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httos://archive.org/details/gardensbulletins425unse

THE

GARDENS’ BULLETIN

SINGAPORE

Volume XXV
1969-1971

A periodical reflecting the interests
and activities of the Botanic Gardens

Singapore

EDITOR:
Director, Botanic Gardens

Singapore

Published by Authority

PRINTED BY LIM BIAN HAN, GOVERNMENT PRINTER, SINGAPORE
1972

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CONTENTS
Vol. XXV

PART I — 4th October, 1969
CHEW, W.-L.: A Monograph of Dendrocnide (Urticaceae) -

Ho.trum, R. E.: A new genus and new species of Malesian
Orchids - - - - - - -

Lim, G.: Calostoma sarasini from Malaya - - -

CHEW, W.-L.: A Monograph of Laportea (Urticaceae) -

PART II — 21st December, 1970

PROwSE, G. A. and RATNASABAPATHY M.: A Species List of
Freshwater Algae from the Taiping Lakes, Malaysia -

STONE, B. C.: Materials for a Monograph of Freycinetia
Gaud. (Pandanaceae) V. Singapore, Malaya and Thailand

Materials for a Monograph of Freycinetia Gaud.
(Pandanaceae) VI. Species of Borneo - -

KENG, HsuaN: Further Observations on Ancistrocladus
tectorius (Ancistrocladaceae) - - = ¢

Ho.tTumM, R. E.: The Genus Orchidantha (Lowiaceae) -

RIDSDALE, C. E.: The Arborescent Naucleae of New Guinea
and Solomon Islands - = 2 Bs is

FuRTADO, C. X.: The Identity of Hyphaene natalensis Kuntze
Asian Species of Hyphaene - - - - -

A New Search for Hyphaene guineensis Thonn. - -

PART Ill — 15th October, 1971

TIxIER, P.: A Contribution to Bryological Knowledge =
Fraser’s Hill Station (Malaysia) - - -

Corner, E. J. H.: Merulioid Fungi in Malaysia - -
REVIEW: Grasses of Malaya (H. Keng) - - -

OsiTuaRY: Juraimi bin Samsuri - - - :

INDEX - - - - E " a r

Pages

1-104

105-108
109-110
111-178

179-187

189-207

209-233

235-237
239-246

247-281

283-297
299-309
311-334

aay 35
355-381
383-384
385

387-394

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THE
| 9
GARDENS’ BULLETIN
SINGAPORE
en VOL XXY, Part I 4th October, 1969
CONTENTS
| PAGE
Cuew, W.-L.: A Monograph of Dendrocnide (Urticaceae) - 1
Ho.trum, R. E.: A new genus and new species of Malesian
Orchids . - . - : - . 105
Lim, G.: Calostoma Sarasini from Malaya - - - 109
Cuew, W.-L.: A Monograph of Laportea (Urticaceae) - 111

To be purchased at the Botanic Gardens, Singapore
Price: $9

DEC 2 2 1969

LIRRARY

_ Published by Authority

Printed by Lim Bian Han, Government Printer, Singapore

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THE

GARDENS’ BULLETIN

SINGAPORE

Vol. XXV, Part I 4th October, 1969

A Monograph of Dendrocnide
(Urticaceae)

by
W.-L. CHEW (CHEW WEE-LEK)

Dendrocnide was established by Miquel in 1851 to accommodate,
as the epithet implies, the tree Nettles. It was, however, considered
congeneric with Laportea by Weddell who had it reduced
accordingly. This resulted in Laportea becoming heterogeneous
and consequently defineable only by elimination.

Having studied the whole group on a monographic basis,
I conclude that Dendrocnide is quite distinct from Laportea
and that its status should be re-established. In my preliminary
paper in 1965, when the transfer of the species was accomplished,
reasons for my decision were given. It suffices here to mention
briefly that Dendrocnide distinguishes itself on the following
score: (a) perennial ligneous trees with irritant hairs, (b) leaves
petiolate, simple, alternate or spiral, (c) stipules entirely connate,
intrapetiolar, (d) flowers 4- or 5-merous, free or in small
glomerules, perianth not fleshy at maturity, (e) achenes usually
reflexed, with ligulate stigmas. Thus it comes very close to
Urera from which it differs in the last two of the points just
mentioned. ,,

I could. thirty-six species for this genus which is widely
distributed in South-east Asia and the Pacific as essentially
lowland trees. It is particularly abundant, species-wise, in the
Philippines and New Guinea.

This monograph is based on the materials deposited in the
following herbaria supplemented by field observations made in
the Malay Peninsula, Borneo and Java:

Amsterdam, Holland (AMD). Berlin, Germany (B).
Arnold Arboretum, U.S.A. (A). Bogor, Indonesia (BO).

‘ Brisbane, Australia (BRI).
Bangkok, Thailand (BKF). British Museum, U.K. (BM).

Berkeley, U.S.A. (UC). Brussells, Belgium (BR).

Cambridge, U.K. (CGE).
(NBG)

Cape Town, South Africa

Chicago, U.S.A. (F).

Copenhagen, Denmark (C).

Edinburgh, U.K. (E).
Florence, Italy (FI).

Gardens’ Bulletin, Singapore — X XV (1969)

Melbourne, Australia (MEL).
Michigan, U.S.A. (MICH).
Missouri, U.S.A. (MO).
Munich, Germany (M).

New York, U.S.A. (NY).

Paris, France (P).
Pondicherry, India (PON).

Geneva, Switzerland (G).

Kepong, Malaysia (KEP).

Kew, U.K. (K).

Kuching, Sarawak (SAR).

Lae, New Guinea (LAE).

Leiden, Holland (L).

Lisboa, Portugal (LISC,
LISJC and LISU).

Lund, Sweden (LD).

Madrid, Spain (MA).

Manila, Philippines (PNH).

Praha, Czechoslovakia (PRC).
Sandakan, Sabah (SAN).
Singapore (SING).

Stockholm, Sweden (S).
Sydney, Australia (NSW).
Uppsala, Sweden (UPS).
Utrecht, Holland (U).
Washington D.C. U.S.A. (US).
Wrocklaw, Poland (WRSL).
Zurich, Switzerland (Z).

1 would like to thank the directors of these herbaria for their
generosity and co-operation in making available for my study
their rich collections. My gratitude is also extended to the
following botanists for their advice and general assistance:
Dr. J. A. R. Anderson (Sarawak), Mr. H. M. Burkill (U-.K.),
Prof. E. J. H. Corner (Cambridge), Dr. C. X. Furtado (Singapore),
Prof. H. Godwin (Cambridge), Dr. Ding Hou (Leiden), Prof. J.
Leandri (Paris) and Prof. C. G. G. J. van Steenis (Leiden).

Part 1 — General

Habit. They are perennial, evergreen trees commonly not more
than 15 metres high with soft wooded stems. Branching is
sympodial, much like that of Terminalia, with the leaves often
forming rosettes at the end of the twigs. There are two exceptions
to this. In D. mirabilis, the base of the stem is succulent and
in D. excelsa, the trees reach a towering height of 35 metres
with buttresses at the base to 2 metres high.

Leaves. These are simple, petiolate, dorsi-ventral, penninerved.
generally coriaceous, spirally set and usually heavily covered
with cystoliths. On overall similarity, they resemble those of
some species of Ficus and are quite unlike those of Laportea
which have urtica-like leaves.

In some cases, the lamina type is very specific as in D. mirabilis
where it is plicate with half the lateral veins palmately branched,
and in D. kajewskii where it reaches its greatest dimension of
1.5 metres in length. In other cases, certain characteristics of the
leaves distinguish groups of species as in D. elliptica, D. kjellbergii,
D. moroides and D. peltata where they are peltate. Then there
are the species whose leaves are devoid of irritant hairs
(in D. luzonensis, etc.) and one whose lamina is decurrent on
the petiole (D. oblanceolata). As shown in the keys, combinations
of the various characteristics of the leaves afford a very good
way of distinguishing the species.

W .—L. Chew — Monograph of Dendrocnide 3

Stipules. One of the important characteristics that distinguish
this genus from Laportea is in the stipules. Here they are large,
entirely connate, bicarinate at the back, intrapetiolar and not
bifid at the apex. In size, they range from 1 to 10 cm. They are
thus very similar to those of Poikilospermum.

Inflorescences. Without exception these are racemose or pani-
culate, pedunculate, axillary, bracteate and usually with irritant
hairs. Unlike those of Laportea, they are quite uniform throughout
the genus. The two sections, however, differ markedly in the
disposition of the female flowers. In section Dendrocnide, the
sessile female flowers are set usually in a row on flabellately
expanded extremities of the peduncles. Such flabellate receptacles
usually curve over at maturity. In D. subclausa they curve com-
‘pletely over the achenes and may reach a size to | cm. in dia-
meter. At the other extreme, there is the species D. urentissima
where the receptacles are very small and often hardly developed.

In sect. Sarcopus, the female flowers are either borne free as
in D. sinuata or in small fascicles as in D. ternatensis or arranged
distichously at the extremities of peduncular branches as in the
amplissima group of species. Three subsections or series could
have been established to accommodate these groups if there had
been less intergradations between them.

Flowers. They are unisexual, 4- or S-merous and _ generally
small. The males are sessile or pedicellate, 4- rarely 5-tepalled,
with an equal number of stamens opposite the tepals and with
a rudimentary ovary. Being the least variable organ, the male
flower does not figure prominently in the delimitation of the
‘species except in D. longifolia where it has the largest and most
fleshy pistillode in the genus.

The female flowers are more useful as a criterion for species
delimitation. Also small, they range in length from 1 to 1.5 mm.
and are usually half as broad. Basically, they have a single
whorled 4-toothed or 4-lobed perianth, a unilocular ovary ovoid
in shape and topped by a simple, ligulate stigma. They are
also either pedicellate or sessile and are devoid of staminodes.
Variation of this basic structure is fairly great. In sect. Dendrocnide
and the amplissima group of species, they are sessile very rarely
subsessile. In the other section, many species have pedicellate
flowers as in D. sinuata where the pedicel is simple and greatly
elongated. In some species, as in D. photinophylla and D. vitiensis
the pedicels elongate greatly at maturity and are usually fleshy
and reflexed.

The 4-lobed perianth is by far the commonest. The lateral
tepals are usually longer and broader than the medial ones.
This difference is greatest in D. rechingeri where the lateral
tepals are at least thrice the length of the medial ones. In most
species however, they are usually not more than _ twice.
D. torricellensis and its relatives are noted by their perianth
_ being lightly 4-toothed and usually rather small. The extreme

condition is found in D. harveyi where it is either very minute or
entirely absent.

4 Gardens’ Bulletin, Singapore — X XV (1969)

The ovary is rather uniform throughout the genus and is
consequently little used in the discrimination of the species.
The stigma varies only in length. In D. amplissima and
especially D. microstigma, it hardly exceeds 0.2 mm. whereas
in D. oblanceolata it often reaches 8 mm. The average length is.
between 1 and 3 mm.

Achenes. These are sessile, asymmetrically ellipsoid to ovoid.,.
very rarely subrotund.

In some, they are very flat with a smooth pericarp as in D.
peltata. In others they are thick and bloated, with a very warted
pericarp as in D. moroides. In length, they vary from 2 to 5
mm. Their size in relation to the pedicels, perianth, stigmas and
other characteristics provide very useful characteristics in sepa-
rating the species.

Habitat. They are essentially lowland primary forest species
preferring slightly damper and somewhat shady habitats. Some
like D. elliptica prefer limestones and their derived soils. Many
of the New Guinea species seem tolerant of fairly exposed habitats
of secondary forests or abandoned vegetable gardens.

Phytogeography. A genus of 36 species, it is spread very widely
from India, Ceylon, China, through South-East Asia, Malesia,
North-East Australia to the Pacific Islands (fig. 1). If Continental
Asia is taken as a unit, 17 species are endemics; otherwise,
15 species are. Thus endemism is very high for this genus. Of
the two sections, sect. Dendrocnide is the more restricted: being
found from China, parts of South-East Asia, through western
Malesia to as far as the Moluccas. It is particularly well repre-
sented in the Philippines where 8 species are recorded. The other
section, sect. Sarcopus, covers the entire range. Its greatest repre-
sentation is in New Guinea where twelve species are recorded.
Thus, sect. Dendrocnide is centred in the Philippines while the
other in New Guinea. The species in each geographic area are
discussed separately.

Continental Asia. There are four species in this region:
D. basirotunda, D. sinuata, D. stimulans and D. urentissima.
Of these, D. basirotunda and D. urentissima can be considered
as endemics for Continental Asia as a whole. The other two
are very widely spread species.

Malay Peninsula and Sumatra. These two regions are identical
in their Dendrocnide flora: there are two species in each viz.
D. stimulans and D. sinuata, both wide ranging ones.

Java and Lesser Sunda Islands. These two are almost identical.
Both have D. stimulans, D. sinuata, D. peltata and D. microstigma;
but the latter has one more viz. D. carriana.

Formosa. Only one species is found in this island, viz-
D. meyeniana which occurs also in the Philippines.

W—L. Chew — Monograph of Dendrocnide

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Philippines. Ten species are recorded from these islands:
8 belong to sect. Dendrocnide and 2 to sect. Sarcopus. Six species
are endemic here. Only two species are wide ranging: D. stimulans
and D. carriana, Of the remaining two, one, D. elliptica, is also
found in Borneo and the other D. meyeniana in Formosa as
mentioned above. Thus the Philippines Dendrocnide flora is
distinctive.

Borneo. This island has three species: D. stimulans, common
in Western Malesia, D. elliptica which is also recorded from the
Philippines and D. oblanceolata, the only endemic. As this last
species is a very close relative of D. stimulans, one can generally
say that the Bornean Dendrocnide is closely allied to that of
western Malesia.

Celebes & Moluccas. This region, often referred to as
‘“‘Wallacea” is interesting. It is the meeting point of the western
and eastern Malesian elements and at the same time, it contains
species peculiar to itself. It is also the eastern most point of
distribution for sect. Dendrocnide. Of the ten species recorded,
D. stimulans and D. microstigma are the western elements while
D. longifolia and D. ternatensis are the eastern ones. The Tanimbar
Archipelago to the far south has two species in common with
north-eastern Australia, viz. D. moroides and D. cordata. The
species D. amplissima, D. kjellbergii and D. celebica are the ones
confined to “Wallacea”. Only one species D. carriana manages to
“cross” this region from West to East Malesia.

Australia. Four species are found here: D. moroides, D.
cordata, D. photinophylla and D. excelsa. Only the last two can
claim to be endemics; the others have now been recorded from
southern Moluccas.

New Guinea. This region has 12 species, 3 of which are ende-
mics, viz. D. torricellensis, D. corallodesme and D. morobensis.

Bismarck Archipelago & Solomons. The Dendrocnide flora of
this region is overwhelmingly New Guinean. Of the 8 species
recorded, 6 have representatives in New Guinea, one i.e. D. latifolia
is a Pacific element and another viz. D. kajewskii is endemic.

Pacific Islands. These islands have 3 species viz. D. vitiensis,
D. harveyi and D. latifolia. The first two have not been recorded
elsewhere while the last has been found in the Solomon Islands.

Affinities. That Dendrocnide belongs to the tribe Urereae,
there is no doubt. Within this tribe, I consider the genus to
be very closely allied to Urera with which it has numerous
characteristics in common such as habit of growth, the form of
the female perianth and achene etc. From Laportea it is now
separated by Discocnide. A few species, such as D. schlechteri
and D. luzonensis come very close to Poikilospermum in vegetative
as well as some floral characters, but these can at once be
distinguished by the presence of irritant hairs on the flowers in
D. luzonensis and by the different female flowers in D. schlechteri.

W —L. Chew — Monograph of Dendrocnide 4

Part 2. — Systematic

DENDROCNIDE

Mig. Plantae Junghuhnianae, 29 (1851); Chew in Gard. Bull.
Sing. 21: 201 (1965). — Urticastrum Heist. ex Fabric. sensu
O.K. Rev. Gen. Pl. 634 (1891), pr. p., exclu. typ. sp.

Dioecious perennial shrubs or trees with irritant hairs. Leaves
petiolate, simple, coriaceous, crenulate, undulate to smooth, spirally
arranged. Stipules entirely connate, intrapetiolar, coriaceous.
Inflorescences pedunculate, racemose, bracteate, axillary, with
flowers freely borne or in small fascicles or in flabellate receptacles.
Male flowers mainly 4-merous, rarely 5-merous, with pistillode
and stamens opposite tepals. Female flowers 4-merous, sessile to
pedicellate, flabellately or distichously arranged or in_ loose
fascicles; ovary ovoid, unilocular, usually with ligulate stigmas;
ovule erect, orthotropous; staminodes absent, pedicels simple or
swollen. Achenes compressed or bloated, ovoid, not chartaceous,
usually strongly warted.

Distribution: China, India, Ceylon, through South-east Asia,
Malesia, Australia and the Pacific islands.
Lectotype species: Dendrocnide costata Mig. [| = D. stimulans
(L.f.) Chew].
sect. DENDROCNIDE

Laportea sect. Dendrocnide (Miq.) Wedd. in Arch. Mus. Hist.
Nat. Paris 9: 133 (1856) & DC. Prodr. 16 (1): 85 (1869).

Characteristics: Female flowers sessile very rarely subsessile,
borne usually in one row (rarely two) on often fleshy flabellate
receptacles.

Lectotype species: Dendrocnide costata Mig. [= D. stimulans
(L. f.) Chew]. Species are: nos. 1 to 10.

sect. SARCOPUS

(Wedd.) Chew, comb. novy.—Laporiea sect. Sarcopus Wedd. in
Arch. Mus. Hist. Nat, Paris 9: 129 (1856) & DC. Prodr. 16 (1): 82
(1869), basionym.

Characteristics: Female flowers sessile or pedicellate, borne
freely, in loose fascicles or distichously at the extremities of pedun-
cles; pedicels often fleshy.

Type species: Laportea gigas Wedd. [= D. excelsa (Wedd.)
Chew]. Species are: nos. 11 to 36.

l1—a
b
2—a
b
3—a
b
4-4
b
5S—a
b
6—a
b
J—a
b
8—a
b
9—a
b
l1—a
b
2—a
b
3—a
b

. Lamina profoundly cordate, broadly ovate, both sides dense with

Gardens’ Bulletin, Singapore — X XV (1969)

Key to Species of Sect. Dendrocnide

. Lamina peltate, obovate, both surfaces dense with irritant hairs

(Borneo & Philippinesietnicycereetice..-i.....e oe .-.: elliptica
. Lamina .non-peltate- variously shaped vcisucd.2-. Sea 2

atvitant hairs :si0t4...oncutycoabttel . senus.<ch ees. tes 0 eee

. Lamina at most lightly cordate variously shaped, the upperside
4

usually’ subslabrous’ to Piatra... -«--.2-. <2: reset oeeegs cee

. Lamina at least twice as long as broad, margin dentate; male

flowers,, 4-tepalled (Philippines). ..: --cn.crnsrren-- ecbeceeeps crassifolia

. Lamina as long as broad, margin smooth; male flowers 5-

tepalled (Chima” & Vietingaigr’.”. ic. rece ace urentissima

. Lamina broadly ovate to oblong-ovate base broadly cordate ...

ee ee ee

. Lamina elliptic, obovate to narrowly ovate, base round to cuneate,

Hot COPGSIE 6. oeckociecDe - clea Sh: ade ee

. Lowerside of lamina glabrous the lowerside of petiole dense with

long irritant, hairs CPWwuIpeiites) ... are. 0 ea eene soya - pos snet densiflora

. Lowerside of lamina pubescent to wooly, petiole evenly covered

with short irritant hairs (Formosa & Philippines) ...... meyeniana

. Lamina obovate, decurrent on the short petiole; margin often

dentate: (Borne@) acu)... hee... 2a ee oblanceolata

. Lamina variously shaped but not decurrent on petiole; margin

often! ‘smeothisyii....w.i8.)... Si... OU... eee

. Lamina very rigid coriaceous, elliptic to ovate, usually 25-40 cm.

long male flowers surrounded by bracts 2-3 cm. long, female
receptacles dense with Tong irritant Hairs .CAAW. Sea.

. Lamina normally coriaceous, elliptic to obovate, usually less than

25 cm. long male flowers with very small interfloral bracts, female
receptacles with .very/‘Shontiairritast (haigiig.. 2... oo. nen nats adnan 9

. Flabellate receptacle greatly enlarged and curved over at fruit.

almost enclosing achenes; lowerside of lamina light pubescent
(Philippings):4 beeiorbatt! Dyce Ry ee ews subclausa

. Flabellate receptacle not entirely curved over; lamina entirely

glabrous! (Phidippines)22.97,. 229.40. OL6MAA.. 2a rigidifolia

. Female inflorescences shorter than leaves, female flowers very

shortly pedicellate; receptacle greatly fleshy at fruit (Philippines)
venosa

ee ee aa’

. Female inflorescences longer than leaves, female flowers entirely

sessile; receptacle not very fleshy at fruit (S.E. Asia, Malesia) ...
stimulans

Ce ee ee i ee ee ees

Key to Species of Sect. Sarcopus

. Shrubby tree with succulent base; lamina ovate; plicate. the

lower lateral veins palmate, the upper penninerved; the upper-
side of lamina often with 2 fleshy appendages (New Guinea &

sa cane geal» ill eis NR en TD i a mirabilis
| INOL UNIS COMMINANION. Of CHATACIEES, ......°.<..0..-..°:..+acce matte as 2
.. Amina, 'pefiate'™).. Yc hts.. Pe ah, SEF OR 3
Lamina ' gon-peltatel 1k. Y2UORIUALD.I0. BALRLORS. 2a OL. vis 5

. Lamina oblong, just peltate; female flowers entirely sessible. with

@-deniate perianth (Celebes) «vicars: dopteasser- ae kjellbergii

. Lamina ovate to almost round usually fairly deeply peltate:
4

female flowers pedicellate, with 4-tepalled perianth ...............

W —L. Chew — Monograph of Dendrocnide 9

4—a.

Stas

10 — a.

11 —a.

12 — a.

Lamina generally ovate, strongly dentate both surfaces dense
with irritant hairs; achenes ovoid, bloated warty, almost covered
by the lateral tepals, borne on greatly swollen pedicels (Southern
IMoimecas Ge JAMIA) | 62...) Seb rs taiet- dag rer - reer moroides

. Lamina almost round to ovate minutely dentate, upperside gla-

brous; achenes flat, round, smooth, tepals at the very base, borne
on non-fleshy pedicels (Malesia) ..........................00cee peltata

Lamina ovate, subglabrous to glabrous, with sinuate to dentate
margin; base cordate or rounded; male flowers on long pedicels:
female flowers very shortly pedicellate, perianth very minute or

Pe a) RE Re cr et Ee harveyi
Poe tie Gomibneron Of CATACUEIE <.....2..........-..--4-00ce serene 6
. Lamina 100 to 150 cm. long, ca. 50 cm. broad, the upperside

glabrous, the lowerside pubescent (Solomons) ............ kajewskii
. Lamina less than 80 cm. long, glabrous to pubescent ............ 7
. Lamina narrowly obovate, to 25 cm. long, with rather long

acuminate apex; male flowers pedicellate, each in the axil of a

eects (feo eet MOT P(RLEIOOCS) rece eee see see ee eee celebica
a, Bis COMMIMMAMON: OL CHAFACTETS 25.04 + seasoqucsasl sp - humo «cd ane ns 8
. Female flowers sessile, at most subsessile, sitting in somewhat

opposing rows on peduncular extremities .......................468. 9
. Female flowers pedicellate, usually in fascicles; pedicels often

RENE Cn SPIN CT PTT mn On. rece ne scene eet int etal tarts ate 13

. Lateral veins at least 20 pairs, very closely spaced; male flowers

pedicellate, rather separate from each other (Celebes & Moluccas)
cd baaembnne: 42 iss TA pdaaeass és: Sabsers< PP ek... Re amplissima

. Lateral veins less than 20 pairs, seldom very close; male flowers

Genstie, Vim creme > iaseicles 2 A. OSD) PAE AONB. 10

Pistillode very large, cubical in shape, fleshy, occupying nearly
the entire floral space; stamens very small, lodged at the sides
of ovary; infrutescence often fleshy and convolute (Celebes,
Moluccas through New Guinea to Solomons) ............ longifolia

. Pistillode small, not cubical nor fleshy; stamens larger than

pistillode; infrutescence fleshy but not convolute ............... 11

Lamina elliptic to oblong, tomentulose on the lowerside, base
round to cordate; lateral tepals of female flowers often greatly
developed (New Guinea to Solomons) ..................... rechingeri

. Lamina narrowly elliptic to obovate, lowerside not tomentulose,

base cuneate; lateral tepals of female flowers not much longer
aD ie. SPEER OER c cee. eur) isog,.. GE. sist). QO. ek 12

Lamina entirely glabrous, obovate; petioles also glabrous; lateral
tepals of female flowers slightly longer than the medial ones:
stigma 1-2 mm. long (New Guinea and Bismarck Archipelago)
a he ll schlechteri

. Sides of mid-rib on the lowerside of lamina with long soft hairs,

otherwise quite glabrous; petioles sericeous; lateral tepals of
female flowers not longer than the medial ones; stigma less than
0.5 mm. long (Java, Lesser Sunda Islands, Celebes & Moluccas)
ST SEPMREER AREY oricdgekas evi cg tovyaind (ne asvnnnde vas hMIARTOITe). 2D. A microstigma

Female flowers in loose fascicles; female pedicels not altogether
RMON CME NIUE Flee fre, fren eed. Sere cnn ppecavcccases 14

. Female flowers fasciculate to glomerulate; female pedicel usually

fleshy throughout ,greatly reflexed ................0..cccccc ccc cee eee 15

Lamina very smooth, entirely glabrous; irritant hairs only on
Dewees TUPI re ee luzonensis

. Lamina not smooth, well armed with irritant hairs not confined

tah Somemn Aiido-Baahema): | . pecasols, . Paiile. 0d. i ek sinuata

10
15 —a.
b
16 —a.
b
17 — a.
b
18 — a.
b
19—a.
b
‘20 — a.
b
21 — a.
b
“22 —a.
b
23 —a.
b
“24 —a.
b
25 —a.

Gardens’ Bulletin, Singapore — X XV (1969)

Female perianth very shallowly 4-toothed, becoming a _ loose

envelope at the base of achene at maturity ........................... 16
. Female perianth with well developed tepals enclosing the base of
achene firmly at maturity +. ..6..-... AERA. a. AR 18

Lamina narrowly elliptic, often toothed, very shortly petiolate,
lateral veins short, numerous; male flowers pedicellate; female
inflorescences commonly borne on leafless twigs (New Guinea)
OL AEE. AL ROSE... OE SOIR aCe. . ee. 6 corallodesme

. Lamina ovate, often very broadly so, often tooth, fairly long

petiolate, lateral veins long, at a low angle to the midrib; male
flowers sessile; female inflorescences borne on leafy twigs ...... 7

Lamina light villous on the lowerside; male flowers dense villous
to ‘wooly (New Guinea? . A021... NEE OLN el torricellensis

. Lamina glabrous; male flowers only with irritant hairs, not villous

nor woolly (Moluccas & New Guinea) ..................... ternatensis

Adult lamina entirely glabrous and devoid of irritant hairs ...... 19

. Adult lamina variously hairy or glabrous but always with irritant

iT} ns Lak ee iain es» 2s tia ign Aa Ow. See MR. FI oe kg. 22

Lamina obovate, the basal lateral veins very short; female pedicels
winged, with irritant hairs in two rows (New Guinea & Solomons)
» ob qes on nasnlile tga isetany Mees tegen . A Guia Otis « Ae; Mires De ae Oe nervosa

. Not this combination of . Characters si cccelondd: devcnicees denne es 20

Stipules 1.5-2 cm. long; lamina usually elliptic, more than 20
cm. long; female pedicels not very long, nor greatly fleshy
(Vacromesia: @ DRSIAMGRIB) oa s.uiici ans tcesanecsenesceehaketeee Gee latifolia

. Stipules less than 1.5 cm. long; lamina ovate-pentagonal or

elliptic, usually less than 20 cm. long; female pedicels elongating
greatly at fruit, and greatly .ewollett.is<..2.:4...sstacaeoee: 21

Lamina ovate-pentagonal, base usually rounded, lateral veins
more than 8 pairs, the basal pair not much longer than the
rest (Samoa, Fiji & New Caledonia) ........................ vitiensis

. Lamina base cuneate to rounded; lateral veins 6-8 pairs. the

basal pair reaching to more than half the length of the lamina
(Asisizaliah: (ar je 7 a ee photinophylla

Male flowers 5-merous, sessile; lamina exact ovate, base rounded
glabrous except for some irritant hairs on veins (China, Burma
RTC he ee es cer Tee ee ee a BP basirotunda

. Male flowers 4-merous, pedicellate; lamina of various shapes and

VCRINEUITS .ssiipninnicsaaiteiednorimocennanpeinerdsiavinaiun eR RUG. An URC) Same 23

Lamina broadly ovate, the underside dense woolly, both sides well
covered with irritant hairs; base profoundly cordate, with the two
lobes oftenvoverlapping (Australia) 200!...009..42/..2mer excelsa

. Lamina of various shapes but the underside not woolly: base

rounded to cordate, the two lobes not over-lapping ............ 24

Lamina cordate shaped, margin broadly sinuate; petioles usually
more than 18 cm. long; irritant hairs not on papillate bases (East
Malesia & “Austialia) *.c...5.. chased oe deel ee cordata

. Lamina ovate, elliptic to pentagonal, margin smooth or dentate,

not sinuate; petioles less than 18 cm. long; irritant hairs on
papillaté bases... 20 i. saccesbeeeahl ae ee Ole 2 25

Lamina ovate, with 6 or 7 pairs of lateral veins; the base with
one pair very prominent veins (New Guinea) ............ morobensis

. Lamina elliptic, oblong to ovate-pentagonal, with 8-11 pairs of

lateral veins; the base with a few pairs of very short veins
(Hast Malayera) (0), i, DS Mee eevee Geer carriana

W -—L. Chew — Monograph of Dendrocnide II

(1) Dendrocnide stimulans (L. f.) Chew in Gard. Bull. Sing. 21: 206
(1965).—Urtica stimulans L. f. Suppl. Sp. Pl. 418 (1781); BI.
Bijdr. 505 (1825), includ. var obtusa—U. fruticosa L. f. op. cit.
417 (1781).—D. costata Mig. Pl. Jungh. 1: 30 (1851).—
Laportea stimulans (L. f.) Mig. in Zoll. Syst. Verz. Ind. Arch.
103 (1854).—L. costata (Miq.) Mig. Fl. Ind. Bat. 1(2): 231
1859), exclud. var. contracta.—Urticastrum stimulans (L. f.) O.K.
& U. costatum (Mig) O.K. Rev. Gen. Pl. 2: 635 (1891).—L.
gracilipes Elm. Leafl. Philip. Bot. 3: 876 (1910).—L.
lanaensis C. B. Rob. in Philip. J. Sc. Bot. 5: 483 (1911).—
L. brunnea Merr. in op. cit. Bot. 10: 298 (1915).—L. annamica
Gagn. & L. thorelii Gagn. in Bull. Soc. Bot. Fr. 75: 2 (1928).—
L. stenophylla Quisumb. in Philip. J. Sc. 41: 317, tab. 1 (1930).
—L. hainanensis Merr. & Metc. in Lingn. Sc. J. 16: 189. fig. 4
(1937).—L. sumatrana Merr. in Pap. Mich. Ac. Sc. 24 (1): 65
(1939).—D. stenophylla (Quisumb.) Chew in op. cit. 206 (1965).

Characteristics: (a) lamina obovate, often narrowly and strongly
cuneate at the base, subglabrous to pubescent, (b) inflorescences
long pendulent especially the females, (c) flowers sessile, (d) female
flowers flabellately arranged in one row, (e) flabellate receptacle
curved but not fleshy nor covering the matured achenes, (f)
peduncles, petioles, twigs, fairly densely covered with irritant hairs.

Distribution: Hainan, Laos, Vietnam, Thailand, Malaya,
Sumatra, Java, Lesser Sunda Islands, Formosa, Philippines,
Anambas Islands, Borneo, Celebes, Moluccas.

Dioecious trees ca. 7 metres, branches rather widespread, bark
light grey, lenticellate, otherwise quite smooth. Twigs glabrous,
rarely light pubescent, without irritant hairs; petiolar scars quite
prominent. Lamina (12—) 15-30 (—40) cm. long, (4-) 6-10 (—14) cm.
broad, elliptic to obovate, rarely oblong or ovate, coriaceous, very
light pubescent only on veins of lower surface; irritant hairs on
veins, intercostals and veinlets of abaxial surface, sparse on
adaxial side; cystoliths punctiform, more on abaxial side; base
generally cuneate, sometimes round or emarginate; apex acuminate,
acumen often sharp; margin entire, very rarely lightly dentate;
lateral veins(8—) 10-13 (—15) pairs, basal pair short, quite pro-
minent, intercostals fairly straight and prominent. Petioles (2—) 5—10
(-15) cm. long, dense with punctiform cystoliths, generally some-
what glabrous, sometimes very light pubescent, with irritant hairs.
Stipules 0.75—2 cm. long, ovate, coriaceous in the centre, chartace-
ous towards the margins, pubescent when young, light pubescent on
the outside when matured./nflorescences branched racemes, floral
fascicles 2-10—flowered, 10-55 cm. long, females usually much
longer than males, axillary, usually solitary, rarely paired,
pendulent peduncular barcts minute to as much as 2 cm. long
triangular, pubescent; peduncles light to dense pubescent, usually
armed with irritant hairs. Male flowers sessile, 1-2 mm. long,
6.75—1.5 mm. broad, light pubescent, irritant hairs small and sparse,
4-tepalled; stamens 4, with reflexed filaments; pistillode ca. 1 mm.
long, obpyramidal; interfloral bracts minute to as large as tepals.

12 Gardens’ Bulletin, Singapore — X XV (1969)

variable, triangular, often light pubescent. Female flowers sessile,
slightly sunken in a row on fleshy flabellate peduncular recepta-
cles, ca. 1 mm. long, ca. 0.75 mm. board, pubescent. basal half
usually armed with irritant hairs, 4-tepalled; tepals slightly
decussate-imbricate, light pubescent; ovary at maturity slightly
longer than tepals; stigma ligulate, to 5 mm. long; interfloral bracts
usually less than 1 mm. long, ovate, pubescent, often bordering
the row of sunken female flowers. Achenes 2-4 mm. long & broad,
aimost flat except for the centre, asymmetrical, otherwise round,
rarely asymmetrically elliptic; pericarp smooth to warty; persistent
perianth a minute cupule at the base; persistent stigma curled
over the apex of the achene.

A highly variable species, particularly in the size of the lamina,
this occurs in primary forests from sea-level to about the 1200-
metre contour. In Mt. Kinabalu in Sabah, it is_ particularly
common at about the 1,000 metre line. In Sarawak, it is common
on limestones.

I have seen the types of all the synonyms cited above including
Thunberg’s collections from Java, on which Linnaeus’s U.
stimulans and U. fruticosa are based; and I am convinced that
they belong here.

CHINA:—Hainan: Pak Shik-ling: Ku-Tung village: Lei 584, April

1933 (type of L. hainanensia: A, BO & K). Pak Shek-shaan: Tsang 17487,
June 1928 (UC).

LAOS:—Luang Prabang: Thorel 3487 (syntype of L. thorelii: A & P).
—Muang Huang: Wieng Chan: Kerr 20799, March 1932 (BM & K).

VIETNAM:—Annam: Dyjiring: Chevalier 30985 (type of L. annamica:
P).—Tonkin: Mt. Basei: Balansa 2528 & 3289 (syntypes of L. thorelii:
K &_P),

THAILAND:—Kao Talu: Kerr 11807, Feb. 1927 (BM).—Nakorn
‘Sritamarat Pen.: Kiriwong: Smitinand 5257, July 1951 (A).

MALAY PENINSULA:—Penang: Waterfall Gardens: Curtis 1100,
July 1892 (K & SING)—Selangor: Batu Caves: Ridley 8527, July 1897
(K. & SING). Bukit Lagong: Sinclair 10727, Nov. 1962 (FI & SING).

SUMATRA :—Asshan: Rahmat-si-Boeea 6925 (type of L. sumatrana:
US); 7851 (A, S & US).—Enggano: Lutjeharms 3603, May 1936 (A, K,
L, NY & SING).

JAVA:—sine loc.: Junghuhn s.n. (type of D. costata: L. herb. 908-188-
3206). Thunberg s.n., 1777 (types of U. stimulans & fruticosa in herb.
Montin:. S3 Zollinger, 510, (A, Fi, G, Kye 13):

LESSER SUNDA ISLANDS:—Bali: Kostermans KKSS. 125 & 127,
June 1958 (BO, K & L)—Lombok: Bloembergen 3027, Feb. 1939 (L).

FORMOSA :—Koshum: Wilson 11015 (K).

PHILIPPINES :—Mindanao: Apo Mt.: Elmer 10499, May 1909 (type
of L. gracilipes: A, BO, E, F, G, K, L, MO, WRSL & Z). Lanao Lake:
Clemens 462, April 1906 (type of L. lanaensis: F. & M). Mayo Mt.:
Ramos & Edano 49413, April 1927 (type of L. stenophylla: B, NY & UC).
—Camar: Canayan Valley: Ramos 17559 (type of L. brunnea: K. & US).

BORNEO:—Sabah: Kinabalu: Chew & Corner RSNB. 4986, April
1964 (K & SING). Tawao: Elmer 21115, (A, BR, C, F, K, L, S & Z).—
Sarawak: Bau: Chew CWL. 537, July 1963 (SING). Api: Anderson 4286,
July 1961 (K & SAR)—Kalimantan: Lampoeng: Winkler 3352, Aug. 1908
(BO & L).

ANAMBAS ISLANDS:—Siantan: Henderson SFN. 20281, April 1928
(BO, K, SING & UC).

CELEBES:—Minahassa: Koorders 19393, March 1895 (L); Warburg
15648 (E).

MOLUCCAS:—Buru: Musoena Lawan: Toxopeus 550, Nov. 1921 (BO).

W.-L. Chew — Monograph of Dendrocnide 13

Fig. 2: D. stimulans: A = female twig; B & C = female flowers; D. E &
F = achenes; G = male inflorescence; H = male flowers.

14 Gardens’ Bulletin, Singapore — X XV (1969)

(2) Dendrocnide oblanceolata (Merr.) Chew in op. cit. 204 (1965)—

Laportea oblanceolata Merr. in J. Str. Br. R. Asiat. Soc. 85: 167

(1922),

Characteristics: Close to D. stimulans from which it differs in
(a) lamina decurrent on the short petiole, (b) undersides of lamina
pubescent to light canescent, (c) flabellate receptacles of female
inflorescences fleshly at fruit, (d) stigma up to 8 mm. long.

Distribution: Borneo (endemic! ).

Dioecious trees to 10 metres high. Twigs dense pubescent, soon
subglabrous, irritant hairs sparse; petiolar scars prominent. Lamina
(20-) 25-35 (—50) cm. long, (6—) 10-14 (—20) cm. broad, obovate,
coriaceous, uppersides glabrous, without irritant hairs, lowerside
glabrous to ligh pubescent, with hairs and irritant hairs mainly on
mid-rib and veins, both surfaces very dense with punctiform
cystoliths; base cuneate, decurrent on the short petiole; apex long
acuminate, acumen 1-3 cm. long; margin lightly toothed to sinuate;
lateral veins ca. 12-15 pairs, the basal ones very short and incon-
spicuous; intercostals straight, numerous. Petioles 2-3 cm. long,
winged, fairly thick, dense pubescent on the inside, soon glabrous,
dense with punctiform cystoliths; irritant hairs on the inside.
Stipules 2-3 cm. long, dense pubescent, soon glabrous, coriaceous.
Inflorescences unisexual, branched racemes, females up to 50 cm.
long, pendulent, males to 20 cm. long, axillary, often paired;
peduncle dense with long irritant hairs. Male flowers sessile, ca. 2
mm. long and broad; perianth pubescent, with irritant hairs and
punctiform cystoliths, 4-tepalled; stamens 4, with reflexed filaments;
pistillode ca. 1 mm. long; interfloral bracts 2-3 mm. long almost
linear, ca. 3 to each flower. Female flowers sessile, flabellately
arranged in one row; perianth ca. 1 mm. long, slightly less broad,
4—lobed, subglabrous but armed with irritant hairs and elongated
cystoliths; tepals 4, 3 ca. 1 mm. long, and much smaller; ovary ca.
1.25 mm. long and | mm. broad; stigma ligulate, to 8 mm. long,
Straight; interfloral bracts minute. Achenes to 5 mm. long, ca. 4
mm. broad, very slightly asymmetrically ovoid, fairly thick, peri-
carp warty generally, rarely smoth, nearly half covered by the per-
sistent perianth, reflexed; the fleshly flabellum well armed with
irritant hairs.

An endemic of Borneo, this species is common at about the 1,500
metres contour, particularly on the Pinosuk Plateau of Mt.
Kinabalu. Its short petioled, large obovate lamina and long pen-
dulent purplish female inflorescences dense with irritant hairs are
fortunately very conspicuous in the forest! Vernacular name
Tahipoi.

15

W —L. Chew — Monograph of Dendrocnide

female flowers;

leaf-bases; C =

twig; B =
male interfloral bracts.

achene; E

D =

Fig. 3: D. oblanceolata: A

16 Gardens’ Bulletin, Singapore — X XV (1969)

SABAH :—Mt. Kinabalu: Kiau: Clemens 10074, Nov. 1915 (A, BO &
UC).—Mesilau River: Chew & Corner RSNB. 4985, 7048 & 7094 (K &
SING). Penibukan: Clemens 3103/1, Jan. 1933 (A, BM, BO, K, L, NY &
UC.—Sandakan Distr.: Batu Lima: Agama 1026, Nov. 1920 (type of L-.
oblanceolata: A, K & US).

SARAWAK :—Baram Distr.: Batu Gading: Anderson S. 20242, July
1964 (SAR & SING).—Kapit Distr.: Bukit Raya Anderson 14375, Aug.
1961 (SAR).

KALIMANTAN :—Berau: Kostermans 21756, Nov. 1963 (L).—West
Koetai: G. Palah: Endert 3510, Sept. 1925 (BO).

(3) Dendrocnide elliptica (Merr.) Chew in op. cit. 203 (1965). —
Laportea elliptica Merr. in J. Arn. Arb. 35: 135 (1954).

Distinguishable by the following characters: (a) lamina peltate.
obovate, sericeous on both surfaces, (b) female flowers sessile,
flabellately arranged in one row, (c) achenes smooth, flat, with the
small perianth as a cupule at the base (d) margin of lamina evenly
and minutely dentate.

Distribution: Borneo and Philippine Islands.

Shrubby tree to 10 metres high; trunk ca. 30-45 cm. diam. Twigs
velutinous, especially at apices; with irritant hairs; petiolar scars
very prominent; bark dark grey. Lamina (13-) 15-25 (-35) cm.
long, (7-) 10-15 (—18) cm. broad; broad elliptic to obovate;
chartaceous to slightly coriaceous; sericeous, hairs mainly on the
veins, dense on the abaxial side, less so on the other, intermixed
with irritant hairs; young ones very dense sericeous; cystoliths
punctiform, evenly distributed on the adaxial surface, on the other
surface mainly on the veins if visible; base round, peltate, apex
acute to acuminate; margins evenly and minutely dentate; lateral
veins 12-15 pairs, evenly spaced, arched, quite prominent on the
abaxial side, less so on the other, basal pair less arched; the peltate
base with 3 short pairs of radiating veins; intercostals straight, well
spaced, anastomosing towards the mid-rib. Petioles (4-) 6-9 (—13)
cm. long; dense with punctiform cystoliths; densely sericeous with
irritant hairs when young. Stipules 1-2 cm. long, often early
caducous; bicarinate on the outside; scaphiform; coriaceous; young
ones dense sericeous with irritant hairs. /nflorescence unisexual:
branched racemes; (15—) 30-50 (-60) cm. long; the females longer
than the males; solitary, rarely paired; peduncular bracts unpaired,
up to 0.5 cm. long; peduncles thin, dense velutinous, well armed
with irritant hairs. Male flowers shortly pedicellate; 1-1.5 mm. long
and broad; pedicel ca. 0.5 mm. long, perianth sparsely
pubescent, sparsely armed with short irritant hairs, 4—tepalled:
tepals ovate; stamens 4, with inflexed filaments; pistillode half as
long as tepals; interfloral bracts triangular, very light pubescent on
the outside, ca. 1-1.5 mm. long, 1 to 2 subtending each flower.
Female flowers ssesile; 2-4 mm. long, ca. 0.5 mm. broad; very

W .-L. Chew — Monograph of Dendrocnide 17

4cm

Pr, pf

i
:
AG
Ma ,
= twig; B = female inflorescence; C = female:

Fig. 4: D. elliptica: A
flowers; D = achene

18 Gardens’ Bulletin, Singapore — X XV (1969)

slightly sunken in a row on flabellate peduncular receptacles;
perianth light to dense pubescent, often with irritant hairs,
- 4tepalled; tepals small ca. 0.75 mm. long, ovate, margins frilled;
ovary ca. 1 mm. long, asymmetrically ovoid; stigma 2-3 mm. long,
ligulate, thin; interfloral bracts very small, few, generally triangular.
Achenes ca. 3 mm. long, ca. 2 mm. broad; arranged in a row on
the reflexed flabellate peduncular receptacle; asymmetrically round,
very flattened; pericarp very smooth, very rarely slightly warty;
- edge smooth; apex abruptly acute, with long thin persistent styles;
perianth a minute cupule at the base.

It occurs in primary as well as in secondary forests from sea-
level to about 400 metres in altitude. Rather common at the base
of limestone hills in Sarawak. Vernacular names are: Hanjarapai
-(Dusun: North Borneo), Repai (Iban: Sarawak) and Tubun-
mumpaha (Ifugao: Philippines).

This species is typified by two collections of Elmer from North
Borneo viz., 21472 and 2/433. (The first collection was distributed
under the wrong number 2/1/72 by Elmer and the other was cited
erroneously as 1/433 by Merrill in the original description). I have
chosen as lectotype a specimen of the former ‘collection now
- deposted at the Arnold Arboretum.

Because of the peltate lamina, Merrill had this species confused
with D. peltata of Java. I have found the similarities between
these two species to be entirely superficial. The closest relative of
this species is actually D. crassifolia of the Philippines from which
it can quite easily be distinguished by the following characters: (a)
lamina smaller and obovate in this species and (b) teeth of lamina
also smaller than those of D. crassifolia, (c) base peltate, rounded.
whereas in D. crassifolia, they are cordate but not peltate.

NORTH BORNEO:—Kudat: Battah & Anthony SAN. 37678, Oct.
1963 (SING)—Lahad Datu: Muin Chai SAN 15536, July 1961 (L.).—
Ranau: Meijer SAN. 26442, June 1961 (K & L); Sinclair et al. 9253,
June 1957 (L & SING).—Semporna: Puasa 7662, Aug. 1937 (K, L, &
SING).—Tawau: Brand SAN. 24494 (K); Wood SAN. 17168, Feb. 1955
(BRI & L); Gibot SAN. 37035, Dec. 1963 (SING); Elmer 21433, Oct.

1922/March 1923 (A, BO, BR, C, F, K, L, M, MO, S, SING, U & Z),
21472, Oct. 1922/March 1923 (type: A, BO, K, L, SING & UC).

SARAWAK:—Bau: Anderson 12580, July 1960 (SAR).—Lundu:
Haviland 991 (K, SAR & SING).—Tiang Bekap: Anderson 12535, June
1960 (K, L, SAR & SING); Chew CWL. 647, July 1963 (A, K, L &
SING)—Mt. Braang: Haviland 153 (K), 347, Oct. 1889 (SAR).—Mt. Poi:
Clemens 20227, Sept. 1929 (A, K, MO, SAR & Z).

KALIMANTAN :—Mt. Pamathas: Korthals s.n. (L herb. 908-188-3266).
—Mt. Njapa: Kostermans 21302, Oct. 1963 (BO, L & SING).—South-east:
Winkler 3478 (L).

PHILIPPINES:—Luzon Island: Banaue: Conklin & Buwaya PNH.
80485, May 1963 (L).

(4) Dendrocnide crassifolia (C.B. Rob.) Chew in op. cit. 203 (1965).
—Laportea crassifolia C.B. Rob. in Philip. J. Sc. Bot. 5: 480

(1911).
Characteristics: (a) lamina ovate-cordate on long petioles; both

surfaces well covered by irritant hairs and dense punctiform cysto-
liths, particularly on mid-rib, (b) margin serrate, (c) apex shortly

W.—L. Chew — Monograph of Dendrocnide 1

acuminate, (d) inflorescences densely pubescent, (e) male flowers.
sessile to subsessile, dense with short irritant hairs, 4-tepalled, with
long interfloral bracts, (f) female flowers sessile, flabellately
arranged, (g) peduncular bracts also on flabellum.

Distribution: Philippine Island (and Talaud Islands?).

~ Dioecious tree to ca. 6 metres high; trunk ca. 15 cm. diam. Twigs

dense pubescent to velutinous, dense with irritant hairs; petiolar
scars very prominent. Lamina (23—) 30-38 (-40) cm. long, (15-)
20-25 (-30) cm. broad, broadly ovate, coriaceous; the upper surface
well covered with short stiff irritant hairs, the mid-rib denser;
punctiform cystoliths very dense; the lower surface sericeous to
velutinous, the mid-rib dense pubescent, intermixed with long
irritant hairs, similarly covered with punctiform cystoliths; base
cordate, the two basal lobes often touching; apex acuminate,
acumen 1-2 cm. long; margin serrate; lateral veins 11—13 pairs,
evenly spaced, rather closely set, quite prominent, the basal pair
not more prominent, reaching to about 4 the length of lamina
only; intercostals fairly straight, very numerous, quite prominent.
Petioles (7—) 12-15 (—23) cm. long, dense pubescent, well armed
with fairly long irritant hairs; cystoliths not visible. Stipules 2-3
cm. long, often woolly, well armed with irritant hairs, early cadu-
cous. Inflorescences unisexual, branched racemes, usually 15 cm.
long and broad, the males longer than the females, the latter not
pendulent, usually solitary; penduncles dense with fairly long
irritant hairs; peduncular bracts elliptic to linear, to 1 cm. long,
unpaired, often present on flabellum. Male flowers subsessile to
very shortly pedicellate, ca. 1 mm. long and broad, densely covered
with short irritant hairs, 4-tepalled; tepals broad ovate, slightly
ridged; stamens 4, with slightly reflexed filaments; pistillode ca.
0.5 mm. long, less broad; interfloral bracts ca. 1 mm. long, elliptic
to linear, also with irritant hairs, 3 or 4 to each flower: pedicel less
than 0.3 mm. long. Female flowers sessile, borne flabellately in one
row; perianth ca. 0.6 mm. long, ca. 0.5 mm. broad, 4-lobed, well
armed with irritant hairs, one much smaller than the other three;
ovary ca. 0.7 mm. long, ca. 0.4 mm. broad; stigma ligulate, to
1.5 mm. long; interfloral bracts absent. Achenes 2.5 mm. long,
orbicular-ovate, compressed, its surface more or less tuberculate:
flabellate receptacle at fruit to 1.2 cm. across.

I have not seen Clemens’ fruiting specimen. My description of
the achenes is therefore based on that of Robinson.

Merrill’s D. elliptica from Borneo and the Philippines is very
close to this.

H.J. Lam’s collection from the Talaud Islands (Lam 3007) comes
very close to this species. It differs in being less virulent and has a
slightly differently shaped lamina. His collection is referred to this
with reservation.

MINDANAO ISLAND—Davao Prov.: Mt. Mayo: Edano PNH.
11389, April 1949 (A, L & PNH).—Lanao Prov.: Mataline Falls:
Hutchinson 3925, March 1906 (type: K & US).

20

Gardens’ Bulletin, Singapore — X XV (1969)

Ne ep
rs &y DY ee A SS Ma
tx PT NEP IE
ef NX FY IY ys Fae hie
y IS | oy
Ao) _ :
tess >

2 Tae TO y Z Ss 2
raf. %) f ra 3 xa
a> iy, 6 as ES juraimi
EN ea . /; ard.
<< Nm YA 2

Fig. 5: D. crassifolia: A = twig; B

= female inflorescence; C = female
flowers.

WL. Chew — Monograph of Dendrocnide 21

(5) Dendrocnide meyeniana (Walp.) Chew in op. cit. 204 (1965).

Characteristics: (a) the lamina being oblong-ovate, base broadly,
shallowly cordate, often appearing subpeltate, the upper surface
covered sparsely with short irritant hairs, the lower surface pube-
scent to subglabrous, (b) the lateral veins often arising vertically
from the mid-rib, then arching out, (c) petioles, stipules and male
flowers dense pubescent, or subglabrous, (d) female flowers flabel-
lately arranged but often not in one plane.

Distribution: Formosa & Philippines.

(a) forma meyeniana—Urtica umbellata Blco. Fl. Filip. 696
(1837), non Bory (1802).—U. meyeniana Walp. in Nov. Act. Nat.
Cur. 19. Suppl. 1: 422 (1843).—U. ferox Blco. op. cit. ed. 2. 484
(1845), non Forst. f. (1786).—Laportea gaudichaudiana Wedd. in
Arch. Mus. Hist. Nat. Paris 9: 137 (1856) et in DC. Prodr. 16(1):
86 (1869).—L. pterostigma Wedd. in DC. Prodr. 16(1): 87 (1869).
—Urticastrum gaudichaudianum (Wedd.) O.K. & pterostigma
(Wedd.) O.K. op. cit. (1891).—L. meyeniana (Walp.) Warb. in
Perk. Fragm. Fl. Philip. 168 (1904).—L. diffusa C.B. Rob. in
Philip. J. Sc. Bot. 5: 482 (1911).—L. leyiensis C.B. Rob. in op. cit.
484 (1911).—D. diffusa (C.B. Rob.) Chew in op. cit. 203 (1965).

This form distinguishes itself by the undersides of the lamina
being pubescent to densely so.

Dioecious trees 5-7 metres high. Twigs light pubescent, with
short irritant hairs particularly at the nodes; petiolar scars large
and prominent. Lamina (12—) 20-25 (-50) cm. long, (8—) 10-15
(-30) cm. broad, oblong-ovate to oblong, rarely narrow ovate,
coriaceous, the upper surface well covered with very short
appressed irritant hairs, very dense with punctiform cystoliths, the
mid-rib with long and short irritant hairs, the lower surface dense
pubescent to woolly: base broadly shallowly cordate, appearing
subpeltate; apex acuminate, acumen ca. 2 cm. long: margin broadly
undulate, occasionally bluntly dentate; lateral veins ca. 13-17 pairs,
evenly spaced, arising almost vertically from mid-rib, then arching
out, very prominent, the basal pair not more promenent, reaching
to ca. + the length of the lamina; intercostals very numerous, close-
ly set. Petioles 5-20 cm. long, closely pubescent, with irritant hairs.
Stipules ca. 1 cm. long, appressed pubescent, rather foliaceous at
the margins, early caducous. Inflorescences branched racemes, up
to 25 cm. long, ca. 20 cm. broad, the females usually much nar-
rower and pendulent, solitary, rarely paired; peduncles pubescent,
well armed with irritant hairs; peduncular bracts triangular, to 1
cm. long, unpaired. Male flowers sessile, ca. 1 mm. long & broad,
densely pubescent, with few irritant hairs, 4-tepalled, stamens 4,
with very slightly reflexed filaments; pistillode ca. 0.5 mm. long,
0.3 mm. broad, obconical; interfloral bracts ca. 0.75 mm. long,
somewhat obovate, dense pubescent on the outside, 3 or 4 to each
flower. Female flowers sessile, flabellately arranged, perianth ca.
0.75 mm. long, ca. 0.5 mm. broad, 4-tepalled, the laterals much
broader than the medial ones, well armed with irritant hairs:
ovary ca. 0.6 mm. long, 0.4 mm. broad; stigma ligulate, ca. 2 mm.

22 Gardens’ Bulletin, Singapore — X XV (1969)

long; interfloral bracts 0.2 mm. long, often absent; the receptacle
usually curving at the sides, causing flowers to appear arranged
in more than one row. Achenes 2 mm. long and broad, sigmoid to
circular, somewhat flat, with lightly tuberculate pericarp, the lateral]
tepals narrow, at most half the length of the achene.

Having examined a large quantity of collections both from the
Philippines and Formosa, I find it impossible to maintain the
specific stati of the species diffusa, pterostigma and leytensis. They
are simply of one species whose leaf size varies quite considerably.

A paratype of Robinson’s L. densiflora (Merrill 998) has been
found to belong to this species. As the protologue of the former
species does not take Merrill’s collection into account, this transfer
does not involve any nomenclatural changes.

FORMOSA :—Takow: Henry 720 (A, K, NY & US).—South-west:

Swinhoe 37 (type of L. pterostigma: K)—Bankinsing: Faurie 606, Feb.
1914 (A).

PHILIPPINES:—Luzon Island: Benguet Prov.: Williams 1548, Nov.
1904 (A, K, NY & US). Laguna Prov.: Elmer 18235 (A, BO, F, K, MO,
NY & US). Tarlac Prov.: Curran 5149, Sept. 1906 (type of L. diffusa:
US). Sine loc.: Cuming 567 (type of L. gaudichaudiana: CGE & K).—
Mindoro Island: Baco River: Merrill 998, April 1903 (paratype of L.
densiflora: K & NY).—Leyte Island: Palo: Elmer 7351 (type of L.
leytensis: A, BO, E, FI & K).—Umingan: Pangasinan: Merrill Sp. Blco.
401, May 1914 (A, BO, F, K, L, MO, NY & US).

(b) forma subglabra (Hay.) Chew, comb. nov.—Laportea minda-
naensis Warb. in Perkins, Frag. Fl. Philip. 168 (1904).—L. batanen-
sis C.B. Rob. in op. cit. 481 (1911); Li in Pac. Sc. 7: 182 (1953).—
L. subglabra Hay. in J. Coll. Sc. Tokyo 30(1): 278 (1911).—L.
platyphylla Merr. in Philip. J. Sc. Bot. 11: 176 (1916).—L.
kotoensis Hay. ex Yamam. Suppl. Ic. Pl. Formos. 1: 2 (1925).—
L. pterostigma Wedd. forma subglabra (Hay.) Li, Woody FI.
Taiwan 135 (1963), basionym.—D. batanensis (Rob.), D. platy-
phylla (Rob.), D. subglabra (Hay.) Chew in op. cit. 202-206 (1965).

This form differs from the type in the lamina being subglabrous
to glabrous beneath.

In lamina size, this taxon is perhaps the most variable in the
genus. It ranges from about 8 to 45 cm. In vestiture, it varies from
absolute baldness to somewhat puberulous. All the species here
reduced have been very thoroughly investigated and have been
found to merge into each other so very imperceptibly that no
discontinuity can be detected between them.

H.-L. Li (1963) had this correctly reduced to a form of L. ptero-
stigma (which is in this work considered conspecific with D.
meyeniana). I accept his treatment.

FORMOSA :—Koshun Prov.: Kurasu: Wilson 11015, Nov. 1918 (US).
—Taito-cho: Kotosho: Tanaka 10406, March 1931 (UC).

PHILIPPINES:—Batan Islands: Fenix 3719, May-June 1907 (US).—
Catanduanes: Ramos & Edano 75581, July-Sept. 1928 (NY, SING &
UC).—Jolo Island: Ramos & Edano 44364, Sept. 1924 (A, NY & US).—
Luzon: Mt. Bulusan: Elmer 15608, Dec. 1915 (A, BO, C, FI, L, MO,
NY, S, U, UC, US & Z).—Mindanao: Taumu: Warburg 14702 (type of
L. mindanaensis: NY)—Mindoro: Mt. Yagaw: Conklin PNH. 17554,
May 1953 (A & PNH).—Polilo Island: Fox PNH. 8993, Nov. 1948 (A,
L & PNH).—Panay: Libaco: Martellino & Edano 35387 (K & US).—
Samar: Ramos 24406 (type of L. platyphylla: K).

W—-L. Chew — Monograph of Dendrocnide 23

imm

‘Fig. 6: D. meyeniana f. meyeniana: A = twig; B = male flower; C =
female flower; D = achene.

24 Gardens’ Bulletin, Singapore — X XV (1969)

(6) Dendrocnide densiflora (C.B. Rob.) Chew in op. cit. 203 (1965).
—Laportea densiflora C.B. Rob. in Philip. J. Sc. Bot. 5: 479
(1911).—L. subpeltata C.B. Rob. in op. cit. 485 (1911).—D. sub-
peltata (C.B. Rob.) Chew in op. cit. 206 (1965).

This species is notable by its mid-rib on the underside of the
lamina and the petiole being covered with a dense row of long
irritant hairs and the female flowers flabellately borne in more
than one row.

Distribution: Philippines (endemic!).

Dioecious tree to 6 metres. Twigs closely brown pubescent, with
few irritant hairs; petiolar scars prominent. Lamina 20-35 cm.
long, 10-20 cm. broad, oblong rarely ovate, the upper surface
dense with punctiform cystoliths, with irritant hairs on mid-rib
and veins, the lower surface with irritant hairs on the midrib; base
usually broadly shallowly cordate, often appearing subpeltate; apex
acuminate, acumen to 0.5 cm. long; margin faintly dentate to
sinuate; lateral veins 13-15 pairs, arising almost vertically from
mid-rib, then arching strongly outwards, the basal pair not more
prominent, usually giving off two more faint pairs at the base:
intercostals numerous, closely set. Petioles to 15 cm. long, closely
pubescent, the lower side with a dense row of long irritant hairs.
Stipules to 2 cm. long, somewhat foliaceous at the margins, pube-
scent, early caducous. Inflorescences branched racemes, pendulent,
to 30 cm. long, axillary, often paired; peduncle dense with long
irritant hairs. Male flowers sessile, ca. 1.5 mm. long and broad,
puberulous, with few irritant hairs and punctiform cystoliths, 4-
tepalled; stamens 4, with slightly reflexed filaments; pistillode ca.
0.3 mm. long, obconical; interfloral bracts to 1 mm. long, some-
what triangular, ca. 2 to each flower. Female flowers sessile,
flabellately borne, often in more than one row, often appearing
congested; perianth ca. 0.5 mm. long, 0.3 mm. broad, 4-tepalled,
well armed with hyaline irritant hairs and elongated cystoliths;
tepals almost equal-sized; ovary ca. 0.5 mm. long and 0.3 mm.
broad, ovoid; stigma ligulate, ca. 1-2 mm. long, straight; interfloral
bracts minute. Achenes unknown.

Robinson kept densiflora and subpeltata apart on the grounds
that only in the former species, are the female flowers borne in
more than one row, a characteristic which I have now found also in
the latter species. Besides, this phenomenon is not always observ-
able in every individual receptacle.

This species is close to D. meyeniana particularly forma sub-
glabra from which it can easily be distinguished by the presence
of a dense row of rather long irritant hairs on the petioles and
mid-ribs.

Of the two collections cited by Robinson under this name, the
paratype Merrill 998 does not belong here: it is D. meyeniana
forma meyeniana.

MINDANAO :—Misami Prov.: Mt. Malindang: Mearns & Hutchinson
4586, May 1906 (type of L. subpeltata: NY & US).—Zamboanga Prov.:
San Ramon: Hallier 4714, Feb. 1904 (type of L. densiflora: L & NY).

NEGROS OR.:—Lake Balinsasayao: Britton PNH. 19599, Sept. 1953
(L & PNH).

W—L. Chew — Monograph of Dendrocnide

aK
SAS Bs ay
is

Fig. 7: D. densiflora: leaf.

25

26 Gardens’ Bulletin, Singapore — X XV (1969)-

(7) Dendrocnide rigidifolia (C.B. Rob.) Chew in op. cit. 205 (1965).
—Laportea rigidifolia C.B. Rob. in Philip. J. Sc. Bot. 5: 483
(1911).

Characteristics: Close to D. subclausa from which it can easily
be distinguished by the following: (a) lamina narrow elliptic,
glabrous on both surfaces, (b) flabellate receptacle of female
inflorescences not entirely curved over.

Distribution: Philippines (endemic! ).

Dioecious shrubby trees ca. 3 metres high. Twigs subglabrous,
very sparse with irritant hairs; petiolar scars very prominent.
Lamina (30-) 40-55 cm. long, 7-11 cm. broad, narrowly elliptic, .
rigid coriaceous, the upper side glabrous, very dense with puncti-
form cystoliths, the lower side sparse with very short irritant hairs,
otherwise also glabrous, the punctiform cystoliths only on veins;
base cuneate; apex very long drawn acute; margin smooth; lateral
veins ca. 10 pairs, distantly spaced, the basal pair quite prominent,
reaching to one-third the length of the lamina; intercostals nume-
rous, prominent, fairly distantly spaced. Petioles 6-13 cm. long
dense with punctiform cystoliths, here and there with few very
rigid irritant hairs, otherwise quite glabrous. Stipules to 3 cm. long, .
rather broad and somewhat foliaceous at the base, light puberu-
lous to glabrous. Inflorescences branched racemes, to 22 cm. long,
pendulent, axillary, solitary; peduncular bracts at the lower axils
to 1.5 cm. long; peduncle covered with rigid irritant hairs. Male
flowers shortly pedicellate, arranged loosely in pseudo-capitula of
ca. 8 flowers, ca. 1 mm. long and broad, well covered with irritant
hairs and punctiform cystoliths, 4-tepalled; stamens 4, slightly
reflexed at the apex; pistillode ca. 0.75 mm. long, 0.5 mm. broad,
obovoid: pedicels at most 0.5 mm. long; interfloral bracts 2-3 mm.
long, elliptic, arranged loosely around the pseudo-capitulum.
Female flowers sessile, flabellately arranged in one row; perianth
ca. | mm. long, ca. 0.75 mm. broad, 4-lobed, well covered with
irritant hairs and cystoliths, tepals equal-sized; ovary ca. 1 mm.
long, 0.5 mm. broad, ellipsoid: stigma ligulate, ca. 5 mm. long,
Straight; interfloral bracts minute; the flabellate receptacle not
strongly curved over, bearing very numerous long irritant hairs on
one side and long peduncular bracts on the other side. Achenes
obliquely orbicular, compressed, 5 mm. diam., minutely tuber-
culate.

