphylls and sterile acrophylls. The latter are in my
opinion typical of L. sinuata and Dr. Posthumus writes ‘‘the
material belongs to one and the same plant’’. The bathyphylls
are very similar to those of the unnamed Java species, and it
might be difficult to distinguish them with certainty.

Bathyphylls. Stipes about 4 cm. long, fronds commonly to
22 cm. long and 7 cm. wide; pinnz about 12-jugate; lowest pinnz
short and somewhat deflexed; apical pinna articulate, the base
almost equal and the apex rather acute; middle pinne usually
largest, commonly about 3.8x1.2 cm., the lower base very
narrowly cuneate, the upper subtruncate but suddenly narrowing
near the very base and so forming a sigmoid curve, the edges
slightly and rather distantly toothed, especially towards apex,
the apex rather bluntly rounded, texture thin, venation distinct,
of two series of areoles; wing of the rachis rather unusually
pronounced with an auricle below each pinna; scales fairly
numerous on rachis, a medium brown colour.

The bathyphylls differ from those of the unnamed Java
species in the sigmoid curve of the upper base, the less toothed
margin, and the blunt apex; also perhaps in the rather large wing
to the rachis and paler scales.

The specimens show another feature not mentioned in the
original description, namely that the sterile acrophyll pinnz are
not always so long-stalked as described. The smaller acrophylls,
and often the upper pinnz of larger ones, would perhaps hardly
be described as stalked, but they all have the sigmoid curve of
the upper base described in the bathyphyll, only in a more
pronounced degree. In the larger pinne this curve is exagger-
ated, and combined with a similar but much flatter curve in the
narrow lower base gives a winged stalk. The lowest pinne of
large acrophylls are rather short (not much more than half the
length of the longest ones), and the lower base much broader,
sometimes almost equal to the upper base, the winged stalk still
present. In the pinna from the type collection in the Buitenzorg
herbarium the winged stalk is only 5 mm. long, and this seems
a common length.

Gardens Bulletin, S.S.

217

Another correction of the original description (so far as
complete specimens examined by me are concerned) is that the
pinnze may be up to 22-jugate.

CELEBES. N. Celebes: Kaudern 73 (type, as cited above).
S. Celebes: Kosali-Porema, 650 m. Kjellberg 2608 (B).

BORNEO. B.N. Borneo: Menetendok, Kinabalu, 2,500
-ft., Clemens s.n. 1-3-33 (S); above Koung, 2,500 ft., Clemens
51308 (M).

JAVA. Res. Fasoeroean, Tangkil Zuidergeberte, 400-500 m.
Koorders 23554b (B). Res. Probolinggo, Mt. Lamongan, 450 m.,
Posthumus 3975 (S).

NEW GUINEA. Papua: Veiya, sea level, Carr 11677 (S).
Dutch N. Guinea: von Romer 216 (B).

Lomagramma sumatrana v.A.v.R. Bull. Jard. bot. Buitenz. III.
Ser. 2: 158. 1920. Plate 16.

Leptochilus cuneatus R. Bonap. Notes Pterid. XIV: 453.
1923.

The original description of this species was as follows :—

L. lomarioide@ J. Sm. affinis sed pinnis sterilibus integerrimis
vel ad apicem solum serratis, in sicco viridi-vel fusco-olivaceis, subtus

quam supra pallidioribus, pinnis fertilibus anguste linearibus, 4-5 mm.
latis subtus unique soriferis basi interdum sterilibus dilatatisque.

Forest; to 5 m. high scandent; alt. 350-500 m.—Originally
determined by me as L. pteroides J. Sm.

SUMATRA (Deli, Sibolangit, J. A. Lorzing nos. 5564, 5623, 2oth
March-8 April 1918).

It is evident that by L. lomarioides in the above description
the author meant the unnamed lowland species of Java, not the
true L. lomarioides (=L. abscondita v.A.v.R.), which has quite
entire pinnz. I have examined the type of L. sumatrana, and
believe it to be identical with the lowland species of the Malay
Peninsula, which is well represented in the Singapore herbarium
by bathyphylls as well as acrophylls. The following full des-
cription is based on Peninsula specimens so far as bathyphylls
are concerned, and on both Sumatran and Peninsula specimens
for the acrophylls.

Bathyphylls. Stipes 3-10 cm. long; lamina to about 40 cm.
long and 9 cm. wide. Pinne 20 to 30 pairs, lowest rather
reduced, more distant and reflexed, upper pinne gradually
smaller, the uppermost grading into lobes on the apex of the
frond which is not pinna-like and not articulated to the rachis.
Largest pinnee about 5 cm. long by 1.2 cm. wide, commonly about
4cm. by 0.9 cm., at right angles to the rachis, upper base broadly
subtruncate subauriculate, lower base half as wide, rounded or
sometimes nearly cuneate, sides almost parallel for two-thirds of
their length, than gradual'y tapering to acute or subacute apex;
edges shallowly crenate with one or two sharper teeth at the
distal end of each lobe, the teeth or groups of teeth 3-5 mm.

Vol. IX.- (1937).

218

apart. Texture thin, colour usually green when dry but some-
times brown; midrib pale, raised above and below, veins distinctly
prominent on both surfaces when dry, concolorous with the
surfaces, slender.

Sterile acrophylls. Stipes to 20 em. long; frond to at least
90 cm. long and 25 cm. wide. Lowest pinnee somewhat reduced
and more distant, uppermost gradually smaller, apical pinna
articulate and same shape as others. Middle pinne to about
14 cm. long and 2 cm. wide (occasionally to 2.5 cm.), at right
angles to the rachis or slightly oblique, sessile or stalked less than
i mm., base broadly cuneate above, rounded to cuneate below
and about half as wide, edges usually distantly toothed towards
apex, otherwise entire, sides approximately parallel for *% of
their length, apex acuminate; texture thin but firm, veins raised
zbove and below, in 3 series of areoles, the costal areoles not
considerably larger than the next series.

Fertile fronds. Pinnz 2.5 to 3 cm. apart, to about 15 cm.
long by 5 mm. wide, stalked to 2 mm., the base unequal; upper
vinnee gradually reduced to about 4 cm. long; venation in 2 or
three series of areoles, usually prominent above; lower surface
soriferous throughout except for the midrib and the thin pale
edges.

The characteristic features of this species are the long narrow
bathyphylls with many pinnz and the apex not articulate, the
broad sterile acrophyll pinnz, thin in texture and slightly toothed
at most towards the apex, and the broad slightly stalked fertile —
pinne.

I have not seen Brooks’s specimen from southern Sumatra
which was described by Bonaparte as Leptochilus cuneatus, but
from the description (especially of the broad fertile pinnze) it
appears almost certainly to belong here. Bonaparte quotes also
under Leptochilus cuneatus a specimen of Raciborski from Java
(Salak). It seems possible that this is a paratype of L. abscon-
dita (L. lomratoides) but if so the description would not apply
to it. I have seen no specimens from Java to which the
description of L. cuneatus would apply.

In his description, v.A.v.R. has the phrase ‘‘basi interdum
sterilibus dilatatisque’’, referring to fertile pinne. The type
collection has a frond which is partly fertile and partly sterile,
and at the transition region some pinna are sterile at the base
with fertile apical half. This is of course not a character of the
species, but just a chance occurrence.

L. sumatrana occurs in the low country and foothills of the
Main Range in the Peninsula, and, so far as recorded, in low
country also in Sumatra. The mountain species is L. perakensis,
which is very different at all stages.

SUMATRA. Sibolangit, 475 m., Loérzing 5564 (B, S, M),
5623 (B). Benkcelen, Lebong Tandai, Brooks 4678 (type of
Leptochilus cuneatus R. Bonap., not seen).

Gardens Bulletin, S.S.

219

MALAY PENINSULA (all S). Selangor: Ginting Simpah,
1,500 ft,, S.F.N. 26104 (Holttum). Negri Sembilan: Ulu
Bendul, S.F.N. 9859 (Holttum). Pahang: Pulau Tioman, W.
of Joara Bay, S.F.N. 1138 (Burkill). Perak: Chanderiang,
500-1,000 ft., King’s Collector 5778 (distributed as Acrostichum
perakense Bedd.).

Lomagramma sp.

As noted by van Alderwerelt van Rosenburgh, there are two
species of Lomagramma in Java (in addition to L. sinuata, which
has been twice collected in East Java); van Alderwerelt accord-
ingly decided to provide a name for the second species. Owing
to the fact that he evidently never saw the type of L. lomarioides,
he gave a new nate to that species instead of the other, which
therefore still lacks a name. ‘The unnamed species is evidently
a fairly common one in some parts of Java, but curiously enough
it has never been collected in a fertile condition. It is very
similar to L. Copelandii, and might possibly be identical with
that species. In view of this uncertainty, I do not name it here,
but give a description of the bathyphylls and sterile acrophylls.
The bathyphylls are rather similar to those of L. sinuata, but the
pinnee are much more toothed, more inclined to have an acute
apex, and the scales are more numerous and darker. The
acrophylls lack the characteristic base of L. sinuata and are
distinctly toothed towards the apex.

Bathyphylls. Smallest fronds seen, with lamina 12 cm.
long, have the apical pinna largest and articulated like the others.
This condition persists in bathyphylls up to 25 cm. long; in
larger fronds the largest pinne are about one-third from the apex,
the apical pinnee somewhat reduced. Middle pinnz of typical
fronds about 4 cm. long and 1.1 cm. wide, the upper base truncate
to broadly cuneate, the lower base very narrowly cuneate, the
apex gradually tapered to an acute point (or rounded in the
smaller pinnee), the midrib slightly falcate, the margins slightly
lobed, the lobes about 4 mm. wide, each with 2 or 3 teeth, the
basal ones sometimes not toothed; lowest pinnze with lower base
more broadly rounded, and apex rounded. Pinnz to about 18-
jugate, usually separated by about half their own width; texture
thin; veins raised on both surfaces, areoles in two series.

Sterile acrophylls. Stipes to 15 cm. or more; frond to 60 cm.
long and 20 cm. wide; pinne to at least 20-jugate. Largest
pinnze about 12 cm. long and 1.8 cm. wide, midrib slightly
faleate, upper base broadly, lower more narrowly cuneate, sides
parallel for more than half their length, entire except for the
apical third which is distantly toothed (teeth 4-5 mm. apart),
apex acuminate. Texture thin; veins raised on both surfaces,
usually pale on the lower surface, areoles in about 3 series, costal
ones somewhat larger than the next. Lowest pinnz somewhat
reduced, the base more broadly cuneate.

Vol. IX. (1037).

220

Fertile fronds not seen.

This species is certainly very like L. Copelandii. The
youngest plants differ from that species in having the apical
pinnze articulate from a very early stage and larger than the
others; larger bathyphylls differ in having the lower base of the
pinnze broader, and in a larger number of basal pinnz reduced
and reflexed; acrophyll pinnz differ in being distinctly toothed
towards the apex.

JAVA: Res. Paseerean: veet G. Smerce, 400 m. Posthumus
1613 (P); helling G. Smerce, 700 m. Posthumus 1621 (P); bij
Waterval Scemberwangi, 300 m. Posthumus 1508 (P, S); helling
G. Smerce 825 m. Posthumus 549 (P, B); helling G. Smerce
400 m. Posthumus 1608 (P). Res. Batavia: Buitenzorg 240 m.
Bakhuizen fil. 3123 (B); Sitcehiang 500 m. Bakhuizen 6452 (B,S);
Kloof v/d Salak, Bakhuizen 6304 (B); Gcenong Batce, 250 m.
Bakhuizen fil. 3664 (B); Botanic Gardens, Buitenzorg (B),
Posthumus s.n. 1925 (P).

SUMATRA. Aer Telani Kloof, 11 km. N. van Banging
Agoeng, Ranau N. W., 400 m. van Steenis 3951 (B).

8. SPECIES DUBIA

Lomagramma sorbifolia ( Willd.) Ching, Lingnan Sci. Journ. 12:
566. 1933.
Aspidium sorbifolium Willd. Sp. Pl. 5: 23. 1810.

This species was described by Willdenow as follows :—

A. frondibus pinnatis, pinnis lanceolatis alternis obtusiusculis,
serratis, basi superiore truncato-subauriculatis, inferiore cuneatis,
rachi paleacea. W.

Habitat in India orientalli.

Caudex filiformis repens glaber, Stéfes pollicaris glaber. Rachis — .
paleis brevibus obsita. F/zondes quinquepollicares pinnate. Pinne
mediz pollicares, inferiores et superiores sensim breviores, lanceo-
late obtusiuscule remote serrate serraturis 5 vel 6 in utroque latere,
basi superiore truncate, vix manifeste auriculate, inferiore cuneate.
Costa media paleis parvis ventricosis obsita. sSovos perfectos non
vidi. W.

Prof. Diels has very kindly sent me a photograph of the
type of the species, which is in the Berlin herbarium. As one
would anticipate from the description, the specimen consists of
bathyphylls from a young plant. The pinne are similar in many
respects to the lowland species from Java and also to L. sumat-
rana, L. grosseserrata, and L. Matthewii. These species however
differ very markedly in the apex of the bathyphyll. In the Java
fern the apical pinnee are similar to the others, subequal, and
articulated. In the other species the pinnz are gradually reduced
to the apex of the frond, which consists of a narrow lamina
continuous with the rachis. Willdenow’s specimen has the latter
character, and is quite unlike any specimen I have seen from

Gardens Bulletin, S.S.

221

Java. It is probably one of the other three species, but our
present knowledge is not adequate to say which. ‘The only
course therefore is to ignore Willdenow’s name for the present.

9. DOUBTFUL SPECIMENS

There are several doubtful specimens, some of which may
represent new species, but the material is inadequate for a full
description to be possible. Some of the specimens may represent
bathyphylls of known species; but there is no definite evidence
of this. It is clear from the foregoing descriptions that for a full
characterisation of a species of Lomagramma all stages of
development must be available; species resembling each other at
one stage may differ strikingly at another.

LUZON. Rizal Province, Loher 23150 (M). ‘This specimen
consists of bathyphylls about the same size as those of L. Cope-
landii, but differing in the shape of the pinnee, which are more
oblique and falcate, the upper base subauriculate, the lower base
more narrowly cuneate, the edges not lobed and only irregularly
toothed towards the apex. Judging by the bathyphylls of those
species in which a number of specimnens are available, I do not
think it likely that this specimen comes within the range of
variation of L. Copelandii. It is probable therefore that there is
another Philippine species and to it may possibly belong some of
those specimens which I have referred to L. pteroides or L. Cope-
landii, but until full collections of all stages are available nothing
further can be said.

NEW GUINEA. N. New Guinea, H. J. Lam no. 1200,
13-9-1920 (B). ‘This specimen consists of very small bathyphylls,
all of about the same size, to about 7 cm. long (including the
stipe) and 2.8 cm. wide. ‘The collector noted that the fern was
very abundant on the ground and on roots (of trees presumably)
but that no fertile fronds could be found. ‘This specimen might
perhaps be a very young stage of L. novoguineensis, or even of
L. sinuata, but there is no character in the bathyphylls of Lam’s
specimen to give a definite connecting link with either species.

CELEBES. Manado Resid., Tomeolama, 700m. C. A. Wisse
no. 71 (B). ‘This specimen consists of four bathyphylls rather
similar to those of the specimen from Luzon above quoted, but
the pinne more regularly toothed. It might well be the same
species.

TAHITI. M. Vesco 1847 (M, ex herb. Mus. Paris.). This
specimen is the apical half of a sterile acrophyll, which is very
near the unnamed Java species in appearance.

Pol. 1X. (1937).

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_ STRAITS SETTLEMENTS |

9th October, 1937 Part 3
. CONTENTS
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THE

GARDENS’ BULLETIN

STRAITS SETTLEMENTS

Vol. IX 9th October, 1937 Part

3

# A COMMENTARY ON THE LAWS OF BOTANICAL

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NOMENCLATURE
By ©... Fperano,

Botanic Gardens, Singapore

Introduction

Definitions of the Terivs. Valid Lid Risivinate

Kinds of Invalid Names, and Nameless ike tunic
Groups

Provisional Names

Kinds of Illegitimate Names or Dpsmatiy ms

Legitimate Names ;

Three Customary Procedures Pte Niue Art. oe

What a New Combination or a New Name is

Alternative Names $f

Rival Names of Equal Age and Brad Hank

Isonvms of Unequal Age Based on Basinyms of Equal
Age : ets" ea an

Varietal Names :

Nomen Dubium or Misinma Eomienicitee
Tautonyms and Homonyms

Linnean Genera aah

Orthography of Names

Validating a Name by Beene t toa Masapplication
Citation of Misapplications of Names
Nomenclatorial Expressions

Nomina Specifica Conservanda

Effectiveness of Advance Separates

Representative Botanical Institutions aiid: ee 36
Summary

1. Introduction

Page
223
224

227
230
232
234
237
239
239
240

241
242
244
249
255
256
258
261
262
267
271
273
280

Since the main object of the International Rules of Botanical
Nomenclature, ed. 3 (1935) is to ensure simplicity and finality
in the names and to prevent more than one interpretation to a

given name sensu stricto,

it is unfortunate that there should

exist a good deal of misconception and dissatisfaction over the
But, for the existence of this state of affairs, the Rules

Rules.

223

224

themselves cannot be entirely exonerated from blame. The
terms valid and legitimate with their respective derivatives,
which have a great importance in botanical nomenclature, have
been defined and used equivocally in the Rules; and some
nomenclatorial points, where definite instructions are absolutely
needed to avoid confusions and equivocations, are left
ambiguous. ‘The result is that practices contrary to the spirit of
the Rules seem admissible under the letter of the Rules.
Already in a small footnote in the GARDENS’ BULLETIN,
S.S. Vol. VIII, 1935, p. 341, I first drew the attention of botanists
to the need of making a clear distinction between the two terms,
legitimate (=legal) and valid. In the same publication (l.c.
p. 336), I pointed out the necessity of prohibiting the acceptance
of a newer interpretation of a name, the type of which is not
extant, without showing any adequate botanical reasons for the
rejection of earlier interpretation. But as the nomenclatorial
laws are of very great importance to systematists, and as in
neither of my previous communications have I developed my
ideas on the subject to any great length, I make no apology to
revert to it here.

2. Definitions of the Terms Valid and Legitimate

A name of a taxonomical group is invalid because certain
factors existing in the name itself, or in its botanical descrip-
tion, or in the circumstances affecting the publication of the
book or periodical in which the name is published, violate the
Rules; that is, the name does not satisfy the conditions prescribed
as basic for the words constituting the name, for the words or
references constituting its taxonomical description, for the nature
of the book or periodical wherein it is published, or for the means
taken to make the name known to systematic botanists. Failure
to comply with the basic conditions prescribed in the Rules
makes a taxonomic name invalid; that is, it does not count, and
has no status, in systematic botany; and as such it should not
be listed in works like INDEX KEWENSIS.

Examples of invalid names are all nomina nuda; tautonyms;
names published since January st, 1935, without Latin
diagnosis; or published before the starting points prescribed in
Art. 20, or in an ineffective publication; names which cannot
be considered as belonging to the binomial system; etc. All such
names may be conveniently classed as NOMINA INVALIDA.

Taxonomic names that satisfy every one of these conditions.
are valid; and as these are the only ones that count in systematic
botany, they must all be registered in works like INDEX
KEWENSIS. Valid names, however, may have been published
in violation of another set of conditions or laws that have
nothing in common with those mentioned above, and which may
be termed secondary or accessory. ‘These conditions are laid

Gardens Bulletin, S.S-

225

down with the object of ensuring uniformity and finality in the
valid botanical nomenclature so as to render it more easily
followed by the interested public, without, at the same time,
losing the trend of the history of the individual group from its
earliest valid origin. Valid names that satisfy all these accessory
conditions are legitimate, while those that do not satisfy any one
of these conditions are illegitimate.

Though several names may have been validly proposed for a
given taxonomic group (i.e. a group in a given sense, in a given
position and in a given rank), the secondary rules make
it impossible for the group to bear more than one
legitimate name and regard all the other rival names as illegiti-
mate. Hence an author, having defined his taxonomic group
and assembled together all the rival names of the group, should
consult the accessory rules in finding out the correct, that is, the
legitimate, name for his group. Should opinions differ on a
given taxonomic group with regard to its limits, rank or position,
so as to permit it being regarded as one according to one opinion,
or as two of the same rank, or as one of a different rank, according
to another, then an illegitimate name to one botanist will become
a legitimate name to another, and vice-versa. The main object
of the Rules is, therefore, not to interfere with the taxonomic
opinions of botanists, but to instruct them as to which of the
existing names or epithets should be legitimately used for a
given taxonomic group in a given sense, or what conditions should
be complied with in order that a certain name may be admitted
first as valid and then as the legitimate.

Illegitimate names owe their origin: to their authors’
ignorance of the previous systematic work on the subject; to the
wrong supposition that the previous illegitimate names are invalid;
to faulty judgement or to differences of opinions about the
limits of the taxonomic groups involved; to misinterpretation of
previously published names; to oversight when in one and the
same work the same name is adopted for more than one species, or
more than one name for the same species; or also to special
decisions of the Congress against the oldest validly published
names that are not later homonyms.

Unlike the invalid names, illegitimate names cannot be
disregarded in systematic botany, but, like the legitimate names,
must be registered in INDEX KEWENSIS, because they
effectively preclude one from applying them again legitimately
to another and later taxonomic group of equal rank; and also
because their status is often a matter of taxonomic opinion so
that, as said above, an illegitimate name to one botanist may be
legitimate to another. Illegitimate names constitute the
synonymy of the legitimate names, and, when cited as such, serve
to show how, and possibly, why, the nomenclature of each

Vol. IX. (1937).

226

individual group has undergone a series of changes from its
earliest valid origin as well as to indicate how the opinions of the
botanists, who cite them, differ from those of others on the same
subject.

Later names of a taxonomic group and later homonyms, if
not conserved, are illegitimate. The following two examples
may be cited to illustrate the point in question :

(a) Desmos chinensis Lour. (1790) was published simultaneously
with D. cochinchinensis Lour. (1790), but the author himself thought
them to be doubtfully distinct and suggested the possibility of the
former being a variety of the latter. Though there are botanists
who regard the species as distinct, in which case both names are
legitimate, there are other botanists who consider the species as
specifically equivalent. As the names are of the same age, it becomes
difficult to decide in this case which of the two should be regarded as
illegitimate and which legitimate. The new Rules prescribe to follow
the botanist who first decided on the matter; and as in this case it
happens that the botanist, who was first to consider these two species
definitely as equivalent, had cited D. chinensis Lour. as a synonym
of the other, D. chinensis will have always to be considered as
illegitmate whenever it is regarded as being identical with D.
cochinchinensis Lour.

(6) The next example is of two valid genera, the oldest of which
is arbritarily made illegitimate by the decisions of the Congress:
Homalocenchrus Meig. (1760) and Leersia Swartz (1788) were
published independently, but based on the same type species PAalaris
oryzotdes L. Here though the first name is older of the two, the
second is conserved by the Rules and is made thereby legitimate for
the group. But as this conservation makes the first name only
illegitimate and not invalid, the name Homalocenchrus cannot be
legitimately used for any other genus in which the type species
Phalaris oryzoides is not included. In such cases as this there is also
the possibility of future decisions reversing the previous ones and
making legitimate what is now illegitimate, and vice versa.

In order to follow the Rules better, therefore, read:
LEGITIMATE for Valid in Arts. 16 and 58; ILLEGITIMATE
for illegitimate in Arts. 60 (1-partly), 60 (3), 61, and 69 (partly);
VALIDLY for Validly in Arts. 19, 37-40, 42, 45, 48 and 58;
VALIDATE for Validate in Arts. 41 and 43, VALID for
legitimate in Arts. 20, 45, 56, 60 (2), and 69; VALIDATE for
legitiniize in Art. 38; INVALID for illegitimate in Arts. 2
(partly), 60 (4-5), 67 and 68; using VALID and LEGITIMATE
with their respective derivatives as defined in this paper; though,
to make the Rules precise and unequivocal, it would be
necessary to state the Rules afresh, changing completely the
sections into which they are divided and the order in which the
Articles are arranged.

Articles affecting the validity of the names of taxonomic
groups are Arts. 19, 20, 36-44, 45 (partly), 59 (partly), 60 (4-5),
67 and 68. All the remaining Rules [i.e. Arts. 21-35, 45
.{partly), 46-58, 59 (partly), 60 (1-3), 61-66 and 69-74] and
Art. 16 (partly) affect the legitimacy.

Gardens Bulletin, S.S. |

oka

227

3. Kinds of Invalid Names, and Nameless Taxonomic Groups

Botanical legislators will do well to recognize the following
kinds of invalid names, as such a recognition will make the
nomenclatorial laws more precise and unambiguous :—

I. A name in an ineffective, and therefore invalid,
publication.
Botanically ineffective publications are those that are
published in contravention to Art. 36.

In my opinion here should also be included all those
publications that ordinarily do not reach systematic botanists
e.g. modern Encyclopedias and Dictionaries on gardening
and on the uses of plant products; seed-lists, seed-catalogues;
non-botanical and non-systematic periodicals and books;
botanical and systematic books meant to be used as text-
books for schools, colleges, etc. As a decision making
non-botanical, and non-systematic books and_ periodicals
systematically ineffective may have undesirable effects,
time should be fixed (say January rst, rg41) after which such
publications would be considered ineffective.

II. A naine in an effective, but invalid, publication.

In this class will come all botanical works that have
been effectively published, but which, owing to the starting
points fixed in Art. 20, cannot be taken into account for
systematic purposes; or such works published after the
starting points, but where the binary system is not con-
sistently followed throughout, cf. Art. 68 (4).

Should any one in future publish any systematical work
without adopting in it the binomial system, that work
should, in my opinion, be regarded not only as invalid but
also as ineffective, so that it would not be possible to vali-
date new binomials by referring to the descriptions in such
works.

The question of validity and effectiveness of works of
early modern authors not following consistently the binomial
system is sub judice, a Committee having been appointed by
the last Congress to investigate this matter.

III. A name in an effective and valid publication.
Such a name may fall into any one of the following
categories :—
(a) A validable name but without adequate systematic,
or with invalid, description :—

1. A name without any botanical description or
without a reference to another effectively
published description or plate.

2. A name without a description, but with refer-
ences not admitted by the Rules.

pot. EX. (1037):

228

3. A name with a description, but not in the
language prescribed.

(b) A validable name with adequate description or
references but proposed for adoption in the future
should certain expectations come true. Such a
description should in my view be regarded as a
non-validable description. (Provisional names).

(c) A non-validable name but with a valid description
(i.e. a description that would have been valid had
the name been also valid).

1. A name not intended by the author to be a
taxonomic name cf. Arts. 67 (1) and 68 (1-2);
or coinciding with technical terms to be
rejected under Art. 67 (2).

2. A name not belonging to the binary system
published as an exception in a book where
the binary system is consistently employed
cf. Art. 68 (4).

3. A name belonging to the binary system, but a
tautonym.

IV. Nameless Taxonomic Groups.

This class includes all those taxonomic groups which,
though recognized as new and accompanied with valid
descriptions, are published without any vernacular or
scientific names. They are sometimes separated with
difficulty from the Class III-c (1). Such species are some-
times distinguished by the terminology ‘‘Species No. I’’,
“Calamus cf. anomalus’’, ‘“‘Calamus anomalus?’’, etc.

Failure to grasp these distinctions by the legislators has
caused much confusion regarding the proper attitude to be taken
towards these invalid names. It is a general, though not
apparently universal, custom not to validate a taxonomic name
by referring to invalid literature, or to the descriptions of invalid
names in valid literature, exception being made to plates and
figures im effective but invalid literature in Class II and to valid
descriptions in Classes III-c (1-3) and IV and also to species in
Linnaeus’s Species Plantarum ed. 1 and 2. Where reference
has been made to a description in an effective but invalid
botanical publication (Class II), it has been customary to take
this as a reference to the type specimen of the invalid species.

I think a great number of Nomina ambigua, confusa vel
dubia would disappear if all botanists were to adopt this attitude
in interpreting a name validated by references to effective but
invalid publications. References to invalid literature or species
do not produce a new isonym (vide infra for the definition of this
term, section 5), but a new species. Hence despite the citation
of an invalid name and even the adoption of its trivial epithet,

Gardens Bulletin, S.S.

229

the author can, if he likes, choose as the type of his species any
specimen, even one that is not included in the invalid references
cited, provided the specific description is drawn from it.

Owing to the unintentional omission of significant words,
Arts. 37, 42, 43 and 44 of the Rules seem to permit validation
of names of taxonomic groups by referring to any sort of invalid
names other than nomina nuda, provided the reference has
satisfied the conditions imposed in Art. 36 for its effectiveness;
while Arts. 43 and 44 seem to admit references to plates invalidly
but effectively published. This means that one can validate a
naine of a species or genus by merely referring to plates (issued
before 1908), and descriptions, that have appeared in popular,
non-systematic and non-botanical books or periodicals, or to
species that have been published after 1934 without any
diagnosis in Latin, provided the periodicals and books have com-
plied with the conditions laid down for the effectiveness of their
publication.

Such a procedure, if admitted as valid, would violate the
very spirit with which the Arts. 37-44 have been inserted in the
Rules. In fact I think that it would be to the advantage of
systematists in general if, henceforward, only citations of valid
species, or of invalid names but with valid descriptions in
Class III-c and Class IV should be permissible references for
validating a name of a taxonomic group; in future all references
to any other invalid names or descriptions, even those in Class II
to which reference has been permitted, should be made invalid.

This defect in the Rules can be remedied by inserting the
word validly after effectively in lines 3 of Arts. 37 and 44, and
in line 2 of Art. 42; and the word effectively before the last
phrase before January 1st, 1908 in Arts. 43 and 44; as well as by
taking the word effectively in the sense suggested under invalid
names of Class I and Class II. But should there be any objections
to give the word effectively the sense proposed above, it is
necessary to define valid publications as being those that
ordinarily reach systematic botanists, a provisional exception
being made to periodicals like the Indian Forester, Journal of
Royal Asiatic Society, Malayan Branch, Gardeners’ Chronicle,
etc. which are not purely botanical periodicals. But publications,
which their titles indicate to be purely zoological, ethnological,
geological, medicinal, etc. and which should therefore have
nothing to do with systematic botany, or which belong to the
class of modern dictionaries and encyclopedias on gardening or
plant-products, botanical text-books for schools and colleges, and
seed-lists, should, for the purpose of systematic botany, be
declared invalid, even when they are effectively published. It
would then be a very simple matter to rule out as invalid all
taxonomic names published in invalid publications, as well as
such.as have been published merely by references to descriptions
or figures in invalid publications. A ruling like this would, for

Vol. IX. (1937).

230

instance, invalidate the alga, Rhopalodia vermicularis O. Mull
forma recta Fl. Rich in Journ. Linn. Soc., Zoology, XX XVIII
(1933) 262.

The Rules are ambiguous as to the status of the names
published in the direct contravention of Arts. 54, 59 and 69, so
that it is not possible to decide whether the names thus published
are valid but illegitimate, or invalid. In this matter Art. 60 is
also ambiguous (vide my remarks in Section 6, p. 235). A
definite ruling on this matter is necessary as it affects the status
of homonyms and consequently the legitimacy of certain names
rejected as later homonyms.

4. Provisional Names

A new amendment made to the Rules at the last Congress
(1935) regards provisional names as invalid, but no adequate
description or examples are given to show exactly what a pro-
visional name is. ‘The clause regarding the provisional names
reads as follows :—‘‘A name of a pees group is not validly
published unless it is definitely accepted by the author who
published it. A name proposed provisionally (nomen provi-
sorium) in anticipation of the eventual acceptance of the group,
or of a particular circumscription, position, or rank of a group,
or merely mentioned incidentally is not validly published’’.

Now the effect of the first sentence of this decision will have
far reaching results, and, if put into practice, may produce
chaos and confusion, requiring a lengthy list of nomina generica
et specifica conservanda.

(1) Eptpremnum humile Hook. f., Fl. Brit. eH. VI (1893) would
become invalid because of the author’s following remarks on the

genus: ‘‘This genus [£p7premnum] had better be merged in
Pad beahore. ”

(2) Plectocomiopsis dubius Becc., which has been described at
some length in Ann. Roy. Bot. Gard. Calc. XII (1918) 56, will suffer
the like fate because of the following observations: ‘“‘It is a very
imperfectly known and doubtful species———__We may even suppose
that P. dubius represents a juvenile stage of P. Wzrayi——————In
conclusion, P. dubius established on Ridley’s n. 12119 must be con-
sidered as yet as a doubtful species, but not rie £2 referable to
P. geminiflorus as has been stated by Ridley

(3) Vesmos chinensis Lour. will cease to have any valid status
Reo the author had added to the description the following words:
*‘fortasse illius varietas’’.

In each of these cases, the author may be interpreted to mean
that there are reasons to doubt the delimitation of the genus or
species concerned, but that he leaves the subject to future
botanists, whose attention he wishes to draw by his remarks.

At present he is content with the names he has adopted and ~

described.

A parallel case is also of Cyrtophyllum? lanceolatum DC. sp.
nov. Prodr. IX (1845) 31. Here the author means that he is
pretty sure that the species as represented by the specimen he has

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described is a new species; and that though his material is either
aberrant or insufficient to say definitely whether it belongs to
Cyrtophyllum or to any other closely allied genus, the author
assigns it to the genus Cyrtophyllum: Let the future investi-
gators transfer the species to another genus if they disagree with
his disposition.

If botanists are to reject the above species on the plea that
the author in each case did not accept definitely the genus or
species, then this rejection is going to create a very bad precedent
which, as said above, will have to end in a creation of a lengthy
list of nomina generica et specifica conservanda to prevent chaos
and instability in nomenclature.

Similarly I consider that the name Neurotecoma K.
Schumann is validly published, though the author considered
the material at his disposal insufficient for a thorough investiga-
tion. If it is invalid, as proposed in Briquet’s Recueil
Synoptique p. 41, 1930, and declared by the last Congress (1935),
because of certain words contained in its description, then
Spirotecoma Baill. ex Dalla Torre et Harms is also invalid because
the last two authors did not make any exclusion from Schum-
man’s description, but meant that Spirotecoma Baill. was
synonymous with Neurotecoma Schum. Further, as explained
above, the practice of validating a name by referring to an invalid
description, or to a description that, though published in a valid
publication, does not belong to Class III-C and Class IV, is not
supportel by usage and therefore needs condemning. If in the
description or remarks there were certain words that made the
genus Neurotecoma invalid, the words were not excluded by
Dalla Torre and Harms in their citation. Had they described the
genus, then the description, and not the citation, would have
validated the genus Spirotecoma. ,

A parallel may be found in the case of nomina confusa. If
a nomen confusum is used as a basinym without showing
exactly what portions from the description and what specimens
are to be excluded from the basinym*, then its isonym* becomes
also a nomen confusum. If this interpretation of the provisional
names is not accepted, then it will be possible to validate a name
by referring to an invalid description; and it will also mean that
invalid provisional names will become valid by merely
registering them in works like the Index Kewensis.

According to my view, the names in the following sentences
aie invalid and cannot be used as basinyms :—
(1) “If this section [Mesembryanthemum sect. Minima Haw.]

proves to be a genus, the name of Conophyton would be apt?’
(Haworth, Rev. Gen. 1821 p. 82).

(2) ““The anther-appendage is analogous to that of Doryphora;
consequently this Eupomatia [E. Belgraveana F. Muell.] might sub-

_ generically or perhaps even generically be separated (as Himantandra)’’

(Australas. Journ. Pharm. Jan. 1887). (cf. also Alternative names).

* These terms are explained on p. 233 under exact synonyms.

Pot. EX. \(1037).

232

(3) Should Beccari’s supposition that Piychosperma angustifolium
Bl. is identical with Coleospadix oninensis Becc., accord with actual
fact, then Coleospadix angustifolius (Bl\.) would ‘be the correct name
(hypothetical case).

(4) I make the new combination Coleospadix angustifolius (Bl)
to be used instead of C. oninensis Becc., should Beccari’s surmise
that this species is identical with Ptychosperma angustifolium Bl.
turn out to be correct (hypothetical).

(5) If Curtis’s specimen from Penang proves to represent a
species different from FPothos Maingayi Hook. f., then it might be
called P. Curtisiz. The latter may be described or distinguished as
LT ii Aa IE Re ay: (hypothetical).

But if, instead, a different wording was used so that not the
adoption, but the rejection, of the names proposed was made
dependent on future investigations, I think the names could be
regarded as valid, and could be used as basinyms for making
new binomials. These sentences (1)-(5) might be turned in this
way so as to validate the new names :—

(1) I think this section should be raised to a generic rank. I
propose to name it Conofhyton. [In this case Conophyton Haw.

would have been an alternative name to Mesembryanthemum sect.
Minima Haw. See below my remarks on alternative names].

(2) Eupomatia Belgraveana F. Muell. is so distinct that it would
be better, I think, to make it the basis of a new genus Himantandra.
The anther-appendage of this species is analogous to that of
Doryphora, and difiers from other Eupomatia species

(3) Beccari thinks that Coleospadix oninensis Becc. is identical
with Ptychosferma angustifolium Bl. Though this cannot now be
proved beyond all doubt, for there is no type specimen of the latter,
I think it is better to follow Beccari in this matter. I therefore make
the new combination Coleospadix angustifolius (B)}).

(4) On the supposition that Beccari’s opinion is correct that
( oleospadix oninensis Becc. and Ptychosperma angustifolium Bl. =
the same species, I make the combination c.. angustifolium (Bl).
be used as the leg gitimate name for C. omimensis (Becc.).

(5) Curtis’s specimen resembles Pothos Maingayi Hook. f. so
very closely that I hesitate to describe it as specifically different, but
fiowers are more than twice as large as in that plant
I name it provisionally as Pothes Curtisaa Hook f. Further studies
might show it to be a mere form or variety of P. Maingayt.

or: this specimen, though it resembles very much Pothos Maingayi
Hook. f., might be specifically separated under the name P. Curtisii,
for it differs from P. Mamgayi in 2

5. Kinds of Illegitimate Names or Synonyms

Some botanists have tried to subdivide illegitimate names
or synonyms of a taxonomic group into names that are valid (or
legitimate) and invalid (or illegitimate). But they have been
unable to dissociate from these terms also the sense given in this
paper and have succeeded in creating a great deal of confusion
and equivocation on the matter; and as the Rules of Nomenclature
have not escaped from these confusing notions, it seems useful
before proceeding further to pause a while and inquire into the

Gardens Bulletin, S.S.

233

exact nature of the three main classes into which synonyms may
be subdivided :— ;

tr. ABSOLUTE OR EXACT SYNONYMS are illegitimate
names based on the same specimen, or a duplicate from the same
type collection or from the same type plant, as the one on
which the legitimate name was based. For the oldest name on
which all new names and new combinations have been based I
propose the term BASINYM, and for all the new names and new
combinations based on the basinym the term ISONYM.
Taxonomic groups published independently as new species, but
based on the type specimen of a valid name, or on a
duplicate of the type specimen, are called TYPONYMS. A
legitimate name may itself be a basinym or an isonym: if the
former, it will have at least one isonym as its absolute synonym;
if the latter, it will have one basinym as a synonym, though it
may also have in its synonymy one or more isonyms. In either
case there may also be one or more typonyms, in relation to which
the legitimate name itself might be regarded as a typonym.

The following two examples will show how these terms are
applied :—

(a) Both Unona cochinchinensis DC and Unona Desmos Dunal
were based on Desmos cochinchinensis Lour., the second as a new
name and the first as a new combination for the third name. VD.
cochinchinensis is, therefore, the basinym of the first two, both of
which are its isonyms. In this instance Vesmos cochinchinensis Lour.
is also the legitimate name under the genus Vesmos, the genus Unona
being reserved for American species. Had the generic name Unona
been applicable legitimately, Unona cochinchinensis (Lour.) DC would
have been the legitimate name, while the other two binomials (one
an Isonym and the other a basinym) would have to be quoted in the
synonymy.

(6) Actinophloeus cuneatus Burret (1931) was based on a specimen
derived from a plant cultivated in the Botanic Gardens, Buitenzorg,
Java, under the gardens’ no. V-—I-16. Quite independently of this
name, Ptychosperma tenue Becc. (1934) was described as a new
species having for its type another specimen derived from the above
mentioned plant from the Buitenzorg Botanic Gardens. Later on,
with the first species for its basis, the new combination, Ptychosperma
cuneatum (Burr.) Burr. (1935) was made. Here Actinophla@is
cuneatus Burr. is the basinym of P. cumeatum Burr. and conversely
P. cuneatum Burr. is the isonym of the former. Both these are
sande of P. tenue Becc., and vice-versa. (see also the example
in 2).

2. EQUIVALENT SYNONYMS have more or less the
same particular circumscription or concept as the legitimate
name, but are not based on the same type specimen, or its
duplicate. Some of these may have been published originally as
distinct from those to which they are now united, and possibly
may have been based on the paratypes or the lectoparatypes of
the species to which they are now reduced. Though they may
have been generally accepted as identical or equivalent synonyms,
some authors, particularly ‘‘splitter’’ botanists, may recognize in
them as many species in the Jordanian sense as there are names.

Vot. IX: (1637).

234

Such names, even when cited in the synonymy, have been called
legitimate names in the example given under Art. 60 (1) of the
Rules; but it is evident from the definitions given in this paper,
it is a misnomer to class them as such. They may be legitimate
if they are maintained as independent species; otherwise they
must be considered as illegitimate names.

Examples :—

(a) If, in the case of Cucubalus treated in the example under
Art. 60 (1) of the Rules, one follows Miller and considers C. latifolius
Mill. (1768) and C. angustifolius Mill. (1768) as two separate species
distinct from C. Behen L. ( 1753), then Miuller’s binomials are
legitimate; but if one follows other authors and regards Miller's
species as synonyms of C. Behen L., then both Miller’s names become
illegitimate for C. Behen L. and equivalent synonyms of it.

(6) Blume, working out the plants cultivated in the Botanic
Gardens, Buitenzorg, published Calla silvestris Bl. (1823), which later
became the basinym of Raphidophora silvestris Engl. (1879). Hooker,
studying the flora of the Malay Peninsula, found a Raphidophora
plant that was not credited for the region under his purview, and
which he described as 2. Wvayi Hook. f. (1893). Later Ridley,
examining the species of Raphidophora from Borneo, detected one
that was not recorded for the country, and proposed R. nigrescens
Ridl. (1905) as a name for it. Now it has been found out that,
though the types of each of these species were derived from three
different collections made in three different countries, nevertheless
all the three species represent one and the same species. As Calla
silvestris Bl. is the oldest of the three: binomials published as new
species, its isonym, fF. silvestris (Bl.) Engl., becomes the legitimate
name for the group under Raphidophora, with R. Wrayi Hook. f. and
R. nigrescens Ridl. as equivalent svnonyms and with C. szlvestris Bl.
as the absolute synonym.

3. Besides the two classes enumerated above there are
synonyms which consist of names based on monstrosities; and of
nomina dubia, confusa vel ambigua. These together with such
syhonyms of the above three classes as have been rejected by
the special decisions of the Congress (nomina rejicienda) may be
termed NOMINA EXCLUDENDA.

6. Legitimate Names (Arts. 69 and 60)

To the followers of the modern, so-called binomial, system
of plant nomenclature, there is no Article more important in the
new Rules of Botanical Nomenclature than Art. 69. It forms
the basis of all the other regulations framed with the view to
laying down the correct procedure to be adopted in putting the
principle of priority in force. Almost all the illegitimate names,
that could have been avoided by authors at the time of publica-
tion, have been published because the questions dealt with in this
Article have been much disputed even by the staunchest
supporters of the priority principle. And it is no exaggeration
to say that this Article, if it had been properly worded and
quoted, would have saved several botanists from being confused
by the Rules and would have perhaps rendered some other
Articles like Arts. 53-55 unnecessary or made them mere

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sub-divisions of Art. 69. Yet students of the Rules note with
vexation that this Article is given a secondary importance and
that it is even ignored in Articles where reference to it is
absolutely due. The chief defects in Art. 69 may be summarized
as foliows:

In the first place this Article, which forms a part or the
basis of Arts. 53—55, is misplaced so as to be easily overlooked by
botanists consulting the Rules for definite instructions on the
retention of names and epithets of groups below the rank of
genus (Sect. 9-11, Arts. 53-58), or even on the RETENTION
OF NAMES AND EPITHETS OF GROUPS BELOW THE
RANK OF GENUS ON TRANSFERENCE TO ANOTHER
GENUS OR SPECIES, which is the title of the Section 9,
Arts. 53-55. In the second place, Art. 69 is not referred to in
Art. 60 (1-5) where illegitimacy of names is defined; consequently
it might easily lead one to the erroneous conclusion that, though
the violation of Art. 54 renders a binomial illegimate [ Art. 60 (2) |,
the violation of Art. 69 does not. In the third place the terms
legitimate and illegitimate are used equivocally both in the text
and the example. And in the fourth place, Art. 69 leaves the
point ambiguous when, after discarding all names regarded as
not available or illegitimate under Arts. 60-68, there are yet
available for the taxonomic group more than one name that is
considered ‘‘not illegitimate’’ under Art. 60 (1).

In order to be able to discuss Art. 69 fully, it may be useful
to transcribe here the text and the examples in extenso :—

‘‘In cases foreseen in Art. 60-68 the name or epithet to be rejected
is replaced by the oldest legitimate name,* or (in a combination) by
the oldest legitimate epithet which will be, in the new position, in
accordance with the Rules. If none exists, a new name or epithet
must be chosen. Where a new epithet is required, an author may,
if he wishes, adopt an epithet previously given to the group in an
illegitimate combination, if there is no obstacle to its employment
in the new position or sense.

“Examples: Linum Radiola L. (1753), when transferred to the
genus Aadiola, must not be called Radiola Radiola (L) Karst., as
that combination is contrary to Art. 68 (3): the next oldest specific
epithet is mwltifiorum, but the name Linum multiflorum Lam. (1778)
is illegitimate, since it was a superfluous name for Linum Radiola L:
under Radiola the species must be called R. linoides Roth (1788),
since dimoides is the earliest legitimate epithet available. The
combination 7alinum polyandrum Hook. (in Bot. Mag. t. 4833: 1855)
is illegitimate, being a later homonym of 7. folyandrum Ruiz et Pav.
(Syst. Fl. Per. I, 115: 1798): when Bentham transferred 7.
polyandrum Hook. to Calandrinia, he called it Calandrinia polyandra
(Fl. Austral. I: 172: 1863). This is treated, not as a new
combination, but as a new name, ©. folyandra Benth. (1863)’’.

If the word legitimate in the text is taken in the sense
defined in the footnote to Art. 45, and, therefore, in the sense
given in this paper, then the Article ceases to be an instruction,
for a taxonomic group can bear only one name that is ‘‘strictly
in accordance with the Rules’’ (=legitimate). Should the Rules

* Italics are mine.

eat, BX :' €1937 Ja?

230

make it possible for a taxonomic group to bear more than one
legitimate name, then the element of simplicity, uniformity,
definiteness and stability will at once disappear from plant
nomenclature. The Rules do not admit, therefore, an old or
older, or the oldest, legitimate names.

If, as in Arts, 20, 56 and 60 (2), the word legitimate is taken
to mean valid as defined here, the first sentence of the text of
Art. 69 will have a meaning, but nevertheless the instructions
will be far from clear. In the first place it will be seen that
the word illegitimate in the text and in the examples cannot be
taken to mean invalid. As it might mean illegitimate in the
sense given to the term in Art. 60 it will be worth while, at this
stage to analyse this Article also.

It is obvious that the word illegitimate, as interpreted
in Art. 60 (2-3) means illegitimate; and as interpreted in Art.
60 (4-5) means invalid; both in the sense given here. But to give
a precise interpretation to it as used in Art. 60 (1) is an impossi-
bility, despite the definitions and examples given. I surmise
that in the first (Cainito) and third (Chrysophyllum sericeum)
examples, an illegitimate mame means either an invalid (or
illegitimate ?) name published in contravention to Art. 59, or a
later typonym, when the earlier name was legitimate; or perhaps
a new natmne published illegitimately when a new combination
was possible.

In the second example (Unisema), illegitimate might be taken
to mean illegitimate as defined in this paper; and in the fourth
example (Cucubalus) the phrase ‘‘ not illegitimate’’ might be
taken to mean not ‘‘not invalid’’ but illegitimate under certain
conditions and legitimate under others; for this is a case of a
synonym belonging to the EQUIVALENT class (example a).

With such different and ambiguous meanings given to one
word and in one Article, how can Art. 69 be clear and
unequivocal? Is Linum multiflorum Lam. referred to in the
first example under Art. 69 an invalid name (which it does not
appear to be) or a later typonym or isonym? Why after the
exclusion of Radiola, is the trivial name linoides ‘‘ the earliest
legitimate epithet available’, and why should the binomial
R. linoides Roth have priority over L. multiforum Lam.? Is
R. linoides Roth an equivalent synonym of L. RadiolaL? Ina
hypothetical case, if the species L. Radiola L,. had to be trans-
ferred again to a new genus where the specific epithet Radiola
could not be used, is the author effecting the transference obliged
to adopt again the later specific epithet linoides and definitely
reject multiflorum? Since from the text and the examples under
Arts. 60 and 69, it is not possible to give definite answers to
these questions, it is no wonder that botanists find the Rules as
confusing as ever, and go on creating new binomials knowing
fully well that many of them will have to be rejected later as
illegitimate.

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7. Three Customary Procedures Affecting Art. 69

I enumerate here three practices that are current among the
supporters of the priority principle, one of which may be legiti-
mized under the Rules. In this exposition I leave tautonyms and
names of unequal rank out of consideration. The reasons for
excluding these are:—(a) According to the definitions given
above, tautonyms are invalid names, and therefore, must not
enter into nomenclatorial considerations. (b) The practice of
taking into account names of unequal rank for the purpose of
priority is almost dead and is moreover, condemned by Art. 58
of the Rules. For the purpose of priority and legitimacy only
valid names of the same rank as the corresponding taxonomic
group under consideration are to be taken into account. The
procedures may be enunciated thus:

When there are available for a taxonomic group several
rival names of equal and corresponding rank, reiect all Nomina
Excludenda because their status is doubtful or ambiguous, or
because they have been excluded once for ail by special decisions
of the legislative body. After having done this, then adopt any
of the following procedures :

(x1) Of the remaining rival names, omit also the later
homonyms; then in the case of the names of a family or a genus,
the oldest name becomes also the legitimate name. In the case
of the subdivision of a genus, or of a species or a variety, the
epithet of the oldest name of corresponding rank becomes the
legitimate; but if this epithet cannot be used in the new, required
position, because of the resulting tautonym or later homonym,
then the oldest name of equal rank already existing in the
required position becomes the legitimate name for the group.
If none exists, the author is free to choose any epithet he likes,
being allowed also to use the epithet from any of the several
synonyms and homonyms, irrespective of their age, provided its
use does not create a tautonym or a later homonym.

(2) For the first part follow the procedure as enunciated in
(1); but in a new combination where the oldest epithet cannot be
legitimized for the taxonomic group below the rank of genus
because’of the resulting tautonym or a later homonym, the author
must adopt the epithet from the next oldest name that can be
used, without producing a tautonym or a later homonym,
regardless of the required position. If none exists, then the
author is free as in (z) to apply any legitimable epithet he
chooses.

(3) The third procedure is to include all valid names that are
not nomina excludenda, so that in the case of a taxonomic group
of the rank lower than a genus, later homonyms are also taken
into account. In the case of the names of families and genera,
the legitimate name is the oldest name of the corresponding rank
that is not a later homonym. In the case of names of the sub-

Vol. IX. (1937

238

divisions of a genus, or of the species, and the varieties, choose
as the legitimate epithet for the group, the one from the oldest
name of corresponding rank, unless the epithet thus chosen either
is in the required position a later homonym, or when transferred
to the required position, produces a tautonym or a later
homonym. If either of these obstacles exists, then the next
oldest epithet that can overcome both these obstacles must be
used as the legitimate epithet for the group. If none exists, then
the author is free to choose any legitimable expithet he likes.

Examples :-—

(a) Assuming Linum Radiola L. (1753), L. multifiorum Lam,
(1778) and Radiola linoides Roth (1788) given in the example under
Art. 69 to be all valid names, F. linoides Roth is the legitimate
name for the species according to the first procedure, because the
oldest specific epithet cannot be used under the genus Radiola, and
because FR. linoides Roth is the oldest name under Radiola. In the
next example Bentham was not obliged to adopt the specific epithet
polyandra when he transferred 7alinum polyandrum Hook. to the
genus Calandrinia because 7. polyandrum Hook. was a later
homonym.

(6) According to the second procedure, one is bound to adopt the
epithet mzultifiorum under Radiola, because L. multiflorum Lam. is
the next oldest name; but one is not obliged to adopt the specific epithet
polyandrum under Calandrinia, for Talinum polyandrum Hook.,
because this was a later homonym.

(c) According to the third procedure, the author is bound not
only to adopt the specific epithet wzwltiflorum under Radiola, but also
to use the epithet polyandra under Calandrinia, because Talinum
polyandrum Hook. is the oldest valid name for the species and the
specific epithet can be legitimized under Calandrinia.

The first procedure is not followed generally by strict
supporters of the principle of priority. It seems probable that,
in drafting Art. 69 of the Rules, the legislators had this procedure
in mind. The second may be considered a reasonable procedure,
but it has never received any great support from the followers
of the priority principle. The third procedure is the most
favoured of all the three practices. And as it follows, from
the provision made at the last Congress (1935) for validating and
legitimizing alternative names (see below for the definition) even
though the basinym is a later homonym, that the practice of
basing a new combination on a later homonym cannot be entirely
condemned, the third procedure appears to me the most natural
outcome of putting the principle of priority in practice.- Besides
under the priority principle the epithet for the correct rank of a
taxonomic group is of greater importance than its disposition in
the correct genus or family.

One of the principal reasons why some botanists object to
the second and the third procedures indicated above is that they
think that it should never be possible to legitimize a name
published in contravention of the Rules. But even if the first
procedure was adopted as a rule, it would not be possible to

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achieve this object in every case, as it will be seen from the
following hypothetical case:

Supposing Linum multifiorum Lam. in the above example was
a valid and legitimate species quite distinct from Linum Radiola L.
and Radiola Linoides Roth, and Roth, in transferring that species to
the genus fRadiola, had used the epithet pauciflorum, rejecting
multtflorum as unsuitable for the species, then Roth’s Radiola
paucifiorum nom. nov. would contravene Art. 59 of the Rules. Now
if someone were to publish the name Radiola multifiorum for a new
species, then #. multifiorum (Lam.) comb. nov. would not be
legitimable, and, owing to the reservation of this binomial for the
new species, 2. pauciflorum nom. nov., although published in contra-
vention of the Rules, would become automatically the legitimate name
for L. multifiorum Lam. under the genus Radiola, unless legislation
ie passed invalidating all the names published in contravention to

rt. 50.

8. What a New Combination or a New Name is

These terms are so familiar to any taxonomist that it might
appear fatuous on my part to try to explain them here. But
as they are not defined in the Rules and as in one instance at least,
they seem to have been mis-stated, it appeared to me that a con-
sideration on the matter would not be out of place.

Both a new combination and a new name are validated by
citation so that both these terms imply the existence of a basinym
on which each is based, though, as explained above, a basinym
may have, for its isonyms, both a new name and a new combina-
tion at the same time. But all basinyms must be valid names;
consequently a species, published by referring to an invalid, or
pre-Linnean, name, figures and description cannot be called
either a new combination or a new name, but must be called a
new species. If the specific epithet in the basinym and in its
isonym is the same, then the isonym is called a new combination;
if the specific epithet in the basinym is different from that of its
isonym, then the isonym is a new name. According to this
definition, Calandrinia polyandra Benth. in the second example
under Art. 69, must be called a new combination based on

Talinum polyandrum Hook., and not a new name as stated in
Art. 609.

9. Alternative Names

Now alternative names (nomina alternativa seu eventualia) are
generally new combinations published simultaneously with the
basinym, though there is nothing in the new provision passed at
the last Congress, to deter one from making alternative names
that are new names (vide infra). In the example of alternative
names discussed at the last Congress, the alternative name
Andropogon Bequertii De Wild. was quoted as nom. nov. of
Cymbopogon Bequertii De Wild. sp. nov.; but the distinction
between new names and new combinations was not considered.

The practice of publishing alternative names has had an
origin which, though not made clear at the last Congress, appears

Vol. IX. (1937).

240

to me quite sound. There are genera like Andropogon and
Cymbopogon, Ptychosperma and Actinophleus, and Pasania
and Lithocarpus, where it is not yet possible to decide definitely
whether it is better to split a genus into two or more genera, or to
keep it entire and divide it into so many sections or subgenera.
Specialists find as many good reasons in favour of the first
alternative as they do in favour of the second. Now if an author
accepted such a genus sensu lato and published new species in
a section of such disputable status, a botanist in search of easy
honours might at once seize the opportunity, whether he had
himself studied the group or not, to make new combinations under
the alternative and disputed genus based on the binomials of the
first botanist; or vice-versa, had the specialist published the new
species under the latter genus, it would not have deterred the
second botanist from making new combinations under the former.
In such a case, though the second botanist might have no opinion
of his own on the genera and might even be utterly unacquainted

with them, the Rules would oblige one, as they do even now, to .

accept the new combinations made by him. It is to curtail such
vexatious activity, and also to keep the honour where it is due,
that the practice of publishing simultaneous isonyms or alterna-
tive names has arisen. Therefore it was but just that it was
validated by the last Congress.

I think it is necessary to make additions to this amendment
prohibiting alternative names that are not new combinations and
also alternative names of a different rank from the basinym.
According to these provisions Conyphyton or Minima could not
be proposed as an alternative name to Mesembryanthemum sect.
Minima, nor Cymbopogon fluminense var. riparium to Cymbo-
pogon riparium, as these names belong to different ranks; neither
Cymbopogon riparium or Andropogon riparium could be proposed
as alternative to Cymbopogon fluminense, because the trivial
epithets are different. Without such an amendment, new names
or new combinations of equal and unequal rank could be proposed
as alternative names.

19. Rival Names of Equal Age and Equal Rank
(Arts. 56 and 61)

When for one species there are available two or more valid
names of equal age, then Art. 56 instructs us to follow the
botanist who first preferred one of them as the legitimate name
for the species. It should have been added here that, if the
name first chosen cannot be legitimized, then the author who is
first to choose the next legitimable name must be followed.

When one and the same name is used simultaneously for
more than one species, according to the new addition to Art. 61,
“the first author who adopts one of them, or substitutes another
name for one of them, must be followed’’.

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In cases foreseen in Arts. 56 and 61 I think it is necessary
to give new combinations a precedence over new names; and
these over new species. Again when homonyms of equal age
are isonyms of equal kind, those which have older basinyms
should be preferred. The reason for this preference is that,
though the age of a binary or ternary name is counted from the
time it is published in its correct position and rank in a genus
or a species, the age of the epithet itself is counted, according
to the priority principle, from the date of its first publication in
the same rank for the group but independently of the position
regarding its family, genus or species, so that when a new species
is older than a homonymous new combination, the former takes
precedence; but when they are of equal age, the principle of
priority should oblige one to give precedence to those isonyms
which have the oldest basinyms.

Thus, whenever Colocasia esculentum (L. 1753) Schott (1832) and
C. antiquorum Schott (1832) isonym of Arum colocasia L. (1753)
are regarded as synonymous, the former should be chosen as the
legitimate name for the taxonomic group thus constituted. So also
whenever one of these is to be reduced as a variety of the group, it
would be the new species, and not the new combination, that would
have to be relegated to the varietal rank.

Further it is also necessary to add here that the choice made
under Arts. 56 and 61 holds effective so long as the species or
names remain in the same genus under which the choice was
made; but on their transference to another genus, the author who
first made the distinction in the new position must be followed,
provided this distinction does not conflict with the rule of
precedence.

Thus, even when someone had discovered that Avum colocasia L.
(1753) and A. esculentum L (1753) were synonymous and reduced the
latter to a synonym of the former, Schott (Syn., 1856, p. 41) would
not have been justified in reducing A. esculentum L. (1753) to a
synonym or variety of Colocasia antiquorum Schott (1832)=isonym
of A. colocasia L. (1753); for such a reduction is inadmissible under
the priority principle, as it would mean giving to a younger epithet
precedence over an older one, when the latter is legitimable in the
required position. Hence Hubbard and Rheder (Bot. Mus. Leafl.
Harvard Univ. I, 1932, No. 1 p. 5) are justified in publishing the
combination, Colocasia esculenta (LL) Schott (1832) var. antiquorum
(Schott) Hubb. et. Rhed. (=Colocasia antiquorum Schott) without
first inquiring into the history of their respective basinyms, A7vum
esculentum L. and A. colocasia L., But it seems to me that the
combination Colocasia esculenta (L) Schott. var. colocasia comb. nov.

(=A. colocasia L.) would not have been contrary to the Rules as
they stand. (see also the examples in the next section 11).

11. Isonyms of Unequal Age based on Basinyms of Equal Age

As proposed above, under equal circumstances isonyms that
are new combinations should have precedence over new names.
But the case where the basinyms are of equal age but their
isonyms are of different ages, has so far remained untreated.
From a study of the recent works of eminent taxonomists, it is
evident that there is no fixed procedure for dealing with the

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case; but, as a general rule, I think that the isonym which is
the older or the oldest should take precedence. Thus, of names
Bruguiera gymnorrhiza (1,.) Lam. (1797) and B. conjugata (L.)
Merrill (1914) which were based on basinyms of equal age,
Rhizophora gymnorrhiza L. (1753) and R. conjugata L. (1753),
botanists should, in my opinion, be obliged to adopt B.
gymnorrhiza (L.) Lam. as the legitimate name whenever the two
species are regarded as conspecific. At least that has been the
procedure long current among the strict supporters of the
priority principle, though some botanists of high reputation have
now begun to use the younger of the isonyms as the legitimate.

However this rule of procedence, if followed in every case, might
seem to give rise to difficulties. Suppose, before Lamarck made his
combination, B. gymnorrhiza, a botanist had detected the identity of
the two species and reduced 2. gymnorrhiza L. as a synonym of
R. conjugata L., then under Art. 56, R. conjugata L. has to be
retained as the legitimate name whenever the two are retained under
the genus Rhizophora and are considered as conspecific; but it does
not say anything if Lamarck, in transferring the species to Bruguiera,
had inadvertently or deliberately, made the new combination with
R. gymorrhiza L. as the basinym and with or without 2. conjugata L.
as asynonym. To some this combination might seem illegitimate and
they might be tempted to consider, under such circumstances, B#.
conjugata (L) Merr. (1914) as the legitimate name. But not only
under the rules of precedence but also under the most common pro-
cedure affecting the priority principle, Bruguiera gymnorrhiza (Linn.)
Lam. (1797), being the older of the two isonyms of the basinyms of
equal ages, is the legitimate name under Aruguiera.

Such cases mentioned here and in section ro are extremely
rare; but they show the effectiveness of the strict application of
the rule of precedence in preventing usages that are contrary to
the accepted principles of nomenclature and in ensuring simpli-
city and uniformity in the solution of such complicated problems.

12. Varietal Names

Of late there has been a tendency to regard a specific name
(even when the species itself was originally described from a
single type specimen) as a conglomeration of varieties and then
to give each variety a separate name. In naming the varieties
two common procedures exist: the first is to call var. genuina,
typica, primitiva, communis, etc. the form that the author, who
makes the separation into varieties, considers as the most
primitive or the most common of the varieties, it being
immaterial whether this form agrees with the type of the species
or not. If the type form does not agree with the so-called
primitive or common form, then the type form receives another
varietal name. ‘The second procedure is to call var. typica,
genuina or originaria the type form (i.e. the form that agrees
with the type specimen) and to give new names to other forms.
The latter procedure is approved in the Recommendation XVIII
of the Rules, while the first, though not expressly condemned,
is being abandoned by botanists. But I consider that even the

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second procedure is wrong and, if admitted as valid, gives rise
to combinations that are contrary to the spirit of the Rules
themselves and that might not be in accordance with either of
the procedures mentioned above.

Thus Colocasia esculenta (L) Schott var. antiquorum (Schott)
Hubbard et Rheder (1932), being a later isonym of equal rank for
C. antiquorum Schott var. tyfica Engl. (1879), would have to be
called C. esculenta (L) Schott var. tyfzca (Engl.) com. nov., a com-
bination that might not be applicable either to the type of the species
or to the most primitive or the most common form. Further since the
Rules themselves do not condemn the procedure which admits giving
a new varietal epithet to the type specimen of a species, while using
such terms as typizcus, primitivus, or communis for the form which
is either the commonest in the wild state or the most primitive, it
appears that one cannot consider invalid the trinomial Aglaonema
angustifolium N. E. Brown var. undulatum Ridley (Mat. Fl. Malay.
Pen., Monocot., III, 1907, p. 21), although the varietal name is given
exactly to the same form as represented by the type of A.
angustifolium N. E. Br. (1895).

Engler reduced Aglaonema angustifolium N. E. Br. (1895) to a
synonym of A. Schottianum Mig. (1856) and named the form
represented by Brown’s type (A. angustifolium var. undulatum
Ridl.) as A. Schottianum Mig. var. Brownti Engl. (Pflanzenreich,
hft. 64, 1915 p. 21). But under the priority principle this varietal
name of Engler’s cannot be the legitimate one, since of the two
varietal names, uwudulatum (1907) and Brownii (1915), the former is
both legitimable under A. Schottianum Mig. and earlier.

Further Ridley mentions the ‘‘typical form’’ (z.e. the common
form) of the species, A. angustifolium N. E. Br., but does not give
it a name. Had he called it var. typizcum, then on its transference
to A. Schottianum Miq., we should have, side by side with A.
Schottianum Miq. var genuinum Engl. (1915) containing the type of
the species, A. Schottiamum Mig. var. typicum comb. nov., not
containing the type, and with the possibility that neither of the
varieties represented the commonest form of the species when viewed
in the broad sense.

The other alternatives are either to repeat the specific epithet
(recommended under the Rules, Rec. XVIII), or to regard
C. antiquorum Schott as equivalent to C. antiquorum Schott,
subsp. antiquorum Schott, var antiquorum Schott, forma
antiquorum Schott. In the last case the Recommendation
XXXVI (2) will automatically become a rule and oblige botanists
to choose antiquorum as the legitimate epithet whenever
C. antiquorum is reduced to a variety or a subdivision of another

species. This would necessitate an alteration in the text of
Art. 58.

But to admit such names as Colocasia antiquorum var.
antiquorum or C. antiquorum var. genuinum, and Aglaonema
angustifolium var. augustifolium, A. angustifolium var.
undulatum, or A. angustifolium var. typicum (or genuinum etc.)
all referring respectively to the type of Colocasia antiquorum
Schott and Aglaonema angustifolium N. EK. Br. would be tanta-
mount to adopting ternary specific names in modern botanical
nomenclature and to making the present obligation to use the
binary names for species a farce or non-obligatory. Further, an

Kot. BX) Cias7)

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acceptance of such ternary names will, in turn, have to be
followed by the removal of the ban on all those books that are
now condemned for not using the binary specific names in their
nomenclature.

In my opinion, therefore, all such ternary names, when
applied to the type of the species and retained under it, should
be condemned as being contrary to the binomial system of
nomenclature. Though Aglaonema angustifolium N. E. Br.,
could in sensu lato be used to include all the subspecies and
varieties described under it, in sensu stricto it must be applicable
the Holotype and the plants represented by it: all trinomials
with A. angustifolium as the first part being invalid for this form.

13. Nomen Dubium or Mixtum Compositum* (Arts. 52, 62-65)

When a species is to be divided into two or more species,
Art. 52 of the Rules provides for the retention of the specific
epithet to the type designated by the author. But in the case

* A Taxonomic Name must indicate only one element of its
rank. A Mixtum Compositum is a taxonomic name which by
definition covers more than one element of the same rank, or to which
more than one element of the same rank has been assigned in the
original description. All plants belonging to one form, or variety,
or species, or genus (etc.) make, in the opinion of a botanist, one

ELEMENT of the same rank. There are two kinds of Mixta
Composita:

(1) NEGATIVE MIxXTUM COMPOSITUM: a name whose description
is SO vague as to be applicable to more than one element. These may
be subdivided according to their types as follows:

(a) types represent more than one element, but the holotype is
indicated ;

(6) types represent. more than one element, but the holotype is
not indicated (=xomen dubium) ;

(c) types are not extant (=nomen dubiun).

(2) Positive MIxTUM COMPOSITUM: either (A) a taxonomic
name whose description includes discordant characters, each set being
applicable to one element, and whose types are either

(a) not extant (=nomen confusum) ;
(6) or if extant, then either a holotype is not indicated (=nomen
confusum) ; or
(c) a holotype is indicated.
or (B) a name whose description, though not containing discordant
characters, agrees only partly with the indicated holotype.

If in (1) all syntypes represent one element, then such a
taxonomic group is excluded from the category of a mixtum com-
positum; if in (1) and (2) the types are extant and a holotype is
indicated, the Rules oblige us to interpret the name with reference
to the indicated holotype, even when this type is not the one that
agrees best with the description. If types are extant and do not
agree with the description at all, then the probability is that some
types are lost, or that the author has used a wrong terminology to
describe the characters of the type specimens. It is only when the
primary types are not extant, or when extant one of them is not
indicated as a holotype, that the difficulties arise in interpreting
satisfactorily a mixtum compositum. In common parlance a mzxtum
compositum means a name of any of the two classes: (2-A-a) and
(2-A-0).

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245

where the type has not been designated, regulations for choosing
the lectotype have not yet been made; so that it is quite easy at
present to reject a species whose syntypes represent more than
one species, as a nomen confusum, dubium vel ambiguum
(Arts. 62-64).

Yet if one were to apply these provision (Arts. 62-64) in every
case, where a specific description was drawn from syntypes
representing more than one species, I think the majority of the
specific names from the time of Linnzeus down to the end of the
last century will have to be discarded, and a similar fate might
also await several names published during the last thirty or forty
years.

The usual procedure in such cases has been to retain the name
for that one of the syntypes that agrees best with the description,
and to exclude from the species all other specimens that do not
correspond with the selected syntype (=lectotype). Should
there be more than one such specimen having equal claims to be
selected as the type and should they represent more than one
species, the first author to choose one of them as the type must,
in my opinion, be followed. This principle is also in accordance
with the provisions made in Arts. 56 and 61 for selecting one
name as the legitimate from a number of rival names of equal
age.

If this procedure be legitimized, a large number of species
having now a disputable status will become clear and stable,
thereby diminishing the number of the nomina ambigua, dubia
vel confusa. In fact I think that the Appendices for Nomina
Ambigua and Nomina Confusa should not be prepared unless
this procedure, and the one recommended for interpreting the
species validated by references to invalid but effective literature
(Class II), are found unworkable (which might be only in a very
limited number of cases). The principle if adopted will induce
also a better research into the status of old species and it will
avoid a haphazard rejection of species on the basis of our
ignorance.

Examples :-—

(z) Under Cerbera Manghas L. (1753) Linnzus cites several
entirely discordant elements which have made some botanists either
reject the species as mixtum compositum, or interpret it with reference
to one or other of the discordant elements that do not agree with the
description. It is true that Linnzus cited several references that are
not identical with the type on which the description was based; but
in doing this Linneus was keeping up the procedure of the times
which, perhaps with the intention of helping the students of the local
floras, made botanists include in the synonymy all the references
that they not only considered as equivalent, but even suspected as
approximate, synonyms. In my opinion the ambiguity or confusion
has arisen on this Linnean species owing either to the failure to
notice that the lengthy specific description of Linnzus was derived
from Osbeck’s specimen from Java, or to the imperfect acquaintance
of some botanists with the genus; for it is obvious that the Linnean
description is not applicable to C. odollam Gertn. at all but is

Vol. IX. (1937).

246

exactly C. lactaria Ham. ex DC. Much harm would result if, owing
to imperfect acquaintance with genera or species concerned, or owing
to failure to account correctly for the type specimen, one were to

reject a good species like C. Manghas L. as a nomen ambiguum, —

confusum, vel, dubium, or to conserve another name against it.

(2) Under Poa amboinica L., Mantissa II, 1771, p. 557, Linneus
gave a specific description which does not agree either with the plate
or with the description of Phanix amboinica montana Rumph. (Herb.
Amboin. VI p. 19 t. 7 fig. 3), a reference given by Linnzus in the
synonymy of the species. In 1786 Retzius pointed out these dis-
crepancies and excluded the Rumphian species from P. amboinica L.,
retaining this name for a South Indian plant which was distributed
under the name by Keenig, and with which the Linnean description
agreed. It is not known whether Retzius had seen the actual type
specimen, or its duplicate, on which Linnzus based his description,
nor does he give any reasons why he interpreted the species by
reference to Koenig’s specimen. There is also no specimen in the
Linnean herbarium to decide this matter. But seeing that Retzius
was a contemporary of Linnzus and Koenig, and that Koenig corres-
ponded with several botanists in Europe including Linnzus and
Retzius, it is not unreasonable to suppose that Retzius had clues, now
lost to us, that warranted his interpretation by reference to Koenig’s
specimen. Mr. C. E. C. Fischer, who inquired into this subject, also
supports this conclusion. (Kew Bull. 1934 p. 398). But the absence
of any obligation for botanists to follow the first botanist (in this
case Retzius) who selected the lectotype or neotype of Poa amboinica L.,
has made some take the Rumphian figure as the type of the Linnean
species, causing thereby instability and lack of uniformity in the
nomenclature of the two species, and has made others favour the

rejection of P. amboinica L. either as nomen ambiguum or nonten
confusum.

The most difficult problem that arises in stabilizing plant
nomenclature is in regard to species, the types of which are not
extant and the original description of which seems to be
inadequate for the correct specific identification. In such cases
it is customary to identify the species by interpreting their
botanical descriptions on plant-geographical considerations. But
if, by such a procedure, one cannot definitely accept or reject
a given interpretation of such a species, then one should be
obliged to follow the earliest interpretation that is not obviously
discordant with the original description. It should be prohibited
to interpret the species merely on the vernacular names, and the
economic uses, enumerated in the original descriptions, and to
make changes accordingly in the specific nomenclature. The
particulars about the vernacular names and the economic uses
of a species may supply a clue to its ultimate identification on
a scientific basis, but they cannot be accepted as decisive, or
even as a part of the botanical description. In this field it is
desirable to restrict the activities of those who rely on such
means because their activities are often a source of great confusion
in plant nomenclature and lead to instability in the nomenclature
of the species that had otherwise definite and stable botanical
names.

I enumerate here a list of principles which have proved
useful to me in the interpretation of a mixtum compositum, or

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247

a doubtful species whose types are not extant. It might be worth
while to give these a trial in order to test their universal utility :

A. When a description covers more than one species or
varieties (or element), then give more importance to:

1. the characters mentioned both in the body of the
description and in the remarks than to those
mentioned under the remarks only and not in the
description, when under the remarks the author
recognized more than one form of the species sensu
lato. The probability is that the characters men-
tioned only in the remarks have been deliberately
excluded from the body of the description because
they belong to one of the subordinate groups, not
to the typical form;

2. the characters mentioned in the body of the description
than to those mentioned in the remarks, if the
remarks refer to a form or variety different from
the one described in the body (e.g. Croton polot
Burm.f.);

3. the characters mentioned in the description than to
those not mentioned in it but figured in the plate
published simultaneously with the description, even
though these characters have been cited in the
remarks as belonging to one of the subordinate
groups constituting the main group;

4. the characters recognizable by the senses of touch and
vision than to those recognizable by the sense of
taste and smell;

5. the characters referring to twigs, leaves, flowers and
fruits than to those referring to the entire plant,
and observable only in the field;

6. the characters visible in the dry state (and referring to
twigs, leaves, flowers and fruits) than to those
observable in the field or in the living state, except
when there is a clear evidence to show that the
author studied the plants in the field and not in the
herbarium, in which case both sets of characters
could be given an equal value (as in Blanco’s Flora
de Filipinas, Loureiro’s Flora Cochinchinensis);

and 7. characters observable on the plant itself than those
referring to its economic uses (cf. also C. 2).
B. Attach equal importance to:

1. the specimens cited but not figured and to those that
are both cited and figured. Very often the
specimens studied are not the ones that are figured;
and very often, too, figures are not drawn by the

Vol. IX. (1937).

248

authors themselves, but by their draughtsmen. Out
of these specimens choose as the lectotype the
specimen that agrees best with the description;

2. the specimens cited but not figured and the specimens
not cited but figured, provided there is clear evidence
to show that the description covers both. (e.g.
Polypodium sessilifolium Hk., the type of which
is taken to be the specimen figured, although it was
not cited; see Gard. Bull. S.S. IX p. 134.)

and 3. the characters observable in the field like the colour,
height of the plants and the thickness of the main
trunk and those observable in small specimens
consisting of twigs, leaves, flowers, and fruits,
provided there is a clear evidence te show that the
author drew his specific description in the field and
not in the herbarium.

C. Disregard altogether any information or particulars
regarding :

I. vernacular names;

2. such economic uses as do not give any clue as to the
nature, size, and colour of the plants;

and 3. any part of the plant if there is a definite evidence to
show that the author derived that bit of information
from others or from books and herbarium labels;

but these actions are to be taken only when the particulars on
any of these points introduce the discordant element in the
description, or render a decisive interpretation impossible.
These particulars, if taken into account, must not be made the
principal basis for rejecting an accepted interpretation of a
species.

D.—(1) If a mixtum compositum (positive) covers two or
more species or elements of a rank higher than that of species,
then follow the earliest interpretation that takes into account all
the characters of greater importance, even if this interpretation
does not agree with the characters of lesser importance. In the
interest of stability in the names of families, genera, and species,
a consideration of characters of lesser importance (such as those
observable in the living state only and as undergo change during
drying) must not be admitted as a plea of sufficient importance
for rejecting an earlier interpretation because such a rejection
might affect profoundly the long accepted nomenclature of
several species, genera, etc.

(2) If a mixtum compositum (positive) consists only of
different varieties or of forms of lower rank than species, then
accept the earliest interpretation that takes into account both
the important and the less important botanical characters.

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(3) If itis a question not of a (positive) mixtum compositum,
but of an apparently indifferent description that seems applicable
to more than one element, and whose types are not extant, or if
extant represent more than one element, then the botanical charac-
ters of lesser importance must also be considered, and the earliest
interpretation that fits in with all the botanical characters given
in the original description and with plant geographical considera-
tions must be accepted. A ruling to this effect might discourage
interpretations of old, doubtful species without an adequate study
and reason.

E. If the name of a mixtum compositum, positive or
negative, has been long overlooked by botanists, or if it has been
excluded as a species of doubtful affinity or as a nomen dubium,
or confusum, it should not be used for any taxonomic group
without adequate botanical reasons, or proofs (vide infra), when
the adoption of such a name leads to changes in the accepted
nomenclature of species and genera. The very fact that the
name of a doubtful species has long been cited in the synonymy
must not be taken as a reason of sufficient importance to adopt
the name under the principle of priority. Such names have been
frequently cited in the synonymy of a species, not because the
mames are identifiable, but because the authors citing them
suspect them to be identical and hope the citation may form a
clue to the ultimate identification of the mixta composita on
botanical grounds.

NOTA: By PROOFS or ADEQUATE BOTANICAL
REASONS are meant the reasons why the name must be clinched
to a particular taxonomic group and not to any others, or how
the description published under the name agrees with that of the
taxonomic group identified with it. Thus I would consider
Mischophleus vestiaria (Giseke) Merrill, Interpr. Rumph. Herb.
Amb. (1917) 121 and Ormosia pinnata (Lour.) Merr. in Lingnan
Sei. Journ. XIV (1935) 12 as examples of interpretations un-
accompanied by any reasons except by the writer’s statement of
his convictions, while that of Cerium spicatum Lour. and Aidia
cochinchinensis Lour. by Moore in Journ. Bot. LXIII (1925) 250
as examples of interpretations accompanied by proofs or adequate
botanical reasons.

14. Tautonyms and Homonyms

The regulations concerning tautonyms are embodied in the
following extract of Art. 68 (3):

‘Specific epithets are illegitimate in the following special cases

and must be rejected.————(3) When they exactly repeat the
generic name with or without the addition of a transcribed symbol
(tautonym)—————_Examples—————_(3) Linaria Linaria, Nasturtium

Nasturtium-aquaticum.”’

From this it is evident that tautonyms come under the
regulations enacted as essential for the words constituting a

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250

specific name, and, therefore, are to be regarded as invalid names.
And so far as the wording goes, this rule appears to be complete
in itself, though it is not clear under what provision the binomial
Nasturtium Nasturtium-aquaticum is to be regarded as a
tautonyin, unless of course aquaticum is considered as a symbol.
But if one considers this Article in conjunction with the regula-
tions on the orthography of names (Art. 70), there appears to be
much room for equivocal interpretations.

That the reasons for asserting this may be better followed
I transcribe here some parts of Art. 70 that are to the point :

‘When the difference between two generic names lies in the
termination, these names must be regarded as distinct, even though
differing by one letter only. This does not apply to mere orthographic
variants of the same name.”’

‘‘Note 3: In deciding whether two or more slightly different
names should be treated as distinct or as orthographic variants, the
essential consideration is whether they may be confused with one
another or not: if there is serious risk of confusion, they should be
treated as orthographic variants. Doubtful cases should be referred
to the Executive Committee.”’

‘‘Note 4 :—————Epithets bearing the same meaning and differing
only slightly in form are considered as orthographic variants. The
genitive and adjectival forms of a personal name are, however,
treated as different epithets (e.g. Lysimachia Hemsleyana and
L. Hemsley)’.

Further we are told that generic names like Rubus and Rubia
Peponium and Peponia, Iris and Iria, Gerrardina and Gerardiina
Desmostachys and Desmostachya, or Peltophorus and Peltophorum
are not pairs of homonymous names. This appears surprising,
because one would think that they should come under the
provision made in Art. 70, Note 3, which permits one to treat
a name as an orthographic variant even wher a name or epithet,
owing to its close similarity with another of the same rank but
of different origin, is liable to be confused with the latter. Yet
we are told that such words as napaulensis, nepalensis and
nipalensis; heteropus and _ heteropodus; Bradlea, Bradleia,
Bradleja and Braddleya; trinervis(-e) and trinervius(-a, -um); are
groups of orthographic variants. The first two groups of epithets
are regarded as orthographic variants, apparently because they
mean the same thing, and because they are derived from the
same word, or because they refer to the same place; the third
group of words because they all refer to the same person and
because each of them has been spelt by subsequent authors both
as Bradleia and Bradleya, and this in spite of a ruling that changes
in the original spelling are not to be recognized under Art. 70;
but the last one, though it has the same meaning, has two
different roots and belongs to two different declensions in Latin.

If Bradlea Braddleya is a tautonym because the second
epithet is an orthographic variant of the first, the question arises
whether such combinations as Desmostachys Desmostachya,
Gerrardina Gerardiina, Cherefolium cerefolium L,., and Kandelia

Gardens Bulletin, S.S.

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251

Candel (I,.) Druce are tautonyms? The principles regulating
the homonymous generic names, if applied to these cases, would
admit these as valid names and not reject them as tautonyms,
though under usage, and perhaps under the provision in the
Note 3 of Art. 70, one would be able to reject many such names
as tautonyms. Further if Nasturtium Nasturtium-aquaticum is
a tautonym, are Radiola Radiolaris, Leontice Leontopetaloides,
Ficus syco-morus and Pinanga Pinangifolia also tautonyms? If
such names as Desmostachys Desmostachya or Gerrardina
Gerardiina are not regarded as tautonyms, despite the provision
in Art. 70, Note 3, I see no reason for outlawing such binomials
as Bradlea Braddleya, and Radiola Radiolaris, or even Radiola
Radiola.

In this matter usage, too, has been very erratic, even amongst
botanists who have shown manifest adhorrence for tautonyms;
for, while they readily discard as tautonyms such binomials as
Normanbya Normanbyi (F. Muell.) L. H. Bailey (1935) and
Sullivantea Sullivantii Britton (1894), they admit without any
protests binomials like Cuminum Cyminum L., Kandelia Candel
{(L.) Druce (1914) and Carum Carvi Ll. (the last being also spelt
as Carum Carui.).

Similarly the regulation pertaining to homonyms is also
unsatisafctory. Article 61 defines a later homonym as a naine
which ‘‘duplicates a name previously and validly published for
a group of the same rank based on a different type. Note:
Mere orthographic variants of the same naime are treated as
homonyms, when they are based on different types.—See Art 70.”’

Does this means that binomials like Baccaurea malayana and
B. malayensis, or Rhizophora Candel L. and Rk. Candellaria DC
ate to be treated as two pairs of homonymous names, on the
ground that they bear the same meaning and differ but very
slightly in form (Art. 70, Note 4)? If trivials epithets like
Ceylanica (from Portuguese Ceylao), Zeylanica (from Dutch
Zeylan) and Ceylonica (from the English Ceylon), or like trinervis
(third declension in Latin) and trinervius (second declension in
Latin) are to be taken as homonymous, because they mean the
same thing, are epithets like Ceylonica, Singalensis, Singalana,
Sinhalensis, Singalica, Cingalensis and Ceylonensis; Brownet,
Brownti, Morenii and Brunonis; Sinica and Sinensis; Hispanien-
sis, Hispanus and Hispanicus; Javana, Javensis, Javanensis and
Javanica; Aquillaria and Agalocha; van-der-Brecke and
Paludanus; Sahare and Saharana (both referring to the country
Sahara); Anamitica and Annamensis; Celebica, Selebica and
Celebesiana; to be taken as homonymous groups? ‘These
questions do not appear to have been settled by the Rules, and
the examples given in the Rules do not apply to such cases.
The recent usage on this point is also very contradictory.

Vol. IX. (1937).

252

The treatment of homonymous generic names in Kew
Bulletin (1935) 341-544 also shows a complete absence of
uniformity in the principles adopted by different contributors to
the Bulletin, thus confirming my contention that the Rules are
equivocal on the subject of homonyms and tautonyms.

_ Thus Prof. Alfred Rheder (Arnold Arboretum) regards Andr-
zetowskia Reichb. (1823) and Andreoskia DC. (1824) as orthographic
variants, because both genera were named in honour of A. Andr-
zeiowski. If this principle is adopted as a general rule, it would
appear that the Rules do not permit naming more than one genus
after the same person.

But Miss M. L. Green (Kew) considers Wendiza Meyen (1834)
not homonymous with Wendia Hoffm. (1814), though the latter name
was ‘corrected’? by De Candolle to Wendtia in 1830, that is, four
years before Wendtia Meyen was published, and though both W endtia
(1834) and Wendia (1814) appear to have been published in honour
of the same person. Apparently Miss Green has overlooked the case
of Bradlea, Bradleia, Bradleja and Braddleya discussed in the Rules,
where we are told to consider these four generic names as mere
orthographic variants, not only because they commemorate the same
person, but also because each of them has been spelt by subsequent
authors indifferently both as Bradleia and as Bradleya, and ‘‘only
one can be used without serious risk of confusion.”’

Further Dr. R. Mansfeld (Berlin) treats Pentaceros Mey (1818),
spelt later as Pentaceras by Roem. et Schult. (1819), as if it is
homonymous with Pentaceras Hook. f. (1862); Pentagonia Benth.
(1844) with Pentagonium Schau. (1843); and Psilanthus Hook f.
(1873) with Psilosanthus Neck. (1790). And yet, in the Rules, Pepo-
nium and Peponia, Symphostemon and Symphyostemon, Elodes and
Elodea, are cited as examples of distinct, non-homonymous names.

Then Prof. A. Rheder is not sure whether Augusta Leandro (1819)
and Augustia Klotzsch. (1854) are homonymous names, despite their
different derivation, but proposes the conservation of "Augusta Pohl
(1829), also spelt later as Augustea, though it is a monotypic genus
having a legitimable valid synonym; and without waiting for the
decision of the Congress, he makes the new combination A. longifolia
(Spreng.) Rheder, apparently in the sure belief that this proposal will
be accepted by the Congress.

In contrast, Mr. C. A. Wheatherby (Gray Herbarium) considers
Conocephalus Bl. (1825) and Conocephalum Wiggers (1780) as
homoymous names. He strikes Conocephalus Necker (1790)* out of
consideration, because Necker did not publish a new genus of that
name, but merely changed the spelling of Conocephalum Wigg.,
which change, he maintains, is inadmissible in the Rules. (That
such changes are some times admissible for the purpose of homonyms
may be seen in the example of Bradlea and Braddleya discussed under
the Rules). Further he says that ‘‘Conocephalus Bl. was unanimously
employed up to 1934’’, when Merrill abandoned it to take up
Kotkilospermum Zippel apud Miq (1864). He admits that even
Merrill has not examined the type of the genus Pozkilospermum, the
botanical affinity of which is yet uncertain, and that further changes
will be necessary if Pozkilospermum is segregated generically from
Conocephalus Bl. proper. He acknowledges that ‘‘the least dis-
turbance in nomenclature would have resulted from conserving
Conocephalus Bl. against its earlier homonym [Conocephalum

* According to the view of Dr. A. J. WILMOTT (Kew Bull. 1935
Pp. 92) Conocephalus NECK., being a name of NECKER’s ‘‘Species
Naturalis’’, would be an invalid name, though all NECKER’s
‘“Species Naturales”’ correspond to LINNEAN Genera.

Gardens Bulletin, S.S.

253

Wigg]———. Since, however, the number of species concerned
is not very large (about 35), it is probably better to accept Merrill’s
treatment of the matter.’”’ This might bd paying respect to the person

of Merrill, but not to any principles given in the Rules, for the Rules
have expressly instructed us to take the generic names like Peltophorus
and Peltophorum as different, and not homonymous, names. Besides
changes in the nomenclature of thirty-five species may seem insignifi-
cant to a non-Malayn botanist, but to Malayan botanists who have
been accustomed to see Conocephalus Bl. used in the literature of a
_ hundred years for a group of common forest plants, changes in the
names of thirty-five species are rather a big and vexatious alteration,
even more than to American botanists is the rejection of the name of
the monotypic genus Augusta Pohl (1829), the conservation of which
requires a new combination (now made by Prof. Rheder), because
an earlier specific epithet is available for the species than the
one hitherto used under the genus.

These cases also show how arbitrary is the present
procedure of comserving or rejecting the so called homony-
mous generic names, without having first defined precisely
what a homonym is.

In view of this it is highly desirable that the regulations
on this point should be clarified so as to define, if possible, both
the tautonym and homonym in the same set of general rules;
for, strictly speaking, a tautonym ought to be defined as a specific
name in which the trivial epithet is a homonym of the generic
name. If it is impossible to frame general laws which shall be
applicable equally to tautonyms and to homonyms, because of
some difference in the principles appertaining to them, or because
of the mighty nomenclatorial changes that may result from the
application of any such uniform rules to all homonyms, then it
is necessary to show in the Rules how and why these principles
differ, so that the student of the Rules will be able fully to
appreciate the regulations and, when occasion arises, to decide
for himself whether or not a certain name is a homonym or a
tautonym. If any extensive changes may result from the adop-
tion of such principles in the generic nomenclature, then it will
be advisable to conserve such generic names as exceptions to be
mentioned in the Rules, where they will be handy. General
laws are much better for the guidance of botanists than
obligations to call now and then upon the Executive Committee
to give its arbitrary decisions on the matter.

If attempts are not made to clarify this situation as soon as
possible, then it is to be feared that the opinion, favouring
tautonyms as legitimable valid names, will soon gain ground,
and we may have to witness the present regulations against
tautonyms and homonyms vetoed, or at least considerably
modified, by future congresses.

If general laws, that give due regard to the derivation of the
words, are not possible, then artificial rules such as the zoologists
have, will be better adapted to the needs of systematists so that,
apart from the generic names which, to prevent extensive
changes, may require different rules, specific epithets, even of

Vol. BX: 61037 ):

254

different origin, may be considered as homonymous if they
resemble each other very much, at least in the main part of the
word. Besides now that the Rules do not acknowledge mis-
applications and miscitations as validly-published new binomials
as was done formerly, there is no need to attach any great
importance to the derivation and meaning of the words, provided
serious risks of confusing one name with another are avoided and
a uniformity in the rules is secured. Thus if Gerardiina and
Gerrardina are not homonymous, there is no reason why Bradlea
and Braddleya should be.

For this purpose the following groups of letters might be
considered as equivalent :

(A) Terminations of generic, tribal, or sectional names:
a=t¢=ca—cya; us—yus—ys—1s=— e5=— 0S; Ae
on=oon.

(B) Terminations of epithets of species and of groups
below the rank of species:
os=us (-a, -um)=ius (-ia, -ium)=is (-e)=yus
(-ya, -yum)=ys=es=on=ela = eva = ea=aea;
i—M— pe a
(C) Letters standing in the middle of the epithets denoting
genera, sections, tribes, species, varieties, etc. :

o=a; es=us; v or w (as vowels)=u; g=gh=gu;
c=cc=ck=k@=kk=ch (hard); c (followed by
i, @, ¥, ae, oe, except when ae is the genitive
of the words ending in a, in which case c is
hard) =z=s=ch (soft)=ss; d=dd=dh; p=pp;
t=tit=—th=ct;: ph=pf =f=f/—7=2.- (oneee
and w as consonants); m=mm; n=nn=nh=m
(before b, p, m, or n); r=rr=rh; l=ll=lh; wski
(Polish) =wiski; drze=dreze; ae=oe=ue=oy
= ey=e==1 =j = y= yi= 4 = tae = ene (exces
when ae is the genetive of words ending in ca,
ga, and ka, or is followed by 1).

(D) Letters standing at the beginning or in the first
syllable of the epithets or names of genera, sections,
tribes, species, varieties, etc. :

(x) value of the consonants given in (C);

(2) e=ae=0e; 1=79;

(3) a vowel preceded by h is equivalent to the
vowel without the. preceding h.

For the purposes of generic homonyms, the rules of termina-
tions of the generic names and those for the middle and the
beginning of the words (i.e. A, C and D) should be taken into
account : while for the purpose of tautonyms, and homonyms of
specific names and of names below the rank of species the
regulation for the terminations of the specific epithets and for

Gardens Bulletin, S.S.

255

the valency of letters occurring in the middle and at the begin-
ning of the words should be taken into account. If it is desirable
to regard such epithets as malayana and malayensis; javana,
javensis, javanensis and javanica; philippensis, philippinensis and
thilippina; timoriensis and timorensis; or anamiticum and
annamense, as homonymous, a paragraph could be added making
adjectives ending in (a) nus (-a, -um)=ianus (-a, -um) =i (m) ensts
{-e)=ensis (-e)=(it) icus (-a, -um) as homonymous.

This rule would make Gerrardina and Gerardiina, or
Anadendron and Anodendrum, as pairs of homonyms; but not
Chonemorpha and Conomorpha; but if it is desirable to keep the
first two pairs as distinct, then they could be included as nomina
generica comservanda, or as exceptions to the general rules.

As an extreme case, tautonyms might be defined as a specific
name in which the specific epithet either is homonymous with
the generic name (Radiola Radiola), or is an adjective or genitive
(masculine or feminine) derived from it or its root (Normanbya
Normanbyi, Radiola Radiolaris), or is itself the root of the
generic name (Kandelia Candel), or still is the generic name with
another word or particle added, with or without the change of the
terminal ending of the generic name (Nasturtium Nasturtium-
aquaticum; Pinanga Pinangifolia; Pinanga pinangioides:
Araucaria araucana).

However it would be undesirable to put a ban on all suck
names. I think tautonym-phobes would be satisfied if only
absolute tautonyms were invalidated, by defining absolute
tautonyms as specific names where the trivial epithet, or its root,
is homonymous with the generic name, or its root. This would
outlaw all such names as Normanbya Normanbyi, Carum Carui
L., Chaerefolium cerefolium L., and Radiola Radiola L., but not
Volvaria volvacea, Kandelia Candel, Nasturtium Nasturtium-
aquaticum, Radiola Radiolaris, Cuminum Cyminum L., and
Brownea Brunonis.