I am not entirely satisfied with the status of this species. The
differences between this and D. subclausa, as pointed out by Robin-
son, are no doubt observable; but they are of such a nature that
they are perhaps manifestations of developmental stages of one
species. However, as this species is known only by the two collec-
tions cited here, a solution to this problem will have to await.
further investigation in the field.

LUZON ISLAND :—Benguet Prov.:—Sablang: Fenix 12748, Nov.-Dec.

1910 (BO, E, F, L, NY & US). Mt. Santo Tomas: Williams 991, June 1904.
(type of L. rigidifolia: US).

WL. Chew — Monograph of Dendrocnide af

imm

Wiv Awe
Or eyreres
:

SE
=, SEF

6cm

Fig. 8: D. rigidifolia: A = twig; B = male flower; C = female flower.

28 Gardens’ Bulletin, Singapore — X XV (1969)

(8) Dendrocnide subclausa (C.B. Rob.) Chew in op. cit. 206 (1965).
—Laportea subclausa C.B. Rob. in Philip. J. Sc. Bot. 5: 486
(1911).—L. pendula Merr. in Philip. J. Sc. Bot. 13: 270 (1918).

Characteristics: (a) lamina rigid coriaceous, glabrous on the
upperside, pubescent on the lowerside, (b) male flowers shortly
pedicellate, in pseudo-umbels subtended by bracts to 1.5 mm. long,
(c) female flowers flabellately borne in one row, (d) achenes large,
pyriform, with a smooth pericarp, almost enclosed by the enlarged
receptacle densely covered with long irritant hairs.

Distribution: Philippines (endemic! ).

Dioecious shrubby trees to 3 m. Twigs glabrous to dense pube-
scent. sparsely covered with irritant hairs; petiolar scars prominent.
Lamina (20—) 25-35 (—53) cm. long, (7—) 9-15 (—23) cm. broad,
elliptic or ovate, coriaceous, the upper side glabrous, very dense
with punctiform cystoliths, the lower side pubescent to sericeous,
with punctiform cystoliths on the veins; irritant hairs mainly on
the lowerside; base round to cordate, rarely truncate; apex shortly..
bluntly acuminate; margin smooth; lateral veins (10—) 13-15 pairs,
widely spaced, the base usually 5—veined; intercostals prominent,
straight, numerous. Petioles (S—) 9-15 (—23) cm. long, dense with
punctiform cystoliths, subglabrous and sparse with irritant hairs.
Stipules 1-3 cm. long, scaphiform, often cordate at the base, mar-
gin rather foliaceous, light pubescent. Inflorescences branched
racemes, females up to 75 cm. long, pendulent, males much
shorter but broader, axillary, solitary, peduncular bracts at the
lower axils to 1.5 cm. long, peduncle dense with long rigid irritant
hairs. Male flowers shortly pedicellate, arranged in pseudo-capitula
of 8-10 flowers, ca. 1 mm. long and broad, well covered with long
irritant hairs and punctiform cystoliths, 4-tepalled; stamens 4,
slightly reflexed at the apex; pistillode ca. 0.75 mm. long, 0.5 mm.
broad. obovoid; pedicel ca. 0.5 mm. long; interfloral bracts to 2-3
mm. long, elliptic, encircling the pseudo-capitulum. Female flowers
sessile, flabellately arranged in one row: perianth ca. 0.75 mm.
long and broad, 4-tepalled; lightly covered with irritant hairs and
elongated cystoliths, tepals equal sized; ovary ca. 1 mm. long, 0.5
mm. broad, ellipsoid: stigma ligulate, 6-8 mm. long, quite straight;
interfloral bracts minute; the fleshy flabellate receptacle well curv-
ed, bearing irritant hairs only on the outside. Achenes to 4 mm.
long, 3 mm. broad, bloated ellipsoid to ovoid, with rather smooth
pericarp, on a very short stipe, almost totally enclosed by the
fleshy, greatly recurved receptacle up to 1 cm. in diameter.

This species has a very wide range of ecological tolerance. It
occurs along roadsides at 500 metres in altitude to mossy forests
well above 1,000 metres. The flabellate receptacle finds its greatest
development in this species. Reaching a size of 1 cm. in diameter,
it is so greatly curved over that it almost entirely encloses the
flowers and achenes, bearing on its outside numerous long irritant
hairs.

I find no significant difference between Merrill’s L. pendula and
this species the closest ally of which is perhaps D. rigidifolia.

W -—L. Chew — Monograph of Dendrocnide 29

Fig. 9: D. subclausa: A = leaf; B = female inflorescence; C = male
flowers.

30 Gardens’ Bulletin, Singapore — X XV (1969)

CATANDUANES ISLAND :—sine loc.: Ramos 30229, Nov.-Dec. 1917
(type of L. pendula: K & US).

LUZON ISLAND :—Isabela Prov.: San Mariano: Clemens 16703, April
1926 (UC).—Laguna Prov.: Mt. Maquiling: Elmer 17972, July 1917 (A,
BO, C, F, FI, K, L, MO, NY, S, U, UC, US & Z); Bterill 6290, Feo.
1909 (type of L. pendula: K, NY & US).—Sorsogon Prov.: Mt. Bulusan:
Elmer 14974, Nov. 1915 (A, BM, C, F, K, L, S, US & Z).

MINDANAO ISLAND:—Agusan Prov.: Tungao: Anonuevo PNH.
13733, May 1950 (A, L & PNH).

PANAY ISLAND:—Capiz Prov.: Jamindan: Ramos & Edano 30876,
April-May 1918 (L).

(9) Dendrocnide venosa (Elm.) Chew in op. cit. 207 (1965).—
Laportea venosa Elm. Leafl. Philip. Bot. 3: 878 (1910).—L.
monticola Merr. in Philip. J. Sc. Bot. 10: 299 (1915).—L. tripli-
nervia Merr. in op. cit. 300 (1915).—L. conduplicata Elm. Leafi.
Philip. Bot. 9: 3237 (1934).

Characteristics: A species notable for its shortly petiolate leaves
with few veins; inflorescences shorter than the leaves; male flowers
4-tepalled, subsessile; female flowers borne flabellately in one row
with receptacles becoming very fleshy and greatly curved at
maturity.

Distribution: Philippines (endemic! ).

Dioecious trees less than 10 metres high. Twigs glabrous, with
punctiform cystoliths usually at the nodes, almost deviod of irritant
hairs; petiolar scars very prominent. Lamina (10—) 15-20 (-23)
cm. long, 5-8 cm. broad, elliptic to ovate, somewhat coriaceous,
‘the upper-side glabrous, dense with punctiform cystoliths, the lower
side well covered with very short appressed irritant hairs and
‘elongated cystoliths; base broadly cuneate; apex acuminate, acumen
ca. 0.5 cm. long; margin slightly undulate; lateral veins 6-8 pairs,
‘the basal pair not more prominent, reaching to one-third the length
‘of the lamina; intercostals numberous somewhat anastomosing.
Petioles 3-5 cm. long, glabrous, well covered with punctiform
cystoliths, almost devoid of irritant hairs. Stipules 0.5 cm. long,
sdense pubescent, very early caducous. Inflorescences unisexual,
branched racemes, up to 10 cm. long and broad, axillary, paired;
peduncle thin, sparse with appressed irritant hairs, otherwise quite
glabrous. Male flowers subsessile to sessile, ca. 1.5 mm. long and
broad, subglabrous, with few appressed irritant hairs, 4-tepalled;
stamens 4, with slightly reflexed filaments; pistillode ca. 0.5 mm.
long, obpyramidal; interfloral bracts minute. Female flowers shortly
pedicellate, somewhat flabellately arranged in one row, few in each
group; perianth ca. 0.8 mm. long, 0.4 mm. broad, 4-tepalled, lightly
covered with very short irritant hairs, the laterals broad ovate,
greatly concave, the medials narrow ovate; ovary ca. 1 mm. long
and broad; stigma ligulate, ca. 5 mm. long; interfloral bracts very
-small; the lower portion of the pedicels fused to form an irregular-
ly shaped somewhat flabellate receptacle. Achenes ca. 3 mm. long
and broad, almost round, flat, with warty pericarp, only the base

-covered by the tepals, greatly reflexed; pedicels fleshy, the lower
fused portion (somewhat flabellate receptacle) becoming fleshy
cand greatly curved.

W.—L. Chew — Monograph of Dendrocnide

achene.

= female flower; C =

ponies
twig; B

Fig. 10: D. venosa: A

32 Gardens’ Bulletin, Singapore — X XV (1969)

I have examined the type materials of the four names enumerat-
ed above and have found no difference of any significance between
them. They are referable to one species sufficiently characterised
by the greatly enlarged and fleshy receptacles which are not quite
flabellately shaped. In this attribute, this species approaches D.
urentissima of China and Tonkin from which it differs in numerous
other characteristics such as the number of tepals in the male
flowers, the shape, size and vestiture of the lamina. In vegetative
characteristics, this species can perhaps be mistaken for D. stimu-
lans, but a closer look will tell them apart.

LUZON :—Ifugao Prov.: Mt. Polis: McGregor 19765, Feb. 1913 (type
of L. monticola: US); Sandkuhl 298, Jan. 1915 (A).—Tayabas Prov.:
Mt. Pulor: Ramos 19462, Jan. 1913 (type of L. triplinervia: L & US).—

Sorsogon Prov.: Mt. Bulusan: Elmer 17152, Sept. 1916 (type of L.
conduplicata: A, BO, C, F, FI, K,L, MO, NY, S, U, UC, US & Z).

MINDANAO :—Davao Prov.: Mt. Apo: Elmer 11948, Oct. 1909 (type
of L. venosa: A, BO, E,:F, FI, K, L, US, WRSL & Z).

NEGROS ORIENTAL:—Lake Balinsasayao: Britton PNH. 19604, Sept.
1953 (L & PNH).

(10) Dendrocnide urentissima (Gagn.) Chew in op. cit. 207 (1965).
—JLaportea urentissima Gagn. in Bull. Soc. Bot. Fr. 75: 3 (1928)
et in Lecomte, Fl. Gen. Indo-China 5: 856 (1929); Wu in Acta
Phytotax. Sinica, 6 (3): 276 (1957).—L. chingiana Hand.-Mazz.
in Sinensia 2 (1): 1 (1931).—Dendrocnide chingiana (Hand.-
Mazz.) Chew in op. cit. 202 (1965).

Characteristics: (a) lamina usually broadly ovate, as long as
broad, cordate base, upper surface with short dense irritant hairs,
lower surface dense velutinous, (b) lateral veins 8 pairs, (c) inflore-
scences long, pendulent; peduncles dense pubescent and with dense
irritant hairs, (d) female flowers subsessile due to partial fusion of
pedicels to form flabellum that becomes fleshy at fruit, (e) male
flowers sessile, 5-tepalled, dense pubescent with irritant hairs, (f)
achenes round to sigmoid, warted, slightly bloated, with perianth
as a cupule at the base.

Distribution: China & Vietnam.

Dioecious trees to ca. 4 metres high. Twigs dense pubescent, or
lightly so, with irritant hairs; petiolar scars very prominent.
Lamina (14-) 16-22 (—27) cm. long, (14—) 16-22 cm. broad, broadly
ovate, coriaceous, upper surface dense with short irritant hairs,
lower surface dense velutionous, both sides with punctiform cysto-
liths; base usually strongly cordate; apex acuminate, acumen to
about 1 cm. long; margin very slightly and faintly toothed: lateral
veins ca. 8 pairs, the basal pair usually slightly more prominent:
intercostals straight, numerous. Petioles (8—) 10-15 (—20) cm. long,
dense pubescent; irritant hairs short, fairly numerous. Stipules to
1 cm. long, pubescent, caducous. Inflorescences unisexual, branched
racemes, 20-30 cm. long, ca. 10 cm. broad, pendulent, axillary.
solitary or paired; peduncle dense pubescent, irritant hairs long
and numerous. Male flowers sessile to subsessile, ca. 2 mm. long
and broad, 5-tepalled: perianth fairly dense pubescent, with few

W —-L. Chew — Monograph of Dendrocnide dd

Fig. 11: D. urentissima: A = leaf; B = female inflorescence; C = female
flower; D = male flower.

34 Gardens’ Bulletin, Singapore — X XV (1969)

irritant hairs at the apex: tepals ovate; stamens 5, with slightly
reflexed filaments: pistillode half the length of tepals; interfloral
bracts minute, irregular in number. Female flowers subsessile,
somewhat flabellately arranged, 3-5 in each group, perianth ca. 1
mm. long and broad, 4-lobed, pubescent and well armed with
irritant hairs; tepals 4, one large, ca. 1 mm. broad, 2 medium, ca.
0.5 mm. broad and 1 small, ca. 0.3 mm. broad; ovary 1 mm. long
and 0.5 mm. broad; stigma ligulate, to 3 mm. long, generally
curled; interfloral bracts ca. 0.3 mm. long and broad, ca. 2 to each
flower: free parts of pedicels to 0.3 mm. long, glabrous. Achenes
to 3.5 mm. long and broad, round to somewhat sigmoid, fairly
thick; pericarp warty; perianth as a minute cupule at the base;
stigma greatly reflexed; achene reflexed over the slightly fleshy
flabellum.

It occurs as riverine, trees in lower montane forests of South-
east Continental Asia about 500 metres in altitude. It has been
collected from limestones.

I agree with Wu (1957) that L. chingiana is similar to this species.

Being closely related to D. basirotunda, also of Continental
Asia, this species links section Dendrocnide to Sarcopus. The
partial fusion of the female pedicels into tiny flabella which become
slightly fleshy at maturity puts this species actually right in
between these two sections. I place this in the former section
because, it seems to me, the flabella are slightly more noticeable
than is the succulence of them.

CHINA:—Yunnan: Jenn-yeh Hsien: Wang 80006, Oct. 1936 (A).—
Kwangsi: Sin-shu: Ching 7329, Sept. 1928 (type of L. chingiana: A).

Vietnam:—Tonkin: Dong-dang: Balansa 585 (type of L. urentissima:
K & P). Van Linh: Petelot 6689, Dec. 1940 (US).—Annam: Phanrang
Prov.: Canna: Poilane 8930, Nov. 1923 (A & P) & 12505, Oct. 1925 (A).

(11) Dendrocnide basirotunda (Wu) Chew, comb. nov. —
Laportea basirotunda Wu in Acta Phyiotax. Sinica, 6 (3):
276 (1957), basionym.

Characteristics: Close to D. urentissima from which it differs
in: (a) lamina ovate, slightly longer than broad, both surfaces
almost glabrous except for some irritant hairs on veins, (b)
inflorescence as long as broad, not pendulent, (c) peduncles light
puberulous, with long stiff irritant hairs, (d) male flowers sub-
glabrous, (e) female flowers fasciculate, pedicels elongating and
swollen pedicel.

Distribution: China, Burma and Thailand.

Dioecious trees to 15 metres high. Twigs glabrous, almost
devoid of irritant hairs; petiolar scars very prominent. Lamina
(12-) 16-20 (—24) cm. long, (9—-) 12-14 (-17) cm. broad, ovate,
coriaceous, both surfaces almost glabrous except for some irritant
hairs on veins; cystoliths punctiform; base rounded, sometimes
slightly cordate; apex shortly, bluntly acuminate, acumen to
1 cm. long; margin slightly and irregularly sinuate, otherwise
quite entire; lateral veins 8-9 pairs, the basal pair not more

W —L. Chew — Monograph of Dendrocnide

Fig. 12: D. basirotunda.

36 Gardens’ Bulletin, Singapore — X XV (1969)

prominent; intercostals straight, numerous. Petioles (4-) 8-10
(-13) cm. long, puberulous; irritant hairs short, not numerous.
Stipules to 1 cm. long, velutinous, caducous. Inflorescences
unisexual, branched racemes, up to 15 cm. long and broad, not
pendulent, axillary, solitary or paired; peduncle light puberulous
to glabrous but with plenty of long, stiff irritant hairs. Male
flowers sessile, ca, 1.5 mm. long and broad; perianth subglabrous,
with few irritant hairs and punctiform cystoliths at the apex,
S-tepalled; stamens 5, filaments reflexed slightly at the apex;
pistillode half length of tepals; interfloral bracts ca. 0.3 mm.
long, triangular, 3 or 4 each flower.Female flowers very shortly
pedicellate, about 3 flowers in each fascicle; perianth ca. 1 mm.
long and slightly less broad, 4-lobed, light pubescent, well armed
with short irritant hairs; tepals 4, 3 ca. 1 mm. long, | ca. 0.5 mm.
long, all rather narrow; ovary ca. 1 mm. long, half as broad;
stigma ligulate, ca. 2-3 mm. long, rather straight; interfloral bracts.
minute. Achenes to ca. 3.5 mm. long, ca. 2.5 mm. broad,
asymmetrically ovoid, sigmoid to almost round, fairly thick;
pericarp warty; perianth as a minute cupule at the base; stigma
greatly reflexed; achene reflexed over the greatly elongated and
swoolen pedicel.

This species occurs as shade trees in dense primary forests
near streams above 500 metres in altitude. It is said to be
commonest at about the 1000-metre contour.

- Wu cited four collections of C. W. Wang as types for this
species, viz. Wang 78665, 79201, 79935 and 26. I have selected
a duplicate of Wang 79935 deposited at the Arnold Arboretum
as the lectotype.

CHINA:—Yunnan Prov.: Jenn-yeh Hsien: Wang 79935, Oct. 1936
(lectotype in A). Che-li Hsien: Wang 78665 & 79201, Sept.-Oct. 1936 (A)-

BURMA:—Shan State: Keng Tung: Rock 2040, Jan. 1922 (A, UC &
US).

THAILAND :—Klang-dong: Larsen 9420, Jan. 1962 (SING)—Phu
Krading: Dee 452, Oct. 1951 (BKF).

(12) Dendrocnide sinuata (BI.) Chew in op. cit. 206 (1965). —
Urtica ardens Bl. Bijdr. 504 (1825), non Link (1822). —
U. sinuata Bl. Bijdr. 505 (1825). — U. crenulata Roxb. FI.
Ind. 3: 591 (1832), non Sev. (1785). — U. pulus Steud. Nom.
Bot. ed. 2. 2: 735 (1841). — U. obnoxia Hassk. Cat. Hort.
Bog. Alt. 305 (1844). — Laportea crenulata Wedd. in Arch.
Mus. Hist. Nat. Paris 9: 133 (1856) et in DC. Prodr. 16 (1):
85 (1869). — L. sinuata (Bl.) Mig. Ann. Mus. Bot. Lugd.
Bat. 4: 301 (1869). — Urticastrum sinuatum (Bl.) O.K. op. cit.
(1891). — L. ardens J. J. Smith in Meded. Dept. Landb. 10:
684 (1910). — L. pustulosa Ridl. in J. As. Soc. Str. Br. 82:
194 (1920). — L. integrifolia Wu in Acta Phytotax. Sinica,
6 (3): 277 (1957). — D. pulus (Steud.) Chew in op. cit. 205
(1965).

Characteristics: (a) lamina elliptic, oblong to rhombic, glabrous

except for the rather long, slightly appressed irritant hairs, (b)

margin of lamina sinuate to dentate, rarely smooth, (c) flowers

W -L. Chew — Monograph of Dendrocnide 37

35cm \

Fig. 13: D. sinuata: leaf-form (typical).

Gardens’ Bulletin, Singapore — X XV (1969)

Fig. 14: D. sinuata: fruiting twig; leaf-form atypical.

38

W .—-L. Chew — Monograph of Dendrocnide 39

pedicellate, the male 4-tepalled, (d) achenes 3-5 mm. long, bloated,
warty, pyriform, on non-fleshy pedicels, the base slightly covered
by the persistent perianth.

Distribution: India, Ceylon, China through South-east Asia to
Malay Peninsula, Sumatra, Java and Bali.

Trees ca. 10 metres high. Twigs glabrous, with irritant hairs
pointing downwards. Lamina (15—) 20-65 cm. long, 7-35 cm.
broad, elliptic to rhombic, rarely ovate or obovate, juvenile ones
often cordate, coriaceous, usually glabrous; irritant hairs sparse
on the adaxial surface, more on the veins of the other surface:
cystoliths punctiform; base cuneate, truncate to profoundly
cordate; apex long to short acuminate; margin sinuate to dentate:
lateral veins 10-15 pairs, the basal pair slightly more prominent.
Petioles (3—) 6-20 (-30) cm. long, with punctiform cystoliths, .
sparsely to densely covered with downward pointing irritant hairs.
Stipules ca. 2 cm. long, chartaceous, caducous. /nflorescences
unisexual, branched racemes with flowers in loose fascicles, males
5—15 cm. long, females 10-30 cm. long, axillary, solitary, rarely
paired; peduncle often dense with long irritant hairs, with minute
peduncular bracts. Male flowers pedicellate, 2-3 cm. long, 1-2 mm.
broad, lightly covered with irritant hairs, 4-tepalled; stamens 4,
on reflexed filaments; pistillode small. Female flowers very shortly
pedicellate; perianth 1-1.5 mm. long, 0.5 mm. broad, sparsely
covered with irritant hairs, 4-lobed; ovary ca. 1.5 mm. long;
stigma ligulate, ca. 2-5 mm. long; pedicels well armed with
irritant hairs. Achenes 3-5 mm. long, 2.5-4.5 mm._ broad,
asymmetrically pyriform, bloated, with warty pericarp, the base
slightly covered by the small perianth, rarely reflexed; pedicels
elongating greatly at fruit.

My concept of this species departs markedly from that of
J. J. Smith. I stretch it to include L. ardens which I consider
to be a juvenile stage of this species as observed in the field
by Koorders. I d onot agree that this is perhaps similar to either
Gaudichaud’s latifolia from the Pacific or Poiret’s gigantea, a
doubtful species of uncertain origin, although, I concede, it is.
closely related to the former.

The type material of L. pustulosa and L. integrifolia have
been examined and found to be minor variants of this very
variable species which is known throughout west Malesia as
the “‘pulus” or “‘pulutus’’.

INDIA:—Assam: Barnard MR. 33, Jan. 1936 (BM).—Bengal: Cowan

s.n., Nov. 1923 (E). Cult. Calcutta: Wall. Cat. 461] (BR, CGE, K & M).—
Sikkim: Clarke 36225C, Oct. 1884 (G).

CEYLON :—Nugatenne Gap: Alston 259 (C, K, MEL & UC).—sine
loc.: Thwaites 2200 (CGE, G, K & P).

BURMA:—Kachin Hills: Myitkyina: Lace 5553, Nov. 1911 (E & K).

CHINA:—yYunnan: Luh shuen-hsien: Wang 8/058, Nov. 1936 (type
of L. integrifolia: A)—Hainan: Lokwi: Lau 28363, Dec. 1936 (A). Ma
On Shan: McClure 7747, Oct. 1921 (K, MO & P).

VIETNAM:—Mt. Dinh: Pierre 4708, Dec. 1866 (A, BO, NY & P).—
Thua-Thien: Eberhardt 1260 (P).—Tonkin: Bon 90 (P).

THAILAND:—Doi Puka: Kerr 4906, Feb. 1921 (BM).—Patani:
Gwynne-Vaughan 373 (CGE).

40 Gardens’ Bulletin, Singapore — X XV (1969)

MALAY PENINSULA:—Perak: Allen 4879, Jan. 1963 (A, C, CGE,
G, K, L, NY & SING); Father Scortechini 1817 (type of L. pustulosa: K).

SUMATRA:—Tapanoeli: Rahmat si Toroes 5309, Aug. 1933 (A, NY &
US).—Sibolangit: Nur 7408, Aug. 1921 (BO, K, NY & SING).

ance ISLAND:—Flying Fish Cove: Ridley 158, Oct. 1901
(SING).

JAVA:—West: G. Tjermai: Backer 4820 (BO).—Semarang: Koorders
38849, Nov. 1900 (BO & L).—-sine loc.: unknown s.n., L. herb. 908—188-
3235 (type ? of U. sinuata: L); unknown s.n., L. herb. 908-188-3275 (type
? of U. ardens: L). :

BALI:—G. Kelatakan: Sarip 133 (BO & L).

(13) Dendrocnide luzonensis (Wedd.) Chew in op. cit. 203 (1965).

Characteristics: Close to D. sinuata from which it differs in:
(a) lamina smooth, entirely glabrous, narrowly elliptic to
oblanceolate, (5) flowers pedicellate, in fascicles, (c) irritant hairs
present only on flowers, (d) achenes ca.4 mm. long, warted bloated,
reflexed, (e) female pedicels curved, somewhat swollen progressively
towards the apex.

Distribution: Philippines (endemic! ).

(a) var. luzonensis — Laportea crenulata (Roxb.) Wedd. var.
luzonensis Wedd. in Arch. Mus. Hist. Nat. Paris 9: 133 (1856).
— L. luzonensis (Wedd.) Warb. in Perk. Fragm. FI. Philip-
168 (1905). — L. agusanensis Elm. Leafi. Philip. Bot. 8: 2860
(1915). — L. latilanceolata Elm. op. cit. 9: 3238 (1934).

Dieoecious trees. Twigs finely puberulous, without irritant hairs;
petiolar scars prominent. Lamina (9-) 15-20 (—25) cm. long,
(3—) 5—8 (-12) cm. broad, narrowly elliptic, elliptic to oblanceolate,
coriaceous, entirely glabrous, without irritant hairs, cystoliths of
adaxial surface punctiform, those of abaxial surface punctiform
and linear; base broad cuneate to rounded; apex acuminate,
acumen to 2 cm. long; margin smooth, rarely undulate; lateral
veins 8-10 pairs, usually chocolate coloured when dry, the basal
pairs only slightly less prominent; intercostals slightly anastomosing,
faint on the adaxial side. Petioles (2—) 3-5 (—10) cm. long,
without cystoliths, light puberulous. Stipules up to 1 cm. long,
coriaceous, caducous, puberulous. J/nflorescences unisexual,
branched racemes, up to 10 cm. long, slightly less broad, axillary,
solitary; peduncular bracts minute, ca. 0.5 mm. long; peduncle
glabrous, with punctiform cystoliths; irritant hairs absent. Male
flowers pedicellate, ca. 1.5 mm.long and broad; perianth glabrous
except for ca. 10 irritant hairs at apex, 4-tepalled; stamens 4,
with slightly reflexed filaments; pistillode thin; interfloral bracts
ca. 2 to each flower, small scaphiform; pedicels often greatly
elongated at anthesis. Female flowers pedicellate, 3-5 flowers
at each fascicle; perianth less than 1 mm. long, 4-lobed, with
punctiform cystoliths and long, broad irritant hairs; tepals 4,
2 large, 1 small and 1 minute; ovary ca. 1 mm. long; stigma
ligulate, up to 2 mm. long, slightly flattened at the broad base
of style, generally curled; interfloral bracts small, irregular in
number; pedicel ca. 1 mm. long, dilated at apex at maturity.

41

W -L. Chew — Monograph of Dendrocnide

JURAIM! DEL.

Fig. 15: D. luzonensis var. luzonensis: A = twig; B = male inflorescence,

achene

achene; E

female inflorescence, part; D

in cross-section.

part: ©

42 Gardens’ Bulletin, Singapore — X XV (1969)

Achenes up to 4 mm. long, 3 mm. broad, asymmetrically ovoid
pyriform, with warted pericarp, rather bloated, with persistent
perianth as a cupule at the base; persistent stigma reflexed over
side of achene; achene reflexed over fleshy pedicel.

An endemic of the Philippines Islands, it occurs in forested
valleys, usually near streams below 2,000 metres in altitude.

Vernacular names: Bungabung, Lipa, Lupa, Sagai and Sagay
de Bungabung.

Having examined the type collections, I conclude that Elmer’s
agusanensis and latilanceolata are both conspecific with Weddell’s
luzonensis.

LUZON ISLAND:—Balaan Prov.: Mt. Mariveles: R. Meyer 2631,
Feb. 1905 (BO, K, NY, SING & US).—Sorsogon Prov.: Mt. Bulusan:
Elmer 15206, Nov. 1915 (type of L. latilanceolata: A, BO, C, F, FI, K,
L, MO, Ss... UC. “Os"e Z).

MINDANAO ISLAND:—Agusan Prov.: Mt. Urdaneta: Elmer 14019,
Oct. 1912 (type of L. agusanensis: BO, E, F, L, MO & US).

SINE LOC.:—Cuming 522 (type of L. crenulata var. luzonensis: C
CGE, G, G-DC, K, L, M, MEL, MO & UPS).

(b) var. anacardioides (C. B. Rob.) Chew, stat. nov. — Laportea
anacardioides C. B. Rob. in Philip. J. Sc. Bot. 5: 478 (1911),
basionym. — D. anacardioides (C. B. Rob.) Chew in op. cit.
202 (1965).

This variety distinguishes itself from the type variety by the
lamina being sharply cuneate at the base and the basal pair of
lateral veins very short.

LUZON ISLAND:—Rizal Prov.: San Andales: Edano PNH. 48786,
Dec. 1926 (A,LB--NY %& UC).

MINDANAO ISLAND:—Davao Prov.: Santa Cruz: Williams 2766,
May 1905 (type of L. anacardioides: US).

>

(14) Dendrocnide peltata (Bl.) Mig. Jungh. 1: 30 (1851); Chew
in op. cit. 205 (1965).

Characteristics: (a) lamina peltate, ovate, densely villose to
glabrous on the lowerside, (b) female flowers pedicellate, in
fascicles, (c) matured achenes flat, smooth, with the small persistent
perianth as a cupule at the base.

Distribution: Christmas Island, Java, Bali and New Guinea.

(a) var. peltata. — Urtica peltata Bl. Bijdr. 496 (1825). —
Laportea laxiflora Wedd. & vriesiana Wedd. in DC. Prodr. 16 (1):
81 (1869). — Urticastrum peltatum (Bl.) O.K., laxiflorum (Wedd.)
O.K. & vriesianum (Wedd.) O.K. Rev. Gen. Pl. 2; 635 (1891).
— Laportea peltata auct. non Decne.: Wedd. in Arch. Mus.
Hist. Nat. Paris 9: 126 (1856); Mig. Fl. Ind. Bat. 1 (2):
230 (1859); Wedd. in DC. Prodr. 16 (1): 80 (1869); Mig. Ann.
Mus. Bot. Lugd. Bat. 4: 302. tab. 10. fig. 1 (1869); J. J. Smith
in K. & V. Bijdr. 12: 678 (1910); Backer & Bakh. v./d. Brink,
Pl Java' 2: 38 (1965).

Distribution: Java, Bali and New Guinea.

Dioecious trees to 30 metres. Twigs puberulous, pubescent to
tomentose, with minute irritant hairs. Lamina (13—) 16-25 (—30)
cm. long, 8-) 12-16 (-20) cm. broad, generally broad ovate to

W —L. Chew — Monograph of Dendrocnide 43

0
Ro, DAS
eA
\
SS
(NYE!
q

QO al
(> on
che i AB
a
BQ Le Vee 2,
VA,
6%

Fig. 16: D. peltata var. peltata: A = twig; B = female flower; C = achene.

44 Gardens’ Bulletin, Singapore — X XV (1969)

subrotundate, coriaceous, the upper surface almost glabrous, with
few dispersed irritant hairs, rarely pubescent, the lower surface
dense pubescent to villous, irritant hairs profuse; base distinctly
peltate; apex very shortly acuminate, acumen to 0.5 cm. long:
margin denticulate, becoming smooth at the base; lateral veins 7-10
pairs above the prominently tri-nerved base, rather prominent;
intercostals numerous, straight, fairly prominent. Petioles (3-)
8-12 (—15) cm. long, cystoliths not visible, tomentose, very rarely
glabrous, irritant hairs very sparse. Stipules 1-2 cm. long.
coriaceous with chartaceous undulate margins, pubescent on the
outside. Inflorescences unisexual, branched racemose, flowers in
few-flowered fascicles, (8—) 10-18 (—20) cm. long, (4) 8-10 (-15
cm. broad, axillary and behind the leaves, solitary, rarely paired:
peduncular bracts small to minute; peduncles pubescent, rarely
brown tomentose, with fairly long, rigid irritant hairs. Male
flowers pedicellate, ca. 1.5 mm. long, 1 mm. broad, pubescent,
with few irritant hairs at the apex, 4-tepalled; stamens 4, with
reflexed filaments; pistillode small; interfloral bracts to 0.25 mm.
long, narrow ovate, ca. 4 to each flower; pedicel to 1 mm. long
at anthesis, light puberulous. Female flowers pedicellate; 5—7
flowers in each fascicle; perianth with 4 decussate-imbricate very
narrow tepals, the two laterals ca. 0.75 mm. long, the third slightly
shorter and the fourth ca. 0.25 mm. long, very light puberulous,
with few irritant hairs; ovary ca. 1 mm. long, ca. 0.5 mm. broad;
stigma ligulate, ca. 1 mm. long; interfloral bracts ca. 0.25 mm.
long, narrow ovate, ca. 4 to each flower; pedicel to 1 mm. long,
almost glabrous. Achenes ca. 2 mm. long & broad, almost
round, flattened, smooth, persistent perianth forming a minute
cupule at the base, reflexed; pedicel not longer than 2 mm. at fruit.

This variety occurs from sea-level to about 1,300 metres, and
seems particularly common at about the 500-metre contour. In
New Guinea, it is found in both primary and secondary forests.

Carr recorded the use of the bark of this variety for loin cloths
by the Papuans! | |

Vernacular names: Boeroenjah, Lateng-lade; Galamo.

I have examined the type collections of this variety as well as
those of Weddell’s species cited above, and have found that
J. J. Smith was quite right in reducing the latter species.

This variety has hitherto been regarded by most botanists to
be the same as the Timor species Laportea peltata Gaud. ex
Decne. Having examined the holotype of Gaudichaud’s species
in the Paris herbarium, I conclude that while Blume’s species
shares with Gaudichaud’s the common characteristics of the
peltate hairy lamina, the two differ from each other quite clearly
in the achene. In the former species, the achenes are very flat
and smooth, while in the latter, the achenes are rather bloated
and warty. Besides, in Blume’s species, the fruiting pedicels are
not swollen as is the case in Gaudichaud’s Timor plant. Actually,
Gaudichaud’s plant is conspecific with D. moroides, an Australian
species.

WL. Chew — Monograph of Dendrocnide 45

JAVA:—West: Bandoeng: van Steenis 1686 (BO).—Central: Semarang:
Koorders 28026, June 1897 (BO & L).—East: Besoeki: Koorders 22426
(A, BO & L).—-sine loc.: unknown coll. 1345 (type of U. peltata: L); de
Vriese 6 (type of L. vriesiana: L & P); Zollinger 3136 (holotype of L.
laxiflora: G-DC).

BALI:—Gunong Kelatakan: Sarip 108, July 1918 (BO, L & VU).

NEW GUINEA:—West: Habbema Lake: Brass & _ Versteegh
10485, Oct. 1938 (A & BO). Nabire: Patema: Kanehira & Hatusima
11908, Nov. 1940 (A & BO). Vogelkop Peninsula: Kebor Valley: Koster
BW. 7097, 7138 & 7177, Oct. 1958 (L & LAE).—East: Morobe Distr.:
Bulolo: Hartley 10900, Nov. 1922 (L); Havel & Kairo NGF. 17150 (BRI,
L, LAE, NSW & SING).

PAPUA :—Kanosia: Carr 11127, Feb. 1935 (A, BM & L.)—Veiya: Carr
11723, March 1935 (A & BM).

(6) var. murrayana (Rendle) Chew, stat. nov. — Laportea
murrayana Rendle in Andrews, Monog. Christ. Isl. 189 (1900),
basionym. — Dendrocnide peltata subsp. murrayana (Rendle)
‘Chew in op. cit. 205 (1965).

Distribution: Christmas Island.

This variety differs from the type variety in the lamina which
though distinctly peltate is subglabrous to glabrous.

I have examined the holotype of. this taxon and am quite
Satisfied that this is a variant of D. peltata.

CHRISTMAS ISLAND:—Flying Fish Cove: Andrews 147, Feb. 1898
(type in BM & K).—Smith Point: Ridley 157, Oct. 1904 (K & SING).

(15) Dendrocnide carriana Chew, spec. nov.

Characteres: D. peltatae affinis, sed a qua laminis haud peltatis
subtus glabris vel pubescentibus haud villosis differt.

Distribution: Philippines, Lesser Sunda Islands, Molucas and
New Guinea.

Dioecious trees to 27 metres high. Twigs light pubescent, soon
glabrous, without irritant hairs. Lamina (8-) 10-15 (-17) cm. long,
(6—-) 8-10 (-14) cm. broad, elliptic, oblong to pentagonai-ovate,
coriaceous, with mounted irritant hairs mainly on mid-rib and
Jateral veins, rest of lamina glabrous, rarely pubescent; base
cuneate to cordate; apex acuminate, acumen to | cm. long;
margin minutely, irregularly dentate to crenate; lateral veins
8-11 pairs, prominent, the basal pair more prominent, arising
slightly above the base, reaching to half the length of the lamina,
the base with 2 or 3 more pairs of very short veins arising from
the same point. Petioles (5—) 8-12 (—14) cm. long, covered with
numerous very short appressed irritant hairs and a few long
mounted ones. Stipules less than 1 cm. long, entirely glabrous,
caducous. Inflorescences to 9 cm. long, 7.cm. broad, axillary,
solitary; peduncles pubescent, with few mounted irritant hairs,
with minute peduncular bracts. Male flowers pedicellate; perianth
ca. | mm. long, 0.8 mm. broad, pubescent, often densely, 4-tepalled:
stamens 4, with reflexed filaments; pistillode ca. 0.6 mm. long,
obconical; interfloral bracts minute; pedicel to ca. 1 mm. long,
not fleshy. Female flowers pedicellate, ca. 7 flowers in each

46 Gardens’ Bulletin, Singapore — X XV (1969}

Fig. 17: D. carriana: A = twig; B = male flower; C = female flower;
D = bisexual flower; E = achene.

W—L. Chew — Monograph of Dendrocnide 47

fascicle; perianth ca. 0.8 mm. long, 0.4 mm. broad, 4-tepalled,
lightly covered with irritant hairs, the lateral tepals narrow,
twice as long as the. medial ones; ovary ca. 0.6 mm. long,
ca. 0.4 mm. broad; stigma ligulate, ca. 1.3 mm. long, curved;
interfloral bracts minute; pedicel ca. 0.5 mm. long. Achenes
2 mm. long, 1.5 mm. broad, ovoid, flat, with smooth pericarp,
shortly stipitate, sometimes surrounded by 2-3 stamens with
long filaments; perianth about as long as achene; pedicel to
2 mm. long, not fleshy.

Amongst the characteristics that distinguish this species, the
most interesting is the occasional development of stamens in the
female flowers! This phenomenon, which has no parallel in the
genus, seems to occur only after fertilization, especially when
the achene is almost matured.

Like D. peltata, with which it is closely related, this species
also has a very puzzling disjunct distribution. I have examined
the collections very carefully and am convinced that they are
conspecific despite their odd distribution.

PHILIPPINES:—Mindoro: Mt. Yagaw: Conklin PNH. 18933, Sept.
1953 (A, L & PNH).

LESSER SUNDA ISLANDS:—Bali: Mt. Batukau: Kostermans & al.
KKSS. 129, June 1958 (A, BO, L & SING).—Lombok: Sembami: Rensch
313 (BO)—Timor: Nasimetan: Neth. Ind. F. Service bb. 27110, March
1939 (SING).

MOLUCCAS:—Ternate: Batjan: de Haan 139, Sept. 1937 (L).

NEW GUINEA (NORTH-EAST) :—Morobe Distr.: Sambanga: Clemens
7153, Sept. 1937 (A & B). Mt. Shungol: Hartley 12475, Dec. 1963 (L).

NEW GUINEA (PAPUA):—Northern Distr.: Alola: Carr 13644, Dec.
1935 (holotype in K; istoypes in A, L & SING). Isuarava: Carr 15573 &
15775, Feb. 1936 (A, BM, C, K, L, NY & SING). Lala River: Carr
13969, Dec. 1935 (A, BM, C, K, L & SING).—Milne Bay Distr.: Mt.
Dayman: Brass 22881, June 1953 (A, L & LAE).

(16) Dendrocnide ternatensis (Mig.) Chew in op. cit. 207 (1965).
— Laportea ternatensis Mig. Ann. Mus. Bot. Lugd. Bat. 4:
302 (1869). — L. glabra Ridl. in Trans. Linn. Soc. Bot. 9: 150
(1916). — L. mammosisetosa Winkl. in Bot. Jahrb. 57: 507
(1922). — L. crenulata Wedd. var. nitida Wink]. in op. cit.
506 (1922). — D. nitida (Winkl.) Chew, op. cit. 204 (1965).

Characteristics: (a) plant glabrous except for the long irritant
hairs often with papillate bases, (b) lamina oblong to broad
ovate, often very light puberulous on the mid-rib, otherwise quite
glabrous; margin distantly coarsely serrate at the apical half,
often sinuate, (c) male flowers sessile, 4-tepalled, very light pube-
rulous, with short irritant hairs, (d) female flowers long pedi-
cellate; perianth lightly 4-toothed, dense with irritant hairs and
punctiform cystoliths; pedicel curved, dilated progressively
towards apex, bearing irritant hairs on the dorsal spine, (e)
achenes bloated, with the loose, lightly 4-toothed perianth at the
base.

Distribution: Moluccas, Tanimbar Islands and New Guinea.

48 Gardens’ Bulletin, Singapore — X XV (1969)

Fig. 18: D. ternatensis: A = twig; B = female flower; C = achene.

49

W—L. Chew — Monograph of Dendrocnide

Fig. 19: D. ternatensis: 2 leaf-forms.

50 Gardens’ Bulletin, Singapore — X XV (1969)

Dioecious trees to ca. 10 metres high. Twigs glabrous, without
irritant hairs, usually heavily covered by punctiform cystoliths,
especially at the nodes. Lamina (18—) 20-30 (-42) cm. long, (7-)
10-20 (—32) cm. broad, broadly ovate or oblong, very coriaceous,
glabrous, occasionally light puberulous on the mid-rib, without
irritant hairs; base shallowly cordate to truncate; apex acuminate,
acumen to 2 cm. long; margin sinuate to coarsely and irregularly
serrate at the apex half, the teeth usually becoming longer toward
the aptex; lateral veins (9—) 12-16 pairs, closely set, the basal
pair usually further away from the other veins; intercostals very
numerous, not very straight. Petioles (4—) 8-15 cm. long, glabrous,
densely covered with punctiform cystoliths, irritant hairs absent.
Stipules 2-3 cm. long, puberulous, almost devoid of irritant hairs.
Inflorescences unisexual, narrowly to very broadly branched
racemes, 5—6 flowers in each fascicle, solitary or paired, up to
15 cm. long, 12 cm. broad; peduncular bracts to 1 mm. long,
0.5 mm. broad, elliptic; peduncles puberulous to subglabrous,
well armed with short irritant hairs. Male flowers sessile, 1.5
mm. long, 1.25 mm. broad, well covered with short irritant hairs
and punctiform cystoliths, 4-tepalled; stamens 4, with slightly
reflexed filaments; pistillode ca. 0.75 mm. long, obconical to
obpyramidal; interfloral bracts ca. 0.3 mm. long, narrow elliptic,
1 to each flower; flower at most subsessile at anthesis. Female
flowers pedicellate; perianth ca. 1 mm. long and broad, 4-toothed,
well covered with irritant hairs and punctiform cystoliths; teeth
of perianth equal-sized; ovary ca. 1 mm. long and broad; stigma
ligulate, to 1.5 mm. long, very slightly curled; interfloral bracts
minute often absent; pedicel to 2 mm. long, curved, the dorsal
spine with large irritant hairs on papillate bases. Achenes ca.
3 mm. long, 2 mm. broad, often to 3.5 mm. long, 3 mm. broad,
obovoid or ovoid, bloated, particularly at the base, with warted
pericarp, the shallowly 4-toothed perianth loosely covering the
base; stigma reflexed; pedicel at fruit to 3 mm. long, dilated
progressively to the apex, strongly arched, the dorsal spine with
large irritant hairs on papillate bases.

A tree species of primary as well as secondary forests, it has
been recorded (Saunders 323) to have attained a height of 25
metres with buttresses up to 2 ft. x 2 ft. Occurring mainly below
800 metres altitude, it sometimes forms ‘‘an important constituent
of seral shrubberies on muddy river banks’? (L. J. Brass 7434).

Winkler (1922) kept Ridley’s glabra and his own mammosise-
tosa aS separate speices closely allied to Miquel’s ternatensis.
The types of these have been examined by me in great detail
and have been found to belong to one species which varies
considerably not only in leaf form but also in the presence or
absence of the papillate bases of irritant hairs which are supposed
to be a distinctive feature of mammosisetosa. D. nitida, based
on Winkler’s variety under L. crenulata, has now been found
after further investigation to belong to this speices. I have
examined the holotype, deposited at Wrocklaw, and am convinced
that my present treatment of it is correct.

W —L. Chew — Monograph of Dendrocnide 51

This together with D. torricellensis and D. corallodesme form
a group of closely related species charatcerised by the shallowly
4-toothed perianth of the female flowers becoming a small but
loose cupule at the base of the achene.

MOLUCCAS:—Geram: Loa Loa Mt.: Eyma 3218, Sept. 1938 (BO,

BRI, L, PNH & SING).—Ternate: Teysmann HB. 5185 (type of L.
ternatensis: BO & U).—Tidore: Vriese & Teysmann s.n. (L).

TANIMBAR ISLANDS:—Jamdena: Borssum-Waalkes 3240, April
1956 (K & L).

NEW GUINEA (WEST):—Alkmaar: Versteeg 1547, July 1907 (BO).—
Hollandia: Bier River: Iwanggin BW. 9077, June 1959 (L).—Nassau
Range Distr.: Mt. Carstenzs: Kloss s.n., Nov. 1912 (type of L. glabra:
K).—Vogelkop Peninsula: Soron: Beccari herb. 9493 (FI).

NEW GUINEA (NORTH-EAST):—Madang Distr.: Gogol River:
Lauterbach 1153, Dec. 1890 (holotype of L. crenulata var. nitida: WRSL.)
Ramu Valley: Saunders 385, July 1955 (BRI, L & LAE).—Morobe Distr. :
Markham Valley: Brass 32555, Nov. 1959 (A, K, L, LAE, NY & US).

PAPUA:—Western Distr.: Fly River: Brass 7434, Aug. 1936 (A, BO &
L).—Central Distr.: Sogeri area: Forbes 403 (lectotype of L. mammosise-
tosa: L). Anga River: Brass 5490, Sept.-Nov. 1933 (A, BO, BRI, NY &
US).—Milne Bay Distr.: Goodenaugh Island: Brass 24382, Sept. 1953 (A).

(17) Dendrocnide torricellensis (Laut.) Chew in op. cit. 207 (1965).
— Laportea torricellensis Laut. in K. Sch. & Laut. Nachtr.
Fl. Schutzgeb. 251 (1905).

Characteristics: Closely related to D. ternatensis from which
it distinguishes itself clearly in (a) lamina light villous on the
lowerside, (b) male flowers dense villous to woolly.

Distribution: New Guinea (endemic! ). .

Dioecious trees to 15 metres high. Twigs glabrous, without
irritant hairs; petiolar scars prominent. Lamina 17-30 cm. long;
8-13 cm. broad, elliptic to ovate, coriaceous, upperside with few
hairs on the mid-rib, otherwise glabrous and without irritant
hairs, lowerside pubescent to sparsely so, with few short irritant
hairs; cystoliths punctiform, dense on both surfaces; base cuneate
to rounded; apex acuminate, acumen to 5 mm. long; margin very
imperceptibly crenate, toothed to almost smooth; lateral veins
16 pairs, closely set, parallel, the basal pair slightly shorter but
otherwise as prominent as the others; intercostals very numerous,
closely set, straight. Petioles to 5 cm. long, puberulous to almost
glabrous, with very few short irritant hairs, dense with punctiform
cystoliths. Stipules ca. 1.5 cm. long, pubescent, with few irritant
hairs. Inflorescences unisexual, branched racemes, flowers loosely
arranged in short rows, solitary, 13 cm. long, 5 cm. broad,
peduncular bracts small; peduncles puberulous to subglabrous.
Male flowers sessile, ca. 1.5 mm. long and broad, dense pubescent
to villous, 4-tepalled; stamens 4, with slightly reflexed filaments:
pistillode ca. 0.7 mm. long; interfloral bracts minute. Female
flowers pedicellate; perianth to 0.75 mm. long, ca. 0.5 mm. broad,
very shallowly 4-toothed, with long irritant hairs on the dorsal
side, well covered with punctiform cystoliths; ovary ovoid, covered
by the perianth; stigma ligulate, ca. 1 mm. long; interfloral bracts
very minute; pedicel ca. | mm. long, with long irritant hairs on

52 Gardens’ Bulletin, Singapore — XXV (1969)

Fig. 20: D. torricellensis: A = twig; B = female flower; C = achene.

W.—L. Chew — Monograph of Dendrocnide 53

the dorsal side. Achenes to 2 mm. long, 1.5 mm. broad, very
slightly asymmetrically ovoid, with smooth pericarp, slightly
bloated, the lower portion covered by the lightly 4-toothed loose
perianth; stigma reflexed; achene reflexed over the progressively
thickened pedicel.

Said to be a species of subcanopy trees in rain forests between
100 and 500 metres in altitude.

Indeed, there is a resemblance between this and D. microstigma
as pointed out by Lauterbach who referred to the latter as
contracta. This resemblance, however, is quite superficial: the
closest relative is D. ternatensis from which it differs in the male
flowers being woolly and the undersides of the lamina pubescent.

NEW GUINEA (NORTH-EAST):—Sepik Distr.: Mt. Torricelli:
Schlechter 14385, April 1902 (holotype in WRSL; isotype in BO).—
Morobe Distr.: Oomsis: Brass 29235, April 1959 (A, L, LAE, NY & US).

(18) Dendrocnide corallodesme (Laut.) Chew in op. cit. 202
(1965). — Laportea corallodesme Laut. in K. Sch. & Laut.
Nachtr. Fl. Schutzgeb. 252 (1905).

Characteristics: Closely related to D. ternatensis from which
it can easily be distinguished by (a) the leaves being smaller,
shortly petiolate, narrowly elliptic, with numerous, short lateral
veins, (b) the male flowers pedicellate and (c) the female inflo-
rescences commonly borne behind the leaves.

Distribution: New Guinea (endemic! ).

Dioecious trees to ca. 6 metres. Twigs glabrous, without irritant
hairs; petiolar scars prominent. Lamina (13-) 16-20 (—26) cm.
long, (3—) 4-7 (-9) cm. broad, narrowly elliptic to elliptic, coria-
ceous, glabrous, without irritant hairs; base cuneate, often very
sharply so; apex acuminate, acumen to 2.5 cm. long; margin
usually smooth, sometimes very broadly and shallowly toothed,
rarely strongly so; lateral veins 13-15, sometimes to 19 pairs,
closely set, the basal pair usually shorter and less prominant
than the others; intercostals anastomosing, rarely straight. Petioles
2-3 (-5) cm. long, glabrous, without irritant hairs, densely covered
with punctiform cystoliths. Stipules usually 1—-1.5 cm. long, rarely
longer, lightly puberulous, soon glabrous, without irritant hairs
Inflorescences unisexual, broadly branched racemes, 5—6 flowers
in each fascicle, usually paired, up to 10 x 10 cm. spread;
peduncular bracts minute; peduncle puberulous, with very short
irritant hairs; rami- and cauliflorous. Male flowers pedicellate:
ca. 1 mm. long and broad, fairly dense with short irritant hairs
and punctiform cystoliths, 4-tepalled; tepals ovate, fairly strongly
keeled; stamens 4, with slightly reflexed filaments; pistillode
obpyramidal, to 0.4 mm. long, 0.3 mm. broad at the apex; inter-
. floral bracts to 0.3 mm. long, triangular, 1-3 to each flower:
pedicel to 1 mm. long at anthesis, often with a collar just below
the perianth. Female flowers pedicellate; perianth 1.5 mm. long,
1 mm. broad, 4-toothed, well armed with irritant hairs and
punctiform cystoliths; teeth of perianth equal-sized; ovary 1.5

54

2mm

imm

2cm

Gardens’ Bulletin, Singapore — X XV (1969)

\ ; i jeraum
del.

Fig. 21: D.corallodesme: A = leaf; B = cauliflorous female inflorescence;

C

male flower; D = achene.

W —L. Chew — Monograph of Dendrocnide 5

WA

mm. long, 0.8 mm. broad, ellipsoid; stigma ligulate, to 3 mm.
long, very slightly curled; interfloral bracts, minute, somewhat
triangular; pedicel to 1.5 mm. long, curved, the dorsal spine with
large irritant hairs sometimes on papillate bases. Achenes usually
3 mm. long, 2.5 mm. broad, often larger, obovoid or ellipsoid,
bloated, particularly at the base; pericarp slightly warted; the
shallowly 4-toothed perianth loosely covering the base; stigma
reflexed; pedicel at fruit to 2 mm. long, dilated progressively to
the apex, curved, the dorsal spine with rather large irritant hairs
sometimes borne on papillate bases.

Said to be a common small tree along rivers in primary forests
below 500 metres in altitude. Vernacular names: Ambu, Kongru,
Kwi, Segre and Wel.

NEW GUINEA (WEST):—Vogelkop Peninsula: Manokwari and
vicinity: Koster BW. 10817, Feb. 1960 (KEP, L, LAE & SING): Koster-
mans 2915, July 1948 (A, BO, K, L & SING).

NEW GUINEA (NORTH-EAST):—Madang Distr.: Ramu _ River:
Hoogland 5042, July 1955 (A, L & LAE); Schlechter 13840, Jan. 1902
(type: WRSL).

PAPUA:—Milne Bay Distr.: Kwagira River: Peria Creek: Brass
24200, Aug. 1953 (A & LAE).

(19) Dendrocnide moroides (Wedd.) Chew in op. cit. 204 (1965).
— Laportea peltata Gaud. ex Decne. in Nouv. Ann. Mus.
Hist. Nat. Paris, ser. 3. 3: 490 (1834) et Herb. Timor. 162
(1835), sens. str. — L. moroides Wedd. in Arch. Mus. Hist.
Nat. Paris 9: 142 (1856) et in DC. Prodr. 16 (1): 88 (1869).
— Urticastrum moroides (Wedd.) O.K. op. cit. (1891).

Characteristics: (a) shrubs or trees densely armed with irritant
hairs, (6) lamina broadly ovate, base cordate-peltate, margin
sharply toothed, (c) inflorescences bisexual, flowers pedicellate,
10 to 20 female flowers around each male, (d) achenes warty,
bloated, almost covered by densely hairy lateral tepals and borne
on greatly swollen pedicels.

Distribution: Southern Moluccas and North-eastern Australia.

Monoecious shrubs or trees capable of reaching 10 metres high.
Twigs pubescent, dense with irritant hairs; petiolar scars rather
prominent. Lamina (10—) 14-20 (—38) cm. long, (9—) 12-18 (—30)
cm. broad, broadly to very broadly ovate, coriaceous to charta-
ceous, both surfaces very dense with irritant hairs and punctiform
cystoliths, young ones dense canescent; base cordate, peltate;
apex acuminate, acumen 1-2 cm. long; margin dentate, often
coarsely; lateral veins 6-8 pairs, prominent on the lower surface,
the basal pair most prominent, reaching to more than half the
length of lamina; intercostals very numerous, straight. Pefioles
(8—-) 12-18 (—22) cm. long, dense to very dense with irritant hairs;
cystoliths not visible. Stipules ca. 1 cm. long, pubescent, armed

56 Gardens’ Bulletin, Singapore — X XV (1969}

with minute irritant hairs, caducous. /nflorescences bisexual, loosely
branched racemes, with flowers in fascicles; the male flowers
sparely intermixed with the females; ca. 15 cm. long, 8 cm. broad,
axillary. usually paired; peduncular bracts elliptic, to 1 mm. long:
peduncles pubescent and very dense with long irritant hairs. Male
flowers pedicellate; perianth ca.1.5 mm. long and broad, puberulous,
with very few irritant hairs, 4-tepalled; stamens 4, filaments
reflexed; pistillode ca. 0.75 mm. long, half as broad; interfloral
bracts minute, often absent; pedicel to 1.5 mm. long. Female
flowers pedicellate, ca. 10 flowers at each fascicle with one male
flower in the centre; perianth ca. 0.75 mm. long, 0.5 mm. broad,
4— lobed, dense with irritant hairs and punctiform cystoliths,
one tepal much smaller than the other three; ovary ca. 0.5 mm.
long, half as broad; stigma ligulate, to 2 mm. long; interfloral
bracts small, one to each flower; pedicel short. Achenes to 2 mm.
long, 1.5 mm. broad; ovoid to triangular, bloated, with warted
pericarp, almost entirely covered by the lateral tepals, reflexed;
pedicel elongated and very swollen at fruit.

This species, the most viciously stinging in the genus, is unique
in having bisexual inflorescences in which the few male flowers.
are dispersed in the inflorescence surrounded by numerous females.
If the other characteristics, like the warty achene enclosed by
lateral tepals and borne on swollen pedicels, are taken into account,
it cannot possibly be confused with any other in the genus. The
closest relative of this is perhaps D. cordata.

This was actually first described under the name Laportea
peltata by Decaisne (1834) who based it on a collection of
Leschenault from Timor. Unfortunately Weddell (1856) had this
confused with the Javanese peltate speices which Blume had
described as Urtica peltata. This was followed by all subsequent
authors with the result that D. moroides never got properly
identified with the Timor plant. Happily, the present clearing-up
does not involve the loss of the use of the epithet moroides
because the other epithet peltata has already been used by Blume
for the Javanese species just mentioned (see notes under D.
peltata).

MOLUCCAS:—Timor: sine loc.: Walsh 84 (BO) & Leschenault s.n.

(holotype of L. peltata Decne.: P)—Tanimbar Isls.: Jamdena: Inge:
Borssum Waalkes 3366, April 1956 (L).

QUEENSLAND (AUSTRALIA) :—Cook Distr.: near Atherton: Volck
& Dansie 1471, Jan. 1958 (BRI). Cairns: Helms 1171, June 1922 (C).
Wongabel: Smith 10175, Sept. 1957 (BRI).—Endeavour River: Cunning-
ham s.n. (holotype of L. morcides Wedd.: G).—Mowbray River: Brass
1981, Jan. 1932 (A, B & BRI).—North Kennedy Distr.: Rockingham Bay:
Dallachy s.n. (K & MEL).

NEW SOUTH WALES (AUSTRALIA) :—Drake: Crawford 16 (MEL).
-—Nimbiro: near Lismore: Cheel NSW. 77928, Sept. 1925 (NSW).—Tweed:
River: Bauerlen NSW. 77926 (NSW).

W.—L. Chew — Monograph of Dendrocnide x

IRA! DEL IKE

Fig. 22: D. moroides: A = twig: B = female flower; C = achene.

58 Gardens’ Bulletin, Singapore — X XV (1969)

(20) Dendrocnide cordata (Warb. ex Winkl.) Chew in op. Cit.
202 (1965). — Laportea cordata Warb. ex Winkl. in Bot. Jahrb.
57: 503 (1922). — L. cordatifolia L. S. Smith in Proc. Roy.
Soc. Queens]. 70 (5): 31 (1959). — L. gaudichaudiana auct non
Wedd.: K. Sch. & Hollr. Fl. Kais. Wilh. Land. 38 (1889).

Characteirstics: Closely related to D. moroides from which it
differs in (a) plant altogether less irritant, (6) lamina cordate not
peltate, (c) inflorescences unisexual, rarely bisexual as well.

Distribution: Timor, Tanimbar Islands, New Guinea, New
Britain (Bismarck Archipelago) and Queensland (Australia).

Monoecious shrubs or trees to 10 metres high. Twigs puberu-
lous, dense with irritant hairs; petiolar scars prominent. Lamina
(10-) 18-25 (—35) cm. long, (10—) 16-22 (—30) cm. broad, broadly
to very broadly ovate, coriaceous, both surfaces well armed with
short irritant hairs and very dense with punctiform cystoliths,
young ones pubescent to puberulous: base cordate, not peltate;
apex acuminate, acumen 0.5 to | cm. long; margin dentate to very
broad crenate; lateral veins 8-10 pairs, the 2 lower ones arising
directly from the base ,the second pair most prominent and
reaching to half the length of lamina; intercostals numerous,
straight, prominent. Petioles (10—) 18-22 (—26) cm. long, generally
puberulous, irritant hairs present; cystoliths not dense. Stipules
1.5 to 2 cm. long, pubescent, sparsely armed with minute irritant
hairs, caducous. Inflorescences unisexual, rarely bisexual as well,
loosely branched racemes, up to 20 cm. long and 10 cm. broad,
axillary, usually paired; peduncular bracts small; peduncles
pubescent and well covered with irritant hairs. Male flowers
shortly pedicellate, ca. 1-1.5 mm. long and broad, puberulous,
with few irritant hairs, 4-tepalled; tepals ovate; stamens 4, fila-
ments reflexed; pistillode ca. 0.5 mm. long, half as broad; inter-
floral bracts minute; pedicel ca. | mm. long or less. Female
flowers very shortly pedicellate, in fascicles, very rarely with a
male flower in each group; perianth to 1 mm. long, 0.5 mm.
broad, 4-lobed, fairly dense with irritant hairs and punctiform
cystoliths: 1 tepal slightly smaller than the others; ovary 0.75 to
1 mm. long, half as broad; stigma ligulate, ca. 1-2 mm. long;
interfloral bracts very minute. Achenes 2 mm. long, 1.5 mm.
broad, almost triangular, slightly bloated, with warty pericarp,
almost entirely covered by the lateral tepals, reflexed; pedicel
elongated and swollen at fruit.

It is very remarkable that this species should have female
flowers and achenes almost identical with those of D. moroides
when these two species differ greatly in the sexuality of the inflores-
cence as well as in the type of leaf base.

L. S. Smith was quite right in pointing out the closeness.
between his cordatifolia and this species. I have examined the types
of both and have found them to be ocnspecific. The differences
that Smith pointed out are not sufficient from a monographic

W—L. Chew — Monograph of Dendrocnide 59

Fig. 23: D. cordata: A = twig; B = male inflorescene.

60 Gardens’ Bulletin, Singapore — X XV (1969)

point of ivew to warrant their separation. They are probably the
effect of altitude as cordatifolia occurs at 1,200 metres while cordata
in the lowlands.

Vernacular names: Gabu, Salat & Sisik (Papuan); Gympie
(Australia); Leswatan Alas (Timor).

TIMOR:—Kapan: Bloembergen bb 27110, March 1939 (BO).
TANIMBAR ISLANDS:—sine loc.: Buwalda 4162 & 4292, March 1938
(L).

NEW GUINEA (NORTH-EAST):—Sepik Distr.: Sepik River: Con-
stantinhafen: Hollrung 513 (type of L. cordata: BO, K & MEL).—-
Morobe Distr.: Oomsis Creek: Hartley 10032, March 1962 (L & LAE).

NEW GUINEA (PAPUA):—Central Div.: Kanosia: Carr 11207, Feb.
1935 (A, BM, C, K, L & SING).

BISMARCK ARCHIPELAGO:—New Britain: West Nakai: Cape
Hoskin: Floyd NGF. 3498, July 1954 (LAE).

AUSTRALIA (QUEENSLAND) :—Cook Distr.: near Atherton: Smith
10132, Sept. 1957 (type of L. cordatifolia: BRI); Volck & Dansie 1470,
March 1958 (BRI).

(21) Dendrocnide photinophylla (Kunth) Chew in op. cit. 205
(1965). — Fleurya photinophylla Kunth in Ann. Sc. Nat. ser.
3. 7: 183 (1847). — Urera leichardiana Wedd. in op. cit. ser.
4. 1: 178 (1854). — Laportea photinophylla (Kunth) Wedd.
in Arch. Mus. Hist. Nat. Paris 9: 138 (1856). — Urticastrum
photinophyllum (Kunth) O.K. op. cit. (1891).

Characteristics: (a) plant entirely glabrous, (56) lamina shiny,
elliptic to ovate, glabrous with slightly sinuate margin, (c) lateral
veins 8 pairs or less, (d) irritant hairs only on flowers and pedun-
cles. (e) pedicels elongating greatly at maturity, the female ones
swollen and curved, (f) achenes flat, half covered by persistent
perianth.

Distribution: Australia (endemic! ).

Dioecious trees to 20 metres high. Twigs glabrous, young ones
with few irritant hairs; petiolar scars prominent. Lamina (6-)
8-12 (-18) cm. long, (3—) 4-6 (-10) cm. broad, ovate or elliptic,
coriaceous, entirely glabrous, devoid of irritant hairs, dense with
punctiform cystoliths on both surfaces; base cuneate to rounded;
apex bluntly acuminate, acumen 0.5—1 cm. long; margin entire
to sinuate, bluntly dentate in the juvenile ones; lateral veins 6-8
pairs, prominent, the basal pair straight, more prominent, reaching
to more than half the length of the lamina; intercostals rather
straight, faint. Petioles 3—7 cm. long, without cystoliths, glabrous,
devoid of irritant hairs. Stipules to 1 cm. long, coriaceous, cadu-
cous. puberulous when young, devoid of irritant hairs. /nflores-
cences unisexual, branched racemes, ca. 6 cm. long and broad,
axillary, solitary, with flowers in loose fascicles; peduncular bracts
minute; peduncle puberulous to almost glabrous, with dispersed
short irritant hairs. Male flowers shortly pedicellate, ca. 1.5 mm.
long and broad, subglabrous, 4-tepalled; stamens 4, with reflexed
filaments; pistillode ca. 0.5 mm. long; interfloral bracts minute:
pedicel elongating greatly at anthesis, then slightly swollen and

W —L. Chew — Monograph of Dendrocnide 61

Fig. 24: D. photinophylla: A = twig; B = juvenile leaf; C = female
flower; D = achene.