It may be observed here that zoologists have already removed
the ban on tautonyms, even on absolute ones, and that the
admission of absolute tautonyms as legitimable valid names would
simplify much the procedure prescribed for new combinations.

15. Linnean Genera (Arts. 20 and 42)

Art. 20 has the following note on the genera in Linnzus’s
Species Plantarum:

“Tt is agreed to associate generic names which appear in Linnzus’s
Species Plantarum ed. 1 (1753) and ed. 2 (1762-63) with the first
subsequent descriptions given under those names in Linnzus’s Genera
Plantarum, ed. 5 (1754) “and ed: 6 (1764)’’.

From this one might conclude that, though the two editions
of the Genera Planiarum are respectively later than the two first
editions of the Species Plantarum, the ed. 5th and ed. 6th of the
Genera Plantarum, are to be regarded for the purposes of Arts.

Vol. IX. (1937).

256

41 and 42, as having been published simultaneously with the two
editions of the Species Plantarum, and that the genera described
in the 5th edition of the Genera Plantarum are to be given
precedence over those in the 6th edition. Since, except in the
case of descriptio generico-specifica (Art. 43), a genus is not
validated by a description or mention of the included species
(Art. 41), this would also mean that a generic name like Cycas
used in the two editions of the Species Plantarum, but not
described in either of the two editions of the Genera Plantarum,
will have to be regarded as invalid.

But from the note made in Art. 42 this does not appear to be
the intention of the legislators. The note runs as follows:

‘““An exception is made for the generic names published by
Linnzus in Species Plantarum ed. 1 (1753) and ed. 2 (1762-63), which
are treated as having been validly published on these dates.’’

This would mean that Linnzeus’s genera in Species Plant-
arum ed. 1 and 2, have to be interpreted with regard to the
species described in those two books, in which case the generic
descriptions in Genera Plantarum ed. 5 and 6 could be taken as
giving only clues to the correct interpretation of the genera.
Should there be a divergence between the descriptions given in
the Genera Plantarum and those indicated by the species or by
the lectotype species adopted under the Rules, then the descrip-
tions of the species should alone be taken as decisive, no changes
in the conception of the Linnean genera or in the lectotype
species being allowable on the basis of the Linnean generic
descriptions.

16. Orthography of Names (Rec. XLIII)

This note is provoked by the following remarks made by
Dr. A. B. Rendle in the review of the late Dr. A. S. Hitchcock’s
Manual of the Grasses of the United States: ‘‘And why does
Dr. Hitchcock who is a leading supporter in America of the
International Rules of the Botanical Nomenclature, desecrate
personal trivial names with a small letter?’’ (Journ. Bot.
BX XT) 1636, ps 26):

In the first place the regulation regarding the use of a capital
letter for personal trivial names is embodied not in a Rule but
in a Recommendation, so that the use is not an obligation. The
recommendation reads as follows :—

‘Specific (or other) epithets should be written with a small initial
letter, except those which are derived from names of persons
(substantives or adjectives) or are taken from generic names
(substantives or adjectives). Examples: Fucus indica,——————
Lythrum Hyssopifolia, Brassica Napus, #3

At first the recommendation seems very simple; yet on a
closer examination it will be found that the recommendation,
besides being very illogical, is also extremely difficult to follow.

Logically one cannot write Napus in Brassica Napus L,.,
Hyssopifolia in Lythrum Hyssopifolia L., and Radiola in Linum

Gardens Bulletin, S.S.

— a a anh st

257

Radiola I,. (the last is mentioned in the example under Art. 69
of the Rules) with initial capital letter because all the species
concerned are Linnzeus’s, who did not use any of these names
for a genus. They were raised to the generic status long after
Linnzeus: Napus in 1829, Hyssopifolia in 1852, and Radiola in
1788. They may have been used as generic names before
Linnzeus, but under the Rules such names are invalid. If made
valid especially for the purpose of following this recommendation,
then practically all vernacular names, deprived of their qualifying
adjectives, would also become valid for the purpose, because
they are also generic names, though not in the Linnean sense.
In fact, Linnzeus wrote with the capital letter all vernacular
names used as trivial epithets, but in the Rules such names are
printed with a small letter, e.g. Gluta Renghas L. in Art. 70.*

But even when cases like those mentioned above are
excluded, the Recommendation ‘is still difficult to follow. The
specific name say Victoria or Victoriana has, according to this
recommendation, to be spelt with a capital letter, if it refers to
the Queen or a lady named Victoria, and with a small letter if it
refers to a place. This means that a botanist who wishes to
follow this recommendation faithfully, will have to waste a good
deal of his time in looking up the original references, and even
then he might find the matter not easy to decide upon, because
in the original reference the derivation of the name is not clearly
stated. A rather lengthy list might be made of such names as
are applicable at the same time equally to persons and to places.

Then a doubt arises whether such names as Princei, Con-
naughtii, etc. should be spelt with a capital when they are taken
not from personal names but from titles e.g. Prince Albert, Duke
of Connaught, etc. Further why should the supporters of the
recommendation not begin with a capital letter such names as
van-Steenisii, van-Houtteana, von-Muelleri, etc.; and again why
include a capital in the middle of such names as MacArthurii,
MacDonaldiana, etc.? Under the recommendation Van-steenisii,
Van-houtteana, Von-muelleri, Macarthurii, Macdonaldiana, etc.
(with or without the middle dashes) should be the correct
latinized epithets.

All this shows that the recommendation is contrary to the
spirit of the Rules which contains the following remarks in
Art. 2:

“‘The recommendations deal with subsidiary points, their object

_ being to bring about greater’ uniformity and clearness in future

nomenclature : names or forms contrary to a recommendation cannot
on that account be rejected, but they are not examples to be followed’’.
(italics are mine).

Hence this recommendation, which proposes to introduce
into the Rules an element of illogical diversity, and of arbitrari-
* New amendments recommend the use of capitals for epithets

derived from vernacular names also, but do not consider the faults
pa Rec. XLIITI.

Vole 1X... (1937).

258

ness, at the sacrifice of simplicity and uniformity, should not
have been adopted by the legislators even as a Recommendation.

No wonder, therefore, that such staunch supporters of the
Rules as the late Dr. Hitchcock are found to disregard the
recommendation for the adoption of a capital letter for certain
trivial names and to use a small letter uniformly for all, a proce-
dure that has also the approval of practically all non-botanical
systematists, and that is being logically followed by the majority
of American botanists.

17. Validating a Name by Referring to a Misapplication

There are cases where botanists have looked upon misinter-
pretations of a species as constituting valid but illegitimate
homonyms. ‘This faulty view has apparently had its genesis at
a time when the type-basis concept was not generally accepted,
while the faulty way of referring to a misapplication has been
mainly responsible for making it widely current amount botanists
and for obscuring the actual difference between a misapplied
name and a homonym.

But now that the type-basis concept has been made obligatory
in modern plant-nomenclature, there is no reason whatsoever why
misapplications should be confused with homonyms, even in the
manner of referring to them in literature. Thus to cite Tsuga
Mertensiana (Bong.) Sarg. for T. Mertensiana (Bong.) Carr. sec
or emend Sarg., or T. Mertensiana Carr. for the misapplication
Carriére made in transferring Pinus Mertensiana Bong. to the
genus Tsuga, has been declared irregular by the last Congress
(1935).

Fortunately, however, whenever a misinterpreted specimen
has been found to be a new species, botanists holding the
erroneous view have not tried to validate the new species by
merely referring to the misapplication; but in every case I have
seen they have included at least either a valid description for
their new species or a reference to a validable figure.* Were it
not for the description, or the reference to the figure, the new
binomial would have been a nomen nudum, even if the references
to misinterpretations contained descriptions applicable to new
species; for all misapplied descriptions, however detailed, are
intended by their author to be nothing but complementary or

* After writing the above, I came across Rendle’s ‘‘Gonolobus
jamaicensis nom. nov.”’ (Journ. Bot. LXXIV, Dec. 1936, p. 345)
which is a nomen nudum, as it is based on the misapplication
G. rostratus (Vahl) Roem. et Schult. by Schlechter (Urb. Symb. I,
p. 284), J.e. on G. rostratus (Vahl) Roem. et Schult. sexsu Schlechter
partim, excl. specimen typicum. Rendle himself has not given any
description of the species, nor has he referred to any validable
description or plate, though he has indicated the type. A name
accompanied by an indication of the type but not by a description
or by a validable reference, is not admitted as valid under the Rules,
é.g. several names in Wallich’s Catalogue.

Gardens Bulletin, S.S.

259

supplementary to the existing descriptions of their misapplied
name, and therefore they do not change the type of the misinter-
preted species, nor even do they succeed in changing the status
of the misapplied name so as to make it a new homonym. All
that these misinterpretations do is to mislead some botanists as
to the exact identity, and the range of distribution, of the species.

Since a misinterpretation creates neither a new name, nor a
new type, that can be called its own, it cannot be called a
homonym, which has both a name and a type of its own. And
since, according to the type-basis concept, no one is obliged to
consider the specimens wrongly cited under a species, even when
these specimens have formed the basis of a detailed misinterpreted
description, it follows that one is not bound to look up references
to misinterpretations or to miscited specimens, unless the mis-
determined specimens or figures are made the types of an original
description of a new species. In the latter case it is not the
reference to a misinterpretation, but the fresh description given
of the specimen or the figure under the new specific name, that
transforms the specimen or the figure into the type, irrespective
of any amount of references to its previous misinterpretations or
miscitations. Similarly in the case of a validable figure, it is the
reference directly to it, rather than to any of its previous
misinterpretations, that validates the description of a new species
and makes the figure the type of the species.

Thus Calophyllum Jacquinit Fawcett et Rendle (Fl. Jam. V,
1926, p. 200) is validated not by references given under it to the
various misinterpretations of C. Calaba L. sensu Jacq. (Sel. Stirp.
Amer. 1763, p. 269), but by the several plates quoted under it and
by the original description given by the authors. It is right in this
case to consider C. Jacquinii not as nomen novum (as regarded by the
authors) but as a species nova, and therefore to look upon all the
specimens cited by the authors as its syntypes. Had C. Calaba in
Jacquin’s work (Sel. Stirp. Amer. 1763, p. 269) been a later homonym
of C. Calaba L. (1753), then C. /acquinii Fawc. et Rendl. would
have been a new name so that its syntypes would have consisted of
the syntypes of Jacquin’s species only, and not any other plate or
specimens cited by Fawcett and Rendle.

However Clerodendron nutans Wall. ex Don, Prodr. Fl. Nepal
(1825) 103 and C. nutans Wall. afud Hook. in Bot. Mag. (1831) t.
3049 are to be regarded not as two different misinterpretations, but as
two different homonyms, of C. mutans Wall. ex Jack in Malay. Misc.
I, part 1 (1820) 17, for they were all published independently of each
other. It is true that in each case the author tried to interpret
Wallich’s manuscript species, but manuscript species and manuscript
types have no status in modern systematic botany so that a botanist,
who wishes to validate a manuscript species, is free to choose any
type he pleases. Hence the type of C. mutans Jack (1820) [I prefer
this formula to C. nutans Wall. ex Jack, which is recommended
under the Rules] is a Penang specimen that was not seen by Wallich,
that of C. mutans Don (1825) is a specimen from Wallich’s collection
from Assam studied by Don, and of C. nutans Wall. (1831) is
Wallich’s specimen described in Bot. Mag. t. 3049 which might be
one of the specimens listed in Wallich’s Catalogue no. 1793 [The
specimen figured in the plate is different and was not seen by Wallich].
It might be noted here that C. Jackianum Wall., Cat. (1828) n. 1704

Wot. [X. €1937).

260

is an absolute synonym of C. nutans Jack (1820), although Wallich
intended to apply that name to a species represented by a specimen
he had collected in 1822 in Penang and distributed under n. 1794.
This specimen cannot be included as a syntype of C. Jackianum Wall.
because the binomial was validated by citing C. mutans Jack. as its
basinym.

If, on the other hand, this way of reasoning is not accepted
and it is allowed to validate a new species by referring to a case
of a misinterpretation or misapplication without the type speci-
men and the type description of the misinterpreted species, then
it would follow that one could validly publish by citation more
than one new binomial by referring not only to more than one
misinterpretation of a single species, but also to different
elements originally included under one and the same species.

Thus supposing two different elements were included in
Pinus Mertensiana Bong., one of them would be the type
(=lectotype) of the species and the other a case of simultaneous
misapplication of the name. One new binomial, Tsuga Merten-
siana (Bong.) Carr., would be valid by citing its basinym,
P. Mertensiana Bong. quoad specimen lectotypicum, and the
other, say T. Bongardii nom. nov., by quoting its basinym,
P. Mertensiana Bong. excl. specimen lectotypicum. ‘This means
that compilers of such works as Index Kewensis would have to
list every single case of misapplication because it would in effect
be equivalent to a homonym, thus defeating the very purpose
for which the works came into existence. Herbaria all the world
over would also be burdened with the responsibility of bestowing
as much care on the miscited specimens as they bestow on the
type specimens.

It must be noted here also that misinterpretations come
neither under the category of such names as are accompanied
with valid descriptions but are invalid because of some faults in
their names [cf. Invalid species of Class III (1-3)], nor under
the category of the species which, though recognized as new and
accompanied with valid descriptions, are published without any
name of their own (cf. Class IV). Taxonomic groups of the
latter categories (nameless or with invalid names) differ from
misapplications in that they are published as new and are accom-
panied with valid descriptions which are based on their own
types, and, unlike misapplications, are not attributed to earlier
valid names. Had such groups been given each a validable
name, then the name would have been accepted as valid in
botanical nomenclature. Long usage, and also the new Rules,
has made it possible to validate names by referring to the
descriptions of the taxonomic groups of these two classes. ‘Thus
Arenga Griffithii Seeman (Pop. Hist. Palms, 1856, p. 61) has
been validated by a reference to a valid description published
without any name by Griffith in Palms of British India, 1850.
p. 168. (See also Section Io).

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261

18. Citation of Misapplications of Names

The current practice of citing misapplications of names or
cases of misinterpreted species is very unsatisfactory. It makes
no distinction whatsoever between a misapplication and a true
synonym or a homonym. Despite the lively discussion reported
to have occurred at the last botanical Congress (1935) on the
necessity of adopting a clear procedure for citing misapplications,
the point was unfortunately left undecided. Some think that
misapplications are better left out altogether from the synonymy
of a species, and a proposal to that effect was even suggested at
the last Congress. The principal reason for adopting this
attitude by some botanists is, I think, that they find the present
way of citing misapplications very vexatious and misleading,
and also because no good way has so far been suggested for
distinguishing between actual synonyms and misapplications.
And yet references to misapplications are very necessary to show
how the views of one botanist differ from another, especially
in the case of misapplications which, owing to the great influence
of the botanist who first misapplied them, have become widely
current among systematists in general. In view of this I take
this opportunity to make some suggestions on the matter, or
rather to defend the procedure I have been following for some
time.

During the discussion at the last Congress, it was pointed
out that to cite “Pinus inops Ait. sensu Bongard; non Ait.’’
would seem ‘‘sheer nonsense’’, since a species cannot be Aiton’s
and not Aiton’s at the same time. But in my view to cite “‘Pinus
inops Ait. sensu Bongard’’ leaving out the misleading phrase
‘‘non Ait.’’, would make sense, since it would mean that Aiton’s
binomial Pinus inops was taken in a different sense, misapplied
or wrongly interpreted by Bongard. As the Rules have expressly
recommended the use of such particles as ex, in, apud and
secundum in order to show clearly the authorship or the inter-
pretation of certain species, I do not see any reason why the use
oi sensu should be condemned in citations which, without its use.
would cause much trouble and nuisance to botanists. The
following two examples will illustrate my view on the matter :

(1) Hedyotis monocephala Hook. f., Fl. Brit. Ind. III (1880) 63,
non Mig.

(2) Hedyotis connata Merr., Enum. Philipp. Pl. III (1923) 409,
non Hook. f.

At first sight the reader might assume that both these
binomials are later homonyms, and, therefore, they are to be
rejected; but this interpretation is true only in the first case,
the second being an example of a misapplication. To avoid such
a conclusion, or to save the reader the trouble of consulting the
original references in order to find out what the author actually
means by his citations, I adopt the above manner for citing only

Vol. IX. (1937).

262

homonymous names, using sensu to indicate misapplications.
ihus:

‘“‘Hedyotis monocephala Hook. f., Fl. Brit. Ind. III (1880) 63,
non (or nec, haud, not, etc.) Miq.’’

would mean that Miquel and Hooker are separately authors
of two ‘different species, but both bearing the same name. If a
botanist wishes to say that Hooker’s species is a later homonym,
he has only to add the year (1869) after Miq., and any reader
would at once understand what is meant. Similarly if Miquel’s
name had been neglected by botanists in favour of Hooker’s
and one wanted to emphasize the fact that legitimately Hooker’s
binomial had no precedence over Miquel’s, it could be done thus :

‘“‘“H monocephala Miq., Ann. Mus. Bot. Lugd. Bat. IV (1869) 225,
non Hook. f. (1880)’’.

I cite a misapplication thus:

“‘Hedyotis connata Hook. f. sensu Merr., Enum. Philipp. Fl. Pl.
III (1923) 4096’’.

while I would have cited the species as:

“Hf. connata Hook. f., Fl. Brit. Ind. III (1880) 62, non sensu
Merrill (1923)’’.

if I considered Merill’s interpretation as erroneous but
widely accepted and, therefore, wanted to warn the reader against
following it. In practice, however, the last procedure need not
be used unless the references have to be very limited. Where
there is some latitude for giving references, once can always
exclude a misapplication by showing what portion is excluded
from, or included in, it, thus:

‘“‘“Hedyotis connata Hook. f., Fl. Brit. Ind. III (1880) 62; Merr.,
Enum. Philipp. Pl. III (1923) 496 quoad specimen Wallichianum”’ or
‘excl. specimina malesiana’’.

But in the case of misapplications where the type specimens
are to be excluded, the use of the word sensu in the manner
indicated above makes the statement both concise and explicit.

However this is not the only way of citing misapplications
clearly; and other ways may occur to readers. But such
formule as

Hedyotis connata (vix Hook. f.) Merr., Enum. Philipp. Pl. III
(1923) 406.
and

Hedyotis connata Merr., |. c. p. 496, vix Hook. f.

are not recommendable, as they are also confusing like the
previous ones discussed above. (See also Section 19).

19. Nomenclatorial Expressions
The argument of the preceding sections 17 and 18 rests on
the principle that a taxonomic name is a nomenclatorial expres-
sion the meaning of which is not self-evident. Taxonomists
therefore must agree upon the way in which a nomenclatorial
expression is to be expounded, i.e. its denotation and its

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263

connotation, in order to prevent ambiguity. A nomenclatorial
expression should, in fact, have the exact significance of an
algebraic expression. But however obvious this principle may
be and however explicit the Recommendations in the Rules, the
fact remains that the same class of nomenclatorial expression
may carry a different meaning not only in the works of different
botanists but in the works of the same botanist.

The meaning of a nomenclatorial expression is dual: it
denotes a certain kind of plant, and it connotes the class of the
expression and its author, and/or the opinion of one or more
botanists. The denotation has recently been fixed by adoption
of the type-concept. It is the connotation that is uncertain and,
if this is not remedied, there will be great confusion. ‘The
following example will explain the point:

The taxonomic name Podocarpus javanicus Merrill is a
nomenclatorial expression of the binomial class at the first intent.
Therefore it should connote that that species of Podocarpus was
first described or validly instated by Merrill. But such an
expression is also used to connote:

(a) a binomial of the second intent, that is, one that has
been transferred from one genus to another, e.g.
Agathis alba Foxworthy (=Dammara alba Lam.),
Artocarpus champeden Spreng (=Polyphema
champeden Lour.).

(b) a misidentification or misinterpretation of the species
of one author by another, the name of the second
author being attached to the binomial of the first,
e.g. Hedyotis connata Mertr. is a misinterpretation
by Merrill of H. connata Hook. f.

(c) an interpretation of the species of one author by
another whether rightly or wrongly, with or without
proof, but in the absence of exact denotation by a
type specimen, e.g. Bridelia monoica Merr. is
Merrill’s interpretation of Cluytia monoica Lour.;
Calamus usitatus Merr. and C. usitatus Becc, are
respectively Merrill’s and Beccari’s interpretations
of C. usitatus Blanco.

All these three uses are common. Uses b and c mean that
one is permitted to make any number of homonyms on a basinym
according to the number of botanists who have made wrong
identifications, or different interpretations, of the original species;
yet all these homonyms are invalid.

Actually Podocarpus javanicus Merrill connotes a mis-
identification of Podocarpus javanicus (Burm. f.) Merr., which
is a binomial of the second intent based on Thuya javanica
Burm. f. and identified by Merrill with Podocarpus imbricatus Bl.

Vol. IX. (1937).

264

I consider that the more usual nomenclatorial expressions
should have the meaning I give below. These expressions may
be subdivided into two main classes: (A) Expressions that denote
a taxonomic group and connote the class of the taxonomic name,
and the author who first described it; and (B) Expressions that,
in addition to the previous meaning, connote either an inter-
pretation given to the name, subsequent to the original
description, and the author of this interpretation; or the specimens
that have to be retained or excluded under the later interpreta-
tions. Under (C) I have included both those expressions which
connote the person through whose offices an author secured the
validation of his expression, and his description of it, and those
expressions which I consider misleading.

The expressions of the class (A), though they need not
indicate the type, must be interpreted according to the type,
even when there are grave errors in the description given and
in the specimens cited. If no type has been designated in the
original description, or if it is not extant, then the species must
be typified and the errors excluded (see Section 13). Ex-
pressions of the class (B), referring to errors in description or
citation, cannot constitute a valid description or a valid name,
because a name or a description with its type excluded is
inadmissible in botanical nomenclature. In fact the use of
nomenclatorial expressions to indicate errors and exclude types
seems to be an anomaly, but since they serve a useful purpose
in showing the portions excluded from a mixtum compositum
and those retained as the lectotype, I think no harm is done by
retaining their conventional use in botanical nomenclature,
provided any chance of their being confused with other ex-
pressions is elimated. However it must be understood that,
since such names indicate errors in description or citation, they
cannot be used for validating a new name or a new nomenclatorial
expression (see Section 17).

Class A:

1. Juniperus silvestris Roe denotes a certain species of plant
and connotes a new taxonomic binomial described by Roe. Here
the type is the specimen or the plant studied by Roe.

2. Thuya silvestris (Roe) Doe denotes the same species, and
connotes two ideas, namely, a taxonomic name originally
published by Roe, but later transferred to another genus by
Doe. ‘The type here is the specimen or plant studied by Roe.

3. Thuya silvestris Doe denotes a certain species described
by Doe and connotes that the new name was given to the species
by Doe. Here the type is the specimen or plant studied by Doe.

Class B:

1. If J. silvestris Roe was a mixtum compositum (positive),
the resolution of the species into various components, or the

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265

subsequent correction of it by other authors, may be indicated
thus :

(a) J. silvestris Roe quoad specimen lectotypicum = J.
silvestris Roe emend. Doe = J. silvestris Roe
sensu restricto = J. silvestris Roe.

(b) J. silvestris Roe excl. specimen lectotypicum is merely
a conventional expression to indicate the element
that had introduced errors in the original descrip-
tion and citation of the species. It has no valid
taxonomic or nomenclatorial status.

2. If J. silvestris Roe was not a mixtum compositum, but
had been subsequently misinterpreted by Doe, then

J. silvestris Roe sec. or sensu Doe = J. silvestris Roe
quoad interpretationem Doeanam et excl. typum.

A description given by Doe, cannot be considered as valid
in taxonomic botany because, according to the type-basis concept,
a description must be interpreted by its types, and the very fact
that Doe gave his description under J. silvestris Roe means that
he tacitly admitted Roe’s type as the type of his description.
The above expression therefore indicates the errors that were
introduced by Doe, and so a new name cannot be validated
merely by referring to J. silvestris Roe sensu Doe. (See also
Section 17).

3. If J. silvestris Roe emend. Doe was the basinym of
Thuya silvestris (Roe) Doe, then this fact may be expressed
thus :

Thuya silvestris (Roe emend. Doe) Doe.

4. If this emendation was done by Doe after he had made
the combination and not previously or simultaneously, then the
expression to indicate this fact is

T. silvestris (Roe) Doe emend. Doe (=T. silvestris (Roe)
Doe quoad specimen lectotypicum.).

5. If Doe, in transferring J. silvestris Roe to Thuya, had
misinterpreted it, then according to the Rules, T. silvestris (Roe)
Doe must be interpreted according to the type of its basinym
J. silvestris Roe. The elements of this misinterpretation can be
resolved thus:

(a) T. silvestris (Roe) Doe sensu typico, originali or
basynimico = T. silvestris (Roe) Doe quoad basinym
= T. silvestris (Roe) Doe = J. silvestris Roe.

(b) T. silvestris (Roe) Doe excl. basinym = T. silvestris
(Roe) Doe pro parte non-typica, non J. silvestris
Roe.

The latter has no valid status in taxonomic botany.

Vol. IX. (1937).

266

6. If Juniperus silvestris Roe is a polymorphous species,
then

(a) J. silvestris Roe sensu stricto = the type form.

(b) J. silvestris Roe sensu lato = both the type and other
forms. |

7. If J. silvestris Roe was a negative mixtum compositum,
and if one were to follow Doe’s interpretation of it, without being
sure whether or not Doe had made any good case for his
interpretation, then this idea could be expressed thus:

J. silvestris Roe teste/fide Doe.

Class C:

Other expressions used by authors, or recommended or
prescribed in the Rules.

1. ILEX (Tournefort) Linn. is a misleading expression for
ILEX Linn., though the former is according to the Recommenda-
tion XXXII. Here the genus need not be interpreted with
reference to Tournefort’s genus at all, the latter being pre-
Linnean and invalid. ILEX Linn. must be interpreted after
the plant studied by Linnaeus and not Tournefort.

2. Scindapsus argyreus Hort. ex Engler is a misleading
expression for S. argyreus Engler, as the application of the name
by Engler may prove to be quite different from one that was
current among the horticulturists of the time. The former
expression is prescribed in the Rules (Art. 48), though the latter
is also permitted.

3. Clerodendron nutans Wall. ex Jack is a misleading
expression for C. nutans Jack. It is true that Jack adopted the
name given by Wallich to plants in herbarium or in the garden,
but the plant to which Jack reserved the name might be different
from the one to which Wallich had given it. Here the type is
the specimen studied by Jack and not the one studied by Wallich.
The first expression is prescribed in Art. 48 of the Rules, though
the second one is allowed whenever an abbreviated form is
desired.

4. Anodendron manubriatum (Wall.) Merrill is a misleading
expression for A. manubriatum [Wall. ms. nomen nudum]
Merrill = A. manubriatum Merrill. The first two expressions
seem to be the natural extension of the procedure of the Recomm.
XXXII, though the third is the correct expression for the plant.
The type is the plant studied by Merrill and not one studied by
Wallich.

5. Clerodendron nutans Wall. in Don means that the name
and the description of Wallich was published in a work or a
paper edited by Don. If the author of the nomenclatorial ex-
pression is the same as the editor, then omit both the word in
and the editor’s name. But this in cannot be omitted if the work

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267

edited by Don is a periodical, though often in such cases the
editor’s name may be omitted. (vide infra).

6. Clerodendron nutans Wall. apud Don means that Wallich
is the author of the name and the description published in a
monograph, or a revision, prepared by Don. Don may have
concurred with Wallich’s opinion and therefore allowed Wallich
the credit of his work; but still Wallich alone must be considered
as the author of the species. If Don had also added some
citations to the ones given by Wallich, they must be excluded
from the consideration in typifying the species.

N.B.—The use of these two words in and apud is not very
clearly stated in Art. 48 of the Rules. Thus in the case of
Viburnum ternatum Rehder in Sargent, Trees and Shrubs II
(1907) 37, Sargent is considered as a part of the title of the
book, whereas from the treatment it appears to be the name of
the author, or of the general editor, of the book, Rehder being
a contributor to the work. ‘The following three examples will
illustrate the distinction I have made above: (a) Iguanura
brevipes Hook. f. apud Beccari et Hook. f., Palmeze in Hook. f.,
Flor. Brit. Ind. VI (1892) 416, or merely I. brevipes Hook. f.,
Flor. Brit. Ind. VI (1892) 416. (b) Iguanura parvula Becc. apud
Becc. et Hook. f., Palmeze in Hook. f., Flor. Brit. Ind. VI
(1892) 417 or I. parvula Becc. in Hook. f., Flor. Brit. Ind. VI
(1892) 417, (c) Globba Gibbsie Ridl. in Gibbs, Contrib. Flor.
Mount Kinabalu in Journ. Linn. Soc., Bot. XLII (1914) 162,
or G. Gibbsie Ridl. in Journ. Linn. Soc., Bot. XLII (1914) 162.

20. Nomina Specifica Conservanda

The enforcement of the priority principle and the type-basis
concept is causing radical changes in the popular nomenclature
of some plants. But this is inevitable since certain adjustments
have to be effected so that, in order to conform with the Rules,
all misapplications and all illegitimate names may be rejected
without any regard to the influence and position of the botanists
who put these misinterpretations and illegitimate names in
circulation. In complying with these provisions we support the
laws that have been made with the express purpose of bringing
about ultimate simplicity, uniformity and definiteness in plant
nomenclature.

But time is needed in order to discover all misapplications
and illegitimate names; and to do this work satisfactorily a
thorough research in the types and also a monographic revision
of the plants concerned, not in regard to one small region, but
of the entire area where they and their allies occur have to be
undertaken so that one and the same species may not receive
different names in different regions or political countries. It is
natural, therefore, that a work of such vastness is not to be done
haphazardly, nor by one or a few botanists. Normally the more

Vol: 1X. (1937).

268

botanists work and try to specialize in families and genera in
co-operation with field workers and plant collectors, the shorter
will be the time to reach the goal envisaged by the enforcement
of the two principles mentioned above.

Thete are several cases yet to be investigated of the nature
of Raphidophora silvestris (Bl) Engl., R. Wrayi Hook. f., and
R. nigrescens Ridl. mentioned above (p. 234), where the species,
which in the three different regions passed as three different
species, are actually only one. In this case the botanists and
others concerned in two of the countries will have to adapt
themselves to the changes in the nomenclature of the species,
despite the popularity of the rejected names. ‘Then there are
cases where opinions are not agreed over the identity of species
or over the limits of taxonomic groups. In such cases it is but
natural that the legitimate name of one botanist should become
illegitimate for another. "To prevent such oscillations in names
is impossible unless legislation is passed fixing the limits of the
taxonomic groups concerned and preventing any difference of
opinions on the matter. Such a legislation is not within the
province of the nomenclature laws which, being concerned only
with the names, leave the delimitation of taxonomic groups to
the judgement of the individual botanist; and, if actually passed,
it would be tantamount to forcing the taxonomic opinions of
one set of botanists on another and to hindering the progress
of the taxonomic science.

And yet in a Memorandum submitted at the last Congress
supporting a proposal for nomina specifica conservanda we read
many unjust criticisms against unavoidable changes and against
systematists in general, as if systematists effect these changes out
of sheer pleasure. ‘The criticisms also reveal a misapprehension
as to the actual state of affairs.

One critic for instance, writes: ‘‘Foresters greatly resent the
changing of names of well-known forest trees without any
reference to the persons mostly concerned’’ as if the correct
application of a name depended upon the views of some particular
persons using the names. Another says: ‘‘Practically all the
evidence of priority must have been ferretted out by botanists
by now, and if some obscure references do turn up in future,
there should be some limitation’. This may be true in some
few cases only, especially in case of the native flora of some parts
of Europe where systematic studies have been carried on very
intensively for a number of generations by botanists who had
opportunities to study not only the plants in the field but also
the types in the herbarium; but this is not so in the great majority
of cases. Besides, the priority principle and type-basis concept
have been very recently adopted by botanists and so an intensive
study is necessary before all misapplications and synonyms can

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269

be detected. No doubt legitimate nomenclature changes will
cause some troubles at present, but in the end the procedure,
when worked out in full, is expected to result in stable
nomenclature except in species where the status might remain
still a matter of taxonomic opinion.

But some people are impatient, and it is this impatience,
coupled with inadequate understanding of the results expected
from the adoption of the priority principle and type-basis concept,
and with the lack of precision in some articles of the Rules, that
is, in my opinion, the pricipal reason why some people clamour
for nomina specifica conservanda. In fact we read in the
Memorandum referred to above that the Inspector-General of
Forests to the Government of India has stated the following :
“Tf these motions are not passed I shall have to consider the
appointment of a committee to deal with nomina specifica
conservanda as far as Indian forestry is concerned, irrespective
of what names European botanists may give to our trees’’.

It is to be feared that, if such a committee ever gets to work
in India without any reference to European herbaria where most
of the types of Indian plants are preserved, many misapplications
will be perpetuated; and, if this is done without any regard to
the nomenclature of the plants that also occur both in India and
in countries beyond the control of the Inspector-General of the
Forests in India, chaos and confusion of vast magnitude will be
the result. Besides what guarantee can the Inspector give that a
conserved misapplication will not be again misapplied, necessitat-
ing another committee decision to satisfy the demands of the
future users of the second misapplication ?

Judging from the way the name Rhizophora conjugata L.
(Sp. Pl., 1753, p. 443) was being used only a few years ago both
in Asia and Europe, it is almost certain that, if votes were taken
then, the name would have been conserved for a plant which
Linnzeus did not intend, and which should actually be called
R. apiculata Bl. (1827) with R. candelaria DC. (1828) as a
synonym. Yet Trimen who was well acquainted in Ceylon with
Rhizophoraceous plants in the field and had also examined the
type drawing in Hermann’s herbarium (apparently the sole basis
of the Linnean species), showed that R. conjugata I. was con-
specific with Bruguiera gymnorrhiza (L.) Lam. (1797) (Journ.
Linn. Sec. Bot. XXIV, 1887, p. 142). Rhizophora conjugata L.
as until recently understood and as even now used in some recent
Asiatic works, is therefore nothing but a misapplication to be
rejected under the Rules in order that its correct name R.
apiculata Bl. might be used, and in order that R. conjugata L.
might be retained for the plant for which Linnzeus had published
the name. If now because of some trouble that may be caused
to Indian foresters, the Inspector-General’s committee retains the
Linnean name for R. apiculata Bl., then there will be three

Val TX... (4937).

_

270

names, R. conjugata L. sensu the Inspector-General’s committee,
R. conjugata L,., and R. apiculata Bl. current for two plants.

It is almost certain that the nomenclature of the Indian
forestry department will not be followed by systematists and other
workers not only in Europe and America, but also in Malaya,
Indo-china, Netherlands Indies, Philippines, Formosa, China,
etc., and perhaps even by systematists in India itself. In the
end it might be found that no one, outside the Indian forestry
department, would be able to follow the plant nomenclature
adopted by the Indian foresters, unless a detailed synonymy were
given; and conversely the Indian foresters might not be able to
keep in touch with the work that is being done by persons not
connected with the Indian forest department but in subjects
concerning Indian forest plants. Thus the work of the Inspector-
General’s committee might hinder rather than aid the progress
in the knowledge of the Indian plants themselves.

In my view, therefore, the time is not vet ripe for proposing
a list of nomina specifica conservanda. Some time is necessary
so that the nomenclature that was in use may be adjusted to the
principles of priority and the type-basis concept adopted by the
Rules. Detailed investigations are, moreover, necessary in order
that misapplications may be detected. Further the methods
proposed for interpreting a mixtum compositum or for rejecting
nomina ambigua and confusa have also to be worked out. It is
only when these methods and investigations have failed to make
the nomenclature stable that the nomina specifica conservanda
could be undertaken, if actually desired. Otherwise an adoption
of a nomina specifica conservanda would mean the perpetuation
of errors circulated through careless work and the discouraging
of thorough investigations 1n systematic groups; it would mean
that any influential groups of botanists could circulate any
amount of misinterpretations and, should anyone less influential
but more careful contest the interpretations on the basis of the
types, the former could resort to votes in order to perpetuate
their misapplications by means of legislation. In the end we
may get the Congress degraded into a sort of political parliament
where parties would meet, each with the intent of defending,
right or wrong, an interpretation proposed by its members.

Instead of trying to influence the botanical nomenclature of
Indian plants of forestry importance, it would be better if the
Inspector-General of Forests were to use his weight in order to
develop the vernacular nomenclature of the plants to be used
both by the members of the different forest departments under
his charge and by the traders interested in the Indian forest
products. Such an attempt might in the long run bring about
more clearly certain differences among the species that are now
unperfectly known, and thereby might help botanists to clear the
status of the species concerned.

Gardens Bulletin, S.S.

271

21. Effectiveness of Advance Separates

Article 36 of the Rules of Botanical Nomenclature deals with
the conditions regulating the effectiveness of a systematic
publication, and it reads as follows:

‘‘Publication is effected, under these rules, by sale to the general
public or to botanical institutions, of printed matter or indelible
autographs, or by distribution of these to specified representative
botanical institutions.*

‘‘No other kind of publication is accepted as effective : communi-
cation of new names at a public meeting, or the placing of names in
a collection or gardens open to the public, does not constitute
effective publication.’’

But at the last Congress (Amsterdam 1935) there was added
to the foregoing Art. 36 a further clause, which reads as follows:

‘Where separates from periodicals or other works placed on
sale are issued in advance, the date on the separate is accepted as the
date of effective publication.’’ (Proceedings I, 1936, p. 343).

Now this additional clause is so worded that, I suggest,
it makes it possible to secure effectiveness for a publication in a
way which the foregoing text of Art. 36 was meant to prevent.
It weakens and lessens the effectiveness of the text by not making
it obligatory for the authors of separates to distribute these to
any specified persons or institutions, a distribution which, in the
original text, is the alternative to placing the printed matter or
indelible autograph on sale. The effect of the clause is to
encourage the printing of works in parts but the distribution only
in completed volumes; and yet, because of the separates issued
in advance, to legitimize the claim to make the date on the parts
or separates (and not the later date on which the volumes were
distributed) as the date of their effective publication.

Already the existence of this practice of distributing some
institutional publications in volumes, instead of in parts as and
when issued, is causing much confusion and inconvenience even
to the institutions which maintain an exchange with, or subscribe
to, such publications; in as much as the latter institutions are
prevented from keeping in touch with the recent advances and
progress in systematics of the countries in which they are most
interested, being obliged to wait for the information till the
completion of the volumes. There is no fixed period within
which a volume of such a periodical might be completed and be
made available to the public; and it is not uncommon to find
botanists accumulating their separates before finally distributing
them, simply in order to save themselves the expense of frequent
distribution. As a consequence of this lack of immediate distri-
bution of systematic papers several names may be published
which later may have to be discarded as superfluous or
illegitimate.

* Only a tentative list has been compiled of the representative
botanical institutions to be recognized under this Article, but no final
at has yet been reached as to the names to be included in
this list.

Vol. IX. (1937).

272

If now, encouraged by this new clause, this practice otf
distributing departmental publications only in completed volumes
and not in parts as issued, or of distributing in one year periodi-
cals dated in a previous year, becomes general, one can well
imagine the evil effects of the proposed new rule. What amount
of unnecessary inconvenience and confusion would have resulted
if the sale of the FLORE GENERALE DE L’INDOCHINE was begun
only on the completion of each volume, instead of, as hitherto,
on the issue of each fascicule, and yet, because of the separates
issued in advance as authors’ copies, the validity of the publica-
tion was to begin on the date on which each fascicule was
printed? Some volumes of this flora have actually taken twenty
years or more to complete.

Hence if the work is to be effectively published by means of
advance separates, these separates should be regarded as works
not offered for sale, for such they are in fact; and conse-
quently, the clause governing such works should be made to
operate. The mere fact that a work is regarded as an advance
separate from a regular periodical to be placed on _ sale
should not obscure the fact that these separates, at the time
of their issue, were not placed on sale; and so they should
not be exempted from the rule and be allowed to secure effective-
ness in publicity without making the work available to the
botanical public, principally to those institutions which are most
interested in it. Should some accident or economic upheaval
destroy the issue of the periodical or cause it to be suspended
before the intended completion of the volume, then the papers
issued in advance as separates may never become available for
purchase. In such an event there might be all the elements to
pronounce the work ineffectively published, though the new
modification to Art. 36 would make its publication effective.

Moreover if the principle is admitted of legitimizing a
publication at a date when it is issued for private distribution,
what is the use of penalizing others who issue independent works
and‘ circulate them in a similar manner, simply because no
promise has been made to publish them at a later date in a regular
periodical? Again, what if the author of such a private work
were to announce in it that he would issue at a later day an
edition for the public? Which date should systematists then
take for the purpose of reckoning the effectiveness of the
publication? For, to say that a certain, privately circulated
work is an advance separate of a regular periodical is no more
than to promise to publish the work later in the periodical which
will be released at an indefinite date. Hence the new clause to
Art. 36 would be applicable in this case and would make the first
issue of the work effective.

To anticipate the date of the systematic validity of a work
on the expectation of certain events to take place at an indefinite
future is, I submit, an undesirable procedure. If the date of the

Gardens Bulletin, S.S.

273

effectiveness of a publication is to be judged, it should be on the
action already taken by its author or publisher to make the work
available to the systematic public interested in it, and not on the
action that the author or the publisher will take, or has promised
to take, at some future day.

If an institution finds difficulty in printing and distributing
the results of its research work as it progresses, it could publish
these results from time to time in those privately owned periodi-
cals which are only too willing to accept them for publication,
and the institution could order reprints from the publishers for
the purpose of exchange.

From the foregoing it will be seen that the additional clause
as it stands introduces a very disquieting element into the Rules
and makes Art. 36 practically valueless. The legislators might
have been led to adopt the clause owing to its misleading
phraseology, for the clause might be easily mistaken for an
amendment to Art. 45, which deals with the dates of effective
publications. Had the following proposal been made, its
undesirability would have been at once apparent; yet it has the
same significance as the clause adopted :

‘‘Publication can also be effected, under these Rules, by means
of printed matter or indelible autographs not placed on sale, and not
distributed to the specified institutions, if they are regarded, at the
time of the issue, as advance separates from periodicals or other
works placed on sale, it being immaterial whether or not they are
actually reprinted in the periodicals or other works’’.

However the new clause can be made to conform with the
text of Art. 36 by the addition of some such words as the
following :

‘‘provided that immediately following the issue of separates they be
duly distributed to the specified representative botanical institutions.’’

22. Representative Botanical Institutions under Art. 36

The discussion held at the last botanical Congress in
Amsterdam, though short, showed that the tentative list of the
representative botanical institutions prepared by Dr. T. A.
Sprague and Miss M. L. Green for recognition under Art. 36 was
not satisfactory. To be effective the list should conform : firstly,
to the needs of the institutions working in the area about which
the papers to be validated deal; and, secondly, to the needs of
those institutions which, though not situated in the area, are yet
directly interested in the studies of the flora. On the other hand,
certain institutions neither want, nor have any place to
accomodate, systematic papers concerning regions, in the floristic
studies of which they are not even remotely interested. Con-
sequently it would be worse than useless to send such papers to
them, as these institutions might destroy the papers immediately
on their receipt and without keeping any record.

In order to draw up a satisfactory list, it would seem advisable
therefore, to divide the entire globe into about a dozen regions,

Vol. 1X.\(6.1937):

274

which need not necessarily be botanical regions, each having
the following subdivisions: (a) including the important botanical
institutions working in the divisions with which the work deals;
(b) including the important institutions of the neighbouring
countries which may be directly or indirectly interested in the
flora of the ‘region; and (c) including the institutions which,
though situated in countries far away from the regions dealt by
the work, are interested in the floristic study of the region, as
well as the institutions of those countries which have colonial
dependencies in the region.

The total number of institutions thus enumerated should
not exceed 20-25, and in no country should more than one
institution be named, unless the extra institutions in any single
country are sufficiently important to be included under (a).

With such a list determined upon, it should be made
obligatory for an author, desiring to make his work effective for
the purpose of systematics, to send copies of his work to the
relative institutions under a, b and c.

In the case of an author’s work embracing an area falling in
more than one regional group, he should send at least to one of
the most important institutions in each country named under (a)
and (b) in the groups affected. This latter procedure would
meet the case of the author who might be uncertain into which
regional group his work should exactly fall; for example a work
dealing with the whole of France or with Himalayan regions,
or with Tropical America, or Indo-China. This suggested
procedure would oblige authors who wish to describe, in one
single paper, new species and genera from all parts of the world,
to send their papers to an enormous number of institutions. But
botanists who would think of preparing such a work for private
distribution only should be thus penalized in the interest of
science.

It would be further desirable to make this distribution to
the specified institutions through a central committee, which
should be responsible so to mark every copy thus distributed that
botanists consulting any one of the copies will know at once
that the work, though not placed on sale, has been effectively
published. Such an obligation would render difficult to pass
undetected any future offence against Art. 36. Besides without
such a committee to keep a check on this distribution, how is one
to know, without making a detailed inquiry, that certain
privately distributed work sent to one institution is not sent to
others specified under the Rules? In view of the difficulties to
decide immediately on this matter, a botanist might be induced
to accept an ineffective publication as valid and to make
accordingly several nomenclature changes, resulting in invalid
names passing not only as valid but even as legitimate. If
several years elapse before the mistake is discovered, then the

Gardens Bulletin, S.S.

275

rejection of the invalid names, though circulated erroneously as
legitimate, might be resented because of their being long in use.
There is also the possibility of some institutions losing the copies
sent to them, and, if the institutions are unable to produce
records showing whether they had actually received the copies
or not, the loss might vitiate any subsequent inquiry into the
effectiveness of the work. All such drawbacks would disappear
if the distribution of the publications not placed on sale has
perforce to be done through a central committee.

A fourth clause (d) could be added by way of recommenda-
tion including those large institutions or scientific libraries
which, though not actively interested in the flora of the region,
are willing to accomodate works for the purpose of making them
available to research workers, provided these institutions are not
already included in a, b and c.

To illustrate all that I have said above, I append the
following tentative list (omitting the considerations of
palzobotanists who should have their own separate list) which
might form a basis for future discussions :—

im Vol. IX. (1937).

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Gardens Bulletin, S.S.

277

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Vol. IX. (1937).

280

23. Summary

In this paper I have attempted to make clear and precise some
tules that were obscure, misleading, or defective.

The greatest fault in the Rules is the equivocal use of the
words valid and legitimate and their respective derivatives. As
defined here, invalid taxonomic names are those that have no
status in modern botanical nomenclature, all those that have
such a status being defined as valid.

Invalid names and nameless taxonomic groups can be
arranged into four main classes :—

(1) those that are ineffectively published;

(2) those that are published in effective but invalid
publications;

(3) those that are published in valid publications, but
either containing defects in (a) the names them-
selves, and/or (b) their descriptions; or (c) con-
sisting of only names without any descriptions; and

(4) those that are published in valid publications and with
valid descriptions, but without any names
whatsoever.

The present usage allows the validation of a taxonomic name
by a reference to a valid description of invalid names in Class 3
and of nameless taxonomic groups in Class 4; also permits
references also to plates in publications that are rendered invalid
under Art. 20; but the wording of Arts. 37, 42 and 44 of the
Rules seems to allow references to descriptions and plates
published in any books, even popular and non-botanical books
and written in any language, provided the conditions for effec-
tiveness as laid down in Art. 36 are complied with. The Rules
have, therefore, to be amended so as to exclude references to
such non-botanical publications and invalid descriptions. It
would indeed be advantageous to admit in future as valid only
references to valid descriptions, and plates, in valid publications,
defining valid publications to mean such publications as are
effectively published and as ordinarily reach systematic botanists.

Provisional names are invalid because of the defects that make
their descriptions invalid. Since one may not validate a name of
a taxonomic group by referring to an invalid description, it
follows that references to provisional names or to their
descriptions are not valid.

Valid names may be legitimate or illegitimate. An author,
having defined his taxonomic group and assembled together all
the rival names of the group, should consult the Rules in finding
out the correct name for his group. Under the Rules it should
be possible for only one name to be correct for a given taxonomic

Gardens Bulletin, S.S.

281

group (i.e. a group in a given sense, in a given position and in a
given rank): this name is the legitimate name for the group
ri defined by the author, all others being regarded as illegitimate
or it.

Illegitimate names are cited as synonyms, which may be
divided into two classes (1) absolute synonyms, consisting of
typonyms, isonyms, and/or basinyms, and (2) equivalent
synonyms. A third class of synonyms consists of names doubt-
fully referred to the species or of names based on monstrosities
of the species.

As the present wording of the Rules is obscure and equivocal
regarding the procedure to be followed in selecting the legitimate
name out of the several rivals of unequal ages, three usages are
aisles only one of which will have to be adopted in the

ules.

The difference between a new combination and a new name
is explained.

Alternative names are simultaneous isonyms; but those
simultaneous isonyms that are not new combinations of equal
rank should be prohibited.

The Rules oblige one to follow the botanist who first
preferred one of the rival names of equal age and equal rank for
one species (Art. 56), and one of the homonymous names of
equal age (new addition to Art. 61); but it is necessary to add
to both these Articles that this preference holds effective so
long as the names remain in the same position or genus under
which the choice was made. On the transference of any of the
names to another position or genus, the author who first made
the distinction in the new position should be followed. But
in every case the botanist exercising his privilege under Arts.
56 and 61 should see that a new combination gets precedence
over a new name, and this over a new species; and also that,
among the homonymous names that are isonyms of equal age
and equal kind, the one that has the oldest basinym gets the
precedence.

Varietal names given to the type form of a species (as in
Rec. XVIII) and retained as varieties of it are equivalent to
ternary specific names and cannot have a place in modern plant
nomenclature. Examples are given to show that such a
procedure may also lead to serious confusion.

When a species is to be divided into two or more species
and it is not possible to assign the original description definitely
to any of its syntypes, then the first author to choose one of them
as the lectotype or to transfer the other specimens to different
-species should be followed. If the syntypes of a species are not
extant and the original botanical description combined with

Vol: TX. 4937 J:

282

phytogeographical considerations admit more than one interpreta-
tion that is not obviously discordant with the original description,
then botanists should be obliged to follow the earliest of the
interpretations; or if the species has been previously excluded as
doubtful, then the name should not be adopted to any taxonomic
group, without any adequate botanical reasons or proofs. In
my opinion the time is not yet ripe for making a list of nomina
ambigua, dubia vel confusa: suggestions for stabilizing names
without such lists should first be given an opportunity to do their
work in full.

The Rules regarding tautonyms and homonyms appear to be
very confusing. It might be better to frame some general rules
without attaching any importance to the meaning and origin
of the words, provided serious risks of confusing one name with
another are avoided. Suggestions are made showing how this
rule could be framed.

Since according to Art. 42 generic names in Linnzus’s
Species Plantarum, eds. 1 and 2, are validly published, the
purport of the provision under Art. 20 seems to make the generic
descriptions in Linnzus’s Genera Plantarum, eds. 5 and 6,
merely supplementary.

It seems logical to write all trivial epithets with a smail
initial letter, whether or not they are the names of persons or
of genera, or their derivatives.

Misapplications cannot be used to validate new species. It
is therefore important to adopt a method of citation which will
clearly distinguish misapplications from illegitimate homonyms.
It is argued that the use of the word sensu for this purpose is a
rational one.

A brief discussion on the different kinds of nomenclatorial
expressions is introduced in order to further clarify the funda-
mentals underlying them. Though the use of nomenclatorial
expressions to indicate errors seems to be an anomaly, yet their
conventional use is retained, as they help to refer to errors both
concisely and precisely. But they should be used in the way
suggested, so that they might not be confused with expressions
that do not refer to errors. It is shown again that expressions
referring to errors cannot be used as a basinym of a new name.
Expressions with in, ex, and apud are also discussed.

The principle of priority and the type-basis concept have
been adopted as universal rules only very recently. Time is
therefore needed so that plant nomenclature may adjust itself
to these principles. It would be detrimental to the interests of
systematic botany if nomina specifica conservanda were adopted
at this stage.

The additional clause voted to Art. 36 by the last Congress
is against the spirit of the article itself, as it will legitimize

Gardens Bulletin, S.S.

283

practices of securing effectiveness in publication which it is the
principal object of Art. 36 to prevent.

A principle is here enunciated by which unanimity and
satisfaction might be secured in recognizing representative
botanical institutions under Art. 36. Reasons have been
adduced to show the need of distributing the copies of a publica-
tion not placed on sale through a central committee. A
tentative list of these institutions is also given.

Vol. IX. (1937).

LIST

OF BINOMIALS DISCUSSED, AND OF NEW
USED,

284

IN THE PAPER.

TERMS

Agiaonema : Element, definition of, 244 adnot.
angustifolium N. E. Br. 243 Epipremnum humile Hook. f., es
angustifolium var. undulatum Equivalent synonyms

Ridl. 243 Eupomatia Belgraveana F. Matt
Schottianum Mia. 24+ 231
Schottianum var. Brown1i Exact synonyms 233

Engl. 243 :

Schottianum var. genuinum Layo eeed Mae” ah

Engl. 243 :

Absolute Synonyms 233 Isonym 233

Actinophlceus cuneatus Burr; 233 are as

Andreoskia sc : :

Linum multiflorum Lam 235

Andrzeiowskia icuien: adinla 2

Andropogon Bequertii De Willd. oot einen De 235

239 Mesembryanthemum sect. Minima

Arenga Griffithii Seem. 260 Haw. 231

Arum colocasia L. 241 Mixtum Compositum, definition of

ie esculentum L. 241 244 adnot

ugusta 252 :

Augustia 253)0)' Neraae sect ae sa

aciuom si Nomina invalida 224

Bruguiera conjugata (L) Mer>. 242 Pentagonia 252

Bruguiera gymnorrhiza (L) Lam Pentagonium _ 252

242 aoa al. oryzoides L. 226

entaceras 252

Calandrinia polyandra Benth 233, Pentaceros) ; 252
230 Plectocomiopsis dubius Bec. 230

Calla silvestris Bl. 234 Poa amboinica L. 246

Calophyllum Calaba L. 259 Poikilospermum 252

Calophyllum Jacquinii Fawce. et. Psilanthus 252

Rendl. 259 Psilosanthus 252

Cerbera Manghas L. 245 Ptychosperma cuneatum Burr. 233

Cerbera lactaria Ham ex DC. 246 Ptychosperma tenue Becc. 233

Cerbera odollam Gertn. 245 Radiola linoides Roth. 235

Clerodendron Jackianum Wall. 259 Radiola Radiola Karst. 235

Clerodendron nutans Don. 250 Raphidophora nigrescens Hook. f.

Clerodendron nutans Jack 250 234

Colocasia antiquorum Schott 241 Raphidophora silvestris Engl. 234

op easls antiquorum var. typica Raphidophora Wrayi. Hook. f. 234
ngl. 243 Rhizophora apiculata BI. 269

Colocasia esculenta Schott. 241 Rhizophora candelaria D.C. 269

Colocasia esculenta var. antiquorum Rhizophora conjugata L. 242, 269

(Schott) Hubb et Rhed. 241, 243 Rhizophora gymnorrhiza L. 242, 260
Colocasia esculenta var. colocasia Rhopalodia vermicularis O. Mull
(A) Be, : 241 forma recta Fl. Rich. 230
ae el 2 Spirotecoma Dalla Tore et Harms

Conophyton Haw. Ss pide

ynonyms, absolute 233

Cymbopogon Bequertii De W ll. Synonyms, equivalent ° aaa

Cyrtophyllum? lanceolatum D.C. Syenyins, Seat 233

230 Talinum polyandrum 235,230

Cucubalus angustifolius Mill. 234 Typonym 233

Cucubalus Behen L. E 234 Unona cochinchinensis DC. 233

Cucubalus latifolium Mill. 234 Unona Desmos Dunal 233

Desmos chinensis Lour 225 Valid publications 2-8

Desmos cochinchinensis 226, 233 Wendia 252

Wendtia 252

Gardens Bulletin, S.S.

285

4

THE NOMENCLATURE OF TYPES

By C. X. Furtapo, B.Ag.,

Botanic Gardens, Singapore.

Page
1. Introduction ss a oe 0 285
2. Varieties of Types a Fis yes 2. ee
3. Explanation of Terms, and Examples ey, it 289
4. The Labelling of Type Plants in the Garden 153) ee
5. The Labelling of Type-Specimens in the Herbarium ... 306
6. Keferences... tes - Je ... 3808
7. Index bi ms: Ad “ ccnp 809

1. Introduction

Not many years ago, when I first became interested in
systematic botany, I experienced considerable difficulty in
comprehending the full implication of the terms ‘‘Type’’ and
““Co-Type’’. And although systematic botanists have now
accepted the principle of the ‘‘Type basis Concept’’, a perusal
of recent taxonomical papers combined with a personal inspection
of the labelling of specimens in some of the well-known herbaria
in Europe leads me to conclude that systematists are not yet
agreed on a uniform system of naming both type specimens with
their duplicates as well as those specimens that are almost as
valuable to taxonomists as the types themselves. This disagree-
ment is all the more surprising when one realises that systematists
would be the first to admit the value of uniformity in the naming
of type specimens for taxonomic reference. So great, however,
is the confusion at present reigning concerning the nomenclature
of some kinds of types that it more than bears out the remark
made by Dr. Agnes Chase, the well-known agrostologist, with
reference to plant nomenclature in general: ‘‘Paradoxical as
it may sound, it is the effort made during the last twenty years
or so to bring about stability and uniformity in the use of names
that has caused such a bewildering diversity’’.

Now in the following summary I have gathered together
most of the important terms used to designate type-specimens in
the hope that this compilation may provoke the interest of
herbarium-keepers in the matter. Obviously, by adopting a

Vol. IX. (1937).

286

uniform system of naming the type material in their charge,
herbarium-keepers would increase considerably the value of the
specimens. For the sake of clearness I have had to define again
a few old terms and add a few new ones.

One apology I have to make: I cannot now cite authority
for every one of the terms adopted in this paper.* This is because
the compilation was originally for my own instruction only.
However, most of the terms here discussed can be found in
Dr. B. D. Jackson’s Glossary of Botanical Terms, 4th Ed. (1928).

2. Varieties of Types

Nowadays a description of a new species is based on certain
specimens, some or all of which are preserved in a herbarium.
It is with the uniform naming of these specimens and their
duplicates that I am most concerned. But because palezeonto-
logical botany has its own system of type-classification,
paleontological types will not be considered in this paper.

For accuracy in taxonomic work, especially in the revision
of species, one must know the status of each specimen: whether,
for instance, it was part of the original material on which the
species was founded; how it came to be regarded as authentic or
by whom it was identified; whether it was selected by the author
as typical of his species or if it supplied any particular informa-
tion. And in the case of duplicates, many other problems arise.
For instance, if the original specimens on which an author
founded his species are in one herbarium, can one consider the
duplicates in another herbarium as equivalent? Did the author
examine all the duplicates of each collection of the original
material before they were distributed, or had he only one
specimen of each at the time he drafted his description.

For taxonomic purposes equivalent duplicates must have
been gathered at the same time from the same plant and must
have the same collection number (cf. Isotype and Clastotype
below). But it is common for collectors to take their duplicates
from several plants of apparently the same species and perhaps
at widely different dates, and to give these various specimens an

* After completing my paper I had an opportunity to consult
Dr. D. L. Frizzell’s compilation entitled TERMINOLOGY OF
TYPES, published in The American Midland Naturalist XIV, 1933,
pp. 637-668, and obtainable in a reprint form from the University
of Notre Dame, Indiana, U.S.A. Though some terms like Isotype,
Isoholotype, Isolectotype, etc. as used in systematic botany have not
been recorded, the booklet is of special value as it gives full particulars
regarding the authorship of the terms and the place of their publica-
tion, as well as references to the subsequent use of the terms in a
sense different from the original. There are dealt in it 233 terms or
phrases of which the word type, or.its variant, forms a part, but only
about half of this total number of terms explained pertains to the
nomenclatorial types in biology, the others being non-nomenclatorial.

Gardens Bulletin, S.S.

287

identical field number.* In some cases, as some fungi, herbs
and small, little-branched palms, it will be evident from the
material whether or not the duplicates are equivalent, but in
most other cases it will not be so self-evident; and, should the
collection be of some large and difficult genus like Eugenia, Ficus,
Eria, Calamus, Polvpodium, Sphagnum or Sargassum, one might
doubt the collector’s ability to distinguish every species in the
field. A collection of such duplicates may be found after a
critical study in the herbarium to comprise more than one species
so that the duplicates are not equivalent; and if they are
distributed before the mistake is realised, much confusion will
result.

To avoid such doubts, and for reasons which will be
apparent, I propose the following terms for the labelling of type-
specimens so that in most cases there can be no doubt of the
status of a herbarium specimen and its taxonomic value:

TYPE SPECIMEN: in a strict sense, is a herbarium
specimen (a) that has been cited in the original description as
having been seen by the author; or (b) if not cited, any original
specimen on which the description of the species was based. For
convenience all secondary types, which may be either specimens
chosen as standards subsequent to the original description of the
species or merely specimens matched with the type specimens,
and all the duplicates of the type specimens sensu stricto and of
the secondary types, are called the Type Specimens sensu lato.
Type specimens sensu lato may be divided as follows:

A: Primary Types or Proterotypes (Proteros=first, former):
are the type specimens sensu stricto. These are subdivided
as follows :—

1. Ho.otypPre (Holos=entire): The specimen chosen
by the author and cited in his original description as that
by which the species must be interpreted.

2. PARATYPE (Para=beside): any proterotype other
than the holotype.

* One cannot condemn too strongly the practice of lumping
together specimens collected from different localities or/and on
different dates and then giving the same field or distribution number

as if they had been collected on one and the same day and from one
and the same field; for under field conditions it could not have been
possible for the collectors to compare the specimens, or the plants
from which the specimens had been taken, in a living state. Further
the specimens so distributed will convey a wrong idea of both the
flowering and fruiting seasons of the plants as well as their distribution
in the district. This will hold good even when the different localities
whence the specimens were obtained lie in the same political or
botanical district.

Collectors therefore should make it their rule, as far as possible,
to collect from the same bush or plant all the specimens to be included
under one field number; or, if this is not possible, to collect
simultaneously from different plants growing in the same locality
so that they may be compared in the living state in the field by the
collector himself.

Vol. IX.\(€1637):

288

3. SYNTYPE (Syn=with): Any proterotype, if no
holotype has been chosen.

B: Secondary Types or Heterotypes (Heteros=other): These
are specimens chosen or studied subsequent to the original
description. They may be divided as follows :—

1. LECTOTYPE or LECTOHOLOTYPE (Lektos=chosen) :
A syntype selected as a holotype. If the selection is made
by the author of the species himself, then the Lectotype
may be termed an IDEOLECTOTYPE. (/deos=peculiar).

2. LECTOPARATYPE: A syntype not chosen for the
Lectotype.

3. APOTYPE (Apo=from): A specimen on which is
based the first correct interpretation of a species of which
the description is too inadequate and the proterotypes are
too imperfect or abnormal to furnish a clue for its correct
identification. If both the types and the specific descrip-
tion cannot be identified except by an historical inquiry
into the use of the specific name, then the specimen that
supplies the historical clue becomes the Logo-Apotype
(Logos = discourse).

4. TopotyPE (Topos=place): A specimen from a
type locality agreeing with the type.

If it is from the Holotype, Paratype, Lectotype, etc.
locality, then the specimen must be called accordingly a
Topoholotype, Topoparatype, Topolectotype, etc.

5. Icotypr ([kos=what is like): A specimen agree-
ing with the accepted standard of the species, but not
from any Proterotype locality.

6. NEOTYPE (Neos=new): A _ specimen agreeing
with the original description of a species, and selected as
the type when no material from the proterotype collection
exists.

According to the locality from which the Neotype is
obtained, the specimen would be called Neotopotype (from
the Holotype or Lectotype locality), Neoparatype, (from
the Paratype locality), or Neoicotype (not from any
Proterotype locality). (See also under Proteromerotype
below).

7. IDEOTYPE or IDIOTYPE ([dios=peculiar or own):
A specimen identified by the author as typical of his
species. . |

According to the place whence the specimen was

collected, it could be Ideotopotype, Ideoparatype, etc. (see

also under Lectotype and Proteromerotype).

8. PROTEROMEROTYPE (Meros=a part): A specimen
taken from the holotype plant because no herbarium
material was preserved at the time of description.

Gardens Bulletin, S.S.

289

If the specimen was taken by the author of the species,
then the specimen may be called Ideoproteromerotype or
simply Ideomerotype (cf. also Merotype below).

C. Duplicate Types or Antitypes (Anti=against; instead of):
These are specimens purported to have been collected by the
same collector at the same time and in the same place, and
bearing the same number (if any) as the primary or the
secondary types; or else fragments taken from types; or
specimens derived from plants raised vegetatively or by seed
from the type plants. They may be classified as follows :—

1. Isotype (Jsos=equal to): A specimen collected
from the same plant as the type, at the same time, and
bearing the same collection number.

2. HaptotypPE (Hapto=I fasten upon): A specimen
purported to have been collected simultaneously with the
type and bearing the same collection number, but not
certainly from the type plant.

3. MEROTYPE (Meros=part): A specimen taken
from a type plant but not simultaneously with the type,
and consequently with a different number (cf. Protero-
merotype).

4. CLONOTYPE (Klon=a young shoot): A specimen
from an asexual offspring of a type-plant.

5. SPERMOTYPE (Sperma=seed): A specimen from
the seed or spore offspring of a type plant.

6. CLASTOTYPE (Klastos=broken): A fragment of
a type specimen. ~-

Note.—According to the status of the type-plant (vide
infra), or type-specimen, from which these duplicates were
obtained, so these prefixes can be added to define the duplicates
more precisely e.g.: Isoholotype, Iso-paratype, Iso-apotype,
Iso-neotype, Haptoneotype, Mero-holotype, Clonoparatype,
Spermolectotype, Clastoproteromerotype.

3. Explanations of these Terms and Examples

1. PRIMARY TYPES :—In many older and also in some
modern works on botany the specimens actually studied and on
which the authors based their species are not cited. But from
the range of distribution given for the species and from the
history of the specimens together with any notes by the persons
who studied the specimens, it is often possible to find out the
specimen or specimens on which the species was based. Such
specimens, therefore, though not cited in the original description,
are also to be included in the Primary Types as defined herein.

Example 1:—Demonorops pseudomirabilis was described by

Beccari as a new species but with the following particulars under
its habitat or range of its distribution: ‘‘Cultivated at Buitenzorg,

Moe. IX \Otos7):

290

from Palembang in Sumatra” (Rec. Bot. Surv. Ind. II, 1902 p. 226).
Here no definite specimen is cited, though the holotype is in Beccari’s
herbarium, now forming the part of Webb’s Herbarium in Florence.

Example 2: the Catalogus Plantarum in Horto Botanica
Bogoriensi Cultarum, I1, 1844, p. 58, Hasskarl published a new aroid,
Scindapsus pictus, with a brief specific description only, without citing
any specimens whatsoever or giving the range of the distribution of
the species. Should there exist any specimens on which Hasskarl
based the specific description of his species, those specimens will
have to be included under the Primary types. If only one, it will be
the Holotype of the species.

2. HOLOTYPE and PARATYPE :—Botanists frequently
call the Holotype the ‘‘Type’’, but it is convenient to have the
term Type (i.e. Type-specimen) with a more general meaning to
indicate any specimen which is recognised as typical of the
species. It is not essential to indicate directly the Holotype,
indirect indication being quite sufficient. But, in order to be
valid, indications of the Holotype, whether direct or indirect,
must be made in print and simultaneously with the publication of
the species itself. As for Chirotypes (types indicated in herbarium
only and not in literature or types of manuscript species), these
cannot be taken into account to the prejudice of published types.
If only one specimen was cited by the author of the species, or
if there was only one specimen for study and that was not cited,
then that specimen becomes the holotype.

Example 1:—In the case of Demonorops pseudomirabilis (see the
Example zt under the Primary Types) only one specimen derived from
the plant cultivated in the Buitenzorg Botanic Gardens, was studied
by Beccari. That specimen therefore is the holotype of the species.

Example 2:—-F. von. Mueller, in erecting a new species, Liviston2
Marie, made the following observations : ‘“‘Descriptio fructus, gue
species precipue limitat solummodo e speciminibus occidentalibns
pendet quum tantummodo foila hujus palmz superbe ex Australia
centrali possideo, sed plantam ex utraque regione conspecificam
arbitror’’ (Fragm. Phyto. XI, 1878, p. 54). Since the delimitation
of the species was said to have been based chiefly on the fruits, it
means that the West Australian specimen which bore fruits, and not
the Central Australian one which was sterile, is the Holotype of
Mueller’s species. (The Central Australian is a paratype). When,
therefore, Mueller recognised that the Central Australian plant was
specifically distinct from the West Australian, he could not reserve
the name, L. Marie, for the former and give the latter (the Holotype
of L. Maria) the new name, L. Alfrediz, (Vict. Nat. XI, 1892 p. 112).
If a new species had to be erected to distinguish between two kinds
of palms, then the paratype of ZL. Marig (but never the holotype)
could have been made the Holotype of the new species (cf. also
Art. 52 of the Internat. Rules Bot. Nomencl., 1935).

Example 3: (in
Journ. Roy. Asiat. Soc. Straits Br. XXXIV, 1900, p. 98,) the author
of the species gave the following particulars: ‘‘Bismarck Archipelago
(Micholitz). Flowered in the Botanic Gardens, Singapore, Feb.
1900’. The description itself was obviously derived chiefly from a
living plant. Hence only the specimen taken from the plant
cultivated in the Singapore Gardens and studied in detail by Ridley
must be regarded as the Holotype. Should it be necessary, at any

Gardens Bulletin, S.S.

291

future date, to regard Micholitz’s own specimen from the Bismarck
Archipelago as distinct from the one that was derived from the plant
cultivated in the Singapore Botanic Gardens, then, the name 4d.
terminale Ridl. should be retained for the specimen derived from the
cultivated plant.

Example 4:—In REINECKE, Fl. Samoa-Ins. in Engl., Bot.
Jahrb. XXV_ (1898) 588, Warburg published a new binomial under
Cybhokentia thus:

“‘C samoensis Warb. (prob.=Clinostigma samanse Wendl.)
Bonplandia X (1862) 106)
“Upolu: Mai 1894 (n. 322)”’.

Now this binomial of Warburg is being taken not as a new
combination based on Wendland’s species cited above in the synonymy,
but as a new species with Reinecke’s specimen from Upolu in Samoa
as the holotype. But from Warburg’s remarks on the species it 1:
evident that the binomial is an isonym of Wendland’s species (2.e.
Wendland’s species is the holotype of Warburg’s binomial) and that
the word ‘‘prob.’’ following the binomial was added to indicate that
Reinecke’s specimen was referred to the species with some doubt.
I transcribe here the portion of Warburg’s remarks that is most
pertinent to the point in question :

“Dass diese Art mit Clinostigma samanse identisch ist, erscheint
mir nach WENDLAND’s Beschreibung (Bonplandia 1862, p. 196) zwar
sehr wahrscheinlich aber nicht ganz zweifellos; nach derselben ist z.
B. die Rachis der Hauptverzweigungen des Bliitenstandes viel dicker
ies-a0 emi, berranserem, Exemplar nik: 9—8)-5. caccis..csccvegersncuecanssseecns
RRR Ree MIS 28 REET. foi VD a clk AEN s desde aed Lo daatlarenpiiceeneas) pales 1. DESORE
mit Recht, die Verwandtschaft der samoanischen typischen Art mit
Cyphokentia, und ich sehe in der That nicht den mindesten Grund,
Warum unsere samoanische Art, sowie auch der WENDLAND’ sche
Typus der Gattung Clinostigma nicht zu Cyphokentia gehoren soll,
namentlich nicht in der weiteren Fassung Drude’s in ENGLER-PRANTL.

‘“‘Ausser dieser Art werden noch 2 Arten zur Gattung Climostigma
gebracht————_[ Warburg then refers to an undescribed species
and Cl. Mooreanum, but not to Cl. samanse Wendl.].

Further, whereas in Reinecke’s paper above quoted (cf. of.
cit. Pandanacez pp. 578-581; Palmz pp. 588-592; Ficus pp. 613-617)
Warburg. published a new species invariably with a diagnosis in
Latin and with an ‘‘n. sp.’’ at the end of the binomial, he did
neither of these things in the case of Cyphokentia samansis, being
content to quote Wendland’s species as a synonym and to show how
Reinecke’s specimen differed from Wendland’s description of Cl.
samense Wendl.

This procedure of interpreting a new isonym (=new combination
or new name) is also in accord with the new International Rules of
Botanical Nomenclature, ed. 1935 (cf. Art. 54), which prohibit the
adoption of a new holotype for the species which has been transferred
from one genus to another, the prohibition being valid whether or not
this transference is accompanied with a new description drawn from
a different plant or species (The case discussed in Examéfle 2 under
SYNTYPES stands on a different footing).

In view of the above, one has to treat Cyphokentia samensis
Warb. not as a new species, but as an isonym of Clinostigma samanse

Wendl.
_ Hence to avoid future complications and misinterpretations of the
binomial Clinostigma Warburgii Becc. in Atti Soc. Tose. Sci. Nat.

Vol. IX. (1937).

292

XLIV, 1934, p. 44,* I asked the late Count Professor Martelli to
insert the words ‘‘quoad, icon. et specimen a Reinecke lectum”’
immediately after its synonym, Cyphokentia samaensis Warb., and to
add the diagnosis in Latin, though in his manuscript Beccari had
considered C. samansis Warb. as a new species independent of
Wendland’s, and treated Cl. Warburgii Becc. only as a new name
for the former.

Example 5:—/xora nigricans R. Br. made its appearance as a
nomen nudum in Wallich’s Catalogue (1828) n. 6154 and in Wight’s
Catalogue (1833) n. 1335. In 1834, however, it was published with a
description in Prodromus Flore Peninsula Indie Orientalis Vol. I,
p. 428 by Wight and Arnott, who after quoting the references to the
two catalogues, make the following remarks:

‘‘This was sent to Dr. Wallich by Dr. Wight, but is not noticed
under any of the letters of n. 6154; it perhaps, therefore, forms
another species in the List, although we cannot see any difference
between it and /. nigricans’’.

It is evident from this quotation that, though Wight and Arnott
seem to take /. nigricans R. Br. in Wall. Cat. as a validly published
species, yet the description they have given in the Prodromus is
entirely of Wight’s specimen. That this conclusion is correct is
further confirmed by the observations Robert Wight has made in his
/cones, which consist of figures of the Indian plants described in the
Prodromus and in Wight’s J/llustrations of Indian Botany. After
quoting bodily the description (with one or two verbal additions)
Wight writes as follows:

‘‘Some points of the character do not accurately correspond with
the figure, which may be accounted for by the former being taken
from dried specimens, the latter from the fresh plant; the divisions
of the stigma do not separate at first and the mature berry is globose
and purple, not unlike a small black cherry, but changes in drying.”’
(Icones I, 1840, t. 318).

Now since Arts. 36 and 37 of the botanical Rules ed. 1935, do
not recognize any of the names in the two above quoted Catalogues,
even though these names had been placed on herbarium duplicates
and thus made available to several botanical institutions, it follows
that the binomial /. migricans became systematically valid only with
the botanical description published by Wight and Arnott; and as this
description was based entirely on Wight’s specimen n. 1335, this
specimen is therefore the holotype of the species, which ought to be
quoted as /. nigricans Wight and_Arnott, Prodr. Ind. J. (1834) 428,
though the Rules also permit one to quote it as /. migricans R. Br. ex
Wight and Arnott, despite the fact that the type of the species is not
any of the specimens that R. Brown had seen.

* It may be noted here that, owing to ill-health of the late Count
Professor Martelli, I was entrusted to frame all the diagnoses in
Latin for Beccari’s new genera and species published in the Aftz so
that the genera and species may be validated according to the
resolutions passed by the International Botanical Congress, Cambridge,
1930. But before the paper was out and when it was too late to make
any changes in it, a modification to this resolution was announced
(Journ. Bot. LXXII, 1934 Suppl.) whereby the diagnoses in Latin
were rendered unnecessary since Beccari’s’ diagnoses were all in a
modern language (Italian) and published before the end of the year
1934—two conditions that required to be satisfied in order to
legitimize any new taxonomical unit published in any language other
than Latin.

I mention these facts so that systematists may know which views
to follow should they notice any discrepancies between the Latin and
the Italian descriptions.

Gardens Bulletin, S.S.

|

sie

BAN crt tl aha

293

From this it is obvious that any interpretation of the species by
a reference to any of the specimens distributed by Wallich under
6154 A-D is not permissible, though these specimens may be included
as the paratypes of the species.

According to King the specimens distributed under Wallich’s
Cat. n. 6154 are of more than one species. (Materials Fl. Malay.
Pen. IV, 1904 p. 149). Probably the failure to recognize this, and
also to accept Wight’s specimen as the holotype of /. nigricans has
contributed to a great deal of confusion over the species.

3. SYNTYPE:—If the author has not indicated either
directly or indirectly the Holotype specimen and if there are
more than one Proterotype, then all the Proterotypes are called
Syntypes of the species. No species, therefore, can have both
the Holotype and Syntypes, or Paratypes and Syntypes. In its
original sense the term Co-type* (Co = Cum = with, together)
is also synonymous, but because Cotype has been used loosely
for Paratype, Isotype, Haptotype or Merotype as well, I propose
to drop the term in favour of Syntype, which is now becoming
popular (See also Lectotype).

Example 1:—In Philipp. Journ. Sci. 58 (1935) 14, Danser quoted
the specimen, /eg. Ramos and Endano n. 38740 in the Manila
herbarium as the type (=holotype), and its duplicates in the herbaria
of Berlin, Leiden and Buitenzorg as the cotypes, of his species,
Amylotheca amplexicaulis. As all the three latter specimens were
definitely cited by Danser as examples of his species, the term
‘“cotypes’’ was used here to indicate the specimens that are both
paratypes as well as haptoholotypes (=duplicates of holotypes) of
the species.

In another case (op. cit. Vol. 58, p. 23) Danser indicated the
specimen leg. McGregor n. 43873 from the Manila herbarium as _ the
type (=holotype), and its duplicates in the herbaria of the New York
Botanic Gardens and of the University of California, Berkeley, as
the cotypes, of Loranthus cordilimbus Merr. (synonym of Amylotheca
saccata Dans.). But as Merrill has not referred in his description to
the duplicates of the holotype as examples of his species (op. cit. 30,
1926 pp. 389 and 3o91), it follows that by the term ‘‘cotypes’’ Danser
merely means haptoholotypes.

Example 2:--In Prodr. Flor. Nov. Holl. I (i810) 225, Robert
Brown published the new binomial, Heleocharis capitata, to include
a species which he had collected in Australia, with the following
remarks :

“Scirpus capitatus Linn. sp. pl. ed. Willd. I p. 294 (secundum
synonyma Brownei et Sloanii, sed a planta virginiana Herb. Gron.
diversus ; hzec autem, quoniam Linnzus, cum primum Sp. pl.
edebat, nullum hujus exemplum in herbario suo habebat, istius
speciei unica auctoritas est.).

_ Scirpus culmo nudo, spica_terminatrice subrotunda. Hort.
Cliff. 21 (fide speciminis Herb. Cliff)’’.

Here Brown does not state that his species is either S. capitatus
L. sensu Willd. or S. capitatus Willd. excl. syn Linn. et. Gronov.,
but that the Linnean species with its type specimen from Virginia
in the Gronovian herbarium [2.e. Clayton’s plant] is not his
Heleocharis capitata, that S. capitatus L. sensu Willd. is a mixed
species, and that, as far as is known to him, only Willdenow’s
synonyms referring to Patrick Browne and Sloane are to be included
in his species.

* As a rule Zoologists use this term correctly to mean syntypes.

Val. LX.\63037).

294

Consequently it is illegitimate to treat Heleocharis capitata R. Br.
as a new combination based on Scirpus capitatus Linn., Sp. Pl. |
(1753) p. 48, and thus make precisely the plant that R. Brown excluded
from his binomial the holotype of Brown’s species. Despite the
current practice, therefore, it is right to look upon H. cafitata R. Br.,
not as new combination, but as an entirely new species, having for its
syntypes the following specimens: R. Brown’s specimen from
Australia [Gulf of Carpentaria]; Clifford’s specimens; and the
Jamaican specimens from Sloane’s and Browne’s herbaria: all of
which were referred by R. Brown in his description or remarks.
The Caribean (West Indian) specimens that Willdenow referred to
S. capitatus are not to be included among the syntypes of Brown’s
species, nor should Willdenow’s name be added in brackets after
Brown’s binomial and before Brown’s initials (as it has been done
by Dr. O. A. Farwell in Rhodora XXXII, 1930, p. 181) (cf. Example
4 under Lectotype).

Note:—Several botanists including C. B. Clarke (Journ. Linn.
Soc. Lond. XXX, 1894, p.'310), Dr. S. F. Blake (Rhodora XX, 1918,
p. 24) and Dr. O. A. Farwell (Rhodora, XXXII, 1930, p. 181) have
ignored altogether the remarks made by R. Brown to the effect that
at the time of the first edition of the Species Plantarum (1753) no
specimens of Scirpus capitatus existed in Linnzeus’s herbarium, and
have accordingly caused a good deal of confusion over the interpreta-
tion of S. capitatus and Heleocharis capitata, Clarke and Blake going
to the length of treating the former binomial as the ‘“‘basinym’’ of the
latter. It is true that there are now in the Linnean herbarium two
specimens both named as S. capitatus by Linnzus himself; but at
the same time there is no valid reason to disregard R. Brown’s note
about their non-existence there in 1753. On the contrary the fact
that neither of the specimens was listed by Linnzus in the three
enumerations of his herbarium supports R. Brown’s contention.
Mr. J. E. Dandy of the British Museum, London, who has looked
up the matter, at my request, informs me that one of the specimens
identified by Clarke (l. c.) as H. ovata was received from Kalm, and
that the other representing H. caribea was from P. Browne.*
Mr. Dandy has also verified that H. capitata R. Br. is identical
with H. geniculata (L) Roem. and Schult. quoad var #, the diagnosis
from Hort. Cliff. quoted by R. Brown being according to the original.
This means, therefore, that H. carib@a (Rottb.) Blake and JZ.
capitatus R. Br. are synonyms of A. geniculata (L) R. and S. as here
interpreted (vide Example ¢ under Lectotype) and also that the
species that pass as H. geniculata and S. capitatus should be known
respectively as H. elegans Roem. Schult., and A. fliformis (Lam.)}
Kunth=AH,. tenis (Willd.) Schult.

Example 3:—In Florz Siamensis Enumeratio II, pt. 2 (1934) 147,
the late Prof. W. G. Craib published a new variety, /xora affinis
Wall. ex G. Don var. arguta Craib comb. nov., by the following
citations :

/. arguta R. Br. ex King in Journ. As. Soc. Beng. LXXII,

2 (1904) p. 74. /. nigricans R. Br. ex Wight et Arn. var. arguta

Hook. f., Flor. Brit. Ind. III (1880) 149 p70 parte.

After enumerating several specimens from the Peninsula Siam,
Prof. Craib indicates that, outside that region, this variety occurs
only in the Malay Peninsula and that its type (=holotype) is from
Penang; but as no specimen from Penang was quoted by Hooker,
who was the first to make the valid use of the term avguta in the
synonyms quoted above, it seems worth while to investigate the exact
status of the types of all the three names.

~

* Linneus acquired Browne’s herbarium in 1758. Browne
published his Natural History of Jamaica in 1756.

Gardens Bulletin, S.S.

295

Under /. nigricans var. arguta, Hooker cited only two specimens,
both of which are from India, one from Heyne (the only one cited in
Wallich’s Catalogue n. 6157) and the other from Ritchie; and, though
the species proper is said to occur in Burma and the Malay Peninsula,
Hooker has not made any definite statement as to the occurrence of
the variety outside India.

Now King. Z.c. discovered that Wallich’s n. 6154 A-D, to which
the manuscript name /. mzigvicans R. Br. was first attached though
not, validly published, represented more than one species and
concluded that Hooker had made a muddle of /. migricans in having
quoted for it Wallich’s n. 6154 without any distinction of letters,
King having overlooked the fact that /. migricans was validated much
earlier than Hooker by Wight and Arnott in their Prodromus / (1834)
428, a reference also. given by Hooker, and that, therefore, the
species had to be interpreted by a reference to Wight’s n. 1335 (wide
Example 5 above under Holotype). In the hope of getting over this
confusion erroneously attributed to Hooker, King restored /. ?2arguta
R. Br. in Wall. Cat. n. 6157 for the Malayan plant referred by Hooker
to L. nigricans, but quoted at the same time /. nigricans var. arguta
Hook. fil. as a synonym and made the following observations on the
types :

‘‘The type specimens of this [species] in the Wallichian Catalogue
[n. 6157] consist of two sheets distinguished by the letters A. and B.
The former was received from Herb. Heyne, and is therefore supposed
to have come from Southern India. Sheet B is attributed to Penang.’

Now according to Art. 58 of the botanical Rules, King was not
obliged to adopt the trivial name avguta for his species, even though
the species was to be validated by citing /. migricans var. arguta
Hook. f.. But having adopted the name, cited the variety in the
synonymy, and quoted at least one of the two syntypes of the variety
as a type of the species, King’s species cannot be regarded except as
a new combination based on Hooker’s variety. One is not at liberty
to assume that King had drawn his specific description entirely from
the new set of specimens and that he had not personally seen any of
the syntypes of Hooker’s variety, regarding Heyne’s specimen seen
by King as the duplicate of Hooker’s syntype (cf. Arts. 18, 49, 50-52).
In Wallich’s Catalogue only one specimen is listed under n. 6157
without any distinctive letter, and that one is from Heyne, which is
also quoted as the syntype of Hooker’s variety. As Hooker did not
cite any specimen from Penang, Wallich Cat. n. 6157—-B from Penang
cannot be included among the syntypes of King’s /. arguta, unless
it can be satisfactorily shown that the omission was clearly a printer’s
error or a lapsus calami. If it is proved that Ritchie’s specimen
cited by Hooker is distinct and cannot be included under the variety,
then Heyne’s specimen will become the Lectoholotype of /. argztu
King as well as of /. nigricans var. arguta Hook. f.

Now if Prof. Craib was of the opinion that neither of the syntype2s
of Hooker’s variety corresponded to the Malayan specimens referred
by King to /. arguta and still considered that his own variety /.
affinis var. arguta should refer only to the Malayan specimens cited
by King, and not to any of the two Indian specimens referred by
Hooker, then he could not validate the variety by citations. He could
have however quoted the synonyms and shown how the syntypes of
Hooker’s variety were to be excluded. But since he has not followed
this procedure, it means that, despite the use of fro parte after
Hooker’s trinomial quoted in the synonymy of /. affiis var. arguta
Craib, the latter variety has to be considered as a new combination
entirely based on Hooker’s variety, and that the syntypes of the
former are precisely those of the latter and not any more or less.
In new combinations the Rules do not allow a person to choose a new

Vol. IX. (1937).

296

set of syntypes from specimens that are not the primary types of the
Original species or variety on which the combination is based (Arts.
18, 490, 50-52 and 35).

4. LECTOTYPE:—Where there is reason to doubt whether
the syntypes are specifically identical it is useful to distinguish
one specimen as the Lectotype. And it is advisable that a
competent systematist should make this selection. It is futile
merely to choose the specimen first cited in a description where
the order of citation has no such preferential significance.
Besides no specimen is likely to be uniformly accepted as the
standard (Lectotype) of a species unless cogent reasons are
brought in favour of the selected specimen. Where there are
equally good botanical reasons in favour of more than one
specimen being chosen as the Lectotype, systematists must, in
the interest of stability of the nomenclature, accept as the
standard of the (original) species the first published choice of a
Lectotype.

Note:—I wish to protest against unnecessary alterations
being made in the status of the syntypes even when the changes
made are by the author of the species himself. If an author
desires to bind the interpretation of his species to any one
particular syntype, then to make the binding valid he must
indicate, at the time of publication of the species, that specimen
as the holotype (unless of course the omission from the original
description is a lapsus calami or the fault of copyists or printers,
in which case correction should follow as early as possible).
Later when the specific binomial has become, by publication,
an entity in the dominion of taxonomical science, the author has
as much or as little right as anybody else to make valid alterations
in any part of the species. If, subsequent to the publication of
the species, an author who has omitted to name the holotype in
the original description, doubts the specific identity of the
syntypes or fears a misinterpretation of his species, he should
investigate the entire species again and choose a lectotype; or,
if that is not possible, he should indicate both the specimens on
which he had based the most important portion of the specific
diagnosis, as well as those which appear to him to be not typical
of the species, though not cited in the original description.
But a lectotype should not be chosen unless it is really necessary;
in other words, if a botanist examining the syntypes of a species
cannot find any specific or varietal difference among them, he
should not distinguish one as a lectotype. This proeedure will
not only prevent a systematist from making an unnecessary or
haphazard distinction among the several syntypes and avoid
thereby a misdirection of future studies regarding the species
and the types, but also oblige him, before selecting the Lectotype,
to show the need for distinguishing one syntype from another.

Example 1:—In 1931 Dr. E. Hultén and Dr. H. St. John jointly

published a new aroid, Lysichitum americanum, without indicating
the Holotype (Svensk Bot. Tidskr. XXV, 1931 pp. 153-464). Though

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297

the paper was published jointly, they elaborated their work in two
different places, the senior author in Europe and Asia, the junior
author in America ; and they admitted that even at the time of
publication they had not had an opportunity to complete the revision
side by side. Later, apparently they met in America and jointly
selected a specimen which, for several reasons, seemed to them best
“fitted to stand as the type of the new species’. (Of. cit. XXVIII,
pt. 2, 1934). Though this selected specimen _ is proposed as the
Holotype, it is evident from the definitions proposed above that it may
not be accepted except as a Lectotype, and, being chosen by the authors
of the species themselves, it can be labelled as an ldeolectotype.
Should, in future, the species prove to be composite, it is questionable
whether one should abide by this Ideolectotype. An objection that
becomes at once apparent is that, up to the time of the publication
of the species, the senior author had not seen the specimen later
chosen as Ideolectotype, while it is not sure whether the junior had
based on the specimen any portion of his conclusions regarding the
new species. But, if no serious objection is found against this
Ideolectotype, preference should certainly be given to it in any future
investigation concerning the status of the species.

Example 2:—Having shown that the syntypes of Demonorops
propinguus Becc. belong to more than one species, I retained this
specific name for Griffith’s fruit-bearing specimen (a syntype of VD.
propinquus) and transferred Forbes’s specimen (another syntype) to
another species (2. confusus) then described in Gard. Bull. S.S. V/I/,
1935 Dp. 347 and 348. Griffith’s specimen becomes accordingly the
Lectotype of DV. propinquus Becc. It should not be permissible to
disregard this first selection of the Lectotype on such trivial grounds
that the specimen was not the one cited first in the original description
or on the ground that it was Forbes’s specimen which had suggested
to Beccari the specific epithet ‘‘propinquus’’. Or else the nomencla-
ture of both the species (2. propinguus and LD. confusus) will become
unstable.

Example 3:
(1786) 20 that the Linnean species, Poa amboinica, agreed in its
description with a specimen from India, distributed under that name
by Koenig, while the Rumphian species, Pha@nix amboinica montana,
cited in the synonymy by Linnzus was totally a different species both
as regards its description and as regards its illustrations. Retzius
therefore excluded the Rumphian species from the synonymy of
P. amboinica L., giving, at the same time, a fuller description of the
species represented by Keenig’s plant, to which Retzius exclusively
reserved the Linnean binomial. It is not known whether Retzius had
with him Linnzus’s type; but from historical ground it appears
certain that Linnzus had based his specific description of P. amboinica
on a specimen collected by Keenig in S. India. Kcnig’s specimen in
Retzius’s herbarium (in Lund) becomes either the Lectotype of the
species if the specimen was the same as the one studied by Linnzus ;
or, if not, the specimen is a Neotype or, to be more precise, a
Neotopotype, and it will remain so unless the specimen actually
studied by Linnzus is discovered, when the specimen in Retzius’s
herbarium would become merely a Topotype.

Note:—Though the fact that Retzius had excluded the Rumphian
species from the synonymy of P. amdboinica L. was at times overlooked,
Retzius’s interpretation of the species was almost universally accepted
until Dr. E. D. Merrill (Interpr. Rumph. Herb. Amb., 1917, p. 88)
made a new combination Andropogon amboinicus (L) Merr., on the
supposed grounds that P. amboinica L. was entirely based on
Phenix amboinica montana Rumph.. After an extensive inquiry into
this subject, Mr. C. E. Fischer (Kew Bull., 1934, pp. 398-400) came
to the conclusion that Linnzus had based his description of P.

Vol. IX. (1937).

298

amboinica on a specimen from §S. India sent to him by Keenig and
that the Rumphian species quoted in its synonymy is altogether a
different species, belonging neither to the genus Poa in the Linnean
sense, nor to Eragrostis to which Keenig’s specimen and the Linnean
description of P. amboinica are now referred. In other words, Mr.
Fischer admits that the specimen on which Linnzus based the ’ chief
part of his specific description was not the Rumphian species, but a
specimen from Kenig. But because the Rumphian species had
suggested to Linnzus the trivial name for Koenig’s plant, Mr. Fischer
concludes that the specific epithet in P. amboinica L. should be reserved
for the Rumphian species referred to the genus Andropogon by
Merrill. Thus Mr. Fischer maintains that the Rumphian species
must be accepted as the Lectotype of P. amboimica L., so that
Andropogon amboinicus (L.) Merr. (1917) is a legitimate combination,
and that P. amboinica L. emendavit Retzius (1786)= Eragrostis
amboinica [amboinensis] (L. emend. Retz) Trinius (1840) is illegiti-
mate as a name for Poa riparia Willd. (1803)=Evagrostis riparia
(Willd.) Nees (1829). The reasons given by Mr. Fischer for
“(pomp ae Retzius’s legitimate interpretation of ‘P. amboinica L. are
botanically of minor importance (cf. Example 2 above); and so in
the interest of stability in the nomenclature of the species included
in. the original Poa amboinica L. the newer interpretation by
Dr. Merrill and Mr. Fischer should not be possible.

Though Art 52 of the International Rules of Botanical Nomencla-
ture, 3rd Ed. (1935) is incomplete, as the Appendix / which promises
to regulate the choice of Lectotypes is not yet published, one can
successfully maintain that Linnzeus had not fully studied the figures
and description of Phanix amboinica montana Rumph., _ (pre-
Linnean), and that the only plant he had studied in detail and on
which, moreover, he had based his diagnosis was a specimen either
sent to him by Keenig, or raised by Linnzus from the seeds sent by
Koenig. Therefore the type of Poa amboinica L. must be Keenig’s
plant and not the Rumphian species which Linnzeus had cited
presumably as an approximate synonym of P. amboinica; if the type
has been lost, Retzius has provided a Neotopotype. But even
granting that both Keenig’s plant and the Rumphian species have
each an equal claim to be chosen as the Lectotype, it seems but right
to allow the author who first recognised the mixed composition of a
species to select the Lectotype. A parallel may be found in A7zt. 56
of the Rules which confers full rights on the author to choose, when
uniting two or more species of the same date, any one of the specific
epithets as legitimate for the group resulting from the union of the
species, unless of course the resulting binomial becomes a tautonym
or a later homonym. Such reasons as the unsuitability of the trivial
pe cannot be successfully adduced against an already published
choice.