62 Gardens’ Bulletin, Singapore — X XV (1969)

fleshy. Female flowers pedicellate, 6-8 flowers in each fascicle;
perianth ca. 1 mm. long, 4-lobed, very light puberulous with few
short irritant hairs, dense with punctiform cystoliths, 2 tepals
half the length of the other 2; ovary 1 mm. long, 0.5 mm. broad;
stigma ligulate, ca. 1.5 mm. long, curled; interfloral bracts small,
irregular in number; pedicel to 1 mm. long. Achenes to 2 mm.
long and broad, somewhat sigmoid, flattened, with smooth peri-
carp, half covered by persistent perianth, reflexed over elongated
and swollen pedicels 3-4 mm. in length.

Common in dense “rainforest”? on deep sand in the Rockhamp-
ton and Moreton Districts of Queensland where it is known as
the Mulberry Stinging Tree and Nettle Tree.

This was described by Kunth from materials cultivated at
Goettingen in Germany, the type subsequently deposited at the
Berlin Herbarium. As this Herbarium was destroyed in the last
World War, the holotype can be considered as having been
destreyed with it. White’s collection no. 5600 from Nudgee is
therefore here made a Neotype. D. vitiensis of Samoa and Fiji
and D. morobensis of New Guinea seem close to this (see notes
there).

QUEENSLAND :—Brisbane Distr.: Nudgee: White 5600, March 1929
(neotype in A; duplicates in BRI & NY).—Darling Downs: Kilarney:
Boorman 31, Feb. 1905 (NSW)—Moreton Distr.: The Creek: Leichard 26
(holotype of U. leichardiana: P). Mt. Tamborine: White 12726, May 1945

(BO & K).—Rockhampton Distr.: Rockhampton: Dietrich 605, 1438,
1889 & 2192 (MEL); Thozet 408 (MEL).

NEW SOUTH WALES:—Brisbane River: sine loc.: Cunningham s.n.
(CGE & K).—Hastings Distr.: Walcha: Crawford 16, May 1887 (MEL).—
North Coast Distr.: Richmond River: Fawcett 1879 (MEL).—Tooloon
Range: sine loc.: Vickery s.n., April 1953 (NSW herb. 77931).

(22) Dendrocnide vitiensis (Seem.) Chew in op. cit. 207 (1965).
— Laportea vitiensis Seem. Fl. Vit. 239. tab. 60 (1868).

Characteristics: Almost similar to D. photinophylla except for
the following: (a) lamina larger and ovate-pentagonal in shape,
(b) lamina with more than 8 pairs of lateral veins, (c) peduncles and
flowers fairly densely covered with irritant hairs.

Distribution: Samoa, Fiji & New Caledonia.

Shrubby trees to 10 metres high. Twigs very light puberulous at
apex only, other parts quite glabrous, without irritant hairs; petiolar
and stipular scars prominent. Lamina (7—) 12-16 (—20) cm. long,
(3—) 6-8 (-10) cm. broad, somewhat ovate-pentagonal, coriaceous,
entirely glabrous, devoid of irritant hairs; cystoliths punctiform,
very dense on the upper surface, elongated on the lower surface:
base rounded to cordate, often somewhat truncate; apex acute to
shortly acuminate; margins very often sinuate; lateral veins 8-10
pairs, prominent, basal pair slightly more prominent, almost
straight; intercostals prominent, not numerous. Petioles 2-4 cm.
long, with elongated cystoliths, entirely glabrous, devoid of irritant
hairs. Stipules early caducous, ca. 0.5 cm. long, very lightly
puberulous, devoid of irritant hairs. Inflorescences unisexual,
branched racemes, with flowers in loose fascicles, solitary, usually

ed on ieee eS

W —L. Chew — Monograph of Dendrocnide

63

Fig. 25: D. vitiensis: A = twig; B = male flower; C = achenes.

64 Gardens’ Bulletin, Singapore — X XV (1969)

up to 8 cm. long and broad, rarely more; peduncular bracts
unpaired, small; peduncles thin, densely covered with short irritant
hairs. Male flowers sessile to subsessile, pedical elongating to as
long as perianth at anthesis; perianth to 2 mm. long and broad,
closely puberulous, often somewhat glabrous at anthesis, with few
irritant hairs, 4-tepalled; tepals narrow ovate; stamens 4, with
reflexed filaments; anthers large; pistillode thin, ca. 0.75 mm. long;
inter-floral bracts minute. Female flowers pedicellate; perianth to
0.75 mm. long and broad, 4-lobed, densely covered with irritant
hairs, tepals 4, 2 longer and 2 slightly smaller; ovary ca. 1 mm.
long, 0.75 mm. broad, asymmetrically ovoid; stigma ligulate, to
2 mm. long; inter-floral bracts small, irregular in number; pedicel
elongating greatly at fruit. Achenes ca. 3 mm. long, 2 mm. broad,
asymmetrically ovoid pyriform, with slightly warty pericarp,
flattened, almost half covered by the narrow persistent perianth;
stigma greatly displaced to one side of achene; achene reflexed
over swollen pedicel.

Mainly a shade tree along rivers in primary forests, occurring
below 1,000 metres in altitude. Vernacular names are: Salato,
sometimes Mbolavatu.

This species is very closely related to the Australian species
D. photinophylla. In fact, the resemblance is so great that Weddell
had them lumped into one species in 1869. I have compared these
two very carefully and have found them to be quite distinct on the
characteristics enumerated in the diagnosis above.

SAMOA:—Upolu: Tanumalala: McKee 3014, Aug. 1955 (K & L).—
sine loc.: Reinecke 91, Oct. 1894 (BO, K, US, WRSL & Z).

FIJI:—Koro: Main Range: Smith 985, Jan. 1934 (BO, GH, K, NY, S,
UC & US).—Levuka: Ovalau: Gillespie 4425, Jan. 1927 (K).—Vanua
Levu: Thakaundrove: Smith 359, Nov. 1933 (BO, GH. K, NY, S, UC &
US).—Viti Levu: Tholo North: Degener 14797, March 1941 (A, K,
MICH, MO, NY, S & UC).—sine loc.: Seemann 427 (type of L. vitiensis:
GH, K & MEL).

NEW CALEDONIA:—Lifa Island: Thiebault (collection not seen).

(23) Dendrocnide excelsa (Wedd.) Chew in op. cit. 203 (1965). —
Urera excelsa Wedd. in Ann. Sc. Nat. ser. 4. 1: 178 (1854). —
U. rotundifolia Wedd. in op. cit. 177 (1854). — Laportea gigas
Wedd. in Arch. Mus. Hist. Nat. Paris 9: 129. tab. 3 (1856) et in
DC. Prodr. 16 (1): 82 (1869). — Urticastrum gigas (Wedd.) O.K.
op. cit. (1891).

Characteristics: (a) large tree to 35 metres high, frequently
buttressed at the base, (b) lamina ovate, base cordate, the two
lobes often overlapping, the underside dense woolly, the upperside
subglabrous but with irritant hairs, both surfaces dense with
punctiform cystoliths, (c) male flowers rather long pedicellate at
anthesis, 4-tepalled, with irritant hairs, (d) female flowers shortly
pedicellate, 4-tepalled, perianth densely covered with long white
hairs, (e) achenes flat, strongly tuberculate, on long fleshy pedicels
armed with few large irritant hairs; the lateral tapals narrow, half
the length of the achene.

Distribution: Australia (endemic! ).

W —L. Chew — Monograph of Dendrocnide 65

Fass

y

Fig. 26: D. excelsa: A = twig (after Weddell); B = female flower; C =
achene,

66 Gardens’ Bulletin, Singapore — X XV (1969,

Dioecious tree to 35 m. high, the base of the trunk usually
highly buttressed. Twigs usually with irritant hairs, with large
and very prominent petiolar scars. Lamina (10—) 13-20 (—25) cm.
long, (S—) 10-12 (-18) cm. broad, those of juvenile plants up to
45 cm. long, 30 cm. broad, ovate, coriaceous, the upperside sparse
with short irritant hairs, the lowerside dense woolly, both surfaces
dense with punctiform cystoliths; base cordate, often very strongly
and with the two lobes overlapping; apex shortly and very bluntly
acuminate, acumen to ca. 7 mm. long; margin very shallowly and
broadly dentate, crenate, gnawed to somewhat entire; lateral veins.
6-8 pairs, the basal pair more prominent than the rest, reaching to
half of the length of the lamina; intercostals somewhat straight,
quite prominent. Petioles 5-10 cm. long, dense pubescent to woolly,
well armed with long thick irritant hairs. Stipules 1-2 cm. long,
somewhat coriaceous, pubescent, caducous. /nflorescences uni-
sexual, branched racemes, flowers in close fascicles, up to 12 cm.
long, 6 cm. broad, axillary, paired, rarely solitary, peduncular
bracts at the lowest axils to 3 mm. long, triangular, peduncles
pubescent, armed with short irritant hairs. Male flowers, subsessile
to shortly pedicellate; perianth ca. 1.5 mm. long and broad, well
covered with short irritant hairs, 4-tepalled; stamens 4, on reflexed
filaments; pistillode ca. 0.5 mm. long, obpyramidal; interfloral
bracts minute; pedical elongating to 1.5 mm. at anthesis. Female
flowers pedicellate; 6-8 flowers in each fascicle; perianth ca.
0.8 mm. long, 0.6 mm. broad, 4-lobed, dense with white hairs as
well as irritant ones; ovary ca. 0.8 mm. long, 0.5 mm. broad,
ovoid; stigma ligulate, ca. 1 mm. long; interfloral bracts triangular,
ca. 0.4 mm. long, 2 or 3 to each flower; pedicel to 0.5 mm. long,
with few stiff irritant hairs. Achenes 2 mm. long, 1.5 mm. broad,
ovoid, flat, with strongly tuberculate pericarp, the lateral tepals
narrow, at most half the length of the achene; pedicel to 5 mm.
long, very fleshy, bearing 1 or 2 large irritant hairs 1-1.5 mm.
long, curved.

Judging on floral characters, this species of giant trees seems:
closest to D. photinophylla. The female flowers of these differ
only in one character viz. the presence of very long rigid irritant
hairs on the pedicels in D. excelsa. They however, can easily be
distinguished from each other by the leaves which are glabrous in
D. photinophylla but dense woolly on the underside in this.
species.

The specific epithet gigas is nomenclaturally superfluous. When
published in 1856 by Weddell, it included the names excelsa and
rotundifolia either of which, being quite legitimate, should have
been used by the author.

My interpretation of this species is based entirely on that of
Weddell. Collections of Cunningham and Verreaux authenticated.
by this author under the name gigas have been examined by me.

W.—L. Chew — Monograph of Dendrocnide 67

QUEENSLAND :—Darling Downs Distr.: Bunya Mts.: White 9145,
Aug. 1933 (A, BRI & NY).—Moreton Distr.: Mt. Glorious: Smith 10540,
April 1959 (BRI).—Mt. Tamborine: White 12728, May 1945 (BO).—Wide
Bay Distr.: Imbil: Petrie BRI 57800, Sept. 1922 (BRI).

NEW SOUTH WALES:—Beaury State Forest: White 12781, May 1945
(A, BO & K).—Liverpool Ranges: Caird’s Gap: Johnson & Constable
NSW. 32026, Oct. 1954 (NSW).—Cambewara Range: Rodway 1803, July
1935 (A &K).—Five Islands Country: Cunningham s.n. (type of L. gigas
Wedd.: K).—South Coast: Kiama: Pullen 4065, April 1964 (L & NSW).—
Singeton, N.-E.: Story 7174, March 1960 (K & NSW).—Sydney Distr. :
Bulli Pass: Einar & Du Rietz 4073, July 1927 (S)——New Holland:
Verreaux 559, May 1845 (P).

(24) Dendrocnide morobensis Chew, spec. nov.

Characteres: D. photinophyllae primo adspectu simillima, sed
pilis stimulantibus basi papillatis, femineis pedicellis haud suc-
culentibus et tepalis lateralibus angustioribus brevioribus differt.

Distribution: New Guinea (endemic! ).

Dioecious trees to 25 metres high, Twigs glabrous, without
irritant hairs, with very large petiolar scars. Lamina 6-17 cm.
long, 3-10 cm. broad, ovate, often pentagonal, coriaceous, very
sparse with mounted irritant hairs, very dense with punctiform
cystoliths; base round, truncate to very shallowly cordate; apex
acuminate, acumen to 1 cm; margin crenate to sinuate; lateral
veins ca. 6-7 pairs, the basal pair reaching to half the length of
the lamina; intercostals numerous. Petioles 2-3 cm. long, entirely
glabrous. Stipules less than 1 cm. long, coriaceous, entirely
glabrous, caducous. /nflorescences branched racemes, ca. 7 cm.
long, 5 cm. broad, axillary, solitary; peduncular bracts minute;
peduncle subglabrous to pubescent, with mounted irritant hairs.
Male flowers pedicellate, perianth 1.3 mm. long and_ broad,
glabrous, 4-tepalled; stamens 4, with strongly reflexed filaments;
pistillode ca. 0.5 mm. long, narrowly obconical; interfloral bracts
minute; pedicel elongating to 1.5 mm. long at anthesis, not fleshy.
Female flowers pedicellate, 5-8 flowers in each fascicle; perianth
ca. 0.6 mm. long, 0.4 mm. broad, 4-tepalled, lightly covered with
irritant hairs, the lateral tepals narrow, about twice as long as
the medial ones: ovary ca. 0.8 mm. long, 0.4 mm. broad; stigma
ligulate, ca. 1.5 mm. long, curved; interfloral bracts small,
irregular in number; pedicel to 0.6 mm. long. Achenes 2 mm. long,
1.5 mm. broad, somewhat ellipsoid, with minutely warted pericarp,
very shortly stipitate, only the very short stipe covered by the
persistent perianth; pedicel elongating to ca. 1.5 mm., not fleshy.

Known so far from the highlands of the Territory of New
Guinea (above 2,000 metres in altitude), this species appears very
similar to the Australian D. photinophylla with which it has
numerous characteristics in common such as the lamina shape and
venation, small glabrous stipules and the general glabrousness in

both species. Upon a closer examination however, these two can
easily be distinguished as the New Guinean species is armed with
irritant hairs borne on papillate bases and has female flowers
borne on simple non-fleshy pedicels. Besides, the lateral tepals
_ are very narrow compared to those of D. photinophylla.

68 Gardens’ Bulletin, Singapore — X XV (1969)

4cm

Fig. 27: D. morobensis: A = twig; B = male flower; C = female flowers;
D .=) achene,

a

— oo

W —L. Chew — Monograph of Dendrocnide 69

Vernacular names are: Nungindankil and Nunts. The specific
epithet is derived from the name of the district where the holotype
was collected, viz. Morobe.

NEW GUINEA (NORTH-EAST):—Morobe Distr.: Mt. Sarawaket:
Clemens 6282, May 1937 (A & B). Above Wau: Womersley & van Royen
NGF. 5896, June 1954 (holotype in L; isotypes in A, BRI & LAE).—
Eastern Highlands: Fatima River: Womersley NGF. 24558, May 1965
(SING). Upper Omahaiga River Valley: Pullen 526 & 569, Sept. 1957
(L & LAE).

(25) Dendrocnide nervosa (Winkl.) Chew, comb. & stat. nov. —
Laportea crenulata Wedd. var. nervosa Winkl. in Bot. Jahrb.
57: 507 (1922), basionym.

Characteristics: (a) lamina obovate, glabrous, without irritant
hairs, the upperside very dense with punctiform cystoliths, the
lowerside less so, (b) male flowers pedicellate, 4-tepalled, (c)
achenes slightly bloated, with the 4-tepalled perianth at the very
base, on winged pedicels with large irritant hairs borne on two
LOWS.

Distribution: New Guinea and Solomon Islands.

Dioecious trees to 12 metres, the trunks often buttressed. Twigs
glabrous, without irritant hairs; petiolar scars very prominent;
cystoliths usually absent. Lamina (8—) 12—20 (—25) cm. long, (3-)
5-8 (-11) cm. broad, obovate, often narrowly, sometimes narrowly
elliptic, coriaceous, glabrous, without irritant hairs; base cuneate,
very rarely rounded, apex acuminate, acumen usually 1.5 cm. long;
margin smooth, sometimes sinuate; lateral veins (6—-) 8-10 (—11)
pairs, fairly distantly set, strongly arching, the basal pair usually
straighter, shorter and less prominent; intercostals few, anastomos-
ing. Petioles 3—5 cm. long, glabrous, without irritant hairs, densely
covered with punctiform and elongated cystoliths. Stipules ca.
1 cm. long, puberulous, devoid of irritant hairs. /nflorescences
unisexual, well branched racemes, with few flowers in each fascicle,
usually paired, up to 10 cm. long and broad, the males usually
much smaller; peduncular bracts triangular, ca. 0.5 mm. long,
pubescent; peduncles puberulous to pubescent, with numerous
very short irritant hairs. Male flowers pedicellate, ca. 2 mm. long
and broad, punctiform cystoliths sparse, 4-tepalled; tepals narrow
ovate, the apex with a broad filmy extension, stamens 4, with very
slightly reflexed filaments; pistillode ca. 1 mm. long and broad at
the apex, obpyramidal; interfloral bracts triangular, ca. 0.2 mm.
long, 2 or 3 to each flower; pedicel to | mm. long at anthesis.
Female flowers pedicellate; perianth to slightly less than | mm.
long and broad, 4-tepalled, well covered with irritant hairs and
cystoliths; tepals somewhat equalsized, rather narrow, the sides
not touching each other; ovary ca. 1 mm. long and 0.8 mm.
broad; stigma ligulate, to 2 mm. long, slightly curled; interfloral
bracts very narrow ovate, ca. 0.5 mm. long, irregular in number;
pedicel ca. 1 mm. long, rather fleshy, usually with two rows of
irritant hairs. Achenes to 3.5 mm. long, 2.5 mm. broad, ovoid.

70 Gardens’ Bulletin, Singapore — X XV (1969)

Fig. 28: D. nervosa: A = twig; B = male flower: C achene.

W —L. Chew — Monograph of Dendrocnide 71

sometimes almost spherical, bloated; pericarp often warted; the
4-tepalled perianth at the base usually reflexed; stigma reflexed;
pedical to 3 mm. long, usually developing lateral wing-like exten-
sions, the edges bearing rather long irritant hairs.

A species of medium sized trees of dense primary as well as
secondary forests and old abandoned gardens. It seems to be able
to tolerate an altitudinal range from sea-level to about the 1300-
metre contour. Brass recorded it as substage tree in the Castanopsis-
oak forest in the Eastern Highlands Districts. Achenes ripen
orange to coral red and most probably distributed by birds.
Vernacular name: Ambu, Kili, Poto, Sarat, Tetasab and Wet (New
Guinea).

Whatever this species has in common with D. sinuata (= L.
crenulata) is entirely superficial. The coral-redness of the mature
achenes with their winged pericels has no parallel in the genus.
Vegetatively this is somewhat similar to some short-leafed
individuals of D. schlechteri with which it is obviously related.

NEW GUINEA (NORTH-EAST):—Madang Distr.: Finnisterre Mts.:
Hoogland 5168, Sept. 1955 (L, LAE & US). Ramu Valley: Schlechter
18377, Oct. 1908 (lectotype in B; duplicates in BR & UC).—Eastern High-
lands: Kassam: Brass 32462, Nov. 1959 (NY, LAE & US).—Morobe
Distr.: Markham Valley: Umi River: Brass 32593, Nov. 1959 (A, K, L,
LAE, NY & US).

PAPUA:—Northern Div.: Tufi: Hoogland 4540 & 4541, Aug. 1954
(L & LAE).

SOLOMON ISLANDS:—Choiseul Island: Whitmore BSIP. 5684, April
1964 (L, LAE & SING).—Guadaloanal Island: Dennis RSS. 2127, July
1965 (SING).

(26) Dendrocnide latifolia (Gaud.) Chew in op. cit. 203 (1965). —
Laportea latifolia Gaud. Bot. Voy. Bonite, tab. 81 (1844). —
L. saipanensis Kaneh. in Bot. Mag. Tokyo, 45: 277 (1931). —
L. kusaiana Kaneh. in op. cit. 46: 449 (1932). — D. saipanensis
(Kaneh.) Chew and D. kusaiana (Kaneh.) Chew in op. cit.
203, 205 (1965).

Characteristics: (a) lamina elliptic generally, entirely glabrous,
with 8-13 pairs evenly spaced lateral veins, (b) petioles dense
pubescent, (c) peduncles with patches of irritant hairs, (d) lateral
tepals of female flowers broad ovate, with irritant hairs, much
larger than the medial ones, (e) achenes flat, smooth, at most
half covered by the perianth, usually on curved, non-fleshy short
pedicels.

Distribution: Pacific Islands (Micronesia and Melanesia).

Dioecious trees to 12 metres high. Twigs puberulous to
pubescent, soon unevenly glabrous to glabrous, with very few
irritant hairs; petiolar scars very large and prominent. Lamina

72 Gardens’ Bulletin, Singapore — X XV (1969)

(13-) 20-26. (—33) cm. long, ca. 5-15 cm. broad, narrowly
elliptic to ovate, sometimes obovate, coriaceous, entirely glabrous,
devoid of irritant hairs, dense with punctiform cystoliths on
both sides; base narrowly to very broadly cuneate, rarely
rounded; apex acuminate, acumen 1-2 cm. long; margin
irregularly sinuate, bluntly dentate in juvenile ones; lateral veins
10-14 pairs, evenly spread, the basal pair usually less prominent.
Petioles (3—) 5-8 (-13) cm. long, usually dense pubescent and
with irritant hairs, soon unevenly glabrous to glabrous. Stipules
1.5—2 cm. long, dense pubescent, caducous. Inflorescences uni-
sexual, branched racemes, ca. 10-15 cm. long and broad, axillary,
solitary, with minute peduncular bracts; peduncle puberulous,
with numerous unevenly distributed irritant hairs; flowers in loose
fascicles. Male flowers sessile to subsessile, ca. 1.5 mm. long and
broad, light pubescent, 4-tepalled; tepals broad ovate, ridged on
the outside, with few irritant hairs; stamens 4, with reflexed
filaments; pistillode 0.5 mm. long; interfloral bracts very small.
Female flowers shortly pedicellate to subsessile, 2 or 3 in each
fascicle; perianth to 1 mm. long, 4-lobed, with irritant hairs; the
lateral tepals ovate, not more than twice the length of the medial
ones; ovary ca. 0.75 mm. long, 0.5 mm. broad; stigma ligulate
to ca. 2 mm. long, slightly curled; interfloral bracts small,
irregular.in number; pedicel to 1 mm. long, with irritant hairs.
Achenes to 2.25 mm. long, 2 mm. broad, asymmetrically obovoid,
flattened, with smooth to tuberculate pericarp, at most half
covered by the lateral tepals; stigma reflexed over achene: pedicel
very short, non-fleshy; extremities of peduncles not fleshy at
maturity.

Being so widely spread over the Pacific, it is natural that it
should have a fairly wide range of variation not only in leaf form
and size but also in the vestiture of the petioles and stipules. The
lamina vary from the very small, narrowly elliptic ones with
absolutely glabrous petioles in Solomon Islands to the large
ovate or even obovate one with dense pubescent petioles found
in the Marianna Islands. That Kanehira should have two species
described from this great variation is therefore not surprising.

Again, in the Solomon Islands, there is another population
which has such obovate laminas and glabrous petioles that they
are almost indistinguishable from D. schlechteri of New Guinea.

Weddell and J. J. Smith considered this species similar to D.
sinuata. I have compared the type of sinuata carefully with
Gaudichaud’s plate of latifolia but cannot find any valid grounds
for this opinion.

This species has been cultivated under the name Laportea
schomburgkii var. vesicolor.

73

W —L. Chew — Monograph of Dendrocnide

Fig. 29: D. latifolia: A & B = twigs: C = female inflorescene, part; D

female perianth; E = female flower (after Gaudichaud).

74 Gardens’ Bulletin, Singapore — X XV (1969)

MARIANNA ISLANDS:—Saipan Island: Hosokawa 7595, July 1934
(US). Kanehira 887, July 1930 (syntype of L. saipanensis: A).—Rota
Island: Kanehira 1754 (K), 1776, July 1932 (NY) & 1783 (US)—Guam
Island: Stone 5096, April 1964 (SING).

CAROLINE ISLANDS:—Ponape Island: Mt. Tortom: Kanehira 1555,
July 1931 (NY).—Kusaie Island: Kanehira 1304, June 1931 (syntype of
L. kusaiana: US).

SOLOMON ISLANDS:—Bougainville Island: Corner NGF. 13561, Oct.
1960 (CGE, L, LAE & SING).—Choiseul Island: Whitmore BSIP. 5409,
March 1964 (L, LAE & SING).—New Georgia Island: Maenu’u BSIP.
6073, June 1965 (SING).—Santa Ysabel Island: Beer BSIP. 6675, April
1966 (SING).—Malaita Island: Lipageto BSIP. 3447 & 3466, Dec. 1963
{L, LAE & SING).—Guadalcanal Island: Kajewski 2526, April 1931 (A,
BM, BO, BRI & L).—San Cristobal Island: Whitmore RSS. 6130, July
1965 (K & SING).

NEW HEBRIDES:—Vanua Lava Island: Kajewski 42], July 1928
(A, BRI, K & NY).—Espiritu Santo Island: Schmid 318 & 319, May 1965
({NOU).—Vate Island: Schmid 320, May 1965 (NOU).

LOYALTY ISLANDS :—Lifu Island: Kepare: Sarasin 814, April 1912
(Z).

NEW CALEDONIA:—Noumea: Franc 1229, Jan 1908 (UC).—Sarra-
mea: Guillaumin 8504, Nov. 1950 (A, NY & US).

Cult. ex Hort. Kew., May 1875 (CGE, E, FI, K, L, MEL, UPS, US &
Z). :

(27) Dendrocnide rechingeri (Wingkl.) Chew in op. cit. 205 (1965 .
Laportea rechingeri Winkl. in Bot. Jahrb. 57: 506 (1922).

Characteristics: (a) lamina elliptic to oblong, apex short
acuminate, base round to cordate, rarely broad cuneate, (b) under-
‘sides of lamina tomentulose, upper-side glabrous except for some
irritant hairs on the mid-rib, (c) lateral veins ca. 13-15 pairs,
(d) female flowers sessile, in fascicles; perianth deeply 4-lobed,
dense with irritant hairs which are sometimes hyaline, (e} achene
flat, lightly tuberculate, sessile, the lateral tepals often greatly
developed.

Distribution: New Guinea, Bismarck Archipelago and Solomon
Islands.

Diocecious trees to 14 metres high. Twigs pubescent at the apex,
soon subglabrous, with very few irritant hairs, dense with
punctiform cystoliths; petiolar scars very prominent. Lamina (20-)
30-38 (-48) cm. long, (10-) 15-25 (—28) cm. broad, elliptic to
oblong, coriaceous, the upperside with few short irritant hairs on
the mid-rib, otherwise glabrous, the lowerside tomentulose with
some short irritant hairs, both surfaces very dense with punctiform
cystoliths; base rounded to shallowly cordate, rarely cuneate; apex
sharply but shortly acuminate, acumen to 0.75 cm. long: margin
entire to very broadly sinuate; lateral veins 13-15 pairs, rarely
more, fairly distantly spaced, the base usually with 2 pairs of
less prominent veins; intercostals straight; very numerous, quite
prominent. Petioles 6-15 (—20) cm. long, well covered with short
irritant hairs, fairly dense with punctiform cystoliths. Stipules
3-5 cm. long, coriaceous, pubescent, with few short irritant hairs.
Inflorescences unisexual, branched racemes, to 15 cm. long and
broad, axillary, paired or solitary, with triangular peduncular
bracts to 5 mm. long: peduncle dense with irritant hairs; flowers

W —-L. Chew — Monograph of Dendrocnide

eet 5 2 !
4, Ss s
7 7 4» y (
¢ Pha ¥ / / /
eS Pe ot ‘ffi {| pe
ee > ve hehe i F vl
al 1 : 3 4
Z “eZ ea / / [ hift seal in %
\ Hf hi peck a ; ‘ /
\ | Age tote /
Fy os re OE | 7 cA,
Mak Cada .
ey nae ds
Nf . ue ce
~ ae ae | |
( j r

jo aim
del,

= achene.

Fig. 30: D. rechingeri: A = twig; B

76 Gardens’ Bulletin, Singapore — X XV (1969)

in somewhat congested fascicles. Male flowers sessile, ca. | mm.
long and broad, well covered with short irritant hairs and
punctiform cystoliths, 4-tepalled; tepals ovate; stamens 4, on
slightly reflexed filaments; pistillode ca. 0.5 mm. long, obpyramidal;
interfloral bracts ca. 0.3 mm. long, triangular, ca. 3 to each flower.
Female flowers sessile, rarely subsessile, in loose fascicles; perianth
ca. 0.6 mm. long, ca. 0.4 mm. broad, 4-lobed, covered with irritant
hairs, often long and hyaline, and cystoliths;: the lateral tepals
much longer than the medial ones; ovary ca. 0.75 mm. long and
0.5 mm. broad; stigma ligulate, to 1.5 mm. long, straight; interfloral
bracts minute. Achenes ca. 2 mm. long, 1.5 mm. broad, ellipsoid,
flattened, with very finely tuberculate pericarp, sessile on non-fleshy
peduncles, with the lateral tepals often developing to as long as the
achene, the medial ones remaining very small; stigma reflexed;
achene very slightly reflexed.

Though I have not seen the type materials, I am able to identify
this species without much difficulty because of the excellent
‘descriptions Winkler supplied. It is found to be somewhat
intermediate between D. latifolia, mainly of the Pacific Islands
-and D. schlechteri of New Guinea in numerous characteristics
such as the shape of peduncular extremities, shape of lamina etc.
Had this species not possessed certain other characteristics peculiar
to itself, I would not have hesitated to have this reduced to
D. latifolia.

This seems to be common at about the 500 metre contour.
Vernacular names: Bandoei (Indonesian New Guinea), Akoako,
Giruppa and Latu (Solomon Islands).

NEW GUINEA (WEST):—Manokwari vicinity: Waren: Kanehira &
Hatusima 14211, April 1940 (A & BO).

PAPUA:—Soron: Beccari herb. 9492 (FI).

BISMARCK ARCHIPELAGO :—Duke of York Island: Rabaul: Bliss
.610, Nov. 1937 (BRI & K).

SOLOMAN ISLANDS:—Bougainville Island: Kugumaru: Kajewski
1943, July 1930 (A, BO & BRI).—Guadalcanal Island: Kavua Stream:
Whitmore & Womersley BSIP. 709, Nov. 1962 (L, LAE & SING).—Santa
Ysabel Island: Maringe Lagoon: Whitmore BSIP. 2288, Oct. 1963 (L &
LAE).

(28) Dendrocnide schlechteri (Winkl.) Chew in op. cit. 205 (1965).
— Laportea schlechteri Winkl. in Bot. Jahrb. 57: 505 (1922).

Characteristics: Very closely related to D. rechingeri from which
it differs in (a) plant completely glabrous and quite without irritant
hairs, (b) flowers sessile, without irritant hairs, but with punctiform
cystoliths, (c) lamina consistently narrow obovate with a strongly
cuneate base and abruptly acuminate apex, (d) achene half or more
covered by the lateral tepals, sitting on swollen peduncular extre-
mities devoid of irritant hairs.

Distribution: New Guinea & Bismarck Archipelago.

Dioecious shrubs or trees to 10 metres. Twigs entirely glabrous
and quite without irritant hairs, dense with punctiform cystoliths,
particularly at the nodes; petiolar scars fairly prominent. Lamina

WL. Chew — Monograph of Dendrocnide 77

Fig. 31: D. schlechteri: A = twig; B = female flowers; C = achene.

78 Gardens’ Bulletin, Singapore — X XV (1969)

(12-) 18-25 (—33) cm. long, (6-) 7-9 (-17) cm. broad, narrowly
obovate, very rarely elliptic-oblong, coriaceous, entirely glabrous,
devoid of irritant hairs, both surfaces dense with punctiform
cystoliths; base cuneate, to very narrowly so; apex bluntly acu-
minate, acumen ca. 1.5 cm. long; margin entire; lateral veins
10-13 (-15) pairs, fairly distantly spaced, the basal pair very short
and less prominent than the others; intercostals numerous, straight, ©
faint. Petioles (4-) 6-12 (-19) cm. long, entirely glabrous, devoid
of irritant hairs, densely covered with punctiform cystoliths.
Stipules 3-4 cm. long, narrow, coriaceous, fairly persistent, entirely
glabrous, without irritant hairs. Inflorescences unisexual, branched
racemes, usually 10 cm. long, 6 cm. broad, sometimes to 15 cm.
long, axillary, paired or solitary; peduncular bracts minute, few;
peduncle glabrous, without irritant hairs, well covered with
punctiform cystoliths; flowers in rather congested fascicles. Male
flowers sessile, ca. 1.5 mm. long and broad, entirely glabrous,
devoid of irritant hairs, with punctiform cystoliths, 4-tepalled;
tepals ovate, smooth; stamens 4, on reflexed filaments; pistillode
ca. 0.5 mm. long and broad at the apex, obpyramidal; interfloral
bracts very minute. Female flowers sessile, 5-7 flowers in each
fascicle; perianth ca. | mm. long, 0.5 mm. broad, 4-lobed, quite
glabrous and without irritant hairs, with punctiform cystoliths; the
lateral tepals slightly longer than the medial ones; ovary ca. 0.5
mm. long, ca. 0.3 mm. broad; stigma ligulate, to 2 mm. long, fairly
straight; interfloral bracts minute, irregular in number. Achenes
ca. 2 mm. long, 1.5 mm. broad, symmetrically ovoid to broad
ellipsoid, slightly flattened, with tuberculate pericarp, ca. half or
more covered by the lateral tepals, the medial tepals remaining
very small; stigma reflexed; achenes slightly reflexed, borne sitting
on swollen peduncular extremities at maturity.

A species of small slender trees, sometimes rather shrubby,
occurring mainly near streams in primary forests up to 1,200
metres in altitude.

This species is interesting. Firstly it is somewhat intermediate
between D. rechingeri and D. longifolia as far as the female
inflorescence is concerned. The peduncular extremities are swollen
but not to the same extent as that found in the latter species. A
cew collections have been encountered in which these receptacles
are so very slightly fleshy as to be almost indistinguishable from
those of D. rechingeri. Nevertheless, this species still possesses a
number of characteristics peculiar to itself so that I have no choice
but to retain it specific status as established by Winkler.

Secondly, this species bears some resemblances to some New
Guinea species of Poikilospermum. This resemblance is especially
marked in the vegetative structures as well as in the branching of
the inflorescence. Still, one can easily fit this into the present
genus despite the absence of irritant hairs.

WL. Chew — Monograph of Dendrocnide 79

NEW GUINEA (WEST):—Cycloop Mts.: Koejaboe River: Lam 7771,
Oct. 1954 (L, PNH & SING).

NEW GUINEA (NORTH-EAST) :—Sepik Distr.: Leitre Village: Sayers
NGF. 19509, March 1964 (L & SING).—Madang Distr.: Djamu-Klamn:
Schlechter 16615, Oct. 1907 (type of L. schlechteri: P & UC).—Morobe
Distr.: Bewapi Creek: Henty NGF. 14819, Sept. 1962 (BO, L & LAE).—
Oomsis: Corner NGF. 12477, Aug. 1960 (A, BRI, CGE, L, LAE & SING).

PAPUA:—Central Div.: Magarida: Buderus NGF. 24077, Sept. 1965
(SING).

BISMARCK ARCHIPELAGO:—New Britain: Jacquinot Bay: Mair
NGF. 1861, April 1945 (L & LAE).

(29) Dendrocnide longifolia (Hemsl.) Chew in op. cit. 203 (1965).
— Laportea longifolia Hemsl. in Hk. f. Ic. Pl. 26: tab. 2559
(1898). — L. salomonensis Rech. in Fedde, Repert. 11: 182
(1912). — L. warburghii Winkl. in Bot. Jahrb. 57: 509 (1922).
—— D. salomonensis (Rech.) Chew in op. cit. 205 (1965). — D.
warburghii (Winkl.) Chew in op. cit. 207 (1965).

It distinguishes itself by the following combination of characters:
(a) lamina very large, obovate, glabrous, densely covered with
punctiform cystoliths, (b) stipules large, 6-10 cm. long, (c) male
flowers sessile, 4-tepalled, with a very large and fleshy cubical
pistillode, (d) female flowers sessile, in one or two rows around
fleshy peduncular extremities, (e) irritant hairs only on peduncles
and flowers. ,

Distribution: Celebes, Moluccas, New Guinea, Bismarck Archi-
pelago and the Solomon Islands.

Dioecious trees 8-10 metres high. Twigs few, stout to massive,
spreading, with rosettes of large leaves at the apices, with wide pith,
eventually hollow at old age, lightly pubescent, soon glabrous;
lenticels large; petiolar scars very prominent. Lamina (25—) 50-60
(-75) cm. long, (13—) 20-30 (—35) cm. broad, usually obovate, very
coriaceous, glabrous, with very few irritant haris near the mid-rib,
both surfaces very dense with punctiform cystoliths; base broadly
cuneate; apex rather shortly and bluntly acuminate, acumen less
than 1 cm. long; margin sinuate to entire; lateral veins 15—20 pairs,
rather distantly spaced, the basal pair short and less prominent;
intercostals straight, very numerous, rather faint. Petioles 6-10
(—15) cm. long, often very thick, glabrous, devoid of irritant hairs,
dense with punctiform cystoliths. Stipules 6-10 (-15) cm. long,
very coriaceous, usually glabrous, devoid of irritant hairs, dense
with punctiform cystoliths, quite persistent. Inflorescences uni-
sexual, shortly branched racemes, branches mainly borne in one
plane, to 20 cm. long and broad, the females much smaller and
more compact, axillary, usualiy paired, the first branch with
peduncular bracts to 3 cm. long; peduncle pubescent, fairly dense
with short irritant hairs. Male flowers sessile. ca. 1.6 mm. long

80 Gardens’ Bulletin, Singapore — X XV (1969)

Fig. 32: D. longifolia: A = twig & infrutescence; B = pistillode & stamens
of opened male flower; C = female flowers; D = achene.

W —L. Chew — Monograph of Dendrocnide 8]i

and broad, well armed with irritant hairs and punctiform cystoliths,
4-tepalled; tepals ovate; stamens 4,with very short straight fila-
ments, at most as long as pistillode; pistillode cubical, notched at.
the sides, ca. 1.5 mm. cube, almost occupying the floral cavity;
interfloral bracts ovate, ca. 2 to each flower. Female flowers.
sessile. set bifariously along the edge, slightly on the upperside of
the somewhat flattened extremities of peduncles; perianth ca.
1 mm. long, 0.5 mm. broad, 4-tepalled, well armed with short.
irritant hairs; the lateral tepals longer than the medial ones; ovary
ca. | mm. long, ovoid; stigma ligulate, ca. 1 mm. long, fairly
straight; interfloral bracts equilateral-triangular, ca. 0.5 mm. long
and broad at the base, dense with short irritant hairs. Achenes
ca. 3 mm. long, 2 mm. broad, symmetrically ovoid to ellipsoid,
rather flat, with very finely tuberculate pericarp, at least half
covered by the lateral tepals, slightly reflexed, borne sitting on
very fleshy and greatly swollen peduncular extremities.

Hitherto known only from a plant introduced into cultivation:
at Kew from New Guinea, this species is here considered to be:
con-specific with the species of medium-sized trees with massive:
pithy branches common in the Morobe district of New Guinea.
and the Bougainville Island and which Winkler described as.
warburghii and Rechinger as salomonensis. These species have:
in common the characters enumerated in the diagnosis above. The:
long, very narrowly elliptic shape of the lamina of Hemsley’s.
type is entirely abnormal and is no doubt due to cultivation in
an artificial environment.

Various authors have had this species confused with D..
amplissima whose resemblance to this species is quite superficial.

CELEBES :—sine loc.: de Vriese s.n. (L. herb. 908-188-2367; -2370 &
—2392).

MOLUCCAS :—Ambonia: Robinson Pl. Rumph. 310, July-Nov. 1913 (BO,,.
K, L & US).—Ceram: Teysmann 5017 H.B. (U).—Halmaheira: Weda:
Anang 639, May 1938 (L)—Morotai: Tiai: Kostermans 1678, June 1949:
(BO, K, L, PNH & SING).—Ternate: Sasa Besar: Beguin 1437, Feb. 1921
(BO & L).

NEW GUINEA (WEST):—Sorong: Beccari s.n., 1872 (FI. herb. 9492A).

NEW GUINEA (NORTH-EAST):—Morobe Distr.: Oomsis: Corner
NGF. 13501, Sept. 1960 (L & LAE).—Lae Botanic Gardens: Corner NGF.
12454, Aug. 1960 (A, CGE, K, LAE & SING).

BISMARCK ARCHIPELAGO :—New Britain: near Rabaul: Kanehira
4092, Feb. 1937 (A). .

SOLOMON ISLANDS:—Bougainville Island: Crown Prince Mt.:
Womersley NGF. 13383, Oct. 1960 (CGE & SING).—Kolombangara:
Shoulder Hill: Gafui BSIP. 8679, Jan. 1968 (SING).—San Cristobal
Island: Waimamura: Brass 2668, Aug. 1932 (BO & L).—Santa Ysabel
Island: Ulavu village: Hunt RSS. 2919, Oct. 1965 (SING).

CULT: ex New Guinea:—Veitch s.n. (holotype of L. longifolia Hemsl.:
K).

82 Gardens’ Bulletin, Singapore —- X XV (1969)

(30) Dendrocnide microstigma (Gaud. ex Wedd.) Chew in op. cit.
204 (1965). — Laportea microstigma Gaud. ex Wedd. in Arch.
Mus. Hist. Nat. Paris 9: 138 (1856); J. J. Smith in Koord. &
Valet. Booms. Java, 12: 690 (1910), exclud. syn. U. amplissima
Bl. — Urticastrum microstigma (Gaud. ex Wedd.) O.K. op. cit.
(1891).

Characteristics: (a) lamina narrowly elliptic, glabrous except for
the soft long hairs at the sides of mid-rib on the lower surface,
with 14-16 pairs of short, closely spaced lateral veins, (b) stipules
and young petioles dense sericeous, (c) inflorescences shorter than
leaves, compact, (d) male flowers 4-tepalled, sessile, (e) female
flowers subsessile on opposing rows along swollen peduncular
extremities, (f) stigma very short, (g) female perianth shallowly and
broadly toothed. ,

Distribution: Java, Lesser Sunda Islands, Celebes and Moluccas.

Dioecious trees to 10 metres high. Twigs sericeous when young,
well armed with irritant hairs. Lamina (10—) 16-20 (—35) cm. long,
(4-) 6-8 (-10) cm. broad, narrowly elliptic, coriaceous, with some
sericeous hairs at the sides of the mid-rib on the lower surface,
dense with punctiform cystoliths; base cuneate; apex long acu-
minuate, acumen 2-4 cm. long; margin smooth, often slightly
sinuate; lateral veins 12—16 pairs, short and closely set, the basal
ones very short and inconspicuous; intercostals numerous, faint.
Petioles (2—-) 3—5 (—10) cm, long, dense sericeous when young, with
short irritant hairs. Stipules 1-1.5 cm. long, dense sericeous,
coriaceous, caducous. /nflorescences unisexual, branched racemes,
10-15 (-20) cm. long, 5-10 cm. broad, axillary, solitary,
peduncle pubescent. Male flowers sessile, ca. 1 mm. long and broad:
perianth puberulous, dense with minute punctiform cystoliths, with
some short irritant hairs, 4-tepalled; stamens 4, filaments reflexed;
pistillode ca. 0.5 mm. long; interfloral bracts minute. Female
flowers sessile, becoming subsessile at maturity, arranged somewhat
in two opposing rows along extremities of peduncles; perianth ca.
0.75 to 1 mm. long, ca. 0.5 mm. broad, 4-lobed, dense with
punctiform cystoliths intermixed with few irritant hairs; tepals
about equal-size, with rounded apices; ovary ca. 0.5 mm. long,
ca. 0.3 mm. broad; stigma very short ligulate, ca. 0.2 mm.; inter-
floral bracts minute. Achenes ca. 2 mm. long, 1.5 mm. broad.
ovoid to ellipsoid, often sigmoid, slightly bloated, with warted
pericarp, half covered by the perianth, reflexed over the very
short pedicel.

Closest to this is Blume’s D. amplissima with which J. J. Smith
considered the present species to be conspecific. I have made a
detailed study of the types of these two species and now conclude
that though rather similar, the two can easily be separated on at
least three characteristics.

W.-1.. Chew — Monograph of Dendrocnide 83

Wy \
/ I 7 4 \
| | AN !
3cm i by \.
\ “a /

| /
PAOANN /

Fig. 33: D. microstigma: A = twig; B = female flower; C = achene.

84 Gardens’ Bulletin, Singapore — X XV (1969)

JAVA :—Besoeki Prov.: Djember: Koorders 9474, 9511, 9512, 9513,
9516, 9519, 9520, 12813 & 39927 (BO & L). Mt. Raung: Jacobs 4810, May
1957 (L, LAE & SING).—sine loc.: Leschenault 468, or 558?, (holotype of
L. microstigma: P).

LESSER SUNDA ISLANDS:—Bali: Poeloekan: van Steenis 7691,
April 1936 (BO)—Lombok: Rindjani Ma.: Elbert 1679, June 1909 (A, BO,
K & L)—Sumbawa: Mt. Batulanteh: Kostermans 18359, April 1961 (L).—
Sumba: Taboemsoeng: Teysmann 5939 (BO).

CELEBES:—Minahassa: Tompaso: Reinwardt s.n., (L. herb. 908-188-—
2790).—S. —-E. Celebes: Buton Island: Limbo: Elbert 2768, Aug. 1909
(BO & L).

MOLUCCAS :—Sula Islands: Mangole: Bloembergen bb. 29840, Sept.
1939 (A, BO, L & SING).—Buru Island: Nal Besi: Toxopeus 248, July
1921 (BO & L).—Ceram: sine loc.: de Vriese 153 (L).

431) Dendrecnide amplissima (Bl.) Chew in op. cit. 202 (1965).
— Urtica ovalifolia Bl. Bijdr. 504 (1825), non Stokes (1812). —
Urera amplissima Bl. Mus. Bot. Lugd. Bat. 2: tab. 22 (1857).
— Laportea amplissima (Bl.) Mig. Fl. Ind. Bat. 1 (2): 232
(1859) et Ann. Mus. Bot. Lugd. Bat. 4: 301 (1869); Wedd. in
DC. Prodr. 16 (1); 87 (1869); Merr. Inter. Rumph. Herb. Amb.
200 (1917), partim. — Urticastrum amplissimum (BIl.) O.K. Rev.
Gen. Pl. 635 (1891).

Characterised by: (a) lamina elliptic to slightly obovate, with at
‘least 20 pairs of lateral veins, the upper surface entirely glabrous,
the lower sparsely covered with very short, appressed irritant hairs,
(b) male flowers pedicellate separated from each other (not
glomerulate), 4-tepalled, with 2—3 rather long irritant hairs, other-
wise glabrous, (c) female flowers sessile, distichous on the pedun-
cular extremities, with a rather short stigma.

Distribution: Celebes & Moluccas.

Dioecious tree to ca. 8 metres high. Twigs light puberulous, later
glabrous, without irritant hairs; petiolar scars very prominent.
Lamina (20—) 30-35 (-45) cm. long, (12—) 15—20 (—25) cm. broad,
elliptic to very slightly obovate, coriaceous, glabrous except for
the few very short, appressed irritant hairs on the lower surface,
both surfaces dense with punctiform cystoliths; base cuneate; apex
very short acuminate, acumen narrow, sharp, usually less than
1 cm. long; margin sinuate; lateral veins 22—25 pairs, very rarely
less, very closely set, the basal pair very short; intercostals very
numerous, straight, faint. Petioles (4-) 6-8 (-10) cm. long,
puberulous when young, later entirely glabrous, dense with puncti-
form cystoilths. Stipules 4-7 cm. long, light pubescent with very
few irritant hairs, caducous. Jnflorescences branched racemes, ca.
23 cm. X 35 cm. spread, females generally much smaller, axillary,
usually solitary; peduncular bracts at the lower axils to 1 cm. long,
those at the apex minute, narrow scaphiform, unpaired; peduncles
fairly dense with short irritant hairs. Male flowers pedicellate, ca.
1.25 mm. long and broad, glabrous except for a few rather long
irritant hairs, dense with puctiform cystoliths, 4-tepalled: stamens
4, with short very slightly curved filaments, the connective-filament
joint placed on the inside; pistillode obpyramidal, thin, less than

W.-L. Chew — Monograph of Dendrocnide 85

4om del. ;

Fig. 34: D. amplissima: A = leaf; B = male inflorescence, part; C =
opened male flower; D, E & F = female flowers (arter Blume).

86 Gardens’ Bulletin, Singapore — X XV (1969)

0.5 mm. long; interfloral bracts to 1 mm. long, triangular, 2 per
flower, pedicel to 1 mm. long; flowers borne loosely, not glomeru-
late. Female flowers subsessile to sessile; perianth ca. 1 mm. long,
0.5 mm. broad, 4-tepalled, with a few longish irritant hairs, other-
wise glabrous, well covered with punctiform cystoliths; ovary ca.
0.8 mm. long, 0.5 mm. broad, ovoid; stigma ca. 0.5 mm. long,
somewhat brush-like; interfloral bracts minute; flowers distichous
on peduncular extremities. Achenes to 3 mm. long, 2.5 mm. broad,
broadly ellipsoid, flat, pericarp smooth; only the base covered by
the perianth; slightly reflexed, still sessile; peduncular receptacle
not fleshy.

There is a very superficial resemblance between this species and
D. longifolia, also from the same region, from which one can
easily distinguish on closer examination. Merill (1917) had these
two species confused — of the four collections of Robinson from
Amboina which he studied, no. 310 is found to belong to D.
longifolia and not this species.

This species is very close to D. microstigma. Both have rather
similar female flowers which are subsessile and have very short
stigmas. Vernacular names: Poeloes djalatang, Daun gattal and
Polot-mera (Indonesian).
iste ha Reinwardt 539, L. herb. 908-188-2793, Oct.

MOLUCCAS:—Amboina: Robinson Pl. Rumph. Amb, 311 (A, L, NY &
US), 312 (L & US),. 313 (BO, F, L, NSW, SING & US).—-sine loc.:
Zippelius s.n. (type of U. amplissima: L & UV).

JAVA (cult. in Hort. Bot. Bogor): Blume s.n., L. herb. 908-189-1970
(type of U. ovalifolia: L); Teysmann s.n. (K, L, MICH & U).

(32) Dendrocnide kjellbergii Chew, spec. nov.

Characteres: D. amplissimae similis sed floribus femineis
omnino sessilibus, et perianthio quadridentato, dentibus latis vix
altis arcte dissimilis.

Distribution: Celebes.

Dioecious trees ca. 7 metres. Twigs thick, pubescent at the apex,
with few irritant hairs. Lamina 25-35 cm. long, 15-25 cm. broad,
oblongish elliptic, rarely broad ovate or obovate; the upper
surface with few irritant hairs on the mid-rib, otherwise entirely
glabrous, dense with punctiform cystoliths; the lower surface
pubescent to woolly, with irritant hairs on mid-rib and veins;
base round, truncate to slightly cordate, often slightly peltate; apex
very shortly acuminate, acumen to | cm. long; margin very finely
distantly dentate; lateral veins ca. 12-14 pairs, prominent, the
basal pair reaching to 4 the length of the lamina; intercostals
numerous, not very prominent, not straight. Petioles 10-15 cm.
long, dense pubescent, with irritant hairs, soon subglabrous; well
covered with punctiform cystoliths. Stipules 5-10 cm. long,
puberulous to pubescent, dense with punctiform cystoliths, quite
persistent. Inflorescences branched racemes, 20-25 cm. long, 7-10
cm. broad, axillary, usually solitary; peduncular bracts at the
lower axils less than 1 cm. long, unpaired; peduncles pubescent,

EEE

W -L. Chew — Monograph of Dendrocnide 87

Fig. 35: D. kjeilbergii: A = twig: B = male inflorescence: C = female
flowers: D = male flower; E = achene.

88 Gardens’ Bulletin, Singapore — X XV (1969)

well armed with irritant hairs. Male flowers pedicellate, ca. 1.5
mm. long and broad, pubescent, with rather long irritant hairs here
and there, sparsely covered with minute punctiform cystoliths,
4-tepalled; stamens 4, with rather long, strongiy reflexed filaments;
pistillode obconical, ca. 0.5 mm. long, 0.3 mm. broad; interfloral
bracts ca. 0.5 mm. long, triangular, 2 or 3 to each flower; pedicel
to 1 mm. long, puberulous; few flowers in each glomerule. Female
flowers entirely sessile, perianth ca. 0.5 mm. long and _ broad,
shallowly 4-toothed, sparse with very short irritant hairs, other-
wise giabrous, with punctiform cystoliths; the teeth of the perianth
almost equal-sized; ovary.ca. 0.5 mm. long, ca. 0.3 mm. broad,
protruding greatly from the perianth; stigma ligulate, ca. 0.6 mm.
long, somewhat brush-like at the base, interfloral bracts absent;
flowers somewhat distichous on peduncular extremities. Achenes
ca. 2 mm. long, 1.5 mm. broad, ovoid, flattened, with smooth
pericarp, very shortly stipitate, the perianth remaining as a very
shallowly 4-toothed plate hardly covering the very short stipe.

The somewhat distichous disposition of the female flowers and
the shortness of the stigma put this easily into the D. amplissima
group of species from which it distinguishes itself by the female
perianth being shallowly 4-toothed and the achene definitely,
though very shortly stipitate. On the other hand, the sessile nature
of the female flowers puts this amongst the D. schlechteri group
of species. However, judging on the overall characteristics, I
prefer to place this species in the former group ‘with which it
definitely has closer affinities.

CELEBES :—Menado Residency: Poso subdistr.: Eyma 3536 (A & L) &
3734 (L).—Makassar Peninsula: Bantimoeroeng: Buwalda 3809, Feb. 1938
(BO & L). Lombasang: Bunnemeyer 11108, April 1921 (BO & L).—Lake

Towuti Distr.: Liasa Kjellberg 2258, Sept. 1929 (holotype in S; isotype
in BO).

(33) Dendrocnide harveyi (Seem.) Chew in op. cit. 203 (1965). —
Laportea harveyi Seem. Fl. Vit. 238. tab. 59 (1868).—L. milnei
Seem. op. cit. (1868).—Urticastrum harveyi (Seem.) O.K. &
U. milnei (Seem.) O.K. op. cit. (1891).—D. milnei (Seem.) Chew
in op. cit. 204 (1965).

Characterised by: (a) lamina broad to very broad ovate, margin
sinuate to tooth, base round to cordate, glabrous to sub-glabrous
on both surfaces, (b) male flowers long pedicellate, pedicel at
least as long as perianth even before anthesis, (c) perianth hardly
deevloped in the female flower, (d) achenes exposed completely,
stipitate, warty, generally flattened and borne on swollen pedicels
almost without irritant hairs.

Distribution: Fiji, Tonga, Samoa and Cook Islands.

Dioecious tree ca. 10 metres high, often to 15 metres. Twigs
lightly puberulous, soon quite glabrous; without irritant hairs:
petiolar scars very prominent and raised. Lamina (13-) 20-26
(—30) cm. long, (9—) 12-20 (-25) cm. broad; broad ovate; coria-
ceous; mid-rib on the underside light puberulous, otherwise
entirely glabrous and without irritant hairs; cystoliths punctiform.

W.—L. Chew — Monograph of Dendrocnide 89

WIS TF 4.

=

aE ROU teee

O

BN RAR ESS SS

7
=

imm

15ram

Fig. 36: D. harveyi: A = twig; B = male flower; C = female flowers;
D = achene.

90 Gardens’ Bulletin, Singapore — X XV (1969)

dense on both surfaces; base rounded to cordate; apex acute to
short acuminate; margins coarsely dentate, irregularly crenate to
unevenly sinuate; lateral veins 8-10 pairs, the basal pair very
prominent, reaching to a third the length of lamina; intercostals
numerous, prominent. Petioles 8-12 (—15) cm. long; very light
puberulous, with few minute irritant hairs; cystoliths punctiform,
dense. Stipules 1-3 cm. long; puberulous; without irritant hairs;
well covered with elongated cystoliths. /nflorescences unisexual;
branched racemes; to 15 cm. long and broad; often paired;
peduncular bracts small; peduncles well armed with short irritant
hairs; flowers in loose fascicles. Male flowers pedicellate; perianth
ca. 1.5 mm. long and broad, puberulous to subglabrous, with very
few short irritant hairs; 4-tepalled; tepals ovate; stamens 4, with
very slightly reflexed filaments; pistillode to nearly | mm. long:
interfloral bracts small, irregular in number; pedicels to about
2 mm. long. Female flowers shortly pedicellate; perianth hardly
developed to at most 0.2 mm. long, 0.1 mm. broad, faintly 4-
toothed. well covered with punctiform cystoliths; ovary ca. 0.75-1
mm. long, half as broad, the base hardly covered by the minute
perianth; stigma ligulate, to 2 mm. long; interfloral bracts minute,
irregular in number; pedicel elongating at fruit. Achenes ca. 2 mm.
long and 1.5 mm. broad, very shortly stipitate; asymmetrically
broadly ellipsoid; pericarp minutely warted, flat, only the very
short stalk covered by the minute perianth; stigma displaced to one
side; achene reflexed over the fleshy pedicel.

I have examined the holotype of D. milnei and have found it
to be conspecific with this species. Although the specimen is a
scrape, it contains a portion of the female inflorescence with
female flowers quite typical of D. harveyi.

Reported to be a dominant tree species in canyons in primary
forests below 800 metres in altitude. Vernacular names: Salato
(Fiji, Tonga); Magiho (Cook Islands).

This is the only species in the genus where the female perianth
is hardly developed: in fact in many flowers, it is absent.

FIJI:—Kandavu Island: near Ngaloa Bay: Smith 45 (BO, K, NY. S,
UC & US)—vVanua Levu: Taviuni: Seemann 426, May 1860 (A, K &
MEL).—Viti Levu: Mba: Smith 4428 & 5282 (A, CRI, K, L, S & US).—
sine loc.: Harvey s.n., Nov. 1855 (type of L. harveyi: BM); Milne s.n.,
1853 (type of L. milnei: BM).

TONGA :—Eua Island: Fuai: Yuncker 15329, March 1953 (A, U & US).

SAMOA :—sine loc.: Powell 205 & 329 and Whitmee 271 (K).

COOK ISLANDS :—Niue Island: Lakepa village: Yuncker 9927, Jan.
1940 (A & UC).

(34) Dendrocnide mirabilis (Rechinger) Chew in op. cit. 204
1965). — Laportea mirabilis Rechinger in Denkschr.. Akad.
Wiss. Math.-Natur. 89: 543. tab. 1 (1914).

Characterised by: (a) shrubby tree with a succulent base, (5)
lamina ovate, plicate, the upperside glabrous, with 2 erect fleshy
appendages in the middle, the lowerside with very few short
irritant hairs, lateral veins palmate and penninerved, (c) male
flowers sessile, 4-tepalled.

W.—L. Chew — Monograph of Dendrocnide

Fig. 37: D. mirabilis: A = twig; B = male flowers.

91

92 Gardens’ Bulletin, Singapore — X XV (1969)

Distribution: New Guinea and Solomon Islands.

Tree ca. 4 metres high, succulent at the base. Twigs entirely
glabrous, without irritant hairs nor cystoliths. Lamina 18-23 cm.
long, 15-18 cm. broad, ovate to somewhat round, coriaceous,
the upperside entirely glabrous, bearing 2 fleshy appendages at
the point of origin of the palmate veins, the lowerside sparsely
covered with short irritant hairs and rather pubescent on the
mid-rib, particularly at the point of origin of the palmate veins,
both surfaces densely covered with punctiform cystoliths; base
decurrent unequally along petiole; apex very shortly acuminate,
acumen ca. 5 mm. long; margin greatly sinuate; lateral veins ca.
12 pairs, the 2nd to the 6th pairs arising palmately from the
mid-rib about 4 cm. from the base, the rest of the veins penni-
nerved; intercostals, particularly those at the distal half, rather
straight. Petioles 8-12 cm. long, covered with short irritant hairs
and somewhat elongated cystoliths. Stipules 6-8 cm. long, to 1.5
cm. broad, fairly coriaceous, persistent, pubescent. /nflorescences
unisexual, branched racemes, ca. 10 cm. long, 8 cm. broad, axillary,
paired; peduncular bracts at the lower axils oblong, ca. 1.5 mm.
long, | mm. broad; peduncle closely covered with very short
irritant hairs; flowers in glomerules of 7 to 10. Male flowers
sessile, ca. 1.5 mm. long and broad, well covered with short
irritant hairs and punctiform cystoliths, 4-tepalled; tepals ovate:
stamens 4, on slightly reflexed filaments; pistillode ca. 0.7 mm.
long and broad at the apex, obpyramidal; interfloral bracts minute.
Female plant unknown.

The peculiar leaf form makes this species the easiest to recognise
in the genus. In fact the distinction is so great that the description
and photo illustration supplied by Rechinger are enough to enable
one to fix this species without recourse to examining the type
material.

Rechinger had only a sterile specimen to base this species on
while the other collection cited here, Darbyshire 1016, is fortunately
a staminiferous specimen. Until we have the female plant, it is
useless to speculate on the relationship of this plant to others
in the genus.

Known so far from cultivation by the villagers as an ornamental
because of the interesting leaf-form. Vernacular name: Apungeh
(Mekeo language in Maipa).

NEW GUINEA (PAPUA):—Central Distr.: Kairuku Subdistr.: Akaifu
River: Maipa village: Darbyshire 1016, Sept. 1962 (LAE).

SOLOMON ISLANDS:—Bougainville Island: Djup: Rechinger 4940
(photo seen!).

(35) Dendrocnide celebica Chew, spec. nov.

Characteres: D. rigidifoliae affinis, sed bracteis maximis, floribus
masculis axillaribus singulatim gerentibus differt.

Distribution: Celebes (endemic! )

Dioecious trees ca. 2 metres high. Lamina 25 cm. long, 7 cm.
broad, narrowly obovate, the upperside glabrous, densely covered

W.—L. Chew — Monograph of Dendrocnide 93.

imm |].

Fig. 38: D. celebica: A = twig; B = male flower with bract.

94 Gardens’ Bulletin, Singapore — X XV (1969)

with punctiform cystoliths, the lowerside with few short irritant
hairs on the veins and veinlets and with few dispersed elongated
cystoliths; lateral veins ca. 13 pairs; apex sharply acuminate to
somewhat caudate, acumen to 4 cm. long; base broadly cuneate;
margin lightly dentate, especially towards the apex. Petioles to
10 cm. long, dense with punctiform cystoliths, sparse with long
thin irritant hairs. Stipules to 5 cm. long, well covered with
irritant hairs and punctiform cystoliths. Inflorescences unisexual,
weak, poorly branched racemes, to 12 cm. long; peduncle dense
with short irritant hairs. Male flowers pedicellate; perianth ca.
0.5 mm. long, 0.3 mm. broad, with few long irritant hairs and
punctiform cystoliths, 4-tepalled, with 4 stamens; pistillode ca.
0.4 mm. long, club-shaped; pedicel ca. 0.4 mm. long; each flower
in the axil of a boat-shaped bract 2.5 mm. long, | mm. broad.
Female plant unknown.

I am certain that this represents an undescribed species well
characterised by the male flowers being borne each in the axil
of a large bract and the shape and size of the lamina.

In the former character, this species approcahes D. rigidifolia
of the Philippines with which it is obviously closely related. In
the latter character, it comes very near to a narrowly-leafed form
of D. sinuata found in Sumatra with which it probably has but
superficial resemblance.

CELEBES:—Batoe Watoewila: G. Kjellberg 1041, 25 March 1929
(holotype in S; isotype in BO).

(36) Dendrocnide kajewskii Chew, spec. nov.

Characteres: Ab omnibus speciebus adhuc cognitis laminis
giganteis subtus pubescentibus haec species bene differt.

Distribution: Solomon Islands (endemic! )

Dioecious trees 5-6 metres high. Lamina 150 cm. long, 50 cm.
broad, the upper side glabrous, dense with punctiform cystoliths.
the lower side pubescent, well covered with numerous irritant
hairs; margin broadly toothed. Petioles to 25 cm. long, closely
covered with very short irritant hairs. /nflorescences branched
racemes, axillary, solitary, to 25 cm. long, 10 cm. broad, with
peduncular bracts to 3 cm. long; peduncle dense with irritant
hairs. Male flowers pedicellate; perianth ca. 1.5 mm. long, 2 mm.
broad, 4-lobed, with 4 stamens on long filaments, well armed with
irritant hairs; pistillode small; pedicel ca. 1 mm. long; interfloral
bracts to 1.5 mm. long, numerous. Female plant unknown.

The immense size of the lamina together with the other
characteristics here mentioned set this well apart from all the
other species of Dendrocnide. D. longifolia is perhaps the
closest to this, whose relationship cannot be properly determined
until its female plant is found.

SOLOMON ISLANDS :—Bougainville Island: Buin: Kugumaru: S. F.
Kajewski 1974, July 1930 (holotype in A; isotypes in BO & BRI).