Example 4: I (1753)
48, with description, Scirpus culmo tereti nudo, subglobosa terminal,
was based on at least three syntypes: (a) Scirpus culmo nudo, spica
terminali [terminatrice | subrotunda L., Hort Cliffort. (1737) wis
(b) Juncus aquaticus geniculatus, capitulis equiseti, major Sloane,
Cat. Pl, Ins. Jam: (1606).37° Voy.) Jamogeat. Past. ST ligegh rer 81
fig. 3; and (¢) Juncus aquaticus geniculatus, capitulis equiseti, minor
Sloane, zd@.. Cat. (1606) 37; #2. Voy. (1707) .122 t. 75 fig. 2. [There is
ge a specimen in Linnzus’s herbarium named by Linnzus himself

s “Scirpus geniculatus 3’’, which, according to Mr. Dandy, was not
in the Linnean herbarium in 1753 ‘and so it has not status as regards
the typification of S. geniculatus: its spike is long and cylindric
and not subglobose as described by Linnzus. It has long been
referred to Heleocharis interstincta (Vahl) R. Br. (cf. C. B. Clarke,
in Journ. Linn. Soc., Lond. XXX, 1894, p. 309)].

Gardens Bulletin, S.S.

299

The types of the Sloane references are still extant, and Mr; E. J.
Dandy, of the British Museum, London, informs me that the type of

Juncus—————major pertains to the long-spiked species commonly
known as 4. geniculata, and that of /wuncus————-minor, like
that of Scirpus—————subrotunda Hort. Cliff. 21, represents a small

round-headed species, identical with WH. cafitata R. Br. (=H.
caribeaa Blake) and agreeing with the Linnean description of S.
geniculatus. Since the type of /ncus—————wmayjor, Sloane, does
not accord with the Linnean description, it also cannot be chosen for
the Lectotype of the species. This means that the current interpretation
of the species Heleocharis geniculata (L.) Rom. and Schult. 1s
erroneous, being applicable to the species that should be known as
Hf. elegans (Humb. and Kunth) Rem. and Sch.

These two exclusions leave still two more syntypes, out of which
the Lectotype of H. geniculata must be chosen. In this connection,
it must be noted that, though Sloane’s references seem to have
suggested to Linnzus the specific epithet ‘‘geniculatus’’, the Hortus
Cliffortianus supplied the bulk of the specific description that
accompanied the publication of S. geniculatus in Linn., Spec., Pl. I
(1753) 48, and since this Hortus Cliffortianus reference also alludes to
the syntype with which Linnzus was best acquainted (Linnzus having
examined and described it in the Hortus), the specimen in Clifford’s
herbarium should, therefore, be considered as the Lectotype of A.
geniculata (L.) Rem. and Schult. as emended here. Sloane’s type
of Juncus—————minor, with which Linnzus was less acquainted,
and on which the specific description was not based, becomes, thereby,
its Lecto-paratype. cf. Note on the syntype: Example 2.

5. APOTYPE :—From the definition given above it will be
seen that an Apotype is not merely a specimen that helped to
complete the description of a species; rather it is one that helped
to supply the chief diagnostic or identification clue lacking both
in the original description and the proterotypes of the species.
This definition slightly alters the meaning given in Jackson’s
Glossary where the term Apotype is defined as a ‘‘supplementary
type aiding the completion of descriptions’’—a definition that

would include also the Topotypes and Icotypes as defined here
(vide infra).

Example 1:—In this category will come any specimen that helped
first to identify definitely the species, which, like Scimdapsus pictus
Hassk., Scindapsus argyreus Engl. or Pothos pinnatus Wall., were
based on juvenile forms or sterile specimens.

Example 2:—Pinanga maculata Porte ex Lem. (1863)—a native of
the Philippines—was founded on a juvenile mottled form cultivated
in Europe. As the mottling occurs frequently in the juvenile forms
of several Pinanga palms, the species was regarded as unidentifiable.
Since the species was described from living specimens cultivated in
Europe, I undertook in Europe an inquiry into this subject and found
that the Philippine plant cultivated there as P?. maculata was identical
with P. Copelandii Becc. (Fedde, Repert. XXXV, 1034, p. 281).
The specimen that gave the most important clue to this identification
was Vilmorin’s specimen in Wendland’s herbarium. This specimen

should, therefore, be called the Apotype or Logo-Apotype of P.
maculata,

6. TOPOTYPES and ICOTYPES :—Even when the Pro-
terotype is characteristic, then similar specimens from the
Proterotype, or even from other, localities are of great value,

Vol. IX. (1937).

400

especially to the herbaria that possess only imperfect material,
or none at all, of the Proterotype collections. Moreover such
specimens may serve to supplement the details concerning either
the plant organs not represented in the original specimens, or the
full range of variation in the species. Hence the need to
distinguish Topotypes and Icotypes from Apotypes.

In Jacksons’ Glossary, Icotypes are defined as ‘‘Types
serving for identification, but not previously used in literature’’.
According to that definition, Icotype specimens, as soon as they
are cited in literature, would cease to be Icotypes and would
therefore have to be relabelled—a thing not always easy to do
when most botanical institutions do not have a staff enough to
cope with such a work. Besides Jackson’s definition would also
include the Topotypes that are not used in literature. To avoid
this I have modified the definition of Icotype to include only
those specimens that agree with the original type though not
from type localities, and irrespective of the fact that they are or
are not cited in literature.

Note:—According to the status of the locality from which
the Proterotype was obtained, the prefix Topo can be added to
define the newer collections more accurately; e.g. Topoholotype,
Topoparatype, Topolectotype, Topoapotype, etc. Similarly the
prefix Ico will indicate the status of the type specimen with
Which the Icotype has been compared: e.g. Icoholotype,
Icoparatype, Iconeotype, etc.

Example:—In the case of Poa amboinica L. emend. Retzius,
discussed under Lectotype (Example 3), Mr. Fischer mentions, besides
the specimen (the Neotype or Lectotype) in Retzius’s herbarium in
Lund, two other specimens of Evagrostis amboinica [amboinensis] (L.}
Trin., that are preserved in the Smithian and the Natural History
Museum herbaria in London. One of these specimens was collected
from the Lectotype locality in South India, while the other was grown
in England from the seeds collected from the same locality. Both
these specimens will have to be called Topolectotype, or Topo-
neotype.

7. NEOTYPE :—

Example 1:—Scirpus glaucescens Willd. (the basinym of Heleo-
charis glaucescens Schultes) was described as having had compressed
stems and trifid styles, but the only specimen with that name in
Willdenow’s herbarium has been identified by specialists as Heleocharis
palustris (L.) R. and S., a species with non-compressed stems and
bifid styles. Now should the specimen in Willdenow’s herbarium be
considered the holotype of AH. glaucescens (Willd.) Schult? If that
specimen has been correctly identified as H. palustris, it cannot be
H. glaucescens (ex descriptione), and so one may presume that the
holotype of the latter has been lost or mislaid; but, if it really has
compressed stems and trifid styles, it can be taken as the holotype
of H. glaucescens. As the type plant of H. glaucescens was growing
in the Berlin Botanic Gardens, it is possible, that Willdenow, the
director of the institution, did not preserve the actual specimen which
he dissected and from which he drew his specific diagnosis, and that
later, in order to fill up the gap in the herbarium, he took a specimen
of what appeared to him the type plant or a plant identical with it.

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301

Dr. O. A. Farwell (Rhodora, XXXII, 1930) shows that, excepting
perhaps the one phrase glauco-virescens, Willdenow’s description is
quite applicable to AH. acuminata (Muhl) Nees and that ‘“‘his
characters of three styles and compressed stems clinch the identity.
No other species of the genus known to me so well fills the bill;
certainly not the plants in America masquerading as H. glaucescens,
for they have two stigmas and the stems are not compressed.”’
If Farwell’s identification of the species is correct, then the Holotype
of H. acuminata becomes the Neotype of H. glaucescens (Willd.)
Schultes. [But whatever may be the status of Willdenow’s specimen
(Pseudotype or Holotype), it is evident that Willdenow’s description
does not apply to H. palustris (L.) R. and S., and so Willdenow’s
species (ex descriptione) cannot be regarded as a synonym of the
latter] (see also the Example under Merotype and Proteromerotype).

Example 2:—cf. Example 3 under Lectotype.

8. IDEOTYPE :—According to Jackson’s Glossary, an
Ideotype is ‘‘a specimen identified by the describer, but not
from the original locality’’, while a specimen identified by the
describer from the original locality is called a Metatype (-Meta =
with; after). But since a heterotype chosen or identified by an
author may fall in any of the several heterotype categories, I
modify this definition so that the prefix Ideo shall denote a
heterotype identified or chosen by the author of species. Thus
there could be Ideolectotype, Ideotopotype (Metatype), Ideo-
proteromerotype, Ideoneotype, Ideoicotype, etc.

Examples:—cf. Example z under Lectotype and Examfle r under
Merotype and Proteromerotype.

9. MEROTYPE and PROTEROMEROTYPE :—For
taxonomical purposes, the plant that supplied a type specimen
of the species, is known as the type-plant (vide infra, on the
labelling of Type-plants); and any specimen of this plant
collected before or after the Proterotype-collection, is known as
a Merotype. According to the status of the proterotype or
secondary type derived from the plant, the Merotype can be
more precisely described by such combinations as Meroholotype,
Merolectotype, Mero-apotype, Meroneoholotype, etc. The
definition of this term in Jackson’s Glossary is not the same
as that given by Dr. W. T. Swingle, the originator of the term,
but it is applicable to Swingle’s Clonotype—a term not included
by Jackson.

If none of the original specimens studied by the author
of a species was preserved, no proterotypes can _ exist.
But a herbarium specimen from the type-plant, if collected
subsequently by the author or someone else, would naturally fall
into the Merotype category. In order, however, to indicate that
no Proterotypes of the species exist, I distinguish as a Protero-
merotype the Merotype first proposed as the standard of the
species.

As, however, neither a Merotype, nor a Proteromerotype,
can have entered into the original description of a species, they
cannot be considered as Holotypes, Paratypes or Syntypes. For
instance, take the orchids or the herbs which have been described

Vol. IX.. (1937).

302

from living plants without preservation of herbarium material.
Anyone, even the author of the species, who wished subsequently
to take herbarium material from the type-plant, might have been
misled into taking a specimen from the wrong plant, though
somewhat similar to the one described. Now should a subsequent
worker notice discrepancies between the herbarium material and
the original description, the prefix Mero or Proteromero would
at once put the worker on guard against the specimen. Besides,
during the interval that must elapse between the taking of the
original material and the Merotype, the type-plant itself may
have grown more fully and have exhibited certain characteristics
that were not visible in the original material and which, there-
fore, were not noted in the original description. In such a case,
too, the prefixes, Mero and Proteromero, might prevent erroneous
interpretation of the specific description and indicate the extent
of the variability in the plant (vide also under Clastotype; and
also the example under Isotype).

Example:—In the case of Scirpus glaucescens Willd., the basinym
of Heleocharis glaucescens Schultes, mentioned in Example zs under
Neotype, it has to be noted that the species was based on a plant
growing in the Berlin Botanical Gardens. Presumably Willdenow
drew his specific description from a living specimen which he did
not preserve, and later he took, for his herbarium, material from
a wrong plant mistaking it for the type-plant. If this surmise is
correct, this second specimen should have been labelled as the
Proteromerotype, or more precisely Ideoproteromerotype—a name that,
had it been used by Willdenow himself, would have saved many
controversies over the exact status of the species. For, as pointed
out in the previous discussion of the case, (cf. under Neotype) the
specimen that in Willdenow’s herbarium ‘bears Willdenow’s specific
name has been identified with a species (Heleocharis palustris) that
has neither compressed stems nor trifid styles, characters which
Willdenow expressly mentioned in the specific description of his
S. glaucescens..

10. ISOTYPE :—A botanist, working critically on rare or
taxonomically difficult plants, will naturally prefer to give all
specimens taken simultaneously from one individual plant one
field number and to the specimens taken in a similar manner
from another plant another field number; even though both plants
are of the same species and grow side by side, and both collections
are taken on the same day and at the same hour. Now should
a specimen from any of these collections become a Holotype,
Lectotype, Apotype, Neotype, etc. respectively, the remaining
specimens from the same collection will be of special value in
any subsequent critical study of the species or to the other
herbaria not possessing any of the original types, since they were
all collected at one and the same time and from one and the
same type-plant. Such duplicates are called by Swingle
‘“‘Synchronous Merotypes’’, but I propose instead to restrict to
them the use of the word Isotype and to call them respectively
Isotype, Isoapotype, Isoneotype, etc. as the case may be. The
term Isotype, although originally proposed by Wilmott in the

Gardens Bulletin, S.S.

303

sense of Swingle’s Merotype, has recently come to be used rather
indifferently to include all duplicates herein distinguished as
Merotypes, Isotypes, Haptotypes and Clonotypes. (vide also
under Clastotype).

Example:—In order to clarify the case of Merotypes and Isotypes
further, I would refer to the system organised by Dr. F. H. Endert
of the Boschwezen (Forest Department) in the Netherlands Indies,
in order to secure adequate herbarium material of precisely such large
forest trees as are likely to be omitted by ordinary ‘extensive’
collectors. A forest official resident in a district selects and numbers
a tree which is apparently typical of a species in a division of his
district. In the flowering season the tree is climbed and enough
material for 5 herbarium specimens is collected. Later, when the
fruit is formed, the tree is felled and from it five fruiting specimens
together with timber sections are taken. Now the question arises:
Should both the flowering and fruiting specimens receive the same
field number? Though both collections must receive the same free
number, they should certainly not receive the same fe/d number since
they were not collected simultaneously, or, if given the same field
number, then a letter or other sign should be added to distinguish
them. Thus if flowering specimens bear the field No. 165, the fruiting
specimens could be given the field No. 165-A.

But whether these two collections are given two different field
numbers or the same tree number, or both, care should be taken to
indicate that, in each collection, the duplicates are of the Iso-Class,
while the fruiting specimens are Mero-class duplicates of the
flowering specimens so that, should any of the specimens become,
say the Holotype, Neotype, Apotype, or a Syntype of a species, the
institutions in possession of the duplicates of the type will know
exactly the relation existing between the type and the duplicates:
that is, if a flowering specimen from this collection were to become
say the Holotype of a species, all the other specimens from the same
collection will be Isoholotypes and the specimens in fruit Meroholo-
types; or vice versa, if a specimen in fruit become the Holotype, all
the duplicates in fruits become Isoholotypes and the specimens in
flowers Meroholotypes. (see also note under Haptotypes below).

11. HAPTOTYPES :—Collectors generally, but specially
those who require to make many duplicates, cannot give as much
care in preparing their duplicates as can a botanist making his
Isotypes. Collectors often give the same field number to
specimens collected in the same locality from a number of
different plants, that is to say, from plants that appear to the
collector to be conspecific, but which may not be. Now it has
already been pointed out in the introductory discussion of the
Varieties of Types that in some cases, as of certain palms, herbs,
orchids, grasses, or fungi, such procedure may be the only means
to secure specimens in sufficient quantities to make many
duplicates and, at the same time, to prevent the numbering in
the field from becoming excessive and valueless. In these
circumstances it should not be surprising if some duplicates
belong to another species than the Proterotype. When, therefore,
there is no means of ascertaining whether the specimens bearing
one and the same number were derived from the same individual
plant (or clump in the case of tufted plants), the duplicates
should be regarded with suspicion and should never be called

Vol. IX. (1937

304

Isotypes. Therefore I propose the term Haptotype for such
duplicates. Only a Haptotype certainly collected simultaneously
from the type-plant should be called an Isotype.

Note:—Even when all the specimens of a collection have
been derived simultaneously from a single clump or tuft, it is
advisable to look upon the duplicates with suspicion unless such
duplicates have been prepared carefully by a compétent botanist.
i make this note after having noticed several discrepancies in the
case of duplicates of palm specimens derived from the plants
cultivated in the Botanic Gardens, Buitenzorg, Java. There the
authorities in charge of the gardens aim to keep the numbering
of the cultivated plants as accurate as possible; yet despite their
care they are not able to prevent the seeds of one species from
growing in clumps of another closely related species. In
consequence one may sometimes see a clump labelled as of one
species containing stems of another closely related species that

~ has grown there from self-sown seeds.

Example 1:—In Flora Capensis V, Section 1 (1910) 246, N. E.
Brown published the new species Orthosiphon Thorncrofti N. E. Br.
with Thorncroft’s specimen in Kew herbarium numbered 3123 as a
syntype. Dr. Maurice Ashby, who has made this species the type
(=basinym) of his new combination, Hemizygia Thorncrofti (N. E.
Br.) Ashby, has shown that a specimen bearing this number in the
herbarium of the Transvaal Museum at Pretoria does not correspond
with the syntype in the Kew herbarium but represents Hemizygia
PRE RAL CRESES (Schlecht.), Ashby (Journ. Bot. LX XIII, 1935, p. 340
adnot.).

Example 2:—Ridley gave the number 5123 to some Demonorops
specimens collected from more than one clump, no doubt in the
belief that these clumps were all of the same species. Subsequently
he submitted one specimen to Beccari, who determined it as VD.
angustifoliuns Mart. Accordingly, Ridley entered this name on all the
remaining duplicates of the MHapto-class numbered 5123 in the
Singapore herbarium. It now appears that most of these are D.
grandis Mart. Presumably, Ridley regarded all these duplicates as
equivalent like the duplicates of Iso-class, and thus was led to
misinterpret Y. angustifolius Mart. in both his Materials II (1907)
and his Flora V (1925).

12. CLONOTYVPE aad SPERMOTYPE -—Whesm =. tpe
collection is small and when it is difficult to transplant the type
plant to a garden, every opportunity should be taken to propagate
the type-plant vegetatively or by seeds and to take duplicates
from its progeny. This practice is not uncommon in many
botanic gardens, though the specimens thus derived rarely bear
precise labels. To be accurate a specimen from such an offspring
should be labelled either a Clonotype, if the offspring was raised
vegetatively, or a Spermotype if it was raised from seed or spore.
These terms, Clonotype and Spermotype, are useful also because
they may enable one to decide with herbarium material whether
or not a variation from the type is hereditary in the species.

U3e CLASTOTYPE :—Taxonomically considered, the only
equivalent duplicates are Clastotypes and Isotypes. No other

Gardens Bulletin, S.S.

395

duplicates can be regarded as taxonomically equivalent since
they were not certainly gathered at the same time as the original
type and from the same type-plant. Merotypes and Clonotypes,
although they stand next in importance as duplicates, may have
the disadvantage of showing marked variations from the
Proterotypes and Isotypes or they may have been taken from
a wrong plant owing to a mistake in the identity of the type-
plant. Particular care should be taken, therefore, to prevent
such important duplicates as the Clastotypes, Isotypes, Mero-
types, and Clonotypes becoming mixed during drying, transit,
mounting, etc. To prevent mixing, numbered tags should be
attached to the larger parts of specimens, and numbered capsules
should include seeds, fruits, flowers, and- other small
components.

4. The Labelling of Type Plants in Gardens

Botanical gardens often have type-plants or their offspring
in cultivation. It would be useful if these type-plants were
correctly labelled in the garden and the particulars of their history
and place were kept filed in the office records. If this course
were followed, the importance of the plants would become at
once apparent both to the curators and visiting botanists.
Besides, if the field labels of a type-plant were lost, the record
file would enable one to locate the plant.

This, of course, is merely a suggestion. Each institution
has its own methods, adapted to its particular requirements.
The suggestion is put forward merely to indicate one way at least
whereby type-plants growing in a garden might be made more
useful to systematists and less liable to be forgotten. As an
instance, I may point out that the seed progeny of type-plants
of Livistona Woodfordti, Ptychosperma Sanderianum and
Borassus Machadonis are presumably all growing in the Botanic
Gardens, Singapore, but owing to inadequacy of labels and
records it is extremely difficult to identify with any degree of
certainty the plants that were raised from the seed directly
obtained from the Holotype-plant.

As to the terminology of the type-plants one can adopt with
advantage the one proposed above for labelling the type-
specimens. The term Holotype, Paratype, Syntype, Lectotype,
Neotype, Apotype or Ideotype, when applied to a type plant,
would show the nature of the type specimen which this type-
plant had supplied. Furthermore, a plant collected together
with the holotype-specimen, and which is believed to be of the
same species as the holotype, might be called a Haptotype plant,
or more precisely a Haptoholotype-plant. The offspring of a
type-plant, would also be called by the designation of their
respective parents with the added prefix Spermo or Clono,
according to the derivation of the offspring by sexual or asexual
propagation. Finally, by adding the number of the propagation,
as I, 2, 3, etc., the exact generation of any one of the progeny

Vol. DX Cro 37).

306

would be ascertainable at a glance. Thus ‘‘Spermoholotype 3’’
would mean that the plant so labelled was of the third sexual
generation from the holotype plant.

5. The Labelling of Type Specimens in the Herbarium

In conclusion, I would remark on the labelling of type-
specimens in the herbarium. Several botanists seem to favour
strongly a tiny label with a printed inscription to indicate each
kind of type, while others go further and recommend special
colours for the slips, or special coloured ink for the inscriptions
(one colour for each kind of type). In fact these different kinds
of printed slips are many times found included with the
unmounted duplicates of types (antitypes) sent in exchange from
other herbaria.

To my mind these tiny slips are often nothing but a great
nuisance. They are easily dislodged not only during the transit
of unmounted specimens but also later when the specimens are
being consulted in the herbarium, poisoned or mended. Should
any of the slips fall out of a bundle, what clue is there to its
right place? Should any be misplaced in the herbarium, what
indication is there even for a botanist to detect the error? And
one has to bear in mind that, while these slips are intended to
aid the trained botanists, all the work of mounting, poisoning,
mending, etc. may be entirely in the hands of persons who
scarcely comprehend “‘all the fuss that botanists make over wee
bits of printed paper’’, which can be had in ample quantities
in the stores, and which they can hardly distinguish from the
ones sent by other herbaria to indicate that a specimen has
already been poisoned. Needless to say, the work of mounting
and mending herbarium specimens should be rendered as simple
as possible, so that, in the event of labels falling out of a packet,
the right place could be found with little difficulty even by the
personnel of the mounting department; or when, as sometimes
happens, a label is pasted on a wrong sheet, the mistake may be
recognised and corrected by anybody consulting the specimens.
But reliance on printed slips, coloured or not, only introduces
into such instances a most discouraging confusion; and possibly
a confusion that not even the botanist himself would find it
easy to straighten out, while in some instances an initial error
would most certainly go undetected unless the botanist
consulting the specimen were himself to undertake a revision of
the species paying particular attention to its history and that of
its types.

On these grounds alone there is in my opinion a serious
objection to the u&Se of tiny printed slips. Instead, the terms
Holotype, Paratype, Lectotype, etc. should be written by hand
on the specimen sheet itself or on the label that bears the name
and number of the specimen. But there are other grounds why
the simple and direct method of writing down by hand should
be encouraged, when recording the status of a specimen in a

Gardens Bulletin, S.S.

307

herbarium. For one thing, writing is easier and quicker than
searching for the appropriate slip, which is a labour likely to
discourage botanists from recording on the specimen sheet any
discovery they may make regarding its relation to a published
species or plate, or even the fact that it was compared with any
of the existing types. For another thing, these tiny printed
slips have no place for date or even the initials of the person
who made the comparison or study; and in a large number of
cases, as with Lectotypes, Apotypes, Neotypes, etc., for instance,
it is important to know who made the selection of the Lectotype
from the Syntypes, or who compared a herbarium specimen with
the Holotype, etc. But if the method of writing directly by
hand is adopted and even if the writer has omitted his initials,
it is often easy to recognise the handwriting. On the other hand,
the bare printed slip may have been put by anyone without any
expert observation through a mistake: it carries on the face of it
no personal guarantee. Finally, how obscure might have been
the history of many a classical type-specimen in the herbaria of
Linnaeus, Lamarck, Jussieu, Banks, Delessert, Retzius, Wallich,
Griffith and the rest and how different the interpretation of the
species now ascertained to have been based on those specimens,
were it not for the handwriting of the persons who studied the
types? .

No doubt to preclude entries directly on the herbarium sheets
by incompetent persons, there is in force in many herbaria a
regulation that all annotations should be made on separate slips
of paper or determination-labels to be pasted or pinned after-
wards on the sheet. These labels could obviously be used also
to indicate the nature of the types. As for the unmounted
duplicates to be distributed, the information about their nature
could be entered directly on each label that accompanies the
duplicate giving particulars about its collection number, collector,
locality, date, etc., though even in these cases the above
mentioned determination-labels would also be far preferable to
the tiny printed slips.

When recording particulars of the nature of the type on
slips or determination-labels, it is advisable to head the label with
the collector’s name and the collection number (or, if without
number, the locality and date) followed immediately by the
name of the species, e.g.

Koenig, South India

Poa amboinica L. emend. : : : :
in Retzius’ Herbarium in

Ts Retz. ca
Neolectotype. sec. Retz. :
initials :
Vilmorin, from cultivated
specimens

Lem. barium in Berlin.
Logo-apotype sec. Furtado

Pinanga maculata Porte aa’ in Wendland’s _ Her-
|
gals: Cex. ¥ . 15-11-33. ||

Vet. EXC 01837).

308

Ridley, gogo
Areca bongayensis Furtado sp.

ipa in British Museum

Haptoholotype Herbarium.

initials: C.X.F. 15-2-34.
Ridley, on Matang, in July,
1903
Calamus conjugatus Furt. sp. in Herb. Singapore.
nov.
Holotype.
Wallich 4441
{ Raphidophora peepla (Roxb.)
| Schott.
Neotopotype. sec. Furtado
Wallich, 4441 partim
{ Rhaphidophora peepla (Roxb.)
Schott.
Hapto-Neotopotype or
*Topotype sec. Furtado
C.X.F. 18-2-34.
These entries will serve the double purpose of guiding the
personnel of the mounting department in finding the correct
sheet for a label, should it have become detached and fallen
out of the specimen cover; and they will enable the systematist
to detect mistakes at once should labels have been glued on
wrong sheets.

in British Museum
Herbarium.

in Herb. Kew.

_—_—_~_-————

REFERENCES

1. CHASE, A.—Some Causes of Confusion in Plant Names—
Journ. For. XVII (1919) 159—162.

2. JACKSON, B. D.—A Glossary of Botanic Terms with their
Derivation and Accent. London, 1928 (4th ed.).

3. MERRILL, E. D.—Type, Cotype and Topotype Labels—T7orreya
AVIL" (2677). Fs:

4. SWINGLE, W. T.—Merotypes as a Means of Multiplying
Botanical Types—/ourn. Wash. Acad. Sct. II (1912) 220.

. -SWINGLE, W.. T..—Glastotypes Clonotypes and Spermotypes,
Means for Multiplying Botanical Type Specimens—
Op. cit II (1912) 344.

Un

Gardens Bulletin, S.S.

309

INDEX

New terms in clarendon type

Amomum terminale 290
Amylotheca amplexicaulis 203
Amylotheca saccata ~ 293
Andropogon amboinicus 207
Antitypes 289
Apotype 288, 299
Borassus Machadonis 305
Clastotype 288, 304
Clinostigma :
samoense 291
Warburgii 291
Clonotype 288, 304
Cotype 293
Cyphokentia samoensis 291
Demonorops:
angustifolius 304
confusus 297
grandis 304
propinquus 207
pseudomirabilis 2&9, 299
Duplicate types
Eleocharis, vide Heleocharis _
EnpertT,: Dr. FF. H: | Intensive
system of Collecting plants 303
Eragrostis :
amboinensis 298, 300
amboinica 298, 300
riparia 298
Haptotype 288, 303
Heleocharis :
acuminata 301
capitata R. Br. 293, 299
caribea 294, 290
elegans 294, 299
filiformis 204
peuscetata./(L.) KR. ~and -S:
emend. 294, 208
glaucescens 300, 302
interstincta 298
ovata 204
palustris 300, 302
tenuis 204
Hemizygia:
Thorncroftii 304
transvaalensis 304
Heterotypes 288
Holotype 287, 290
Icotype 288, 209
Ideolectotype 207
Ideotype 288, 301
Idiotype 288
Isotype 289, 302

Vol. IX. (1937).

Ixora affinis var. arguta Craib 294
I. arguta R. Br. ex King 204

I. nigricans R. Br. 292
I. nigricans var. arguta Hook f.
204

Labelling of Type-plants 305
Labelling of Type-specimens 306
Labels for Type-specimens 306
Lectoholotype 288, 296
Lectoparatype 288
Lectotype 288, 296
Livistona :

Alfredii 290

Mariz 290

W oodfordii 305
Logo-apotype 288, 290
Loranthus cordilimbus 293
Lysichitum americanum 296
Merotype 288, 301
Merotype, Synchronous 302
Metatype 301
Neotype 288
Orthosiphon Thorncroftii 304
Paratype 287, 290
Phoenix amboinica 297
Pinanga maculata 299
P. Copelandii 209
Poa:

amboinica 297, ,300

riparia 2908
Pothos pinnatus 299
Primary Types 287, 289
Proteromerotype 288, 301
Proterotypes 287, 289
Ptychosperma Sanderianum 305
Scindapsus :

argyreus 209

pictus 290, 201)
Scirpus:

capitatus 203

geniculatus 290, 208

glaucescens 300, 302

Specimens, how to number them in

the field 287, 303
Spermotype 288, 304
Syntype 288, 203
Topotype 288 299
Type-plants, labelling of 305

Type-specimens and their labels 306

Type-specimen, definition of 287
Type-specimens, labelling of 306
Types, varieties of 286

310

ASTERISKS IN LINNAEUS’S SPECIES PLANTARUM

By C. X. Trieeapo,

Botanic Gardens, Singapore

Page

1. Introduction and Conclusions by ae ii, ae
2. Notations in the Species Plantarum : Ve ban Gk
(1) Doubtful Taxonomic Groups st ve eee

(ii) Imperfectly Investigated Taxonomic Groups ... 312

(111) Synonyms of Doubtful Identity re Liste Re

(iv) Doubtfully Separable Taxonomic Groups <a ee

(v) Species Containing Asterisked References 2 Ba

1. Introduction and Conclusions
In the Journal of Botany, xxv, 1637, p: 78, Mro Tl A.

Spracug and Mr. A. W. Exen. have conjointly published a
note drawing the attention of botanists to the significance of
asterisks in LINNAEUS’S Species Plantarum ed. 1 (1753).
They have shown that the use of asterisks was explained by
LINNAEUS himself in the following paragraph in the preface of
the second edition of his work:

“DESCRIPTIONS * tantum in obscuris adhibere necessem fuit,
easque sine ambagibus, ut obtinerem compendium tironibus gratum’’.

This paragraph, but without the asterisk, is the verbatim reprint
from the first edition of LmnNAEuUS’s Species Plantarum.
Consequently Mr. SprRaGuE and Mr. EXELL assume that the
asterisk was accidentally omitted after the word DESCRIP-
TIONES in the first edition of the work. They translate the
paragraph as follows:

‘‘Only in doubtful cases was it necessary to cite descriptions, and

those, straightforward ones, so that I might keep the handbook
suitable for beginners.’’

While agreeing with the explanation of the omission of the
asterisk from the first edition of the Species Plantarum, I find
myself compelled to dispute this translation and, therefore, to
reconsider the conclusion of these authors on the significance
of the asterisk. The translation appears to me misleading,
because it is too free and is removed from its context, and
is therefore decidedly ambiguous. LINNAEUS cited descrip-
tions for nearly every species, but no one imagines that
he considered them all doubtful. Is ‘‘compendium’’ really a
‘‘handbook’’?? And what can be meant by the indefinite phrase
‘‘doubtful cases’? Are we to suppose that LINNAEUS was

Gardens Bulletin, S.S.

Sl) 6h hCUs lL eee

CO ———————

4

3a1

doubtful of every species in the synonymy of which he employed
an asterisk, or that he was merely doubtful of the unasterisked
citations under it; or that he considered the citations with an
asterisk to some extent doubtful though supplying a straight-
forward description, while those without an asterisk were not
at all doubtful? It is clear that the translation of the phrase
‘‘in obscuris’’ and the interpretation of the asterisk may be
matters of considerable taxonomic importance, for on them will
depend the procedure for interpreting a Linnean species with an
asterisked citation.

In view of this uncertainty I have undertaken an inquiry
into the problem. My conclusions are these:

1. The paragraph should be read:

‘Only in the case of obscure plants was it necessary to
indicate with an asterisk such descriptions as were straight-
forward, so that I might keep the compendium suitable for
beginners.”’

Note:—(a) ‘‘obscure’’ does not mean doubtful (see section 2,
subsection V, p. 314). (b) ‘‘Compendium’’ means
the list of synonyms or citations, i.e. the synopsis
of the species under consideration (see section 2,
subsection V, p. 315).

z. LINNAZUS did not mean to invest the asterisks with
the force of indicators for discriminating between the synonyms
(citations). The asterisks were meant as a guide for beginners,
and all synonyms, whether with or without an asterisk, were
equally important for the experienced botanist.

The evidence for my conclusions is set forth below.

2. Notations in the Species Plantarum

In order to view the problem of asterisks in their proper
light, I have investigated the meaning of several other notations
used by Linnaeus for similar purposes. I shall therefore divide
the problem under the following heads :—(i) Doubtful Taxonomic
Groups; (ii) Imperfectly Investigated Taxonomic Groups; (iii)
Synonyms of Doubtful Identity; (iv) Doubtfully Separable
Taxonomic Groups; and (v) Species Containing Asterisked
References.

Except when otherwise stated, all paragraphs and examples
quoted below are from LINNAEUS’sS Species Plantarum ed. 1
(1753), and the quotations without page references are from the
preface of the same work.

Shoe eT vax ONO MILE (GRD UPs

LINNAEUS, in his preface to the Species Plantarum, informed
the reader that, having himself been many times deceived by
the specific descriptions given by others, he had left out of his
book all species of which he could not examine actual specimens
or drawings, so that what was uncertain might not get mixed

Vol. EX. (1937).

312

with what was known for certain. The sentence in question
runs thus:

“NON VISAS plantas heic omisi, toties elusus ab auctoribus,

ne dubia certissimis miscerem”’,
It is obvious that LINNAEUS meant by the word plantas not only
specimens but also drawings for it is a well-known fact that
LINNAEUS based several species merely on drawings. Nor is
this contradicted by the fact that his previous experience in the
matter of published descriptions, unsupported by drawings or
specimens, led him to adopt CAESALPIN’S dictum :

‘“‘Ignoto Genere proprio nulla descriptio, guts accurata tradita,
certam demonstrat, sed plerumque fallit.”’
which motto LinnaEuS placed on the reverse of the title-page
of his book. Nor is it contradicted by the further fact that,
despite all his efforts to keep out species of which he could not
examine drawings or specimens, there are evidences to show
that he has not been able to ignore entirely the descriptions of
his contemporaries and friends. ‘Thus, for instance, he published
an unnamed Trientalis (i.e. without a trivial name); but here he
was careful to mark the reference with an obelisk (dagger) and
make the following observation :

‘“Plantam hanc non vidi, ex auctoritate Clariss. D. Royeni
adposui.’’ (p. 344).

It is evident therefore that by the phrase ‘‘in obscuris’’
LINNAEUS did not mean dubious species (plante dubie), i.e.
those that were not known to him from specimens or reliable
drawings.

nu. PePEerer FECT ye Invweeritc..ae
TAXONOMIC GRo UF's

It is understandable that, in such a comprehensive work,
LINNAEUS had not sufficient opportunities or material to
investigate fully every one of the species which he described.
He therefore indicated insufficiently investigated species with an
obelisk so that others might examine them in more detail. The
use of obelisks is explained in the following sentence :

“‘si vero aliquando contigerit non sufficienter inspexisse plantam,
vel specimen imperfectum obtinuisse, signo+hoc notavi, ut alii
eandem accuratius examinent.”’

The dagger is employed perhaps more with the purpose of
directing botanists to a further inquiry into species than with
the purpose of indicating his own imperfect inquiry. Thus the
reference cited under the unnamed Trientalis sp. is, as noticed
above, daggered, though LinNAEUS himself had not had any
opportunity of seeing the plant. An unnamed Phyllis sp. (p. 232)
is daggered and contains the following remarks: ‘‘Flores hujus
mihi ignoti sunt’’, while the following species, although not

Gardens Bulletin, S.S.

313

daggered, have attached to them remarks which show that
LINNAEUS had imperfect specimens for examination :

(a) Cassia javanica p. 239: ‘‘Flores mihi non protulit’’.

(b) Laurus Winterana p. 371: ‘‘Flores ipse non vidi’’’.

(c) Eranthemum capense p. 9: This is not daggered neither
in the first edition nor in the second edition of Species Plantarum,
but the second edition contains the following note: ‘‘Habitus
Chironie, unicum specimen minus perfectum vidi, ideoque
accuratius examinanda.’”’

The cases are very few where the species containing
asterisked references also contain remarks to show the species
were imperfectly investigated, and so far I have come across
only one species where both an obelisk and an asterisk occur
(i.e. Ammi glaucifolium p. 243). |

I conclude from this that the phrase ‘‘in obscuris’’ does not
refer to such cases as were imperfectly investigated by LINNAEUS.

i. SYNONYMS. OFUDOUBTFUL IFDENTIT Y

That LINNAEUS did not generally allow himself to be guided
by descriptions alone, unaccompanied by specimens or drawings,
has been explained above. It is therefore to be presumed that
he did not cite any synonyms of which he had not examined
drawings or specimens. Further it is to be presumed that
LINNAEUS did not give each and every synonym, even when its
type specimen or drawing was available for inspection. On the
title-page of Species Plantarum, LINNAEUS speaks of Synonymis
Selectis and, in the following paragraph of the preface, he informs
readers, of the reasons that led him to give more synonyms to
non-European plants than to the European plants:

“SYNONYMA paucissima in EUROP£ZIS plantis adhibui,

contentus C. Bauhino & Iconographo prestantiore; in EXOTICIS
vero plura, quum difficiliora minusque trita sint.’’
But despite this selection of synonyms, LINNAEUS had _ cases
Where he was not sure of the correctness of his determination
of them; and such synonyms he has shown with a question
mark. Such queried references have, in my opinion, to be
excluded in typifying a Linnean species.

Examples of queried references will be found under A pluda
mutica p. 82, Aristida adscensionis p. 82, and Queria canadense
D. OO.

There are, however, cases like Alpinia racemosa (1753) p. 2
and (1763) p. 2, which contain references that are queried not
in the first edition of the work, but only in the second. But
these references stand on a different footing and should be
included in typifying a Linnean species.

‘It is evident that the phrase ‘‘in obscuris’’ and asterisks do
reas pad to the synonyms of whose identity LinNAEus had still
oubts.

Viol. EX: 01637).

314

iv. DOUBTFULLY SEPARABLE
TAXONOMIC GROUPS

From time to time it happens that an author cannot decide
whether certain taxonomic groups should be separated as species,
quite distinct from others, or should be subordinated to other
groups. LinNAEUsS himself made a point of indicating under
each species or variety any such uncertainty. And that LINNAEUS
did not use the asterisks to indicate such uncertainty is obvious
because, firstly, most of the species accompanied by asterisked
references have no remarks about uncertainties; and secondly,
most of the species having such remarks bear no asterisked
references. But the conclusive evidence that ‘‘in obscuris’’ and
asterisks do not refer to this kind of uncertainty is the fact that
LiInnaEts established several monotypic genera accompanied by
asterisked references to the type species but without remarks
throwing doubts upon the status of the type species. And what
makes this evidence so strong against the view that ‘‘in obscuris”’
relates to such uncertainty on the part of LINNAEUS, is that the
work of LINNAEUS is not such as to lay him open to the charge
of inconsistency in having established a new genus without being
sure of the only species included under it.

(a) Examples of Species containing remarks but no asterisks:

1. Prunus sibirica p. 474: ‘‘An hec satis distincta a precedenti ?”’
2. Cistus pilosus p. 528: ‘He tres. @. 8. ¥. vel varietates vel
valde affines certiora determinent autopte2’’.
Asarum canadense p. 442: ‘‘An varietas przcedentis; sic
suadent Folia bina; dissuadent alia’’.
4. Hyacinthus monstrosus p. 318: ‘‘An sequentis sola varietas ?”’
5. Leontice Leontopetaloides p. 313: The diagnosis is obelisked,
and the remarks are as follows: ‘‘Habitus hujus generis, at flos

videtur satis diversus, an sufficienter pro distincto generi determinent
autopte.”’

_ 6. Sanicula canadensis p. 235: ‘‘Structura ita precedentis, ut
vix differentia, sed planta decupulo sepe omnibus partibus major.’’
7. Gentiana campestris p. 231: ‘‘Precedenti nimis affinis &

fere varietas aut hybrida.’’

(b) Examples of Species in monotypic genera containing
asterisked references:

Maranta arundinacea p. 2; Eranthemum capense p. 9; Amethystea
cerulea p. 21; Muntingia calabura p. 509; Nama zeylanica p. 226;
Heliocarpus americana p. 448; and Gethyblis afra p. 442.

Each of the last two species contains one reference, which
is to the type specimen itself. There is no diagnosis independent
of this reference, yet one finds this only reference is marked with
an asterisk. .

Vv. SPECIES CONTAINING ASTERISKED
REFERENCES
I submit that the foregoing discussion has made it amply
evident that the phrase ‘‘in obscuris’’ does not refer to any cases
when Linnaeus had doubts either about a species, or about a
description, or about a synonym.

Gardens Bulletin, S.S.

315

Looking therefore to the accepted meanings of the word
obscurus I find that they can be divided into two classes: The
first is original and objective ‘‘obscure, not manifest to the eye,
dificult for observation’’ i.e. the object itself present some
difficulties, which come in the way of one’s seeing it clearly or
of discriminating between its various parts; the second class is
figurative and subjective and refers to the difficulties experienced
in getting to know about the object or forming a definite opinion
about the status of the object. Difficulties (or obscurities) of the
first class affect the apprehension of the characters of the thing
observed, even when the observer has full opportunity to examine
it; those of the second class arise from insufficiency or
inaccessibility of material or from inability to comprehend the
characters already apprehended. Difficulties of the second class
were dealt with by LINNAEUS in the four different procedures
discussed above. I surmise therefore that LINNAEUS used the
word obscurus to express not his own doubts, but the obscurities,
intricacies, or difficulties which the species themselves presented
to an observer. LINNAEUS has used the word obscurus, or its
derivative, in such a sense in the following note under Cistus
apenninus p. 529: ‘‘Cistorum historia maxime omnium obscura
ob varietatum copiam, illustrabitur si determinent in singulis——

)

Now in order to determine what sort of objects or cases are
implicit in the phrase ‘‘in obscuris’’, it is necessary to consider
the entire paragraph with reference to the one immediately
preceding it. The two paragraphs are:

“SYNONYMA paucissima in EUROPEIS plantis adhibui,

contentus C. Bauhino & /conographo prestantiore; in EXOTICIS
vero plura, quum difficiliora minusque trita sint.

““DESCRIPTIONES * tantum in obscuris adhibui necessum fuit,
e€asque sine ambagibus, ut obtinerem compendium tironibus gratum.’’

From this context it appears that the word plantis is understood
after ‘“‘in obscuris’’, so that we get actually three consecutive
sentences where the words planta and adhibere, or their
derivatives, occur or are understood, viz. :

1. Synonyma paucissima in Europzis plantis adhibui,

2. In Exoticis [plantis] plura [synonyma adhibui]

3. Descriptiones tantum in obscuris [plantis] adhibere necessum
fuit ;
This preceding paragraph also explains the word
““compendium’’ as referring to the ‘‘synonyma’’. It can hardly
be translated as ‘‘handbook’’ because four paragraphs earlier
LinnakEvs referred to his work as ‘‘Libello’’ with a capital letter
(“‘compendium’”’ is written with a small letter). Further the
original meaning of the word ‘‘compendium”’ is ‘‘a hanging
together’’ and its immediately derived meaning is ‘‘a putting
together’’ with the idea of effecting a saving, and therefore
“‘a shortening’’. Now LINNAEUS put together the synonyms

Vol. IX. (1937).

316

under each species and effected a saving in space by not quoting
all the descriptions given by other authors. Hence I maintain
that word ‘‘compendium”’ is employed by LINNAEUS to mean
rather the ‘‘compendium synonymorum’’ (putting together the
previously described species in a synoptic form) than a shortening
of descriptions or of his book (as Mr. SpraGueE and Mr. EXELL
imply), of which LINNAEUS did not speak. But whether ‘‘com-
pendium’’ is translated as I suggest here or whether it is
translated as “‘handbook’’, the interpretation of the significance
of asterisks and of the signification of the phrase ‘‘in obscuris’”’
remains unaffected.

In view of the foregoing reasons I suggest that the entire
paragraph containing the phrase ‘‘in obscuris’’ might be best
translated as follows:

‘“Only in the case of obscure plants was it necessary to indicate
with an asterisk such descriptions as were straightforward, so that
I might keep the compendium suitable for beginners’’.

This means that LinNAEUsS asterisked references only in the
case of species that presented difficulties for observers or
investigators, the object of asterisks being to lighten the task
of beginners by directing them primarily to straightforward
descriptions. Other references LINNAEUS left unasterisked not
because there was any doubt as to their identity or status, but
either because the plants were not obscure or not difficult to
study, or because the descriptions in cited references were thought
to be not straightforward.

From this it follows that asterisks were not intended by
LINNAEUS to indicate any distinction between diagnoses or
synonyms and descriptions, as Mr. SPRAGUE’s and Mr. EXELL’s
note seems to suggest; nor do the asterisks indicate that LINNAEUS
considered the descriptions given in Species Plantarum merely
diagnoses, or inadequate descriptions. LINNAEUS himself gave
long descriptions even when he had asterisked some references,
as can be seen from the following examples:

Cerbera manghas p. 208; Leontice thalictroides p. 312; Vaccinium
frondosum p. 351; Origanum syriacum p. 590; Gerardia purpurea
p. 610; Malva tomentosa p. 687; Trifolium tomentosum p. 771;
Hyoseris virginica p. 809; Carduus helenioides p. 825; and Cacalia
suaveolens p. 835.

The majority of references asterisked are from LINNAEUS’S
previous works, but other references also have been asterisked,
even to RHEEDE’S Hortus Malabaricus (e.g. Curcuma rotunda
p. 2), though descriptions from this last work are usually regarded
as ‘‘obscure’’ by modern botanists. Further if two or more
references are found to contain straightforward descriptions of
one and the same synonym LINNAEUS asterisked both the
references (e.g. Eranthemum capense p. 9, Euphorbia Cyparissias
p. 461). There are also instances where references that are not
asterisked in the first edition of Species Plantarum, are marked
with asterisks in the second edition (e.g. Justicia adhatoda and

Gardens Bulletin, S.S.

317

J. ecbolium, ed. 1 p. 15 and ed. 2 p. 20). The probable reason
for this is that the marks were inadvertently omitted in the first
edition of the work, or that LINNAEUS did not realize at the time
either how ‘‘obscure’’ was each of the species, or how straight-
forward were the descriptions.

This view of the asterisks in LINNAEUS’S Species Plantarum
makes the interpretation of asterisked references clear, not only
in cases where they are the only references given under a genus
or species (references which may be considered as the sole basis
of the genus or species) and in cases where a lengthy description
is given as well, but also in the case of Ammi glaucifolium
p. 243, which has a reference marked with both an obelisk and
an asterisk. Similarly this view enables us to understand why
under Valeriana tripteris p. 32, not only the reference to Bauh.
Prodr. p. 86 (specially noted by Mr. SpraGugk and Mr. EXE 1),
but even other references given under the same species, are not
asterisked; or why the only reference given in Grislea secunda
D- 248 or in Allobhyllus zeyvlanicus p. 348 is asterisked, while
that in Gethylis afra p. 442 or in Heliocarpus americana p. 448
is not, though in each case the reference is to a Linnean work,
and the species is the type of a monotypic genus and is not
accompanied with a diagnosis or a description.

Vol. IX. (1937).