W —L. Chew — Monograph of Dendrocnide 95

Species Incognitae

(1) Dendrocnide contracta (Bl.) Chew in op. cit. 202 (1965). —
Urtica contracta Bl. Bijdr. 505 (1825). — Laportea contracta
(Bl.) Miq. Fl. Ind. Bat. 1 (2): 232 (1859), pro syn. — L. costata
(Mig.) Miq. var. contracta (Bl.) Miq. op. cit. — L. stimulans
(L.F.) Mig. var. contracta (Bl.) Miq. Ann Mus. Bot. Ludg. Bat.
4: 302 (1869).

Type specimen: — U. contracta: Blume from Tjiandjoer, West

Java (in Leiden herb.?).

In the herbarium at Leiden, there are numerous early collec-
tions from Celebes which have been equated with this species
by previous authors. I have studied and found them to belong to
D. microstigma. Amongst these collections, there is one, collected
by Reinwardt from Tompaso in Celebes (Leiden herb. 908—-188-
2790), which bears the name U. contracta Bl. in Blume’s hand-
writing. If this is, the type material of Blume’s species, then
D. microstigma should be conjoined to this species. I am
inclined however to the view that this is not the type (as the type
was collected from Tjiandjoer in West Java) but one that was
erroneously named by Blume subsequent to the publication of
this species. What this species is then, | do not know.

(2) Dendrocnide gigantea (Poir.) Chew in op. cit. 203 (1965). —
Urtica gigantea Poir. Encyc. Meth. Bot. Suppl. 4: 225 (1816).
— Urera commersoniana Wedd. in Ann. Sc. Nat Ser. 3. 18: 202
(1852).

Distribution: Melanesia.

This is suspected by many to be similar to D. sinuata. | have
examined the holotype in Jussieu’s Herbarium in Paris. It has
such a small and immature inflorescence that its identity cannot
be determined with certainty. Judging from the leaves, I would
say it comes very close to D. latifolia.

As for its origin, Poiret had it erroneously recorded as from
the district of Magellan (South America). The small label attached
to the holotype bore a note by an unknown person to the effect
that the specimen was “found amongst a bundle of Magellan
materials’. | am quite certain it came from some islands in
Melanesia.

_ Pacific Islands: — Melanesia: sine loc.: Commerson s.n. (in
Herb. Juss. Cat. no. 16880: P).

(3) Dendrocnide sessiliflora (Warb.) Chew in op. cit. 206 (1965).
— Laportea sessiliflora Warb. in Bot. Jahrb. 13: 292 (1891);
Winkl. in Bot. Jahrb. 57: 508 (1922).

Distribution: New Guinea and Bismarck Archipelago.

Winkler (1922) clearly pointed out the doubtful nature of
this species which is based on a mixture compositum. The des-
cription supplied by Warburg, though detailed, is unfortunately
lacking in diagnostic characters. As I have not seen the type
(destroyed in Berlin) or any of the collections identified by
Winkler with this, I have no choice but to leave this uninterpreted.

‘96 Gardens’ Bulletin, Singapore — X XV (1969)

Species Excludenda

Dendrocnide coerulea (Bl.) Mig. Pl. Jungh. 1: 31 (1851). —
Urtica coerulea Bl. Bijdr. 495 (1825).

Weddell found this to belong to Cypholophus and had it
transferred to C. lutescens Wedd. I have examined Junghuhn’s
collection and agree with Weddel in its interpretation.

Bibliography

Backer, C. A. and Bakhuizen v. d. Brink, Jr. (1965). Flora of Java.
2: 38-39.

‘Chew, W.-L. (1965). Laportea & Allied Genera. Gard. Bull Sing.
21: 195-208.

Gagnepain, F. in Lecomte. (1929). Flore Generale de I’ Indochine.
5: 854-860.

Gaudichaud, C. in Freycinet. (1830). Voyage Autour du Monde
PEPOE LY: Uraine (Bot.) 498.

Hooker, J. D. (1888). Flora of British India. 5: 549-550.

Li, H.-L. (1963). Woody Flora of Taiwan. 134-135.

Miquel, F. A. W. (1851-1857). Plantae Junghuhnianae. 29-31.
Miquel, F. A. W. (1859). Flora Indiae Batavae. 1 (2). 229-232.

Rechinger, K. (1913). Botanische u. Zoologische Ergebnisse.
Denkschriften der Math.-Natur. Kl. Kaiser. Akad. Wissen.
Wissen. 89: 542-544.

Ridley, H. N. (1923). Flora of the Malay Peninsula. 3: 359-360.

Robinson, C. B. (1911). Philippine Urticaceae. Philip. J. Sc. (Bot.)
5: 477-487.

Smith, J. J. in Koorders and Valeton. (1910). Boomsoorten op
Java. 12: 675-692.

Weddell, H. A. (1856). Monographie de la Famille des Urticees.
Arch. Mus. Hist. Nat. Paris, 9: 121-142.

Weddell, H. A. (1869). Urticaceae. DC. Prodr. 16 (1): 77-87.

Winkler, Hub. (1922). Die Urticaceen Papuasiens. Bot. Jahrb.
57. 501-509.

W —L. Chew — Monograph of Dendrocnide 97

Index to collections studied
Abbreviations used :—

amp = amplissima. mic. = microstigma.
bas. = basirotunda. mir. = mirabilis.
car. = carriana. morob. = morobensis.
cel. = celebica. moroi. = moroides.
coral = corallodesme. ner. =! nervosa.
cord. Fast COR sata, obl. = oblanceolata.
cata Fi =F pigie. pel.-mur. = peltata var. murrayana.
den. = densiflora.
» pel.-pel. = peltata var. peltata.
ell. = ,elliptica. be hu donbula
NY = 1 =
exc. =) eR CElSa. P P hi a
Tec! = rechingeri.
har. = harveyi. : y 459 a¥,
kaj. = kajewski. rig. = rigidifolia.
kje. = kjellbergii. sch. = schlechteri.
lat. = |atifolia. sin. =m siruata.
lon. = longifolia. sti. = stimulans.
luz.-ana. = luzonensis var. Lab Oe ulselausa)
anacordioides. :
luz.-luz. = luzonensis var. ter. = ternatensis.
luzonensis. tor. = torricellensis.
mey.-mey. = meyeniana f. ure. = urentissima.
meyeniana.
mey.-sub. = meyeniana f. has a ce |
subglabra. vit. = vitiensis.

Aban & Singh: SAN. 30072 = sti. — Acuna, R. A.: 23125
= sti. — Aed, E.: NSW. 77936 = pho. — Aet (exp. Lundquist):
421 = ter. — Agama, J.: 1026 = obl.; 9891 = sti. — Ah Wing,
J.: 35391 = obl. — Alambra & Caules: 23214 = sti. — Allen,
Biv M.c..4493) = sestine 4667.0 = —sin.ye 4718) \& 472807 sti.;
4846 = sin.; 4877 & 4878 = sti; 4879 = sin.; 4880 = .
4887 = sin.; 4888 = sti. —:.Alston, A. H. G.: 259. = sin;
14463 = sti. — Alvins: 871 & 1109 = sti. — Amdjah: 172 &
637 = sti. — Ampuria, J.: SAN. 25271 & 35281 = sti. — Anang
(exp. de Haan): 639 = lon. — Anderson, J. A. R.: 1258, 4286,
42794 Gi 9786 = stie 12535°—ell.7°12580 &:12913 — stu; 14375
= obl.; 15282, 15359 & 24045 = sti.; 20242 = obl. — Andrews,
C. W.: 147 = pel.-mur. — Anonuevo, P.: PNH. 13733 = sub. —
Anwar: SAN. 37936 = sti. — Aris: Ja. 3353 = mic. — Ashton,
Pe Sieune3236. =! Si: $1\46719 \HSobl.i§. 22085 = sti.

Backen..G. As Jeli 5t, Gy, 26lla-so28 = Sti.;.4820 —-sin;
COL? &. /aVJ, = stirs 9475 =~ gun; 120264—' pel.-pel:;.,12263 &
LOZI4, = St. 10042.= sin.) 4/293) '= sti.:’ 18268 = pel-pel;
18491, Za095s, A3742 OC 23907 = sti; 25098 = pel-pel.; 25232
= Sin vlene ee gig = S304) =; ncas31155 = sth —
Baki. Vv. a, Bonk Jr: 1630,= sit,— Balansa; B.: 585 = wure,;
2528 & 3289 = sti. — Barnard: MR. 33 = sin. Bartlett, H. H.:
6849, 7734 & 8186 = sti; 13453 = mey.-mey.; 15631 & 15741
= sub. — Battah & Anthony: SAN. 37678 = ell. — Bauerlen,
W.: 493 = ter.; NSW. 77926 = moroi. — Baumann, M::
14819 = lat. — Beccari, O.: 218 & 992 = sti. — Becking: 118

98 Gardens’ Bulletin, Singapore — X XV (1969)

= mic. — Beer: BSIP. 5107. 6675, 0iJ24d220 & TIGS = Tat.
— Beguin, V. M. A.: 778, 1232, 1437 & 1782 = lon. — Betche,
E.: 317 = vit. — Beumee; J.: 12 = sin; 19 = sti.; 4872 = ‘sin.;
6081 & 6753 = sti. — Bliss, G.: 610 = rec. — Bloembergen, S.:
3027 = sti.; bb. 27110 = cord.; bb. 29840 = mic. — Boerlage:
564 = lon. — Bolster, F.: 184 = mey.-mey. — Bon: 90 = sin.:
2744 & 2843 = ure.; 5838 & 6051 = sin. — Boorman, J.:
31 = pho. — Borssum-Waalkes, J.: 571 = sin.; 3240 = ter.:;
3366 = moroi. — Brand, D.: SAN. 24494 = ell. — Brass, L. J::
690: = ter.po198hice. morons-.2668 = lon. 2357S"= fec.;: 466
= lat.; 5490, 7434 & 7434A = ter.; 22881 = car.; 24146 &
24200 =ele@rald).24382 = ten, 29235 = tor: 37402 = mer.
32555 ‘& 325600 ters; 32593 54 nér.: 32688! =“sch.; 32699"
32719 = ner. — Brass & Versteegh: 10485 & 12594 = pel.-pel.
— Brinkman, R.: 644 = sti.; 650 = sin.; 816 = sti. — Britton,
B. B.: PNH. 19599 = den.; 19604 = ven. — Brown, R.: 3094
= exc. — Bruggeman, M. L. A:: 527 & 841 = sti. — Bruderus,
J.: NGF. 24077 = sch. — Bumkoed: 89 = sti. — Bunnemeyer,
H. A. B.: 439, 1305, 3089, 3542, 3598, 3776, 5041 & 8646 = StL.
8769, 9028, 9383, 9427 & 10472 = sin.; 11108 = kje. — Burger,
D.: 2083, 6655 & 6672 = pel-pel. — Burkill, I. H.: SFN. 6347
= sti. — Burton, R. E.: 165 = lat. — Buwalda, P.: 3809 = kje.;
4162 & 4292 — cord.; 7581 = sti..— Buoysman’ Mi: “29Sce
435 = sti.

Carolyn & Frake: PNH. 35995 = sti. — Carr, C. E.: 11127
= pel.-pel; 11207 = cord.; 11723 = pel-pel.; 13644, 13969,
15573 & 15775 = car.; 16061 = pel.-pel.; SFN. 27088 = sti. —
Chai, P.: S. 19766 = sti. — Cheel, E.: NSW. 77928 = moroi.
— Chevalier: 30985 = sti. — Chew Wee-Lek: CWL. 537 = sti;
647 = ell. — Chew & Corner: RSNB 4066 = sti.; 4985 = obl.:
4986 & 6025 = sti.; 7048 = obl.; 7070 = sti; 7094.= ob —
Chew et al.: RSNB. 29, 234 & 1474 = sti; 1955 = obl.; 2850
= sti. — Ching, R. C.: 7329 = ure. — Christophersen, E.:
302 = vit. — ‘Clarke, GB: 36225 =«sin. —~ Clason-Laarman:
111 = st — Clemens; (244 '& 335 —?rseh.; 462 = sisvel0heS
ter.; 6282 = morob; 7153 = car.; 8453, 8585 & 8660 = ner.;
8684 = sch.; 10074 = obl.; 10402 = sti.; 10475 = obl.; 10774
= cord.;-11094 = ‘sch; 416703 —.-sub.; -17035.5 = mey--meys
17409 & 17410 = sti.; 20227 = ell.; 21615, 28478, 29330 &

29933 soe §21031%S 32791 —_obl: 34095 \— st: 40748 — ok,
41462 & 41584 = ner.; 43662 = pho. — Colfs, P.: 338 = mic.
— Conklin, "FC.: “PNHO 17554 = mey.-sup.; 18933" =" Car:

18985 = luz.-luz. — Conklin & Buwaya: PNH. 80485 = ell. —
Copeland, E. B.: 608 = mey.-sub. — Copeland, H. F.: 78 = sub.
— Coppe, D:: 77 = lat. —“Comer, FJ. a. NG, 12400, —
lon.; 12464 = ter.; 12466, 12472 & 12477 = sch.; 13501 = lon;
13561 = lat. — Cowmeadow: BSIP. 2518 & 4774 = lat. —
Cuadra, Aus” A. ‘1430 =" st. —“Cumune: S22" Tiz.-luz.) S67
= mey.-mey. — Curran, H. M.: 5149 = mey.-mey. — Curtis.
Ce AGC ae oD ee

W —L. Chew — Monograph of Dendrocnide 99

Peer wt 2... a). = St..— Daman Er. 76 = bar. —
Danser. .. 0040" = Sit’ — Darbyshire, P-’J.:° 629 = cord.;
1016 = mir. — Darbyshire & Hoogland: 7980 = coral.
Darling, F. W.: 18685 = luz-luz. — Darnton, [.S.: 539 = sti.
— Dee: 452 = bas. — Degener, O.: 14797, 14822, 15034 &
15479 = vit. — Degener & Ordonez: 13951, 14203 & 14246 =
vit. — Dennis, G.: 248C = vit.; RSS. 2127 = ner. — DeVore
& Hoover: 182 = sti. — Dietrich, A.: 605, 1438, 1889 & 2192
= pho. — Docters van Leeuwen 11086 & 11226 = ter.

Ebalo, L. E.: 744 = sti. — Eberhardt: 1260 = sin. — Edano,
G. E.: PNH. 11389 = cra.; 11934 & 26792 = mey.-mey.; 48786
= luz.ana.; 48816 = luz.luz. — Einar & DuRietz: 4073 = exc.
— Elbert, J.: 780, 1679 & 2768 = mic. — Elmer, A. D. E::
6464 & 7351 = mey.-mey.; 7442 & 8087 = sti.; 8273 & 8932 =
mey.-mey.; 10499 = sti.; 11948 = ven.; 13324 = mey.-sub:;
13966 = sti; 14019 = luz.-luz.; 14516 = sti.; 14974 = sub.;
15206 = luz.luz.; 15547 & 15548 = sti.; 15608 = mey.-sub.;
£71529 = were 17262. V=— “amey-sibi:: 1 7300.x= —ver.p° 17340 V&
Pree) == Si U2. == sub; 18235, =. mey.-mey.; 20859, 20887
om 2111) = St, 29455 & Zier zZ = ell. — Endert; ‘F.H.°*2378
=str.; 3510''= obi; 46265'52933°5297 & ‘5298 =" sti, —‘Enggoh:
O70 = su yma, ‘PS POS2I8 =" ter 35396 &*3754! "hie

Faber, F. C.: 3215A = sti. — Faurie: 606 = mey.-mey. —
Fawcett: 1879 = pho. — Fenix, E.: 214 mey.-mey.; 3719 =
mey.-sub,,..12583 s: mey;-mey,; 12748..— -rig.; 15151. = sti;
26044 = mey.-mey. — Fischer, C. E.: 2488 = sin. — Floyd, A.:
NGF. 3498 = cord.; 7281 = ner. — Floyd & Hoogland: 3819 =
ner. — Forbes, H. O.: 403 = ter.; 1053 & 1054 = sti. — Forest-
Guards: 7405 = sin. — Fox, R. B.: PNH. 4883 = mey.mey.;
8993 = mey.-sub. — Foxworthy, F. W.: 12233 = mey.-mey.;
25712 = sti. — Frake, C.: PNH. 35995 = sti. — Franc, L:
1229 = lat. — Franck, C. W.: 1004 & 1400 = sti. — de Fretes:
5577 & 5578 = amp.

Gadoh: KL. 1279 = sti. — Gafui, I. H.: BSIP. 8679 = lon. —
Gamble, J. S.: 2409A, 7662 & 15940 = sin. — Gammill, J. A.:
236 = mey.-mey. — Garber: 780 = vit. — Gardner: 812 = sin.
— Gibbs, L. S.: 4104 = sti. — Gibot, A.: SAN. 35835 = stL;
37035 = ell.; 60789 = sti. — Gillespie, J. W.: 2610 = har.;
2864 = vit.; 3076 & 3383 = har.; 3706 & 3756 = vit.; 4163 =
har; 4425 & 4621 = vit.; 4705 = har. — Glassman, S. F.: 791
= mey.-mey.; 2685 & 2714 = lat. — Goy & Smith: 317 = pho.
— Greenwood, W.: 74 = har.; 142, 248 & 292 = vit.; 766 =
har.; 1234 = vit. — Griffith herb.: 1285 & 4513 sin. —
Guillaumin, A.: 8504 = lat — Gwynne-Vaughan: 373 sin.

Haan, G. de: 139 = car.; 20la = lon. — Hagerup, O.: 48 =
sti. — Hallier, H.: 163 & 478 = sti.; 2775 = obl.; 4714 = den.
— Hamilton, F. B.: 2004 = sin. — Haniff, Md.: SFN. 10340 =
sin. — Hartley, T. G.: 9946 = ner.; 10032 =cord.; 10038 = sch.;
10900 = pel.-pel.; 12475 = car. — Havel & Kairo: NGF.

100 Gardens’ Bulletin, Singapore — X XV (1969)

17150 = pel.-pel.; 17282 = car. — Haviland, G. D.: 153, 347
& 991 =, -ell.;. 1222 & 1230K -— sti134! = obl. Hela:
1171 = moroi. — Henderson, M. R.: 11498, 20281 & 21707 =
sti. — Henry, A.: 720 = mey.-mey. — Henty, E. E.: NGF. 9828
= lon.; 10674 = ner.; 14399 = coral.; 14819 = sch. — Henty &
Sayers: NGF. 20572 = ner. — Hollrung, M.: 513 = cord. —
Holstvoogd, C.: 178 = sti. — Hoogland, R. D.: 4540 & 4541
= ner.; 5042 = coral.; 5168 = ner. — Horne, J.: 152 & 384 =
vit. — Hose, C.:'5 = sti! -S0Hosokaway 77197595 *=cdap
Huitema, W. K.: 54 = sti: 115 = sin. — Hunt, P. F.: RSS.
2651 & 2906 = lat.; 2919 = lon. — Hurlimann, H.: 156 &
614 = har. — Hutchinson, W. J.: 3925 = cra.

Iboet: 43, 169 & 228 = sti. — Ilias-Paie: S. 16361 = sti. —
Iwanggin, G. Th.: BW. 9077 = ter.

Jacobs, M.: 4454 & 4808 = sti.; 4810 = mic. — Jansen, E.:
4660A = sin. — Jeoward, D.: 81 = vit. — Johnson & Constable:
NSW. 32026 = exc. — Junghuhn, F. W.: 160, 473 & 476 = sti.

Kairo, A.: NGF. 17445 = pel.-pel. — Kajewski, S. F.: 421 =
lat.;,.1943,. =. recip ck974.=ndeaqeo@d260>—. dat; 2582)5— nee.
KalshovennrLs, Grek: .6¢c= gpelppel.; a79n—— asin,: | 80 pee;
81 = pel.-pel. — Kanehira, R.: 5, 887, 1304, 1555, 1754, 1776,
1783 & 2218 = lat.; 2504 & 2551 = sti.; 4092 = lon. — Kanehira
& Hatusima: 11414 = coral.; 11908 = pel.-pel..; 13360 = coral.;
14211 = rac. — Karta: 12 = sin.; 14 & 19 = sti. — Kartodi-
hardjo: 2040 = sti.; 2062 = sin. — Kaudern, W.: 21 = sti. —
Keith: 289 = sti. — Kere, F.: BSIP. 4928 & 5008 = lat. —
Kern, J. Hi’ 8527°= sto] Ker, A. FG’ 150F-]"shn.~ aoa.
6422 & 6641 = sin.; 7811 & 11807 = sti; 12157 = sin.; 12553
= sti; 15590 = sin.; 16058, 17168, 18367 & 20799 = sti. —
Kiah: SFN. 23958 = sti. — King’s coll.: 5593, 7142 & 8565 =
sti. — Kjellberg, G.: 1041 = cel.; 2258 = kje.; 2345 = mic.; 2730
= amp. — Kloss, C. B.: SFN. 6829 & 18657 = sti. — Kochumen,
K. M.: 75989 = sti. — Koens, A. J.: 338 & 365 = sti. — Kondo
& Edano: PNH. 36806 = mey.-sub.; 39000 = sti. — Koorders,
S. H.: 3215 = sti. — 9467 = pel.-pel.; 9468 & 9469 = sin.; 9470
= pel.-pel.; 9471 = sin.; 9472 = pel.-pel.; 9473 = sti.; 9474 =
mic.; 9475 to 9484 = pel.-pel.; 9485, 9488, 9489 & 9490 = sin.;
9491 to 9500 ="sti.; 95017 =“sin; 9502 & 9503 ="stii> 95a =
sin | “OS03 0 207 = St. 9508 =" si. S509 a S10 St; oe
to 9573 "=", 9914 & 9515 = “stl, 9910 10.9920, = iii, ee
=i pel.-pel:' W0se3," 11007 & ‘TTIST “= "str, “11607 “Ac> Thea
sin.;’ 12372 2 Si. TOTS =* pelApe oizels =e. ee
14799) S30 Slee Stt.; 19309 tee = «6, ee
mic,; 20163" ="Stl-7" 20022 = pel.-per, 20js0.—— sie., aa
pel.-pel.; 20859 = sin.; 20953 = pel.-pel.; 21343 = mic.; 21344
=|, sin.:), 21345 «= -pelepel.; ; 213460 amic:; 21344,'=..) pel.-pen;
21349; = paic.; 21350)=.sin.; 21351.&:29352 = apel_-pel; 21354
= sin,; 21355 =, sti.; 22425 =.mic.; 22426 & 22666;—)pel--pel:
22773 .=,; sti; 23028,= pel.-pel.; 2323)1,c& 233991. sti; (23706

W.—L. Chew — Monograph of Dendrocnide 101

& 23760 = pel.-pel.; 24624 & 25769 = sti.; 26102 = sin.; 28020:
= sti.; 28026, 28832 & 29442 = pel.-pel.; 29445 = sti.; 29454,
29456 & 29819 = pel.-pel.; 30029 & 30148 = mic.; 30884 =
sin.; 33667, 34197, 34702, 35773 & 36240 = sti.; 36782 = sin.;
38338 = mic.; 38340, 38352, 38368 & 38369 = pel.-pel.; 38452
& 38478 = sin.; 38492 & 38498 = pel.-pel.; 38554 = mic.; 38819
& 38820 = pel.-pel.; 38849 = sin.; 39333 = sti.; 39906 = sin.;
oe) ==. pel.-pel.; 39926.— sin.; 39927 =mic.; 39962.= sin.;
40042 = mic.; 40057, 40062 & 40066 = pel.-pel.; 40256 & 40459
= sti; 41022 = pel.-pel.; 41910, 41974, 43979, 44081, 44121,
44266, 44559 to 44561 = sti. — Koster, Ch.; BW. 317 = ter.;
320, 7097, 7138 & 7177 = pel.-pel.; 10817 = coral. — Koster-
mans, A.; 59| = rec.; 1346.& 1678, = lon.;'2723 & 2915 = coral.;
4877 = ell.; 5338, 6206, 6238, 13454 & 13855 = sti.; 18359 =
mic.; 21302 = ell.; 21521 = sti.; 21756 = obl. — Kostermans et
al; KKSS. 125 & 127 = sti:;,129 = car..— Kunstler, H.: 4027
= sti. — Kuntz, R. E.: 063 = mey.-mey. — Kuntze, O.: 4911 &
wid, =: sti.;, 6983, =. sin.

Peace, J. H.2"5555 *=" Sin) —*Lajangah, ‘J. “K's SAN) 33122 &
aes == Si amy rd; SU0T —"'tla.. 3025 = lon? FIT) =
sch. — Lane-Poole, C.: 633 = coral. — Langlasse, E.: 58 = cra.;
94 = sub. — Lanjouw, L. R.: 184 = sti. — Larsen, K.: 9420 =
bas, — Lan.s. K.°.28363. = sin. —- Lauterbach, C.: TI53"=
ter.; 2554 = coral. — Lei, C. I: 584 = sti. — Leichardt: 26 =
pho. — Lesger, B.: 150 = sin. — Lipageto, Z.: BSIP. 3363,
3447 & 3466 = lat. — Loher, A.: 5008 = mey.-mey.; 5009 =
mey.-sub.; 6892 = mey.-mey.; 6933 = sub. — Lorzing, J. A.:
521 & 649 = sin.; 795, 1711 & 2887 = sti.; 3805 = sin.; 4198
= sti.; 4213 = sin.; 4298 = sti.; 4654 = sin.; 5398 & 14946 =
sti. — Lorzing & Jochems: 7452 = sti. — Ludlow et al.; 7282.
= sin. — Lutjeharms, W. J.: 3603 & 4941 = sti.

Maenu’u, L.: BSIP. 6073 & 6461 = lat. — Maingkom: bb.
7540 = mic. — Mair, K.: NGF. 1861 = sch. — Martellino &
Edano: 35387 = mey.-sub.; 35732 & 35760 = sti. — McClure,
F, A.: 7747 = sin. — McGregor, R. C.: 11256 = mey.-mey.;
19765;= ven.; 22906..= sub..— McKee,.H. S.: 3014 = vit. +
Mearns & Hutchinson: 4586 = den. — Meebold, A.: 5843 &
13011 = sin.; 16512 = har. — Meijer, W.: 2109, 5811 & 5812
= sti; 26442 = ell.; 29242 = sti.; 49826 = obl. — Mendoza,
D.: PNH. 18555 = mey.-mey. — Mendoza & Convocar: PNH.
10653 = mey.-sub. — Mendoza & Steiner: PNH. 41368 = luz.-
luz. — Merrill, E. D.: 988 & 998 = mey.-mey.; 1818 & 4838 =
sti.; 6290 = sub.; 8148 = sti.; 11649 = ven.; Ph. Pl. 294 = ©
sub.; Sp. Bl. 401 = mey.-mey. — Merritt, M. L.: 11379 = sti.
— Meyer, R.: 2631 = luz.-luz.; 2850 = sti. — Mikil, G.: 35755
= obl.; 41833, 46720 & 56298 = sti. — Millar, A. N.: NGF.
9288 = lon.; 14510 = ner. — Moran, R.: 4454 & 4535 = lat. —
Motley, J.: 346 = sti. — Mousset, J. P.: 280 = sti. — Muin-Chai:
Sees 15586, = ‘ell,;.31633. =» sti.

102 Gardens’ Bulletin, Singapore — X XV _ (1969)

Native coll.; 51 = sti. — Neth. Ind. For. Ser.: bb. 27110 =
car. — Nielsen, E.: 1159 = sti. — Nieuwenhuis, A. W.: 293 =
sti. — Nur, Md.: -7201'= sti.;°7408 =sin!; 41193’ ="stu

Parks, “H. (E./ "T6002, "=" Nar,, 20991 =" Vil, -—- "Feary, 1 ee
870A = sin. — Persieh: 257 & 492 = moroi. — Petelot, A::
6689 = ure. — Pierre, L.: 4708 = sin. — Pijl: 435 = pel.-pel. —
Pleyte, D. R.: 105 = lon. — Poilane: 8930 & 12505 = ure. —
Popta, S.: 1540 = sti. — Posthumus: 925 = sti. — Powell, Th.:
205 & 329 = har. — Prain’s coll.; 367 & 756 = sin. — Pringgo-
Atmodjo, R. M.: 407 = sin.; 411 = sti.; 487 = sin. — Puasa:
7662 = ell. — Pulle: 3098 = sti. — Pullen, R.: 526 & 569 =
morob.; 1078 = sch.; 1219 & 1706 = ter.; 1784 = cord.; 4065
= ORG

Quisumbing, E.: 7995 = mey.-mey.

Rahim Ismail: 98867 = sti. — Rahmat-si-Toroes: 1229, 3222,
4638 & 4941 = sti.; 5309 = sin.; 6925 & 7851 = sti. — Ramos,
M.: 438 = mey.-mey.; 1247 = obl.; 12142 = mey.-mey.; 14677
= mey.-sub.;.’17559 = ,sti.; 19462 =. ven.; 24061 = Juz.-luz.;
24274 & 24406 = mey.-sub.; 30229 = sub.; 33001 = mey.-sub.;
33175 = luz.-luz.; 43180 = sti.; 80306, 80461 & 80591 = mey.-
sub. — Ramos & Edano: 26362 = mey.-sub.; 26493 = mey.-
mey.; 30876 = sub.; 31101 = mey.-sub.; 36649 = sti.; 44364
= mey.-sub.; 49080 = mey.-mey.; 49285 = sti.; 49361 = cra.;
49413 = sti.; 75581 = mey.-sub.; 84935, 84936 & 84941 = sti.
— Rant, A.: 847 = lon. — Reillo, J.: 16446 = den. — Reinecke:
Sl) = vit 96 har. oe ie Reinwardt, C.: 539 = amp.
— Rensch, I.: 313 = car.; 704 = mic. — Richards, J.: NSW.
77929 = moroi. — Richards, P. W.: 1088 = sti. — Ridley, H. N.:
157 =~ pel-“mur.:- 158/ 159 °Sc "160 = "sin $8527 1494 SS
— Robinson, C. B.: 9731 = sub.; Pl. R. Amb. 310 = lon.; 311,
312 & 313 = amp* Rel? Robi 1920 =" ampi*== Reck,- #4
2040 = bas. — Rodway, F. A.: 1803 & NSW. 75174 = exc. —
Royen, P.: NGF. 20067 = lon. — Rudolph: 19 = cord. — Rutten:
427 =) tee 1718, = amps -1399.— ster.

Saanan (exp. Hulstyn): 103 = amp. — Sablaya, M.: 20 =
mey.-sub. — Saint-John, H.: 18228 = har.; 18255 = vit. —
Sandkuhl, H.: 298 = ven. — Santos, J. V.: 4156 = sti.; 4889
= mey.-mey.; 5232 = mey.-sub. — Santos, P.: 4156 = sti. —
Sarasin, F.: 466 & 814 = Jat. — Sarip: 108 = pel.-pel.; 133 =
sin. — Saunders, J. C.: 213, 323 & 385 = ter.; 462 & 557 =
ner. — Sayers, C. D.: NGF. 19509 = sch. — Scheffer, R. H.: C.
17 = sti. — Schlechter, R.: 13840 = coral.; 14385 = tor.; 16615
= sch.; 16885 = ner.; 16894 = coral.; 18377 = ner. — Schmid;
M.: 318 to 320 = lat. — Schram, F. A. W.: 2784 = ter.; 7858
& 7892 = pel.-pel. — Scortechini: 643 = sti; 1817 = sin. —
Seemann, B.: 426 = har.; 427 = vit. — Shanesy, J. O.: 24 =
pho. — Sidek b. Kiah: S. 36 = obl. — Sijde, H. A.: BW. 5534
= coral. — Sinclair, J.: 10727 = sti. — Sinclair & Edano: 9605
= sti. — Sinclair et al.: 9253 = ell. — Singh, J.: SAN. 28271

W.—L. Chew — Monograph of Dendrocnide 103:

= obl. — Singh et al.: 48869 = sti. — Sinnangul, H. T.: SAN.
39973, 56172, 57262, 57314 & 57334 = sti. — Smith, A. C.: 45
= thar; 359,985 .& 1595,=—. wit.;,4428 = whar.; 4450, 5033 &
2) a= ile 262 = har. S42 =_vit,; 5626 =.har.; 6313 =
Vit.;:722). = har.; 7835, 8315. é& 8658. .= vit. — Smith, E.:- 181
= sti..— Smith, L. S.;-10132, = cord.; 10175. = moroi; 10540
= exc. — Smitinand, T.: 1046 & 5257 = sti. — Snan: 669 =
sti. — Soegandiredjo: 79 & 174 = sti. — Soegeng-Reksodihardjo,
W.: 171 = sin. — Sow et al.: A. 1682 = obl. — Steenis, C. G. G.
Pere et St LOG0 = pel-pel." 25/5 35192" sti’; S297 -&
3431 = sin.; 7691 = mic.; 8088, 8304, 9433 & 10830 = sti. —
Stern, W. L.: 2175 = sti. — Stone, B. C.: 2387, 4138 & 5096
=Jaty — Story; R:: 7174-= exc. — Streimann &, Kairo: NGF.
21174 & 21177 = ner. — Strugnell, E. J.: 12670 = sti. — Sulit,
M. D.: PNH. 6924 = sti.; 8363 = mey.-mey.; 9864 = sti.; 11692
= mey.-mey. — Sulit & Conklin: PNH. 16861 = mey.-mey. —
Swain: 202 = pho. — Swinhoe, R.: 37 = mey.-mey. — Syming-
ton, C. F.: 22821, 22972 & 23013 = sti. — Synge, P. M.: 1088
='su:

Tabot de Matalban: 157 = sti. — Tabualewa, A.: 15640 =
vit. — Tanaka, T.: 10406 = mey.-sub. — Teo, L. E.: TP. 49
= sti. — Teona, R.: BSIP. 4861 & 6304 = lat. — Teysmann:
On? pra omne 3/5185 e—oter.375939°0— mic; 6756 = coral; /1171lt &
13794 = mic.; 16371 = sin. — Thorel: 3487 = sti. — Thorenaar,
A.: 70 = sti. — Thozet, A.: 408 = pho. — Thwaites: 2200 =
sin. — Tjokrosoendjojo: 1662 = sti. — Tothill, J. D.: 785 =
har. — Toxopeus, L. J.: 248 = mic.; 550 = sti. — Treutler: 802
= sin. — Tsang, W. T.: 17487 = sti.

Uliee: $,= tmic.esiibimesco: 73..=. sin;

Vaupel: 295 = vit. — Verhoef: 32 = pel.-pel. — Verreaux:
559 = exc. — Versteeg, G. M.: 1401 & 1547 = ter. — Versteegh,
Chr.: BW. 7519 = pel.-pel. — Vesterdal, A.: 22 = sti. — Vickery,
or Naw. 77918 to 77920 = exc.: 77931, 779348 7294 = pho.
— Vidal: 3879 & 3880 = sti.; 3881 = mey.-mey.; 3884 & 3885
= luz.-luz. — Vieillard: 2161 = lat. — Volck & Dansie: 1470
= cord.; 1471 = moroi. — de Vriese: 6 = pel.-pel.; 40 & 153
=, "MIC,

Wagiman: 4087 = sti. — Walangitang: 57 = mic, — Wall.
Cat.: 4611 = sin. — Walsh, M. E.: 84 = moroi. — Wang, C. W.:
78665 & 79201 =°bas:; 79495 = sin.; 79935 = bas.; 80006 =
ure.; 80794 & 81058 = sin. — Warburg, O.: 14702 = mey.-sub.;
Pore = su. —-* Waternouse, J. TH Ls °Y.'155'. =‘ rec.; 322 =
lon. — Waturandang, A. G.: 2, 187, 406 & 492 = mic. — White,
ot eas ee. ae = er.’ SoU = pho.; 6059, 9145 .&
cae) -—— Onc, taico = ple, 12728 &°12/8l = exc., 12995 =
pho. — White et al.: NGF. 1530 & 1655 = ner.; 1662 = cord.;
1675 = lon. — White, K. J.: NGF. 10477 = ner. — Whitmee,
S. J.: 271 = har. —- Whitmore, T. C.: BSIP. 798 & 2288 = rec.;
2857, 2863, 2956, 4752 & 5409 = lat.; 5684 = ner.; FRI. 3399

104 Gardens’ Bulletin, Singapore— XXV (1969)

= sti.; RSS. 6048 & 6130 = lat. — Whitmore & Womersley:
BSIP. 709 = rec. — Wight, R.: 459 = sin. — Williams, J. B.:
J. 41 = pho. — Williams, R. S.: 991 = rig.; 1548 & 2058 =
mey.-mey,;..2706 =, Juz.-ana.: “2ol0- _sti. Wilson, E. H.:
11015 = mey.-sub. — Winckel, W. F.: 1610 = sti. — Wind, R.:
2510 = sti. — Winkler, Hub.: 2983, 3330 & 3352 = sti.; 3478
= ell. — Wirawan, N.: 34 & 326 = sti. — Witkamp, H.: 1 &
2) =) sth: 3,.=.510,; 3.= Stow Womersiey. 2 &.. NGF. S0lg
ner.; 13383 = lon.; 19422 = ner.; 24558 = morob. — Womersley
-& Royen: 5896 = morob. — Wood, D. D.: 676 = obl. — Wood,
G. H. S.: SAN. 17168 = ell. — Wullen: bb. 14136 = mic.

Yates, H.’ S:: 603, 1029; 1360'& 2526 = sti) 25532 = ve
Yuncker, "T. G.:' 9927 & 15329" = War.

Zain, M.: 61307 = sti. — Zollinger: 510 & 1312 = sti.; 3007
= sin.; 3136 = pel.-pel.; 3807 = sin. — Zwickery, A. L.: 547
<= icra!

Index to Dendrocnide species

kajewskii, 94. stimulans, 11.

. kjellbergii, 86.

. latifolia, 71.

. longifolia, 79.

. luzonensis, 40.

. Meyeniana, 21.
. microstigma, 82.

subclausa, 28.

. ternatensis, 47.

. torricellensis, 51.
. urentissima, 32.
. venosa, 30.

D. amplissima, 84. CLC. mirabilis, 90.

D. basirotunda, 34. D. morobensis, 67.
D. carriana, 45. LC. moroides, 55.
D. celebica, 92. D. nervosa. 69.

D. contracta, 95. D. oblanceolata, 14.
D. corallodesme, 53. D. peltata, 42.

D. conden, JS: D. photinophylla, 60.
D. ciassmial 18. D. rechingeri, 74.
eae Taeiditolia ta
ee D. schlechteri, 76
D. excelsa, 64. ; ke =o
Deusipadiza. 95) D. sessiliflora, 95.
D: haiveyi, 88. D. sinuata, 36.

D. D.

D D.

D D

D D

CD D

D D

Cc D

. vitiensis, 62.

A new genus and new species of Malesian Orchids:

by

R. E. HOLTTUM

Recently I have received specimens and photographs of orchids
from Mr. K. C. Cheang (Penang) and Mr. A. G. Alphonso:
(Singapore). These indicate that corrections should be made to
some statements in my book Orchids of Malaya (3rd edition, 1965),.
and involve the recognition of a new genus and a new species.
From Singapore I have also received specimens of a further new
species originating in Sabah. The new genus and new species are:
described in the present paper, with appropriate name-changes.

Smitinandia, a new genus of Sarcanthine Orchids

Smitinandia, genus novum, habitu. Sarcantho Lindl. simile;
inflorescentia horizontale vel nutans, simplex vel ramosa; flores
parvi, carnosi, diu persistentes; labellum calcaratum, trilobum,
lobis lateralibus parvis, carnosis, lobo intermedio fere quadran-.
gulari, carnoso, antice tenui et rotundato, ore calcaris callo e basi
lobi intermedii nascente occluso; pollinia 4, in paribus valde
inaequalibus disposita, stipite brevi sursum dilatato suffulta.

Species typica: Saccolabium micranthum Lindl.

In Orchids of Malaya (\st ed. p.731, 3rd ed. p.735) I included
Saccolabium micranthum Lindl. with doubt in the genus Ascocen-
trum Schltr, remarking that I knew of no other genus to which it
could be referred. In recent years several plants of this species have
been sent to Kew from Thailand and have flowered; I have also:
received flowering material in alcohol, and photographs, from
plants in cultivation at the Botanic Gardens at Singapore and
Penang.

This new material shows the pollinia clearly. They are not, as I
had supposed, like those of Ascocentrum, which has two pollinia,
each with a small cleft; the species micranthum has four pollinia,
in two pairs, the two in each pair very unequal, as they are in
Sarcanthus and Thrixspermum. Viewed from the front, the smaller
members of each pair are hidden, and so are not shown in
Pantling’s drawing, on which I largely based my description, as
I had no fresh material when I prepared the book. The shape
of the labellum differs greatly from that of Sarcanthus, and the
callus which closes the entrance to the spur is an outgrowth from
the base of the midlobe of the lip, not from the back of the spur
as in Sarcanthus. As pointed out by Seidenfaden and Smitinand,
Saccolabium humile Ridl., agrees exactly in floral structure with
S. micranthum, and I unite the two species in a new genus, as these
authors suggest, naming it in recognition of their excellent work on
the orchids of Thailand.

106 Gardens’ Bulletin, Singapore— XXV (1969)

Key to the species of Smitinandia

Inflorescence unbranched, scape much shorter than rachis; dorsal
sepal 3 mm. long. S. micrantha

Inficrescence branched, scape as long as one of the branches:
dorsal sepal hardly 2 mm. long. S. humilis

Smitinandia micrantha (Lindl.) Holttum, com nov. — Saccola-
-bium micranthum Lindl., Gen. et Sp. Orch. (1833) 220. — S. fissum
Ridl., Journ. Linn. Soc. Bot. 32 (1896) 361; Flora Mal. Penin. 4
(1924) 173. — Cleisostoma micranihum (Lindl.) King & Pantl.,
Ann. R. Bot. Gard. Calcutta 8 (1898) 234, t.312. — Ascocentrum
micranthum (Lindl.) Holtt., Rev. Fl. Malaya 1 (1953) 731; Seiden-
faden & Smitinand, Orch. Thailand pt. IV, 1 (1963) 595, fig. 446. —
Cleisostoma tixieri Guillaum., Bull. Mus. Hist. Nat. Paris, ser. 2,
32 (1959) 369; Seidenfaden & Smitinand, Orch. Thailand Pt. IV,
2 (1964) 826.

Type specimen: Wallich 7300, Sylhet and Nepal (K).

Distribution: Sikkim Himalaya southwards through Tenasserim,
Laos, Cambodia and Thailand to Langkawi and Kedah.

As noted in Holttum 1953, the distribution of purple markings
in the flowers varies somewhat, those from northern Malaya having
least purple, but the shape of the flowers varies little. The scape of
the inflorescence is often very short (under 1 cm.) and is much
thicker than that of S. humilis. Old inflorescences bearing fruits are
sometimes longer than the leaves, but usually they do not exceed
the leaves.

Smitinandia humilis (Ridl.) Holttum, comb. nov. — Saccolabium
humile Ridl., Journ. Str. Br. R. Asiatic Soc. no. 59 (1911) 198:
Seidenfaden & Smitinand, Orch. Thailand pt. IV, 1 (1963) 616,
617, fig. 465 A, B.

Type specimen: Curtis 3028, Poongah (Phang-nga), Thailand
(S. dupl. K).

Distribution: Eastern and Peninsular Thailand, Laos.

Seidenfaden and Smitinand publish good illustrations, and (p.
616) point out the relationship of this species to S. micrantha. I
have not seen a specimen of Cleisostoma poilanei Gagnep. (Bull.
Soc. Bot. France 79: 34, 1932) but from the description it seems
likely to be synonymous with S. humilis, rather than with S.
micrantha as suggested by Seidenfaden and Smitinand (p. 595).

A new species of Hylophila Lindley

Hylophila cheangii Holttum, sp. nov. — AH. lanceolata sensu
Holttum, Rev. Fl. Malaya 1 (1953) 119, p.p.

Hylophila habitu H. lanceolata (BI.) Miquel similis, sed ab ea
inflorescentia omnino hirsuta atque aurantiaca, appendiculis
stigmatis nullis, distincta.

Stem 25 cm or more long, basal part creeping and rooting at the
nodes, internodes 2.5-3.5 cm long: leaf-blade 9-11 x 2.5-3.0 cm,
widest in the middle, narrowed to base and slightly acuminate apex,

Holttum — Malesian Orchids 107

sheath and petiole 2.5 cm long (closed sheath 8 mm). Inflorescence
terminal, scape to 3 cm; rachis 2.5-4.5 cm long, hairy, reddish
orange: bracts 12 mm. long, narrowed to a slender tip. hairy as
rachis and of same colour; dorsal sepal orange, 10 mm long, 4 mm
wide near base when flattened, gradually narrowed to apex, hairy
on outer surface; lateral sepals 10 mm long, 5 mm wide near
base; petals as long as sepals, 3 mm wide, widest in middle, hair-
less, clear yellow; labellum saccate, 6 mm long, orange yellow,
its base joined to the sides of the column, with a very small
aperture in front of the column, its blade very narrow, 5 mm long.
a deeper orange; column less than 2 mm long, bearing a distinctly
bilobed swollen stigma in an arc across its front, the two halves
together forming a crescent with ascending arms just beneath the
rostellum; anther elongate and bent forwards almost at right
angles to the column; pollinarium 7 mm long, the narrow viscisium
5 mm long; rostellum deeply lobed.

Type: coll. K. C. Cheang, s.n., terrestrial by streamside, Fraser’s
Hill, Pahang, 27 June 1968 (K). The following specimen in Herb.
Kew also agrees: Moulton 6694, Gunong Temabok, Upper Baram
Valley, Sarawak, 3,000 ft. (distributed as Dicerostylis lanceolata
BI.).

I believe Mr. Cheang’s specimens to be identical with those few
others from Malaya which have passed as Hylophila lanceolata
and are so described in my book on the orchids of Malaya. When
preparing my description for the book, I saw no fresh material, and
accepted J. D. Hooker’s identification of Malayan specimens with
Blume’s Java species (see Hook. fil. in Fl. Brit. India 6: 110). I
took details of colour from J. J. Smith’s revised description of the
Java species, made from fresh specimens in 1921. Ridley’s previous
description of the same old Malayan specimens available to me
(he never found any himself) included the statement “flowers
white”, which he probably copied from Blume. My description, as
regards dimensions of flowers and their parts, was evidently taken
from dried Malayan specimens, and agrees well with the new
specimen found by Mr. Cheang. The only discrepancy is the
absence of stigmatic appendages on Mr. Cheang’s specimen; I
suspect that I took these on trust, copying Hooker’s statement.
Hooker saw only a single specimen, collected in Perak by Scor-
techini about 1885, and the Kew sheet bears sketches of floral
details, made by Hooker. The sketches show two appendages at
the base of the column, but comparison with the floral fragments
left after Hooker’s dissection inclines me to think they may have
been parts of the lip remaining attached to the column. I tried
to dissect two further flowers-buds from Scortechini’s specimen, but
both had been so damaged by an insect that details could not
be seen. Whatever may be the truth about this Scortechini speci-
men, I have no doubt that Mr. Cheang’s is quite distinct from
Blume’s species Dicerostylis lanceolata which was illustrated by
him in Coll. Orch. (1859) 116, pl. 38, fig. 1.

108 Gardens’ Bulletin, Singapore — XXV (1969)

So far as I know, the only other record of this species from
Malaya is a specimen (in Herb. SING.) collected by Curtis on
Bujong Malacca in the 1890s. Thus Mr. Cheang’s specimen is the
first known to have been found for about 70 years.

It may be that Hylophila and Dicerostylis should be distinguish-
-ed by the shape of the stigma itself rather than by the two appen-
dages on the coiumn which are not part of the stigma and which
were regarded as distinctive of Dicerostylis by Blume. The type
species of Hylophila is H. mollis Lindl. When preparing my book
I saw fresh material of this, and described the stigma as “large,
convex”; I have tried to check this from dried specimens but
find it very difficult to see details clearly on them. If a bilobed
stigma is regarded as characteristic of Dicerostylis and a simple
‘one of Hylophila, then H. cheangii would belong to Dicerostylis;
but in view of the close similarity of all other parts of the flower
I think it not unnatura in any case to unite the two genera, as was
‘done by Miquel.

A new species of Trichoglottis from Sabah.

Trichoglottis appendiculifera Holttum, sp. nov.

Caulis ad 40 cm vel ultra longus; folia 4-6 cm longa, 11-16 mm
lata, apice sat profunde inaeaualiter biloba, lobis rotundatis, costis
subtus (praesertim basin versus) carinatis, carina in vaginam decur-
rente et illic alam 1 mm latam translucentem efficiente, varginis
leviter imbricatis. Inflorescentia per vaginam erumpens, uniflora.
‘Flores pallide viridelutei, sepalis petalisque fasciis transversalibus
‘subro-aurantiacis irregulariter ornatis. Sepala petalaque 13 mm
longa, sepalum dorsale 34} mm, sepala lateralia 44 mm, petala 24
mm lata. Labellum 9 mm longum; lobi laterales pallide brunnei,
erecti, fere oblongi basi 24 mm longi, 14 mm alti, intus breviter
pilosi; lobus intermedius labus, striis pallide violaceis ornatus, 64
mm longus, e basi sensim ampliatus, ambitu anguste triangularis
desuper visus, carina tenui alta antice abrupta declivi praeditus et
supra apicem appendiculam angustam tenue fere 2 mm longam
oblique ascendente gerens, carina omnino pilis horizontaliter paten-
tibus ac basi pilis ascendentibus ornata. Gynostemium 3 mm altum,
minute papillosum; pes gynostemii vix 2 mm longus; ligula as basin
-gynostemii enata 2 mm longa, angusta, apicem leviter bilobatam
versus ampliata, marginibus ciliata.

Type: cult. Hort. Bot. Singap., origin Sook Plain, Sabah, coll.

A. G. Alphonso, no 1/110/66 (Singapore Introd.), specimen in
Herb. Kew.

The peculiar complex structure of the labellum of this species
:seems to be auite distinctive.

Calostoma sarasinii from Malaya

by
G. LIM

Botany Department,
University of Singapore,
Singapore

The genus Calostoma Desv. comprises a small group of puff-
balls (Gasteromycetes) that are not familiar to many. Although the
genus is widely distributed, the species may be divided into two
natural groups characterised by the form of the spores: an eastern
group with globose spores, and a western group with elliptical
‘spores (Massee, 1888-89). Massee (1888-89) distinguished these
species with their areas of distribution as follows — six eastern
species: C. berkeleyi Massee, Ceylon, C. insignis (Berk.) Massee,
Ceylon, C. junghuhnii (Schlechtend. et Miiller) Massee, Ceylon,
Java, Sikkim Himalayas, Sumatra, C. orirubra Cooke (orirubrum),
Larut in Perak, Malaya Peninsula, C. pachystelis (Ces.) Massee,
Borneo, Ceylon, and C. viridis (Berk.) Massee, Sikkim Himalayas,
Sinchal, Tonglo; four western species: C. cinnabarinum Desv.,
Massachusetts to Carolina, New Granada, C. fusca (Berk.) Massee,
S. Australia, Tasmania, C. lurida (Berk.) Massee, Australia, and
C. ravenelii (Berk.) Massee, mountains of Carolina. Burnap (1897)
and Webster (1899) described three species found in America, C.
cinnabarinum, C. lutescens (Schw.) Burnap and C. ravenelii, and
of these, C. lutescens is the only western species described as
having globose spores.

Boedijn (1938) who studied the Malaysian materials in the
herbarium at Buitenzorg, distinguished six species: C. berkeleyi, C.
insignis, C. junghuhnii, C. orirubra, C. retisporum Boedijn and C.
sarasinii (Henn.) van Overeem. These were recorded from the
following regions — Borneo, Celebes, Java, Moluccas, New Guinea
and Sumatra. All six species posses globose spores.

Thus, among the species mentioned in literature, only C. oriru-
bra was recorded from the Malay Peninsula and was included in
Saccardo’s compilations (1891). The description given for this
species does not fit the specimens collected recently from Max-
well’s Hills, Perak, Malaya. However, the collection in external
morphology and spore size and form matches closely the excellent
account and drawings given by Boedijn (1938) for C. sarasinii.
The only slight difference lies in the spore warts which were
described by Boedijn as ‘strongly truncate at the apex’ but which
in the recently found specimens, did not appear quite as truncate.
Since in all other details the fungus resembles the material studied
by Boedijn, it is suggested that the recent Malayan collection
belongs to C. sarasinii. This is a new record of the species for
Malaya.

110 Gardens’ Bulletin, Singapore — X XV (1969)

The specimens were found growing on the ground in primary
jungle, at 4,000 ft. altitude, in Maxwell’s Hills, Perak, in August
1967. Nine fruiting bodies of varying ages and sizes, occurred in a
clump. This appears to be an uncommon fungus and has not been
observed again in spite of several return visits.

It is of interest that with this report, only two species of
Calostoma are thus recorded for Malaya and both were collected
from the same district, Larut, in the State of Perak.

REFERENCES

BoEDUN, K. B. (1938). The genus Calostoma in the Netherlands
Indies. Bul. du Jardin Bot. de Buitenzorg. ser. III. 16, 64—75.

BurnaP, C. E. (1897). Note on the genus Calostoma. Bot. Gaz.
23, 180-192.

MASSEE, G. (1888-89). A monograph of the genus Calostoma
Desv. (Miiremyces Nees). Ann. Bot. 2, 25-45.

SACCARDO, P. A. (1891). Sylloge Fungorum. 9, 270-271.
WEBSTER, H. (1899). Notes on Calostoma. Rhodora. 1, 30-33.

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A Monograph of Laportea (Urticaceae)

by
W.-L. CHEW (CHEW WEE-LEK)

Laportea was large and mixed. It defied definition and could
only be identified by a system of elimination.

Now, with the ligneous species removed and Fleurya and
Sceptrocnide added to it, it is more natural, and can be defined
thus: (a) herbs, rarely shrubs, with irritant hairs, (b) leaves
petiolate, simple, alternate: lamina generally chartaceous with
margin variously toothed; (c) stipules partially connate, bifid at the
apex, intrapetiolar, (d) flowers unisexual, 4- or 5-merous, free or
in loose glomerules, perianth not fleshy at maturity, (e) female
pedicels winged, (f) achenes usually reflexed, often stipitate and
sometimes on small gynophores, with ligulate stigmas, dispersed
usually with the perianth attached.

A total of 22 species are accounted for in this genus which has
two sections: sect. Laportea and sect. Fleurya. The former section,
which contains 10 species, distinguishes itself from the latter,
with 12 species, in the female pedicels being winged laterally.
Essentially African, this genus of weedy herbs has a few species
almost pan-tropical in distribution while two others have extremely
wide ranges of climatic tolerance.

This work is based on the materials of the following herbaria
supplemented by some field studies in South-East Asia:

Amsterdam, Holland (AMD). Lisboa, Portugal (LISC, LISJC &

Arnold Arboretum, U.S.A. (A).
Bangkok, Thailand (BKF).
Berkeley, California (UC).
Berlin, Germany (B).
Bogor, Indonesia (BO).
Brisbane, Australia (BRI).
British Museum, U.K. (BM).
Brussels, Belgium (BR).
Cambridge, U.K. (CGE).
Cape Town, S. Africa (NBG &
SAM).
Chicago, U.S.A. (F).

Claremont, U.S.A. (POM & RSA).

Copenhagen, Denmark (C).
Edinburgh, U.K. (E).
Florence, Italy (FI).
Geneva, Switzerland (G).
Kepong, Malaysia (KEP).:
Kew, U.K. (IK).

Kuching, Sarawak (SAR).
Lae, New Guinea (LAE).
Leiden, Holland (L).

LISU).
Lund, Sweden (LD).
Madrid, Spain (MA).

_Manila, Philippines (PNH).

Melbourne, Australia (MEL).
Michigan, U.S.A. (MICH).
Missouri, U.S.A. (MO).
Munich, Germany (M).
Noumea, N. Caledonia (NOU).
New York, U.S.A. (NY).
Paris, France (P).

Praha, Czechoslovakia (PRC).
Sandakan, Sabah (SAN).
Singapore (SING).

Stockholm, Sweden (S).
Sydney, Australia (NSW).
Uppsala, Sweden (UPS).
Utrecht, Holland (VU).
Washington D.C. U.S.A. (US).
Wrocklaw, Polland (WRSL).
Zurich, Switzerland (Z).

112 Gardens’ Bulletin, Singapore — XXV (1969)

I am grateful to the directors of these herbaria for their genero-
sity in making their valuable collections available for my work.
I am also grateful to the following botanists for their general
assistance at various times on many matters: Prof. E. J. H. Corner
(Cambridge), Dr. C. X. Furtado (Singapore), Prof. J. Leandri and
Prof. R. Letouzey (both of Paris), Prof. C. G. G. J. van Steenis
(Leiden) and Sir George Taylor (Kew).

My gratitude is also extended to the Royal Society of London
for their generosity in granting me a Royal Society — Nuffield
Foundation Commonwealth Bursary which enabled me in 1964
to visit. and consult the rich collections in the herbaria in London,
Paris, Geneva, Leiden and Utrecht.

Lastly, I would like to thank my wife, Sigrid, for her translations
of the French and German works consulted and for her general
assistance in the listing of the collections.

Part 1 — General

Habit. Most species are well branched annual herbs less than
2 metres high. The stem is usually weak and often woody only at
the base. L. ruderalis, L. cuneata and L. caffra are the smallest in
the genus. At the other end of the scale, we have L. decumana a
species of tall perennial shrubs with woody stems. Generally
erect, L. peduncularis and L. ovalifolia occasionally send out
repent shoots which are claimed to be capable of scrambling over
fairly great distances. Laportea therefore contrasts quite clearly
from Dendrocnide a genus of perennial woody trees up to 35 metres
high.

Leaves. Generally, they are simple, petiolate, dorsiventral, pen-
ninerved, chartaceous, spirally or alternately arranged and with
variously toothed margin. Overall, the similarity is with the leaves
of Urtica rather than the other genera of the family.

The lamina offers good characteristics for the discrimination
of the species. In some cases the form and dentition of the lamina
are very specific as in L. perrieri, which has teeth 7—20 cm. in
length, L. ruderalis whose very small lamina is completely glabrous
and devoid of irritant hairs and L. macrostachya with an ovate
lamina which is caudate-emarginate at the apex. In L. decumana
and L. floribunda, the lamina is bullate and densely covered with
erect irritant hairs, and the margins very finely denticulate. Only
in one species is the lamina polymorphic i.e. in L. cuneata where
its margin varies from almost entire to doubly serrated at different
stages of its life.

Petioles. Only the vestiture of these is used to separate some
species. In L. grossa and L. mooreana, they are covered with long
mounted irritant hairs. In others, these hairs are not mounted but
are either erect or appressed.

Stipules. These differ fundamentally from those of Dendrocnide.
Here they are intrapetiolar but only partially connate. As a result,
all are bifid at the apex. In L. ovalifolia the bifid portion is as
long as the lower connate part. Most species have them to about
1 cm. in length, but L. violacea has stipules less than 0.3 cm. in
length while in L. decumana they reach 2 cm.

W.-L. Chew — Monograph of Laportea ges

Inflorescences. They are more variable here than in Dendrocnide.
{In most species, they are unisexual, paniculate and fairly well
branched. The females are usually borne in the upper axils and
are often more widespread than the males. L. canadensis, L. bulbi-
fera, L. alatipes etc. illustrate this type of inflorescence well. In the.
Madagascar species as in L. amberana or L. perrieri this general
pattern is slightly modified: branching occurs mainly at the top
of the inflorescence and then weakly so. This reduction in the
lateral branches is greatest in the L. macrostachya — ovalifolia
group of species where they become somewhat elongated with
the flowers glomerulated at intervals along the seemingly un-
branched peduncle. Such ‘‘interrupted”’ inflorescences can be either
unisexual as in L. macrostachya or bisexual as in L. interrupta.
In L. caffra the inflorescence is very small, branched once at the
top and usually bears less than 10 flowers. The most distinctive
inflorescence is in L. cuneata. Here the males are apedunculate
and are borne at the base of the corymbose female inflorescences.

Male flowers. These are somewhat similar not only to those of
Dendrocnide but also to others in the tribe Urereae. They are
small, 4- or 5-merous, sessile or pedicellate. Stamens are equal in
number to the tepals and are always borne opposite to them on
rather long reflexed filaments. The pistillode is always present as
a thin often hyaline structure in the centre of the floral cavity.
The tepals are of two sorts — corniculate or non-corniculate.
This characteristic is extremely useful in the delimitation of the
species if it is considered together with the other characteristics
like number of tepals and presence or absence of the pedicel.

Female flowers. These are very important for the delimitation
not only of the species but also of the two sections as well as of
the genus as a whole. Basically they are 4-tepalled, with the lateral
tepals much larger than the dorsi-ventrals, and pedicellate. The
evary is simple, asymmetrically ovoid, compressed and has a
ligulate or somewhat linear stigma. Staminodes are absent. The
“wings” on the pedicels distinguish this genus from Dendrocnide.
Within the genus, the two sections differ from each other in the
nature of these “wings”. In section Laportea, the pedicels are
winged laterally and symmetrically; whereas in sect. Fleurya, they
are winged dorsi-ventrally and therefore somewhat asymmetrically.
On the specific level, the nature of the dorsal tepal and stigma is
found to be quite useful in distinguishing some species. L. inter-
rupta and L. ovalifolia for example, are the only species with
3-fid stigmas. In length, the stigma is shortest in L. cuneata and
ruderalis where it seldom exceeds (0.2 mm. and is consequently
brush-like. At the other extreme is L. decumana whose stigma
may reach 3-4 mm. The dorsal tepal may be greatly geniculate
as in L. mooreana, L. cuneata and many others or non-geniculate
as in L. ovalifolia in which case it is often only slightly shorter
than the lateral tepals. In all species, the ventral tepal tends to be
the smallest and in L. mooreana it is often absent.

114 Gardens’ Bulletin, Singapore —XXV (1969)

Achenes. These are small, ranging from | to 5 mm. in length
compressed, often greatly asymmetrical with the stigma displaced
to one side. In sect. Laportea, they are somewhat larger, always
stipitate and usually borne on gynophores. They are thus dis-
persed without the perianth. L. perrieri has the largest achenes
up to 5 mm. in length and 3-4 mm. across. L. disepala has the
shortest stipe being less than 0.2 mm. long. The longest pedicel
(at fruit) is found in L. amberana where it reaches 10 mm. In
shape, they are somewhat semi-circular in the canadensis — alatipes
group of species and variously ellipsoid, obovate and almost
-circular as in L. amberana.

In sect. Fleurya, on the other hand, the achenes are slightly
smaller, mostly sessile and not borne on gynophores. They are
usually dispersed with the perianth attached. Excepting L. ovali-
folia which has achenes to 4 mm. long, the rest of the species have
them up to 2 mm. in length. In the stipitate species, L. cuneata
and L. ruderalis, the stipes vary between 0.2 and 0.4 mm. In shape,
they are generally ovoid excepting L. floribunda where they are
obovoid. Despite the exceptions, the achene does provide good
characters for the discrimination of the two sections generally.

Distribution. This genus is Africa-Madagascar centred. Of a
total of 22 species, 12 are confined to this region (five being
endemics of Madagascar). The north American continent has one
species confined to that region i.e. L. canadensis which has been
recorded from as far south as Mexico. The West Indies, broadly
speaking, have also one species peculiar to these islands viz. L.
cuneata which has so far been recorded from Cuba and His-
paniola. Continental Asia has three species endemic in that region
viz. L. disepala, L. violacea and L. macrostachya. Malesia has one
endemic species viz. L. decumana and another which it shares with
the Pacific islands i.e. L. ruderalis.

The remaining 3 are about the widest distributed species in the
genus: these are L. aestuans, L. interrupta and L. bulbifera.
The first two are entirely tropical; and together they encircle the
tropics! The last, L. bulbifera, has about the greatest climatic
tolerance in the genus. It extends from the cold temperate regions
of Manchuria and North Korea through South-east Asia to the
hot tropics of Sumatra and Java, although in the latter countries
they prefer the highlands above the 1,000-metre contour.

Affinities. Also a member of the Urereae tribe, Laportea was
thought to be closest to Sceptrocnide, an Asiatic genus of herbs.
After further study, the latter has now been found to be actually
congeneric with the present genus. The phenomenon of the stamens
being adnate to the tepals in Sceptrocnide has now been found to
be an exception rather than a rule. Similarly, the stipules have been
observed to be also intrapetiolar and partially connate as in
Laportea and not free and lateral. This leaves Nanocnide (an
Asiatic genus of small herbs) as the nearest relative of Laporiea.
This genus is similar to Laportea in almost all respects except that
its achenes are erect (and not reflexed as in Laportea) and the
stipules free and lateral.

W.-L. Chew — Monograph of Laportea 115

Part 2 — Systematic

LAPORTEA, nom. cons.

Gaud. in Freyc. Voy. Monde Bot. 498 “1826” (1830). — Urti-
castrum Heist. ex Fabric. En. Pl. Helmst. 204 (1759), nom. rej.;
O.K. Rev. Gen. Pl. 2: 635 (1891), p. p. — Oblixilis Rafin. Fl. Tell.
3: 49 (1837). — Laportea sect. Sclepsion Wedd. in Arch. Mus.
Hist. Nat. Paris 9: 123 (1856) et in DC. Prodr. 16 (1): 78 (1869),
D. Pp.

Monoecious,. very rarely dioecious, herbs or shrubs with irritant
hairs. Leaves petiolate alternate, simple, chartaceous with variously
toothed margin. Stipules intra-petiolar, incompletely connate, bifid
at the apex. Inflorescences generally paniculate, pedunculate, axil-
lary, with unisexual flowers in loose glomerules; peduncular bracts
very small. Male flowers 4- or 5-tepalled, with an equal number
of stamens opposite the tepals; filaments reflexed; pistillode pre-
sent, small. Female flowers 4 tepalled, the lateral tepals equal-
sized, the dorsi-ventral ones greatly unequal-sized, much smaller
than the laterals, devoid of staminodes; ovary ovoid; style generally
linear, rarely very short & brush-like. Achenes generally com-
pressed, ovoid to semi-circular, sessile or stipitate, 1 to 4 mm.
long, usually reflexed upon either dorsi-ventrally or laterally winged
pedicels.

Distribution: Pan-Tropical.

Lectotype species: Laportea canadensis (L.) Wedd.

sect. Laportea

This section distinguishes itself by the pedicels of the female
flowers being winged laterally and symmetrically; and the achenes
articulated on pedicels.

Type species: Laportea canadensis. (.) Wedd.
Species are: nos. 1 to 10.

sect. Fleurya

(Gaud.) Chew in Gard. Bull. Sing. 21: 199 (1965). — Fleurya
Gaud. in Freyc. Voy. Monde, Bot. 497 ‘1826’ (1830). —
Schychowskia Endl. in Ann. Wien Mus. 1: 187 (1836). —
Fleuryopsis Opiz in Lotos 3: 240 (1853). — Sceptrocnide Maxim.
in Bull. Acad. Petersb. 22: 230 (1877).

This section differs from sect. Laportea in (a) pedicels of female
flowers winged dorsi-ventrally and asymmetrically, rarely simple,
(5) achenes not articulated on pedicels.

Lectotype species: Fleurya spicata Gaud. |= Laportea interrupta
(L.) Chew].

Species are: nos. 11 to 22.

116
l—a
b
2—a
b
3—a
b
4—a
b
5—a
b
(0. — 4
b
7J—a
b
8—a
db
9—a
b
10 —a.

Gardens’ Bulletin, Singapore —XXV (1969)

Key to the Species of Laportea

. Stigma 3-fid, the central branch longer than the lower two:

inflorescences appear unbranched due to great reduction of lateral
branchéstt.:,. Sa7.ja20.57.. AS, PRONE ONE + ONSET, Vie fs

. Stigma simple, without branches; inflorescences variable ......... 3

. Plant without stolons; inflorescences bisexual, axillary; stigmas

less than 1 m.m. long; male perianth with 4 corniculate tepals
(Central & East Africa to Australia & Pacific) ...... L. interrupta

. Plant with stolons; inflorescences unisexual, often produced from

stolons; stigmas 2-3 mm. long; male perianth with 5 non-
comiculate tepals (tropical Africa). ~.....,.-c<-..-.- L. ovalifolia

. Plant less than 60 cm. high, completely glabrous and quite without

irritant hairs; inflorescence paniculate, well branched, shorter than
leaves; male flowers 4-merous, tepals corniculate; stigma almost
brush-like, less than 0.2 mm. long (East Malesia & Pacific
Islands), 4.2 ).3..h. . Ceo Bete Clete ee eee L. ruderalis

Not this. combrmation) of. characters: <c200 20. do See, bees 4

. Inflorescences unisexual; the females corymbose, axillary; the

males apedunculate, at the base of the female inflorescence; small
herbs less than 60 cm. high dense with long hyaline irritant hairs;
stigma brush-like less than 0.2 mm. long. (Cuba & Hispaniola)

L. cuneata
+Net thiss combination of charasters)1io.ii0a4. col. iow.. tees 5
. Lamina rugose, densely covered with erect irritant hairs, with
lipelly denticulgte migrg@in ....5 0). 2.cr.. -.-<eeeeteaesy +: aoe eens oe 6
. Lamina smooth, with appressed irritant hairs, margin serrate or
dentate, sometimes coarsely to profoundly ©..)....2..0.2.2..0220". 7

. Lamina elliptic; male flowers sessile, 5-merous; stigma less than

0.5 mm. long; female pedicel not winged laterally (Madagascar)
L. floribunda

. Lamina ovate, rhombic to trullate; male flowers pedicellate,

4-merous; stigma 2 to 3 mm. long; female pedicel winged laterally
(Borneo: &' HastoMalesia) ¢.. :20)4,, 9 ea... oe L. decumana

. Lamina almost rotund, base cordate, margin very profoundly

dentate with few teeth each to at least 7 cm. long and 3 cm. broad
at the base; female inflorescences with very stout peduncle, much
longer than the leaves (Madagascar) ........................ L. perrieri

. Not this combination of Charactess!. 2)... ..acin..:a2e. See 8

. Teeth of lamina ca. 3 cm. long, 2 cm. broad at the base, the apex

of lamina often profoundly emarginate, with a long narrow cauda;
female inflorescences elongated, spike-like, mainly in the upper
axes (Jaoan, China; Buta). 2: 2.9:05.0..20s. Oe. 2s L. macrostachya

. Teeth of lamina smaller, the apex of lamina acuminate. sometimes

acute but not caudate-emarginate; if unisexual, female inflore-
seences not elongated’ and, Spike-like. |. .!....tig.2.-hAt een 9

. Teeth of 'amina small very numerous, not exceeding 0.5 cm. long;

anmial Nerds. ........ 04.» aes doee ss hac Dae ee 10

. Teeth of lamina larger, fewer in number, usually between 0.7 cm

& 2 cm. long; annual herbs or perennial shrubs ..................... 18

Femaie inflorescences 2 to 3 times longer than the leaves (inclusive
of petioles), with a stout elongated peduncle branched sparsely
towards the apex; margin of lamina irregularly toothed
(Niddeeaséar) ..i0..: 2... 2s..R0 See. eee L. septentrionalis

. Female inflorescences shorter to at most slightly longer than leaves,

well branched panicles; margin of lamina regularly toothed ... 11

11 —a.
b
42—a.

15 —a.

b

16 —a.

b

| 17 —a.

19 —a.

20 —a.

‘21 —a.

b.

. Not this combination of characters

W.-L. Chew — Monograph of Laportea 117

Herbs with glandular hairs (tropical America, Africa, India to
CS ee eee ee ee L. aestuans

Peres Witt MiAMGMar GNA coe. | aco en come wget a> oa sees mn- 12

Herbs less than 50 cm. high; inflorescences very small, frail,
branched once at the apex bearing very few flowers; lamina ovate
fess than 3 ‘cm-‘long (South Arica) el A e.. L. caffra

. Herbs more than 60 cm. high; inflorescences much larger, with

stout branches, bearing numerous flowers; lamina variously shaped,
ER TENN NINE ns a 3 ok Ge Ps ah pga vcnggig ses 13

Plant often creeping or scrambling; female pedicel winged dorsi-
ventrally, especially at fruit; achene sessile not borne on gyno-
hores, (East South-East & South Africa) ..........4 L. peduncularis

. Plant erect; female pedicel winged laterally; achene stipitate, borne

EN AACE ND I as car SLES isos bins Sc Aiashers deere as fers. SOR spy tdyiie o vy 14

. Male flowers with 4 non-corniculate tepals; achene ovoid, the

lateral sides depressed and warted (Thailand & Indo-China)
eT ek eee. 5 8 OAL eR A SY L. disepala

. Male flowers 5-tepalled, or if 4-tepalled, then tepals corniculate;

achene often semi-circular in shape, the lateral sides not warted,
ME VAR RE. TE Clea Whee, 8. OR Recs ete cee ede np ecee Bde ceese 15

Male flowers sessile to subsessile, with 5 non-corniculate tepals;
base of lamina truncate; achene warted at the sides (China,
Whratlaind Gemini) 1 Bao. dao dew ce ses cee ceeeeeeens L. violacea

D, Gen erential, - { lmorpetta’ Setegte: 16

Woody bulbils often present in the axils of leaves; male flowers
Wwith-S<non-cormculare tepals Gistay: 2 Pe kl). L. bulbifera

. Woody bulbils absent; male flowers with corniculate tepals ... 17

Male flowers with 5 tepals; achenes semicircular (North America)
Bs AS Ld a Ed id Mae ete OV IIIROTCL. | L. canadensis

. Male flowers with 4 tepals; achenes somewhat pyriform (Africa)

— ee P a S Ae: F a n ee e L. alatipes

Achenes completely covered by lateral tepals at maturity, very
slightly reflexed; the dorsi-ventral tepals equal-sized; female
pedicels not winged laterally; lamina narrowly ovate to elliptic
(Contra! .6c~ Hast.) Adsica) fics cen... vaeriesh. seein 2.8. L. lanceolata

. Achenes not completely covered by lateral tenals at maturity,

strongly reflexed; the dorsi-ventral tepals greatly unequai-sized;
female pedicels winged laterally; lamina triangular to ovate, rarely
UE a, Oa: en oe eee ee a eS 2 ee rane a 19

Lamina generally triangular, base truncate; petioles armed with

long mounted irritant hairs; female pedicels winged dorsi-
ventrally 20

. Lamina ovate to very broadly so; base rounded or profoundly

cordate; irritant hairs of petioles not mounted, female pedicels
I kn nan eeds
Inflorescences usually longer than leaves; male flowers with 4
corniculate tepals; lamina with marginal teeth ca. 1.5 cm. long,
0.5 cm. broad at the base: intercostals numerous, closeiy set,
rather prominent (East, Central & West Africa)... L. mooreana

. Inflorescences usually shorter than leaves; male flowers with 5

non-corniculate tepals; lamina with marginal teeth ca. 1 cm. long,
1 cm. broad at the base; intercostals few, anastomosing, not
prominent (South-East & South Africa) ................... L. grossa

Lamina very broadly ovate; base profoundly cordate; margin very
irregularly toothed (Madagascar) .....................0.. L. amberana

Lamina ovate; base rounded; margin rather regularly toothed
gaclneaee cet ng IP. egy septal l a da L. weddellii

118 Gardens’ Bulletin, Singapore — XXV (1969)

(1) Laportea canadensis (L.) Wedd. in Ann. Sci. Nat. Ser. 4. 1: 181
(1854), in Arch. Mus. Hist. Nat. Paris 9: 123. tab. 2c, fig. 24
(1856) & in DC. Prodr. 16 (1): 78 (1869). — Urtica canadensis
L. Sp. Pl. 985 (1753).'— U. divaricata L: op. cit. (1753). —
Fleurya canadensis (L.) Benth. in Hk. Niger. Fl. 517 (1849). —
U. pustulata Liebm. in Vidensk. Selsk. Skr. 5 (2): 294 (1851).
— L. pustulata (Liebm.) Wedd. in Arch. Mus. Hist. Nat. Paris
9: 140 (1856). — Urticastrum divaricatum (L.) O. K. Rev. Gen.
Pl. 2: 635 (1891). — L. divaricata (L.) Lunell in Amer. Midl.
Nat. 4 (7): 301 (1916).

Diagnosis: Annual monoecious herbs with tuberous roots: the
female inflorescences mainly subterminal, the males at the lower
axils; flowers pedicellate, the males 5-tepalled, strongly corniculate.
the females with 4 free tepals, a linear unbranched stigma and
glabrous laterally winged pedicels; the achenes semi-circular, sti-
pitate, on gynophores, smooth, without wings.

Distribution: Canada, U.S.A. (temperate parts) and Mexico
(highlands).

Monoecious annual herb, 1 to 2 m. high, with tuberous roots.
Stem slightly woody at the base, rarely branched, usually covered
with long white irritant hairs, especially on the apex, without
glandular hairs; stipular scars inconspicuous, petiolar scars con-
spicuous. Lamina (7—) 10-15 (—20) cm. long, (3—) 7-9 (12) cm.
broad, ovate to broadly ovate, rarely elliptic, chartaceous, without
glandular hairs; irritant hairs of the upper surface very short.
appressed, scattered, those of the lower surface mainly on midrib,
especially the basal part; cystoliths of both surfaces minute puncti-
form; base generally rounded to slightly cuneate; apex acuminate,
often sharply, 1-2 cm. long; margin dentate, rarely serrate, teeth
to 0.5 cm. long with base to 0.5 cm. broad; lateral veins 6-8 pairs,
prominent, the basal pair reaching to about half the length of
lamina; intercostals numerous, straight, rarely very prominent.
Petiole (3—) 5-10 (-15) cm. long, usually covered with fairly long
irritant hairs, especially the distal half. Stipule to 0.6 cm. long,
connate, intrapetiolar, bifid, the two tips as long as the lower
connate part, triangular, chartaceous, with very few short irritant
hairs, very rarely caducous. /nflorescence unisexual, paniculate,
well branched, axillary, usually solitary, without bulbils, with
minute peduncular bracts, with peduncle usually densely covered
with irritant hairs; the male (5—) 9-10 cm. long, 5—7 cm. broad,
usually in the lower axils; the female 10-15 cm. long, 5—7 cm.
broad, usually in the upper axils or subterminal; the male usually
wide spread, the female narrower and longer. Male flower pedicel-
late; perianth ca. 1 mm. long and broad, glabrous to well covered
with irritant hairs at the apex; tepals 5, strongly corniculate,
cymbiform; stamens 5, with slightly reflexed filaments to 1.5 mm.

W.-L. Chew — Monograph of Laportea 119

JURAIMI DEL.

pedicel.

EB;

Fig. 1. L. canadensis: A, twig; B, male flower; C, female flower; D, achene;

120 Gardens’ Bulletin, Singapore — XXV_ (1969)

long; pistillode ca. 0.5 mm. long, as broad at the apex, obpyrami-.
dal; interfloral bracts numerous, minute, narrowly triangular;
pedicel to 1 mm. long, articulated at about midpoint, glabrous.
Female flower pedicellate; perianth with 4 free tepals, the two
lateral tepals ca. 1 mm. long and broad, asymmetrical, enclosing
the ovary, glabrous; the dorsal one ca. 0.3 mm. long and broad,
strongly geniculate, often with few short irritant hairs; the ventral
ca. 0.25 mm. long, narrow; ovary ca. 1 mm. long and broad,
asymmetrically ovoid; stigma linear, unbranched, slightly kinked
at the base, to 2 mm. long; interfloral bracts minute, numerous,
narrowly elliptic; pedicel ca. 1 mm. long, winged laterally, usually
glabrous. Achene 2.5 mm. long and broad, stipitate, with stipe to
0.5 mm. long, on gynophore ca. 0.4 mm. long and broad, semi-
circular in shape, smooth, without wing, ridge nor depression,
one third covered by the lateral tepals, greatly reflexed; pedicel
to 3 mm. long; strongly winged laterally, glabrous; dispersed free
from the perianth.

A fairly common weed in rich Beech-Maple-Hemlock woods.
Vernacular names are Wood Nettle or Canadian Nettle (English);
Ortega (Mexican).

As I have not seen the types of canadensis and divaricata, I
base my interpretation of this species on that of Weddell. A native
of the cold temperate regions of North America, it extends to the
tropics in the south, only as highland plants, occurring/in Mexico:
at an altitude of about 2,200 metres.

This and the following two form a very closely allied group
of species which differ from each other in rather minor characters.

CANADA:—Manitoba Prov.: Marshall 21, July 1940 (UC).—Ontario
Prov.: Bruce Peninsula: Stokes Bay: Krotkov 8976, Aug. 1934 (A).—
Quebec Prov.: Gatineau Co.: Gatineau Park: Groh 1631, Sept. 1941 (S).
—New Brunswick :Restigouche & Malapedia rivers: Rousseau & Bovin:
32033, July 1929 (A).—Nova Scotia: Hants Co.: Five-mile River: Pease
& Long 21040, July 1920 (A).

UNITED STATES:—Alabama State: Clay Co.: Earle 892, Aug. 1828
(NY).—Arkansas State: Desha Co.: McDaniel 1401, Sept. 1959 (NY).—
Connecticut State: Bridgeport: Emes J, Sept. i895 (A).—Distr. of
Columbia: Potomac River: Morris 3lla, Sept. 1899 (F).—Indiana State:
Jenings Co.: Friesner 21785, Aug. 1947 (BR, MICH & S).—Kansas State:
Woodson Co.: Lathrop 1780, Sept. 1955( NY).—Louisiana State: St.
Landry: Correll 9472, July 1938 (A).—Massachusetts State: Hampden Co. :
Brookside: Seymour 17, Sept. 1913 (A, MO & NY).—Michingan State:
Bernien Co.: Benton: Voss 7551, July 1958 (MICH).—Nebraska State:
Niobrana River Valley: Tosted 652, Aug. 1936 (A).—North Carolina State:
Alamance Co.: Bowner 60, Sept. 1957 (MEL).—North Dakota State:
McHenry Co.: Boivin & Dore 8149, July 1961 (NY).—Oklahama State >
Comanche Co.: Stevens 1361], June 1913 (POM).—Washington State:
Falcon Valley: Suksdorf s.n. (L). :

MEXICO :—Nuevo Leon State: Sierra Madre Oriental: Meyer & Rogers
2824, July 1948 (MO, U & UPS)—Tamaulipas State: Gomez Farias
Region: Martin H.32, May 1953 (MICH).

W.-L. Chew — Monograph of Laportea 12]

(2) Laportea bulbifera (Sieb. & Zucc.) Wedd. in Arch. Mus. Hist.
Nat. Paris 9: 139 (1856) et in DC. Prodr. 16 (1); 82 (1869.—
Urtica evitata Wall. Cat. 4588 (1832), nom. nud. — U. bulbifera
Sieb. & Zucc. in Abhandl. Math.-Phys. Kl. Baier. Akad. 4 (3):
214 (1846). — Laportea terminalis Wight, Ic. Pl. Ind. or. 6: 9.
tab. 1972 (1853). — L. oleracea Wedd. in op. cit. 141 (1856) et
87 (1869). — Fleurya bulbifera (Sieb. & Zucc.) Bl. ex Miq.
Ann. Mus. Bot. Lugd. Bat. 3: 132 (1867). — L. evitata Wedd.
in op. cit. 79 (1869). — Urticastrum bulbiferum (Sieb. & ZUCC.)
O. K., evitatum (Wedd.) O. K., oleraceum (Wedd.) O.K. Rev.
Gen. Pl. 2: 635 (1891). — L. sinensis Wright in J. Linn. Soc.
26: 474 (1899). — L. dielsii Pampan. in Nuov. G. Bot. Ital.
17: 255 (1910). — Boehmeria bodinieri Leveille in Fedde,
Renert,.F)9 2550 (1913).

Diagnosis: Closely related to L. canadensis from which it differs
in the male flowers with non-corniculate tepals, the axils of leaves
normally with woody bulbils and the roots seldom tuberous.

Distribution: Japan, Korea, Manchuria, China, Tibet, Nepal,
Sikkim, Bhutan, Assam, India, Ceylon, Burma, Thailand, Vietnam,
Sumatra and Java.

Monoecious annual herb to 2 m. high, occasionally with tuber-
ous roots. Stem slightly woody at the base, rarely branched,
sparsely to densely covered with long white rigid irritant hairs,
especially at the nodes and apex, old parts becoming glabrous
without glandular hairs; stipular scars inconspicuous, petiolar scars
conspicuous; often with woody bulbils at the leaf axils. Lamina
(5—) 10-15 (—23) cm. long, (3—) 5—8 (—12) cm. broad, usually ovate
to broad ovate, chartaceous, without glandular hairs; irritant hairs
of the upper surface well distributed, slightly more on midrib
and veins, those of the lower surface mainly on midrib and veins,
especially the basal; cystoliths very minutely punctiform, numerous
on the upper surface, often absent from the lower surface; base
rounded to slightly cuneate, rarely cordate; apex acuminate, 1—2
cm. long; margin serrate to dentate, teeth to 0.5 cm. long, with
base to 0.5 cm. broad; lateral veins 4-6 pairs, prominent, the basal
pair reaching to more than half the length of the lamina; inter-
costals numerous, usually anastomosing, often rather faint. Petiole
(2-) 4-8 (-10) cm. long, sparsely to densely covered with irritant
hairs sometimes to 3 mm. long. Stipule 0.5—-1 cm. long, connate,
intrapetiolar, bifid, the two tips about half the length of the lower
connate part, often even shorter, ovate cymbiform, chartaceous,
often with short appressed irritant hairs, early caducous. /nflores-
cence unisexual, paniculate, well branched, axillary, solitary, very
rarely paired, with or without bulbils, with peduncular bracts to
1 mm. long, peduncle very sparsely covered with short appressed
irritant hairs to densely armed with long rigid ones; the male
4-6 (-10) cm. long, 3-9 cm. broad, usually in the lower axils; the

JURAIMI DEL.

Fig. 2. L. bulbifera: A, twig B, tuberous roots; C, male flower; D, female
flower: E, female perianth; F, achene.

W.-L. Chew — Monograph of Laportea 1823

female (7—) 10-20 (—30) cm. long, (2—) 5-10 (—15) cm. broad,
usually in the upper axils or subterminal; the male usually wide
spread, the female usually narrower and longer. Male flower pedi-
cellate; perianth ca. 1 mm. long and broad, almost glabrous to
densely pubescent with irritant hairs; tepals 5, non-corniculate,
cymbiform; stamens 5, with reflexed filaments to 1.5 mm. ‘ong;
pistillode ca. 0.5 mm. long, as broad at the apex, obpyramidal;
interfloral bracts to 1 mm. long, irregular in number, narrowly
elliptic; pedicel to 1 mm. long, articulated near the apex, glabrous
to light pubescent. Female flower pedicellate; perianth with 4 free
tepals, the two lateral tepals ca. 1 mm. long and broad, asymmetri-
cal, enclosing the ovary, with short appressed irritant hairs, the
dorsal one ca. 0.4 mm. long and broad, strongly geniculate. with
short irritant hairs, the ventral ca. 0.25 mm. long, narrow cymbi-
form; ovary ca. | mm. long and broad, asymmetrically ovoid;
stigma linear, unbranched, reflexed, to 3 mm. long; interfloral
bracts minute, irregular in number: pedicel ca. | mm. long, slightly
winged laterally, often glabrous. Achene to 3 mm. long and broad,
Stipitate, stipe 0.5 mm. Jong, on gynophore to 0.3 mm. long and
broad, semi-circular in shape, smooth, without wing, ridge nor
depression, one quarter covered by the lateral tepals, greatly
reflexed; pedicel to 2.5 mm. long, strongly winged laterally, gla-
brous; dispersed free from perianth.

This species is indeed very close to L. canadensis. Both have
somewhat the same habit of growth, leaf-form, inflorescence and
in fact similarly shaped achenes. The main difference between them
is in the male flowers whose tepals are not corniculate in this
species.

Again as in the preceding species, L. bulbifera has also a very
great latitudinal distribution, extending from cold temperate Man-
churia right into the hot tropics in Java, although in the tropical
countries it occurs as montane plants.

The types of sinensis, dielsii and bodinieri have been examined
and found to be conspecific and referable to this species.

The following are a small selection of well over 100 collections
studied. | }

JAPAN :—Hokkaido: near Nopporo: Dorsett & Morse 1043, Sept.
1929 (NY).—Hondo: Mt. Gaki-dake: Furuse 18966, July 1951 (S).—
Yokohama: Maximowicz s.n. (BM, FI, M & US).—Nagasaki: Maxmowicz

un (BM, BO, C, K & S).—sine loc.: Siebold s.n. 1842 (type of U. bulbifera:
).

KOREA :—North: Kirin Prov.: Komarov 514, Aug. 1896 (A, BM,
FI, K, LD, NY & P).—South: Mt. Odae Chung 2970, July 1946 (F &
MICH). .

MANCHURIA :—near Kyaolinzy: Litvinov 2162, Aug. 1902 (A & NY).

CHINA :—Kweichow Prov.: Bodinier 1748, Aug. 1897 (type of B.
bodinieri: E).—Hupei Prov.: Silvestri 409, Oct. 1906 (types of L. dielsii:
Fl).—West Szechuan: Pratt J/] (syntypes of L. sinensis: BM & K).—
Yunnan: Henry 11198 (E, K, MO & NY).

TIBET :—Konbo Prov.: Ludlow et al. 5289 & 5738, July 1938 (BM & EB).

SSM :—sine loc.: Hooker f. s.n. (type of L. oleracea: A, BM, G-DC
& P).

124 Gardens’ Bulletin, Singapore — XXV_ (1969)

INDIA :—Avalangy: Wight herb. 2681, Oct. 1852 (type of L. terminalis:
C, K, L, M & MEL).—Kodaikanal: Fischer 3079 (K).

CEYLON :—sine loc.: Thwaites 3336 (BM, CGE, G-DC & K).
BURMA:—Shan States: MacGregor 713, Sept. 1909 (E).

THAILAND :—Chiengmai: Garrett 1100, Oct. 1938 (A, BO, K & L).
—Doi Sutep: Kerr 2729, Oct. 1912 (BM. & K).

VIETNAM :—Tonkin: Poilane 12797 (P).

ete a al, Kerintji: Bunnemeyer 8812, March 1920 (B, BO &

JAVA:—Pasoeroean: van Steenis 7312, June 1935 (A, BO, BRI, L &
SING).—Mt. Malabar: Hochreutiner 1308, July 1904 (L & Z).

(3) Laportea alatipes Hk. f. in J. Linn. Soc. Bot. 7: 215 (1864);
Wedd. in DC. Prodr. 16 (1): 79 (1869); Letouzey, in FI.
Cameroun, 8: 117. pl. 18 (1968). — Urticastrum alatipes (Hk.
f.) O.K. Rev. Gen. Pl. 635 (1891). — Fleurya urticoides Engl.
in Bot. Jahrb. 33: 122 (1902): var. glabrata Rendle in Prain,
Fl. Trop. Afr. 6 (2): 248 (1917). — Girardinia marginata Engl.
in op. cit. 123 (1902). — F. alatipes (Hk. f.) Brown in Dyer, FI.
Cap. 5 (2): 547 (1925).

Diagnosis: It distinguishes itself from the preceding species by
the following characters: (a) male flowers 4-tepalled, (b) the
achenes somewhat pear-shaped and not semi-circular, (c) stipe of
the achene up to 1 mm. long.

Distribution: Kenya, Tanganyika, Nyasaland, Southern Rhodesia,
South Africa, Belgium Congo and Cameroun.

Monoecious annual (?) herb to 1.5 m. high, very rarely with ©
stolons. Stem slightly woody at the base, usually unbranched
covered with rather long irritant hairs, without glandular hairs;
stipular scars fairly conspicuous, petiolar scars quite conspicuous.
Lamina (4-) 8-15 (—20) cm. long, (2—) 4-8 (—13) cm. broad, ovate,
rarely narrowly, chartaceous, without glandular hairs; irritant hairs
of the upper surface dispersed, those of the lower surface mainly
on midrib and veins; cystoliths of both surfaces punctiform; base
generally rounded, sometimes slightly cordate, very rarely cuneate;
apex acuminate, often sharply, usually less than 1.5 cm. long;
margin serrate, very rarely serrulate or dentate, teeth to 0.5 cm.
long, with base to 0.5 cm. broad; lateral veins 5—7 pairs, pro-
minent, the basal pair reaching to more than half the length of
lamina; intercostals numerous, fairly straight and prominent.
Petiole (2.5—) 5-12 (—16) cm. long, usually well covered with irritant
hairs. Stipule to 1.5 cm. long, connate, intrapetiolar, bifid, the two
tips shorter than the lower connate part, triangular, chartaceous,
almost glabrous. /nflorescence unisexual, paniculate, well branched,
axillary, solitary, without bulbils, with minute peduncular bracts,
peduncle usually well armed with irritant hairs; the male 3-10 cm.
long, 2-6 cm. broad, usually in the lower axils; the female 10-20
cm. long, 5-10 cm. broad, usually in the upper axils or subter-
minal; the male very small compared to the female. Male flower
pedicellate; perianth to 1 mm. long and broad, rarely without
irritant hairs at the apex, usually lightly pubescent; tepals 4,

125-

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WD +

W.-L. Chew — Monograph of Laportea

126 Gardens’ Bulletin, Singapore — XXV (1969)

usually corniculate; stamens 4, with reflexed filaments to 1.6 mm.
long; pistillode ca. 0.4 mm. long, ca. 0.3 mm. broad, obconical;
interfloral bracts to 0.8 mm. long, ca. 0.2 mm. broad, narrow ~
ovate, irregular in number, fringed with short irritant hairs; pedicel —
to 1 mm. long, articulated near the apex, glabrous. Female flower —
pedicellate; perianth with 4 free fringed tepals, the two laterals
tepals to 1 mm. long, ca. 0.4 mm. broad, enclosing the ovary,
very light pubescent with few short irritant hairs, the dorsal
one up to 0.5 mm. long, geniculate, with a few long irritant hairs,
the ventral ca. 0.4 mm. long and broad, broadly ovate; ovary ca.
0.7 mm. long, ca. 0.6 mm. broad, asymmetrically ovoid; stigma
linear, unbranched, slightly reflexed, up to 2 mm. long; interfloral
bracts ca. 0.6 mm. long, ca. 0.3 mm. broad, narrow ovate, irre-
gular in number, glabrescent; pedicel ca. 1 mm. long, winged
laterally, usually glabrous. Achene ca. 1.6 mm. long, ca. 1 mm.
broad, stipitate, with stipe to 1 mm. long, on gynophore 0.4 mm.
long and 0.3 mm. broad, somewhat pyriform, often smooth, with
a very narrow wing, the lateral sides often raised, with tepals at
the base, greatly reflexed; pedicel to 1.5 mm. long, strongly winged
laterally, with few very long irritant hairs, otherwise glabrous;
dispersed free from the perianth.

A highland or montane species occurring above the 1,500 m.
contour, this is said to be very common in clearings in Podocarpus
forests. Cooper recorded its use to punish naughty boys in the
Congo!

Native names: Kwisa (Nyasaland), Itusa & Hou (Congo) and
Kenjie (Cameroun).

KENYA :—Aberdare Mt.: Fries & Fries 708, Jan. 1922 (BR, S, UPS &
Z).—Kakamega Forest: Drummond & Hemsley 4757, Oct. 1953 (BR, FI

& S).—Nakuru Distr.: Maas Geesteranus 5904, Aug. 1949 (BR, F, G, L,
MO, Sed UIC),

TANGANYIKA :—Kilimanjaro Mt.: Schlieben 4906 (BR, G & Z).—
Proto Mts.: Richards 9707, May 1957 (BR).—Kwegoka: Drummond &
Hemsley 2711, May 1953 (BR, LISC & S).

NYASALAND:—Mlanje Distr.: Mt. Mlanje: Brass 16572, July 1946
(MO, NY & US).

te RHODESIA :—sine loc.: Wild 4437, March 1954 (MO &
PS).

SOUTH AFRICA :—Griqualand east: Zuurberg: Tyson 1772, Feb. 1884
(SAM).—Mt. Curie: DFO. 579 (Z).

BELGIUM CONGO :—Olbert National Park: Lebrun 8715, Nov. 1937
(BR).—Mikeno Mt.: Linder 2390, March 1927 (A).—Ruwenzori: Bequaert
4258, May 1914 (BR).

CAMEROUN :~—-sine loc.: Mann 1973 (type of L. alatipes: K); Preuss
‘916 (type of F. urticoides: K).

(4) Laportea disepala (Gagn.) Chew in Gard. Bull. Sing. 21: 200
(1965). — Fleurya disepala Gagn. in Bull. Soc. Bot. Fr. 75: 556
(1928).

Diagnosis: This differs from all others of sect. Laportea in the
following characters: (a) the lateral sides of the achene with a
slightly raised ridge enclosing a warty depression, (b) the lateral
tepals displaced to one side at the base of the achene.

Distribution: Thailand, Cambodia and South Vietnam.

i)
~!)

W.-L. Chew — Monograph of Laportea |

a \
if.

cae “ay
SH

JURAIMI DEL.

Fig. 4. L. disepala: A, twig: B, male flower; C, female flower; D, achene:
E, pedicel.

128 Gardens’ Bulletin, Singapore — XXV (1969)

Monoecious annual herb to 60 cm. high. Stem slightly woody
at the base, unbranched, well covered with long and short irritant
hairs especially at the apex, without glandular hairs; petiolar scars
conspicuous, stipular scars not so. Lamina 5-14 cm. long, 5-11
‘cm. broad, broad ovate, chartaceous, without glandular hairs;
irritant hairs of the upper surface fairly dense, those of the lower
surface mainly on midrib, veins and veinlets; cystoliths punctiform,
‘often very minute, more on the upper surface; base cordate; apex
acuminate, to 1 cm. long; margin serrate, teeth to 0.5 cm. long,
with base to 0.5 cm. broad; lateral veins 4-5 pairs, the basal pair
reaching to more than half the length of the lamina; intercostals
numerous, fairly straight, not very prominent. Petiole to 15 cm.
long, often longer than the lamina, well covered with irritant hairs.
Stipule ca. 0.5 cm. long, connate, intra-petiolar, bifid, the two tips
longer than the lower connate part, narrowly triangular, charta-
ceous, glabrous, early caducous. Inflorescence usually unisexual,
paniculate, well branched, axillary, solitary, very rarely paired, with
few very minute peduncular bracts, peduncle well covered with
rather long irritant hairs; the male ca. 7 cm. long, 3 cm. broad, at
the lower axils; the female to 16 cm. long, 10 cm. broad, in the
upper axils or subterminal; the male much smaller than the female.
Male flower pedicellate; perianth ca. 1 mm. long and broad,
densely covered with short to very long irritant hairs; tepals 4,
non-corniculate, broad cymbiform; stamens 4, with reflexed
filaments to 1.5 mm. long; pistillode ca. 0.4 mm. long. ca. 0.2 mm.
broad at the apex, obpyramidal; interfloral bracts narrow elliptic,
up to 0.5 mm. long, 2-3 to each flower, with few irritant hairs;
pedicel to 1 mm. long, articulated at the apex, glabrescent. Female
flower pedicellate; perianth with 4 free tepals, the two lateral tepals
ca. 0.8 mm. long and ca. 0.4 mm. broad, almost symmetrically
ovate, enclosing the ovary, with 2-4 irritant hairs on the midrib,
with elongated cystoliths; the dorsal one ca. 0.4 mm. long, ca.
0.2 mm. broad, fairly strongly geniculate, with long irritant hairs:
the ventral ca. 0.2 mm. long, 0.1 mm. broad, lightly pubescent:
ovary ca. 0.4 mm. long, ca. 0.3 mm. broad, slightly asymmetrically
ovoid, stigma linear, unbranched, bent, ca. 0.5 mm. long; interfloral
bracts to 0.4 mm. long, narrowly ovate, ca. 2 to each flower; pedicel
ca. 0.4 mm. long, slightly winged laterally, glabrescent. Achene ca.
1.5 mm. long, ca. 1.2 mm. broad, stipitate, with stipe to 0.2 mm.
long, on gynophore ca. 0.2 mm. long and broad, ovoid, without
wing, on each lateral side a slightly raised ovate ridge enclosing a
warted depression, with the lateral tepals at the base and displaced
to one side, reflexed; pedicel to 3.5 mm. long, strongly winged
laterally at the apex, glabrous; dispersed free from the perianth.

This rare species, confined to the limestones in the countries
enumerated, bears some superficial resemblance to L. aestuans of
sect. Fleurya. Gagnepain’s comment that this is related to Fleurya
cordata is therefore not surprising. In my preliminary list in 1965,
this was erroneously referred to sect. Fleurya to which it does not
belong. The species closest to this is L. violacea (see notes there).

W.-L. Chew — Monograph of Laportea 129

The specific epithet disepala is a misnomer. The female perianth
has 4-tepals, not 2; but as in the other species, the medial ones are
so small that they are not readily visible to the hurried observer.

Thailand:—Loei: Pha Nuk Khao: Shimizu et al. 7.8702, Sept. 1967

(SING).—Raohasima: Pak Chawng: Smitinand 4836, Sept. 1958 (BKF).—
Saraburi: Mak Lek: Smitinand & Sleumer 1324, Oct. 1963 (L & SING).

CAMBODIA:—Mt. Kereer: Pierre 4655, April 1870 (type of F.
disepala: A & P).
SOUTH VIETNAM :—Cochinchina: Pierre 4806 (A & P).

(5) Laportea violacea Gagn. in Bull. Soc. Bot. Fr. 75: 4 (1928). —
L. vitifolia Hand.-Mazz. Symb. Sinic. 7: 117 (1929).

Diagnosis: Monoecious herb or subshrub less than 2 metres;
the lamina broad ovate, 6-12 cm. long, truncate at the base; the
male flowers sessile, 5-merous, with non-corniculate tepals.

Distribution: China, Vietnam (North & South) & Thailand.

Monoecious herb or subshrub, usually ess than 2 m. high. Stem
woody at the base, well branched, glabrous to glabrescent, irritant
hairs often absent, without glandular hairs; petiolar scars conspi-
cuous, stipular scars not so. Lamina 6-12 cm. long, 4-10 cm.
broad, broadly ovate, chartaceous to sub-coriaceous, without
glandular hairs; irritant hairs of both surfaces short, appressed and
dispersed, very rarely dense; cystoliths of both surfaces punctiform:
base usually truncate, smooth, rarely slightly cordate; apex
acuminate, usually sharply, ca. 1 cm. long; margin serrate, teeth to
0.4 cm. long, with base to 0.6 cm. broad; lateral veins 4—5 pairs,
prominent, the basal pair most prominent and reaching to more
than half the length of the lamina; intercostals rather straight and
prominent. Petiole (3—) 6—9 cm. long, well covered with minute irri-
tant hairs. Stipule to 0.2 cm. long, connate, intrapetiolar, bifid,
the two tips very short, narrow ovate, chartaceous, light pubescent.
Inflorescence unisexual, paniculate, fairly well branched, axillary,
solitary, with minute peduncular bracts, peduncle densely covered
with short irritant hairs; the male up to 6 cm. long, ca. 3 cm.
broad, usually much smaller, mainly:at the lower axils; the female
15—20 (-30) cm. long: 5-8 cm. broad, usually at the upper axils or
subterminal: the male generally much smaller than the female.
Male flower sessile to subsessile; perianth ca. | mm. long and
broad, densely covered with short irritant hairs; tepals 5, non-
corniculate, cymbiform; stamens 5, with reflexed filaments to 1.5
mm. long; pistillode ca. 0.2 mm. long and broad, obpyramidal:
interfloral bracts ca. 0.2 mm. long and broad, 4—5 to each flower.
densely covered with short irritant hairs. Female flower pedicellate:
perianth with 4 free tepals covered with elongated cystoliths; the
two lateral tepals to 0.8 mm. long, ca. 0.4 mm. broad, almost
enclosing the ovary, with a few short irritant hairs, otherwise
glabrous, the dorsal one ca. 0.2 mm. long, slightly geniculate, with
a few irritant hairs, the ventral ca. 0.3 mm. long, glabrous: ovary
ca. 0.8 mm. long, ca. 0.4 mm. broad, ellipsoid; stigma Iinear, un-
branched, reflexed, up to 2.5 mm. long; interfloral bracts to 0.3

130 Gardens’ Bulletin, Singapore — X XV (1969)

rapa late , <
CEES

Seen gad,

Seivere tree

CATIONS

JURAIMI DEL.

Fig. 5. L. violacea: A, twig: B, male flower: C, female flower; D, achene:
E, pedicel.

W,-L. Chew — Monograph of Laportea 131

mm. long and broad, 3-5 to each flower, often with irritant hairs:
pedicel to 0.4 mm. long, winged laterally, somewhat glabrous.
Achene ca. 2 mm. long, ca. 1.5 mm. broad, stipitate, stipe ca.
0.4 mm. long, on gynophore ca. 0.2 mm. long and broad, asymme-
trically ellipsoid, the lateral sides warted, often raised, surrounded
by a very narrow wing, greatly reflexed; pedicel up to 6 mm. long,
strongly winged laterally, glabrous; dispersed free from the
perianth.

This species has quite a lot in common with the preceding one
with which it is obviously closely related. Both have somewhat
the same habit of growth, shape of lamina and inflorescence and
are both found in the same habitat. They are, however, easily told
apart as in this species the lamina base is truncate, the male flowers.
5-merous and the achene without a depression on the lateral sides.

CHINA :—Lungchcow: Pan Wang Hills: Morse 218 (type of L. vitifolia:
K).—Kwangsi Prov.: Lan Lon: Ching 6640, Aug. 1928 (NY).

VIETNAM :—Tonkin: Lecomte & Finet 260 (P); Bon 2269 (lectotype
of L. violacea: P)— Annam: Bon 5724 (P).

THHAILAND:—Chumphae: Smitinand 4910, Nov. 1958 (BKF);.
Sorensen et al. 6103, Feb. 1958 (SING).

(6) Laportea amberana (Baker) Leandri in Ann. Mus. Col. Mars.
ser. 6. 7-8: 18 (1950) et in Humbert, Fl. Madagascar, 56: 11.
fig. 2 (1965); Chew in Gard. Bull. Sing. 21: 198 (1965). —
Urera amberana Baker in J. Bot. 20: 267 (1882). — L. humbertii
Leandri in op. cit. 19 (1950) & 11 (1965); Chew in op. cit. 199
(1965). — L. manombensis Leandri in op. cit. 21 (1950); Chew in
op. cit. 199 (1965).

Diagnosis: (a) monoecious shrubs with cordate, coarsely, irregu-
larly and shallowly dentate lamina, (b) irritant hairs appressed, (c)
male flowers dense with irritant hairs, sessile, 5-merous, with non-
corniculate tepals, (d) female flowers pedicellate, with ribbed lateral
tepals, (e) achenes almost circular, stipitate, on strongly winged
pedicels to 10 mm. long.

Distribution: Madagascar (endemic!). .

Moncecious shrubs to 1.5 metres high. Twigs somewhat thick,
subglabrous. Lamina 12-20 cm. long, 9-18 cm. broad, very broad
ovate, chartaceous, both surfaces well covered with appressed irri-
tant hairs and punctiform cystoliths; base profound cordate, the
two lobes sometimes touching; apex acuminate, acumen 1-1.5 cm.
long; margin coarsely, irregularly and shallowly dentate, teeth at
most | cm. long, 0.5 cm. broad at the base; lateral veins 5S—6 pairs,
the basal pair very prominent, reaching to half the length of the
lamina; intercostals numerous, somewhat anastomosing, rather
faint. Petioles to 10 cm. long, lightly covered with appressed irri-
tant hairs. Stipules less than 5 mm. long, caducous. Inflorescences
unisexual, paniculate with very short lateral branches, axillary,
usually in the upper axils; peduncle dense with somewhat erect
irritant hairs. Male flowers sessile; perianth ca. 2 mm. long and
broad, dense with irritant hairs, with 5 non-corniculate tepals, 5

132 Gardens’ Bulletin, Singapore — XXV (1969)

del.

Fig. 6. L. amberana: A, leaf; B, male inflorescence; C, female inflores-
cence; D, achene with pedicel.

W.-L. Chew — Monograph of Laporteu 133

Stamens on reflexed filaments; pistillode small. Female flowers
pedicellate; the perianth 4-tepalled covered with irritant hairs, the
laterals ca. 1 mm. long, 0.5 mm. broad, ribbed in the centre.
enclosing the ovary, the dorsal one ca. 0.5 mm. long, slightly
geniculate and the ventral ca. 0.6 mm. long; ovary ca. 1 mm. long,
ellipsoid: stigma linear, ca. 0.75 mm. long, simple. reflexed;
pedicel ca. 1 mm. long, slightly winged laterally, dense with
irritant hairs. Achenes ca. 3 mm. long and broad, almost circular,
stipitate, with stipe to 0.75 mm. long, on a very short gynophore,
lateral sides minutely tuberculate, with the tepals covering the lower
portion of the achene, greatly reflexed; pedicel to 10 mm. long,
strongly winged laterally especially at the apex. well armed with
irritant hairs; dispersed free from the perianth.

I do not agree with Leandri that L. humbertii is distinct from
this species. The differences pointed out by him are, in my opinion
(when viewed from a monographic angle), insufficient to warrant
their separation. The species L. manombensis was reduced by
Leandri to this in 1965 but the name was inadvertently omitted
from the synonymy.

This and the following three form a related group of species
entirely endemic in Madagascar. Though they differ from each
other rather conspicuously in vegetative characters, they possess
many other fundamental characters in common. Their female
inflorescences, for instance, are usually much longer than the
leaves and are furnished with rather stout peduncles. Their closest
ally outside Madagascar is perhaps L. alatipes of East Africa with
which this group is linked by L. weddellii.

L. amberana grows in rather dry forests on sandstones or lime-
stones. Its specific epithet is a direct derivation from the Mada-
gascan native name “amberana”’.

MADAGASCAR :—West: Manombo: Humbert 19991, Jan.. 1947 (type
of L. manombensis: P). Analavelono: Humbert 14234, March 1934 (type
of L. humbertii: P & SING).—Centre: Betsileo: Baron 321 (type of U.
amberana: P & SING). ;

(7) Laportea septentrionalis Leandri in op. cit. 19 (1950) et 12
(1965); Chew in op. cit. 199 (1965).

Diagnosis: It distinguishes itself thus (a) herbs with very few,
small, cordate, irregularly and coarsely-toothed leaves, (b) both
surfaces of the lamina densely covered with irritant hairs and
punctiform cystoliths, (c) inflorescence about 3 times as long as
leaves. (d) stigma about 3 times as long as flower.

Distribution: Madagascar (endemic! ).

Herbs less than 50 cm. high with few leaves. Lamina ca. 5 cm.
long and broad, very broad ovate, chartaceous, both surfaces dense
with appressed irritant hairs and punctiform cystoliths; base cor-
date; apex acute; margin irregularly, coarsely and bluntly dentate,

134

Gardens’ Bulletin, Singapore — XXV_ (1969)

Fig. 7. L. septentrionalis: fruiting twig.

W.-L. Chew — Monograph of Laportea 135

teeth at most 3 mm. long, 5 mm. broad; lateral veins ca. 4 pairs,
the basal pair more prominent, reaching to more than half the
length of lamina: intercostals anastomosing, faint. Petioles to 10
cm. long, well covered with appressed irritant hairs. Stipules mem-
braneous, caducous. /nflorescences unisexual, paniculate, branched
weakly near the apex, axillary, 15 to 25 cm. long, peduncle well
covered with minute irritant hairs. Male flowers sessile, to 2 mm.
long and broad, with irritant hairs, with 5 non-corniculate tepals
and 5 stamens. Female flowers pedicellate; stigma filiform to 4
mm. long, longer than ovary. Achenes ovate, to orbicular, flatten-
ed, ca. 2.5 mm. long, warted on broadly winged pedicels 6-7 mm.
long.

This species comes very near to L. humbertii (= L. amberana) in
lamina shape, venation and even dentition of the margin. Their
female inflorescences are also somewhat similar in shape. They
differ, however, in the habit of growth, lamina size and the number
of achenes in the inflorescence.

It is recorded to be a very rare species of limestones.

MADAGASCAR :—Diego-Suarez Proyv.: Ankarana: Humbert 19004,
Dec. 1937—Jan. 1938 (holotype in P).

(8) Laportea perrieri Leandri in Ann. Mus. Col. Mars. ser. 6.
7-8: 20 (1950) et in Humbert, Fl. Madagascar, 56: 12 (1965):
Chew in Gard. Bull. Sing. 21: 199 (1965).

Diagnosis: (a) monoecious shrubs with cordate, very pro-
foundly dentate lamina, (b) irritant hairs minute, appressed, (c)
inflorescences unisexual, branched racemes (d) male flowers sub-
sessile, 5-merous, with non-corniculate tepals.

Distribution: Madagascar (endemic! ).

Monoecious shrubs to 3 mertes high. Twigs thick and fleshy,
well armed with short irritant hairs. Lamina to 20-65 cm. long
and broad, somewhat circular, chartaceous, with short appressed
irritant hairs mainly on the lower surface and punctiform cystoliths
mainly on the upper; base cordate; apex long acuminate, acumen
to 6 cm. long; margin very profoundly dentate, teeth 7-20 cm.
long, 3-10 cm. broad at the base: lateral veins 4—S pairs, the basal
pair most prominent, rising to about half the length of the lamina;
intercostals numerous. rather faint. Petioles 20-60 cm. long, dense
with short appressed irritant hairs. Stipules to 5 mm. long. Inflore-
scences unisexual, paniculate, axillary, solitary, rarely paired; the
females in the upper axils, the males in the lower, both wide-
spreading and long; peduncle well covered with short appressed
irritant hairs. Male flowers subsessile; perianth ca. 2 mm. long and

broad, lightly covered with short appressed irritant hairs and punc-
tiform cystoliths, with 5 non-corniculate tepals, 5 stamens on

136

Gardens’ Bulletin, Singapore — XXV (1969)

4cm

Fig. 8. L. perrieri: A, leaf; B, male inflorescence; C, female inflorescence.

W.-L. Chew — Monograph of Laportea 157

slightly reflexed filaments; pistillode small, ca. 1 mm. long, obco-
nical; interfloral bracts minute; pedicel extremely short. Female
flowers pedicellate; the perianth 4-tepalled, the laterals ca. 0.75
mm. long, ca. 0.5 mm. broad, almost enclosing the ovary, with
few erect irritant hairs, the dorsal one ca. 0.4 mm. long, geniculate
and the ventral ca. 0.3 mm. long; ovary ca. 0.75 mm. long, 0.5
mm. broad, ovoid; stigma linear, ca. 0.75 mm. long, simple, reflex-
ed; interfloral bracts minute; pedicel ca. 0.75 mm. long, winged
laterally, with erect fairly long irritant hairs. Achenes ca. 3 (-5)
mm. long, 3-4 mm. broad, often asymmetrically elliptic, stipitate,
with stipe to ca. 0.5 mm. long, on a very short gynophore, lateral
sides very minutely tuberculate with the tepals covering the stipe,
greatly reflexed; pedicel to 4 mm. long, strongly winged laterally,
well armed with long erect irritant hairs; dispersed free from the
perianth.

Related to L. amberana (see notes there).

MADAGASCAR :—near Marovato: G. Cours 5449, Jan. 1960 (K & P).
— Namoroka (Ambongo): Perrier de la Bathie 475 (type: K & P).

(9) Laportea weddellii Leandri in op. cit. 22 (1950) et 14 (1965);
Chew in op. cit. 199 (1965).

Diagnosis: (a) monoecious shrubs to 1.5 metres, (b) lamina
longer than petioles, elliptic to slightly ovate, with erect irritant
hairs, (c) teeth of lamina ca. 7-8 mm. long & broad, (d) female
inflorescences slightly longer than lamina, (e) stigma 3-4 mm.
long.

Distribution: Madagascar (endemic! ).

Monoecious shrubs 1-5 metres high. Twigs bright red, with
leaves crowded near the apices. Lamina 10-15 cm. long, 6—7 cm.
broad, elliptic to slightly ovate, chartaceous, subglabrous on both
surfaces, sparse with erect irritant hairs; base truncate to rounded;
apex shortly acuminate; margin grossly dentate, teeth ca. 7-8 mm.
long and broad at the base; lateral veins 4 pairs, rather prominent,
the basal pair reaching to half the length of lamina. Petioles 5—6
cm. long, well armed with erect irritant hairs to 2 mm. long.
Stipules minute, early caducous. Inflorescences unisexual, panicu-
late, to 15-20 cm. long, axillary, the females slightly larger than
the males. Male flowers subsessile, ca. 2 mm. Jong and broad.
Female flowers with small lateral tepals to 0.5 mm. long, on
pedicels 5-6 mm. long; stigma linear., 3-4 mm. long. Achene 3-4
mm. long, 2.5-3 mm. broad.

This description is based on that of Leandri as I have seen only
a single leaf of the only collection known of this species. As in L.
septenirionalis numerous floral details are not available due to
paucity of collections.

MADAGASCAR :—Manambolo Valley: Humbert 13221 (holotype in P).

138 Gardens’ Bulletin, Singapore —XXV (1969)

rc 3ocm

Fig. 9. L. weddellii: a leaf.

W.-L. Chew — Monograph of Laportea 139

410) Laportea decumana (Roxb.) Wedd. in Arch. Mus. Hist. Nat.
Paris 9: 127 (1856) et in DC. Prodr. 16 (1): 80 (1869). — Urtica
decumana Rumph. Herb. Amb. 6: 47, tab. 20, fig. 1 (1750), ante
Linn.; Roxb. Fl. Ind. 3: 587 (1832). — U. rumphii Kostel. in
Allg. Med. Pharm. FI. 2: 400 (1833). — Boehmeria macrothyrsa
Mig. in Zoll. Syst. Verz. Ind. 100, 103 (1854). — Laportea
armata Warb. in Bot. Jahrb. 13: 293 (1891). — Urticastrum
decumanum (Roxb.) O.K. op. cit. 635 (1891). — Laportea pedun-
culata K. Sch. & Laut. Fl. Deutsch. Sudsee 290 (1901). — L.
humilis Laut. in K. Sch. & Laut. Nachtr. Fl. Deutsch. Sud. 251
(1905).

Diagnosis: Easily distinguished thus: (a) perennial shrubs, sub-
‘shrubs or tall herbs, (b) leaves rugose, dense with long irritant hairs
young ones dense wooly, (c) margin of lamina serrulate to denti-
culate; (d) stipules to 2 cm. long, subcoriaceous, lightly bifid, (e)
stigma to 3 mm. long, (f) male interfloral bracts longer than male
flowers.

Distribution: South Borneo, Celebes to New Guinea.

Monoecious perennial herbs, subshrubs or shrubs to 2 m. high.
Stem woody, wood soft, brittle, well branched, densely armed with
long rigid irritant hairs, especially at the apices, without glandular
hairs; stipular and petiolar scars fairly conspicuous. Lamina (17-)
20-30 (-35) cm. long, (6—) 12-18 cm. broad, ovate, rhombic to trul-
late, rarely elliptic, subcoriaceous, usually strongly rugose, dense
woolly, especially on the lower surface of young leaves, usually
light pubescent to almost glabrous on the upper surface; stiff long
irritant hairs present on both surfaces, more on the lower; cystoliths
-of the upper surface minute punctiform, those of the lower surface
elongated; base round to cordate; apex long acuminate, often sharp-
ly, up to 2 cm. long; margin serrulate to denticulate, teeth rarely
exceed 0.3 cm. long with 0.3 cm. broad base, very often only 0.1
cm. long; lateral veins 7-9 (—11) pairs, prominent, basal pair
reaching to a third the length of lamina, usually less prominent
than the others; intercostals numerous, slightly straight to anasto-
mosing, prominent. Petioles (2—) 5-8 (—10) cm. long, usually with
long rigid irritant hairs, young ones often dense woolly. Stipules
to 2 cm. long, connate, intrapetiolar, very slightly bifid at the apex,
cymbiform, chartaceous to subcoriaceous, with short irritant hairs,
quite persistent. Jnflorescences unisexual, paniculate, well branched,
well branched, axillary, usually solitary, without bulbils, with
peduncular bracts to 0.5 cm. long, with peduncle usually densely
covered with irritant hairs; the male (10—) 16—20 (—25) cm. long,
(4-) 6-10 (-15) cm. broad, usually in the lower axils; female
(20-) 30-40 (-55) cm. long, (4-) 8-15 (—20) cm. broad, usually in
the upper axils or subterminal, the female usually larger than the
male. Male flowers pedicellate; perianth to 2 mm. long and broad,
with long irritant hairs at the apex, otherwise glabrous, with elon-
gated cystoliths; tepals 4, non-corniculate, cymbiform; stamens 4,

140 Gardens’ Bulletin, Singapore —_XXV_ (1969)

Eee tt ea Pa ‘
yr. LE r5> soat Ste 5
TS Ae a ie
Separate tects Sy a | {apa & LS ‘ es
=A eit me: Sees ae
INS SSA are Tee SSPE SS ee ate ZAG wee
: = So
Z d

Sorc oa

Ses

Serta
eh

TARN

oo
ee :
\ kat

{mm JURAIMI DEL.

Fig. 10. L. decumana: A, twig; B, leaf surface; C, male flower; D, female
flower; E, pedicel; F, achene.

W.-L. Chew — Monograph of Laportea 141

with reflexed filaments to 2 mm. long; pistillode ca. 0.75 mm. long,
ca. 0.5 mm. broad at the apex, obpyramidal; interflora bracts
narrow lanceolate, almost as long as flower, sparsely covered with
irritant hairs; pedicel to 2 mm. long, articulated near the apex,
glabrous. Female flowers pedicellate; perianth with 4 free tepals,
the two lateral tepals ca. 1 mm. long and 0.75 mm. broad,
slightly asymmetrical, usually enclosing the ovary, with 2 to 4
long irritant hairs at the base, with elongated cystoliths; the dorsal
one ca. 0.75 mm. long, ca. 0.3 mm. broad, often slightly geniculate,
usually fully armed with long irritant hairs; the ventral ca. 0.5 mm.
long, narrow, almost glabrous; ovary ca. 1 mm. long, ca. 0.75
mm. broad, asymmetrically ovoid; stigma linear, unbranched,
slightly bent, to 3 mm. long; interfloral bracts to 1 mm. long,
narrow ovate to lanceolate, numercus; pedicel ca. 0.5 mm. long,
slightly winged laterally, usually with long irritant hairs. Achenes
to 2.5 mm. long and broad, stipitate, with stipe to 0.75 mm. long,
on gynophore to 0.75 mm. long, pyriform, smooth, without wing,
ridge nor depression, with lateral tepals at the base, greatly reflex-
ed; pedicel to 1.5 mm. long, slightly to strongly winged laterally.
with long irritant hairs at the margin; dispersed free from the
perianth.

I find no difficulty in distinguishing this species from the others
in the genus. The habit of growth, leaf-form, texture and vestiture
and numerous other characters make this easily the most distinc-
tive.

Its affinity is not entirely clear to me. On growth habit, I would
consider this a close relative of the Madagascan group of species!
I see in L. decumana a link, though not a very strong one, between
this genus and Dendrocnide. Its stipules are certainly rather simi-
lar in size and texture to those of the latter genus.

Winkler’s concept of this species is too restricted. Having
examined a vast quantity of collections from various locations, I
decide to stretch it to cover not only L. humilis but also B. macro-
thyrsa which was described from a plant introduced into cultivation
in Java from Amboina. .

I select as a neotype for this species a collection of Roxburgh,
reference Wallich Cat. 4620 in Kew Herbarium.

BORNEO :—South-East: sine loc.: Korthals s.n. (L. herb. 906—-171-31 &
908-188-3265).

CELEBES:—sine loc.: de Vriese s.n. (L. herb. 908-188-3268 & 3278).

MOLUCCAS:—Amboina: Robinson, Pl. Rumph. Amb. 316, July—Nov.
1913 (BO, K, L & US).—Ceram: Eyma 1804, Oct. 1937 (BO, K, L, PNH
& SING).—Tanimbar Islands: Borssum-Waalkes 3366, April 1956 (L).

NEW GUINEA :—North-East: Morobe Distr.: Womersley & van Royen
5884, June 1954 (A, BRI, L & LAE). Sattelberg: Warburg 20764, April
1889 (type of L. armata: BM).—West Manokwari: Kostermans 2703 Aug.
1948 (BO & L).—Papua: Boridi: Carr 12995 & 13012, Oct. 1935 (A, BM,
K & L).

Cultivated :—India: Roxburgh, Wall. Cat. 4620 (neotype in K).—Java:
Zollinger HZ. 1149 (type of B. macrothyrsa: UV).

142 Gardens’ Bulletin, Singapore — XXV_ (1969)

(11) Laportea macrostachya (Maxim.) Ohwi in J. Jap. Bot. 12:
331 (1936). — Sceptrocnide macrostachya Maxim. in Bull. Acad.
Petersb. 22: 238 (1877). — L. grossedentata Wright in J. Linn.
Soc. 26: 474 (1899). — L. giraldiana Pritz. in Bot. Jahrb. 29:
301 (1901). — L. longispica Pampan. in Nuov. G. Bot. Ital. 17:
256 (1901). — L. forrestii Diels in Notes Bot. Gard. Edin. 5:
292: (1912).

Diagnosis: (a) lamina grossly dentate, strongly caudate, with
the long tip arising from the broadly emarginate to truncate apex,
(6) female inflorescences elongated, unbranched, (c) achenes
smooth, without ridge nor wing, totally enclosed by the lateral
tepals strangly ribbed on the outside.

Distribution: Japan, China and Burma.

Monoecious annual herbs, usually less than 2 m. high,
occasionally with tuberous roots. Stem woody at the base, usually
unbranched, often well armed with long mounted irritant hairs,
especially at the nodes and apex, old parts usually glabrous,
without glandular hairs; petiolar scars very prominent, stipular
scars not so; rarely with woody bulbils at the leaf axils. Lamina
(7-) 10-20 (—28) cm. long, (5—) 8-18 (-28) cm. broad, ovate,
broadly to very broadly so, chartaceous, without glandular hairs;
irritant hairs more on the lower than on the upper surface,
frequently sparse on matured leaves; cystoliths very minutely
punctiform, numerous on the upper surface, fewer on the lower
surface; base cordate, often very broadly; apex broadly emarginate
to truncate between the ultimate pair of teeth, surmounted
by a narrowly ovate cauda to 8 cm. long constricted at the base;
margin grossly dentate, teeth up to 3 cm. long, with base to
2 cm. broad, the cauda occasionally with serrate margin; lateral
veins 4-5 pairs, very prominent, the basal pair reaching up to
more than half the length of the lamina; intercostals numerous,
straight to somewhat anastomosing, rather prominent. Pefioles
(4-) 7-15 (-18) cm. long, usually sparsely covered with mounted
irritant hairs. Stipules to 0.4 cm long, connate, intrapetiolar, bifid,
the two tips often as long as the lower connate part, often greatly
split by the developing female inflorescence, triangular,
chartaceous, glabrous, caducous. Inflorescences unisexual, axillary,
solitary, very rarely with woody bulbils, with minute peduncular
bracts; peduncle densely covered with very short irritant hairs
as well as long mounted ones, especially when young; the male
7-16 cm. long, 4-8 cm. broad, paniculate, well branched, in the
lower axils mainly; the female up to 32 cm. long, spike-like,
with the flowers fasciculate at intervals along the usually
unbranched peduncle, mainly in the upper axils or subterminal.
Male flowers pedicellate; perianth ca. 1 mm. long and broad,
with few to numerous short irritant hairs; tepals 5, non-corniculate,
cymbiform; stamens 5, with refiexed filaments to 2 mm. long;
pistillode ca. 0.4 mm. long and broad, obconical; interfloral bracts
up to 4.0 mm. long, triangular, irregular in number; pedicel to
0.5 mm. long, articulated at about the middle, glabrous. Female
flowers pedicellate; perianth with 4 free tepals; the two lateral

W.-L. Chew — Monograph of Laportea 143

JURAIMI DEL.

Fig. 11(a). L. macrostachya: A, twig; B, male flower; C, female flower;
D, pedicel and perianth opened; E, achene.

144 Gardens Bulletin, Singapore — XXV (1969)

\ 7 Cie

ee |

| \

|

7 *
SS ! > Y
a ag saci
JURAIMI DEL. ‘
: y
8cm 4. cm

Fig. 11(b). L. macrostachya: four leaf-forms.

W.-L. Chew, — Monograph of Laportea 145

tepals to 0.8 mm. long and broad, very broadly ovate, cordate
based, enclosing the ovary, sparse to dense with short irritant
hairs especially on the prominent rib; the dorsal one to 0.6 mm.
long, ca. 0.4 mm. broad, strongly geniculate, with numerous
short irritant hairs; the ventral ca. 0.5 mm. long, ca. 0.4 mm.
broad, slightly keeled; ovary ca. 0.8 mm. long, ca. 0.6 mm. broad,
asymmetrically ovoid; stigma linear, unbranched, slightly reflexed,
to 1.5 mm. long, interfloral bracts to 0.5 mm. long, irregular
in number; pedicel to 0.6 mm. long, simple, pubescent only
at the base. Achenes ca. 2 mm. long, ca. 1.5 mm. broad, stipitate;
stipe ca. 0.4 mm. long, on gynophore to 0.5 mm. long, ca. 0.2 mm.
broad; broadly asymmetrically ovoid, smooth, without wing,
ridge nor depression, usually entirely enclosed by the lateral tepals,
reflexed, often greatly so; pedicel up to 1.5 mm. long, simple,
not winged, fairly densely covered with very short appressed
irritant hairs; dispersed with the perianth and the greater portion
of the pedicel attached.

The four species, noted in the synonymy above, are being
assigned to this with confidence. I have seen their types and
have found them to be mere individual variants of one species.

This is probably the linking species between the two sections
of Laportea. In its habit of growth, it comes near to L. bulbifera
of sect. Laportea; but it has female inflorescences very similar
to those of L. interrupta of sect. Fleurya.

JAPAN :—Hakodate: Mohidji: Maximowicz s.n. (type of S.
macrostachya: BM & K).—Hondo: Shinano Prov.: Furuse 19740, Sept.

1951 (A)—Honshu: Mt. Tanigawa: Suzuki 398004, July 1949 (K).—
Yamagata Pref.: Atsumi village: Walker 7356, Sept. 1951 (US).

CHINA:—Hunan: Mt. Yun-schan: Handel-Mazzetti 11,135, July 1918
(C & US).—Hupei: Mt. Ku-tien: Silvestri 410 & 410A, July-Aug. 1906
(syntypes of L. longispica: FI); sine loc.: Henry 6066 & 6077 (syntypes
of L. grossedentata: BM, K & US)—Shansi: Ping Chan: Serre A.53/,
July 1929 (UPS).—Shensi: Kia-po: Giraldi 1349 (type of L. giraldiana:
FI).—Szechuan: Pao-hsing-hsien: Chu 2927, June.1936 (BM & E).—
Yunnan: Valleys of Mekong & Yangtze: Forrest 520, Sept. 1904 (type of
L. forrestii: K. & E).

BURMA:—Nam Tamai Valley: Kingdon-Ward 13259 (BM).