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CONTENTS

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THE

GARDENS’ BULLETIN

STRAITS SETTLEMENTS

Vol. IX 31st March, 1938 Part 4

KOTES ON MALAYAN DIPTEROCARPACEA—IV*

By C. F. SYMINGTON,
Forest Research Institute, Kepong

This paper deals with rather a motley collection of species.
In the first place I have tackled certain problems that have arisen
in the course of preliminary botanical study in anticipation of
wood-technological work on Dipterocarpus and Hopea from the
Malay Peninsula. Thanks to the good work of Dr. van Slooten,
the botany of Dipterocarpus is in reasonably good order, and
with the notes here supplied on D. gracilis Bl. and D. chartaceus
Sym. there is not much more essential critical work to be done
on our species at this stage. With Hopea we are less fortunate.
In this paper I have tackled one of the major problems connected
with H. Pierrei and other species in the section Dryobalanoides,
but there is yet much to be done in this rather difficult section
of the genus.

Next I have given a note on the Richetia group of Shorea,
the object of which is to bring together a group of species of
obvious natural affinity. This involves the transfer of several
species of Balanocarpus to Shorea, with concommitant name
changes. I regret the necessity for these changes, but it is a
necessity, and my apologies should rather be for not having
made the changes sooner.

A problem that is becoming more evident and pressing with
increasing knowledge is that of the delimitation of specific and
lesser groups. If, for example, we consider only the Species
occurring on the west side of the Malay Peninsula, we find the
specific limits are on the whole clearly defined, and the variations
within a species are insufficiently constant to suggest the presence
of definable sub-species, varieties, or other forms. Were we to
extend our investigation to the whole of the Malay Peninsula,

= For previous papers see Gard. Bull. S.S. Vol. DRIES) DP. E207 5
{1933) and Vol. VIII, pp. 1-40 (1934) and 265-292 (1935). dali

319

320

things would not be quite so simple and, were the whole possible
range of distribution of our dipterocarps considered, the
delimitation of species would become a very acute problem.
For example, there are several dipterocarps in Brunei that, on
the usual morphological evidence, are clearly much more closely
related to certain Malay Peninsula species than to any other
described forms. Yet in some character or characters, possibly
vegetative ones usually considered of little importance, they do
differ distinctly and constantly. Should these Brunei trees be
considered as species distinct from their Malay Peninsula counter-
parts; as varieties, forms, or some lesser groups; or are the
differences so slight as to be unworthy of definition? These are
obviously questions which can be adequately answered only in
the light of very extensive knowledge of concepts and vafiations
of groups within the family; premature decisions are likely to
necessitate undesirable alterations in the future. Some problems
with practical issues involved, however, cannot be postponed
indefinitely, and one such problem, concerning Shorea assamica,
is tackled in this paper.

The problem of specific definition arises again in this paper
in connection with Parashorea malaanonan and its variety here
described as var. tomentella. P. malaanonan introduces another
matter of general interest, conveniently described as “‘the treat-
ment of doubtful interpretations’’. For example, I am faced
with deciding whether or not to accept Merrill’s interpretation
of Blanco’s Mocanera malaanonan. I do not consider Merrill
was justified in his interpretation, but the evidence is insufficient
to prove him wrong. In the circumstances, and because the
name Parashorea malaanonan is now well established in botanical
and forestry literature, it seems desirable to accept Merrill’s
interpretation. But the position cannot be considered satis-
factory, and it seems appropriate here to plead with those who
undertake interpretations’ of this nature to treat the mames as
nomina dubia or nomina ambigua unless the evidence is
incontrovertible.

The description of new species from Borneo, commenced in
the last paper of this series, is continued here, the following
species: being described and figured—Shorea acuminatissima,
S. xanthophylla, S. sandakanensis, and S. Smithiana.

The paper is concluded with a description of Vatica
diospyroides (one of several interesting new dipterocarps from
Dr. Kerr’s Siamese collections) and a note on two dipterocarps
collected by the Oxford University Exploration Club in Borneo
but omitted from the collection submitted to me in 1934 (vide
Gard, BallkS SS) ak.

I should like to thank the many botanists and forest officers
who have continued to give me their valued assistance and
co-operation. I am also indebted to the Director of the Bureau
of Science, Manila who afforded me the opportunity of studying

Gardens Bulletin, S.S.

=e ey)

321

the Philippine collections of dipterocarps during a visit to Manila
in December 1935, and to the Director of the Royal Botanic
Gardens, Kew who again extended to me the hospitality of the
Herbarium during my leave in 1936.

Dipterocarpus gracilis Bl., Bijdr. V, 224 (1825); V.SI. in Bull.
Jard. Bot. Buitenz. III, 8: 276 (1927). D. pilosus Roxb.
(1832) sensu Parker in Ind. For. Rec. 13: 15 (1927), non
sensu Brandis in Journ. Linn. Soc. (Bot.) 31: 28 (1895).
D. vernicifluus Blanco, Fl. Philip. ed. 2, 314 (1845). D.
Skinneri King in Journ. As. Soc. Beng. 62, pt. 2: g1 (1893);
Ridl., Fl. Mal. Penins. 1: 214 (1922); V. Sl. in lL.c. 294;
Foxw., Mal. For. Rec. 10: 70 (1932) var. hirtus Ridl. excl..
D. angustialatus Heim in Bot. Tidsskr. 25: 43 (1903);
Fischer in Kew Bull. 1926: 457.

In my endeavour to decide upon the correct name for the
tree we have known in the Malay Peninsula as Dipterocarpus
Skinneri I find it impossible to distinguish our tree from that
known in Burma, the Andamans, and Siam, as D. pilosus, and
from the tree known as D. gracilis in the Netherlands Indies
and the Philippines.* D. pilosus has been considered systemati-
cally by Parker and D. gracilis by van Slooten, both in 1927.
These authors were working simultaneously on different territorial
regions and neither has indicated that union of the two species
might be necessary, although Parker united D. Skinneri King
with his D. pilosus, and van Slooten drew attention to the
similarity of D. Skinneri King and D. gracilis Bl. Moreover,
prior to publication of Parker’s work, the identity of D. pilosus
Roxb. was, as van Slooten (l.c., p. 274) rightly points out,
entirely problematical, although Brandis and some other authors
considered it to be identical with D. Baudii Korth. Parker’s
interpretation of Roxburgh’s D. pilosus is based on field
investigation and a process of elimination. His arguments are
very reasonable, but one cannot overlook the fact that the plant
he calls D. pilosus does not entirely fit Roxburgh’s, admittedly
brief and inadequate, description. I should prefer to consider
Roxburgh’s D. pilosus as a nomen dubium, but as I am solely
concerned at present with finding a name for the Malay Peninsula
tree formerly known as D. Skinneri, a decision on this point is
immaterial. I am satisfied that our tree is specifically identical
with Parker’s D. pilosus and with D. gracilis Bl. as interpreted
by van Slooten. Blume’s name has precedence, even if D. pilosus
sensu Parker is D. pilosus Roxb., because Roxburgh’s name

was not validly - published until 1832, seven years after
D. gracilis Bl.

_ * D. vernicifiuus Blanco is the more common name in Philippine
literature, but this species was united with D. gracilis Bl. by van
Slooten in 1927.

Vol. IX. (1938).

322

/

‘Dipterocarpus chartaceus Sym. nom. nov. D. Skinneri var.
hirtus Ridl., Fl. Mal. Penins. 1: 215 (1922); V.SI. in Bull.
Jard. Bot. Buitenz. III, 8: 295 (in nota sub D. Skinneri).
D. Skinneri sensu Ridl., op. cit. 214 quoad Curtis 3734;
sensu Foxw., Mal. For. Rec. 10: 70 (1932) partim; non
D. Skinneri King.

Ridly first drew attention to this species when he segregated
from his specimens of D. Skinneri a collection from Pahang
(C.F. 3195), describing it as D. Skinneri var. hirtus. He failed,
however, to recognize Curtis 3734 from Province Wellesley as
the same form, citing it under D. Skinnevi. In 1927 van Slooten
drew attention to the fact that these two specimens were
identical, and expressed the opinion that they should not be
considered as a form of D. Skinneri King but as a distinct species.
But in 1932 Foxworthy took the view that D. Skinneri was a
variable species and included both the typical form and Ridly’s
var. hirtus under the one name.

I am returning to van Slooten’s conception of Ridley’s
D. Skinneri var. hirtus, considering it worthy of specific rank.
This species is undoubtedly very closely related to D. Skinneri
King (= D. gracilis Bl.), but in several respects it is quite
distict. The fruit wings in D. chartaceus are relatively broader
and more distinctly 3-nerved than in D. gracilis, the fruit lobes
are much smaller, the leaf is broader and has fewer nerves, and
the texture of the leaves and twigs is distinct. In D. chartaceus
the hairs are longer, sparse and deciduous, the mature leaf being
almost glabrous, shining on both surfaces, and of a brittle papery
texture: in D. gracilis the undersurface of the leaf is covered
with stellate hairs usually forming a close tomentum.

As the name Dipterocarpus hirtus has already been employed
I am renaming this species D. chartaceus; a name descriptive of
the peculiar leaf texture.

Collections examined :—

PENINSULAR SIAM: For Dept. Siam s.n. (10.5.20). Kerr
12245, 14324, 15693, 18250.
MALAY PENINSULA :
Peruis: C.F. 40828.
PROVINCE WELLESLEY: Curtis 3734.
KELANTAN: C.F. 29028.
PAHANG: C.F. 3195 (Auth. spec. of D. Skinneri var.
hirtus Ridl.), 6650, 15645, 24546, 29601, 40416, 40683, 40686,
40705.
NEGRI SEMBILAN: C.F. 24352.
JOHORE: C.F. 17089 (?)

A form of keruing, widely distributed in the Peninsula
although not abundant, has been known by the name of
D. Skinneri. The opinion is here expressed that this is the

Gardens Bulletin, S.S.

<<
EN
ae

=
Sv =
2S a =

Plate 17. Hopea Pierre: Hance.

323

Same species as that known by the name of D. pilosus in Burma
and D. gracilis in the Netherlands Indies and the Philippines.
The species are united and the oldest name D. gracilis Bl. is
adopted in place of our D. Skinneri.

Another rather rare form of keruing was described by Ridley
in 1922 as D. Skinneri var. hirtus. It is considered that this
variety is worthy of specific rank and it is renamed Dipterocarpus
chartaceus Sym.

Hopea Pierrei Hance in (Lond.) Journ. Bot. 15: 308 (1876) et
329 (1877); Brandis in Journ. Linn. Soc. (Bot.) 31: 67 (1893)
Pl. 2, fig. 10, quoad speciminum cambodiorum; Dakkus in
Bull. Jard. Bot. Buitenz. 3, Suppl. 1: 162 (1930); non Hopea
Pierrei sensu Ridi., Fl. Mal. Penins. 1: 238 (1922); non
sensu Foxw., Mal. For. Rec. 10: 133 (1932); non sensu
auct. philip. (1909-1937). ‘‘Hopea micrantha?’’ Hance in
ke: : 242, non H. micrantha Hook. f....Hancea Prerrei
Pierre, Fl. For. Cochinch. 16 (1891) t. 248 (in tabula antem
Hopea Pierrei). Plate 17.

Pierre’s description and drawing of Hcpea Pierrei are
excellent, while his remarks concerning the diagnostic characters
separating it from Hopea Griffithii Kurz, H. micrantha Hook. f.,
H. Beccariana Burck, and H. microptera Dyer (= H. Dyeri
Heim) bear evidence of careful study, and should have been
sufficient to obviate the mistakes of subsequent authors. The
tree, as I know it from the Malay Peninsula, agrees entirely
with Pierre’s description except that I have not noted any
marked development of adventitious stilt roots. This, however,
is a variable character in the section Dryobalanoides. ‘The most
useful diagnostic characters are the short petiole, the midrib
sunk on the upper surface, the numerous but faint main nerves,
the bottle-shaped ovary and stylopodium, and the glandular outer
sepals. The presence of domatia in the axils of the lower
nerves is variable.

This species was first discovered in the island of Phu Quoc
in Cambodia and tentatively identified by Hance in 1776 with
Hopea micrantha Hook. f. Shortly afterwards, however, he
named the species Hopea Pierrei. In 1891 Pierre described and
figured the species (under Hancea) in considerable detail, basing
his observations on specimens numbered 1425 in his herbarium
which are, presumably, those examined by Hance.

Subsequent misinterpretation of Hance’s species has resulted
in Hopea Pierrei being recorded from Borneo, the Philippine
Islands, and the Malay Peninsula, on false evidence. In 1895
Brandis added Borneo and Singapore to the distribution of the
species on the evidence of Bece. 2504 and 3050 from Borneo,
and a collection of Ridley’s from Singapore. A consideration
of leaf characters alone makes it quite clear that Beccari’s

Vol. IX. (1938).

324

collections are not H. Pierrei Hance. In that species the midrib
is depressed above but in the Bornean collections it is distinctly
raised. I consider that the latter are H. Dyeri Heim. I am
not sure to what specimen of Ridley’s Brandis refers when
recording H. Pierrei from Singapore, but in all probability it was
one of the two Kew sheets of H. Griffithii Kurz from that
locality.

In 1922 Ridley continued to record H. Pierrei from the
Malay Peninsula but his citations appear (it is difficult to identify
the specimens with certainty as no numbers are cited) to include
four distinct species none of which is H. Pierrei Hance. Quite
erroneously he gives H. intermedia King as a synonym of
H. Pierrei Hance.

Foxworthy, who continued to record H. Pierrei from the
Peninsula, had a different conception of the species from either
Brandis or Ridley. The specimens he cites are, with the
exception of King 3525 and Holmberg 820 (which may be
H. Dyeri Heim), all H. pedicellata (Brandis) Sym. (see p. 327).

Hopea Pierrei Hance, however, does occur in the Peninsula,
although Brandis, Ridley, and Foxworthy have all recorded it
as a result of mistaken identification. The earliest collection
from the Peninsula was a sterile specimen (C.F. 3950) from
Bentong District of Pahang collected in 1919. It is represented
only in the Kepong herbarium and until now has been un-
identified. Since 1930 several additional collections have been
made by the Forest Department, at Bukit Raka in Bentong
District of Pahang and at Ulu Gombak in Selangor. Un-
fortunately only one of these has flowers, but comparison of
this with Pierre’s specimen shows that it has all the distinctive
vegetative and floral characters of H. Pierrei Hance.

I have already stated that I consider Brandis wrong in
recording H. Pierrei Hance from Borneo. ‘The species has also
been recorded from the Philippine Islands, from 1909 onwards,
as a result of mistaken identification. I shall not here attempt
to unravel the confusion under the name of H. Pierre in the
Philippine literature but, having examined, as far as I know,
all the material concerned, I can say definitely that none is
H. Pierrei Hance. Actually I think seven different species have,
from time to time, been included under the one name. Among
these are specimens of H. malibato Foxw. and H. Foxworthyi
Elmer (Syn. = H. glutinosa Elmer), but the remainder are
probably undescribed.

Hopea Pierrei is found in cultivation in the Botanic Gardens,
Buitenzorg.

Collections examined :—

CAMBODIA: Pierre 1425 (Auth. spec. of Hopea Pierrei Hance);
Guibier No. 2; Foxworthy No. 8.

Siam: Chantaburi; Kerr 9176: N. Rabil 14.

Gardens Bulletin, S.S.

be 9 tGe
an

Hopea Beccariana Burck.

Plate 18.

325

MALAY PENINSULA :
PAHANG: C.F. 3950, 38023, 38035.
SELANGOR : ‘C.F. 23645, 36632, 372092; 37296, 37297.

JAVA: (Cult.) Hort. Bog. VII—B—s8.

Hopea Pierrei Hance is a form of merawan which was
described originally from Cambodia. Brandis, Ridley, and
Foxworthy, have recorded this species from the Malay Peninsula,
but investigation shows that they have applied the name Hopea
Pierrei to the wrong trees. It is here shown, however, that
true Hopea Pierrei does occur in the Peninsula, although its
known distribution is limited to two localities, Bukit Raka near
Bentong and Ulu Gombak in Selangor.

Hopea Beccariana Burck in Ann. Jard. Bot. Buitenz. 6: 240
(1887); Brandis in Journ. Linn. Soc. (Bot.) 31: 68 (1895);
Sym. in Gard. Bull. 8: 28 (1934) in obs. Hancea Beccariana
Pierre, Fl. For. Cochinch., t. 248 (1891) in nota sub H.
Pierrei. Hopea Nicholsoni Heim in Bull. Soc. Linn. Paris,
2: 293 (1891). Hopea intermedia King in Journ. Asiat.
Soc. Beng. 62: 126 (1893) et in Ann. Bot. Gard. Calcutta
5, pt. 2: 156, Pl. 189A (1896) pro maiore parte; Brandis in
Pewse7, Woartinn Boxw-, ‘Mal: .FKor.! Rec..10 >. 334 (1932);
Durant, Rep. Forests Brunei; 40 (1932); Burkill, Dict. Econ.
Prod. Mal. Penins.: 1190 (1935); non H. intermedia sensu
Foxw.o, Mal. Por. Rec. 3° 74 with 2. plates... (1927).
Balanocarpus ovalifolius Ridl. in Journ. F.M.S. Mus. 10:
130 (1920) pro maiore parte; non B. ovalifolius sensu Foxw.,
Mal. For. Rec. 10: 143 (1932). Hopea Pierrei sensu Ridl.,
Flor. Mal. Penin. 1: 238 (1922) partim, non H. Pierrei
Hance et sensu auct. Plate 18.

This species is adequately described under H. intermedia
by Foxworthy (Mal. For. Rec. 10: 134). I would comment,
however, that the stamens are definitely 15 in number and that
a stilt-root habit does frequently develop in old trees. As
remarked by many authors this species is similar to H. Pierrzi
Hance in flower structure. H. Pierrei, however, differs
markedly in having glandular sepals, in the shorter petiole, and
in the midrib of the leaf which is sunk above as opposed to
raised in H. Beccariana.

Hopea Beccariana was founded by Burck on one of Beccari’s
Bornean collections (1177) which has leaves and immature fruits.
Four years later Heim described an almost identical Beccarian
collection (2761) as H. Nicholsoni.

A tree which I consider to be the same species was collected
on Penang Island by Curtis, and described by King in 1893 as
H. intermedia. King cited three collections when describing
the species—Curtis 425 and 1398, and Kunstler 3709 (Perak).
The last is certainly another species and its citation here may

Vol. IX. (1938).

326

have been partly responsible for subsequent confusion. How-
ever, from the excellent figure published by King in the Ann.
Bot. Gard. Calcutta, there can be no doubt which species King
had in mind as H. intermedia.

Brandis followed King but made some original suggestions
as to the affinities of the species. He suggested that H. intermedia
might be established as con-specific with H. Pierrei Hance, a
suggestion upon which Ridley acted in 1922 W then he united the
two species. Brandis was certainly correct in drawing attention
to the similarity in floral anatomy, but vegetatively these species
are so distinct that there can be little justification for the union.
Brandis also erroneously cited Curtis 1397 under H. intermedia:
it is H. pedicellata Sym. (see p. 327).

Ridley’s conception of H. Pierrei is very confused. His
citations include, besides several collections of H. Beccariana,
those of at least two other species none of which is H. Pierrei
Hance. (Ridley has also cited some collections of H. Beccariana
under H. micrantha). In 1920 Ridley described Balanocarpus
ovalifolius from a flowering collection of H. Beccariana (Haniff
3727). He cited also Curtis 426, which had tentatively been
placed under Balanocarpus latifolius by Brandis, but this is
actually. a_ sterile collection of seedling leaves of Shorea
Maxwelliana King. Ridley’s B. ovalifolius has since been
misinterpreted by Foxworthy.

It is clear from the Plates in Mal. For. Rec. 3 that in 1927
Foxworthy’s conception of H. intermedia was not that of King
but in 1932 this misconception was corrected, although a few
wrongly determined collections have been included in the
specimens cited.

We now have ample flowering and fruiting material of
H. Beccariana from the Malay Peninsula, but from Borneo, the
type locality, only Beccari’s original specimens with young
fruits and some recent sterile collections exist as far as I know.
It may, therefore, seem somewhat rash to unite H. intermedia
King (the Peninsula form) with H. Beccariana Burck, but the
vegetative characters of this tree, in particular the long petiole,
make it distinct from most related forms. Moreover we have
additional support for the union in field and wood characters.

Collections examined :—

MALAY PENINSULA: PENANG: Curtis 425,* 1938 .(Auth.
specs. of H. intermedia King), s.n.; S’pore 3460, 3474, 3721,
3727 (Auth. spec. of B. ovalifolius Ridl.), 9140; C.F. 2552,
8149, 9661, 10802, 11218, 11673, 11678, 11731, 27793, 200201
28051.

KEDAH : C.F. 8906, 11467, 17776, 21920.

= + There has been some confusion in tte labelling of some of the
sheets in the Singapore Herbarium: there are four sheets of 4.
Beccariana and two of H. pedicellata all \abelled Curtis 425.

Gardens Bulletin, S.S.

.
*

ae, 4 /
; ee Ey WS
: ag Fo oe a 7
atyheg or cession a a

; I > = <7

: - es _

: MS

ve my

+

5 cm.

,

Hopea pedicellata (Brandis) Sym.

Plate 19.

327

Kevanran C.F. 473851.

PERAK: Ridl. s.n. (Feb. 1891), C.F. 3413, 11040, 14384,
14387, 17066, 27822, 28504, 31075.

Dinpines 2 C.F 25601, 5007; 9857, 16574, 16500, 27842.
SELANGOR: C.F. 30148, 32854, 37290, 40762, 40970.
PAHANG: ©_P) Sr40, 27071, 37319, 42032.

NEGERI SEMBILAN : C.F. 23670, 24292, 26636, 43543.
Maracca: Goodenough 1447; C.F. 41552.

JOmORE C.F ..-£199}, 5852, 5863.

BORNEO: Sarawak: Becc. 1377 (Auth. spec. of H. Beccariana
Burck) and 2761 (Auth. spec. of H. Nicholsoni Heim).
Peunet co 20002, 20018, 37052," 37121.

British NortTH BoRNEO: Villamil 38 (?)

Hopea Beccariana, the merawan batu of our List of Botanical
and Vernacular Equivalents, is a common tree on ridges in
hilly jungle on Penang Island, the Dindings, and elsewhere
throughout the Malay Peninsula. It has coarsely fissured bark,
a tendency to become stilted, and frequently produces prolific
regeneration. The identity of the tree has not been very well
understood but it has usually been given as Hopea intermedia
King. Hopea intermedia King is here considered to be con-
specific with H. Beccariana Burck described from Borneo and
the latter name is adopted for our merawan batu.

Hopea pedicellata (Brandis) Sym. comb. nov. Hopea micrantha
sensu King in Journ. Asiat. Soc. Bengal 62, pt. 2: 124 (1893)
partim; sensu Ridl., FI. Mal, Penins. 1: 237 (1922) partim;
non H. micrantha Hook. f.. “Hopea Griffithii var. pedicellata
Brandis in Journ. Linn. Soc. 31: 69 (1805) specimina
Sudiatrana -excitisa; Ridl. in. op. cit. 238, in nota sub. H.
Pierrei. Hopea intermedia sensu Brandis in l.c. 67 quoad
Curtis 1397, non H. intermedia King. Hopea mengarawan
sensu Brandis in l.c. 70 quoad King 8170, non H. men-
garawan Miq.. Hopea Pierrei sensu Ridl., op. cit. 238,
partim; sensu Foxw., Mal. For. Rec. 10: 133 (1932) et ex
Burkill, Dict. Econ. Prod. Mal. Penins. 1193 (1935), pro
maiore parte; non H. Pietrei Hance. Plate 19.

As Brandis’ original description of this species under
Hopea Griffithii var. pedicellata is inadequate and _ other
descriptions have been confused by the inclusion of other
species, I have prepared the following—

Branchlets grey or purple-black, often conspicuously
lenticellate, usually puberulous towards the ends but later
glabrous. Leaves lanceolate or oblong-lanceolate, caudate-
acuminate at the apex, rounded or acute and occasionally
subcuneate at the base, usually about 6.52.5 cm. to 11.0
4.25 cm., glabrous on both surfaces except for the midrib which

Vol. IX. (1938).

328

is usualiy puberulous above, drying yellow-brown on _ both
surfaces but usually paler on the upper; midrib raised on both
surfaces, puberulous above, prominent and glabrous beneath;
main nerves about 13 to 16 pairs, fine and parallel, inarching
at the leaf margins, frequently furnished with small tufted
domatia in their axils, between each are 1 to 3 slightly shorter
intermediary nerves parallel to them; main and intermediary
nerves faintly visible and slightly sunk on the upper surface,
visible as faint yellow lines on the smooth under-surface;
reticulations invisible to the naked eye; petioles 0.8 to 1.2 cm.
long, sub-terete, dark and glabrescent or fulvous tomentose;
stipules minute, caducous. Panicles axillary towards the tip of
the branchlets or terminal, of solitary or paired racemes, 3.0 to
4.0 cm. long at most, glabrous or very sparsely puberulous;
ultimate branchlets of raceme 5.0 to 10.0 mm. long, each bearing
3 to 5 secund flowers about 1.5 mm. apart. Flowers sub-ovate
in bud, pedicellate, about 4.0 mm. long (including the pedicel);
pedicel about 1.0 mm. long (not thickening gradually towards
the sepal bases as in some related species), glabrous and drying
black. Sepals broadly ovate, shortly blunt acuminate, glabrous
on both surfaces, sparcely papillose outside. Petals linear-oblong,
usually blunt, tomentose on the portion exposed in bud, white
to creamy yellow in throat of corolla (Symington). Stamens
normally 15 (13 and 16 have been counted), the inner row of
10 sub-equal, the outer row (epistaminous on alternate stamens
of the inner row) smaller; filaments broad at the base narrowing
rather abruptly about the middle to the filamentous upper portion;
anthers oval, 4-celled, the anterior cells almost as large as the
posterior; appendage to connective filiform, about as long as the
filament and anther. Ovary and stylopodium broadly cylindrical,
slightly constricted below the stylopodium, truncate at the top;
style erect cylindrical, about as long as the stylopodium, stigma
minute. Fruit glabrous on all parts; stalk small but conspicuous,
about 1.0 mm. long; two long wing-like accrescent sepals
spathulate, blunt, about 4.50.8 cm., thick and woody at the
base, chartaceous on the free portions, about 7-nerved; three
small sepals, rotundate, about 0.7 cm. high, rather woody; nut
embraced in the lower half by the closely appressed imbricate
sepals or sepal bases, ovate-conical, about 10.0x6.0 mm.,
shining, more or less covered with a thin coat of resinous cement,
crowned with a short sharp apicula (persistent style).

The species here named Hopea pedicellata is common on
the hills of Penang and in parts of Perak. It was first mentioned
in botanical literature in 1893 when King referred four collections
in flower (167 and 1397) and fruit (266 and 8170) to Hopea
micrantha Hook. f., a species described from Borneo. Having
examined all the relevant herbarium material I agree with
Brandis in considering King’s identification to be wrong.
Superficially the two species are extremely alike but, as Brandis

Gardens Bulletin, S.S.

329

points out, the characters of the stylopodia are distinct—in
H. micrantha the ovary is surmounted by a cylindrical style
but in H. pedicellata there is a thick truncated stylopodium.

Although Brandis detected the mistaken identification of
King he does not appear to have had a very clear conception of
the species himself; he cites one specimen (Curtis 1397) under
H. intermedia, one (King 8170) under H. mengarawan, and on
a third (Curtis 167) he founded his H. Griffithii var. pedicellata.
The description of this variety is very brief—‘‘Pedicels nearly
as long as calyx. Branches of inflorescence puberulous in the
Sumatran specimens’’—and the citations are—Penang (Curtis
167), Sumatra (Forbes 3205, 3208). The Sumatran specimens
differ so considerably from Curtis 167 both in vegetative and
flower characters that I do not think they can be considered as
the same species; I am restricting Brandis name, therefore, to
the Penang specimen.

In describing this species as a variety of H. Griffithti Kurz,
Brandis has evidently been mainly influenced by the similarity
in stylopodium. I consider, however, that, in view of the marked
differences in other characters, the two plants should be kept
specifically distinct.

In 1922 Ridley referred a specimen (presumably Curtis 1397,
Govt. Hill, Penang, 1200 feet altitude) of this species to
H. micrantha Hook. f. but concerning Curtis 167, upon which
Brandis’ H. Griffithii var. pedicellata is founded, he says ‘‘it is
just possible that it is a hybrid between H. Pierrei and H.
micrantha’’.

In 1932 Foxworthy described H. pedicellata as H. Pierrei*
but the differences are so considerable that there seems no
justification for doing so. (See also notes under H. Pierrei).

Collections examined :—
PENINSULAR SIAM: S’pore 24205.

MALAY PENINSULA: PENANG: Curtis 167 (Auth. spec. of
H. Griffithii Kurz var. pedicellata Brandis), 266, 425, 1397,
a $1.5) POtes475, 3732, 3770; 0137; C.F. 11679, 277060,
28032.

PERAK ; “King 8170; C.F. 25562, 25635, 25874, 28083,
28994, 28995. .

KEeLANTAN = C.F 37667 (?)

TRENGGANU: C.F. 26913.

On the hills on Penang Island there are two common species
of Hopea known as merawan. One is a smooth-barked tree with
marked stilt roots, while the other (our merawan batu) has
fissured bark and is not markedly stilted. There has been much
confusion concerning the botanical identity of these trees. On

* King 3525 and Holmberg 820 should be excluded from the
citations in Mal. For. Rec. 10 : 134.

Vol. IX. (1938).

33°

pp. 325-327 the fissured-barked tree is investigated and it is
shown that it should be known as Hopea Beccariana Burck, while
in the above note it is concluded that the smooth-barked tree
should be called Hopea pedicellata (Brandis) Syin.

The Richetia Group

I have referred in these Notes on Malayan Dipterocarpacee
I and II to the very close relationship existing between certain
species of Shorea and Balanocarpus and in the latter paper (p. 27)
I have identified the wingless-fruited members of this group with
Heim’s genus Richetia. Similarly the winged-fruited members
may be identified with Heim’s section Richetioides of Shorea.
There is, however, no essential difference between Richetia and
Richetioides and there are species with rudimentary wings which
bridge the apparent gap between them. Lumped together they
form a natural group which, for convenience, I refer to as the
Richetia group.

The Malay Peninsula species of this group have been
enumerated by Desch (Mal. For. Rec. 12: 37) whose wood-
anatomical examinations entirely support my grouping based
on external morphological characters. I therefore feel that
there can no longer be any justification for maintaining some
members of this group in Balanocarpus and some in Shorea and
ai here referring all/the species to Shorea. The name-changes
involved are these—Balanocarpus multiflorus (Burck) Sym. in
Gard: Bull: SS. 7: 153 (1933) becomes Shorea multiflora (Burck) -
Sym. comb. nov.; “Balanocarpus coriaceus (Heim) Brandis in
Journ. gate? So 31: 112 (1895) becomes*Shorea Richetia Sym. -
nom. nov.; “Balanocarpus pahangensis Foxw., Mal. For. Rec.
No. 10: 145 (1932) becomes ‘Shorea balanocarpeides Sym. nom.-
nov.; Balanocarpus maximus King in Journ. Asiat. Soc. Bengal
62: 133 (1893) becomes’Shorea maxima (King) Sym. comb. nov.; -
and “Balanocarpus longiflorus (Brandis) Foxw. ex Sym. in Gard.
Bull. S.S. 8: 29 (1934) becomes’Shorea longiflora (Brandis) Sym.
comb. nov. |

To the Richetia group I assign the following species—
Shorea Faguetiana Heim, S. hopeifolia (Heim) Sym., S. multiflora
(Burck) Sym., S. balanocarpoides Sym., S. blumutensis Foxw.,
S. gibbosa Brandis, S. Richetia Sym., S. kalunti Merr., S. resina-
negra Foxw., S. maxima (King) Sym., and possibly S. longiflora
(Brandis) Sym. The last three, however, are in some respects
atypical. Several hitherto undescribed Bornean species, including
S. xanthophylla Sym. and S, acuminatissima Sym. described in
this paper, also belong to the Richetia group.

The diagnostic anatomical, morphological, and field
characters, of the group are many, but I shall defer enumeration
of them until the undescribed members have been more fully
studied. One important point I should like to record here 1s

Gardens Bulletin, S.S.

i

331

that all species have 2-celled anthers. Foxworthy described
S. resina-negra as having 2-celled anthers, but I cannot find
any other record of this condition in other members of the family,
nor in family or generic diagnoses.

I have referred to this group of species somewhat non-
comittally as the ‘‘Richetia Group’’. It will, of course, be
necessary to assign to it a definite botanical rank such as genus,
subgenus, section, or subsection, but I leave such assignation
until a time, not too far distant I hope, when a reliable taxonomic
treatment of the genus Shorea will be produced. Considered
purely from a botanical standpoint there can be little doubt that
the Richetia group merits generic rank, and there are other
groups within the genus, e.g., sections Eushorea Brandis and
Anthoshorea Brandis, that could also quite legitimately be so
treated. A similar condition exists in the genus Hopea. Let us
hope, however, that if and when these genera are next mono-
graphed, full consideration will be given to the claims of those
whose interest in the trees concerned is mainly from the utilization
standpoint. It is alarming to contemplate the effects of a revision
in nomenclature of the Dipterocarpacee under a narrow generic
conception, but as this can quite conveniently be avoided by
the creation of sub-genera rather than genera, it is to be hoped
that this latter method of treatment will be followed.

Shorea assamica Dyer forma assamica Sym. stat. nov. Up deed
assamica Dyer in Hook: f., Fl. Brit. Ind. x: 307 (1874);
Brandis in Journ. Linn. Soc. 31: 85 (1895), et Ind. Trees. 70
(r911); Foxw. in Philip. Journ. Sc. (Bot.) 4: 516 (1909);
Troup, Ind. Woods and Uses 241 (1909), et Silv. Ind. Trees
133 (1921); Gamble, Man. Ind. Timb. 83 (1922); Pearson et
Brown, Commerc. Timb. Ind. 1: 119 (1932); Parkinson in
Ind. For. Rec. (Bot.) 1: 40 (1937). /

/forma philippinensis (Brandis) Sym. stat. nov. Shorea
philippinensis Brandis in l.c. (1895) 88; Foxw. in op. cit. 6:
272 (I9II) partim, et 13:-190 (1918); Merr. Enum. Philip.
Blow: Pl 37 98° (19023)" “Shorea sp. aff. S.°Harmandii
Pierre,’’ Foxw. in op. cit. 6: 272 (1911). Shorea pallida
Foxw. in op. cit. 13: I90 (1908).

forma Koordersiit (Brandis) Sym. stat. nov. [Vatica
celebica Koorders, Mss. in Herb. Koord. (1895). “Shorea
Koordersii Brandis ex Koorders in Meded./’ ’S Lands Plant.
19: 355 (1898); Icon Bogor. 1, Tab. 80 (1901); Heyne, Nutt.
Plant. Ned. Ind. edit. 1, 3: 303 (1917) et edit. 2, 2: 1120
(1927); Endert in Korte Meded. Bosch. 51 (Tectona 28:
288) 1935. J
Vforma globifera (Ridl.) Sym. stat. nov. Shorea globifera
ian, Pl hele: Penins. >: 232 (1922);. Foxw.}. Mal. For:

Vol. IX. (1938).

332

Rec. 3: 35 (1927), et 10: 191 (1932); Sym. ex Desch, Mal.
For. Rec. 12: 27 (1936). Plate 20.

I have here united under one specific head four ‘‘species’’—
S. assamica Dyer (from Assam), S. philippinensis Brandis (from
the Philippines), S. Koordersii Brandis (from Celebes), and
S. globifera Ridl. (from the Malay Peninsula). Having examined
a considerable amount of material of each I do not think that
specific differentiation of these forms is justified, but there are
certain fairly constant definable minor differences such as one
would expect in distinct geographical races of a widely distri-
buted species. I have, therefore, recognized four forms—forina
assamica (from Assam and Burma), forma philippinensis (from
the Philippines), forma -Koordersii (from Celebes and the
Moluccas), and forma globifera (from the Malay Peninsula and
Sumatra).

There has been comparatively little systematic confusion in
this species. The form assamica, the first to be described, was
collected in Assam and described by Dyer as S. assamica in 1874.
This form has since been collected also in Upper and Lower Burma
and ‘Tenasserim.

The form philippinensis was first described from Luzon by
Brandis in 1895 as S. philippinensis, but this form does not appear
to have been very clearly understood in the Philippines. In
1911 Foxworthy cited under S. philippinensis several collections
of other species, and he listed three authentic collections under
the heading of ‘‘Shorea sp. aff. S. harmandii Pierre’’. These
last three were made the basis of a new species, S. pallida Foxw.,
in 1918 but this was rightly reduced to §. philippinensis by
Merrill in 1923.

The form Koordersii was described as S. Koordersii by
Brandis in 1898 from collections from North Celebes. Brandis
drew attention to the similarity between this and his S. philippin-
ensis, and even suggested their possible specific union when more
material of the latter was available.

The Malay Peninsula form, forma globifera, was described
by Ridley from fruiting material as S. globifera in 1923. In
1932 Foxworthy added a description of the flowers. This form
has since been collected in Sumatra by Yates.

The various forms have been fairly adequately described
under the names of S. assamica, S. philippinensis, S. Koordersit,
and S. globifera, but concerning Foxworthy’s description of the
flowers of the last-named, I would remark that I have been unable
to verify the presence of more than 15 stamens, the species being
typical of the section Anthoshorea Brandis.

In the following Key the main differences between the forms

are enumerated.

Gardens Bulletin, S.S.

ass RSS

a globifera (Ridl.) Sym.

}

:
=
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ey
ww
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8
S
“~
S
8
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Plate 20.

AAAI LS

<—S

mee v4 %.

339

(1) Branchlets of raceme usually simple, solitary, and alternate;
stigma distinctly 3-partite; fruit wings up to about 10.0
em. long; appendage to connective about 1% times as
MAS, LAS TAME MCT, .<. 500 4a cjagraistale itd scat obsieie.« « forma assamica

(1) Branchlets of raceme frequently bi-furcate or fascicled
(2) Stigma usually distinctly 3-partite; fruit wings up to

about 10.0 cm. long; appendage to connective as

lone asthe anther: Ae i2 one, a. forma globifera

(2) Stigma simple, flattened, or obsurely lobed; fruit wings
up to about 7.0 cm. long; appendage to connective
about twice as long as the anther .....:...........0... (3)

(3) Leaves usually closely peltate and smooth
PCH tn te Phot n ous cae os forma Koordersii

(3) Leaves usually sparsely soft-stellate-hairy
ipeneatiy .gissh 1). ..0iba ae. forma philippinensis

Collections examined :—

Boor ora dssamica): G. Mann s.n.. (Auth. spee.tof 'S:
assamica Dyer), G. Mann s.n. 1883, G. Mann s.n.7 1890;
to Mill se ole 73°, Siminons s.n.; A. Das: 106R2: VU.
Kanjilal 3414; For. Dept. Assam 13115 aid s.n. 10.12.36.

BURMA (forma assamica): E. M. Buchanan s.n.; S. M. Toppin
6019; Sukce 2239, 7956; Parkinson 1 and 317.

MALAY PENINSULA: (forma globifera): KEDAH AND LANG-
AMO 7050 )17075507 740) LI 7 oy O27, 12218) H2222,
12227, 20735, 27802, 33176.

BUUANTANGS (e227 44. (32745,.°32740;, 32808, 32800,
33266, 33269, 33270, 33430.

PERAK: C.F. 3072, 8308, 10369, 12983, 12985, 16349,
16706, 27226, 29866, 29894, 29924, 31038, 38103, 39070,
39071, 39223, 41613.

TRENGGANU: C.F. 24282.

PAHANG: C.F. 4027, 4052, and 5073 (Auth. specs. of
S. globifera Ridl.); C.F. 797, 856, 20345, 29398, 29908.

SUMATRA (forma globifera) : Yates 1264, 1675.

PHILIPPINE ISLANDS (forma philippinensis) : Luzon: Vidal
983 (Auth. spec. of S. philippinensis Brandis); Bur. For.
12996 (Auth. spec. of S. pallida Foxw.); Bur. For. 6068,
7084, 13924, 17158, 17643, 17652, 22290, 23471, 27007.

Ay aaa Elmer $4175; For. Bur. 24331, 25021, 27710,
au 25x¢
BASIEAN.:) Fors Bur. .18887; Bur. Sc: 15418, 16102.

CELEBES AND MOLUCCAS (forma Koordersii) :

MINAHASA: Herb. Koord. 167348 and 167358 (V. celebica
Koord. ms.); 159828, 167378, 167398, 167428 and
167966 (Auth. specs. of S. Koordersii Brandis).

TALIABU : Herb. Bog. 362.

PuLau Bisa: Herb. Bog. 54 and 55.

Vol. IX. (1938).

334

PuLau Opt Herb. Bog. s.n.

The species which we in Malaya have been accustomed te
call Shorea globifera is here recognized as a geographical race
of a remarkably widely-distributed species occurring from Assam
to the Moluccas. This species has been described separately in
several different localities—as S. assamica in Assam, S. globifeva
in the Malay Peninsula, §. philippinensis in the Philippines, and
S. Koordersii in Celebes. These are here united under the oldest
specific name S. assamica but the various geographical races,
which do exhibit slight differences, are recognized as forme, ot
forms, the former specific epithets being retained to describe
them. Thus our S. globifera Ridl. (meranti pipit) becomes
Shorea assamica Dyer forma globifera (Ridl.) Sym.

‘Parashcrea malaanonan (Blanco) Merr., Spec. Blanco. 271
(1918), et Enum.. Philip. Blow.. Pilioa%* 2100 (n923)2 Vase
Bull. Jard. Bot. Buitenz. HII, 8: 375 (1927) etvex > Mer,
Plant. Elmer. Born. 202 (1929). Mocanera malaanonan
Blanco, Fl. Philip. 858 (1837).. Dipterocarpus malaanonan
Blanco, op. cit. ed. 2: 312 (1845) et ed. 3, Il: 214 (1878).
Shorea malaanonan (Blanco) Bl., Mus. Bot. Lugd.-Bat. 2:
34 (1852); De Candolle, Prod. 16, pt. 2: 631 (1868); Brandis
in Journ. Linn. Soc. 31: -103 (1895); non S, malaanonan
sensu Merr. et Rolfe in Philip. Journ. Sci. (Bot.) 3: 115
(1908), et sensu Foxw. in Philip. Journ. Sci. (Bot.) 6: 270
(r9o11). Parashorea plicata Brandis in l.c. 104 (1895); Merr.
et Rolfe in l.c. 114 (1908); Foxw. in l.c. 280 (1911), et in
Lc. 13.2 £04 (1918);» Whitford, Philip) Burs Ferm Salk re:
63, tab. 60 & 61 (1911). Parashorea Warburgii Brandis in
l.c. 105 (1895); Foxw. in Leafl. Philip. Bot. 6: 1954 (1913)
et in Philip. Journ. Sei. (Bot.).13 : 994 (1ro8Ss)s, Mere 1c:
100 (1923). Plate 21.

Parashorea malaanonan, if the interpretation here accepted
is correct, first appeared in botanical literature in 1837 under the
name of Mocanera malaanonan Blanco. Blanco’s description,
which was not materially amended in subsequent editions of his
Flora de Filipinas, is translated as follows—

Leaves ovate-lanceolate, broad, frequently pointed, whitish
on the under surface. Calyx divided almost to the base in 5
oblong segments which are slightly contorted over the fruit: the 3
outer segments longer and measuring about 3 inches.’ Corolla
oe kn gee a SLAMENS ....0..05.+-.+---- Nut oval, free whemmipe,
surrounded by the calyx, with delicate (fragile) shell crowned
by the long style which bears 3 stigmas, unilocular and one-
seeded, the seed with 3 or 4 lobes.

Large trees, known in Angat by the possibly equivocal name

““jeme’’* in length, wide, and

forefinger and thumb, z.¢. about 8 in. or 20 cm.

Gardens Bulletin, S.S.

—

0h Te xe

nar, ‘
Se

Eee LEI

————

- Bis

Parashorea malaanonan (Blanco) Merr.

Rs ber
edt? ape eae '

dp seays

Plate 21;

exes ke.
Sy

Soo

whitish and smooth (soft) beneath when young. This character
sufficiently distinguishes the species from others in the genus.
The tree also yields on aromatic resin, but in very small quantities.

Blanco neither cited nor left a type specimen. In the
second and third editions of his work the name is changed to
Dipterocarpus malaanonan. In 1852, in a conspectus of
Malaysian species, Blume changed the genus to Shorea. He
cited no specimen, and it would appear that he saw none but
considered that the species described by Blanco, whatever it was,
must be a Shorea. At that time the genus Parashorea was
unknown and Blanco’s description is essentially that of a Shorea,
except that he says that the nut is free when ripe. Similarly De
Candolle, in 1868, and Brandis, in 1895, enumerated Shorea
malaanonan, but they made no attempt to tie this name down to
any extant herbarium material.

The genus Parashorea was described by Kurz in 1870, the
species enumerated being P. stellata from Burma and P. lucida
from Sumatra. In 1895 Brandis added two new species from the
Philippines, P. plicata, based on flowers and young fruits, and
P. Warburgii, based on ripe fruits evidently picked from the
ground.* He did not suspect that either, or both, of these might
be the plant he enumerated as Shorea malaanonan (Blanco) BI.

Merrill and Rolfe were the first to attempt to identify
Mocanera malaanonan Blanco with existing herbarium material.
In 1908 they identified Blanco’s description with a _ species
described by Vidal in 1883 as Shorea polita, citing a number of
Philippine collections. The only supporting argument for this
interpretation is the statement “‘we are of opinion that the above
specimens are referable to Blanco’s species and accordingly his
specific name is accepted, Vidal’s Shorea polita being here
reduced.’’ ‘The numerous discrepancies between their plant and
Blanco’s description are not explained.

This interpretation of Merrill and Rolfe remained in current
use in Philippine literature until r918 when Merrill himself, in
his Species Blancoane, rejected it. In this work he states—‘‘l
am now convinced tnat this interpretation was erroneous; that
Shorea polita is a valid species entirely distinct from Mocanera
malaanonan Blanco; and that Blanco’s species is identical with
the common and widely distributed Philippine Parashorea plicata
Brandis. Among all the Philippine Dipterocarpacee, Para-
shorea plicata Brandis is the only one that agrees with Blanco’s
description in the characters of the leaves, whitish beneath,
which are further described as wide, pointed, and a “‘geme’’ (i.e.
15 to 18 cm.) long, while the pericarp of the fruit is described.

* P, plicata Brandis and P. Warburgii Brandis were united by
Merrill and Rolfe in 1908, and by Foxworthy in 1gir. In 10913 and
1918 Foxworthy, and in 1923 Merrill, separated them again, but in
1927 Van Slooten, with whom I am in agreement, reunited them,

Vol. IX. (1938).

336

as fragile. The description conforms to Parashorea plicata, and
I have not the slightest hesitation in adjusting the synonymy.
The native name cited by Blanco, malaanonan, is valueless in
interpreting the species, as it is a made up one, literally “false.
anonang’’, anonang=Cordia myxa Linn., and, as used to-day,
is very loosely applied, although I have specimens of Parashorea
plicata Brandis from Laguna Province, Luzon, bearing this
name’. Merrill cited an Illustrative Specimen No. 1053.

I agree with this second interpretation of Merrill’s to this
extent—Parashorea plicata Brandis answers more closely to
Blanco’s description of Mocanera malaanonan than does any other
known Philippine dipterocarp. Nevertheless, Blanco describes
the fruits as having a persistent style with three stigmas and
wings only 3 in. long. These characters cannot be reconciled
with Parashorea plicata and, had I been faced with the original
interpretation of Mocanera malaanonan, I should certainly have
regarded it as a nomen dubium. But in deference to the fact
that van Slooten and others have accepted Merrill’s second
interpretation, and because there certainly is not adequate
evidence to prove that he was wrong, it seems desirable to allow
this interpretation to stand.