(12) Laportea interrupta (L.) Chew in op. cit. 200 (1965):
Letouzey in op. cit. — Urtica interrupta L. Sp. Pl. 985 (1753).
— Boehmeria interrupta (L.) Willd. Sp. Pl. 4 (1): 342 (1805).
— U. javanica Bl. Bijdr. 503 (1825). — Fleurya spicata Gaud
in Freyc. Voy. Uraine Bot. 497 (1830). — F. glomerata Gaud.
in op. cit. 498 (1830). — B. spicata (Gaud.) Endl. in Ann.
Wien Mus. 1: 165; 864 (1836). — U. sessiliflora Blco. FI.
Filip. 696 (1837), non Sw. (1785). — U. gaudichaudii Steud.
Nom. Bot. ed. 2. 2: 734 (1841). — U. spicigera Steud. op. cit.
736 (1841). — U. affinis Hk. & W.-Arn. Bot. Beech. Voy. 69
(1841). — B. javanica (Bl.) Hassk. Cat. Hort. Bog. Alt. 79
(1844). — U. capitata Bocl. Fl. Filip. ed. 2: 483 (1845). —
U. lomatocarpa Hochst. ex Steud. in Flora 33: 260 (1850). —
F. interrupta (L.) R. Wight, Ic. 6: 10, tab. 1975 (1853). —
Schychowskia interrupta (L.) Wight in Safford, Contr. U.S.
Nat. Herb. 9: 371 (1905).

146 Gardens Bulletin, Singapore —XXV_ (1969)

Diagnosis: (a) female flowers with 3-fid stigmas, (b) inflorescences
bisexual, the lateral branches greatly reduced with flowers
fasciculate at intervals along the elongated peduncle.

Distribution: Portuguese Guinea (?), Congo, East and South
Africa, Abyssinia, India, Ceylon, Japan, China through South-East
Asia and Malesia to Queensland (Australia) and the Pacific
Islands.

Monoecious annual herbs, 1 to 2 m. high. Stem woody at
the base, with few to many branches, light pubescent, with
irritant hairs towards the apex, without glandular hairs, green
with reddish-chocolate streaks near the apex, reddish-chocolate
at the base; stipular scars inconspicuous, petiolar scars conspicuous.
Lamina (5—) 8-12 (—18) cm. long, (3—) 5-7 (-12) cm. broad,
ovate to broadly ovate, chartaceous, without glandular hairs;
irritant hairs of the upper surface short, appressed, scattered,
not mounted, those of the lower surface mainly on the midrib
and veins; cystoliths elongated, those of the upper surface
scattered, those of the lower surface mainly along midrib,
veins and veinlets; base obtuse, rounded, truncate to slightly
cordate; apex acuminate, 1-1.5 cm. long; margin serrate, teeth
ca. 0.5 cm. long, with base 0.5 cm. broad; lateral veins 3-6 pairs,
prominent, the basal pair reaching to more than half the length
of lamina; intercostals numerous, quite prominent, often straight.
Petiole (3—) 5-8 (-—12) cm. long, rather densely covered with
elongated cystoliths, with evenly distributed irritant hairs,
pubescent when young, later almost glabrous. Stipules 3-5 mm.
long, connate, intrapetiolar, bifid, the two tips shorter than the
hairs on the spines, otherwise glabrous. Inflorescences bisexual,
paniculate, up to 30 cm. long, with flowers fasciculate at intervals
along elongated peduncle consequent upon great reduction of
the lateral peduncular branches, axillary, sometimes borne at
the base of the stem just above soil level, solitary; peduncular
bracts narrow triangular, ca. 1 mm. long, on the side of each
floral agglomeration; peduncle very light pubescent to glabrous,
very sparsely armed with irritant hairs. Male flowers pedicellate;
perianth 1-1.5 mm. long and broad, with few irritant hairs at
the apex; tepals 3 or 4, corniculate, triangular to cymbiform;
stamens 3 or 4, with reflexed filaments ca. 1.5 mm. long; pistillode
ca. 0.5 mm. long, as broad at the apex, obpyramidal; interfloral
bracts lanceolate, ca. 1 mm. long, variable in number; pedicel
to 1 mm. long, articulated near the apex, glabrous. Female
flowers pedicellate; perianth with 4 free tepals, the two lateral
tepals ca. 0.5 mm. long, almost enclosing the ovary, the dorsal
and ventral ones ca. 0.3 mm. long, straight, slightly geniculate,
often with few irritant hairs, otherwise glabrous; ovary ca. 0.75
mm. long, asymmetrically ovoid; stigma linear, trifid, greatly
reflexed, the central branch to 0.5 mm. long, the lateral branches
to 0.25 mm. long; interfloral bracts minute. numerous, narrowly
lanceolate; pedicel to 0.5 mm. long, slightly winged dorsi-ventrally,
articulated near the apex, with few irritant hairs at the apical half,
otherwise glabrous. Achenes ca. 1.25 mm. long and_ broad,

W.-L. Chew — Monograph of Laportea 147

Fig. 12. L. interrupta: A, twig; B, male flower; C, female flower; D, achene;
E, female perianth.

148 Gardens’ Bulletin, Singapore — XXV ( 1969)

sessile, without gynophore, asymmetrically ovoid, completely
surrounded by a narrow membraneous wing, on each lateral side
a triangular ridge enclosing a warty depression, the tepals forming
a small cup at the base, reflexed; pedicel to 1.5 mm. long,
slightly or not winged dorsi-ventrally, with few irritant hairs:
dispersed with perianth and upper portion of pedicel attached.

A species of herbaceous weeds under partial shade in abandoned
gardens, plantations and roadsides, this is the most widely
distributed one in the genus. Yet, throughout its entire range
it remains remarkably constant in its morphological attributes.
No discontinuities of any taxonomic significance can be found
between the various populations from the different parts of its
range.

The only record from Portuguese Guinea cannot be taken
seriously. There is no doubt that the specimen belongs to this
species; but as the collection is a nixtum compositum, 1 doubt
very much of our specimen belongs to the label attached.
It is likely to be a Mozambique material inadvertently mounted
with a Guinean label. I have not seen Letouzey’s record from
the Cameroun.

The nearest relative is no doubt L. ovalifolia of Africa with
which it has many characters in common (see notes there).

The specimens cited here represent but a very small proportion
of a large quantity of collections studied.

PORTUGUESE GUINEA:—Bissau: Pussube: Santo 2403 (Sheet 1),
Sept. 1947 (LISC).

CONGO :—sine loc.: Coomans s.n., March 1906 (Z).

TANGANYIKA :—Lake Nyasa: Stolz 727 (B, BM, C, G, K, S, SAM,
UPS & Z).

KENYA:—Nyika Territory: Wakefield s.n. (K).

MOZAMBIQUE:—Tuhamitonga: Torre 4085, May 1942 (LISC).

SOUTH AFRICA:—Natal: Durban: Schlechter 2791, June 1893 (BM).

ABYSSINIA:—Schimper 1471 (type of U. lomatocarpa: A, BM, G-—DC,
K, PRC & S).

INDIA:—sine loc.: Wight herb. 2680 (A, C, L, NY & S).

CEYLON:—sine loc.: Hermann s.n. (type of U. interrupta: herb.
Hermann in BM).

JAPAN :—Tokyo: Faurie 6425, Aug. 1890 (K).

CHINA:—Yunnan: Che-li Hsien: Wang 77747, Aug. 1936 (A).

BURMA :—Golden-Valley: Thein-Lwin 67, Aug. 1947 (K).

THAILAND:—North Chiengmai: Sorensen et al. 4064, July 1958
(SING).

VIETNAM:—Hue: Squires 45 (BO, BRI, E, MEL & UC).

FORMOSA :—Bankinsing: Henry 386 (K & MO).

MALAYA:—Trengganu: Yapp 344, Oct. 1899 (CGE & K).

PHILIPPINES:—Mindanao: Butuan subprov.: Weber 1090 (A, F,
NSW, UC & US).

BORNEO:—South-Borneo: Banjermassing: Motley 269 (CGE).

SUMATRA:—Bila: Rahmat-si-Toroes 2843 (MICH, NY, UC & US).

JAVA:—sine loc.: Zollinger 563 (L).—Bogor: unknwon coll. herb.
L.908-188-2807 (type of U. javanica).

LESSER SUNDA ISLANDS:—Bali: van Steenis 7743, April 1936 (A,
BO & L).

W.-L. Chew — Monograph of Laportea 149

CELEBES:—Menado: Kolonedale: Eyma 34/8, Aug. 1938 (BO, L.
PNH & U).

MOLUCCAS :—Pisang: Gaudichaud s.n. (types of F. glomerata & F.
spicata: G).

NEW GUINEA:—North-East New Guinea: Weinland 38/ (BO, BRI,
L, NSW & WRSL).

BISMARK ARCHIPELAGO :—New Britain: Talasea subdistrict: White
NGF. 10959, April 1959 (LAE).

SOLOMON ISLANDS:—sine loc. Milne 473 (K).

QUEENSLAND (AUSTRALIA):—Baron River: Bich s.n. (NSW.
77914).

NEW HEBRIDES:—Malekula: Atchin: Layard 59 (NSW).
NEW CALEDONIA :—sine loc.: Franc s.n. (herb. US. 390576).

CAROLINE ISLANDS:—Ponape Isl.: Metaranium: Kanehira 789,
Aug. 1929 (NY).

MARSHALL ISLANDS:—Jaluit: Schnee s.n. (NSW).
SAMOA :—Tau: Wray-Harris 2373-320 (BO, K & P).
FIJI:—Ovalau: Lovoni Village: Smith 7480 (US).
TONGA:—Tafahi: Hurlimann 561, Dec. 1951 (Z).
SOCIETY ISLANDS:—Tahiti: Hillebrand 162 (K).
COOK ISLANDS :—Rarotonga: Cheeseman 120.669 (K).

CORAL ISLANDS:—sine loc.: Capt Beechy s.n. (type of U. affinis:
G & K).

HAWAII :—Honolulu: Anderson s.n. 1852 (S).

(13) Laportea ovalifolia (Schumach.) Chew in op. cit. 201 (1965);
Letouzey in op. cit. 131 (1968). — Haynea ovalifolia Schumach.
Beskr. Guin. Pl. 406 (1827). — Fleurya podocarpa Wedd. in
DC. Prodr. 16 (1): 76 (1869). — F. ovalifolia (Schumach.)
Dandy in Andrews, Fl. Pl. Angl-Egypt. Sudan, 2: 277 (1952).

Diagnosis: (a)plants stoloniferous, (b) inflorescences often
directly from stolons, (c) males long, with flowers glomerulate
at intervals along the seemingly unbranched peduncle, (d) females
very small and frail (e) achenes very large, rarely reflexed, often
geocarpic.

Distribution: Sierra Leone, Liberia, Ivory Coast, Ghana, through
equatorial Africa to East Africa.

Monoecious annual stoloniferous herbs, often repent; erect
shoots up to 2 m. high. Stem slightly woody at the base, with
few branches, densely to lightly covered with short often appressed
irritant hairs, without glandular hairs; stipular scars inconspicuous,
petiolar scars conspicuous. Lamina (3—) 8-10 (—13) cm. long,
(2-) 4-6 (—8) cm. broad, ovate, chartaceous, without glandular
hairs: irritant hairs of the upper surface short, scattered, some-
times absent, those of the lower surface on midrib and veins;
cystoliths of the uper surface punctiform, scattered, those of
the lower surface on veins; base usually rounded; apex sharp
acuminate to slightly caudate, ca. 1 cm. long; margin crenate,
dentate to serrate, teeth ca. 0.5 cm. long, with base ca. 0.5 cm.
broad; lateral veins 4~7 pairs, prominent, the basal pair reaching
up to more than half the length of lamina; intercostals numerous,
straight. Petioles 5-10 (—16) cm. long, glabrous to lightly

150 Gardens’ Bulletin, Singapore —XXV (1969)

r
#

“inmm 2 JURAIMI DEL.

Fig. 13. L. ovalifolia: A, twig; B, male inflorescence; C, male flower; D,
female flower; E, achene; F, female perianth.

W.-L. Chew — Monograph of Laportea 151
covered with short appressed irritant hairs. Stipules usually 1 cm.
long, connate, intrapetiolar, bifid, the two tips often as long as
the connate part, triangular, chartaceous, with few appressed
irritant hairs, otherwise glabrous. /nflorescences unisexual, the
male paniculate, up to 50 cm. long, the lateral branches very
greatly reduced with flowers faciculate at intervals along
seemingly unbranched peduncle, axillary, usually shooting from
stolons, solitary, with minute caducous peduncular bracts; peduncle
subglabrous to densely canescent, especially the floriferous part,
with minute appressed irritant hairs; the female a simple raceme
to a sparsely branched panicle, rarely longer than5S cm., axillary,
usually produced from stolons, solitary or paired, with peduncular
bracts ca. 1 mm. long; peduncle thin and frail, glabrous to light
pubescent, with very few irritant hairs. Male flowers pedicellate;
perianth to 2 mm. long and broad; with few irritant hairsffi tepals
5, non-corniculate, narrow; stamens 5, with reflexed filaments ca.
3 mm. long; pistillode ca. 1 mm. long, as broad at the apex,
obpyramidal; interfloral bracts numerous, minute; pidecel to 3 mm.
long, articulated near the apex, glabrous. Female flowers pedi-
cellate; perianth with 4 tepals joined at the base; the two lateral
tepals to ca. 1.5 mm. long, enclosing the ovary, glabrous, the dorsal
one ca. 1.5 mm. long, not geniculate, with few irritant hairs,
Otherwise glabrous, the ventral ca. 0.6 mm: long; ovary 1.5-2
mm. long, slightly asymmetrically ovoid; stigma finear, trifid, the
upright one 2-3 mm. long, the other two ca. 2 mm. long, usually
slightly directed downward; interfloral bracts minute, narrowly
triangular; pedicel 0.5 mm. long, articulated at midpoint, almost
glabrous. Achenes to 4 mm. long, 2-3 mm. broad, substipitate to
sessile, without gynophore, very slightly asymmetrically ovoid,
almost completely surrounded by a narrow membraneous wing, on
each lateral side an ovate or elliptic ridge enclosing a warty
depression, with the tepals forming a minute cup at the base, not
or slightly reflexed, often geocarpic; pedicel to 2 mm. long, winged
dorsi-ventrally, with few irritant hairs; dispersed with perianth and
upper portion of pedicel attached.

This equatorial African species is unique in that the
inflorescences often arise from underground stolons and the
achenes often geocarpic. Add to these the characteristics of the
occasional repent habit and the large size of the achenes and
one finds this the most distinctive species in the genus.

The preceding species is its closest ally with which it has
the following characters in common: (a) lamina almost similar
in shape, size and venation, (b) the inflorescences are of the
“interrupted’’ type although the sexuality differs in the different
species, (c) stigma 3-fid, ligulate.

The varieties and subspecies maintained by various authors
are not upheld here. I find too much intergradations between
them. Well over a hundred collections have been studied,
particularly from the Republic of Congo. The following therefore
represent only a small proportion of them.

152 Gardens’ Bulletin, Singapore —XXV (1969)

SIERRA LEONE:—Southern Prov.: Mandu: Deighton 1992, July
1931 (K).

LIBERIA :—Weleo Distr.: Nyaaka: Baldwin 6205, Pune 1947 (K).
IVORY COAST:—Mt. Momy: Chevalier 21356 (K).
GHANA:—sine loc.: Thonning s.n. (—holotype of H. ovalifolia: C). —
TOGO :—Warnecke 438 (K). :
NIGERIA :—Angiana: Barter 92 (type of F. podocarpa: K).
CAMEROON :—Efulen: Bates 424, Oct. 1895 (BR, G, K & Z).
FERNANDO PO:—Moka: Boughey 24, Dec. 1951 (K).
GABON :—sine loc.: Druklage 580, Feb. 1890 (K).

CONGO :—Albert National Park: Kalonge: de Witte 8256, Oct. 1952
(BR).—Eala: Corbissier-Baland 977, March 1931 (A, B, K, MO & S).

ANGOLA :—Cazengo: Ndalatando: Gossweiler 4568B, May 1946 (B,
BM & US).—Golungo Alto: Welwitsch 6265 & 6266, Feb. 1879 (BM,
G & K).

SUDAN :—Azza Forest: Andrews A.1405, May 1939 (K).

UGANDA:—Masaka Distr.: Malabigambo Forest: Drummond &
Hemsley 4539, Oct. 1953 (BR & K).

KENYA:—Mt. Kenya: Verdcourt 2051, Dec. 1957 (UPS).
TANGANYIKA:—Mahenge Station: Schlieben 1814, Feb. 1932 (BR).

(14) Laportea peduncularis (Wedd.) Chew in op. cit. 201 (1965).
— Urtica mitis E. Meyer in Drege, Zwei Pfl. Doc. 127, 143
& 228 (1843), nom. nud., non Vell. (1835). — U. peduncularis
E. Meyer in Drege, op. cit. 143 & 228 (1843), nom. nud. —
Fleurya capensis Wedd. in Arch. Mus. Hist. Nat. Paris 9: 117,
tab. 1, fig. A, 7-8 (1856), exclu. spec. typ., non (Thunb.) Wedd.
(1854). — F. mitis Wedd. in Ann. Sci. Nat. ser. 4. 1: 183
(1854), nom. nud. — F. peduncularis Wedd. in DC. Prodr.
16 (1): 75 (1869).

Diagnosis: It distinguishes itself in the following characters:
(a) an annual herb, often creeping or scrambling, (5) inflorescences
well branched, with slender peduncles, (c) male flowers 5-tepalled,
strongly corniculate, (d) achene sessile, without wings, with a
simple stigma; (e) lamina ovate, margin serrate.

Distribution: East, South-east and South Africa.

Monoecious annual herbs, often creeping or scrambling,
generally 1.5 m. high. Stem woody at the base, rarely branched,
young parts covered with short appressed often slightly mounted
irritant hairs, without glandular hairs, glabrous when old; stipular
scars inconspicuous petiolar scars conspicuous. Lamina (5—) 9-
10 (-12) cm. long, (3—) 6-7 (—8) cm. broad, ovate, rarely trian-
gular, chartaceous, without glandular hairs; irritant hairs of upper
surface short, appressed, sparse, those of the lower surface on
midrib and veins, very rarely mounted; cystoliths of the upper
surface very minutely punctiform, often not visible, those on the
lower surface mainly elongated, intermixed with few punctiform
ones; base generally rounded, rarely cordate or truncate; apex
acuminate, 1-1.5 cm .long; margin serrate to dentate, very rarely
grossly, teeth up to 0.5 cm. long, with base to 0.5 cm. broad;
lateral veins 6-8 pairs, prominent, basal pair reaching up to half
the length of lamina; intercostals numerous, straight, prominent.

: W.-L. Chew — Monograph of Laportea 153

‘5mm JURAIMI DEL.

Fig. 14. L. peduncularis: A, twig; B, male flower; C, female flower: D,
achene; E, female perianth.

154 Gardens’ Bulletin, Singapore —XXV (1969)

Petioles (3—) 5-6 (-10) cm. long, very sparsely covered with
mounted irritant hairs to glabrous. Stipules up to 0.7 cm. long,
connate, intrapetiolar, bifid, the two tips shorter than the connate
part. triangular, chartaceous, with few short appressed irritant
hairs on the spines. /nflorescences unisexual, paniculate, sparingly
branched, axillary, solitary, with minute peduncular bracts, with
peduncle sparsely covered with short appressed irritant hairs, —
often glabrous; the male (3—) 4-6 (-9) cm. long, (2—) 3—5 (-6) —
cm. broad: the female 4-6 (-8) cm. long, (1—) 2-3 (—5) cm. broad,
with very thin peduncle i.e. smaller and thinner than the male. —
Male flowers shortly pedicellate; perianth 1—1.5 mm. long and —
broad, with few very short irritant hairs at the base, otherwise —
glabrous; tepals 5, strongly corniculate, cymbiform; stamens 5,
with reflexed filaments ca. 1.5 mm. long; pistillode ca. 0.75 mm. —
long, as broad at the apex, obpyramidal; interfloral bracts nume-
rous, minute, triangular; pedicel up to 1 mm. long, articulated
near the apex, glabrous. Female flowers pedicellate; perianth with
4 free tepals, the two lateral tepals ca. 0.75 mm. long, enclosing
the ovary, gkabrous, the dorsal one half the length of the laterals,
very slightly geniculate, glabrous, the ventral very minute; ovary
0.75-1.00 mm. long, asymmetrically ovoid; stigma linear, un-
branched, rather straight, 0.5-1 mm. long; interfloral bracts
minute, numerous; pedicel to 0.5 mm. long, strongly winged
dorsiventrally, articulated near the base, lightly covered with
minute irritant hairs. Achenes ca. 1.5 mm. long, ca. 1 mm. broad,
sessile, without gynophore, asymmetrically ovoid, only the apex
with a narrow membraneous wing, on each lateral side a triangular
ridge enclosing a warted depression, half or less covered by the
lateral tepals, reflexed; pedicel up to 2 mm. long, strongly dorsi-
ventrally winged, glabrous; dispersed with perianth and upper
portion of pedicel attached.

This species has the lamina form of L. interrupta and occa-
sionally exhibits the repent habit characteristic of L. ovalifolia.
It is, however, quite distinct from them on other essential cha-
racters. Together with the following two species, it forms a very
natural group.

Over a hundred collections of this species have been studied,
I cite here merely a small selection.

TANGANYIKA:—Morogoro Distr.: Mgeta River: Drummond &
Hemsley 1593, March 1953 (BR, K & S).—Mt. Kilimanjaro: Schlieben
4507, Jan. 1934 (B, S & Z)—Usambara: Holst 9019, Aug. 1893 (K & Z).

NYASALAND:—Cholo Distr.: Mt. Cholo: Brass 17733, Sept. 1946
(BR, K, MO, NY, UC & US). )

MOZAMBIQUE:—Lourenzo Marques: Gaza: Torre 7287, Feb. 1948
(LISC).—Chimorio: Serva de Garuzo: Barbosa 1243, March 1948 (LISC).

SOUTHERN RHODESIA:—Chirinda Distr.: Gazaland: Swynnerton
793, May 1906 (K & Z).—Umtali Distr.: Range Hill: Chase 7259, Feb.
1960 (LISC & MO).

SOUTH AFRICA:—Cape: Somerset: Burchell 3231] (A, K & L).—
Natal: Griqualand East: Tyson 768, March 1886 (A, K, SAM, UPS &
Z).—Port Natal: Drege s.n. (lectotype of F. peduncularis: G).

SWAZILAND: Mbabane District: Compton 25881, April 1956 (NBG)
& 28689, March 1959 (NBG).

W.-L. Chew — Monograph of Laportea 155:

(15) Laportea caffra Chew, spec. nov.
Diagnosis: L. pedunculari affinis, a qua multo pusillioribus,

inflorescentiis brevioribus cum floribus perpaucis, tepalis masculis.
haud corniculatis differt.

Distribution: South Africa.

Monoecious annual herbs less than 50 cm. tall. Stem woody
at the base, branching rather broadly, well armed with irritant
hairs, otherwise quite glabrous. Lamina ca. 2 cm. long, 1-2 cm.
broad, broad ovate to triangular, chartaceous, well armed with
irritant hairs, the upper surface with punctiform cystoliths, the
lower with elongated ones; base rounded to truncate; apex acute;
margin serrate, teeth to 2 mm. long; lateral veins ca. 4 pairs, the
basal pair reaching to half the length of the lamina; intercostals.
few, inconspicuous. Petioles to 1 cm. long, armed with irritant
hairs. Stipules minute, puberulous, caducous. /nflorescences uni-
sexual, poorly branched dichasia, axillary, solitary to paired, to
5 cm. long, 1 cm. broad at the most peduncle sparse with irritant
hairs. Male flowers pedicellate; perianth ca. 1.5 mm. long and
broad, subglabrous; tepals 5, non-corniculate; stamens 5, with
slightly reflexed filaments; pistillode small; pedicels ca. 1 mm.
long, glabrous. Female flowers pedicellate; perianth ca. 0.75 mm.
long, 0.6 mm. broad, enclosing the ovary, with few irritant hairs,
the dorsal one not geniculate; ovary ca. 0.7 mm. long, ovoid;
stigma linear, unbranched, 0.6 mm. long; interfloral bracts minute;
pedicel ca. 0.4 mm. long, slightly winged dorsiventrally, articulated
near the base, subglabrous. Achenes ca. 1.5 mm. long, 1.25 mm.
broad, sessile, without gynophore, asymmetrically ovoid, without
wings, on each lateral side a triangular ring enclosing a warted
depression, half covered by the lateral tepals, slightly reflexed;
pedicel to 1 mm. long, dorsiventrally winged, glabrous; dispersed
with perianth and upper portion of pedicel attached.

In size and habit of growth, this species recalls L. ruderalis
of the Pacific Islands. Both are small low branching herbs. How-
ever, here the superficial similarity ends.

The natural alliance of this species seems to be the pedun-
cularis-mooreana group, also of Africa. As mentioned in the
diagnosis above, my new species is closest to L. peduncularis
from which it differs in three points. The specific epithet is derived
from the adjective caffer which means South African.

CAPE OF GOOD HOPE:—Outeniekwaberge Distr.: near George:

R. Schlechter 2331, 14-III-1893 (holotype in Z, isotypes in A, BR, PRC,
S, SAM & UPS).

NATAL:—near Durban: A. Hapstroem s.n., August 1938 (S).

156 Gardens’ Bulletin, Singapore —XXV (1969)

Fig. 15. L. caffra: A, twig; B, achene; C, female inflorescence.

| W.-L. Chew — Monograph of Laportea 157

(16) Laportea mooreana (Hiern) Chew in op. cit. 201 (1965);
Letouzey in op. cit. 127 (1968). — Adicea mooreana Hiern,
Cat. Afr. Pl. Welw. 1 (4): 991 (1900). — Pilea mooreana
(Hiern) K. Sch. in Just, Jahresber. 28 (1): 463 (1900). —
Fleurya mooreana (Hiern) Rendle in Prain, Fl. Trop. Afr. 6
(2): 250 (1917). — F. funigera Mildbr. in Notizbl. Bot. Gard.
Berlin 8: 278 (1923). — F. urophylla Mildbr. in op. cit. 279
(1923).

Diagnosis: It distinguishes itself in (a) male perianth 4-tepalled,
strongly corniculate. (5) dorsal tepal of female flowers strongly
geniculate and greatly separated from the other tepals at fruit,
(c) lamina triangular to deltate, grossly dentate with teeth to 1.5
cm. long.

Distribution: East, Central and West Africa.

Monoecious annual herbs, generally 1 to 2 m. high. Stem woody
at the base, with few branches, usually covered with long mounted
irritant hairs, otherwise glabrous, without glandular hairs; stipular
scars inconspicuous, petiolar scars conspicuous. Lamina (5—) 8
15 (-19) cm. long, (3—) 7-12 (—15) cm. broad, triangular, broadly
triangular to deltate, chartaceous, without glandular hairs; irritant
hairs of upper surface short, rarely mounted, scattered, those of
the lower surface on midrib and veins, often strongly mounted;
both surfaces fairly densely covered with minutely punctiform
cystoliths; base generally truncate; apex caudate, (I—) 1.5-3 (-4)
cm. long; margin grossly dentate, teeth to 1.5 cm. long with base
to 0.5 cm. broad: lateral veins 5-8 pairs, prominent, the basal
pair reaching up to more than half the length of lamina; inter-
costals numerous, straight, fairly conspicuous. Petioles (3—) 5-15
(-17) cm. long, the distal part usually densely covered with long
mounted irritant hairs, otherwise glabrous. Stipules to 1 cm. long,
connate, intrapetiolar, bifid, the two tips shorter than the lower
connate part, triangular, chartaceous, with few short appressed
irritant hairs on the spines. /nflorescences bisexual, paniculate, to
44 cm. long and 10 cm. broad, axillary, solitary; peduncular
bracts minute, caducous; peduncle sparsely covered with mounted
irritant hairs, otherwise glabrous. Male flowers pedicellate; perianth
1 to 1.5 mm. long and broad, with few irritant hairs at the apex;
tepals 4, strongly corniculate, cymbiform; stamens 4, with reflexed
filaments ca. 1.5 mm. long; pistillode ca. 0.5 mm. long, as broad
at the apex, obpyramidal; interfloral bracts numerous, minute,
narrowly triangular; pedicel to | mm. long, articulated near the
apex, glabrous. Female flowers pedicellate; perianth with 3 or 4
free tepals, the two lateral tepals to 0.75 mm. long, enclosing
the ovary, glabrous, the dorsal one to 0.7 mm. long, geniculate,
with 1 or 2 mounted irritant hairs, the ventral ca. 0.25 mm. or
less in length, often absent; ovary ca. 0.5 mm. long. asymmetrically
ovoid; stigma linear, unbranched, slightly reflexed, ca. 0.3 mm.
long; interfloral bracts minute, numerous, narrowly triangular,
pedicel to 0.5 mm. long, slightly winged dorsi-ventrally, articulated
near the apex, lightly covered with minute irritant hairs. Achenes
1-1.5 mm. long, ca. | mm. broad, substipitate to sessile, on

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Fig. 16. L. mooreana: A, twig; B, mounted irritant hair; C, male flower;

, achene.

F

D, achene with perianth; E, female flower;

W.-L. Chew — Monograph of Laportea 159

minute gynophore, asymmetrically ovoid, partly surrounded by
a narrow membraneous wing, on each lateral side a triangular
ridge enclosing a warty depression, more than half covered by
the lateral tepals, reflexed; pedicel up to 2 mm. long, slightly
winged dorsi-ventrally. lightly covered with minute irritant hairs;
dispersed with perianth and upper portion of pedicel attached.

Though closely related to the two preceding species, L.
mooreana approaches L. grossa in a few characters. In both, the
lamina are triangular and borne on petioles armed with rather
long mounted irritant hairs, although in the latter, the teeth of
the lamina are much larger and fewer in number. Also, the dorsal
tepal of the female perianth are rather strongly geniculate in
both species.

This species is reported to be common at about the 1,500
metre-contour.

UGANDA :—Western Prov.: Toro Distr. Taylor 2486, Dec. 1934 (BM
& MO).—Budongo Forest: Loveridge 121, Nov. 1938(A, K& MO).

TANGANYIKA :—Mahenge: Umebung: Schlieben 1856 (BR, K, S
& Z).

MOZAMBIQUE:—Zambezia: Torre 5179, April 1934 (LISC).—
Chipende: Lemos 8035, 8401 & 19881] (K).

NIGERIA :—Northern Div.: H.V.L.P. 739, Sept. 1930 (K).

BELGIUM CONGO:—Albert National Park: de Witte 7844, 7984 &
12084 (BR).—Beni & Irumu: Louis 5534, Aug. 1937 (BR, C, MO & NSW).

ANGOLA :—Androngo: Welwitsch 6256, 6273, 6276 & 6295 (BM &
K).—Capir: Gossweiler 9998 Jan. 1933 (US) & 9999, Jan. 1933 (LISJC).

(17) Laportea grossa (Wedd.) Chew in op. cit. 200 (1965). —
Fleurya grossa Wedd. in Ann. Sci. Nat. ser. 4. 1: 183 (1854),
nom. nud., et in Arch. Mus. Hist. Nat. Paris 9: 119, tab. 1A,
fig. 1-3 (1856), basionym, et in DC. Prodr. 16 (1): 76 (1869);
Brown in Th.-Dyer, Fl. Cap. 5 (2): 545 (1925).

Diagnosis: Close to L. mooreana and L. peduncularis. From
the former, it differs in the male flowers being 5-merous and the
tepals non-corniculate. From the latter it is distinguishable by
the grossly toothed lamina, the smaller inflorescences and the
dorsal tepal of the female flower being strongly geniculate.

Distribution: South-east and South Africa.

Monoecious annual herbs less than 1 metre high. Stem slightly
woody at the base, very rarely branched, young parts sparsely
covered with long mounted irritant hairs, otherwise quite glabrous,
petiolar scars quite prominent. Lamina (4-) 5-8 (—10) cm. long
and broad, triangular, chartaceous, without glandular hairs, both
surfaces sparsely armed with long mounted irritant hairs, and
covered with minutely punctiform cystoliths; base truncate; apex
long acuminate, acumen 1-—1.5 cm. long; margin grossly toothed,
teeth to 1 cm. long and equally broad at the base; lateral veins
3 to 4 pairs, the basal pair reaching to half the length of the
lamina; intercostals few, somewhat anastomosing. Petioles (3-)
4-7 (-10) cm. long, sparsely covered with long mounted irritant
hairs, otherwise quite glabrous. Stipules minute, connate, intra-
petiolar, caducous. /nflorescences unisexual, paniculate, fairly well

160 Gardens’ Bulletin, Singapore — XXV ( 1969)

scm

JURAIMI DEL.

Fig. 17. L. grossa: A, twig; B, matured female flower.

W.-L. Chew — Monograph of Laportea 161

branched, axillary solitary usually; the females up to 10 cm. long,
5 cm. broad, usually borne in the upper axils; the males shorter
and narrower, borne in the lower axils; peduncle subglabrous,
with few mounted irritant hairs. Male flowers shortly pedicellate;
perianth to 2 mm. long and broad, glabrous; tepals 5, very rarely
4, non-corniculate, dense with minute punctiform cystoliths, cym-
biform; stamens 5, very rarely 4, with reflexed filaments up to
2 mm. long; pistillode ca. 1 mm. long, ca. 0.75 mm. broad at
the apex, obconical; interfloral bracts ca. 0.3 mm. long, triangular,
2-3 to each flower; pedicel ca. 0.6 mm. long, articulated near
the apex, glabrous. Female flowers pedicellate; perianth with 4
free tepals, the two lateral tepals to 1 mm. long and 0.6 mm.
broad, completely enclosing the ovary, glabrous, the dorsal one
ca. 0.6 mm. long, strongly geniculate, glabrous, the ventral ca.
0.4 mm. long, somewhat triangular; ovary ca. 0.8 mm. long, ca.
0.5 mm. broad, asymmetrically ovoid; stigma linear, unbranched,
reflexed, to 0.4 mm. long; interfloral bracts minute, not numerous;
pedicel to 1 mm. long, slightly winged dorsi-ventrally, articulated
near the apex, glabrous. Achenes ca. 1.75 mm. long, ca. 1.5 mm.
broad, sessile, without gynophore, asymmetrically ovoid to slightly
semi-circular, without wings, smooth and without warts, half
covered by the lateral tepals, reflexed; pedicel to 2 mm. long,
curved and thickened at the apex.

The shape of lamina and the form of irritant hairs in this
species recall those of L. mooreana with which it has some rela-
tionship. In both, the lamina is triangular but in this species, the
marginal teeth are larger and fewer in number and there are fewer
lateral veins. Also the lamina as a whole is much smaller than
that of L. mooreana.

MOZAMBIQUE:—Niassa Distr.:—Cabo Delgado: Torre & Paiva

11723, April 1964 (LISC).—Mt. Chicungulo: Torre & Paiva 10967, March
1964 (LISC).

SOUTH AFRICA:—Cape Prov.: Nahoon River: Galpin 5681, April
1900 (K). Pondoland: Drege 9839 (K). Uitenhage: Zeyher 3865 (lectotype
in P; diplicates in FI & SAM).—Natal Prov.: Inanda: Wood 1236 (K &
SAM). Oribi Gorge: McClean 465, April 1937 (MO) & 28475 (K).

(18) Laportea lanceolata (Engl.) Chew in op. cit. 201 (1965). —
Fleurya lanceolata Engl. Abh. Preuss. Akad. Wiss. 37 (1894) &
Pflanzenw. Ost.-Afr. 100: 163 (1895).

Diagnosis: It distinguishes itself from all the species of sect.
Fleurya in (a) plants mainly dioecious rarely monoecious, (5) the
pedicel of the female flower not elongating at fruit, (c) achenes
slightly reflexed, at the lateral sides on ovate ridge enclosing
a smooth depression, (d) male flowers 5-tepalled, smooth, (e) in-
florescence at most as long as petiole.

Distribution: Central and East Africa.

Dioecious, rarely monoecious, annual herbs to 1 m. high. Stem
slightly woody at the base, sparingly branched, glabrescent to
well armed with fairly long thin irritant hairs, especially at the
apex, Old parts becoming glabrous, without glandular hairs:
petiolar scars very conspicuous, stipular scars not so. Lamina

162 Gardens’ Bulletin, Singapore — XXV (1969)

JURAIMI DEL.

Fig. 18. L. lanceolata: A, twig; B, male flower; C, female flower; D,
achene.

W.-L. Chew — Monograph of Laportea 163°

(5-) 8-15 (-20) cm. long, (2-) 3-6 (-7.5) cm. broad, narrowly’
ovate to elliptic, chartaceous, without glandular hairs; irritant hairs:
of the upper surface evenly distributed, those of the lower surface:
more numerous, mainly on midrib and veins, especially the basal;
cystoliths of the upper surface dense, minutely punctiform, those
of the lower surface somewhat elongated, also dense; base cuneate
to rounded; apex long acute to acuminate, 0.5—-2.5 cm. long;
margin fairly broadly serrate, teeth to 0.75 cm. long, with base to:
1 cm. broad; lateral veins 4-7 pairs, prominent, the basal pair
more prominent, reaching to slightly more than half the length.
of lamina, usually straighter than the other veins; intercostals
numerous, not straight, not very prominent. Pefioles (2—-) 4-6 (-9):
cm. long, with irritant hairs mainly at the apex. Stipules to 0.3 cm..
long, connate, intrapetiolar, deeply bifid, the two parts joined
only at the base, chartaceous, glabrous, with elongated cystoliths,
early caducous. /nflorescences unisexual, short paniculate, branched
fairly equally, axillary, solitary or paired, with minute early cadu-
cous peduncular bracts, peduncle well armed with short irritant
hairs; the male rarely more than 2 cm. long and broad, the female
to 6 cm. long and 4 cm. broad. Male flowers pedicellate; perianth
ca. 2 mm. long and broad, glabrous; tepals 5, non-corniculate,
broad cymbiform; stamens 5, with reflexed filaments to 3 mm.
long; pistillode up to 1 mm. long and broad, obconical; interfloral
bracts minute, not numerous; pedicel to 2 mm. long, articulated
near the apex, usually with short irritant hairs. Female flowers
pedicellate; perianth with 4 free tepals, well armed with fine
irritant hairs; the two lateral tepals to 1 mm. long, 0.6 mm. broad.,,.
broadly ovate, enclosing the ovary, fringed; the dorsal and the
ventral ones ca. 0.4 mm. long and broad, very broad ovate, not
geniculate; ovary ca. 1 mm. long. 0.6 mm. broad, asymmetrically
ovoid; stigma linear, unbranched, to 1 mm. long, slightly bent;
interfloral bracts to 0.3 mm. long, triangular, very few; pedicel
to 0.4 mm. long, articulated at about midpoint, frequently gla-
brous. Achenes ca. 1.4 mm. long, ca. 1 mm. broad, substipitate to:
sessile, without gynophore, asymmetrically ovoid, only the bottom
corner with a very narrow membraneous wing, on each lateral
side an ovate ridge enclosing a smooth depression, completely
covered by the lateral tepals, slightly reflexed; pedicel ca. 0.5 mm.
long, often 0.4 mm., simple, not winged, glabrous to very slightly
armed with fine irritant hairs; dispersed with only the perianth
attached.

This very distinctive species is perhaps a remote relative of
L. aestuans with which it has a few points in common such as
lamina shape and margin. The erect nature of the female flowers.
brings this species also close to L. ovalifolia.

CONGO :—Semliki Valley: Ituri: Humbert 9001], July 1929 (B & BR).

KENYA:—Marengi Forest Reserve: Mrima Hill: Verdcourt 1940,
Sept. 1957 (BR).

TANGANYIKA :—Kaogwe Distr.: Faulkner 1033 (BR, K & S).—
Kimboso Distr.: Bruce 1030 (BM, BR & K).—Lushoto Distr.:
Drummond & Hemsley 3187, July 1953 (BR, K, LISC & S)—Mahenge
Station: Schlieben 1546 & 1547, Dec. 1931 (BM, K, S & Z).—Morogoro:
Haarer 1859, June 1930 (K); Cchlieben 3802 & 3904, May 1933 (B, BR,
S & Z)—Usambara: Holst 8833, Aug. 1898 isotype of F. lanceolata: K)

164 Gardens’ Bulletin, Singapore — XXV (1969)

(19) Laportea aestuans (L.) Chew in op. cit. 200 (1965); Letouzey
in op. cit. 121 (1968). — Urtica aestuans L. Sp. Pl. ed. 2. 1397
(1763). — U. divaricata Forssk. Fl. Aegypt. Arab. 160 (1775),
non L. (1753). — U. hirsuta Vahl, Symb. Bot. Pl. 1: 77 (1790).
— U. latifolia Rich. Act. Soc. Hist. Nat. Paris 1: 113 (1792). —
U. nemorosa Kunth in Bonpl. & Humb. Nov. Gen. & Sp. PI.
2: 42 (1817). — U. divergens Mey. Prim. Fl. Esseg. 264 (1818).
— U. caravellana Schrank, Pl. Rar. Hort. Monac., tab. 82
(1819). — Fleurya cordata Gaud. in Freyc. Voy. Uraine Bot.
498 (1830). — F. petiolata Decne. in Nouv. Ann. Mus. Hist.
Nat. Paris, ser. 3. 3: 490 (1834) & Herb. Timor. 162 (1835). —
U. cordata (Gaud.) Steud. & U. petiolata (Decne.) Steud. Nom.
Bot. ed. 2. 2: 734, 735 (1841). — U. schimperiana Hochst. ex
Steud. in Flora, 33: 259 (1850). — F. caravellana (Schrank)
Wedd. in Ann. Sci. Nat. ser. 3. 18: 204 (1852). — F. glandulosa

~ Wedd. in op. cit. 205 (1852). — F. aestuans (L.) Mig. in Martius,
Fl. Braz. 4 (1): 196 (1853). — F. ingrata Mig. in Zoll. Syst.
Verz. Ind. Archip. 103, 106 (1854). —- U. tuberculata Anderss.
in Vet. Akad. Handl. Stockh. 1853: 159 (1855). — F. lurida
Bl. Mus. Bot. Lugd. Bat. 2: fig. 21 (1856). — F. perrieri
Leandri in Ann. Mus. Col. Marseille, ser. 6. 7-8: 16, fig. HI
(1950). — L. bathiei Leandri in Fl. Madagascar, 56: 10 (1965).

Diagnosis: Easily distinguished by the following combination of
characters: (a) monoecious annual herbs with glandular hairs,
‘((b) lamina ovate, chartaceous, the margin dentate, teeth ca. 0.5 cm.
long, (c) inflorescences bisexual, widely branched racemes, (d)
male perianth 4 or 5 tepalled, with few long glandular hairs.

Distribution: ‘Tropical America, West Indies, Tropical Africa,
Madagascar, Arabia, India, Sumatra, Java and Lesser Sunda
Islands.

Monoecious annual herbs, generally 1 m. high, rarely to 2 m.
Stem usually fleshy, slightly woody at the base, with few branches,
glabrescent to densely covered with simple irritant hairs and long
glandular hairs; stipular scars inconspicuous, petiolar scars usually
quite conspicuous. Lamina (3—) 10-15 (—30) cm. long, (2-) 8-12
(—22) cm. broad, ovate to very broad ovate, chartaceous, usually
without glandular hairs; irritant hairs scattered on upper surface,
more and mainly on veins of the lower surface; cystoliths of the
upper surface punctiform, densely and well dispersed over the
surface, those of the lower surface elongated and concentrated
on veins;.base rounded, truncate to cordate; apex acuminate, often
sharply, 1—-1.5 cm. long; margin dentate, the teeth less than 0.5 cm.
long with base ca. 0.5 cm. broad; lateral veins 5—8 pairs, rather
prominent, the basal pair reaching up to more than half the length
of lamina; intercostals numerous, rather straight. Petioles (2-) 5-15
(—20) cm. long, usually densely covered with irritant as well as
long glandular hairs, very rarely glabrescent. Stipules up to 1 cm.
long, connate, intrapetiolar, bifid, the two tips usually as long as
the lower connate part, triangular, chartaceous, with few short
irritant hairs on the spines. /nflorescences bisexual, paniculate, up

165

W.-L. Chew — Monograph of Laportea

|
a
5
4
em
=
ry

Fig. 19. L. aestuans: A, twig; B, male flower; C, matured female flower;

D, perianth; E, achene.

166 Gardens’ Bulletin, Singapore — XXV (1969)

to 20 cm. long, 10 cm. broad, axillary, those of lower axils very
rarely all-male, solitary, rarely paired; peduncular bracts absent;
peduncle usually well covered with irritant hairs and long to very
long glandular hairs. Male flowers pedicellate; perianth up to
1.5 mm. long and broad, with few long glandular hairs at the
apex, at most with very few irritant hairs; tepals 4 or 5, slightly
corniculate, cymbiform; stamens 4 or 5, with reflexed filaments up
to 1.5 mm. long; pistillode ca. 0.25 mm. long, as broad at the
apex, obpyramidal; interfloral bracts very numerous, minute, nar-
rowly ovate; pedicel up to 1 mm. long, articulated near the apex,
glabrous. Female flowers pedicellate; perianth with 4 free tepals,
the two lateral tepals ca. 0.5 mm. long, enclosing the ovary,
glabrescent, the dorsal one ca. 0.25 mm. long, geniculate, bearing
3-5 glandular hairs, the ventral very minute; ovary ca. 0.5 mm.
long, asymmetrically ovoid; stigma linear, unbranched, slightly
reflexed, ca. 0.3 mm. long; interfloral bracts minute, numerous,
narrowly triangular; pedicel up to 0.5 mm. long, usually slightly
winged dorsiventrally, articulated near the apex, glabrous. Achenes
1-2 mm. long and broad, shortly stipitate, on very small gyno-
phore, asymmetrically ovoid, partly surrounded by a narrow
membraneous wing, on each lateral side a triangular ridge en-
closing a warty depression, more than half covered by the lateral
tepals, reflexed; pedicel up to 2 mm. long, often winged dorsi-
ventrally, glabrous; dispersed with perianth and upper portion of
pedicei attached.

This occurs as herbaceous weeds prefering semi-shaded areas
along roads, in abandoned gardens and in plantations and is
almost as widely distributed as ZL. interrupta, another weedy
species. Unlike this species, however, L. aestuans varies quite
considerably in lamina form and size throughout its range. Never-
theless, intergradations between the various populations are so
numerous that it is impossible to recognise any infra-specific groups.

In so far as the inflorescence structure is concerned, this species
approaches L. ruderalis of the Pacific Islands; but these two differ
in floral details.

I have examined an enormous quantity of collections of this
species. The following is but a very small fraction of the total
dealt with.

GALAPAGOS ISLANDS :—Abingdon Island: Stewart 1172, Sept. 1905
(A, MO & US).

MEXICO :—Sinaloa State: La Noria: Mexia 317, Oct. 1925 (MO & UC).

GUATEMALA :—Izabal Dept.: Standley 73090 (F).

PANAMA:—Canal Zone: Ancon: Pittier 2751, Feb. 1911 (US).

CUBA:—St. Elena: Poeppig s.n. (PRC).

JAMAICA :—Burlington: Harris 5981, Nov. 1895 (Z).—New Market:
Britton 1415, Sept. 1907 (NY).

HISPANIOLA :—Haiti: Ekman 2667, Nov. 1924 (K & S).—Dominican
Repub.: Ekman 15919, Aug. 1930 (B, S & US).

COLOMBIA:—Bolivar Div.: Turbaco: Killip & Smith 14346, Nov. 1926
(A, NY & US).

PERU :—Iquitos: Asplund 14147, Oct. 1940 (US); Williams 1340, July
1929 (A, F & US).

W.-L. Chew — Monograph of Laportea 167

VENEZUELA :—Caracas: Bailey & Bailey 375, Dec. 1920 (US).—
Merida State: Fendler 1242 (K).

BRAZIL :—Para: Belem: Dahlgren & Sella 259, March 1929 (F).—
Bahia: Barra: Spruce 1507, May 1851 (CGE & K).

LIBERIA :—Central Prov.: Gbarnga: Traub 258, Aug. 1951 (BM, BR,
‘C, F, MO, UPS & US).

NIBERIA :—Southern: Lagos: Dalziel 1173, Oct. 1917 (C & K).

BELGIAN CONGO :—Equateur Prov.: Bikoro: Goosens 2315, Dec.

(BR).—Orientale Prov.: Yambuya: Louis 11334 & 12334, Nov. 1938

MOZAMBIQUE :—Niassa Distr.: Malema: Torre & Paiva 10562, Feb.
1964 (LISC.)

ar toto inne Prov.: Virunga Mts.: Taylor 1943, Nov. 1934
MADAGASCAR :—Ankarana: Humbert & Cours 32498 & 32519,
Jan.-Feb. 1960 (P & SING).
ARABIA :—Agara: Schweinfurth 1017, Feb. 1889 (P).
INDIA :—Shevaroi Hills: Perrottet 93 (K).

SUMATRA:-—Batang Baroes: Teysmann H.B. 747 (BO).—wNorth:
Siberie Island: Docters van Leeuwen 5327, April 1921 (BO).

JAVA :—West Java: P. Maeira: van Steenis 11247, June 1939 (BO).—
Batavia: Backer 31729, 31730, 31731 & 31732 (BO).

LESSER SUNDA ISLANDS:—Timor: Baumata: Bloembergen 3552,
March 1939 (L).

(20) Laportea ruderalis (Forst. f.) Chew in op. cit. 201 (1965). —
Urtica ruderalis Forst. f. Fl. Ins. Austr. Prodr. 66 (1786). —
Fleurya paniculata Gaud. in Frey. Voy. Uraine Bot. 497 (1830).
— Boehmeria paniculata (Gaud.) Endl. in Ann. Wien Mus. 1:
165. no. 863 (1836). — Schychowskia ruderalis (Forst. f.) Endl.
in op. cit. 187. tab. 13 (1836). — U. paniculata (Gaud.) Steud.
Nom. Bot. ed. 2. 2: 735 (1841). — F. ruderalis (Forst. f.) Wedd.
in Ann. Sc.. Nat. ser. 4. 1: 182 (1854); Wedd. in DC. Prodr.
16 (1): 74 (1869); St. John & Mason in Pacific Sci. 7: 166 (1953).

Diagnosis: It distinguishes itself in (a) plants completely glabrous
and quite without irritant hairs, (b) pedicels of female flowers not
winged, (c) stigma somewhat brush-like, not ligulate.

Distribution: Christmas Island, Philippines, Lesser Sunda Islands
through East Malesia to Melanesia, Micronesia and Polynesia.

Monoecious annual herbs, usually less than 60 cm. high, com-
pletely glabrous and quite without irritant or glandular hairs. Stem
slightly woody at the base, with few to many branches, especially
the small plants on coral rocks, almost completely covered with
elongated cystoliths, often reddish chocolate in colour; stipular and
petiolar scars usually very inconspicuous. Lamina (1.5—) 4-7 (—10)
cm. long, (0.9—-) 2-5 (-6) cm. broad, ovate, very rarely broadly
ovate, chartaceous; cystoliths of the upper surface very minutely
punctiform, dense, those of the lower surface elongated, mainly
on the veins and veinlets; base cuneate to rounded, very rarely
cordate; apex acute, rarely acuminate, usually less than 0.75 cm.

168 Gardens Bulletin, Singapore — XXV (1969):

Imm JURAIMI DEL.

Fig. 20. L. ruderalis: A, twig; B, male flower; C, female flower; D, achene;
E, female pedicel and perianth.

W.-L. Chew — Monograph of Laportea 169

long; margin usually crenate, teeth ca. 0.3 cm. long, with base 0.4.
cm. broad; lateral veins 3—7 pairs, often rather faint, the basal pair
reaching to more than half the length of lamina; intercostals
numerous, faint, rarely straight. Petioles (0.5—) 3-6 (—9) cm. long,
usually densely covered with elongated cystoliths. Stipules 1-2 mm.
long, connate, intrapetiolar, bifid, the two tips shorter than the
connate part, cymbiform, chartaceous, very caducous. Inflore-
scences unisexual, rarely bisexual, paniculate, well branched,.
axillary, solitary, rarely paired, with minute caducous peduncular
bracts, with peduncle completely covered with elongated cystoliths;
male up to 7 cm. long and 3 cm. broad, female to 10 cm. long and
3.5 cm. broad. Male flowers pedicellate; perianth 1-1.5 mm. long
and bread; tepals 4, often strongly corniculate, cymbiform; stamens
4, with reflexed filaments ca. 1.5 mm. long; pistillode ca. 0.5 mm.
long, as broad at the apex, obconical; interfloral bracts few, very
minute; pedicel to 1.5 mm. long, articulated near the apex, glabrous.
Female flowers pedicellate; perianth with 4 free tepals, the two
lateral tepals less than 0.4 mm. long, enclosing the ovary, glabrous;
the dorsal one ca. 0.4 mm. long, very strongly geniculate, glabrous;
the ventral very minute, less than 0.25 mm. long; ovary ca. 0.3
mm. long, asymmetrically ellipsoid; stigma somewhat capitate or
brush-like, not linear, unbranched, slightly reflexed, ca. 0.1 mm.
long; interfloral bracts numerous, to 0.4 mm. long; pedicel ca.
0.3-0.5 mm. long, not winged, articulated near the apex. Achenes
1.25 mm. long, ca. 1 mm. broad, stipitate, with stipe 0.2 mm. or
less, on gynophore ca. 0.5 mm. long asymmetrically ovoid, half
surrounded by a narrow membraneous wing, on each lateral side
an ovate ridge enclosing a warty depression, more than half
covered by the lateral tepals, greatly reflexed; pedicel to 2 mm.
long, simple, not winged; dispersed with the perianth and upper
portion of pedicel attached.

A small herb completely devoid of irritant hairs, this has the
inflorescence of L. aestuans but the female flowers of L. cuneata
with which our species is closely related.

I have seen the holotype of F. paniculata and am quite satisfied
that it belongs here.

CHRISTMAS ISLAND (Indian Oceon):—Flying Fish Cove: Andrews
134 (K); Ridley 155 (K & SING).

PHILIPPINES :—Cavilli Island: Merrill 402, Sept. 1910 (FI, K, M, NY,
U, WRSL & Z)—Luzon: Bulacan Prov.: Angat: Ramos 21725, Sept.
1913 (L).

LESSER SUNDA ISLANDS:—Kangean Archip.: Sepandjang: Backer
29013, April 1919 (BO).

CELEBES:—Boetan: Boer bb. 2235, March 1921 (BO).—Kajoedi:
Docters van Leeuwen 1327, May 1913 (BO & UV).

MOLUCCAS :—Aroe Isl.: Buwalda 4878, May 1938 (A, BO, K & L).
Babar Isl.: Borssum Waalkes 3108, March 1956 (L).

NEW GUINEA:—Jappen: Biak: Aet & Idjan 894, Sept. 1939 (A, BO,.
K & L).—Papua: Misima Island: Brass 27604, Aug. 1956 (L, LAE & US)..

170 Gardens’ Bulletin, Singapore — X XV (1969)

BISMARCK ARCHIPELAGO :—Mussau: Uni: Koie & Olsen 1282,
Jan. 1962 (C & SING).

MELANESIA :—Solomon Islands: Owa Riki Island: Brass 3076, Oct.
1932 (A, BO, BRI & L).

MICRONESIA :—Caroline Islands: Nama Island: Anderson 907, Dec.
1949 (L & US).—Marshall Islands: Likiep Atoll: Fosberg 27004, Aug.
1946 (L & NY).—Marianas: Rota: Gaudichaud 20 (type of F. paniculata:
‘G & P)—Gilbert Islands: Onotoa: Moul 8336, Aug. 1951 (A & UC).

POLYNESIA :—Line Islands: Fanning Isl.: Arundel 8 (A & K); Ball
10, July 1922 (BO, BRI & K).—Society Island: Forster s.n. (type of
U. ruderalis: BM, C & P)—Samoa: Powell 40, Sept. 1877 (K).—Tuamotu:
Raroia Atoll: Doty & Newhouse 11005, June 1952 (US).

(21) Laportea cuneata (A. Rich.) Chew in op. cit. 200 (1965). —
Urtica cuneata A. Rich. in Ramon de la Sagra, Hist. Fis. Isla
Cuba II. Bot. 11: 224 (1850). — Fleurya umbellata Wedd. in
Ann. Sci. Nat. ser. 3. 18: 204 (1852). — F. glomerata Griseb.
Cat. Pl. Cuba 58 (1866), non Gaud. (1830). — F. cuneata (A.
Rich.) Wedd. in DC. Prodr. 16 (1): 73 (1869).

Diagnosis: It distinguishes itself from all the other species by the
following unique characters (a) the male infforescence apedun-
culate, with the flowers at the base of the female inflorescence, (b)
the female inflorescence a simple or compound corymb, (c) lamina

often trilobed with strongly serrated margin, (d) branches often
opposite, arising near the base of the stem.

Distribution: Cuba & Hispaniola.

Monoecious annual herbs to ca. 60 cm. high, rarely repent. Stem
‘slightly woody at the base, with few branches often opposite and
arising near the base, the apex with numerous short appressed
irritant hairs, the rest with fewer but long rigid ones, otherwise
glabrous, quite densely covered with elongated cystoliths, without
glandular hairs, stipular and petiolar scars not conspicuous.
Lamina (0.5—) 1.5—-3.5 (—5.5) cm. long, (0.3—) 1-3 (-5) cm. broad,
‘usually broadly ovate, rarely narrowly ovate or rhombic, often
trilobed, thinly chartaceous, without glandular hairs; irritant hairs
very short, minute and appressed to as long as 3 mm., evenly
distributed on the upper surface of lamina, on the lower surface
mainly on the midrib and veins; cystoliths dense on both surfaces,
those of the upper surface punctiform, often very shortly elongated,
rarely visible, those of the other surface elongated, easily visible;
base narrowly to broadly cuneate to rounded; apex acute to
acuminate, very rarely caudate, up to 0.5 cm. long; margin usually
serrate, rarely smooth or grossly serrate, teeth up to 0.6 cm. long
with base ca. 0.5 cm. broad; lateral veins usually less than 5
pairs, usually faint, the basal pair reaching to slightly more than
half length of lamina; intercostals numerous, anastomosing, very
faint. Peiioles (0.3—) 1-3 (—7) cm. long, usually lightly armed with
long irritant hairs, densely covered with elongated cystoliths.

W.-L. Chew — Monograph of Laportea 171

JURAIMI DEL.

Q

Fig. 21. L. cuneata: A, twig; B to P, different leaf-forms; Q, female flower;
R, achene; S, female pedicel and perianth.

172 Gardens Bulletin, Singapore — XXV (1969)

Stipules less than 0.1 cm. long, connate, intrapetiolar, bifid, the
two tips shorter than the connate part, broadly triangular, very
thin chartaceous, glabrous, usually very inconspicuous, very cadu-
cous. Inflorescences unisexual; male apedunculate, axillary, with
flowers fasciculate at the base of the female inflorescence; female
up to 2 cm. long and broad, corymbose, axillary, solitary or paired,.
without peduncular bracts, with peduncle lightly armed with long
irritant hairs, multi-flowered, very rarely one-flowered. Male
flowers very shortly pedicellate; perianth ca. 1 mm. long and broad;
tepals 4, non-corniculate, cymbiform, densely covered with minute
irritant hairs; stamens 4, with reflexed filaments to 1.5 mm. long;
pistillode ca. 0.5 mm. long, as broad at the apex, obconical; inter-
floral bracts absent; pedicel to 0.5 mm. long, articulated near the
apex, glabrous. Female fiowers pedicellate; perianth with 4 free
tepals, the two lateral tepals ca. 0.4 mm. long, enclosing the ovary,
glabrous; the dorsal one ca. 0.3 mm. long, geniculate, often slightly,
with one or two rather long irritant hairs; the ventral less than
0.2 mm. long, glabrous; ovary ca. 0.3 mm. long, ellipsoid, asymme-
trical at the apex; stigma capitate or brush-like, not linear, un-
branched, refiexed, to 0.2 mm. long; interfloral bracts absent;
pedicel ca. 0.6 mm. long, slightly winged dorsi-ventrally, articulated
near the apex. Achenes ca. 1-1.5 mm. long, ca. 0.8-1.2 mm. broad,
stipitate, with stipe to 0.4 mm. long, on gynophore ca. 0.4 mm.
long asymmetrically ovoid or pyriform, almost entirely surround-
ed by a very narrow membraneous wing, on each lateral side a
triangular ridge enclosing a warty depression, at most half covered
by the lateral tepals, greatly reflexed; pedicel of lowermost flower
longest, up to 4 mm. long, progressively shorter towards apex of
corymb, slightly winged dorsi-ventrally; dispersed with perianth
and upper portion of pedicel attached.

The uniqueness of the inflorescence, both male and female, has
no parallel in the genus. This so impressed Weddell that he noted
it 98.2... species optima, characteribus essentialibus constantis-
simis. He was quite correct in placing this next to L. ruderalis of
the Pacific Islands with which it has close affinities. Both have
very similar female flowers and achenes. Besides, they seem to
prefer limestone habitats.
~ CUBA:—Guanimar: Ramon de la Sagra s.n. (holotype of U. cuneata:
P).—Havana: Curtiss 609, Jan. 1905 (BM, C, F, K, L, MO, NY, PRC &
US).—Isla de Pinos: Ekman 12333, Nov. 1920 (F, K, NY & S).—Santa
Clara Prov. Combs 277 (F, K & MO); Howard 6272, Aug. 1941 (F, GH,
MICH, MO, NY, UC & US).—-sine loc.: Wright 2228 (type of F.

glomerata: BM, GH, K, MO, NY & US); Poeppig s.n. (type of F.
umbellata: L & PRC).

HISPANIOLA: Haiti:—Gonaives: Nash & Taylor 1501(NY).—Hinche :
Ekman 6156, May 1926 (S).—Morne Calumnette: Ekman 7321, Nov.
1926 (S & US).

W.-L. Chew — Monograph of Laportea 173.

(22) Laportea floribunda (Baker) Leandri in Ann. Mus. Col. Mar-
seille, ser. 6. 7-8: 24 (1950) & in Humbert, Fl. Madagascar, 56:
15. fig. 3 (1965): Chew in Gard. Bull. Sing. 21: 199 (1965). —
Pilea floribunda Baker in Kew Bull. 280 (1897).

Diagnosis: It distinguishes itself from all the species of sect.
Fleurya by the following characters: (a) dioecious shrubs densely
covered with short erect irritant hairs, (6) lamina elliptic, finely
dentate, slightly bullate, (c) female flowers somewhat flabellately
arranged.

Distribution: Madagascar (endemic! ).

Dioecious shrubs. Stem dense with short irritant hairs. Lamina
to 22 cm. long, 10 cm. broad, elliptic, subcoriaceous, lightly bullate,
both surfaces densely covered with short erect irritant hairs; base
cuneate; apex acuminate, acumen to ca. 1.5 cm. long; margin rather
finely dentate; lateral veins ca. 8 pairs, the basal pair slightly more
prominent, reaching to a third the length of the lamina; intercostals
numerous, somewhat anastomosing. Petioles to 7 cm. long, densely
covered with short irritant hairs. Stipules 1.5 cm. long, 0.5 cm.
broad, pubescent. Inflerescences unisexual, narrowly paniculate,
axillary, solitary, to 20 cm. long, 5—7 cm. broad; peduncle densely
covered with short irritant hairs. Male flowers sessile, ca. 2 mm.
long and broad with 5 tepals; stamens 5, some occasionally rudi-
mentary, filaments slightly reflexed: pistillode slightly peltate with
a hollow at the apex. Female flowers pedicellate; perianth ca. 0.75
mm. long, 0.5 mm. broad, with few irritant hairs, the dorsal tepal
strongly geniculate, the laterals broad, enclosing the ovary, the
ventral! thin, minute; ovary ca. 0.75 mm. long, ca. 0.5 mm. broad,
ellipsoid; stigma linear, very short, unbranched; interfloral bracts
minute; pedicel to 1 mm. long, simple, articulated near the apex,
subglabrous. Achenes ca. 1.5 mm. long, 1.25 mm. broad, shortly
stipitate, on a gynophore to 0.4 mm. long, asymmetrically obovoid,
withcut wings, smooth, half covered by the lateral tepals, greatly
reflexed; pedicel up to 4 mm. long, simple, subglabrous; dispersed
with the perianth attached usually.

I have not seen the type of this species; and my interpretation
of it is necessarily based on that of Professor Leandri.

This is the only species of sect. Fleurya to have bullate leaves
which recall those of L. decumana of Malesia. In fact, not a few
collections of the latter species from New Guinea have been found
to have leaves almost similar to those of this species in shape and
size etc. However, this similarity is entirely superficial because
these two species belong to two different sections of the genus.

The relationship of this species to those of section Fleurya is
very obscure.

MADAGASCAR :—South-East: Valley of Maloto: Humbert 6279
Oct.-Nov. 1928 (K & P).

174 Gardens’ Bulletin, Singapore — XXV (1969)

aww Ny

YY
A jucaimt
At} lel.

0-5mm

Fig. 22. L. floribunda: A, twig; B, female flower; C, achene.

W.-L. Chew — Monograph of Laportea 175:

Qe 52 ee Ga ee ES

Species Excludendae

Laportea agusanensis Elm.

L. amplissima (Bl.) Miq.
L. anacardioides C.B. Rob.
L. annamica Gagn.
L. ardens (Bl.) Smith
L. basirotunda Wu
L. batanensis C.B. Rob.
L. brunnea Merr.

L

. carruthersiana (Hiern)
K. Sch.

. chingiana Hand.-Mazz.
conduplicata Elm.

. contracta (B1.) Miq.
corallodesme Laut.
cordata Warb. ex Winkl.
. cordatifolia Smith

. costata (Miq.) Miq.

. crassifolia C.B. Rob.

. crenulata Wedd.

. crenulata var.
luzonensis Wedd.

Sill oil oil oil all ofl oll ail ails

. crenulata var.
nervosa Winkl.

. densiflora C.B. Rob.

. diffusa C.B. Rob.

. elliptica Merr.

. gaudichaudiana K. Sch.
gaudichaudiana Wedd.
. gigas Wedd.

. glabra Rid.

. gracilipes Elm.

hainanensis Merr. & Metc.
. harveyi Seem.