The first adequate description of Parashorea malaanonan
Merr. is that published by van Slooten in 1927. ‘This is a very
satisfactory description of the species as I know it except con-
cerning the leaves, of which van Slooten says “‘the undersurface
felt-like to the touch by an usually hardly visible stellate
tomentose brownish indumentum, which may have a more or
less intensely greyvish-white hue especially in young leaves and
in leaves of saplings, along the midrib and nerves, however
usually finely adpressed pubescent, afterwards on the whole
undersurface often glabrous or nearly so ...... hbnee gear i am
inclined to think that this gives a wrong impression of the
immature leaves, as I have examined some specimens, obviously
immature, which are almost entirely glabrous. Indeed it is
possible that some of the speciinens with tomentose leaves
examined by van Slooten are not typical P. malaanonan but
what I have described. (vide p. 338) as P. malaanonan variety
tomentella. ‘The tomentose specimens concerned, which Dr. van
Slooten has kindly communicated to me, are unfortunately all
sterile and I do not think it is possible to say more at present than
that they are referable either to P. malaanonan Merr. or P. malaa-
nonan Merr. var. tomentella Sym. A careful field investigation
of mature and immature stages of both forms will be necessary
to decide the point.

Collections examined :—
BORNEO: BRUNEI: C.F. 30531.
BRITISH NORTH BORNEO: Villamil 20 (?); Elmer 21122
(H.B: 9037.26.15), 22739 (H.B, 9037.26.17); For: Depts here

Gardens Bulletin, S.S.-

337

No. 1; Wood 2266 (H.B. 937.26.1); C.F. 36722 (S.H. 3922),
38727 (S.H. 4351), 38738 (S.H. 4362), 38777 (S.H- 4401),
38833 (S.H. 4457), 39019 (S.H. 4643), 41065 (S.H. 4720),
ATOBO (8-1. AQS51), ari22 (S.A. 5Q70).

PHILIFPINE ISLANDS :* Minpanao: Warburg s.n. (Auth.
spec. of P. Warburgii Brandis); Elmer 14066 (H.B.
Q37-20.23), Baker 6232; Bur, Ags. 356. (H.B; 937.26.19) (?);
For. Bur. 9182, 11560, 12355, 24328, 27471, 27646, 27720,
29420.

NeEGROS: Bur. Sc. 1144 and 1616; For. Bur. 11194, 11244,
bieae (E1.. 937.20.25), 12414, 13755, CE | oS 7.2026):
17418,'.17488, 22677 (H.B. 037:26.30), 29466, 20467; 30127.

PANAY: For. Bur. 17838, 22567, 22569.

Tevte: Wenzel 711, For: Bur. 11635 (H-B. 0937.26.24),
12624, 12769.

BILIRAN: Bur. Se. 18489 and 18992.

SAMAR: For. Bur. 21077 (H.B. 937.26.22) and 25934.

MAsSBATE: Bur. Sc. 1673; For. Bur. 12570 and 12501.

CATANDUANES ISLAND: For. Bur. 6679.

PoLtitLo IsLtaAND: Bur. Sc. 10282.

Luzon: Widal 76, 900, and 2033 (Auth. specs. of P.
pblicata Brandis); 1053 (Illustrative specimen of P. malaano-
nan Merr. in Spec. Blanco. : H.B. 937.26.21); Loher 113, 114,
wAZ ZO, Hamer, 17605, (HB. 937.26.20); Bur. Agri. 51, 104
(eres 1037-20-15), 201; Bur. Sc. 1461, 2660, 3289, 11944,
¥22 745) Pore Bur. 4520, 5754, 10010, 10010; TOO59, 1007s,
10170, 80404, 10500 (EL.BY 9327.26.27), 10502, 10506, 10564,
E0004.) 10006,..1070%,. 10730; 11520, 12775, 14336, 15435,
i762, 26251, 20711, 21100; ZIA12, 216904, 21717, 22240, 22250;
22340, 22034, 23.771, 29819.

The following specimens may be either P. malaanonan Merr.
or P. malaanonan var. tomentella Sym. :—

BORNEO : SoutH East BORNEO: Endert 5165 (H.B. 937.26.14);
pho Too46" (TB: 0937.26.90), bb. 11286.(H.B.. 037.26.10),. bb.
TiOom iH a (O37:20.16),) bb.) 12110. (E1.B.,. 0937.26.12) 4) Bb.
he TAS CE iw087 20-1 8), bd T6860, (HB. .637.26.2)5. Db.
$2002 8 h1037.20.3), bby, 18806 ¢H.B. 927 .26.4), bb.
foGg0s4 (ln: 647.20.5), bb. 19131.(H.B.. 937.26.6), bb. 19144
(i028 7.26.5), bo 19140 .(M..B... 937.26: 7).

BritisH NorRtTH BORNEO: Elmer 21194 (H.B. 937.26.16).

PHILIPPINE ISLANDS: MINDANAO: Hutchinson 7563 (H.B.
6937.20.28).

* Some of the specimens listed from the Philippines are apparently
from seedling or sapling trees. The identification of such specimens
should be considered as tentative. Unfortunately, when examining
the specimens I failed to list which had mature ‘leaves and which
immature.

Vol. FX. (1938 ).

338

Luzon : For. Bur. 6047, 10221 (H.B. 0937.26.29), 14936,
29347. |

“Parashorea malaanonan (Blanco) Merr. var. tomentella Syming-
ton, var. nov., ‘‘Parashorea sp.’’, A. V. Thomas in Mal.
Forester 4: 131 (1935) partim; Sym. ex Desch, Mal. For.
Rec. 12: 33 (1936). Plate 22.

Parashorea malaanonan (Blanco) Merr. forma typica valde
affinis, a qua foliis subtus tomentosis, stylis appendicibusque
brevioribus differt.

Branchlets pale fulvous-tomentose, finally becoming dark
and glabrescent, annular stipular scars conspicuous. Leaves
ovate to elliptic-lanceolate, acute or bluntly acuminate at the
apex (the apices of nearly all the leaves of my specimens are
damaged), rounded or sub-cordate (occasionally acute at one
side) and more or less unequal at the base,.very variable in size
from about 8.0 xX 3.0 cm. to 24.0 X 13.0 cm. or more, glabrous and
shining above, covered with a pale fulvous felt-like tomentum
beneath, ridges between the main nerves sometimes conspicuous;
midrib slightly elevate in the lower portion on the upper surface,
prominent beneath; main nerves about 12 to 16 pairs, inconspi-
cuous above, prominent beneath; nervules join the main nerves
in fine wavy sub-parallel lines; petioles 1.5 to 2.5 cm. long, thick,
rugose, tomentose; stipules caducous (not seen), leaving annular
scars. Panicles terminal or in the upper leaf-axils, racemose, lax,
pale cinereous- or fulvous-tomentose, about as long as the leaves;
ultimate branchlets about 3.0 to 6.0 cm. long, somewhat zig-zag,
2 to 6-flowered. Flowers sub-secund, 1.0 to 1.5 cm. apart in
mature panicles, subtended by caducous bracteoles leaving
prominent scars, about 14.0 mm. long in mature bud, about 3.5-
cm. across when expanded, usually said to be white (occasionally
creamy yellow). Sepals pale close-tomentose on both surfaces;
three outer oblong-obtuse; two inner ovate-acuminate, very
slightly shorter. Petals oblong-obtuse, pale tomentose outside
and along one edge inside. Stamens 15, in three rows, the inner
row of 5 slightly taller than the other rows which are sub-equal;
filaments short, flattened, broad at the base, narrow above,
occasionally furnished at the sides with a few fine hairs; anthers
oblong, about 3 times as long as the filaments, sagittate at the
base, cells pointed at the apex the two inner slightly shorter
than the outer; appendage to connective awl-shaped, thick at
the base, about one half to as long as the anther. Ovary ovate-
conical, tapering gradually into the cylindrical style; style about
as long as the sepals in mature flowers, tomentose on the lower
half, glabrous above; stigma minute, flat. Fruit a 5-winged
nut (mature fruit not seen); stalk short or almost absent, stellate
hairy; bases of accrescent calyx lobes shortly united to embrace
the lower third of the nut, also stellate hairy; three outer calyx

Gardens Bulletin, S.S.

5 cm.

b rT, ™ a
Gees Tren TI >

Be NH RH a mpp yoni

Parashorea malaanonan var. tomentella Sym.

22.

oF
are

Pi

339

lobes linear-oblanceolate or spathulate, rounded at the apex,
about 14.0 cm. long and 1.5 to 3.0 cm. broad, g to 12-nerved,
sparsely hairy; two inner lobes linear-acute, about 3.0 to 7.0
cm. long and up to 6.0 mm. broad but usually much narrower, 3
to 6-nerved; nut (immature) pale tomentose, surmounted by the
long style which is glabrous on the upper portion.

Recent study of the timber commonly known as ‘“‘Borneo
white seraya’’ has necessitated botanical investigation of the
identity of the tree, or trees, producing it. It was generally
assumed that the main source of the timber was a single botanical
species, Parashorea malaanonan (Blanco) Merr. Sterile botanical
material, authenticating two logs (from different trees) used for
pilot tests was thought by me (vide Thomas, in I.c.) to belong to
the same species, although the specimens differed considerably
in leaf indumentum. The subsequent receipt at Kepong of
numerous collections of fertile material of Parashorea from British
North Borneo led me to suspect that these specimens represent
two distinct, possibly specifically distinct, forms. Examination
of flowering collections confirms this suspicion. The smooth-
leaved collections are Parashorea malaanonan (Blanco) Merr. and
those with tomentose leaves. I am naming P. malaanonan var.
tomentella. The main differences between these two forms are
as follows—

P. malaanonan—mature leaves smooth and _ usually
glaucescent on the undersurface; style long, exceed-
ing the sepals, tomentose on the lower third only;
appendage to connective longer than the filament
and anther.

P. malaanonan var. tomentella—mature leaves covered
with a close, pale, fulvous, stellate tomentum; style
not or hardly exceeding the sepals of mature
flowers, tomentose on the lower half; appendage to
connective less than half the length of, to as long
as the anther.

I have resisted a strong temptation to describe this tomentose
form as a new species. If one considers the recent collections
of Parashorea from British North Borneo alone, the two forms
appear in sharp contrast and there would appear to be little
justification for not recognizing two distinct, but closely related,
species., Further, Mr. Keith informs me that specific separation
would be justified, from a consideration of field characters.
There are, however, several collections (unfortunately all sterile),
mainly from South East Borneo but some from British North
Borneo, Mindanao, and Luzon, which appear to be intermediate
between the two. These collections are listed as ‘‘either P.
malaanonan Merr. or P. malaanonan var. tomentella Sym.’’ on
p. 338. .In 1911 Foxworthy (in Philip. Journ. Sci. (Bot.) 6: 281)
cited some of these collections from British North Borneo and

Vol. IX. (1938).

a i te et tt A EL LAI CC LLL LIT

340

from Luzon, referring to them as a form of Parashorea plicata
“‘which is pubescent on the voung twigs and on the undersides
of the leaves’’. In 1927 van Slooten cited Elmer 21194 from
British North Borneo and several collections from South Borneo
under Parashorea malaanonan, considering that they represented
« juvenile form of the leaves of that species.

‘These specimens I have segregated from either P. malaano-
nan or the variety tomentella. As I have remarked on p. 337,
further field investigation is required to decide their status, and
when this decision is made the question of raising Parashorea
malaanonan var. tomentella to specific rank should be
reconsidered.

Collections of P. malaanonan var. tomentella examined :—

BRITISH .NORTH BORNEO’: CoP .38745 (8S... 4400), 1 eee
of P. malaanonan var. tomentella Sym.); Otik; Kabili
Forest Reserve, Sandakan, level land; 23rd March, 1935;
flower. Field notes—‘‘vern. name, urat mata; tree 150 feet
high, 65 in. girth; flower white’’

For Dept: B.N.B. No. 2; C.F. 36673 (S.H. No. 3896),

38687 (S.H. 4310), 38726 (S.H. 4350), 28734 (S.H. 4358),
38845 (S.H. 4469), 38854 (S.H. 4478), 38871 (S.H. 4485),
38935 (S.H. 4559), 38072 (S.H. 4506), 41061 (S.H. 4712),
41064 (S.H. 4719), 41087 (S.H. 4950), 41289 (S.H. 7154),

41332 (S.H. 71097).

The main source of the commercial timber called ‘‘Borneo
white seraya’’ has been generally supposed to be a single botanical
species, Parashorea malaanonan, which is known in the jungle
in British North Borneo as urat mata. ‘Two forms of urat mata
have, however, been recognized—one with smooth leaves and the
other with tomentose leaves. The botanical identity and status
ot these forms is here investigated, and it is shown that only the
smooth-leaved form is Parashorea malaanonan in the strict botani-
cal sense. The form with tomentose leaves is described as a
variety of P. malaanonan— var. tomentella. It is considered
that this. variety may be worthy to rank as a distinct species but
further investigation, particularly of immature stages in the field,
is required to decide the point.

“Shorea acuminatissima Symington, sp. nov. Plate 23.

S. Faguetiane Heim valde affinis, sed ramulis novellis
terrugineo-tomentosis et foliis caudato-acuminatis infra scabrius-
culis distincta.

Branchlets densely or sparcely ferruginous-tomentose when
young, later terete, glabrous, and dark. Leaves ovate- or oblong-
lanceolate, caudate-acuminate at the apex, rounded or sub-acute
at the base, margins slightly revolute, about 7.0x2.5 cm. to
17.0 6.0 cm., glabrous and drying yellow- or red-brown above,

Gardens Bulletin, S.S.

ge

*
«

Plate 23. Shorea acuminatissima Sym.

™
‘>

341

similarly coloured but scabrid stellate-hairy on the nerves
beneath; midrib slightly depressed above, prominent and stellate-
hairy beneath; main nerves about 13 to 18 pairs, inconspicuous
above, strongly elevate and stellate-hairy beneath; nervules faint
but visible on both surfaces, usually sparsely hairy beneath;
petioles terete, 1.0 to 1.5 cm. long, ferruginous-tomentose when
young, scabrous and black later; stipules broadly oblong, oblique,
rounded, about 1.00.5 cm., stellate-hairy, caducous. Panicles
exillary and terminal, usually solitary, racemose, shorter than
the leaves, pale tomentose; branchlets about 0.6 cm. apart, 1.0 to
2.0 em. long when mature, each bearing 4 to 6 secund flowers
(in the immature panicle the flower buds are sub-distichous and
subtended by small, frequently bifurcate, puberulous. bracteoles).
Flowers about 2.0 mm. apart, ovate-lanceolate in mature bud,
about 7.0 mm. long including the pedicel; pedicel 1.0 mm. long,
densely pale tomentose. Sepals sub-equal, ovate-rotundate,
obtuse, tomentose on the portions exposed in bud, white (C.F.
38722 & 38723). Stamens 15, pairs alternating with single
stamens, of three sizes; anthers ovate-oblong, 2-celled; filaments
about as long as the anthers, broad at the base, narrowing slightly
towards the top; appendage to connective filiform, about as long
as the anther. Ovary and stylopodium conical, the former
glabrous, the latter densely hairy; style about as long as the
stylopodium, cylindrical, glabrous; stigma capitate, small. Fruit
sessile; calyx lobes chartaceous, sparsely hairy, three outer larger
than the inner two; large wings spathulate, about 5.01.2 cm.;
small wings similar but slightly (or considerably) smaller; bases
of fruiting sepals thickened and woody, closely appressed to the
base of the nut to form a 5-angled stellate-hairy cup, pointed at
the base; mature nut ovate-oblong, rostrate, about 1.5 x 0.7 cm.,
densely fulvous-tomentose.

This is a typical member of the Richetia group of Shorea and
appears more closely related to S. Faguetiana Heim than to any
others with which I am familiar. The leaves are rather distine-
tive, and there should be little difficulty in identifying the species
from vegetative characters alone. ‘There is a distinct resemblance
in leaf to Shorea Forbesti Brandis from New Guinea, but the
natural relationship to it is remote. I suspect that S. Forbesii
is a Hopea.

Collections examined :—

BRITISH -NORI EH: BORNEO:, C.F .. 38723) (S.H. 4347). (lype
of S. acuminatissima Sym.); Ayan; Experimental Station,
8th mile Sandakan, level land in forest; 29th March, 1935;
flower. Field notes—‘‘vern. name, gagil; tree 120 feet high,
53 in. girth; flower small, white, in cluster’’.

C.F. 41075 (S.H. 4886) (Paratype of fruit of S. acumina-
_tissima Sym.); Puasa; Kabuli Forest Reserve, Sandakan, top

Vol. IXY IO 38).

.
)
|

342

of hill; 21st April, 1935; fruit. Field notes—‘‘vern. name
selangan kuning; tree 9o feet high, 60 in. girth; fruit red’’.

pen 38722 (S,H. 4346), 38724 (S.H. 4348), 38702 (S.H.
4416).

BRUNEI: C.F. 30404.

The species here described as Shorea acuminatissima is a
large tree which has been collected several times from British
North Borneo under the names ‘‘gagil’’ and ‘‘selangan kacha’’.
It has also been collected in Brunei. Timber samples have not
been available for study but, from botanical considerations, it
may safely be said that the timber will be classified as ‘‘white
meranti sub-group 2’’ (vide Desch, Mal. For. Rec. 12: 37).

/
oe xanthophylla Symington, sp. nov. Plate 24.

S. multiflore (Burck) Sym. valde affinis sed foliis fructuque
majoribus, petalis pro rata brevioribus, appendicibus ciliatis,
differt.

Branchlets grey- or reddish-brown, pale puberulous towards
the ends. Leaves oblong, acuminate at the apex, rounded or
sub-cordate at the base, from about 19.0 x 6.0 cm. to 37.0 X 10.0
em., glabrous on both surfaces, drying yellow-brown or greenish-
yellow on the lower surface, usually of a greener tinge above;
midrib very slightly elevate above, prominent beneath; main
nerves g to 13 pairs, faint above, prominent beneath; nervules.
joining the main nerves in rather distant sub-parallel lines, invisi-
ble or faint above, rather prominent beneath; petioles 1.0 to 2.0
cm. long, rather thick, dark, rugose; stipules not seen. Panicles
axillary or terminal, solitary or several together, lax, up to about
14.0 em. long, pale tomentose; branches solitary or in pairs,
irregularly branched; ultimate branchlets with 1 to 6 sub-secund
flowers 1.3 mm. apart. Flowers ovate-lanceolate in nature bud,
about 5.0 mm. long (including the pedicel); pedicel 1.0 mm. long,
pale tomentose. Sepals sub-equal, ovate-rotundate, mucronate
rounded or emarginate at the apex, tomentose outside, glabrous
within. Petals oblong, tomentose on the portion exposed in bud,
said to be white (38938), yellow (38742), lemon yellow (36776),
and yellow ochre (38933). Stamens 15, pairs alternating with
single stamens, of three heights; anthers elliptic-oblong, 2-celled;
filaments I to 2 times as long as the anthers, broad at the base,
narrow in the upper half; appendage to connective a curved awn
about as long as the anther, minutely ciliate towards the end.
Ovary ovate-conical, glabrous below, tomentose in the upper
portion; style cylindrical, about % as long as the ovary, glabrous;
stigma minute, apparently simple. Fruit sub-sessile; calyx lobes
sub-equal, broadly ovate, about 1.0 cm.x0.7 cm., decidous
fulvous-tomentose, shortly united at their bases to form a woody
receptacle; nut obovate, pointed, about 2.3 cm. x 1.7 cm., striate,

Gardens Bulletin, S.S-

Shorea xanthophylla

Ren Ba A

n p

Sa
i=

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ins

(~ =
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Plate 25.

Shorea sandakanensis Sym

Ca ee ae.

343

fulvous tomentose, on germination splitting along three lines
at the apex.

This is yet another species of the Richetia group (vide
»y. 330). It is extremely close to Shorea multiflora (Burck) Sym.
but differs markedly in the size of leaf and fruit.

) Collections examined :— ©

BRITISH NORTH BORNEO: C.F. 36776 (S.H. 3908) (Type
of S. xanthophylla Sym.); Puasa; Bettotan, Sandakan, top
of hill; 23rd Sept. 1934; flower. Field notes—‘‘Vern. name,
Karai batu; tree 70 feet high, 18 in. girth; flower lemon
yellow.”’

C.F. 38742 (S.H. 4366), 38933 (S.H. 4557), 38938 (S.H.
4562), 41108 (S.H. 5496).

SARAWAK: For. Dept. Sar. 00277 (Paratype of fruit of S.
xanthophylla Sym.); Bogil; Sungei Babong, Beseri, hill;
6th August, 1935; mature fruit. Field notes—‘“‘tree 4o feet
high{:22 ims. gitth’’;

. Shorea xanthophylla is a ‘‘new’’ species which has been
recorded several times in the neighbourhood of Sandakan in
British North Borneo and once in Sarawak. Our only informa-
tion concerning the tree is that given on the herbarium labels of
native collectors. It appears to be a small to medium-sized tree
of low hills for which the following vernacular names have been
recorded in B.N.B.—selangan babi, selangan kuning, and karai
batu. Pisang-pisang has also been recorded on two occasions,
possibly on account of a supposed leaf resemblance to some
species of Anonacee. ‘Timber specimens have never been
collected but, from botanical considerations, there can be little
doubt that the timber should be classified as ‘‘white meranti
sub-group 2’’ (vide Desch, Mal. For. Rec. 12: 37). The specific
epithet, xanthophylla, refers to the characteristic yellowish colour
of the dried leaves. .

‘Shores sandakanensis Symington, sp. nov. Plate 25.

S. macroptere Dyer affinis, sed foliis majoribus minus
coriaceis, paniculis longioribus laxioribus, floribus mayjoribus,
alis fructus longioribus, nuce subgloboso dense tomentoso differt.

Branchlets terete, tawny-tomentose when young, later
becoming dark. Leaves oblong, acuminate at the apex (acumen
about 1.5 cm. long), truncate or rounded at the base, usually
about 18.0x6.5 cm. but varying from about 9.04.0 cm. to
23.0X9.5 cm., glabrous above, sparsely stellate-hairy on the
nerves beneath, usually drying yellow-brown on both surfaces;
midrib slightly depressed above, prominent striate and sparsely
hairy beneath; main nerves 13 to 15 pairs, inconspicuous above,
prominent beneath; nervules in fine curved parallel lines,
distinctly visible but not very prominent on either surface;

Vol. TX.\\620 38).

344

petioles 1.0 to 2.0 cm. long, rugulose, tawny-tomentose to black;
stipules oblong, less than 1.0 cm. long, caducous. Panicles
axillary and terminal, solitary, when axillary shorter than the
leaves; main branches rufescent-tomentose, pale tomentose
towards the ends; ultimate branchlets regularly alternating about
1.5 cm. apart, usually simple but occasionally bifurcated, 1.0 té
3.0 cm. long, bearing 3 to 7 (usually 5) distichous flowers.
Flowers ‘‘red, no odour’’ (38730), ‘‘olive green’’ (38839),
“lake green’’ (38778), about 5.0 cm. apart, sessile, about I.0 cm.
long in mature bud. Sepals sub-equal, ovate-rotundate, acute,
pale to silvery tomentose outside, glabrous within. Petals
linear-oblong to linear, tawny-villous on the portions exposed
in bud. Stamens 15, of three sizes; filaments flattened, broad at
the base, gradually narrowing upwards; anthers globular, cells
sub-equal; appendage to connective a curved awn about as long
as the anther (slightly shorter in large stamens), at first erect,
later deflexed. Ovary and_ stylopodium cylindric-conical,
glabrous in the lower portion, villous above; style terete,
glabrous, about half the length of the ovary and stylopodium;
stigma very slightly enlarged, obscurely 3-lobed. Fruit a
5-winged nut; sessile; bases of accrescent calyx lobes thickened,
woody, embracing the lower portion of the nut only, minutely
stellate-hairy; three outer wings sub-equal, linear, rounded at the”
apex, auriculate at the base, about 11.0 x 1.5 cm. to 14.0 X 2.0 cm.,
g to 11-nerved, sparsely minutely hairy, red when young (C.F.
41051); two inner wings much shorter, linear, irregular at apex,
about 3.00.5 cm. to 6.00.7 cm., about 5-nerved; nut ovate-
globular, rostrate, about 1.8 x 1.2 cm., densely pale tawny-tomen-
tose; rostellum about 3.0 mm. long, pale tawny-tomentose.

There are several Bornean forms, some as yet undescribed,
closely related to Shorea macroptera Dyer from the Malay Penin-
sula. So close, indeed, are they that some authors would
probably interpret them as variations of a polymorphous species.
One of these forms (S. Bailloni Heim) was actually united with
the Peninsula species by Brandis (Journ. Linn. Soc. 31: Qo
(1895) ). As herbarium material of these forms accumulates,
however, it becomes increasingly evident that the differences are
such as to justify the recognition of several distinct species. The
differences between S. sandakanensis and S. macroptera Dyer are
of degree rather than kind, nevertheless, the sum of these differ-
ences is so constant that I feel confident in separating them
specifically.

Collections examined :—

BRITISH NORTH BORNEO: C.F. 38730 (S.H. 4354) (Type
of S. sandakanensis Sym.); Agama; Kabuli Forest Reserve,
Sandakan, level ground; 16th March, 1935; flower. Field

Gardens Bulletin, S.S.

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Plate 26.

345

Notes—‘‘vern. name, seraya; tree, 100 ft. high, 60 in. girth;
flower red, no odour; this form of seraya has usually short
bole’’.

CE. 38760 (Sia. 4304), C.F. °38765 (S.H.: 4380), CF.
26770 (3.H1. 4402), Car, 30630 (5.11. 4463), CLE. 35893 (6. E1-
e197), <4. 35040. (Sea. 4570), C.F. 41051 (SH: 4676); C.F.
41068 (S.H. 4850: Paratype of fruit of S. sandakanensis
Sym.); Agama; Kabuli—Sepilok Forest Reserve, Sandakan,
level ground; 5th June, 1935; fruit. Field Notes—‘‘vern.
name, seraya; tree 4o ft. high, 32 in. girth; fruit greenish-
red’’.

? Creagh, s.n., July 1896.*

Shorea sandakanensis appears to be common in the
vicinity of Sandakan, on flat land or on low hills. The
majority of the collections have been named serayva, but one
is named seraya puteh, and one selangan bangar. It appears
to grow to great size, one tree of 200 ft. in height and 108 in.
girth being recorded.

“ Shorea Smithiana Symington, sp. nov. ‘Shorea sp. No. 21606,”’
van Slooten ex Merr. Plant. Elmer. Born. 205 (1929).

Plate 26.

S. rugose Heim foliorum ramulorumque habitu similis, sed
staminibus 22-26, antheris late oblongis, connectivorum appendi-
cibus filiformibus, stylopodio oblongo basi constricto distincta.

Branchlets rather coarse, dark, glabresent to fulvous scabrous
stellate-tomentose at the ends. Leaves broadly obovate to
elliptic-oblong, acute or shortly blunt acuminate at the apex,
truncate to cordate at the base, about 10.0 x 6.0 cm. to 17.0 X 8.5
cm., sparsely stellate-hairy and drying dull grey-brown above
fulvous-tomentose on the lower surface; midrib depressed and
stellate-tomentose above, elevate, thick, closely tomentose, and
furnished with coarse scabrous hairs beneath; main nerves 15 to
20 pairs, inconspicuous above, elevate, tomentose, and scabrous
hairy beneath; nervules faint and sparcely hairy above, in con-
spicuous parallel lines, tomentose and scabrous hairy beneath;
petioles 2.0 to 3.0 cm. long, rugulous, fulvous scabrid-tomentose;
stipules about 1.0 cm. long, oblong, subfalcate, scabrid-tomentose,
caducous. Panicles terminal and axillary, in solitary racemes
usually larger than the leaves (up to 23.0 cm. long), fulvous-
tomentose becoming paler at the branchlets; branchlets regularly
alternating, about 1.0 cm. apart, up to 3.5 cm. long, decreasing
i: length towards the end of the raceme, 3- to 8-flowered;
branchlets and flowers furnished with sub-ovate, obtuse, pale

_* I examined this specimen (in young fruit) at Kew and was of
opinion that it is this species, although the leaves have dried a rather
unusual purple-grey colour.

Vol. IX. (1938).

346

tomentose bracts and bracteoles about 7.0 mm. long. Flower»
secund, about 2.0 to 3.0 mm. apart, ovate-lanceolate in bud,
about 1.0 cm. long including the pedicel; pedicel 1.0 mm. long,
tomentose. Sepals sub-equal ovate-acute, the two inner slightly
thinner and sub-acuminate, tomentose outside. Petals linear-
oblong, blunt, silvery tawny-tomentose on the portions exposéd
in bud, pink (Smith), red (Kamis, 41071). Stamens 22 to 26,
of three heights; filaments broad and flattened at the base;
narrowing abruptly to the filamentous upper portion; anthers
broadly oblong, much shorter than the filaments, 4-celled, cells
sub-equal; appendage to connective filiform, about as long as
the filament. Ovary and stylopodium shaped like an hour-glass,
tomentose; style short, cylindrical, glabrous; stigma minute.
Fruit a 5-winged nut; stalk very short, about 2.0 mm. long,
sparsely stellate-hairy; bases of accrescent calyx lobes thickened
and woody, closely embracing the lower half of the nut, sparcely
stellate-hairy; three outer wings linear, obtuse at the apex, up to
about 15.0 2.0 cm., sparsely minutely hairy, 7- to 1o-nerved
with arched or sub-reticulate nervules; two inner wings similar
but about 34 as long as the outer wings and with fewer nerves;
nut ovate-conical, rostellate, about 2.0 x 1.4 cm., pale silky tawny-
tomentose; rostellum about 5.0 mm. long, sharp and slightly
curved.

This species resembles S. rugosa Heim in vegetative charac-
ters but it is quite distinct in details of floral morphology. I can
suggest no closely-related species. The stamens, ovary, and
stylopodium, are of the type seen in the Pierrea group of Hopea
but rare in Shorea. The same combination may be seen in
S. compressa Burck and S. mecistoptervx Ridl., but S. Smithiana
differs from these in the more numerous stamens and, taking into
consideration the sum total of their characters, I do not feel that
the relationship is very close.

Collections examined :—

BRUNEI: C.F. 30472 (Type of S. Smithiana Sym.); J.S. Smith;
Labi Hills, hillside 700 ft.; 25th Nov. 1934; flower. Field
notes—‘‘Tree 150 feet high, 12 feet girth; flower pink; bark
brown with very deep fissures; large buttresses’’.

C.F. 30524 (Paratype of fruit of S. Smithiana Sym.);
J.S. Smith; 23rd Feb. 1935; fruit from same tree as C.F.
30472.

C.F. 30538 and 39646.

BRITISH NORTH BORNEO: C.F. 36727, 30018 (S.H. AG42);
41070 (S.H. 4858), 41071 (S.H. 4860), 41080 (S.H. 4932);
Elmer No. 21606 (Sun Yatsen No. 56097).

This new Shorea was first collected in British North Borneo
by A. D. E. Elmer about 1922, but it was not until 1934-35 that

Gardens Bulletin, S.S.

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Plate 27.

347

adequate material was available for purposes of description.
During 1934-35 the species was collected in flower or fruit several
times in both British North Borneo and Brunei. From the
former territory the names urat mata, seraya, and seraya merah,
are recorded, and from the latter berat (Murut) and mengkabang
vambai (Dyak). It is evidently a good-sized tree, specimens
over 12 feet girth having been recorded; it produces a timber
apparently of the ‘‘light red meranti’’ grade.

I have named the tree after Mr. J. S. Smith, State Forest
Officer, Brunei, from 1933-1936, whose interest in the collection
of botanical material has resulted in invaluable contributions to
our knowledge ot Bornean trees.

7 Vatica diospyroides Symington, sp. nov. Vatica Scortechinii
(King) sensu Brandis in Journ. Linn. Soc. 31: 122 (1895),
sensu Craib, Flor. Siam Enum. 1: 141 (1925), et sensu V.
elm Bull. Jard. Bot. Buitenz.: UF, 9: 108 (1927), quoad
Curtis 2934; non Retinodendron Scortechinii King. Plate 27.

A V. Scortechinii (King) Brandis foliis tenuioribus, stipulis
eaducis, fructu majore, calycis lobis acuminatis recedit.

Branchlets pale, fuscous stellate-tomentose towards the ends.
Leaves oblong or linear-oblong, acuminate at the apex, acute at
the base, from about 12.0x 2.7 cm. to 27.08.25 cm., glabrous
and drying a pale yellow-green on both surfaces; midrib elevate
on both surfaces, particularly the lower, pale yellow; main nerves
about 15 to 2I pairs, prominent beneath, slightly less prominent
above, usually with 2 to 4 short intermediary nerves; nervules
reticulate, prominent on the lower surface; petioles 1.0 to 2.5 cm.
long, rather thick, rugulose above, fuscous-tomentose when
young; stipules not seen. Panicles axillary and terminal, solitary
or 2 to 3 together, racemose, 2.0 to 4.0 cm. long, fulvous flocculent
stellate-tomentose, about 3- to 12- flowered. Flowers lanceolate-
oblong in bud, about 1.5 cm. long (including the pedicel);
pedicel 2.0 mm. long or almost absent, fulvous flocculent
stellate-tomentose. Sepals all similar, slightly imbricate or
pseudo-valvate, deltoid-acuminate or ovate-acute, tomentose on
both surfaces. Petals slightly contorted in bud, oblong-obtuse.
densely: fulvous-tomentose on the portions exposed in bud,
“creamy white, very fragrant’’ (Kerr). Stamens 15 (rarely 13
or 14), of three heights, episepalous pairs alternating with solitary
intermediate stamens; filaments short, flattened, broad at the
base; anthers much larger than the filaments except in the tallest
row of stamens, posterior cells much shorter than the anterior;
appendage to connective short, thick, claviform. Ovary small,
conical, ferruginous-pilose; style about as long as, to twice as long
as, the ovary, ridged, glabrous; stigma sub-conical, ringed at the
base, obscurely 3- to 6-lobed. Fruit: stalk short, 2.0 mm. long,

Vol. .EX.0(21038 ).

348

fuscous-tomentose; sepals united at the base to form a thin
concave disc, iobes ovate, caudate-acuminate, margins sinuate,
about 1.5 x 0.75 cm., sub-verrucose with a sparse pale tomentum;
nut ovate-globose, rostellate, about 2.5 x 2.0 em., fulvous farina-
ceous-tomentose; rostellum slightly curved, about 3.0 mm. long;
pericarp thick, woody, splitting into three equal portions.

This is a very distinct species of Vatica which differs from
most species in the subgenus /sauxis in lacking recurved fruiting
sepals. The fruits in this respect more closely resemble those of
V. Scortechinti (King) Brandis and V. sarawakensis Heim than
any others, but the flocculent inflorescence is more suggestive
of the subgenus Synaptea. ‘The earliest collection was made by
Curtis in Pang-nga in 1893, but this was referred by Brandis to
Vatica Scortechinii (King). The latter, however, is quite distinct
in the broader and thicker leaves, the large persistent stipules,
and the smaller grooved fruits with obtuse calyx-lobes.

Collections examined :—

SIAM: Kerr t1oo1rr (Paratype of flower of V. diospyroides
sym.); A. F. G. Kerr; Bangkok, altitude 5 m.; 27th Jan.
1925; flower. Field notes—‘‘Local name, chan ta paw;
shrub about 3 m. high; cultivated for its fragrant flowers.”

Kerr 20546 (Type of V. diospyroides Sym.); leg. Heya
Wruit Waradern (?); Bangkok; roth July, 1931; mature
fruit. Field notes—‘‘Local name, chan kapaw; tree; culti-
vated for its fragrant flowers’’

Kerr 17045 (Takuapa), 18286 (Surat), 18527 (Pang-nga);
Curtis 2934 (Pang-nga); For. Dept. Siam 819.

This species is evidently a small tree which has been collected
wild in evergreen forest in Peninsular Siam. The type speci-
mens, however, came from Bangkok where the tree is said to be
cultivated for its fragrant flowers. The name Vatica diospyroides
is given because the fruits bear a distinct resemblance to those
of certain species of Diospyros.

Oxford-Borneo Expedition 1932—Additional Dipterocarpacez

Sherea mecistopteryx Ridl. in Kew Bull. 1925: 280; V.Sl. ex
Merr. in Univ: Calif. Pub. Bot. 15: 203 (1920).

Native Collector No. 2269; Dulit, under 300 m., primary
forest; Oct. 18th, 1932; flower. Field note—‘‘Urat Mata, tree,
39 m. high. lJLarge buttresses. Bark 0.8—1.0 cm. thick, reddish
in blaze, surface yellowish grey, smooth, flaking off in large thin
irregular pieces. Not the same species as the tree of same name
at;. Marudi... ‘Timber Class 1 (e)7

Shorea mecistopteryx has been known hitherto only from two
fruiting collections (Taha s.n. (the type) and Elmer 21569) from

Gardens Bulletin, S.S.

349

localities near Sandakan in British North Borneo. The leaf shape
and distinctive soft indumentum are so characteristic that I feel
justified in referring the Dulit collection to this species. Flower-
ing material from the type locality is, however, desirable to put
the matter beyond doubt. The following is a description of the
flowers of the Sarawak plant—

Panicles axillary, stellate-tomentose, ferruginous on the main
stem, grey on the branchlets, racemose, up to 12.0 cm. long;
branchlets simple or branched, about 1.0 to 4.0 cm. long, each
bearing 2 to 6 flowers subtended by sub-persistent bracteoles;
bracteoles oblong, obtuse, sparsely sericeous-tomentose outside,
glabrescent within. Flowers elongate-pointed in bud, up to
1.8 cm. long including the pedicel; pedicel 2.0 mm. long, pale
tomentose. Sepals tomentose outside, minutely so on the upper
half inside; three outer oblong, rounded at the apex; two inner
slightly shorter, acute at the apex. Petals linear-oblong, rounded
at the apex, pale tomentose outside on the portion exposed in
bud. Stamens 15, of two different heights, pairs alternating with
single stamens; filaments broad at the base, thickened above,
then narrowing abruptly to a short filamentous portion, three to
four times as long as the anthers; anthers rotund-ovate, cells
sub-equal in size; appendage to connective filiform, about 1 to 3
times as long as the stamen. Ovary with stylopodium shaped
like an hour glass (as in Pierrea group of Hopea), glabrous; style
short; stigma obscurely 3-notched.

On floral characters this species appears to be related to
S. pinanga Scheff. and S. compressa Burck, species characterized
by a similar ovary and long, filiform connectival appendages.

Van Slooten (op. cit. p. 202) has questioned whether
S. chrysophylla Ridl. (Kew Bull. 1926: 470) differs from S. mecis-
topteryx Ridl. The collection described above lends support to
Ridley’s view. Although the ovary and stylopodium in S. mecis-
topteryx are exactly similar to those of S. chrysophyila, the
indumentum of the leaves is quite distinct.

Cotylelobium malayanum V.S1. in Bull. Jard. Bot. Buitenz. IIT,
12, pt. 1: 43 (1932). Cotylelobium flavum sensu V. SI. in
op. cit. 10: 396, fig. 1 (1930), non Pierre, Fl. For. Cochinch.,
tab. 258A (1801).

P. W. Richards No. 1816; Ulu Koyan, Mt. Dulit, c. 900 m.,
white sand forest; 15th Sept. 1932; flower. Field note—‘‘Resak
peniau or resak durian. Tree c. 25-30 m. high, 33 cm. diam.
Flowers creamy white with strong, sweet, vanilla-like scent.
Anthers bright yellow. Leaves hard and leathery, dark green
above, light brown below. No buttresses. Bark stripping in
thick longitudinal flakes’’.

Distribution.—Malay Peninsula, Sumatra, and Borneo.

V oh.’ 1X .)'G0938)-

Plate

Plate

Plate

Plate

Plate

Plate

Plate

359

Explanation of Plates

17. Hopea Pierrei Hance. 1, flowering twig. 2, sterile twig
showing domatia on leaves. 3, flower bud (immature). 4,=3
with sepals removed. 5 & 6, outer sepals. 7, 8, & go inner
sepals. 10 & 11, petal from flower bud. 12 & 13, stamens.
14, ovary & stylopodium. 15, section through 14. 16, ripe
fruits. (Drawings of 1 & flower parts based on Pierre 1425;
2 rs Foxw. 8; 16 on Guibier 2). Scale applies to 1, 2, & 16
only.

18. Hopea Beccariana Burck. 1, flowering twig. 2, branchlet
of panicle with expanding flower & two buds. 3 & 4, outer
sepals. 5, 6, & 7, inner sepals. 8 & 9g, petal from mature
flower. 10, I1, 12, & 13, stamens from mature flower. 14,
flower with perianth parts removed to show ovary. 15, section
through ovary. 16 & 17, immature fruits. 18 & 19, mature
fruits. (Drawings of 1 & flower parts based on C.F. 9661 &
11673; 16) & 17,.0n C.F. ‘16574; 18, & 19 on.S’p. 3474).- Seale
applies ta 1, 16, 17, 18, & 109 only,

19. Hopea pedicellata (Brandis) Sym. 1, flowering twig. 2,
branchlet of panicle with expanding flower & two buds. 3 & 4,
outer sepals. 5, 6, & 7, inner sepals. 8 & g, petal from mature
flower. 10, 11, 12, & 13, stamens. 14, flower with perianth
parts removed to show ovary. 15, section through ovary. 16,
panicles with immature fruits. 17 & 18, mature fruits. (Draw-
ings of 1 & flower parts based mainly on Curtis 167; 16 on
C.F. 260137 17 & 18 on Sp.” 3474). “Scale apeives ip tae
17, & 18 only.

20. Shorea assamica Dyer forma globifera (Ridl.) Sym. 1,
flowering twig. 2, immature twig from sapling showing
stipules. 3, branchlet of young inflorescence showing bracts.
4, flower bud. 5-9, sepals. 10 & 11, petal from mature bud.
12 & 13, stamens. 14, ovary. 15, 16, & 17, stigmas. 18, stigma
partially disected. 19 & 20, fruits. (Drawings of 1 & flower
parts based on C.F;-.3072; 2 on. C.F. 27226%.3 on (CF. yesae:
19 & 20 on 39071). Scale applies to 1, 2, 3, 19, & 20 only.

21. Parashorea malaancnan (Blanco) Merr. 1, flowering twig.
2, flower bud. 3, 4, & 5, outer sepals. 6 & 7, inner sepals.
8 & 9, petal from mature flower. 10, flower—sepals & petals
removed. 11 & 12, stamens. 13, Ovary. 14, end of the style
showing stigma. 15, section through ovary. 16, immature
fruit. 17 & 18, mature fruit. (Drawings of 1 & flower parts
based on C.F. 39019; 16 on C.F. 30531; 17 & 18 on Elmer 217309).
Scale abflies \to..1,..16, 175 & 48 only.

22. Parashorea malaanonan (Blanco) Merr. var. tomentella Sym.
1, flowering twig. 2, flower bud. 3, 4, & 5, outer sepals.
6 & 7, inner sepals. 8 & g, petal from expanded flower. 10,
flower—sepals & petal removed. 11 & 12, stamens. 13, flower—
sepals, petals, & most stamens removed. 14, end of style &
stigma. 15, section through ovary. 16 & 17, young fruits.
18=17 with sepals removed. (Drawings of 1 & flower parts
based on.C.F. 38745; 17/00 C.F. 388455 26: on) C2 eaaee):
Scale applies to 1, 16, 17, & 18 only.

23. Shorea acuminatissima Sym. 1, flowering twig. 2, flowering
bud. 3, expanding flower. 4 to 8, sepals. 9 & 10, petal from
mature flower. 11 & 12, stamens from flower bud. 13, 14, & 15,

Gardens Bulletin, S.S.

Plate

Plate

Plate

Plate

Vol.

351

mature stamens. 16, flower—perianth removed. 17, section
through ovary. 18, immature fruit. 19 & 20, mature fruit.
(Drawings of 1 & flower parts on C.F. 38723; fruit on C.F.
41075). Scale applies to 1, 18, 19, & 20 only.

24. Shorea xanthophylla Sym. 1, flowering twig. 2, mature
inflorescence. 3, mature flower bud. 4, 3 with sepals & petals
removed. 5 to 9, sepals. 10 & 11, petal from mature bud.
12 & 13, stamens. 14, section through ovary. 15, stigma. 106,
fruit. 17, germinating fruit. (Drawings of 1 & flower parts
based on C.F. 36776; 2 on C.F. 38742; fruits on For. Dept.
Sarawak 277). Scale, applics fo. 1; :2,,.10,& 17 only.

25. Shorea sandakanensis Sym. 1, flowering twig. 2, flower
bud. 3 to 7, sepals. 8 & g, petals from bud. to & 11, petal
from expanded flower. 12, ovary—sepals & petals removed.
13, 14, & 15, mature stamens. 16, section through ovary. 17,
stigma. 15 & 19, mature fruit. (Drawings of 1 & flower parts
based on C.F. 38730 & 38778; fruits on C.F. 41068). Scale
applies to 1, 18, & 19 only.

26. Shorea Smithiana Sym. 1, flowering twig. 2, young
inflorescence with bracts. 3, flower bud. 4, expanding flower.
5 tog, sepals. 10 & 11, petal from mature flower bud. 12 & 13,
stamens. 14, Ovarv—sepals & petals removed. 15, section
through ovary. 16 & 17, fruits. (Drawings of flower parts
based on C.F. 30472; fruits on C.F. 30524). Scale applies to
he. 2, 20, C017 Onky.

27. WVatica dicspyroides Sym. 1, flowering twig. 2, mature
flower bud. 3 to 7, sepals. 8 & 9, petal from mature flower.
10, 11 & 12, stamens. 13, ovary—sepals & petals removed. 14,
section through ovary. 15, stigma. 16& 17, fruits. {Drawings
of 1 based on Kerr roori & 18577; flower parts on Kerr toort;
fruits on Kerr 20546). Scale applies to 1, 16, & 17 only.

EX ul 1038).

INDEX

Names of groups especially dealt with in this paper are shown in
bold type and synonyms in Zfalics.

Page

Anthoshorea Brandis (section) i. pt Et Pee oe
Balanocarpus Auct. nan oe 330
Balanocarpus coriaceus (Heim) ani <t 330
Balanocarpus longifiorus (Brandis) Foxw. ex

Sym. Me a meas fe 330
Balanocarpus maximus King ~~ we 330
Balanocarpus multifiorus (Burck.) Sym. * 330
Balanocarpus ovalifolius Ridl. if: 325, 326
Balanocarpus ovalifolius sensu Foxw. vs 325, 32¢
Balanocarpus pahangensis Foxw. ... =. 330
Cotylelobium flavum sensu V. Sl. ... as 349
Cotylelobium malayanum \. S|. a3 web 349
Dipterocarpus Gertn. f. hei as 310
Dipterocarpus angustialatus Heim Ke 321
Dipterocarpus Baudii Korth. i: =e 321
Dipterocarpus chartaceus S\m. uy tt 322, 323
Dipterocarpus gracilis Bl. ee ae 321-323
Dipterocarpus hirtus Vesque wa apes 322
Dipterocarpus malaanonan Blanco ed 334, 335
Dipterocarpus pilosus Roxb. =o a 321
Dipterocarpus pilosus sensu Parker iad 427i), S25
Dipterocarpus Skinneri King : 321-323
Dipterocarpus Skinneri King var. ‘ates Ridl. 321-323
Dipterocarpus vernicifinus Blanco ... - 321
Dryobalanoides Mig. (section) ss eae 319, 323
Eushorea Brandis (section) ea i 331
Hancea Beccariana Pierre P. at 325
Hancea Pierrei Pierre ... ve es 323
Hopea Roxb. oe ae a. 319, 331, 341, 340
Hopea Beccariana Burck ... = si 323, 325-327, 330
Hopea Dyeri Heim et ote rae 323, 324
Hopea Foxworthyi Elmer ts i 324
Hopea glutinosa Elmer ... be a 324
Hopea Griffthii Kurz... a x Fas 323, 324, 405
Hopea Griffithii var. pedicellata Brandis a: 327, 329
Hopea intermedia King ... nai ue 324-327
Hopea intermedia sensu Brandis ... = 327, 3290
Hopea intermedia sensu Foxw. se es 325, 326
Hopea malibato Foxw. ... ie 42 324
Hopea mengarawan sensu Brandis as 327, 329

Gardens Bulletin, SS.

Hopea micrantha Hance

Hopea micrantha Hook. f. LA a. 323,
Hopea micrantha sensu King

Hopea micrantha sensu Ridl.

Hopea microptera Dyer {(Ms.)

Hopea Nicholsoni Heim

Hopea pedicellata (Brandis) Sym. _..... r7 324, 326,

Hopea Pierrei Hance

Hopea Pierreit sensu Auct.

Hopea Pierrei sensu Brandis

Hopea Pierrei sensu Foxw. se -? 0 SBSs
Hopea Pierrei sensu Ridl.

Isauxis (Arn.) Brandis (subgenus) ...
Mocanera malaanonan Blanco

Parashorea Kurz

Parashorea sp. 4
Parashorea lucida (Miq.) ieee
Parashorea malaanonan (Blanco) Merr.

Parashcrea malaanonan var. tomentella Sym. Sa z 337

Parashorea plicata Brandis

Parashorea stellata Kurz eae ae

Parashorea Warburghii Brandis ... - 334,
Pierrei (Heim) Sym. (‘‘group’’)

Retinodendron Scortechinii King

Richetia Heim (genus) om a

Richetia (Heim) Sym. (‘“‘group’’) ... < 330-331,
kichetioides Heim (section)

Shorea Geertn. f. Ps, = uae “Rh: S5ks 925;

Shorea acuminatissima Sym. ce ee 330,

Shorea assamica Dyer *
Shorea assamica Dyer forma assamica Sym.

Shorea assamica Dyer forma slobifera ead)
Sym.

Shorea assamica Dyer Steet ceaciecus fcaadss)
Sym. :

Shorea assamica ea forma philippinensis
(Brandis) Sym. : —* 2

Shorea Bailloni Heim

Shorea balanocarpoides Sym.

Shorea blumutensis Foxw.

Shorea chrysophylla Ridl.

Shorea compressa Burck

Shorea Faguetiana Heim

Shorea Forbesii Brandis

Vol. IX. (1938).

Page
323
327, 320
327, 328
326, 327
322
325, 327
327-330
323-325
323-325
323-325
325, 320
323-327
348
334-330
335
338, 339
335

334-330

335
335; 337
336, 349

347
341, 343
341, 340

341-242
331-334

346, 349
330, 341

Shorea gibbosa Brandis

Shorea globifera Ridl.

Shorea hopeifolia (Heim) Sym.
Shorea kalunti Merr.

Shorea Koordersii Brandis
Shorea longiflora (Brandis) Sym.
Shorea macroptera Dyer

Shorea malaanonan (Blanco) Bl.

Shorea malaanonan sensu Merr. et Me

Shorea maxima (King) Sym.
Shorea mecistopteryx Ridl.
Shorea. multiflora (Burck) Sym.
Shorea pallida Foxw.

Shorea philippinensis Brandis
Shorea pinanga Scheff.

Shorea polita Vidal

Shorea resina-negra Foxw.
Shorea Richetia Sym.

Shorea rugosa Heim

Shorea sandakanensis Sym.
Shorea Smithiana Sym.

Shorea xanthophylla Sym.
Synaptea (Griff.) Brandis ieee

Vatica celebica Koorders (Ms.)
Vatica diospyroides Sym.

Vatica Sarawakensis Heim 3
Vatica Scortechinii (King) Brandis

Page
330
332-334
339
330
331, 332, 334
330
343. 344
334, 335
334
330
346, 348-349
330, 342, 343
331-333
331-334
349
335

Gardens Bulletin, S.S.

355

A REDEFINITION OF THE GENUS TERATOPHYLLUM

By R. E. Houttum,
Botanic Gardens, Singapore

When Kuhn published the original diagnosis of the genus
Teratophyllum in 1869 he included in it the bipinnate species
previously known as Polybotrya articulata J. Sm. and Polybotrya
Wilkesiana Brack. These differ notably in several characters
from the other species of the genus; so much so that when !
prepared an account of Teratophyllum in 1932 (this Bulletin,
vol. 5, pp. 277-304) I excluded them as possibly referable to a
distinct genus. Copeland has included them in Lomagramma,
but all known species of Lomagramma have characteristic features
of spores and venation which are quite different from those of the
two species in question (this Bulletin, Vol. 9, p. 104). I have
now examined a number of specimens, and have come to the
conclusion that the best treatment is to unite them to Tera-
tophyllum as proposed by Kuhn. His diagnosis however omits
a number of important points, and I have prepared a new one
which includes what seem to me the essential features of the
genus. The genus as so defined consists of two sections. The
first and larger section comprises those species dealt with in my
former paper; the second section has hitherto not received
adequate description, and I give below as full an account of its
species as the material at present available allows.

I am grateful to the authorities in charge of various herbaria
for the loan of specimens, or for permission to examine speci-
mens. ‘The herbaria in question are cited as follows: Brisbane
(Br.) British Museum (B.M.), Buitenzorg (B.), Kew (K.), Philip-
pine National Herbarium (M.), Singapore (S.), U.S. National
Herbarium (W.).

TERATOPHYLLUM Mett., emend. Holttum

Rhizoma paleis peltatis adpressis fugacibus squamosum, alte
scandens, dorsiventrale, frondium duos vel plures ordines ferens.
Frondes prope solum (bathyphylla) minores, multiformes; frondes
superiores (acrophylla) majores, pinnate vel bipinnate. Pinne
acrophyllorum omnes articulate pinnule articulate apicales
exceptee; venulee pinnarum vel pinnularum sterilium libere,
simplices vel furcatee, ante marginem terminate, margines
integree vel serrate, pinne vel pinnule fertiles plus minusve
angustatee, seepe lineares, semper acrostichoideze, nervatio earum
plerumque venulis irregularibus supernumerariis seepe anastomo-
santibus prope paginam inferiore aucta; paraphyses nulle; spore
perisporio vestitz.

1. Euteratophyllum. Rhizoma tenue, frondes semper
biseriatze, acrophylla semper pinnata. Species typica: Lomaria
aculeata Bl.

Vol. IX. (1938).

350

2. Polyseriate. Rhizoma adultum crassius, frondes poly-
serlatee, acrophylla sterilia et fertilia seepe bipinnata. Species
typica: Polybotrya articulata J. Sm.

The section Euteratophyllum is confined, so far as at present
known, to the Malaysian region, inclusive of the Philippines and
New Guinea. The section Polyseriatze does not occur west of
Celebes, but extends into the Pacific as far as Tahiti, also north-
wards throughout the Philippines and southwards to Queensland.

Key to the section Polyseriatx
Fertile pinnee hardly more than 1 mm. broad T. Brightie

Fertile pinne much broader, or fertile fronds bipinnate
Sterile pinnules auricled on upper base T. articulatum

Sterile pinnules not auricled T. Wilkesianum

Teratophyllum articulatum (J. Sm.) Mett. Kuhn in Ann. Mus.

Lugd=> Bat. 42° 207. 1860.

Polybotrya articulata J. Sm. (nomen nudum) Journ. Bot. 3:
AOI. 1841.

Fée, Hist; Acrost. 74, t.37.°1848:

Lomagramma articulata Copel. Phil. Journ. Sci. 3C: 32.
1908.

Lomagramma bipinnata Copel. Phil. Journ. Sci. r1C: 114.
IgI6.

The type of this species is Cuming 296, from Leyte. I have
seen specimens of this collection at Kew and from the U.S.
National Herbarium. I have also examined the specimens listed
below. I have not examined the type of Copeland’s Loma-
gramma bipinnata, but from description it seems to fall well
within the range of variation of the present species. The range
of variation is considerable, and of the same character in widely
separated countries. The essential features of the species may
be summarised as follows.

Rhizome to 1 cm. diameter, longitudinally ridged, at first
covered with small peltate lanceolate almost black scales with
brown more or less fimbriate edges, smaller scales entirely brown.

Stipes to at least 25 cm. long; main rachis not winged, but
2-grooved above, scaly when young.

Fronds to about 80 cm. long and 30 cin. wide, with abouc
15 pairs of pinnee; basal and apical pinnee somewhat smaller than
the rest.

Pinne to 20 cm. long and 3.5 em. wide, with stalks up to
1 cm. long; pinnules about 20 pairs; pinna-rachis winged, the
wing interrupted at the base of each pinnule and expanded into
a narrow lamina at the apex of the pinna.

Gardens Bulletin, S.S.

ome

Sterile pinnules on short winged stalks about 1 mm. long,
variable in size and shape, commonly about 18 mm. long and
7 mm. wide in the middle; upper base truncate, usually 7-8 mm.
wide, forming a prominent auricle, sometimes a distinct basal
lobe; lower base narrowly cuneate; both edges (except the base)
obliquely toothed, the teeth not more than 1 mm. deep, one to
each vein-ending; apex rounded or bluntly pointed; veins mostly
forked, except the vein in the basal auricle or lobe which is
pinnate; surfaces of fully expanded frond naked except for a few
scales on the lower surface, mostly on the midrib, the larger
scales sac-shaped.

Fertile pinnules on winged stalks 2 mm. long, about 4-7 mm.
long and xr mm. wide when dry, the upper base distinctly
auricled, the apex rounded.

Young stages. One specimen examined shows the young
stage of the plant, namely Clemens 1ro1r8 from Mindanao. The
rhizome is very slender and has simply pinnate fronds to about
12 cm. long and 2.5 cm. wide; the fronds each have about 10 pairs
of pinnee and an apex consisting of a narrow lamina continuous
with the rachis. The stipes are not more than 1 cm. long, and
in all cases there are one or two basal unpaired pinnz on the
side of the rachis towards the base of the plant, the side of the
rachis towards the apex of the plant being devoid of pinnz (this
is in agreement with some species of Euteratophyllum). The
pinnee are up to 12 mm. long and 7 mm. wide, the general shape
like the pinnules of the acrophylls but the veins fewer and the
teeth larger and rounded. ‘The lower surfaces bear sac-shaped
scales like those of the acrophylls.

This specimen evidently represents a young stage, and we
have no definite information as to whether the fronds on it are
normal bathyphylls, or whether such may be bipinnate on larger
rhizomes. The resemblance to T. aculeatum is striking, but the
shape of the pinnz and the scales are characteristic of T. articu-
latum, and there can be no doubt that reference to the present
species is correct.

A specimen from Celebes (part of Biinnemeijer 11138) may
perhaps represent more normal bathyphylls. The fronds are
small, bearing pinnules of hardly more than half the usual size,
and are attached to a slender rhizome. In the same collection
are fronds of normal size with larger pinnules.

Distribution: from Celebes and the Philippines through New
Guinea to the Solomon Islands.

PHILIPPINES. Leyte: Cuming 296 (type collection, K.,
W:); Dasami, C. A. Wenzel 89 (M.).-.Luzon: Mt. Masingit,
Lubuagan, Kalinga Subprov., Bur. Sci. 37500, Ramos and Edano
(M., B.); Caraballo Mt., Nueva Vizcaya Prov., Bur. Sci. 13730,
13733, 13716, Loher (M.). Mindanao: Camp Keithley, Lake
Lanao, M.-S. Clemens 1073 (M.), 1018 (M.), s.n. (B.).

Vol. t5;- Ci0%8).

358

CELEBES. Lombasang, N. W. Celebes, Biinnemeijer
11138 (B.). Papepekang, Bonthain, Teysmann 13688 (B.). Mt.
Bonthain, S. W. slope, 1500 m., Posthumus 2720 (S.). Prov.
Minahassa, Koorders 17073 (B.), 17056 (B.), 17032 (B.).

CERAM. 1000-1400 m., Rutten 2213 (B.).

NEW GUINEA. Nassau Geb. 7oo m., Docters van
Leeuwen 10680 (B.). Boridi, Papua, 4,800 ft. Carr. 14212 (S.).
K. Wilhelmsland, Wapao, 600 m., Bamler (Rosenstock, Fil.
Novoguin. 122, M.).

SOLOMON ISLANDS. Bougainville, 1600 m., Kajewski
1711 (M.), 1200 m., Kajewski 2669 (M.).

Teratophyllum Brightiz (F. von. Muell.) Holttum comb. noy.
Plate 28.

Acrostichum Brightie F. von Mueller, Fragmenta 7: 1109.
1870.

The original description of this species is as follows :—

Alte scandens, frondibus simpliciter pinnatis, rachi aptera, fzmzis
cum rachi articulatis, sterilibus fere conformibus membraneis
perangusto-vel falcato-lanceolatis comspicue petiolulatis ad bastim
fere cuneatam inequilateris tenui-acuminatis irregulariter repando-
crenulatis subtus pallide virentibus, venis tenuibus multis inferme
longiuscule simplicibus superne dicladis, aficibus venarum fere
clavato-incrassatis ante marginem evanescentibus, rachi frondis
fertilis squamulis angustis acutis crispulis conspersa, pinnis anguste
linearibus remotiusculis.

In monte Grahami prope Rockingham’s Bay; J. Dallachy.

Caudex ad altitudinem 20’ scandens, angulosus, superne semi-
pollicem crassus. Rachis frondis sterilis fere nuda. Pinnz steriles
elt le 4-7" late; fertiles vix semilinea latiores. Sporangia
avida.

By the courtesy of Mr. C. T. White, I have examined a
number of specimens from the Queensland Herbarium (Brisbane),
and can add the following particulars to von Mueller’s
description.

Rhizome. ‘The acrophyll-bearing rhizome may have at least
four leaf-gaps. The bathyphyll-bearing rhizome has usually two
leaf-gaps as in Eu-teratophyllum. One specimen shows a stout
old rhizome bearing bathyphylls on young lateral branches;
apparently this rhizome had fallen and was starting new growth
again from its fallen position. |

Bathyphylls. These are fully represented in the Brisbane
collections. They vary a good deal in size, the largest approach-
ing the smallest acrophylls in size of pinne. They differ
constantly from the acrophylls in (1) the apex of the frond,
which consists of a narrow toothed lamina, its midrib continuous
with the rachis, and (2) the broadly truncate or even subauriculate
upper base of the pinne. The smaller bathyphylls show a
decided resemblance to the young stage of T. articulatum
described above, or to pinne of acrophylls of T. articulatum.

Gardens Bulletin, S.S.

Gard. Bull. S.S. Vol. IX. Plate 28.

Teratophyllum Brightig. Bathyphylls, 2 nat. size. Eumundi,
Shirley, U.S. National Herbarium.

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Gard Bull. Ss. Vol. 1 Plate 30.

Teratophyllum Wilkesianum. Fertile frond showing condition
intermediate between pinnate and bipinnate. Tahiti, Vesco,
Manila Herbarium. #2 nat. size.

359

The stipes vary in length, but are commonly 3 to 5 cm. long.
The pinnze are about 20 pairs, the lowest:one or two usually
unpaired. The middle pinne have winged stalks 1-2 mm. long,
the upper base broadly truncate, the lower narrowly cuneate,
the edge obliquely toothed, one vein-ending to each tooth, the
apex rounded in the lower pinnz, more or less acute in the upper
ones, veins mostly forked (the lowest on the acroscopic side often
twice forked); size of middle pinnz of average bathyphylls about
2 cm. long by 5 mm wide. The rachis is winged almost
throughout, the wing produced to a tooth below each pinna.

Sterile Acrophylls. All pinnz are articulate; the pinna
stalks 2-3 mm. long; the upper base sometimes very broadly
cuneate, approaching the bathyphyll condition; the apex rather
suddenly narrowed to a toothed subcaudate tip; the margins
toothed very much as in simply pinnate specimens of T.
Wilkesianum from New Caledonia and Samoa.

Fertile acrophylls. Pinna-stalks to 5 mm. long, pinne to
14 cm. long, usually not more than 1 mm. wide when dry; one
specimen has pinnz almost 2 mm. wide.

Distribution: Queensland, extending southwards just beyond
the tropic.

Specimens examined. Eumundi, Bailey and Simmonds 1894
(several specimens, Br.), J. Shirley (W.). Palm Island, F. L.
Bancroft (Br.). Herberton, F. M. Bailey (Br.). Between
Herberton and Cairns, C. J. Wild 1891 (Br.). Range Trinity
Bay, F. M. Bailey (Br.). Atherton Tableland, W. D. Francis,
May 1928 (Br.). Scrub, Cairns District, Watts, July 1913 (Br.).
Johnstone River, W. A. Kefford (Br.). Covering the branches
of trees, below Bon Accord Falls, Blackall Range, C. T. White,
Ap. 1918 (Br.).

§ Teratophyllum Wiailkesianum (Brack.) Holttum comb. nov.
Plates 29, 30.

Polybotrya Wilkesiana Brack. in Wilkes, U.S. Expl. Exped.
16: 80, pl. ro. 1884.

Lomagramma Wilkesiana Copel. Phil. Journ. Sci. 3C: 32.
1908. Maxon, Univ. Cal. Publ. Bot. 12: 28, pls. 3, 4. 1924.

The type of this species is in the U.S. National Herbarium,
and has been kindly lent to me for inspection. It differs rather
strikingly from all other specimens I have seen from Tahiti and
irom New Caledonia, so that it seems to me desirable to give a
description first of the type only, and then to add some notes
on the points in which other specimens differ. It is clear that
Tahiti ferns which are referred to this species are very poly-
morphic, and more field study will be meeded before we can
understand them at all adequately. For the present I retain all
under this species.

The chief characters of Wilkes’ specimen are as follows.

Vol. IX. (1938).

360

Rhizome 1 cm. diameter, vascular cylinder with 8 leaf-gaps.

Stipe 20 cm. long, stramineous, grooved on adaxial side,
naked except for minute scattered dark narrow peltate scales.
Rachis stramineous, when young scaly above with very narrow
pale scales, naked when old, 2-grooved on the adaxial side, not
winged but the outer margins of the grooves shortly papillate
at the base of each pinna.

Pinne apparently about 12 pairs (rachis broken), all articu-
late to rachis, uppermost somewhat smaller than rest. Sterile
pinne to at least 15 cm. long and 3.5 cm. wide, pinnate, lowest
pinnules 8-15 mm. from main rachis; pinnules to about 18-jugate,
about 7-9 mm. apart; pinna rachis distinctly winged, the wing
broken and prolonged into an auricle at the base of each pinnule.

Sterile pinnules sessile or with a very short winged stalk, to
20 mm. long and 5 mm. wide, the base narrowly unequally
cuneate, the apex falcate acute, the margins with oblique teeth
about 1 mm. deep; veins 6-8 pairs, those on the acroscopic side
all forked, on basiscopic side some at least simple, each vein
ending close to the apex of a tooth; texture thin but firm; colour
dark above, paler beneath; surfaces naked except for a few
minute scales; the distal pinnules gradually reduced and adnate,
merging into the apex of the pinna.

Fertile pinnules to about 18-jugate, distinctly stalked (stalks
about 1 mm. long), 15-20 mm. long and hardly more than 1 mm.
wide when dried, the base almost equally cuneate, the apex
rounded. The venation of the fertile pinnules shows a very
remarkable additional vascular supply to the sporangia, consisting
of two rows of closed areoles, almost completely superseding the
normal venation. ‘There are indications of this in other fertile
fronds, notably Franc 662 from New Caledonia, but in none
that I have examined is the additional vascular system anything
like so fully developed.

The other specimens differ from the type in the following
points.

The sterile pinnules are usually a little shorter and wider,
the upper base more broadly cuneate and the apex less falcate
and less acute. The upper base is especially broad, almost
truncate, in some specimens from New Caledonia, and in these
the first vein is twice forked, approaching the condition found in
T. articulatum. ‘The marginal teeth are usually closer than in
the type, but not appreciably deeper, except in some specimens
which appear to be from young plants, or are perhaps bathy-
phylls; in these (both from Tahiti and New Caledonia, but
especially the latter) the margins are deeply incised between the
main lateral veins.

The fertile pinnules are always unequally cuneate at the
base, sometimes pronouncedly so, and nearly all are shorter than
the type, some barely 5 mm. long. The largest fertile pinnules
I have seen on any specimen except the type are on Franc 662

Gardens Bulletin, S.S.

361

from New Caledonia (specimen at Bititenzorg), on which they are
as much as 15 mm. long.

Simply pinnate fronds occur on some specimens, from both
Tahiti and New Caledonia (also Samoa), both sterile and fertile.
Maxon, 1. c., discusses these fronds and gives references to
former literature. There can be no doubt that both bipinnate
and pinnate fronds occur on the same plant. Setchell and Parks
collected a frond showing an intermediate condition (their
no. 512, figured by Maxon), and there is a specimen in the Manila
herbarium (Vesco 1847) showing a similar intermediate condition
in a fertile frond. It seems that the pinnate condition is some-
times but not always attained by plants of this species, and that
the bipinnate condition is more usual. Setchell and Parks report
the two as associated together. It is evident that the bipinnate
fronds are not merely bathyphylls, as Setchell and Parks record
them on a plant climbing high trees; and also there are bipinnate
fertile fronds. It is probably significant however that there are
no small pinnate fronds, and it is likely that this form is only
developed on high climbing stems.

Pinnate sterile fronds. Pinnee about 12-jugate, stalked,
stalks of lower pinne to about 7mm.long. Pinnze of Setchell and
Parks 512 to about 10 cm. long and 2 cm. wide, base unequally
cuneate (broader in lower pinne than in upper), gradually
narrowed from widest part just above base to near apex, apex
rather suddenly narrowed to a linear portion about 1 cm. long,
edges irregularly toothed, teeth short and not acute, one to each
vein-ending, the margins slightly cartilaginous and reflexed;
veins close, oblique, usually once forked. Pinne of simply
pinnate fronds from Samoa and New Caledonia to 15 cm. long,
the bases on the whole more broadly cuneate (especially in the
upper pinnze) and the edges rather more deeply toothed.

Pinnate fertile fronds. Pinna stalks to 7 mm. long; pinne
to ro cm. long and 5 mm. wide, base unequally cuneate, venation
as in sterile pinnz.

As above noted, the specimens from New Caledonia differ
in some points from Tahiti specimens, but I do not think that
the differences are clear enough to warrant the description of a
new species. It is very remarkable that though specimens have
been collected in Samoa, none have been reported from Fiji,
which is nearer to New Caledonia. The Samoa specimens appear
to be nearer to those of New Caledonia than to those of Fiji.

Y TAHITI. U.S. South Pacific Exploring Exp., Wilkes
(Type, W.). Setchell and Parks 211 (W.), 294 (W.), 206 (B., S.,
Weyer iaw W.), Sao tw..). -Grant 4450, (8., WJ). M; Vesco.s:n.
1e47, dist. ‘Paris (M.). N. J: Andersson s.n. '1852,, © dist.
Stockholm (W.).

NEW CALEDONIA. Franc 662, herb. Bonati (B., W.).
Rosenst. Fil. Nov. Cal. exsic. 16, leg. Franc (M., W.) and 113,

Vol. IX. (1938).

362

leg. Franc (W.). Cribs s.n. rg01, dist. Paris (M.). Richards
s.n. 1863, dist. Sydney (W.).
SAMOA. West Savali, Reinecke 87b (B., W.).

Doubtful Species
Lomariopsis Balanse Fourn. Ann. Sci. Nat. V. 18: 271. 1873.

This is described as a simply pinnate species, and there is
nothing in the original description which gives any definite
information as to which of the genera Lomariopsis or Terato-
phyllum is concerned. There is however a specimen at Kew
(Franc 663, Herb. Bonati) which was determined by Christ as
L. Balanse; this is very close to Teratophyllum Wilkesianum,
and apart from shorter stalks of the pinnz I could not see any
definite distinction. The spores have a finely granular perispore.
In the absence of further information this must be regarded as a
doubtful species. Examination of the type to see whether or not
the terminal pinna is articulated would at least decide the genus.

Discussion on the Polyseriatze

As indicated in the diagnosis, the species of this section of
the genus agree together in having the fronds on the climbing
rhizome polyseriate, not biseriate as in Euteratophyllum. It is
probable also that they have certain distinctive points in common
as regards their bathyphylls, but the information at present
available is incomplete. There appear to be no other constant
features in which all of them differ from Euteratophyllum. The
bipinnate acrophylls of T. articulatum and T. Wilkesianum are
certainly different from anything in Euteratophyllum, but the
simply pinnate acrophylls of T. Wilkesianum and T. Brightie,
it parted from their rhizome, could not be distinguished from
Kuteratophyllum. T. Brighti@, with its complete lack of all
bipinnate fronds, is closely similar to Euteratophyllum except
in its bathyphyls.

The bathyphylls of T. Brightie@ (and probably also those of
71. articulatum, judging from a very young stage) differ from
those of all species of Euteratophyllum in having distinct stipes
free of pinnze; all Euteratophylla have the lowest bathyphyll
pinna right against the rhizome and reflexed to overlap the
rhizome. Most Euteratophylla also have much more deeply
dissected bathyphyll pinne, though a few are simply pinnate.

As regards the general question of the relationship of
Teratophyllum to other fern genera, I cannot give any opinion
beyond the suggestions made in my former paper. It seems
probable that Teratophyllum and Lomariopsis are closely related,
but Teratophyllum is a much more specialised genus in many
ways, with a more restricted distribution. I think that they have
probably had independent origins in a group of acrostichoid
ferns with short-creeping rhizomes; possibly from Bolbitis.

Gardens Bulletin, S.S.

THE GARDENS’ BULLETIN STRAITS SETTLEMENTS

INDEX TO

VOLUME IX

New Binomials are printed in bold faced type and synonyms in

italics.

ERRATUM :—For Lomagramma borneensis on plate 8, read Lowma-

gramma Brooksit.
Absolute synonyms, 233.

Acrostichum Blumeanum HK.,
204+ | rae P
Acrostichum Srighti@ F. VON

MUELLER, 358.
Acrostichum cultratum BAk., 203.
Acrostichum speciosum WHILLD.,
110.
Actinophleeus cuneatus, 233.
Agarics in Malaya, The seasonal
fruiting of, 709.
Aglaonema, Index, 284.
Agriculture, Mr. RIDLEY’S work
for tropical, 31.
Alternative names, 30.
Amomum terminale, 290.
Amylotheca, Index, 309.
Andreoskia, 252.
Andropogon amboinicus, 297.
Andropogon Bequaertii, 230.
Andrzeiowskia, 252.
ANONYMOUS :—-
Dedication of the Bulletin to
H. N. RIDLEY, 1

List of plants named in
honour of Mr. H. N.
RIDLEY, 29

Mr. RIDLEY and Forestry in
Malaya, 42
Antitypes, 289
Apotype, 288 ; 299
Arenga Griffithii SEEM, 260
Arum, Index, 284
Aspidium, 121; 137; and 138
Aspidium ciliatuin WALL., 129
Aspidium glandulosum BUL., 127
Aspidium Keckii LUERSS., 137
Aspidium lineatum BL., 128
Aspidium sorbifolium WILLD.,
220
Aspidium
Asplentum
119
Asplenium glaucophyllum v.A.
¥:R., 119
Asplenium subnormale COPEL.
121

stipellatum BL., 130
polystichoides BL.,

’

Vol. [X. (1938).

Asplenium subscalare v.A.V.R.,
121

Asterisks in LINNAEUS’S Species
Plantarum, 310

Athyrium macrocarpum
BEDD., 121

Athyrium nigripes, (BL.) MOoORE.,
121

Augusta, 252

Augustia, 25

(BL.)

BACKER, C. A.; Semeiocardron
ZOLL, a misinterpreted genus of
Balsaminacez, 70

Balanocarpus, Index, 352

Basinym, 233

Bats, the dispersal of plants by
fruit-eating, 58

Bolbitis diversifolia (BL.) SCHOTT,
121

Bolbitis Quoyana (GAUD.) CHING,

122

Bolbitis subsimplex (FEE) CHING,
121

Borassus Machadonis, 305

Botanical Institutions under Art.
36., 273

Bruguiera, Index, 242

Calamus, Index, 181
Calamus inopinatus FURTADO, SP.

nov., 184

Calamus Steenisii FURTADO, sp.
noy., 182

Calandrinia polyandra BENTH.,
235

Calla silvestris BL., 234
Calophyllum, Index, 284
Campium Matthewit CHING, 206
Cerbera, Index, 284

Cheilanthes varians (WALL) HK.,
122

Chilolepton Blumeanum FEE, 204

364

Chonemorpha :

assamensis | URTADO Sp. NOV.,

115

Blancoi MERR., 116

elliptica MERR. & ROLFE, 116

fragrans (MOON) ALSTON, 115

macrophylla G. DON., 115

macrophylla var. grandis

A.D 895

mollis Mig., 116

penangensis RIDL., 116

Rheedei RIDL., 115
Clastotype, 288; 304
Clerondendron, Index, 284
Clinostigma, Index, 309
Clonotype, 288; 304
Colocasia, Index, 284
Commentary on the laws

botanical nomenclature, 223
Conocephalus, 252
Conocephalum, 252
Conophytum HAw, 231
CORNER, E.J.H.: The
fruiting of Agarics in

of

seasona!
Malaya,

79

Cotylelobium, Index, 352

Cotype, 293

CROIZAT, LEON: An early tertiary
relict in Malaya,
Euphorbia Ridleyi
nom. nov. descr.
emend. (£. syna-
denium Ridl.}
and its position
in the Linnean
genus, 145

Cucubalus, Index, 284

Cyclopeltis crenata (FEE) C.
CHR., 122
Cymbopogon Bequaertii DE

WILLD., 239
Cyphokentia samoensis, 291
Cyrtophyllum? lanceolatum Dc.,
230
Cystopteris
METT., 122

tenuisecta (BL.)

Dzmonorops angustispathus FUR-
TADO Sp. NOV., 161
Dzmonorops Curtisii FURTADO spec.
nov., 164
Dzemonorops, Index, 181; 309
Dzemonorops javanicus FURTADO
Sp. MOV., 170
Dzmonorops sepal yar. pahangensis
FURTADO Var. nOV., 179
Davallia dimorpha HoOLtTTuM sp.
nov., 122
Dendrobium :
Paathii J. J. SM., go
singkawangense |. J. SM. sp.
nov., g!

Desmos, Index, 309

Diciocephala chrysanthemifolia
(pl. 1), 69

Diplazium atrosquamosum, (Co-
pel.) C. Chr. 124

Diplazium insigne HOLTTUM
nov., 123

Diplazium latisquamatum HoLTTUM
sp. nov., 124

Diplazium subintegrum HOLTTUM
Sp. mOVv., 125

Diplazium tomentosum BL., 126

Diplazium velutinum HOLTTUM sp.
nov., 126

Dipterocarpaceze
Malayan, 319

Dipterocarpus chartaceus Sym. nom.
nov., 322

Dipterocarpus, Index, 352

Dipterocarpus oblongifolius, BL.,
The epiphytic flora of, 93

Dispersal of plants by fruit-
eating bats, 58

DOCTERS VAN LEEUWEN, W. M.:

IV, Notes on

The dispersal of plants by
fruit-eating bats, 58

Dryopteris brunnea (WALL.}
C. CHR. var. glabrata
(CLARKE), 127

Drvopteris crenata (FORST).
O. KTZE, 127

Dryopteris dissecta (FORST.)
O. KTZE, 127

Drvopteris glandulosa (BL.)
O. KTZE, 127

Dryopteris iridescens V.A.V.R.,
127

Dryopteris lineata (BL.) C. CHR.,
128

Dryopteris malayensis C. CHR.,
127

Dryopteris multisora C. CHR.,
128

Dryopteris oppositipinna v.A.
v.34

Dryopteris Parishii (HK.) O.
KTZE, 129

Dryopteris polita ROSENST., 129

Dryopteris pseudocalcarata C
CHR., 129

Drvyopteris sparsa (Ham.) O.
KTZE, 130

Dryopteris stipellata (BL.) O.
KTZE, 130

Duplicate types, 28

EATON, :
Rubber, 39

Echites fragrans MOON, 115

Echites macrophylla ROXB., 115

Effectiveness of Advance sepa-
rates, 271

Eleocharis: vide Heleocharis.

Mr. RIDLEY and

Gardens Bulletin, S.S.

365

ENDERT, Dr. F.H.: Intensive
system of collecting plants, 303

Epiphytic flora of Dipterocarpus
oblongifolius, 93

Epipremnum humile HOOK.F.,
230

Equivalent synonyms, 233

-ragrostis, Index, 309

Euphorbia Ridleyi CROIZAT nom.
nov., 147

Euphorbia synadenium RIDL.,
145

Eupomatia Belgraveana F.
MUELL., 231

Exact synonyms, 233

Expressions, nomenclatorial, 262

Fagrea fragrans, flowering of,
in Singapore, 73

Ferns, Notes on Malayan, with
descriptions of five new species,
119

Fimbristylis capillaris (pl.1l) 69

Flora of the Netherlands Indies,
Mr. H. N. RIDLEY and, 44

Flowering of TZembusu trees
(Fagrea fragrans ROXB.) in
Singapore, p. 173

Forestry in Malaya, Mr. RIDLEY
and, 42

Fumaroles in the highlands of
Java and Bali, 64

FURTADO, C. X.: Asterisks in
LINNAEUS’S Species Plantarum,
310

FuRTADO, C. X.: A Commentary
on the laws of botanical nomen-
clature, 223

FURTADO, C. X.: Chonemorpha
macrophylla and allied species,
113

FuRTADO, C. X.: Palmz Malesi-
ce :—VI Notes on some De-
monorops of the section Cym-
bospathe, 152; VII Two new
Calami from the Buitenzorg
Gardens, 182

FuRTADO, C. X.: The nomen-
clature of types, 285

Gymnogramme subtrifoliata Hk.,
142

Hancea, Index, 352

Haptotype, 288; 303

Heleocharis, Index, 309

Hemicardion crenata FEE, 122

Hemizvgia, Index, 309

HENDERSON, M. R.: The epi-
phytic flora of Dipterocarpus
oblongifolius BL. (Neram),

Vol, IX. (1938).

HENDERSON, M. R. & C. G. G. J.
VAN STEENIS: Bibliography of
the works of Mr. H. N. RIDLEY,
from 1872 to 1935, 2

Heterotypes, 309

Himantandra F. MUELL, 231

Holotype, 287; 290

HotttumM, R. E.: A _ Redefini-
tion of the Genus’ Terato-
phyllum, 353

Further notes’ on
Stenochlaena,
Lomariopsis and

Teratophy Hum,
1390
= Notes. on "been
Ferns with des-
criptions of five
new species, 119
sil The flowering’ of
Tembusu trees
(Fagraea fragrans
RoOxB) in Singa-
pore, 1928-1935,

x3

3
a The Genus’ Loma-
gramma, 190
Homalocenchrus MEIG., 226
Homonyms, 249
Hopea, Index, 352; plates 17,
18, 19; explanation of plates
350-351.
Hothouses in the tropics at 3,100
metres altitude, Open Air, 64
Hymenophyllum exiguum BEDD.,
138
Sat el, | exsertum
WALL., 130
Hypolepis bivalvis v.A.v.R., 130
Hyptis brevipes (pl. 1), 60

Icotvpe, 288; 209

Ideolectotype, 297

Ideotyvpe, 288; 301

Idiotype, 288

Illegitimate names, 224; 232

Invalid names, 224; 227

Isauxis, Index, 353

Isonym, definition O£;::223

Isonyms of unequal age based on
basinyms of equal age, 241

Isotvpe, 289; 302

Ixora. Index, 309

Labelling, type specimens and
type-plants, 306

Labels for Type-specimens, 306

LaM, H. JjJ.:. Phylogeny of
single features, 08

366

Lastrea calcarata var. sericea
RIDL., 129

Laws of botanical nomenclature,
A Commentary on the, 223

Lectoholotype, 288; 296

Lectoparatype, 288 3296

Lectotype, 288; 296

Leersia SWARTZ, 226

LEEUWEN vide DOCTERS
LEEUWEN

Legitimate names, 224; 234

Leptochilus cuneatws R. BONAP.,
217

Leptochilus decurrens BL., 131

Leptochilus lomarioides BL., 204

VAN

Leptochilus MOVOZUINEENSIS
BRAUSE., 209
Leucostegia immersa (WALL.}

PR., 131
Lindsaya fiabellata var. gigantea
HK., 131
Lindsaya integra HOLTTUM, 132
Lindsaya javanensis BL., 131
Lindsaya Lobbiana Hk., 131
Lindsaya lucida BL., 131
Lindsavya nitens BL., 131
Lindsaya nitida COPEL., 132
Lindsaya recurvata WALL., 131
Lindsaya tenera var. gigantea
HOLTTUM, 131
LINNAEUS’S Species Plantarum,
Asterisks in, 310; notations in,
311
Linum, Index, 284
Livistona, Index, 309
Logoapotype, 288; 299

Lomagramma abscondita v.A.v.
R., 205

Lomagramma articulata COPEL.,
356, 196 2

Lomagramma bipinnata COPEL.,
356, 196

Lomagramma borneensts, (plate
8)=L. Brooksn

Lomagramma_ Brooksii COPEL.,

199 (pl. 8)
Lomagramma Copsiandii HOLTTUM,
ep. nov. (plates 9 & 10) 201
Lowagramma cordipinna HOLTTUM
cp. MOV., 202
Lomagramma ouitrata
HOLTTUM comb. nov., 203
Lomagramma grosseserrata HOLT-
TUM Sp. nov., (plates 10 & 11),

(BAK.)

203

Lomagramma guianensis (Aubl.)
Ching, 196

Lomagramma lomarioides (BL.)
J. Sm. (pl. 12, 13), 204

Lomagramma Matthewii (CHING)

HOLTTUM Comb. nov., 206
Lomagramma melanolepis V.A.
v.R., 208

Lomnagramma Merrillii HOLTTUM
ep. nov., (plate 14), 208

lLomagramma novoguineensis
(BRAUSE) C. CHR. (pl. 15), 209

Lomagramme perakensis BEDD.,
(pl. 16) 210

Lomagramma polyphylla BRACK,
212

Lomagramma pteroides J. SM.,
(pl. 13), 213

Lomuiagramma pteroides var.
negrosensis COPEL., 201

Lomagramma pteroides var.
subcoriacea COPEL., 213

Lomagramma sinuata C. CHR.,
215

Lomagramma sorbifolia (WILLD.)
CHING, 220

Lomagramma subcoriacea
COPEL., 213
Lomagramma sumatrana V.A.V.

R. (pl. 16), 217

Lomagramma, The Genus, 190

Lomagramma Wilkesiana,
COPEL., 350, 196

Lomaria spectabilis, KZE., 141

Lomaria ? juglandifolia PRESL.,

139

Ceimniskens Balausae FOURN.,
362

Lomariopsis Brackenridgei,
CARR., 142

Lomarionsis leptocarpa FEE, 143

Lomartiopsis leptocarpa HOLTTUM,
14!

Lomariopsis
METTY., 141

Loranthus cordilimbus, 293

Lycopodium cernuum (pl. 1), 69

Lygodium borneense yv.A.V.R.,
132

Lysichitum americanum, 2096

spectabilis (KZE)

Madhuca Ridleyi H. J]. LAM NM. sp.,
98; 107, (pl. 7)

Malaysia and Malaya in plant
Geography, On the applica-
tion of the Terms, 187 ©

Merotype, Index, 309

MERRILL, E. D.: Some Malay-

sian phytogeographical _ pro-
blems, 49

Mesembryanthemum Sect.
Minima HAwW., 231

Metatype, 301

Microlepia pilosula (WALL.)
PER:j 132

Misapplication of names, 258;
261

Mixtum compositum, 244 adnot.
Mocanera, Index, 353

Gardens Bulletin, S.S.

367

stematic) : Alter-
Citation of mis-
illegitimate, 224;
232; invalid, 224; 227; legiti-
mate; 224; 234; orthography
of, 236: provisional, 230; rival
of equal age and equal rank,
240; valid, 224; varietal, 241
Neotype, 288
Nephrodium ciliatum CLARKE, 129
Nephrodium Parishii HOOK., 129
Nephrolepis barbata COPEL, 132
Neram, The Epiphytic flora of,

Names (sy
Mative, 230:
applied, 261:

93
Neurotecoma K. SCHUM...
New combination, 239
New name, 2
Nomen confusum, 244
Nomen dubium, 2
Nomenclatorial expressions, 262
Nomenclatural laws, commentary
on, 223
Nomenclature, A commentary on
the laws of botanical, 223
Nomenclature of types, 285
Nomina excludenda, 234
Nomina invalida, 234
Nomina specifica conservanda,
267
Notations in LINNAEUS’S Species
Plantarum, 311

231

Oberonia Latifii J]. J. SM. Mm. Sp.,
Sq

Oldenlandia herbacea (pl. 1), 69

Open-air hothouses in the
tropics, 64

Ophioglossum simplex RIDL., 133

Orchids from the Malayan Archi-
pelago, So

Orthography of names, 255

Orthosiphon Thorncroftii,

Osmunda javanica BL., 133

304

Palmae Malesicae vi: Notes on
Some Daemonoraps of the
Section Cynobospathae, 152;
vil Two new Calamz from the
Buitenzorg Gardens, 182

Parashorea, Index, 353 (pl. 21):
explanation of plates, 350

maiaanonan (BLANCO)
MERR. var. tomentelila Sym.
var. nov., 338 (pl.

Paratype, 287; 200

Pentaceras, 252

Pentaceros, 252

Pentagonia, 252

Pentagonium, 252

Phalaris orvzoides L, 226

Vol. IX. (1938).

Phegopieris oppostpinna V.A.V.
3 82

Phoenix amboinica, 297

Phylogeny of single features, 98

Phytogeographical problems,
some Malayan, 40

Piai lasa, 119

Piai raya, 119

Pinanga, Index, 309

ee adnata (BL.) BEDD.,

Plants ace by bats, 58
Poa, Index, 300;
Poikilospermum, 252
Polybotrya articulata J. SM.,

356

Polybotrya Wilkesiana BRACK,
359

Viskenddiaen albidosquamatum
BL., 133

Polypodium dilataium var.

grandideutatum CES., 134

Polypodium euryphyllum C.
CHR., 134
Polypodium grandidentatum

(CESATI) BAK., 134

Polypodium Hancockii BAK., 134

Polypodium insigne BL., 134

Polypodium malaicum yv.A.v.R.,
134

Polypodium normale DON, 134

Polvpodium sessilifolium WHK.,
134

Polypodium subnormale V.A.V.
R., 134

Polypodium subevenosum Bak.,
135

Polystichum prolificans v.A.V.
R., 135

Polystichum puncticulaium V-A4.
v.R., 137

Pothos pinnatus, 299
Primary types, 287; 289
Procedures affecting Art. 69, 237
Proteromerotypes, 288; 301
Proterotypes, 287, 2890
Provisional names, 230
Psilanthus, 252
Psilosanthus, 252

Pteris platvsora BaK., 135
Pteris scabripes WALL., 136
Pteris venulosa BL., 136
Ptychosperma, Index, 284;

(Hovtrt.)

309

zevlanica
136

Quvercifilix
COPEL.,

Radiola, Index, 284
kaphidophora, Index, 2
Representative botanical
tutions under Art. ms 2
Retinodendron, Index, 353

insti-
73

368

Rhizophora, Index, 284
Rhopalodia, Index, 284
Richetia, 330
Richetioides, 330

RIDLEY, Bibliography of the
works, 2

RIDLEY and Rubber in Malaya,
39

RIDLEY (Mr. H. N.) and the
flora of the Netherlands
Indies, 44

KIDLEY (Mr. H. N.): List of

plants named in honour of, 29
RIDLEY and forestry in Malaya,

HENRY NICHOLAS (pl.

42
RIDLEY,
dedication of the Bulle-

p. I):
tin, 1
RIDLEY’s work for tropical agri-
culture, 31
Rotan, Index, 181

Rubber in Malaya, Mr. RIDLEY
and, 39

Rumohra aristata (FORST.)
CHING, 137

Rumohra puncticulata (vy.A.v.R.)

HOLtTtTuM Comb. nov., 137

Scindapsus, Index, 309

Scirpus, Index, 309

Seasonal fruiting of Agarics in
Malaya, 79

Semeiocardium arriensii ZOLL.,
72 (pl. 2)
Semeiocardium ZOLL., a mis-
interpreted genus of Balsa-

minaceae, 70

Separates, effectiveness of ad-
vance, 271

Shorea, Index, 353, explanation
of plates. 350-351

*S. acuminatissima Sym. sp. nov.,

350 (pl. 23).
assamica DYER forma assa-

mica SyM. stat. nov., 331

S. assamica DYER forma globi-
fera (RIDL.) SYM. Stat. nov.,
331

S. assamica DYER forma Koor-
dersii (BRANDIS) SYM. Stat. nov.,

331

S. assamica DYER forma philip-
pinensis (BRANDIS) SYM. Stat.
nov., 331

S. sandakanensis SyM. sp. Nnov.,
343 (pl. 25)

S. Smithiana SYM. Sp. NOV., 345,
(pl. 26)

S. xanthophylla Sym. sp. nov.,
342 (pl. 24)

SLOOTEN, D. F. VAN: Mr. H.

N. Ridley and the flora of the
Netherlands Indies, 44

SMITH, J. J.: A few orchids
from the Malayan Archipe-
lago, 89

SouTH, F. W.: Mr. RIDLEY’s

work for tropical agriculture,

31
Specimens, Index, 309
Spermotype, 288; 304
Spirotecoma, Index, 284
STEENIS, C. G. G. J. VAN: On
the application of the terms
Malaysia and Malaya in plant

geography, 187

STEENIS, C. G. G: J. VAN:
Open-air hothouses in _ the
tropics at 3100 metres altitude,
64

STEENIS: vide also HENDERSON
and van STEENIS

Stenochlaena fraxinifolia
PRESL., 139

Stenochlaena juglandifolia
PRESL., 139 .

Stenochlaena laurifolia PRESL.,
141

Stenochlaena Milneit UND., 139

Stenochlaena palustris, 140

Stenosemia pinnata COPEL, 137

SYMINGTON, C. F.: Notes on

Malayan Dipterocarpaceae IV,
319

Synonyms, Index, 284

Syntype, 288, 293

Tabernaemontana elliptica
BLANCO, 116

Talinum, Index, 284

Tautonyms and homonyms, 249

Tectaria decurrens (PR.) COPEL.,

137

Tectaria Keckii (LUERSS.)
C. CER: a

Tectaria Maingavi (BaAK.)
G. “CHms, 448

Tectaria ternifolia (v.A.v.R..)
C. CuHre., 138

Tembusu trees in Singapore,
flowering of, 73

Teratophyllum aculeatum (BL.)
METT., 144

Teratophyllum articulatum

(J. SM.) METT., 356
Teratophylium Brightiae (F. Von
MUELL.) HOLTTUM comb. nov.,
358 pl. 28
Teratophyllum
HOLTTUM 142

Clemensiae

Teratophyllum: Discussion on
the Polvseriatae, 362
Teratophylium leptocarpum (FEF)

HOLTTUM comb. nov., 143

Gardens Pulletin, S.S.

309

Teratophyllum luzonicum
HOLTTUM, 142

Teratophyllum Wilkesianum
(BRACK.) HOLTT. comb. nov.,

359 (plates 209, 30)
Teratophyllum

HOLTT., “143
Topotvpe, 288; 2090
Trichomanes exiguum

BAKER, 138
Trichomanes radicans SW., 138
Type-plants, Index, 309
Type specimens, Index, 309
Types, The nomenclature
_ 285
[yponym, 233

Welliamsit

(BEDD.)

of,

Unona, Index, 284

Vol. IX. (1938).

Validating a name by referring
to a misapplication, 258
Valid names, 224
Valid publications, 229
VAN LEEUWEN vide
VAN LEEUWEN
VAN SLOOTEN :
VAN STEENIS: wide STEENIS
Varietal names, 242
Vatica diospyroides Sym. sp. nov.,
347 (pl. 27)

DOCTERS

vide SLOOTEN

Vatica, explanation of plates,
Sot. .

Vittaria’ malavensis HOLTTUM,
138

Vittaria stenophylla COPEL, 138

Wendia, 252
Wendtia, 252

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