. integrifolia Wu.

. kotoensis Yamam.

. kusaiana Kaneh.

. lanaensis C.B. Rob.

. latifolia Gaud.

. latilanceolata Elm.

. laxiflora Wedd.

. leytensis C.B. Rob.

. longifolia Hemsl.

. luzonensis (Warb.) Wedd.

aS eee Perr rrrrrrerrrere

. crenulata var. nitida Winkl.

Pewee repre oe eee eee Boe 2 oe ee Cee

Dendrocnide luzonensis var.
luzonensis.

. amplissima.

. luzonensis var. anacardioides..

D

D

D. stimulans.
D. sinuata.
D. basirotunda

D. meyeniana f. subglabra.

D. stimulans.

Obetia carruthersiana.
. urentissima.
venosa.

. contracta.

. corallodesme.
cordata.

cordata.

stimulans.

. crassifolia.

. sinuata.
luzonensis var. luzonensis..

nervosa.

ternatensis.

. densiflora.

. Meyeniana f. meyeniana.
elliptica.

. cordata.

meyeniana f. meyeniana.
excelsa.

ternatensis.

stimulans.

. stimulans.

harveyi.

sinuata.

meyeniana f. subglabra.
latifolia.

stimulans.

latifolia.

luzonensis var. luzonensis.
peltata var. peltata.

. Meyeniana f. meyeniana.
. longifolia.

. luzonensis.

176

42,
43.
44.
4S.
46.
AT.
48.
49.
50.
51.
52.
53.
54.
55.
56.
57.
58.
59.
60.
61.
62.
63.
64.
65.
66.
67.
68.
69.
70.
ie
“ae
73.
74,
75:
16.
TT,
78.
79.
80.
81.
82.
83.
84.
85.
86.

oe eee Pe eee SEP PPP Pr RPP PPP PP Pe Pe PPP Pee eS eee re

Gardens’ Bulletin, Singapore — XXV (1969)

mammosisetosa Winkl.

. meyeniana (Walp.) Warb.
. microstigma Wedd.

. milnei Seem.

. mindanaensis Warb.

. mirabilis Rech.

. monticola Merr.

. moroides Wedd.

murrayana Rendle.

_ oblanceolata Merr.

. oblongata Mig.

papangensis Leandri.
peltata sensu Wedd.
peltata Gaud. ex Decne.

. peltata sensu Merr.

. pendula Merr.

. photinophylla (Kth.) Wedd.
. platyphylla Merr.

. pterostigma Wedd.

. pterostigma f. subglabra.

. pustulosa Ridl.

. rechingeri Winkl.

. repens Wedd.

. Tigidifolia C.B. Rob.

saipanensis Kaneh.

salomonensis Rech.

. schlechteri Winkl.

. sessiliflora Warb.

. sinuata (B1.) Mig.

. stenophylla Quis.

. Stimulans (L.f.) Miq.
. subclausa C.B. Rob.
. subglabra Hay.

. subpeltata C.B. Rob.
. Sumatrana Merr.

. ternatensis Miq.

. thorelii Gagn.

. torricellensis Laut.

. triplinervia Merr.

. urentissima Gagn.

. venosa Elm.

. ventricosa Gagn.

. Vitiensis Seem.

. vrieseiana Wedd.

. warburghii Wink].

I

CoOoUSDUoToeVUS

. ternatensis.

meyeniana.

. Microstigma.

harveyi.

meyeniana f. subglabra.
mirabilis.

venosa.

moroides.

peltata var. murrayana.
. oblanceolata.
Villebrunea sp.

Urera cligloba var. papangensis.
eltata.

. moroides.

. elliptica.

subclausa.

. photinophylla.
meyeniana f. subglabra.
. mMeyeniana f. meyeniana.
. meyeniana f. subglabra.

sinuata.

COUDUUUOTeeeS

rechingeri.
Urera repens.
. rigidifolia.
. latifolia.
. longifolia.
. schlechteri.
. sessiliflora.
. sinuata. —
. Stimulans.
. Stimulans.

. subclausa.

D

D

D

D

D

D

D

D

D

D. meyeniana f. subglabra.
D. densiflora.

D. stimulans.

D. ternatensis.
D. -stimulans.

D. torricellensis.
D. venosa.

D. urentissima.
D. venosa.
Pouzolzia sp.
D. vitiensis.
D. peltata var. peltata.
D. longifolia.

W.-L. Chew — Monograph of Laportea . 177
_ Bibliography
Chew, W.-L. (1965). Laportea & Allied Genera. Gard. Bull. Sing.
21: 195-201.

Gaudichaud, C. (1826) in Freycinet, Voyage Autour du Monde
(Botanique): 497-498.

Handel-Mazzetti, H. (1929). Symbolae Sinicae, 7: 116-119.

Hutchinson, J. & Dalziel, J. M. (1928). Urticaceae. Flora of West
Tropical Africa, 1 (2): 441-442.

Keay, R. W. J. (1958) in Hutchinson & Dalziel, Flora of West
Tropical Africa, ed. 2. 1; 619-620.

Leandri, J. (1950). Les Urticacees de Madagascar. Ann. Mus. Col.
Marseille, ser. 6. 7-8: 16-26.

Leandri, J. (1965). Urticacees. Flore de Madagascar et des Comores,
56: 7-16.

Letouzey, R. (1968). Urticaceae. Flore du Cameroun, 8: 115-136.

Maximowicz, C. J. (1877). Sceptrocnide. Bull. Acad. Sci. St.
Petersb. 22: 238-240.

Miquel, F. A. W. (1853) in Martius, Flora Braziliensis 4 (1): 195.

Rendle, A. B. (1917) in Prain, D. Urticaceae. Flora of Tropical
Africa, 6 (2): 245-253.

Weddell. H. A. (1854). Revue de la Famille des Urticees. Ann. Sci.
Wat. ser, 4. 1: 181-183.

Weddell, H. A. (1856). Monographie de la Famille des Urticees.
Arch. Mus. Hist. Nat. Paris 9: 109-142.

Weddell, H. A. (1857). Considerations Generals sur la Famille des
WTCOES: o..0.5: Ann. Sci; Nat. ser. 4.7: 353-354.

Weddell, H. A. (1869). Urticaceae. DC. Prodr. 16 (1) 70-82.

178

cot ca)” guia abt gall ogi eh ols aallty m — tc

Gardens’ Bulletin, Singapore — XXV (1969)

Index to Laportea species

. aestuans, 164.

. alatipes, 124.

. amberana, 131.
. bulbifera, 121.

, Calira, “155.
canadensis, 118.

. cuneata, 170.

decumana, 139.

. disepala, 126.
. floribunda, 173.
. Brossa; 159:

ck ane

. interrupta, 145.

. lanceolata, 161.

. macrostachya, 142.
. mooreana, 157.

. Ovalifolia, 149.

. peduncularis, 152.
. perrieri, 135.

. ruderalis, 167.

. septentrionalis, 133.
. violacea, 129.
. weddellii, 137.

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4.3 3 THE

GARDENS’ BULLETIN

SIN GAPORE
. XXV, Part 21st December, 1970
& eas AES OR, ES ~~
} ae ts) OAS
: MAR 8 1971
SONTENTS
. PAGES
Te ROWSE, G. A. AND RATNASABAPATHY M.: A Species List of
§ | F reshwater Algae from the Taiping Lakes - - - 179-187

y os STONE, B. _C.: Materials for a Moiederaph of Freycinetia
q Gaud. are spamcooky V. Singapore, Malaya and Thailand 189-207

Materials for a Monograph of Freycinetia Gaud. ee
-Raceae) VI. Species of Borneo - oa - - 209-233

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_ tectorius (Ancistrocladaceae) - - - - - - 235-237
E OLITUM, R. E.: _ The Genus Orchidantha (Lowiaceae) - 239-246

ae I IDSDALE, C. E.: The ‘Arborescent Naucleae of New Guinea
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i Asian “Species of Hyphaene me - — - - 299-309

ie A ae Search for Hyphaene ae Thonn. - . 311-334

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A Species List of Freshwater Algae from the
Taiping Lakes, Perak, Malaysia

By GERALD A. PROWSE* and M. RATNASABAPATHYT

Introduction

A need for data on the occurrence and distribution of the fresh-
water algae of Malaysia exists. Such information would be of value
particularly to the fisheries biologist, hydrobiologist, ecologist and
plant geographer.

The Taiping Lakes area is situated at the foot of the heavily
forest clad Maxwell’s Hill, the summit of which is about 1243
meters (4,076 feet) above mean sea level. The Hill and Taiping
district generally are noted for their frequent and heavy rains, about
414 to 495 centimeters (163-195 inches) annually. Thus there is a
constant source of water and the lakes never dry out completely.
However their water level may be controlled by means of wooden
sluice gates. The Taiping Lakes consist of about a dozen lakes and
ponds of various sizes, the biggest being West Lake, approximately
805 meters (880 yards) long and 225 meters (246 yards) wide.
Though few of the lakes are completely enclosed, most of them
receive overflow water from the adjacent stream, drainage canals
and lakes. Most of the lakes sampled were those at the south,
south-east and central area (Fig. 1). Their waters were clearly acid,
the pH ranging from 4 to 6; the colour of the water may be bluish
where still and deep, brownish where there is suspended or dis-
solved humic and detrital material, or, rarely green with a ‘bloom’
of euglenoids. The limit of visibility varied from several centi-
meters to a meter or two. The substratum was mostly of soft mud,
clay and silt; in a few instances, especially in the smaller ponds
there was a thick layer of mowed grass, fallen leaves and twigs,
and other decaying vegetation. Among the variety of animals
encountered were Hydra sp., collected here for the first time in
the Malaysian peninsula in December 1954, and subsequently in
1956-58 (Ratnasabapathy, M., unpublished data); Amoeba sp.,
Arcella sp., Paramecium sp., Coleps sp., Halteria sp., nematodes,
rotifers, aquatic insects including Ranatra sp. and nymphs of
Odonata; Cyclops, shrimps and crabs, Aplocheilus panchax,
Rasbora trilineata, Ophiocephalus sp., half-beaks, catfishes; and
tadpoles of frogs and toads. The lake area and surrounding lands
were at one time mined for tin and the lakes are an outcome
of this. The algae listed in this paper were collected during the

* Tropical Fish Culture Research Institute, Batu Berendam, Malacca,
Malaysia.

+ School of Biological Sciences, University of Malaya, Kuala Lumpur,
Malaysia.

180 Gardens’ Bulletin, Singapore — XXV (1970)

period 22nd September 1957 to 11th January 1958. At that time
the lake area was fairly well stabilised and beautifully landscaped
with grass slopes, lawns, flower beds and a golf course for recrea-
tional purposes. In recent years the central jungle has been turned
into a Zoo with introduced as well as native mammals and birds.
Changes in the algal flora are thus expected and future follow-up
collections should be interesting. The senior author has described
earlier (1958, 1960, 1962a, 1962b) several of the Euglenineae and
Bacillariophyta listed here for the Taiping Lakes, and, is currently
working on a monograph of the Malaysian Desmidiaceae where
the species from the Taiping Lakes will be described.

Species List of Algae

BACILLARIOPHYTA

COSCINODISCACEAE

Melosira granulata (Ehrenberg) Ralfs.
Melosira italica (Ehrenberg) Kuetzing.

EUNOTIACEAE

Eunotia maior (W. Smith) Rabenhorst var. emarginata A. Cleve.
Desmogonium rabenhorstianum Grunow.

NAVICULACEAE
Frustulia rhomboides (Ehrenberg) de Toni.
Navicula radiosa Keutzing var. minutissima (Grunow) P. T. Cleve.
Pinnularia braunii (Grunow) P.T. Cleve.

Pinnularia braunii (Grunow) P.T. Cleve var. amphicephala (Mayer)
Hustedt.

Pinnularia brebissonii (Kuetzing) P.T. Cleve var. hybrida (Grunow)
A. Cleve.

Pinnularia brevicostata P.T. Cleve.

Pinnularia gibba W. Smith var. interrupta A. Cleve-Euler.
Pinnularia hemiptera (Kuetzing) Rabenhorst.

Pinnularia legumen Ehrenberg.

Pinnularia mesolepta (Ehrenberg) W. Smith.

Pinnularia parva (Gregory) P.T. Cleve var. parvula (Ralfs) A.
Cleve-Euler.

Pinnularia stauroptera (Grunow) P.T. Cleve var. subparallela
Mayer.

Pinnularia stomatophoroides Mayer var. nuda A. Cleve-Euler.
Pinnularia tabellaria Ehrenberg.

Prowse and Ratnasabapathy — Algae, Taiping Lakes 181

MILES
0 100 200 So WA
—_—_—_—_—_—————_— eee
(182.88 meters)
Sampling station.......¢ Waterfall’, Road
Grass lawn...............G.L.
Foot he rr

BREIELT Sian ces consasascecseonces

FE pe as
‘ -<--..

i CENTRAL JUNGL

ee

~ _ ~ Swan Lake r

West ’ Lake Q

South Lake AS

FIGURE 1. MAP OF TAIPING LAKES

182 Gardens’ Bulletin, Singapore — X XV (1970)

GOMPHONEMATACEAE
‘Gomphonema subtile Ehrenberg.

NITZSCHIACEAE
Nitzschia palea (Kuetzing) W. Smith.

EUGLENOPHYTA

Euglena fusca (Klebs) Lemmermann.

Euglena gracilis Klebs.

Euglena splendens Dangeard.

Phacus curvicauda Swirenko.

Phacus hamatus Pochmann.

Phacus pseudo-nordstedtii Pochmann.

Phacus trifacialis Prowse.

Trachelomonas hispida (Perty) Stein emend. Déflandre.
Trachelomonas similis Stokes var. hyalina Skvortzow.
Trachelomonas superba Swirenko emend. Déflandre.
‘Trachelomonas volzii Lemmermann var. cylindracea Playfair.

CHLOROPHYTA

PALMELLACEAE
Sphaerocystis schroeteri Chodat.

HYDRODICTY ACEAE
Pediastrum duplex Meyen.
Pediastrum tetras (Ehrenberg) Ralfs.
Tetraedron sp.
OOCYSTACEAE

Nephrocytium agardhianum Naegeli.

COELASTRACEAE

Coelastrum microporum Naegeli.

SELENASTRACEAE

Ankistrodesmus falcatus (Corda) Ralfs.
Closteriopsis longissima Lemmermann var. tropica West & West.
Selenastrum bibraianum Reinch.

ee 2

Prowse and Ratnasabapathy — Algae, Taiping Lakes

DICTYOPHAERIACEAE
Dictyosphaerium pulchellum Wood.

Dimorphococcus lunatus A. Braun.

SCENEDESMACEAE
Scenedesmus brasiliensis Bohlin.
Scenedesmus perforatus Lemmermann.

Tetrallantos lagerheimii Teiling.

ULOTRICHACEAE
Hormidium sp.

OEDOGONIACEAE
Oedogonium sp.

ZY GNEMATACEAE
Spirogyra 2 spp.

MESOTAENIACEAE

Netrium digitus (Ehrenberg) Itzigsohn & Roth.

DESMIDIACEAE
Closterium dianae Ehrenberg.
Closterium juncidum Ralfs.
Closterium libellula Focke.
Closterium liebleinii Kuetzing.
Closterium navicula (Brébisson) Luetkemuller.
Closterium toxon West.
Closterium venus Kuetzing.

Pleurotaenium kayei (Archer) Rabenhorst.

183

Pleurotaenium minutum (Ralfs) Delp. var excavatum Scott &

Gronblad.

Pleurotaenium ovatum Nordstedt.

Pleurotaenium nodosum (Bailey) Lundell var. gutwinskii Krieger.

Pleurotaenium verrucosum (Bailey) Lundell.

Euastrum denticulatum (Kirchner) Gay var. quadrifarium Krieger.

Euastrum gnathophorum West & West.

Euastrum longicolle Nordstedt var. capitatum West & West.

184 Gardens’ Bulletin, Singapore — XXV (1970)

Euastrum moebii (Borge) Scott & Prescott.

Euastrum moebii (Borge) Scott & Prescott var. burmense West &
West.

Euastrum pulcherrimum West & West var. menggalense Scott &
Prescott.

Euastrum serratum Joshua.

Euastrum sinuosum Lenorm.

Micrasterias lux Joshua fa.

Micrasterias mahabuleshwarensis Hobson.

Micrasterias pinnatifida (Kuetzing) Ralfs.

Micrasterias radians Turner.

Micrasterias tropica Nordstedt var. polonica Eichler & Raciborski.
Cosmarium askenasyi Schmidle.

Cosmarium obsoletum (Hantzsch) Reinsch var. sitvense Gutwinski.
Cosmarium pseudoconnatum Nordstedt.

Xanthidium urniforme (West & West) Scott & Croasdale.
Arthrodesmus curvatus Turner.

Arthrodesmus octocornis Ehrenberg.

Staurastrum connatum (Lundell) Roy & Biss.

Staurastrum freemanii West & West var. triquetrum West & West.
Staurastrum pinnatum Turner.

Staurastrum pinnatum Turner var. subpinnatum (Schmidle) West
& West.

Staurastrum sexangulare Lundell.
Staurastrum tohopekalagense Wolle.
Staurastrum wildemanii Gutwinski.
Spondylosium planum (Wolle) West & West.
Spondylosium tetragonum W. West.

CYANOPHYTA

Dactylococcopsis sp.
Merismopedia sp.

Spirulina gomontii Gutwinski.
Oscillatoria sp.

Anabaena sp.

Prowse and Ratnasabapathy — Algae, Taiping Lakes 185

Bibliography

Only literature which has been consulted and is relevant is included
here. References cited previously in papers of the senior author are not

Tepeated.

Behre K. 1956.

Bernard Ch. 1908.

. 1909.

Borge O. 1897.

Desikachary T.V. 1959.

Gronblad R. 1945.

Gronblad R., G. A. Prowse
& A. M. Scott. 1958

Gutwinski R. 1902.

Krieger W. 1933.

. 1937/39.

Nordstedt O. 1880.

Die Siisswasseralgen der Wallacea —
Expedition.

Arch. Hydrobiol. Suppl. 23 (1): 1-
104.

Protococcacées et Desmidiées d’Eau
Douce, Récoltées a Java.

Dept. de l Agric. aux Indes Néerl. 230
pp. Batavia.

Algues Unicellulaires d’Eau Douce
Récoltées dans le Domaine Malais.

Dept. de l’ Agric. aux Indes Néerl. 94
pp. Buitenzorg.

Australische Siisswasserchlorophyceen.
Bih. Kongl. Svenska Vetensk. Akad.
Handl. 22 Afd 3: 1-36.

Cyanophyta.
ICAR. New Delhi 686 pp.

De Algis Brasiliensibus, Praecipue
Desmidiaceis.

Acta Soc. Sci. Fenn. n.s.B. 2 (6) : 1-43.

Sudanese Desmids.
Acta Bot. Fenn. 58 : 1-82.

De algis a Dre. M. Raciborski Anno
1899 in Insula Java Collectis.

Bull. Acad. Sci. Cracovie 9 : 575—
617.

Die Desmidiaceen der Deutschen Lim-
nologischen Sunda Expedition.

Arch. Hydrobiol. Suppl. 11 : 129-230.

Die Desmidiaceen Europas in Raben-
horst’s Kryptogamen-flora von Deut-
schlands, Osterreich und der Schweiz
Teil 1, 1937: 1-712.

Tel 2, 1939 : 1-117.

De Algis et Characeis I : De algis
nonnullis praecipue Desmidieis, inter
Utricularias. Musei Lugduno-Bata-
via. Acta Univ. Lund 16 : 120-133.

186 Gardens’ Bulletin, Singapore — XXV_ (1970)

Prescott G. W. &
A. M. Scott. 1945.

shale

Prowse G. A. 1957.

Prowse G. A. 1958.

. 1960.

,. 1962a.

A DG2D.

Ralfs J. 1848.

Scott A. M. &
R. Gronblad. 1956.

Scott A. M. &

G. W. Prescott. 1958.

27 L9OD.

Scott A. M. &

G. W. Prescott. 1961.

Skuja H. 1949.

The Freshwater algae of Southern
United States 3. The Desmid Genus
Euastrum.

Amer. Midl. Naturalist 34 (1) : 231-
2au.

The Algal Flora of Southeastern United
States 5. Additions to Our Know-
ledge of the Desmid Genus Micras-
terias.

Trans. Amer. Microscop. Soc. 71 (3) :
229-252.

An Introduction to the Desmids of
Malaya.

Malayan Nat. J. 2 (3) : 42-58.

The Eugleninae of Malaya. _
Bard. Bull. Sing. 16: 136—204..

New and Unusual Flagellata in Malaya.
In Proc. Centenary and Bicentenary
Congr. of Biol. Singapore. pp 292-
298.

Diatoms of Malayan Fresh Waters.
Gard. Bull. Sing. 19 (1) : 1-104.

Further Malayan Freshwater Flagel-
lata. Gard. Bull. Sing. 19 (1) : 105-
145.

British Desmidieae. 226 pp. London.

Notes on Indonesian Freshwater Algae
I. Staurastrum wildemannii Gutw.
Reinwardtia 3 (3) : 105-112.

Some Freshwater Algae from Arnhem
Land in Northern Territory of Aus-
tralia. Rec. Amer-Austr. Exped.
Arnhem Land 3: 9-136.

Notes on Indonesian Freshwater Algae
IV. Concerning Euastrum moebii
(Borge) Scott & Prescott comb. nov.
and Euastrum turgidum Wallich.

Reinwardtia 5 (3) : 317-334.

Indonesian Desmids.
Hydrobiologia 17 (1-2) : 1-132, 63 pl.

Zur Siisswasseralgenflora Burmas.

Nova Acta Regiae. Soc. Sci. Upsal.
Ser. 4, 14 (5) : 1-185.

SS

Prowse and Ratnasabapathy — Algae, Taiping Lakes 187

Smith G. W. 1950.
West W. & G. S. West.

1897.
—_————.. 1902.

s 1907.

. 1904/11.

West W., G. S. West &
N. Carter. 1923.

Fritsch F. E.

The freshwater algae of the United
States. 2nd ed. New York.

Desmids from Singapore.
J. Bot. 33. :@soG-157.

A contribution to the freshwater algae
of Ceylon.

Trans. Linn. Soc. London, Bot. 6 :
123~125)

Freshwater algae from Burma, includ-
ing a few from Bengal and Madras.

Ann. Roy. Bot. Gard. (Calcutta) 6
(2) : 175-260.

A monograph of the British Desmi-
diaceae. Vol. 1, 1904; Vol. 2, 1905;
Vol. 3, 1908; Vol. 4, 1911; The Ray
Society London.

A monograph of the British Desmi-
diaceae. Vol. 5. The Ray Society
London.

Collection of algae (F.B.A.) — draw-
ings, as microfiche cards 1-1133.

Materials for a Monograph of Freycinetia Gaud.
(Pandanaceae) V. Singapore, Malaya and Thailand

By BENJAMIN C. STONE

School of Biological Sciences, University of Malaya
Kuala Lumpur, Malaysia

Abstract

Eight species of Freycinetia Gaud. are reported from the Malayan
Peninsula and adjacent Thailand and Singapore. This number is not any
Peninsula and adjacent Thailand and Singapore. This number is hardly
different from that given by Ridley (8) in his Flora of the Malay Peninsula;
however he considered 7 of these as endemic species, while after consider-
able comparative study, it now appears that only one of his species is
endemic, and that rather doubtfully. Hence, most of the binomial species
names used in Ridley’s Flora have had to be changed. In this paper keys,
description, and synonymy are indicated; and one new species, and two
new varietal taxa are proposed.

Introduction

There are only eight species of Freycinetia Gaud. (1824) in the
Malayan peninsula, Singapore, and the narrow Kra Peninsula of
Thailand; thus the area stands in considerable contrast to regions
to the east. Borneo has at least 24 species, the Philippines at least
as many, and New Guinea more than 40 species. Furthermore,
only one of the Malayan species is endemic, and that only tenta-
tively. This is also in marked contrast tc the three regions just
mentioned, each of which has a concentration of endemic species.
The Malayan species show the following distribution:

F. javanica— Malaya and the Kra Peninsula of Thailand;
Sumatra; Borneo; Java.

F. angustifolia — Malaya, Sumatra, Borneo, Java, Philippines.
F. corneri — South Malaya (Johore), Singapore, Borneo.

F. sumatrana— Malaya and the Kra Peninsula of Thailand;
Cambodia; Vietnam; Andaman Islands; Sumatra; Borneo; Java;
Philippines.

F. confusa — Malaya; tentatively considered endemic, but ex-
tremely similar to F. distigmata of Sumatra; to be further studied.

F. imbricata — Malaya, Sumatra, Borneo, Java.
F. rigidifolia— Malaya, Andaman Islands, Borneo.
F. kamiana — Malaya, Sumatra.

The names used above, it will be seen, are nearly all different
from those used by Ridley in the ‘‘Flora of the Malaya Peninsula”’
(Vol. 5, 1925). Ridley credited Malaya with eight species of
Freycinetia, seven of which he considered endemic. But this

190 Gardens’ Bulletin, Singapore — XXV_ (1970)

optimistic conclusion has not fared well, and in the light of mono-
graphic studies all but one of these species has turned out to
be previously known and described from different regions. The
synonymy in brief is as follows:

F. javanica B1. (F. lucens Ridl.; F. montana Ridl.).

F. angustifolia B1. (F. angustifolia B1. sensu Ridl.; F. malaccen-
sis Ridl.).

F. corneri Stone (not known to Ridley).

F. sumatrana Hemsl. ((F. valida Ridl.).

F. confusa Ridl. (endemic).

F. imbricata Bl. (F. kingiana Ridl.).

F. rigidifolia Hemsl. (F. acuminata Rid].).

F. kamiana Stone (not known to Ridley).

In this paper a revision of the Malayan species, including those
of adjacent Thailand and Singapore, is presented, with keys, brief
descriptions, citations of specimens, field notes, and illustrations.
Based upon three years of field work in Malaya and a tour of
several European and American herbaria, it forms the fifth in a
series antecedent to a monograph of the genus. I would like to

express my thanks to the curators and directors of the museums
and herbaria mentioned below for their cordial assistance.

A The Arnold Arboretum, Harvard University, Cam-
bridge, Mass., U.S.A.

BISH B.P. Bishop Museum, Honolulu, Hawaii.
BM British Museum (Natural History), London.
CAL Indian Botanic Gardens’ Herbarium, Howrah, Calcutta,

India.

E Royal Botanic Gardens, Edinburgh, Scotland.

FI Istituto Botanica, University of Florence, Florence,
Italy.

K Royal Botanic Gardens, Kew, Surrey, England.

KLU University of Malaya Herbarium, Kuala Lumpur,
Malaysia.

KEP Herbarium, Forest Research Institute, Kepong, Malaysia.

L Rijksherbarium, Leiden, Netherlands.

SING Botanic Gardens, Singapore.

U University of Utrecht Herbarium, Utrecht, Netherlands.

I should also like to express my thanks to the Chairman of the
School of Biological Sciences, University of Malaya for much
assistance, and to a former student, Miss Tan Poey Keng, who
undertook a very useful preliminary study of Malayan Freycinetiae.

Stone — Freycinetia (Pandanaceae V) 19]
Systematic Treatment

Freycinetia Gaudichaud

Ann. Sci. Nat. I, 3: 509. 1824. Solms-Laubach, Linnaea 1878-9.
Warburg, in Engler, Pflanzenreich IV. 9, 1900.

Ridley, Mat. Fl. Malay Penin. 2: 232-5. 1907; Fl. Malay Penin.
5: 82-84, 1925.

Lectotype species: Freycinetia arborea Gaud. 1824 (see Stone
1967).

Woody climbers with clasping adventitious roots. Leaves simple,
arranged in 3 spiralled ranks, linear, oblong, elliptic, lanceolate or
oblanceolate, commonly serrate-dentate along the margins and on
the midrib dorsally. Leaf-base sheathing the stem, usually with
membranous and caducous, or sometimes more or less permanent
but fragile auricles. Inflorescence terminal on a main or a lateral
branch, bracteate, and pedunculate, of 1 or more cephalia arranged
umbellately or sometimes racemosely, often ternate, sometimes
with up to 7 (very rarely more) unisexual cephalia; plants dioe-
cious. Staminate spadix of crowded simple stamens, pistillodia
sometimes present; anthers subglobose. Pistillate spadix of crowded
gynoecia, baccate when ripe, with usually a rigid or coriaceous
pileus; stigma single or up to 12 (rarely more) per berry, minute,
microscopically papillate, often within a low rim. Seeds numerous
on vertical placentae corresponding to each stigma, usually elliptic
or hemi-elliptic to falcate, with a noticeable translucent raphe with
some silvery raphidophorous cells, and sometimes a similar and
opposite strophiole. Embryo minute, within endosperm.

Distribution: Asiatic and Pacific Tropics; west to Ceylon (not
in India proper); Andaman Islands; Malayan Peninsula and
Singapore; Sumatra; Borneo; Java; the Philippine Islands; Viet-
nam; Formosa (Taiwan); Ryukyu Islands; Bonin Islands; Micro-
nesia; Melanesia; New Guinea; Australia (north Queensland);
Norfolk Island; New Caledonia; New Zealand; Polynesia, including
Hawaii, but not Easter Island.

The greatest concentration of species is in the regions of Borneo,
Philippines, and New Guinea (including the Solomon Islands).

A genus of perhaps 150 species (estimated; there are far more
binomials). The name commemorates Captain Louis de Freycinet,
commander of the vessels ““L’Uranie’”’ and ‘‘Physicienne”’ of the
French Exploring Expedition of 1816-1818.

192 Gardens’ Bulletin, Singapore — XXV _ (1970)

se ee, ST

|

JAVANICA

---—o LL

.
'
.
'
wee ee ee,
.
.

Ww9

\

|

T

CONFUSA

RIGHDIFOLIA
i ' |
| SUMATRANA |
| | var | |
PENANGIANA |
: | | | tT ae
oe | | f
‘ !
}
i— eee rs ~~ : Se on a) ats

oat $ -s. teen d. P=

Fig. i. Auricles of Malayan Freycinetiae. a-b : F. javanica. c — F.

sumatrana. d — F. sumatrana vat. penangiana. e — F. rigidifolia.
f — F. confusa. (All from cited specimens). To same scale.

Stone — Freycinetia (Pandanaceae V ) 193

KEYS TO MALAYAN SPECIES OF FREYCINETIA

I. Field Key for Sterile or Staminate Plants Based
Principally on Vegetative Characters

(1) Auricles caducous, relatively short and broad, not or rarely
more than 6 cm long, tapered or rounded at apex, without a free
apical lobe, entire or minutely and sparsely weakly denticulate;
leaves to about 30 cm long.

(2) Leaves elliptic, to oblong or oblanceolate, definitely narrowed
at base, acute-acuminate, not drawn out to a filiform-flagellate
tip, to 30-40 cm long and 3-5-8 cm wide, though usually smaller;
spadices umbellate.

(3) Pedicels 4+ as thick as long; leaves elliptic, hardly or not
Nt? GAVE cURL ld iid. . A ce Qe 1. F. javanica

(3) Pedicels much more slender; leaves oblong, caudate
NR Se RARE nln ico o .o:s i ae & F. kamiana

(2) Leaves narrowly linear, up to 30 cm long but rarely more
than 1 cm wide, apex gradually attenuate-filiform, base not
or scarcely narrowed; spadices racemose ...... 2 F. angustifolia

(1) Auricles subpersistent but fragmenting and eventually lost,
usually more than 3 cm long, sometimes much more (18 cm),
sometimes with a free apical lobe, otherwise with a truncate apex,
usually slightly to manifestly denticulate on the margins and/or
apex; leaves sometimes over | m long.

(4) Robust plants, stems (adults) often exceeding 15 mm dia-
meter, leaves commonly over 50 cm long, up to 1 m (or more);
auricles mostly more than 5 cm long, up to 15-18 cm long,
with a distinct apical lobe to 1 cm long (in F. sumatrana) or a
blunt truncate apex.

(5) Leaf apex acute, not drawn out; auricles to 9 cm long and
1-2 cm wide, truncate or slightly rounded dat apex. Teeth of
basal leaf-margin broadly deltoid (to 2 mm long and 2 mm
wide), rather distinct (c. 1 cm apart). Pistillate cephalia sub-
globose, on glabrous pedicels, berry with 3-8 stigmas .........
cos BRR) OSS ee ee > ee 3 F. corneri

(5) Leaf apex gradually attenuate, subflagellate; auricles to 10
(or to 18) cm long, 1-1.5 cm wide, apex with a narrow free
ascending lobe, usually denticulate on margin; teeth of basal
leaf margin slender, antrorse, usually less than 2 mm long and
1 mm wide. Pistillate cephalia cylindric, on minutely scabrid
pedicels; berry with 2 (rarely 1 or 3) stigmas ..................
a felons Catlett Webasto)! SOS 4 F. sumatrana

(4) Small to medium plants, stems (adults) rarely over 11 mm
diameter, leaves rarely over 40 cm long; auricles rarely over 5
cm long, usually less than 1 cm wide, truncate, and denticulate
or spinulose at apex but never with a free apical lobe.

194

Gardens’ Bulletin, Singapore — XXV (1970)

(6) Median portion of leaf margin denticulate; leaves narrowly
linear, commonly about 30 cm x 5 mm. Auricles narrow,
truncate, denticulate at apex; blade not narrowed at base
5 F. confusa

Sse 'e Bis 6 we ee ©8089 OF 66600068 COO DMO Aw eS se we ae es wees 6. dae A ee oo ©

(6) Median portion of leaf margin unarmed; leaves mostly
over 6 mm (and up to 15 mm or more) in width. Auricles
entire or spinulose-fimbriate. Leaf blade slightly narrowed at
base.

(7) Auricles entire or with few, small inconspicuous teeth.
Leaves commonly about 30 x 1—1.5 cm, mostly flat, rather
lax. Pedicels of pistillate cephalia smooth or very slightly
scabrid. Berry with 2-5 stigmas ............... 6 F. imbricata

(7) Auriciels deeply fimbriate-spinulose at apex (spinules to
5 mm long), margins denticulate; leaves commonly about
20 cm x 9 mm, stiff, the margins often revolute: abruptly
narrowed at base. Pedicels of pistillate cephalia hispidulous.
Derry with 1-2 stisineee a, 42.) seiccsecee oe 7 F. rigidifolia

II. Key to Fertile Specimens

Based on Pistillate Inflorescences and Fruits

(1) Inflorescences (both ¢ and 9) racemose, the 3-5 spadices clearly
distinct, well-separated, pedicellate .................. 2 F. angustifolia

(1) Inflorescences umbellate, usually of 3 crowded pedicellate
spadices.
(2) Stigmas usually 2 per berry, sometimes 1 or 3.

(3) Pedicels stout, minutely roughened (like sandpaper to
touch) commonly 50 x 5-6 mm. Cephalia to 15 cm long;
bracts white. Robust plamts <...:.:.-:..-....-.- 4 F. sumatrana
(3) Pedicels slender, 10-40 x 3-4 mn, scabridulous chiefly
on the angles. Cephalia 14-35 mm long. Rather small plants.

(4) Auricles deeply fimbriate-spinulose at apex, with spinules
to 5 mm long. Bracts red-orange. Leaves stoutly denticulate
on basal margins, and abruptly narrowed at top of sheath;
up te" 10 mm wid 2°. Aaa cag .. 7 F. rigidifolia

(4) Auricles slightly denticulate; spinules not over 2 mm
long. Bracts purplish. Leaves with small teeth; blades about

47am wide: ...........0. oe B.h6!.eeel- ers 5 F. confusa

(2) Stigmas usually 3-5 (8) per berry, very rarely 2.

(5) Pistillate cephalia oblong. Auricles caducous, sparsely and
weakly denticulate or entire.
(6) Leaves oblong, caudate; pedicels slender, 12-16 x 2-3
8 F. kamiana
(6) Leaves elliptic-oblong, commonly 10-30 cm long and
12-30 mm wide, nearly entire, only at apex with weakly
serrulate-denticulate margins and midribs. Pedicels short
and stocky [UO X 3.5—40 Mime ees cer esscet neers I F. javanica

Stone — Freycinetia (Pandanaceae V) 195

(5) Pistillate cephalia subglobose or short-oblong. Auricles
_ subpersistent but fragmenting, entire to weakly or strongly
denticulate. Leaves not as above. |

(7) Robust plants; stems to 2 cm diameter, leaves to 1 m
long and 3.5 cm wide; auricles denticulate, truncate-rounded
a ie So. ne a 3 F. corneri

(7) Smallish plants, stems to 12 mm diameter, leaves less
than 40 cm long, usually 10-15 mm wide: auricles tapered
and entire to weakly denticulate. Stigmas 3-5 ...............
Hien HBO hag Bl). he HIE: --sorcbeteee lat Onda, enbricata

Enumeration of Species

(1) Freycinetia javanica Blume in Rumphia |: 156, t. 41 (1835).
Figure 1, a—b. | |
Synonymy: F, lucens Ridl. in Mat. Fl. Malay Penin. 2: 232

(1907) in Fl. Malay Penin. 5: 80 (1925). F. montana Ridl. in Mat.

Fl. Malay Penin. 2: 232 (1907) in Fl. Malay Penin. 5: 82 (1925).

Type localities and typification: F. javanica: Java (Blume, L).
F. montana: Perak (Ridley 5194, lectotype, SING). F. lucens:
Singapore (Ridley 3703, lectotype, SING).

Distribution: Malaya (including lower Kra Peninsula), Singa-
pore, Sumatra, Java, Borneo.

Stems to 11 mm diameter. Leaves elliptic or somewhat lanceolate
to oblanceolate, abruptly acute, not or scarcely acuminate, nar-
rowed at base, slightly thick-coriaceous, nearly unarmed (only the
apex and rarely a short part of the basal margin denticulate);
auricles membranous, when fresh colorless and translucent, cadu-
cous, tapered or rounded to the apex, entire or weakly and sparsely
denticulate, mostly 3-6.5 cm long, ‘and 1 cm wide. /nflorescence
terminal, umbellate, usually ternate; pistillate pedicels slightly
scabridulous on the angles, about 1 cm long, 3.5-4.5 mm thick.
Cephalia oblong, mostly 2.5-5 cm long. Berry 1—1.5 mm long.
Stigmas 4-8, usually 4-6. Bracts orange-salmon color or ‘“‘Flesh-
color” often yellowish. Staminate cephalia with whitish stamens,
filaments 0.35 mm long, anthers 0.30 mm long.

Habitat: in humid forests, commonly montane, ascending to
6,000 ft., but also lower, to 1,000 ft. or even lower especially in
the south, as in Singapore (similarly in Sarawak) in well-protected
forested humid valleys and along waterfalls. (Low-altitude plants
often have unusually wide leaves, while mountain populations
mostly show rather short and narrow leaves. Two widely different
auricle forms are found, extremes of a continuous series, which
correspond to the situations men‘ ied; see illustrations).

196 Gardens’ Bulletin, Singapore — xXx V (1970)

Ridley’s distinction between F. lucens and F. montana appears
to be purely imaginary; and he did not distinguish either of these
from F. javanica. Perhaps Ridley was unfamiliar with Blume’s
work, although he admitted one of Blume’s species of Freycinetia
(F. angustifolia) in the Malayan flora, despite the clear occurrence
of three of them (originally described from Java) in the Malayan
peninsula.

Malayan specimens:

THAILAND: Pattani, Kao Katakini, Sept. 1923, Kerr 778I
(BM, K, L). Kao Nam Sao, 900 m., Feb. 1927, Kerr 12053 (BM,
K). Kao Luang, N. Sritamarat, 1,200 m., April 1928, Kerr 15488
(BM).

MALAYA: PERAK: Maxwell’s Hill, June 1893, Ridley 5194
(lectotype of F. montana) FI, SING. Same loc., 1892, Ridley
(SING). Larut, Dec. 1881, King’s collector (i.e. Kunstler) 2619
(FI, SING); 4003 (FI, SING); 4091 (FI, SING); May 1883, 4194
FI, SING). Thaiping, Feb. 1886, Kunstler 8523 (FI); May 1885,
Kunstler 7578 (FI, SING); April 1890, Wray 3950 (FI); Pelan
Tujer, April 1888, Wray 1871] (FI). Larut, Fr. Scortechini (FI, &
sheets). Maxwell’s Hill, 3,700 ft., coll.? SFN. 72557 (SING).
Gunong Hijau, 1892, Ridley (SING).

TRENGGANU: Bukit Kajang, Kemaman, Nov. 1935, Corner
(SING).

PENANG: Sungei Telek Bahang, Feb. 1919 (or 1929) Burkill
4555 (CALC).

PAHANG: Lubok Tamang, 3,500 ft., June 1923, Robinson
11009 (SING). Cameron Highlands, Robinson’s Falls, April 1930,
4,400 ft., Henderson SFN. 23454 (CALC, K, SING); April 1934,
Jaamat & Sow CF. 36165 (KEP); May 1965, Stone 5623 (KLU).
Gunong Benom, main NE ridge, 3,200 ft., March 1967, Whitmore
FRI. 3205 (KEP). Fraser’s Hill, 4,000 ft, Aug. 1923, Nur SFN.
10550 (SING); Aug. 1937, Holttum 33162 (K); near Richmond
Bungalow, 1959, Md. Shah 6048 (K); Aug. 1937, Corner SFN.
33162 (SING); Sept. 1922, Burkill & Holttum 8546 (SING); April
1966, Stone 6273 (KLU); July 1966, Stone 6427 (KLU). Sungai
Kuantan, June 1934, Symington & Kiah SFN. 28771 (K, SING).
Gunong Ulu Kali, 1967, Stone s.n. (KLU).

SELANGOR: “The Gap” at 3,000 ft, May 1902, Curtis
(SING). Bukit Hitam, May 1896, Ridley 7657 (FI, SING). Templer
Park, Kanching 1967, Stone s.n. (KLU). Gunong Nuang, Ulu
Langat, May 1940, Symington 51813 (KEP).

NEGRI SEMBILAN: Seremban, April 1892, Edgerton (SING).
MALACCA: no location, 1867-8, Maingay 3274 (K).

JOHORE: Kukut, Easter 1907, Ridley (SING). Mawai, March
1934, Corner (SING). Sungei Sedili, Feb. 1938, Corner (SING).
Pontian, April 1952, Santiago (SING).

Stone — Freycinetia (Pandanaceae V) 197

SINGAPORE: Kranji, Ridley in 1890 (SING); Bukit Mandai,
1890, Ridley 4618 (FI, K, SING). Bukit Timah, 1892, Ridley 3703
(lectotype of F. lucens) FI, K, SING. Seletar, May 1953, Sinclair
SFN. 39616 (SING, K).

MALAYA, without locality: Griffith 6374 (FI); Maingay’
1537 (K).

(2) Freycinetia angustifolia Blume in Rumphia 1: 157, t. 40 (1835)..
Ridley, Fl. Malay Penin. 5: 82 (1925).

Synonymy: F. malaccensis Ridley in Mat. Fl. Malay Penin. 2:
233 (1907); in Fl. Malay Penin. 5: 81 (1925). F. brunoniana
Wallich (nom. nud.) List, n. 3660 (1828-32). F. debregeasiana:
Gaudich. in Bot. Voy. Bonite, Atlas, t. 37, f. 1-11 (1843).

Type localities and typification: F. angustifolia: Java (Blume,.
L). F. malaccensis: Malacca (lectotype, Derry 406, SING). F.
debregeasiana: not stated, but possibly Singapore (Gaudichaud in.
1837; P? fragment in FI). |

Distribution: Malaya, Sumatra, Java, Borneo, Philippines.

Vernacular name: “selingseng akar’’ (Malay; Johore, Corner:
28074).

Stems 6-12 mm diameter. Leaves linear, slender, thin, 17—20:
cm long, 5—12 mm wide, gradually attenuate, very slightly acumi--
nate, unarmed except at extreme apex. Auricles 15-20 mm long,.
caducous, entire, fragile. Inflorescences racemose, rachis some--
times somewhat zig-zag, pedicels well-separated; bracts dark to
pale yellow or yellow-range, sometimes flecked with red on inner
surfaces, innermost bracts creamy, outermost with green tips; pistil-.
late cephalia on pedicels 9-11 x 1.5-2 mm, glabrous, whitish;.
cephalia shortly cylindric, 23-33 mm long, at first cream-colored;
berry 1 mm long. Stigmas 1-3, rarely 4, very rarely 5. Staminate:
spadix 1-2 cm long, white. Seed with broad raphe and no strophiole.

Habitat: low hill forest, chiefly in the southern and western
parts of the peninsula; also in lowland swamp forest; also on
offshore islands.

Ridley’s F. malaccensis was characterised by its relatively broad’
leaves, but examination of numerous specimens shows that con-
tinuous variation, within the limits given above, occurs; and there:
are no other distinguishing features.

MALAYA: PENANG: George Porter in 1823 (““F. brunoniana.
Wall.” n. 3660) Fl. “‘Coast’? March 1887, Curtis 1181-bis (SING),
1171 (K); Dec. 1886, Curtis 1143 (SING). Penang Hill, Dec. 1895,
Ridley (SING). Telok Bahang, Feb. 1919, Burkill 4555-bis (CALC,.
SING).

PERAK: Sonkai, May 1882, Kunstler 3052 (FI, SING). Kele-
dang Saiong, April 1937, Symington 25557 (KEP).

SELANGOR: Klang, Telok Reserve, June 1921, Burkill 6518
(SING).

198 Gardens’ Bulletin, Singapore —- xXx V (1970)

NEGRI SEMBILAN: Pulau Rembia (Sembilan Islands), along
streambed 100 ft. Jan. 1953,: Wyatt-Smith 76493 (KEP). Bukit
Kandong, March 1885, Omabius (? sphalm.) 1117 (SING).

MALACCA: Mt. Ophir, 1892, Ridley 3895 (FI, K, SING).
Bukit Bruang, Feb. 1890, Derry 406 (lectotype of F. malaccensis
Ridley; miscited by Ridley as “‘Ridley 406’’, FI, SING); March
1893, Derry 1197 (FI, SING; det. by Ridley as F. angustifolia).
Merlimau, 1892, Derry 1046 (FI — as “Ridley 1046’’; SING).

JOHORE: Sungei Tebrau, March 1908, Ridley 13212 (SING).
Pengabong May 1908, Foxworthy CF 1200 (KEP, SING). Kangka
Sedili Kechil, June 1924, Corner SFN 28617 (CALC, K, SING).
Sungei Sedili, April 1934, Corner SFN 28074 (K, SING). Kota
‘Tinggi riverbank Dec. 1892, Ridley 4142 (K, SING). Mawai Road
— Kota Tinggi, March 1936, Corner SFN 30784 (SING). Sungei
Sedili between Mawai and Bukit Tiga, March 1937, Corner SFN
32435 (SING). Sungei Kaipu, March 1937, Kiah SFN 32354 (L,
‘SING). Sungei Sedili, March 1940, Corner (SING). Gunong Panti,
1,500 ft., Jan. 1937, Corner SFN 32216 (L, SING). Kuala Sedili
Road in freshwater swamp forest, May 1961, H.M. Burkill 2667
(L).

SINGAPORE: Bukit Mandai, 1890, Ridley 1643 (SING). April
1921, Burkill 6507 (SING). Bukit Timah, 1893, Ridley 5120
(SING). Sungei Buloh, Nov. 1889, J.S. Gamble (SING). Kranji,
1909, Ridley 14176 (CALC, K, SING). Seletar Reservoir, Feb.
1943, Corner (SING); Jan. 1949, Sinclair (L). Without locality,
‘Gaudichaud, in 1837 (Herb. Delessert, Geneva; fragm. FI; P; am
of F. debregeasiana Gaud.).

MALAYA, without definite locality, Maingay 1537/2 (FI, K,
Lz.

(3) Freycinetia corneri B. C. Stone, Brittonia 20 (1968) 198-202.
Plate II.
Type locality: Singapore (Corner).
Distribution: Malaya (Johore, Selangor), Singapore, and Borneo.

Robust climber, stems to 20 mm diameter; Jeaves to more than
1 m long, 2-3.5 cm wide, linear, apex acute, sheathing base 9-10
cm long, blades coriaceous; auricles to 9 cm long (at least), 1-2
cm wide, the apex abruptly truncate-rounded (rarely almost lobate),
the sides usually entire but the apex ciliate-spinulose. Leaf-margin
-stoutly and rather remote dentate, the teeth deltoid and spreading
at right angles to the margin, the lowermost ones nearly 2 mm long,
up to 1.9 mm wide, about 1 cm apart, distally the teeth smaller.
Midrib denticulate from the middle beyond, the teeth at most
up to 1.9 mm wide, about 1 cm apart. distally the teeth smaller.
2 mm long, smaller toward the apex. /nflorescence terminal, ter-
nate, umbellate, the pedicels (9) about 60 x 5-7 mm, plano-convex,

Freycinetia sumatrana in the Ginting Highlands, Pahang.

Plate I.

3

[EPs

%.

<

¥

Pert a

i. White

corner

ight, F.

R

kamiana.

Fs.

’

icles. Left

Freycinetia auri

Plate II

bar is 1 cm long.

Stone — Freycinetia (Pandanaceae V ) 199

subtrigonal, smooth. Bracts deltoid, to 6 cm wide or more, probably
reddish. Pistillate cephalia about 45 x 40 mm, subglobose. Berry
oblong, about 14 x 3 mm, the pileus 4 mm long, narrowly
pyramidal, apex truncate. Stigmas usually 4-8 (very rarely 3),.
minute, hippocrepiform. Staminate spadix on pedicels to 60 mm
long, spikes ternate or quaternate, cylindric, the floriferous part.
about 30 x 8 mm, filaments 1.5 mm long, anthers 0.75 mm long.

Habitat: lowland forests. A species of restricted range, only
discovered among Bornean specimens after being described from
the Singaporean material (see citation of Borneo specimen below)..

MALAYA: JOHORE: near Kota Tinggi, Feb. 1938, Corner
SFN 34672 (K, PNH, SING). Mawai-Jemaluang Rd. 14th mile,
April 1938, Corner SFN 34938 (SING). Gunong Panti, 1780 ft.,.
June 1963, Stone 4834 (L).

SELANGOR: Sungei Lallang F. R., Y. K. Kam s.n. in 1967
(KLU).

SINGAPORE: | Seletar forest, behind vegetable gardens, Nee:
Soon, Jan. 1949, Sinclair (K). Seletar Reservoir, Feb. 1943, Corner
(SING, type! ). |

BORNEO: Central East Indonesian Borneo, W. Koetai, n. 39,.
Mt. Kemoel, 1800 m., 21 Oct. 1925, Endert 4453 (L.). (New distri--
bution record).

To date the species is known only from the specimens cited.

(4) Freycinetia sumatrana Hemsley in Kew Bulletin, 167 (1896)..
Plate I; figures 1, c-d; 2; 3, a—b.

Synonymy: F. valida Ridley in Mat. Fl. Malay Penin. 2: 233:
(1907); in Fl. Malay Penin. 5: 83 (1925). Winkler in Engl. Bot.
Jahrb. 48: 87 (1912); Merrill, Bibl. Enum. Bornean Pl. 34 (1921)..

Type localities and typification: F. sumatrana: Sumatra (Beccari,
K). F. valida: Singapore (lectotype: Ridley 3937, SING).

Distribution: Sumatra, Malaya, Borneo, Philippines, Andaman
Is.

Robust climbers with stems to 18 mm diameter (or more);
leaves linear, gradually attenuate, commonly 45-110 cm _ long,
1-3 cm wide, margins denticulate nearly all along their length
with relatively slender usually antrorse teeth. Auricles purplish,
subpersistent but fragmenting, 4-8 (-18) cm long, 1-2 cm wide,
with a distinct erect-spreading denticulate apical lobe to 1 cm
long; auricle margins entire to denticulate. Inflorescence terminal,
umbellate, ternate; bracts creamy to pure white, outermost ones.
sometimes tinged pinkish dorsally; pistillate cephalia cylindric, up
to 15 x 4 cm, more often 5-10 x 2-3 cm, dull red when ripe, on
pedicels about 40-55 x 46 mm, costulate, finely roughened-
scabridulous. Berry very slender, to 20 mm long. Stigmas as a.
rule 2 per berry, very rarely 1 or 3. Staminate spadix creamy--
_white, spikes 6—7.5 cm long, 1.35 cm wide.

Leaf spectrum of Freycinetia sumatrana var. penangiana. Pro-
phylls and leaves in numbered sequence; Ist prophyll, no. 1;
prophylls 2 through 8 all unarmed; intermediate prophylls 9-11
with denticulate apex; foliage leaves 12-et seq. Leaf bases of leaves

13, 14, 15, and 45, and leaf apex of leaf 45, show

Stone — Freycinetia (Pandanaceae V ) 201

BCSdel69

Fig. 3.

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F. KAMIANA ¢

a—b, Freycinetia sumatrana. a — detail of apex of staminate
spadix. b — details of berry and seed. c — Freycinetia kamiana
(from holotype). Left, leaf with enlarged inset of marginal teeth.
Right, above: segment of fruiting pedicel; profile of immature
carpels; top view of same. Below, pistillate inflorescence, with one
cephalium in longisection.

202 Gardens’ Bulletin, Singapore — XXV (1970)

Habitat: In Malaya, in forests from the lowlands to about 6,000
ft., the high montane forms are smaller, and often have leading
shoots of a deep purplish color throughout. Younger plants and
lowlands plants are often sterile for long periods. Montane plants
may produce fruit of a normal color but devoid of seed.

THAILAND: Kao Katakini, Pattani, Sept. 1923, Kerr 7792
(BM) Kao Katakuan, 200 m. March 1930, Kerr 18510 (B M).

MALAYA: KEDAH: Turton, Nov. 1889, Curtis (SING).

PERAK: Thaiping Hill, Feb. 1904, Ridley 11891 (SING);
Wray 2170 (Fl). Batu Togoh, 300 ft.. Wray (71082) (SING).
Birch’s Hill, March 1924, Burkill & Haniff SFN 12674 (SING).

PENANG: Penang Hill, 2,400 ft., 1901, Curtis 3538 (SING):
Feb. 1892, Ridley (SING).

PAHANG: Cameron Highlands, Lubok Tamang, 3,000 ft.,
April 1937. Nur SFN 32953 (KEP, K, SING). May 1965, Stone
5625 (KLU). Oct. 1966, Md. Kassim (KLU). Fraser’s Hill, Aug.
1937, Corner (SING). April 1966, Stone 6289 (KLU).

SELANGOR: April 1890, Ridley (SING). Ginting Highlands,
3,200 ft., Nov. 1966, Stone 6568 (KLU).

NEGRI SEMBILAN: Pulau Rembia, Sembilan Islands, Jan.
1953, Wyatt-Smith 76442 (KEP). Port Dickson, Sungei Menyala,
1957, Whitmore 300 (K).

MALACCA: Gunong Ledang, 1892, Ridley (SING).

JOHORE: Mukub, 1908, Ridley (SING). Gunong Belumut,
May 1923, Holttum SFN 10822 (K, SING). Kluang, Gunong
Belumut, 3,300 ft., Feb. 1966, Francis Ng, 98059 (KEP). Mt. Ophir,
May 1890, Derry 627 (SING).

SINGAPORE: Bukit Mandai, Aug. 1899, Ridley 10439 (SING).
Gardens’ Jungle, Chua Chu Kang, 1892, Ridley 3937 (F. valida,
lectotype, SING). No locality, 1905, Ridley (SING).

F. sumatrana Hemsl. var. penangiana B. C. Stone, var. nov. Figures
Lad: 2. |

Ab var. typica in auriculis angustioribus attenuatis minute
truncatis nec lobatis differt.

Holotype: SELANGOR: Templer Park, Kanching, June 1965,
Stone 5809 (KLU). ‘

Type locality; Stone 6040 (KLU).

PENANG: Penang Hill, Jan. 1901, Curtis 3538 (SING). Upper
Tunnel Station, 2,000 ft., Feb. 1966, Stone 6/38 (KLU).

KEDAH: Turton, Nov. 1889, Curtis (SING, in part).

ANDAMAN ISLANDS: 1875, Kurz (FI).

This variety may be a local form, or merely a state of growth.
The narrow auricles, commonly 8-10 cm long, but sometimes to
18 cm long, with a long taper but an abrupt though very brief
truncation at the apex, without a lobe, and with minutely denticu-
late margins and a purplish color, seem to be very characteristic.

Stone — Freycinetia (Pandanaceae V ) 203:

(5) Freycinetia confusa Ridley in Mat. Fl. Malay Penin. 2: 234
(1907); in Fl. Malay Penin. 5: 82 (1925). Figure 1, f.

Not F. confusa Elmer 1907 (renamed by Elmer as F. villarii
1908). |

Type locality: Singapore (lectotype, Ridley 4757, SING).

Distribution: Singapore and Malaya, apparently endemic, but
very similar to and possibly not distinct from F’. distigmata Stone
of Sumatra, which is however a later name than F. confusa.

Slender climber with stems to about 5 mm diameter, leaves
linear, gradually attenuate, 30-33 cm long, 4-5 mm wide, the
margins denticulate. Auricles 25-32 mm long, 4 mm wide, slightly
tapered, the apex briefly truncate, minutely spinulose-ciliate; sides.
entire. Inflorescence terminal, umbe!late, usually ternate; pedicels.
of pistillate cephalia scabridulous, 11 x 4 mm; pistillate cephalia.
(immature) 14 mm long. Berry (immature) 1 mm long; stigmas.
1 or 2 per berry. Staminate inflorescence unknown.

Habitat: Lowland forests.

In his original description, Ridley cited Wray 2151 as a syntype
of F’. confusa, but this specimen is actually F. rigidifolia (see under
that species). Because of this Martelli confused the species and in
FI almost all specimens of F. rigidifolia were marked F. confusd.
Ridley cited Wray 215] as 2157, by mistake,

MALAYA: SELANGOR: Changgang Klang, Oct. 1937, Nur
SFN 33982 (SING).

JOHORE: Pengkalan Raja, July 1939, Ngadiman (SING).
Mawai, April 1936, Corner SFN 30886 (SING).

SINGAPORE: Seletar, Feb. 1893, Ridley 4757 (lectotype,,.
SING). Kranji, 1894, Ridley (SING).

(6) Freycinetia imbricata Blume in Rumphia 1: 157, t. 40 (1835).

Synonymy: F. schefferi Solms-Laubach in Linnaea 42: 98.
(1879). F. kingiana Ridl. in Mat. Fl. Malay Penin. 2: 234 (1907);
in Fl. Malay Penin. 5: 82 (1925).

Type localities and typification: F. imbricata: Java (Blume
2066, L). F. schefferi: Java (Scheffer; Korthals). F.. kingiana:
Perak, Malaya (Kunstler 4654, SING).

Distribution: Malaya, Sumatra, Java, Borneo.

Smallish to medium climbers, stems to 12 mm diameter, leaves
mostly 20-35 cm long, 6-15 (-17) mm wide, rarely to 24 mm wide,
narrowly elliptic-oblong to nearly linear-lanceolate, usually slightly
narrowed at base, acuminate-acute at apex; margin denticulate near
base and apex, otherwise entire. Auricles more or less persistent
and clasping the stem, tapered-rounded at apex, membranous,
entire or weakly and sparsely denticulate, usually 2-5 (—7) cm
long, 4-6 (-9) mm wide. /nflorescence terminal or pseudolateral
(i.e. On short side branches with a few reduced leaves intercalated.

204 Gardens’ Bulletin, Singapore — X XV (1970)

below the bracts). Bracts yellow to creamy. Pistillate cephalia
globose or broadly short-ellipsoid, 25-45 mm long or rarely more,
bright orange when ripe; on pedicels 9-10 x 3.5 mm, glabrous or
(in var. hispidula) minutely hispidulous. Berry to 10 mm long,
pileus 1.5-2 mm long, dome-shaped or low-conic, briefly or not
rostrate, areola less than 1 mm wide; stigmas 1-5, usually 3 or 4.
Seeds about 1.3 mm long, falcate-ellipsoid, raphe with shining
raph:dophorous cells, usually broader than the narrow strophiole.
‘Staminate spadix . .

Habitat: lowland forests, sometimes on rocks.

MALAYA: PERAK: Gopeng, Larut, July 1883, Kunstler
4654 (lectotype of F. kingiana, SING; FI). 1880, Kunstler 1056
(FI, SING). June 1886, 400-600 ft., Kunstler 10/65 (FI).

F. imbricata var. hispidula B. C. Stone, var. nov.
Ab var. typica in pedicellis foemineis hispidulis differt.

Holotype: MALAYA: SELANGOR: Genting Sempah, 1,500
ft., on rock, July 1966, Stone 5847-a (KLU).

SELANGOR: Same as type locality, June 1965, Stone 5&47
(KLU). Ginting Highlands, 3,000 ft., Nov. 1966, Stone 6535
(KLU). Ulu Langat, Jan. 1959, Gadoh (for Millard) 1327 (SING).

PAHANG: Fraser’s Hill, 4,000 ft.. Nov. 1965, Stone 6078
(KLU).

JOHORE: Mawai, Sept. 1934, Corner SFN 28181 (K, SING).
Sungei Kayu, March 1937, Kiah SFN 32380 (KK, SING).

I believe that this variety with hispidulous pedicels and perhaps
mostly somewhat broader leaves, is the same as F. schefferi Solms.
It is not possible to keep the taxa as distinct species however.

47) Freycinetia rigidifolia Hemsley in Kew Bulletin, 166 (1896).
Piswre 1, "€.

Synonymy: F. acuminata Ridley in Mat. Fl. Malay Penin. 2:
233 (1907); in Fl. Malay Penin. 5: 82 (1925). F.. bennettii Miq. as
misapplied in Hort. Bogor., in part, excluding ref. to spms. of F.
javanica.

Type localities and typification: F. rigidifolia: Borneo (Havi-
Jand, K). F. acuminata: Malaya, Selangor (Ridley 7656, SING).

Distribution: Borneo, Malaya, Andaman Islands; cult. in Hort.
Bogor.

Slender climber, with stems “‘to 80 ft. long” (fide Kunstler), to
6 mm diameter or more; leaves stiff, 16-24 cm long, 8-10 mm
wide (Malayan specimens), elliptic-lanceolate, acute-acuminate,
narrowed rather abruptly at base just at apex of auricles, margins
often revolute, blades rather stiffly erect. Auricles abruptly rounded-
truncate, deeply fimbriate with spinules 5 mm long or more at apex,

es

Stone — Freycinetia (Pandanaceae V) 205

the sides with shorter teeth; auricles 25 x 5 mm. Blade margin
at base stoutly denticulate; median part unarmed; apex finely
denticulate. Dorsal surface somewhat glaucous. J/nflorescence
terminal, umbellate, ternate; bracts red or red-orange. Pistillate
cephalia greenish-yellow (immature), cylindric, to 35 mm long;
pedicels scabridulous on the angles, about 30-40 x 4 mm. Berry
(immature) 1 mm long; stigmas | or 2.

Habitat: lowland or hill forest (in Borneo on Mt. Kinabalu to
5,000 ft. or higher).

A very distinct and easily recognised species.

ANDAMAN ISLANDS: South Andaman, 1909, Rogers (FI).
Same locality, Kurz (in part; mixed with some specimens of F.
sumatrana; FI).

MALAYA: PERAK: Batu Tugoh, 200 ft., June 1888, Wray
215] (cited — in error as 2/57 — by Ridley as a syntype of F.
confusa Ridl.) (SING). Same locality, 300-800 ft., Nov. 1885,
Kunstler 8300 (FI).

TRENGGANU: Bukit Kajang, Kemaman, Nov. 1935, Corner
SFN 30406 (SING).

PAHANG: Fraser’s Hill, Aug. 1937, Corner (SING).

SELANGOR: Kuang, May 1892, Ridley 7656 (lectotype of F.
acuminata) (SING). Sungei Buloh, April 1929, Sow and Taehan
SEN. 16447 (SING); same locality, March 1919, Long CF 3307
(SING). Bukit Lagong, July 1965, Kochuninen 93141 (KEP).

JOHORE: Mawai-Jemaiuang Road 13.5 miles, May 1935,
‘Corner SFN 29738 (SING).

(8) Freycinetia kamiana B. C. Stone, sp. nov. Figure 3, c; plate Lf.

Holotype: Y. K. Kam s.n., Bukit Lagong, Selangor, Malaya
{KLU).

Stipite erecta ad 2-2.5 m alta et 18 mm diametro, internodts
ad 4 cm longis, simplex vel sparse ramificata, ramis horizontalibus
vel laxe subpendulis vel decumbentis, ad 5-8 mm diametro, inter-
nodiis 5-20 mm longis, nodis vetustioribus cicatricosis. Folia
juvenilia magna ad 40 cm longa et 6 cm lata, oblonga vel oblonge-
lanceolata, basi 3 cm lato, apice abrupte rotundata caudato, caudo
15-28 mm longo; auriculis ad 35 mm longis et 10 mm latis,
rotundato-attenuatis margine minute distaliter denticulato, mem-
branaceis. Folia vetustiora simillima sed parviora ad 18-30 cm
longa et 2-4 cm lata, marginibus (apicem versus in caudo excepta)
inermis. /nflorescentia foeminea terminalis, ternatis, pedicellis 5,
ad 12-16 mm longis et 2-3 mm crassis, supra minute sparseque
hispidulis. Cephalia oblonga vel cylindric ad 5 cm longa, 1 cm
lata, curvata, baccis immaturis congestis 5—6—angulosis, circiter
5-6 mm longis, 1—-1.5 mm crassis, pileo depresso-pyramidato coro-
natis; stigmatibus 3—5-(7). Cetera ignota.

206 Gardens’ Bulletin, Singapore — XXV (1970)

Stems erect and un- or few-branched (at least at first), later with
longish rambling branches; finally (? perhaps) climbing; stems
reddish to yellowish-brown, up to 18 mm diam. with internodes
as much as 4 cm long, or on younger branch-tips much shorter
(5-20 mm); nodes with conspicuous leaf-scars. Leaves oblong or
oblanceolate-oblong; vigorous young shoots with large leaves up
to 40 x 6 cm, at base narrowed to 3 cm (just above the sheath-
auricles), at apex abruptly rounded-caudate, cauda 15-28 mm
long, denticulate on margins and midrib with minute prickles,
the leaf otherwise quite unarmed. Auricles up to 35 x 10 mm,
rounded or rounded-attenuate to almost deltoid, pale to translu-
cent (in life), with about 5—6 well-spaced slender nerves; margins
‘thin membranous, minutely and weakly serrulate-denticulate to-
ward the distal end with tiny prickles less than 0.5 mm long.
Blade with about 48 prominulent longitudinal nerves, when dry
with numerous manifest tertiary reticulations (forming rectangles);
midrib entire and slightly carinate beneath. Older branches (or
plants) with considerably smaller leaves mostly 18-30 cm long
and 2-4 cm wide, of the same shape etc. but with smaller auricles
12-22 mm long. Pistillate inflorescence terminal, ternate, pedicels.
about 12-16 mm long, 2-3 mm thick, smooth below but towards
apex sparsely and minutely hispidulous. Young cephalia oblong
to cylindric, more or less curved, up to 5 cm long, about 1 cm
thick, with (immature) berries rather crowded, about 2.5 mm
long, only the extreme apex free; pileus polygonal, about 1-1.5
mm wide, shallow, bearing 3-7, usually 4—6 stigmas. Spathes and
staminate plants unknown.

Habitat: known so far only from lowland hills in forest.
Distribution: Malaya and Sumatra.
Specimens examined:

MALAYA: SELANGOR: Bukit Lagong, behind Kepong,
hillside forest, 13 April 1969, Yee Kiew Kam s.n. (Type; KLU,
and distrib.). — Same locality and same group of plants, Jan. 1967,
J. Dransfield s.n. (KLU, and distrib.).

SUMATRA: Cultivated plant from Sumatra, said to be from
“Bukit Tinggi’, now in the Hortus Bogoriensis, Java, no. XI-
B-V-128.

This unexpected new species seems to be very rare in Malaya
at least, and John Dransfield’s collection is the first known. This
was made at a time when no fruits were seen, and the large leaves
and short auricles suggested F. javanica, but could not really be
satisfactory as such because of the great size and unusual shape.
When mv student. Miss Kam, revisited the same locality later on

Stone — Freycinetia (Pandanaceae V) 207

she was fortunate to find young syncarps which have permitted
the recognition and description of this species. Because of this,
and because of her valuable work in elucidating the systematic
anatomy of the Pandanaceae, it is a pleasure to dedicate this
interesting Freycinetia to her.

The plants cultivated in the Botanic Gardens in Bogor were
observed on a recent study visit there. They seem to be precisely
this species. However, no fruiting or flowering materials were seen.
There seems no doubt about the source, so that it can be asserted
that the species is a native of both Malaya and Sumatra.

The habit of this species is unusual in the genus, since the
appearance is distinctly that of an erect shrub. The plants have
not, as yet, been seen as climbers. The branches, however, are
rather straggling, and it is quite possible that in due course the
plants will become topheavy and may eventually undertake the
ordinary climbing behaviour of Freycinetiae. The main stems are
however, at least up to 2.5 m high, perfectly straight and relatively
rigid.

References

GAUDICHAUD, C. B. 1824. Descriptions des quelques nouveaux
genres des plantes Ann. Sci. Nat. I, 3: 508-512.

RipLey, H. N. 1925. Flora of the Malay Peninsula. Vol. 5.
London, L. Reeve & Co. (reprint, 1967).

Stone, B. C. 1967. Materials for a Monograph of Freycinetia
(Pandanaceae), I. Gard. Bull. Sing. 22 (2): 129-152.

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Materials for a Monograph of Freycinetia Gaud.
(Pandanaceae)

VI. Species of Borneo

By BENJAMIN C, STONE

School of Biological Sciences, University of Malaya
Kuala Lumpur, Malaysia

Abstract

Twenty-four species of Freycinetia Gaud. (Pandanaceae) are reported
from Borneo, more than twice as many as previously known. Four species
are new to science: F. parviaculeata, F. biloba, F. kalimantanica, and
F. kinabaluana. The following new infraspecific taxa are also proposed:
F. confusa var. minima, F. javanica var. expansa, F. robinsonii var.
meijeri, and F. palawanensis var. andcrsoniana: A brief phytogeographic
discussion is included.

No account of the Bornean species of Freycinetia has appeared
since that of Merrill (1921), which listed 11 species. Since that
time two further species have been described by Martelli (1929).
The accumulation of specimens in various herbaria has not been
accompanied by investigations into the taxonomy of the genus in
Borneo, although other regions, in particular New Guinea and the
Solomon Islands, have been the subject of study by Martelli and
Perry (1939, 1940). Through the generosity of the Forest Depart-
ment Herbaria in Sandakan (Sabah) and Kuching (Sarawak), as
well as that of the Botanic Gardens (Singapore) and the Forest
Research Institute (Kepong), most of the material collected in
Borneo in recent years has been available to me for study. During
academic leave from the University of Malaya in 1968 studies
were undertaken of materials preserved in the herbaria of the
University of Firenze (where both the Beccari collections and the
Martelli Herbarium are kept), in the Rijksherbarium, Leiden, in
the Royal Botanic Gardens, Kew, and also Edinburgh, and in the
British Museum (Natural History). From these studies it has been
possible to gain a much better understanding of the genus and in
particular of the species occurring in Borneo. In this paper, twenty-
four species are reported, many for the first time, and four new
species are proposed. The species are, wherever possible, referred to
sections of the genus. These sections have been described in Part
IV of this series.

Phytogeographic considerations. The relationships of the Bornean
species rest chiefly with the rest of Malesia (i.e. Sumatra, Java,
Malayan Peninsula, and the Philippines) and only to a slight extent
with Eastern Indonesia (Celebes, Halmaheira, Sunda Islands, etc.).
As far as is presently known, there is no species shared by New
Guinea and Borneo. Three species, at least (probably some other
examples will be discovered) occur both in Borneo and in the
Philippines but not elsewhere (F. discoidea, F. gitingiana, and
F. palawanensis). Two species occur in the Philippines, Borneo,

210 Gardens’ Bulletin, Singapore — XXV (1970)

and eastward, in Java, Sumatra, and Malaya. These are F. sumat-
rana and F. angustifolia. (These species are herein first reported
for the Philippines). If, as seems possible, F. imbricata occurs in
the Philippines under another name (possibly F. montalbanica
Martelli) it could be added to the list. There are five species (F.
sumatrana, F.. angustifolia, F. javanica, F. imbricata, and F. rigidi-
folia) which occur in Borneo, in Malaya, and additionally in
Sumatra, Java, or both, or (as to the first and last only) in the
Andaman Islands, or (as to the first and third only) in southern
Thailand.

This means that thirteen species appear to be endemic to Borneo.
On the other hand it is entirely possible that further species will
come to light as the exploration of the rich Bornean vegetation
continues.

Taxonomic characters, with some advice to collectors. As is also
true of Pandanus as represented in herbaria, there is a great pre-
ponderance of pistillate specimens of Freycinetia. There are perhaps
two main reasons for this, one being the fact that the male inflores-
cences are ephemeral, while the developing fruits persist on the
plants over a period of months; and the other reason being that
since the classification of the genus has been based so heavily on
pistillate materials there is little likelihood of staminate collections
being properly identified unless accompanied by pistillate material
of the same species. As will be noticed in the key and descriptions
to follow, the value of certain vegetative characters has proved to
be great, and attention by collectors to the securing of good
vegetative material, especially the delicate leaf-base auricles, will
greatly facilitate the identification of all specimens and in particular
the males. Further distinctive characteristics, such as the color of
the floral bracts and of the auricles, the presence of glaucousness,
and the differences between leaf size and shape in adult and
juvenile plants, should be borne in mind by the collectors and
noted down. At present it is not possible to determine very young
juvenile and seedling plants. The importance of ripe fruit with
mature seeds is also great, and these are generally to be preferred
to the immature fruits or flowers of pistillate plants if the floral
bracts are lacking. The prophylls of developing young shoots are
also of value, at least potentially, in the process of identification.

A cautionary word should be said of the external features of the
pedicels. At present, no species known produces the staminate
spadices on anything but smooth and glabrous pedicels. However,
the presence of epidermal appendages (spine-like processes, aculeate
or straight, or other scale-like structures which give to the pedicel
a scabrid “feel’’ and appearance) is a useful and important diag-
nostic feature, although present only on pistillate inflorescence
pedicels.

The following key to species is of necessity based on the features
of pistillate specimens, since at least 14 of our species are known
only from pistillate collections. It is hoped that careful attention
will be paid to the desirability of obtaining material of the staminate
inflorescences in future collections.

Stone — Freycinetia (Pandanaceae V1) 211

Key to Borneo Species of Freycinetia

1. Leaf apex rather abruptly acuminate-caudate or cuspidate (but
see also F. gitingiana). Leaves mostly relatively short, rarely
much over 40 cm long.

2. Berry with one stigma (rarely 2). Leaves small, mostly 7—9
cm long, 2-3 cm wide, and elliptic 1 F. tenuis

2. Berry with 2 or more stigmas. Leaves mostly larger.
3. Berry with 2 or 3 stigmas.

4. Blades elliptic, proportionally very wide, 7.5 x 2.5 cm
up to 22.5 x 7.5 cm, apex caudate, cauda slightly
acuminate 17 F. hemsleyi

4. Blades oblanceolate, proportionally narrower, about
16 x 1-3 cm, apex abruptly caudate /2 F. borneensis

3. Berry with usually 3—7 stigmas (very rarely only 2).

5. Fem. pedicels glabrous. Leaves about 40 x 2.4 cm.
Stigmas mostly 4-6 per berry
10 F. palawanensis var. andersoniana

5. Fem. pedicels spinulose-hispidulous or glabrous. Auri-
cles denticulate to long-ciliate.

6. Auricles pectinate long-ciliate. Leaves 12-20 cm
long, 2-3 cm wide, the margins at base and apex
ciliate I3 ¥-Cihars

6. Auricles minutely denticulate on margins. Leaves
about 30 cm long, 2.5-3.5 cm wide, nearly entire
except at extreme apex and base where margins bear
few very minute denticulations © Il F, biloba

1. Leaf apex gradually and narrowly.attenuate, subulate, or acute;
leaves sometimes well over 1 m long.

7. Auricles lobed (i.e. upper end of auricle extended upward-
outward like a lobe).

7a. Robust plants; leaves to 120 cm long, 4-6 cm wide.
Cephalia long-cylindric. Stigmas usually 2 (rarely 3) per
berry; pedicels stout, minutely scabrous 5 F. sumatrana

7a. Smaller plants; leaves c. 50 x 1 cm. Cephalia cylindric;
stigmas 2; pedicels slender, distally spinulose 6 F. lucida

7. Auricles entire, at apex rounded, truncate, or tapered.

8. Cephalia usually 4-7 together, spirally crowded. Robust
plants with leaves 80-100 cm long, 2.5 cm wide
7 F. kinabaluana

8. Cephalia usually ternate (3 together in an umbel), rarely
4, sometimes only 2 or solitary.

212 Gardens’ Bulletin, Singapore — XXV _ (1970)

9. Auricles deeply pectinate-spinulose; leaves rigidly ensi-
form, narrow, abruptly narrowed at the auricle and
with revolute margins 9 F. rigidifolia

9. Auricles, etc., not as above.

10. Inflorescences lateral (i.e. below the main cluster of
leaves, produced at the apex of a short lateral
branch bearing prophylls and floral bracts but no
ordinary foliage); or sometimes both lateral and
terminal.

11. Leaves mostly 15-35 cm long, 0.6-1.5 cm wide;
fem. pedicels glabrous or (in one variety) slightly
hispidulous. Cephalia subglobose or nearly short-
oblong, ternate, binate or solitary 4 F. imbricata

11. Leaves mostly 40-60 cm long, 0.6 cm wide, cepha-
lia subglobose, solitary or two 3 F. sarawakensis

10. Inflorescences invariably terminal (except in F.
imbricata)

12. Inflorescence a raceme of small oblong well-
spaced short-pedicellate cephalia. Leaves linear,
less than 1 cm wide 2 F. angustifolia

12. Inflorescence umbellate (or solitary).

13. Robust plants with leaves 60-100 cm long, up
to 3.3 cm wide, thickly coriaceous.

14. Stigmas invariably 2 per berry; berries ex-
tremely long and slender, filiform. Cephalia
oblong-cylindric, borne on minutely but

; densely aculeolate pedicels
8 F. parviaculeata

14. Stigmas usually 4-8 per berry; berries broader
than filiform. Cephalia subglobose, borne on
glabrous pedicels 15 F. corneri

13. Leaves shorter, or narrower, or both.

15. Pedicels of pistillate inflorescences quite
glabrous.

16. Stigmas 3-7 per berry. Leaves commonly
2 cm wide or wider.

17. Leaves mostly (20-) 40-50 cm long,
about 2 cm wide.

18. Leaves acute, unarmed or nearly so;
stigmas mostly 3-5 (—7)
J4 F. concinna

Stone — Freycinetia (Pandanaceae V1) 213

18. Leaves acuminate, conspicuosly
prickly on basal margins; auricles purp-
lish; stigmas usually 2, rarely 3-4
16 F. discoidea
17. Leaves mostly 12-25 cm long, 1-3 cm

wide; in juvenile plants these dimensions
somewhat exceeded. Auricles not purp-

lish. I8 F. javanica
16. Stigmas 3-4 per berry; leaves 40-60 cm
long, 0.6 cm wide 3 F. sarawakensis

15. Pistillate pedicels scabrid or hispidulous.

19. Stigmas usually 2 per berry. Leaves 45-70
cm long, about 1 cm wide, rather stiffly
linear-ensiform and attenuate-subulate.
Auricles purplish-brown, subtruncate, en-
tire. Floral bracts purple

21 F. winkleriana

19. Stigmas 2—5 (—8) per berry. Auricles entire
or somewhat denticulate on margin.

20. Leaves less than 15 mm wide,

21. Leaves 4-10 mm wide, lanceolate or
linear.

22. Leaves linear, 4 mm wide
24 F. confusa var. minima

22. Leaves 7-10 mm wide
4 F. imbricata var. kuchinensis

21. Leaves 10-13 mm wide, long-lanceo-
late, gradually attenuate. Cephalia
cylindric 19 F. robinsonii var.

: meijeri
20. Leaves 18-30+ mm wide.

23. Stigmas usually 2; cephalia slender
cylindric, 4-6+ cm long, yellowish;
leaves with short conspicuous prickles
on margins near apex and base

22 F. gitingiana

23. Stigmas usually 4-8.

24. Stigmas most commonly 4 (but 3-6):
cephalia about 25 mm long; leaves
virtually unarmed, apex acute

20 F. creaghii

24. Stigmas most commonly 6 (but 5-8);
cephalia about 35 mm long; leaves
almost unarmed, apex attenuate

23 F. kalimantanica

214 Gardens’ Bulletin, Singapore —X XV _ (1970)

Enumeration of species

Sect. Oligostigma Warb. em. B. C. Stone

(1) Freycinetia tenuis Solms, Linnaea 42: 87, 1879.

Liana pumila montana silvatica in saxibus occurrens, caulis ad
4-5 mm diametro, internodiis 12-15 mm longis, foliis ellipticis
acuminatis breve-caudatis basi angustatis 7-9 cm longis, 2-2.8 cm
latis, auriculis brevibus ad 8 mm longis (delapsis); marginibus
foliorum et costa, apicem versus excepta, inermibus caudis breve-
denticulatis. Inflorescentia foeminea terminalis ternata pedicellis
gracilis semiteretibus laevibus 16-18 mm longis, 1.5-1.8 mm
crassis. Cephalia subglobosa vel breviter oblongata 1-1.8 cm
longa, 8-10 mm lata. Baccae maturitate ad apicem succulentae,
stigma unicum. Apex baccae rotundatus. /nflorescentia mascula
subterminalis, bracteis ovatis acutis ad 23 x 16 mm. Spadix ad
1 cm longus, parte floriferis 4 x 2 mm.

Type: SUMATRA: Korthals 23 (L!), staminate.

BORNEO: E. Sabah; Tambunan District, Koingaran, creeper
in mountain forest in rocky soil, fruit green, Oct. 1962, G. Mikil
SAN 31851 (SAN! ). Mt. Kinabalu, Dallas, alt. 3,000 ft., Aug.—Sep.
1932, J. & M. S. Clemens 26210 (= 26734) (L!); Penibukan,
4,000-5,000 ft., Jan. 1933, Clemens 31038 (L!), 50137, 32135
(BM!).

This species is easily recognized among the Bornean Freycinetiae
by its small size and unistigmatic gynoecia. The small globose
cephalia and the berries with rounded, succulent apices, place the
species in the Sect. Oligostigma as recently delimited. It is very
similar to F. beccarii Solms, the lectotype species of this section.
The type specimen was staminate; these collections permit, for the
first time, the description of the pistilate characters.

Sect. Racemosiflorae B. C. Stone

(2) Freycinetia angustifolia Blume, Rumphia 1: 149, t. 43. 1835.
Merrill, 1.c. 19721.

Type from Java. Previously reported from Borneo by Merrill
(Foxworthy 91, from Sarawak; Motley 1128, Kalimantan).

SARAWAK: Lundu, unknown collector (but probably a
duplicate of Foxworthy 91), staminate (SAR!). Simanggang
District, Trisa P.F., small climber reaching 5 ft. high, in peat-swamp
forest, Jan 1958, J. A. R. Anderson 14545 (SAR! ).

INDONESIAN BORNEO (KALIMANTAN): Bandjermasin,
Motley 1128 (K!).

This highly characteristic species can now be confirmed from
the Philippines for the first time:

PHILIPPINES: Sibuyan; Mt. Giting-giting, Prov. Capiz, May
1910, Elmer 12426 (labelled with MS name “‘F. baganot Martelli’
— ined.) (L! EDINB!). — Same location, March 1910, Elmer
12071 (det. as F. elmeri Martelli) (K! EDINB! ).

Stone — Freycinetia (Pandanaceae V1) 215

Sect. Sarawakenses B. C. Stone

(3) Freycinetia sarawakensis Martelli, Webbia 3; 179, 1910. Merrill

(1921) p. 34.

Type from Sarawak (Beccari 1829). Also recorded from Kali-
mantan (Jaheri 1017). Hallier 3188 was also listed; there are two
species under this number in Hallier’s collections. Hallier 3188-bis
represents this species, while 3/88 proper represents (and is the
type of) F. lucida Martelli.

KALIMANTAN: C. E. part at W. Koetai n. 40, on Kiau R. in
riverbank forest, alt. 700 m., fairly common, Oct. 1925, F. H.
Endert 4582 (L!). SARAWAK: Sibu, Lassa, 10 ft. alt. May 1961,
Anderson R-19 & (SAR! ).

This unusual species is the type of Sect. Sarawakenses. No
staminate collections are known so far, but in its curious
lateral and solitary cephalia, and its elongated (50-60 cm long)
leaves, only 6-8 mm wide, it is a striking species. The leaf auricles
are about 25 x 3 mm. The pistillate cephalia are subglobose,
about 30 x 25 mm, and are borne on a stout short pedicel
10 x 4 mm. So far this plant has not been found outside Borneo.

(4) Freycinetia imbricata Blume, Rumphia 1: 157, t. 40. 1835.
Figure 1.

F. kingiana Ridl. Mat. Fl. Mal.’ Pen. 2: 232, 1907.

Type from Java (Blume). Type of Ridley’s F. kingiana is
Kunstler 4654 from Perak, Malaya.

var. hispidula B. C. Stone, Gard. Bull. Sing. 25: 204, 1970.
In pedicellis hispidulis et foliis latioribus differt.
F. schefferi Solms, Linnaea 42: 98, 1879.

Holotypus: MALAYA: Selangor; Ginting Sempah, 22nd mile,
alt. 1,500 ft., climber on rock, July 1966, Stone 5847a (KLU! ).

Exsiccatae: MALAYA: Selangor, type locality, Stone 5847
(KLU! ). Ginting Highlands, 3,000 ft. alt.. Nov. 1966, Stone 6535
(KLU! ).

BORNEO: Sarawak; Kuching, Oct. 1905, Hewitt (SAR! ). Siol,
Apr. 1906, Hewitt (SAR!). Puak, bushes by river, Sep. 1903,
Ridley (SING 026534! ). Simanggang, Triso P.F., alt. 10 ft., small
climber, Sept. 1958, Anderson SAR 14549 (SAR! ). Lundu, Sungei
Rengas, small climber in peat-swamp forest, fruit bright orange,
15 ft. alt., Oct. 1961, Anderson et al. SAR 15421 (SAR! ). Without
locality, Ridley in 1915 (L!). Without locality (but perhaps
Labuan), Mr. (Hugh) Lowe (K!). Sabah; Madai, Governor
Creagh (K! ).

var. kuchinensis (Martelli) B. C. Stone, comb. et stat. nov.

F. kuchinensis Martelli, Webbia 3: 167, 1910. (as Kutchinensis,
p. 316). Merrill (1921) 34.

Type from Sarawak (Beccari P. B. 782-bis). Reported also from
Indonesian Borneo (Jaheri 528).

BORNEO: Sarawak: Gat, upper Rejang River, fruit bright
yellow, 1929, J. & M. S. Clemens 21902 (FI! ).

216 Gardens’ Bulletin, Singapore— XXV (1970)

Fig. 1. Freycinetia imbricata Bl. Java, Korthals (L. 908164).

Stone — Freycinetia (Pandanaceae V1) 217

Sect. Auriculifoliae B. C. Stone

(5) Freycinetia sumatrana Hemsley, Kew Bull. 1896: 158.
F. valida Rid]. Mat. Fl. Mal. Pen. 2: 233, 1907.
Winkler, Engl. Bot. Jahrb. 48: 87, 1912. Merrill (1921) 34.

Type from Sumatra (Beccari). Type of F. valida from Malaya
(Ridley).

BORNEO: Sarawak; Simanggang Distr., Triso P.F., Jan. 1958,
Anderson 14544 (SAR!). Sabah; Tawau Distr., Apas Rd. 12.5
miles, epiphyte in primary forest, fruit dark brown smelling of
meat, Jun. 1959, Meijer SAN 19371 (SAN! K! L! SING! SAR!).
Sepilok Forest Reserve, swamp forest, Mar. 1967. Stone & Meijer
6731 (KLU!).

PHILIPPINES: Basilan Island; Zamboanga Prov., Isobela de
Basilan, Land grant of Univ. Philipp., Liborio Ela Ebalo 988
(PNH! distrib. as F. banahaensis Elm.). Vernacular name, ‘‘kaya-
kan” in Yakan (Jan. 1941).

The cited specimen from Basilan extends this species for the
first time into the Philippines. It is certain that the plants referred
to F. auriculata Merr. are of the same species as this.

(6) Freycinetia lucida Martelli, Webbia 3: 168, 1910. Merrill (1921)
34. Figure 2.

Type from Indonesian Borneo (Hallier 3188) in BO!

Liana scandens; caulibus vetustioribus ramuligeris ad 12-15 mm
diametro, internodiis 15-20 mm longis, radicibus aereis emittenti-
bus, junioribus c. 8 mm diametro, internodiis 10-15 mm longis,
foliosis. Folia longe lineari-attenuata, 45-50 cm longa, 9-13 mm
lata, sensim attenuata, marginibus denticulatis, in sicco revolutis;
parte vaginantia c. 4.5 cm longo; auriculis castaneis 4-5 cm longis
4-5 mm latis apice unilobulatis integris vel ciliatis. nflorescentia
terminalis binatis. Cephalia foeminea cylindracea, immatura
48-50 mm longa, 12 mm crassa; pedunculis angustis 25-27
mm longis 2.5 mm latis distaliter minute et subsparse scabridulosis.
Bacca (immatura) 4 mm longa, 1 mm lata; pileus vix 1 mm altus,
late pyramidatus vertice plano. Stigmata 2 (rare 3, rarissime 4).
Staminodia, minutissima, cetera ignota.

BORNEO: East Indonesian Borneo, Kutei, G. Beratus [Summit,
Piek van Balikpapan] alt. 1,200 m, 19 July 1952, W. Meijer 904
(BO! L!).

This very interesting species is known so far only from the type
collection and the recent one cited made by Willy Meijer.

Gardens’ Bulletin, Singapore — XXV (1970)

218

3

——
a:

Fig. 2. Freycinetia lucida Martelli. Borneo. Meijer 904.

Stone — Freycinetia (Pandanaceae V1) 219

In its lobed auricles, tapered leaves, terminal inflorescence,
cylindric pistillate heads, and berries with usually 2 stigmas, F.
lucida corresponds well with the characters of Sect. Auriculifoliae.
From F. sumatrana Hemsl., it differs in its smaller overall dimen-
sions and entire auricles, and in the different form of spinules
which occur at the distal end of the peduncles. From F’.. ceramensis
Martelli, which is undoubtedly its closest relative, the present
species differs in its slightly smaller yet more strongly denticulate
leaves, more slender pedicels with their distal ends, just below the
cephalium, spinulose rather than smooth or minutely scabrous, and
somewhat fewer and less crowded berries.

Sect. Polystachya B. C. Stone

(7) Freycinetia kinabaluana B. C. Stone, sp. nov.

Liana robusta caulis ad 2 cm diametro, foliis anguste attenuato-
subulatis coriaceis 80-100 cm longis 2.5 cm latis, basi vaginantibus,
auriculis 7-9 (-10) cm longis et 7-10 mm latis, marginibus foliorum
basin versus minime et distanter paucidenticulatis (dentibus vix
0.7 mm longis), apicem versus dorso simillime denticulatis, cetera
laevibus. /nflorescentia foeminea terminalis cephaliis pluribus
plerumque 4~7 spiraliter congestis pedicellis 5 cm longis 4-7 mm
crassis subtrigonatis vel semiteretibus ad angulis et in lineis dor-
salibus scabridulis (papillis 0.2 mm longis) in sicco fuscoatris.
Cephalia breve oblongata vei obscure ellipsoidea juventute breviter
cylindracea, ad 5 vel 6 cm longa et 3.5—5 cm lata. Baccae nume-
rosae maturitate flavidae, ad 20-24 mm longae et 3-3.5 mm latae,
clavatae, pileis 5 mm longis anguste pyramidatis, areola stigmatica
c. 1 x 1 vel 1 x 2 mm, stigmatibus (4-) 5—7 (-9); semenibus ad
1.2 mm longis angustis lineari-ellipticis raphe inconspicuis et
strophiolo nullo.

Holotypus: BORNEO: W. Sabah: Mt. Kinabalu, 5,000 ft.,
Mesilau River, Apr 1964, W. L. Chew & E. J. H. Corner (1964
Royal Society Expedition) RSNB 70/3 (K!). Isotypes (SING!
SAR! SAN! ).

Exciccatae: BORNEO: W. Sabah: Mt. Kinabalu, 5,200 ft.,
below Lumu-Lumu, Apr 1933, C. E. Carr SFN 27104 (SING! ).
Bembangan River, 5,000 ft., Feb 1964, Chew & Corner RSNB
4405 (K! SING! SAN! SAR!). Mt. Kinabalu, Tenompok, 5,000
ft., Mar 1932, J. & M.S. Clemens 28711 (=29550) (L!). Mesilau
River, 5,000 ft., Jan 1964, Chew & Corner RSNB 4177 (K!); Feb
1964, Chew & Corner RSNB 4306 (K! ).

A distinctive large species apparently related to F. corneri
B. C. Stone, F. philippinensis Hemsl. and F. multiflora Merr., and
perhaps also to F. whitmorei B. C. Stone of the Solomon Islands.

In its numerous spirally crowded cylindric cephalia this species
appears to belong in Sect. Polystachyae. In a very superficial
way it also resembles F. sumatrana Hemsl. but differs in many
characters. |

220 Gardens’ Bulletin, Singapore — X XV _ (1970)

Sect. Filiformicarpae B. C. Stone

(8) Freycinetia parviaculeata B. C. Stone, sp. nov. Figure 3.

F. valde proxime F. tessellata et F. ponapensis sed pedicellis
omnino dense scabridulis et foliorum marginibus conspicue late
denticulatis.

Scandens, robusta, caulis ad 15 mm diametro. Folia erecto-
patentia basi amplectentia coriacea ad 60 cm longa et 3.3 cm
lata, apice acuto, margine e basi ad apicem denticulata, dentibus
antrorsibus fere 1 mm longis et 3-7 mm separatis; costa media
dorso apicem versus simillime denticulata, cetera laevia; auriculis
ad 10-12 cm longis et 1-1.5 cm latis, membranaceis, margine ut
videtur integris (sed fragmentatis). /nflorescentia foeminea termi-
nalis breve pedunculata, cephaliis oblongo-cylindraceis ad 7 cm
longis et 3 cm latis, pedicellis 20-24 mm longis et 5 mm crassis,
obscure trigonatis, in toto dense aculeatissimis. Baccae 13-15 mm
longae, 2-3 mm latae (in sicco) apice coriaceo pileo angusto elon-
gato 5-6 x 1-2 mm truncato, parte basilari succulenti seminiferosi
rubri, areola stigmatica elliptico c. 1 mm lato annulo brunneo
nitente cincta, stigmatibus 2 rarissime 3 discretis, seminibus falcato-
lunatis c. 1 mm longis rubro-miniatis, raphe alba angusta, strophiolo
nullo.

Holotypus: BORNEO: Sarawak; Miri District, Sungei Dalam,
low altitude, on tree by stream in swamp forest, fruit bright red,
June 1961,Dan bin Haji Bakar SAR 4502 (SAR!). Isotypes in
|

This species is the first of Sect. Filiformicarpae to be reported
from Borneo, and indeed is the first known west of Wallace’s Line.
It is distinguished from its congeners by the broad and conspicuous
marginal leaf-prickles and the densely aculeate pedicels (see illus-
tration), to which the specific epithet refers.

BORNEO: Sarawak: Rejang Mangrove Reserve, Nov. 1968,
Anderson §. 26572 (SAR! ).

Sect. Hemsleyella B. C. Stone

(9) Freycinetia rigidifolia Hemsley, Kew Bull. 1896: 166. Merrill

(1921) 34. Figure 4.

F. acuminata Ridl. Mat. Fl. Mal. Pen. 2: 233, 1907.

Type of Hemsley’s species was from Sarawak (Haviland 436);
that of Ridley’s from Malaya (Ridley 7656).

The species has been reported from Indonesian Borneo (Jaheri
1017).

Recent collections disclose that this is evidently quite a common
species in Borneo. and that it also occurs in the Anambas Islands
and in the Andaman Islands.

Stone — Freycinetia (Pandanaceae V1) 221

anf
t,

Vi

\ ’ j i

iid
Wg

Fig. 3. Freycinetia parviaculeata B. C. Stone. Holotype. Right: leaf
apex. Center: part of fruit pedicel, showing spinules; below,
enlarged view of spinules. Left, above: berry in profile and top
view, with seed in profile to same scale; below, seed enlarged.

222 Gardens’ Bulletin, Singapore — XXV _ (1970)

BORNEO: Sabah: Mt. Kinabalu; Tenompok, 5,000 ft., March
1932, Clemens 26075 (K!); 26378 (K!); 28750 (L!); 29331 (K!):;
30175 (K! L!). Penibukan, 4,000 ft., Sep 1932, Clemens 40510
(L!). Marai Parai, 9,000 ft., Mai 1933, Clemens 32385 (L!).
Mesilau River, 5,000 ft., Feb 1964, Chew & Corner RSNB 4269
(SAN! SING! ). Ranau Distr. along Kambarango Rd, Aug 1963,
Mikil SAN 38635 (K! SAN!). Sandakan Distr. 25 mi SW of
Sandakan, Lungmanis, Mar 1955, G. H. S. Wood SAN A-2922
(SAN! ). Kabili-Sepilok Forest Reserve, Mai 1961, Patrick Ping
Sam SAN 24845 (SAN! ). Ranau, Mesilau trail, on tree NT-980,
4,500 ft., Feb 1965, Sadau SAN 42888 (K! SAN! ). Sarawak; Bau,
limestone hills, 1893, Ridley 11712 (K!). Same location, Jul 1963.
Quisumbing PNH &7630 (PNH!). Bidi cliff, C. J. Brooks (K! ).
Gunong Subis, Niah, 400 ft., on ‘mor’ layer at summit of limestone
hill, bracts bright orange, June 1962, Anderson S. 16426 (K!
SAR! ). Bau District, 700 ft. Aug 1963, Chew 56] (K!). Near
Kuching, Dec, 1892, Haviland 3132 (K! SAR!). Near Long Kapa
Mt. Dulit, Ulu Tinjar, 700-900 m, bracts scarlet, P. W. Richards
1964 (L!) KALIMANTAN: W. Koetai, n. 39, nr. Mt. Kemoel,
1,600 m., Oct. 1925, Endert 4374 (L!); 4136 (L! ).

ANAMBAS ISLANDS: Boengoeran, east side of Gunong
Ranai, Apr 1928, van Steenis 1200 (L!).

ANDAMAN ISLANDS: S. Andaman, Kurz (FI! ).

This very well characterized species seems to be one of the most
common species in Borneo. It is also in Malaya, where it has been
known by the name F. acuminata Ridl. Its closest relative, and the
only other member of Sect. Hemsleyella, is F. pectinata Merr. &
Perry of the Solomon Islands.

Of the specimens cited, a few deserve particular mention. Several
of the Clemens collections, particularly 40510, 40452 and 40453,
all from Mt. Kinabalu, are somewhat unusual in their narrower
leaves (about 17 cm long and 5.5 mm wide) and the short auricular
spinules, differences which however noticeable appear to grade
continuously into those of larger plants. The extreme of size
increase is afforded by the Richards 1964 specimen, which has
leaves 44 cm long and 18 mm wide.

Freycinetiae Imperfectae vel Incertae Sedis

(10) Freycinetia palawanensis Elmer, Leafl. Philipp. Bot. 1: 216,
1907, var. andersoniana B. C. Stone var. nov.

Liana scandens, caulis ad 1 cm diametro, internodiis ad 2 cm
longis, foliis anguste lanceolatis ad 42 (vel ultra?) cm _longis
et 2-2.5 cm latis apice acuminato-caudato, basem versus leviter
angustatis, marginibus inermibus apicem versus excepta; cauda
minute denticulata; auriculis delapsis ad 3 cm longis. /nflorescentia
terminalis breve-pedunculatis binis vel ternis pedicellis brevis c.
9 x 5 mm laevibus. Cephalia foeminea cylindracea ad 9 cm longa

Stone — Freycinetia (Pandanaceae V1)

Fig. 4. Freycinetia rigidifolia Hemsley. Habit (SAN. 59284).

224 Gardens’ Bulletin, Singapore — X XV _ (1970)

et 1.8 cm crassa. Baccae immaturae 6 mm longae lageniformatae
in pileo abrupte angustato 2 mm longo terminatae, areola stigma-
tica 1 mm lato subtruncato, stigmatibus 4-6, parte basilari semini-
feri semenibus 1.5 mm longis angustis sublinearis raphe perangusto
strophiolo nullo.

Holotypus: BORNEO: Sarawak; Baram District, Ulu Sungei
Melinau Paku, 400 ft., climber on riparian hill, Jun 1961, J. A. R.
Anderson SAR 4076 (SAR! unicate).

Exsiccatae: PHILIPPINES: Mindanao; Bukidnon Province,
Mt. Katanglad, Mar 1949, M. D. Sulit PNH 10082 (PNH! ).

This is a very distinct species. The new variety is named in
honor of the Conservator of Forests for Sarawak, who collected
the type specimen. It differs from the commoner Philippine variety
in its broader, longer Jeaves and larger fruiting heads.

(11) Freycinetia biloba B. C. Stone, sp. nov.

Liana scandens caulis ad 8 mm diametro trigonatis, internodiis
1-2 cm longis, foliis erecto-patentibus 23-34 cm longis, 2.5—3.6
cm latis oblanceolatis acuminatis cuspidato-caudatis, fere inermibus
caudo (3-4 cm longo) et marginibus basalibus per brevem spatiam
(1-2 cm) excepta, dentibus minutis 0.1—-0.2 mm longis obtusatis
1-5 mm separatis, costa media cariniformato laeve sed apicem
versus minutissime denticulato. Auriculae ad 25 mm longae et
6-7 mm latae, membranaceae mox caducae leniter rotundata
marginibus creberrima minutissimeque ciliato-denticulatis. Inflores-
centia ternata terminalis, pedunculis communis 20-25 mm longis
ad angulas dense hispidulis, pedicellis 15-20 mm longis 2.5 mm
crassis subteretibus omnino (basem extremam excepta) stramineo-
hispidulis (setis ad 0.5 mm longis antrorsis congestis aciculari-
ventricosis). Cephalia cylindracea immatura ad 30 x 8 mm.
Baccae numerosae, pileis ad 1 mm latis, stigmatibus (3-) 4-8
(-10) vulgo 4-6.

Holotypus: BORNEO: W. Sabah, Beaufort District, Beaufort
Hill, in primary forest, Mar 1961, Meijer SAN 24834 (SAN! ).

Exsiccatae: BORNEO: Sarawak; Sungei Belaga, hillside with
mixed dipterocarp forest, inflorescence chrome yellow, Apr 1963,
P. S. Ashton S. 18229 (K!). — Mt. Mattang, primary forest, fruit
red-orange, 5 April 1968, J. Dransfield 763 (KLU! ).

This species appears similar to F’. ciliaris Martelli, despite the
major difference in facies caused by the much longer and more
numerous spinule-like cilia on the leaf of the latter. F. biloba also
somewhat resembles F. jaheriana Martelli (of New Guinea) but
has berries with more numerous stigmas and considerably narrower
Jeave. However the nearest relative is F. kamiana Stone of
Malaya. The specific epithet is in reference to the two short con-
spicuous auricles.

Stone — Freycinetia (Pandanaceae V1) 225

(12) Freycinetia borneensis Martelli, Webbia 3: 167, 1910. (as
bornensis; corrected, 1.c. 310). Merrill (1921) 34.

Type from Indonesian Borneo (Jaheri 925) at Leleboelan Tepoetz
(1896-97). A fragment in the Martelli Herbarium in Firenze has a
leaf 16 x 2.8 cm, abruptly cuspidate-caudate, with the cauda 2
cm long; basal auricles 24 x 4-5 mm, apparently rounded; imma-
ture berries 5 x 2.5 mm, pileus narrowed abruptly below the short
stigmatic areola; and stigmas 2 or 3 per berry.

BORNEO: Central E. part, W. Koetai, n. 26, L. Temelin, low
riverbank forest, 200 m. common climber, Aug. 1925. Endert
2918 (L!). Z. en O. Afd. Borneo, km. 14, tusschen Bandjermassin-
Martapoera, Oct. 1939, Polak 513 (L!). — Sarawak; G. Matang,
800 ft. alt., primary forest, clay-loam, bracts orange-pink, spikes
pale pink, April 1966, Anderson S. 25102 (SAR! ).

(13) Freycinetia ciliaris Martelli ex Merrill, Univ. Calif. Publ. Bot.
tb te id of ok

Type from Tawao, Sabah (Elmer 21632).

Stems to 5-8 mm thick; leaves oblanceolate, 12-20 cm long,
(1.3-) 2-3 cm wide, caudate-acuminate, acurnen 2-3 cm long,
spinulose; margins (except medially) spinulose; auricles 11—14
mm long, 4-5 mm wide, inarched at apex, pectinate-spinulose
with setae 1-1.5 (or more) mm long, caducous. /nflorescence
ternate, terminal, the pedicels hispidulous, berry with 3-7 stigmas
on a dome-like pileus.

BORNEO: Sabah; Sandakan District, Paitan Forest Reserve,
50 ft., primary forest nr. swamp, Feb. 1963, Ampuria SAN 32085
(K! L! SAN! ). Sepilok Forest Reserve, lowland swamp, uncom-
mon low climber, bracts greenish-white, male spadices 3, oblong,
1 cm long, March 1967, Stone & Meijer 6713 (KLU! ).

One of the most characteristic of the Borneo species, and indeed
very distinct in the genus.

(14) Freycinetia concinna Martelli ex Merrill, Univ. Calif. Publ.
Bot. 15: 16, 1929.

Type from Tawao, Sabah (Elmer 2/596).

BORNEO: Sarawak; Tatau District, Kakus-Pandan Forest
Reserve, on exposed rocky ridge in ‘‘kerangas’”’ forest, inflorescence
vellowish-green, 1,300 ft., Oct. 1959, E. F. Brunig S. 11954 (SAR!).

(15) Freycinetia corneri B. C. Stone, Brittonia 20 (3): 199, 1968.
Type from Singapore (Corner).

BORNEO: Central E. pt., W. Koetai, n. 39, Mt. Kemoel, alt.

1,800 m, mountains in forest, big climber, Oct. 1925, Endert 4453
(L! ).

226 Gardens’ Bulletin, Singapore —XXV (1970)

This species hitherto known only from Singapore and Johore
now appears in Borneo. Among the large robust species it is
quickly distinguished by its smooth pedicels, subglobose cephalia,
4—8-stigmatic berries, and broad acute leaves with the subtruncate
auricles slightly denticulate at the apex.

(16) Freycinetia discoidea Martelli, Leafl. Philipp. Bot. 111 (Art.
60): 1113, 1911. Figure 5.
Type from Sibuyan Island, P.I. (Elmer 12197).

BORNEO: Sabah; Sandakan District, Leila Forest Reserve,
300 ft., Oct. 1951, Patrick Ping Sam SAN 27822 (SAN! ). Sepilok
Forest Reserve, Sepilok Besar, in swamp with Pandanus corneri
Kaneh., March 1967, Stone & Meijer 6733 (KLU! both stam. and
pistill. plants). Happy Valley, near Leila, Apr 1965, Meijer
4828] (K!).

This is a striking species here first reported tentatively from
Borneo. The leaves are markedly glaucous on their undersurface.
about 38 cm long and 2.5 cm wide, broadly linear, with auricles
50 x 5 mm having entire margins though a very few spinules
occur at the truncate apex. The floral bracts are pinkish-orange;
pedicels are smooth, 15—20 mm long; cephalia about 50 x 14 mm;
and the berries have 2—5, but commonly 2-3, stigmas. One dis-
cordant feature is the indication of scabrid pedicels in the original
description, while our specimens have quite glabrous pedicels.
Conceivably this is an important difference.

(17) Freycinetia hemsleyi Warburg, Pflanzenr. IV. 9: 36, 1900.

Merrill (1921) 34.

F. beccarii Hemsl. Kew Bull. 1896: 166, not of Solms 1879.

Type from Sarawak (Beccari P. B. 3598).

This plant has not appeared again in more recent collections.
It is probably a close relative of F. javanica Bl. and perhaps with
that species should form a section. It has however much broader
leaves of more oblanceolate form, and fewer stigmas per berry.
The following brief description, improving somewhat on Hemsley’s
and Warburg’s, is added, with the hope that the species may be
again collected:

Stems to 5-6 mm _ diameter; leaves _ elliptico-oblanceolate,
broadest beyond the middle, about 3 times longer than wide, up
to 22.5 x 7.5 cm, more often about 16 x 4.5—5 cm, slightly cus-
pidate; auricles short, about 11 x 4.5 mm, denticulate along their
margins and abruptly rounded at apex; Jeaf-veins widely spaced,
20-25 per leaf-half, from 1-1.9 mm apart; inflorescence terminal
and ternate, the pedicels about 20 x 3 mm, scabridulous on the
angles, and all around distally; immature cephalia about 15 x 7
mm; berry with 2-4, or rarely 5, stigmas.

Stone — Freycinetia (Pandanaceae V1) 227

~~
SQ

Fig. 5. Freycinetia discoidea Martelli. Habit (P. P. Sam 27822).

228 Gardens’ Bulletin, Singapore — XXV (1970)

(18) Freycinetia javanica Blume, Rumphia 1: 156, t. 41. 1835.
Merrill (1921) 34.
Type from Java (Blume).
var. javanica

BORNEO: Eastern pt., Nunukan, N. of Tarakan, inner forests
at low altitude, with dipterocarps, a climber on Oncosperma, Nov.
1953, Meijer 2100 (L!).

var. expansa B. C. Stone, var. nov.

Ab var. typica in foliis valde latioribus differt.

Holotypus: BORNEO: Sarawak; Stepok Forest Reserve, nr.
Kuching, peat-swamp forest, Apr 1967, Stone, Anderson, & Chai
6773 (KLU! ).

Exsiccatae: BORNEO: Sarawak; Near Kuching, Nov. 1892,
Haviland 1970 (K!); Quop, Apr 1907, Hewitt (K! SAR). Kuching,
Mar 1893, Haviland 3131 (SAR!).

These plants cannot be specifically segregated from Blume’s
F.. javanica, yet they differ from most of the Javan, Malayan, and
Sumatran collections I have seen in their broader leaves. However
it is somewhat doubtful that this feature is maintained throughout
Sarawak; certainly the Meijer specimen from East Borneo appears
to be the typical variety.

(19) Freycinetia robinsonii Merrill, Philipp. J. Sci. Bot. 3: 311,
1908.

Type from Luzon, P.I. (Forestry Bureau 3037, Borden).

Represented in Borneo only by the following variety.

var. meijeri B. C. Stone, var. nov.

Liana scandens, caulis ad 7 mm diametro. Folia anguste lanceo-
lata attenuata 27-41 cm longa 9-13 mm lata, tenuiter coriacea,
basi amplectentia, marginibus, basem apicemque, minute et sub-
sparse denticulata, tenuiter coriacea, dentibus ad 0.4 mm longis,
1-2 mm separatis; auriculis 28-45 mm longis, basem versus 5-6
mm latis, chartaceis, brunneis, attenuatis, integerrimis, lateraliter
fragmentatis. /nflorescentia foeminea terminalis, ternatis, pedicellis
23-28 mm longis, 2—2.3 mm crassis, distaliter ad angulas hispidulo-
scabridis; cephaliis cylindraceis 25-40 mm longis, 7-9 mm crassis
(immaturis), baccis numerosis, pileo sulcato pyramidato areola
stigmatica 1 mm lato, stigmatibus 2-3, plerumque 2. Inflorescentia
mascula simillima sed parviora ternata, pedicellis c. 24 mm longis,
1-1.5 mm crassis, laevibus, spicibus c. 15 x 3 mm staminibus
numerosis filamentis c. 1 mm longis, antheris subgloboso-oblongis,
0.2 mm longis; bracteis Janceolatis.

Stone — Freycinetia (Pandanaceae V1) 229

Holotype: BORNEO: Sabah; Penampang District, Sangai-
sangai, Babagon Rd., c. 4,000 ft. alt., primary forest, near riverside,
fruit green, 17 March 1965, J. K. Laiangah SAN 44362 (L!).
Isotype SAN!

This species is very close to F. vidalii Hemsl. of the Philippines,
differing in the shorter and slightly wider leaves; and also super-
ficially resembles F’. apayaoensis Merr., another Philippine species,
which however has stiffer, more evidently denticulate, lineate,
ensiform leaves. But F. robinsonii is closest to F. multiflora Merr.

BORNEO: Sabah; Tamu Darat, 7 miles on Kota Belud —
Ranau Rd., 200 ft., by river in shade, flowers deep salmon with
yellow anthers, Jan. 1954, S. Darnton 77 (BM! L! Staminate
paratype). — Diwata River 10 mi. WSW of Lahad Datu, “‘porum-
pung” (Dusun Kinabatangan dialect), Feb. 1954, Rapag SAN
A-i816 (SAN). Sandakan District, Sungei Meliau near Kiabau,
50 ft. alt., inflor. orange-red, May 1965, Meijer SAN 49834 (SAN).

This new variety is dedicated to Willy Meijer, formerly Forest
Botanist at Sandakan, who provided much help in the field on the
author’s excursions in 1967.

(20) Freycinetia creaghii Hemsley, Kew Bulletin 1896: 167. Merrill

(1921) 34.

Liana scandens, caulis ad 12 mm diametro. Folia erecto-patentia
tenue coriacea basi amplectenia vaginata, ad 45 cm longa et 3 cm
lata, late linearia sed basim versus paullulo angustata apicem
versus subabrupte acutata non acuminata, marginibus inermibus.
(in base apiceque per brevem spatiam excepta dein perminute
persparse denticulatis); auriculis ad 4 cm longis et 4 mm (vel
ultra?) latis, ut videtur fragmentatis. /nflorescentia foeminea ter-
minalis breve-pedunculata cephaliis ternis subellipsoideis 2.5 cm
longis et 1.5 cm latis, pedicellis 16 mm longis et 2 mm crassis,
minute spinuloso-scabridis. Baccae immaturae 6 mm longae, pileo
anguste obconico 2.5 mm alto in sicco leviter costulato. Areola
stigmatica 0.5-0.8 mm lata stigmatibus 3-5 vulgo 3 bene distincta.

Type: BORNEO: Sabah: Madai, Governor Creagh (K!)
staminate.

BORNEO: Sarawak; Kuching District, 300 ft., Semengoh Forest
Reserve, lowland primary dipterocarp forest, May 1961, Ghazalli
S. 13676 (SAR! ).

The type is a staminate specimen, which may account for the
fact that no other collections have ever been identified as this
species. The above collection permits fuller description and shows
the characters of the pistillate form.

230 Gardens’ Bulletin, Singapore— XXV (1970)

(21) Freycinetia winkleriana Martelli, Webbia 2: 168, 1919.
Winkler, Engl. Bot. Jahrb. 48: 87. 1912. Merrill (1912) 34.
Engl. Bot. Jahrb. 48: 87. 1912. Merrill (1921) 34.

Type from Indonesian Borneo (Winkler 3313).

Stems to 5 mm diameter; leaves slender, linear, rather stiffly
erect, up to 70 cm long, 10 9-11 mm wide, apex long attenuate-
subulate and subflagellate, triquete, leaf-base marginal teeth (just
above auricles) c. 0.5 mm long; median margins unarmed: apex
denticulate, teeth 1-2 mm apart; midrib distally dorsally spinulose.
Auricles slender, up to 45 x 3 mm, entire, slightly rounded to
subtruncate at apex, often brownish-purplish. Inflorescence ter-
minal, cephalia usually 3 but occasionally 4, on pedicels about
20 x 2 mm, closely and finely hispidulous at distal ends; immature
pistillate cephalia cylindric, about 20 x 4.5 mm. Berries usuaily
with 2, sometimes with 3, rarely with 4 stigmas.

BORNEO: Sarawak; Bintulu District, Bukit Urang, common
climber, spathes purple, Apr 1960, Brunig S. 12033 (K! SAR!).
Kuching, 1911, Salub (SING-026532!). Kuching, “‘native collector,”
475 (PNH!). KALIMANTAN; near Mempawa, peat forest
behind Kampong Sungei Koenjit, Oct. 1940, Polak 704 (L!).

SUMATRA: Indragiri uplands, Berapit, primary forest, Apr
1939, Buwalda 6354 (L! ).

The above specimen first extends the range to Sumatra.

(22) Freycinetia gitingiana Martelli, ex Elmer, Leafl. Philipp. Bot.
Sari, Toit.

BORNEO: Sabah; Mt. Kinabalu, Kokolitan, 2,500 ft., Mai
1953, C. E. Carr SEN. 27342 (SiG }. |

This plant seems probably to be of Philippine affinity, and in
particular resembles F. gitingiana Martelli. Because of this, and in
view of the imperfect state of our knowledge of the Philippine
Freycinetiae, I equate it with that species tentatively. The field
notes indicate that it is a species with “‘golden yellow fruits” which
seems to be a rather rare color.

(23) Freycinetia kalimantanica B. C. Stone, sp. nov. Figure 6.

Liana scandens, caulis ad 1 cm crassis; foliis c. 50-60 cm longis,
c. 18 mm latis, attenuatis, paene inermis, marginibus base perpauce
minuteque denticulatis (dentibus 2-3), apice extremam minute
denticulato; nervis c. 40-44; auriculis subpersistentibus, usque ad
12 cm longis, attenuatis, inermibus, brunneis subnitentibus, sectilis
ve] fissis in segmentibus, ultime caducis, base 12-13 mm latis.
Inflorescentia terminalis ternatis, cephaliis oblonge-cylindraceis,
pedicellis 12-15 mm longis, 5 mm crassis, dense minuteque aureo-
brunneo-hispido-scabridis; cephaliis 3.5-4 cm longis, 1.5-1.8 cm
crassis; braccis plurimis immaturis 4.5 mm longis, pileo plano 0.5
mm alto, stigmatibus plerumque 5-8. Cetera ignota.

231

Stone — Freycinetia (Pandanaceae V1)

Fig. 6. Freycinetia kalimatanica B. C. Stone, Holotype, Endert 4184.

232 Gardens’ Bulletin, Singapore — X XV (1970)

Holotype: BORNEO: W. Koetai no 38, 1,500 m alt., 15 Oct.
1925, F. H. Endert 4184 (BO! ).

Very distinct among the Borneo species by the broad, rather
shining brown auricles, which fragment into segments by lateral
splits, and are evidently entire. The leaves are virtually unarmed.

The name is based on the Indonesian name for Borneo, Kali-
mantan.

This species is evidently closely allied to F. corneri Stone but
differs in the densely scabrid 9 pedicels and smaller leaves and
fruits.

(24) Freycinetia confusa Ridley, Mat. Fl. Malay. Pen. 2: 234,
1907. |
Not F.. confusa Elmer 1907 (renamed by Elmer F. villarii 1908).
Type from Singapore (Ridley 4757, lectotype, SING! ).
Represented in Borneo by the following variety.
var. minima B. C. Stone, var. nov.
Foliis brevioribus plerumque 12-16 cm longis.

Holotype: BORNEO: Sarawak; Nanga Balang, extreme head-
waters of Batang Balleh, Kapit District, 950 ft. alt., 20 yds. from
riverbank, dense root-climber to about 20 ft., bracts green tinged
purple at tips, 29 June 1969, J. A. R. Anderson S. 29301 (KLU!
isotypes SAR, K).

The species proper is known from Malaya and Sumatra; this is
its first report in Borneo, but the specimens cited all agree in the
shorter leaves (only about half as long as in Malayan and
Sumatran specimens seen) and a varietal status is thus proposed.

In its very narrow leaves it cannot be confused with other
species except perhaps F. sarawakensis, which has much longer
leaves and lateral inflorescences usually of only one cephalium;
or F. winkleriana, which has longer, slightly wider, more rigid,
and basally purple-tinged leaves with much narrower auricles
(those in F. confusa are pale or stramineous, reach 2-3 mm width,
and are sometimes truncate at apex and minutely ciliolate, while
those of F. winkleriana are only 1 mm wide at apex, entire, and
often purplish). Another similar plant is F. imbricata var.
kuchinensis, but it has somewhat wider, more lanceolate leaves,
with the margins sometimes recurved.

An additional specimen:

BORNEO: Sarawak; ridge between Sungei Balang and Sg.
Balleh in extreme headwaters of Balleh, Kapit District, 1,200 ft.
alt., sandstone-derived soils, small root-climber, 30 June 1969,
Anderson & Ilias Paie §. 28334 (KLU, L, SAR).

Stone — Freycinetia (Pandanaceae V1) 233

References

MARTELLI, U. 1929. Pandanaceae. /n Merrill E. D. 1929. Plantae
Elmerianae Borneenses. Univ. Calif. Publ. Bot. 7/5.

MERRILL, E. D. 1921. A Bibliographic Enumeration of Bornean
Plants. J. Str. Br. Roy. Asiat. Soc. Spec. No. 1-637.
(Cf. pp. 34-37).

MERRILL, E. D. and Perry, L. M. 1939. On the Brass Collections
of Pandanaceae from New Guinea. J. Arn. Arb. 20 (2):
139-186, 2 pls.

——. 1940. Plantae Papuanae Archboldianae II. J. Arn. Arb.
215@):02863—200,,1 pl.

STONE, B. C. 1967. Materials for a Monograph of Freycinetia
Gaud. (Pandanaceae). I. Gard. Bull. Sing. 22 (2):
129-153.

1968. Materials for a Monograph of Freycinetia
(Pandanaceae) IJ. The correct name for the Freycinetia
of Tahiti. Taxon 17: 174-176.

. 1968. Materials for a Monograph of Freycinetia
(Pandanaceae) III. Brittonia 20 (3): 198-203.
—_—. 1968. Materials for a Monograph of Freycinetia

(Pandanaceae) IV. Blumea 16 (2): 361-372.

. 1969. Materials for a Monograph of Freycinetia
(Pandanaceae) V. Malaya and Thailand. Gard. Bull.
Sing. 25 (2): 189-207.

+h

Further Observations on Ancistrocladus tectorius
(Ancistrocladaceae)

By HsuaN KENG

Department of Botany
University of Singapore, Singapore

In a previous paper (Keng 1967), the general habit, flower, fruit,
seed and seedling of Ancistrocladus tectorius (Lour.) Merr.
(Ancistrocladaceae) were described. No explanation was
given to the nature of its hooked branches (see Fig. 1, b and c),
from which the generic name is derived.

In a recent trip (September, 1968) to Batu Limbang, southern
Johore, Malaya, the present writer observed that the sympodial
cirrhate branches of Ancistrocladus tectorius, during that time of
the year, often bear an enlarged tip on the lower segment or
segments. A close examination reveals that each of these enlarged
tips actually represents a reduced flower (see Plate 1, A and B),
with purple perianth and aborted androecium and gynoecium.
Some of these reduced flowers were fully unfolded, but were all
closed and mostly dropped off after bringing back from the field.

A completely developed inflorescence of Ancistrocladus is a
profusely branched dichasium (fig. 1 a). It is likely, therefore, that
the sympodial cirrhate branches represents the modified peduncles
and pedicels of a partially developed inflorescence. Further evi-
dence is found in a rather unusual specimen collected by the
present writer during a still recent (November, 1968) trip to Pulau
Langkawi, Kedah, Malaya. In this specimen (Plate 1, C), the
cirrhate branches are much excessively ramified, comparable closely
to a fully developed inflorescence.

Another well-known example which possesses somewhat similar
structure is Artabotrys (including A. suaveolens Bl. and other
species) of the Annonaceae (Sinclair 1955). In Artabotrys, the
flattened and hooked peduncles bear flowers on the outer surface
of the curved hooks. Although Annonaceae and Ancistrocladaceae
are both characterized by the possession of ruminate endosperm
in their seeds, their flowers, however, are so vastly different. It is
rather difficult to conceive that these two families are closely
related. The more plausible alternative explanation would be that
the modification of peduncles into hook-like structures in these
two groups is a result of convergent evolution. (The presence of
ruminate endosperm in the seed is, however, probably due to the
maintenance of an ancient character).

236 Gardens’ Bulletin, Singapore — X XV (1970)

Figure I. Ancistrocladus tectorius (Lour.) Merr.

a: spurred branches with dichasial inflorescence; b & c: sympodial
cirrhate branches; from the axils of hooks, the foliaged spurred
branches arise; d: flower, note the outer sepals with large glands;
e: half-flower, showing the cone-shaped stylar bases and the inferior
ovary; f: two stamens of different length, the one opposite sepals is
slightly longer; g: samaroid fruit, winged by persistent accrescent
calyx.

LOETRIRY LO. Saris
Sranennnnnnnnntinnnneineoeey

Plate 1. Ancistrocladus tectorius (Lour.) Merr.

A. Terminal portion of an actively growing plant, showing several
sympodial cirrhate branches. In one branch the lower hook (marked
with an arrow) bears an enlarged tip.

B. Portion of A, enlarged (scales in mm), showing the enlarged
tip which is an aborted flower, with normal but smaller perianth and
reduced androecium and gynoecium.

C. Herbarium specimen, the cirrhate branches are excessively
ramified, comparable closely to a fully developed inflorescence.

*

Keng — Ancistrocladus tectorius 237

Incidentally, a further note on phylogenetic consideration of the
family Ancistrocladaceae could be made here. The diplostemonous
androecium in Ancistrocladus (Keng 1967, fig. 1) is probably
fundamentally different from that of Linaceae-Hugoniaceae — the
androecium of the latter group is mostly ob-diplostemonous
(Melchior 1964). Based on their comparative studies of wood,
Gottwald and Parameswaran (1968) suggested a possible relation-
ship between Ancistrocladaceae and Dioncophyllaceae.

Line drawings and photographs, based on and taken from the
fresh and herbarium materials, are presented as Figure 1 and Plate
1. To Mrs. R. S. Keng who prepared the drawings and to Mr.
D. Teow, who took the photographs, the author is indebted.

Literature cited
1. Gottwald H. and N. Parameswaran, 1968
Das sekundare Xylem und die systematische Stellung der

Ancistrocladaceae und Dioncophyilaceae. Bot. Jahrb. 88:
4-69.

2. Keng; H:, 1967

Observations on Ancistrocladus tectorius. Gard. Bull.
Sing: 22: 9113=121.

3. Melchior, H., 1964

A. Engler’s Syllabus der Pflanzenfamilien, 12 Aufl. Berlin:
Gebruder Borntraeger.

4. Sinclair, J., 1955

A revision of the Malayan Annonaceae. Gard. Bull. Sing.
14: 149-516.

“

The Genus Orchidantha (Lowiaceae)

By R. E. HOLTTUM
Royal Botanic Gardens, Kew, U.K.

1. Historical Account

This genus was based on the species Orchidantha borneensis,
described by N. E. Brown in 1886, from a cultivated plant. Only
two days later appeared Scortechini’s description of Lowia
longiflora, a new genus and species from Perak. The two species
are quite distinct, but certainly congeneric. In 1893 Ridley pub-
lished another generic name Protamomum for a third species, P.
maxillarioides, which he found in Pahang. When describing the
latter, and knowing of Scortechini’s species (though apparently
not that of N. E. Brown) Ridley recognized that Lowia and
Protamomum should belong to a distinct family, which he named
Lowiaceae, but no formal description was given. The family is
near Musaceae, agreeing with Musa in having five stamens with a
distinctive large petal on the same radius as the missing stamen,
but in other ways is very different from Musa. In his monograph
of Musaceae K. Schumann (1900) formally established a subfamily
Lowioideae. The family name Lowiaceae did not receive a latin
description until Nakai published one in his conspectus of
Scitamineae (Nakai 1941); his description is based on Ridley, and
is in part inaccurate, in part very vague, and is quite inadequate
to characterize the family though it formally validates the name.

In 1924 appeared the fourth volume of Ridley’s Flora of the
Malay Peninsula, in which two species of Orchidantha are des-
cribed. One of these was called O. longiflora, but the species
described was different from Scortechini’s. Henderson found
Scortechini’s species again in Perak and published a description
of it as a new species O. calcarea in 1933, recognizing that it was
distinct from Ridley’s description and not having seen Scortechini’s
original one.

In 1944 I studied plants of all three Malayan species in cultiva-
tion in the Botanic Gardens, Singapore, and wrote descriptions,
discovering the confusion about the name OQ. longiflora; I also
examined the branching of the inflorescence in these plants, which
had not previously been described. My observations were however
not published.

240 Gardens’ Bulletin, Singapore —XXV (1970)

In 1955 Irwin Lane published a paper in which he discussed
the distinctions between genera in Musaceae (in Schumann’s broad
sense of the family) and gave reasons for the family status of
Lowiaceae. In this paper he published the first description of the
branching of the inflorescence in Orchidantha, agreeing with my
own observations. He also noted that the extension of the ovary,
to which the floral parts are attached, is solid, not tubular as
described by Ridley (N. E. Brown had also noted this in 1886).

In 1959 Tomlinson published detailed observations on the
anatomy of Musaceae and allied families, giving new evidence for
the distinctness of Lowiaceae, and in 1962 a discussion on the
inter-relationships of all the families of Scitamineae (or Zingi-
berales). These observations were the basis of his account of the
group of families (and others) in 1969. Both Lane and Tomlinson
referred to Nakai’s publication above-mentioned, but cited it
inaccurately.

In 1961 Larsen published descriptions of two new species of
Orchidantha, from Thailand and Laos, with illustrations. His
diagram of the branching of the inflorescence of O. laotica does
not agree with my observations and those of Lane.

In 1969 Dr. Hsuan Keng published observations on plants of
O. longiflora in cultivation in Singapore, with drawings and a
photograph. He stated that the labellum remained rigid for only
6-8 hours, “then turned yellow and curled up and withered
immediately afterwards’. My observations on plants of O. fimbriata
and O. maxillarioides (which had flowers on longer inflorescences
than shown by Dr. Keng) were that the flowers lasted two days
in an expanded condition, though they were limp on the second
day. There is an inaccuracy in Dr. Keng’s floral diagram (fig. 1 g)
which shows the missing stamen on the radius of the dorsal sepal:
it should be on the radius of the labellum, its position thus agree-
ing with the missing stamen in Musa.

The following account of the general morphology of Orchidantha
is based on my own observations of living plants in Singapore.
Following it is a key to all known species, based partly on Larsen’s
observations, and a cescription of each species, those of Malaya
being dealt with in detail, the rest more briefly. At least one more
species exists in Borneo; a specimen of this (Keith 10013, Kabili
Forest Reserve; Sandakan) bearing a fruit but no flowers, is in
the Singapore herbarium.

Literature cited above

Brown, N. E. 1886 (23 October). — Orchidantha borneensis, a
new genus of Scitamineae. Gard. Chron. N.S. 26: 519.

HENDERSON, M. R. 1933. — Additions to the Flora of the Malay
Peninsula. Gard. Bull. Str. Settl. 7: 87-128.

.
° = .

Holttum — Orchidantha 241

Kena, H. 1969. — Notes on the flowers of Orchidantha longiflora
(Lowiaceae). Gard. Bull. Sing. 24: 347-349.

Lang, I. E. 1955. — Genera and Generic Relationships in Musa-
ceae. Mitt. Bot. Staatssamml. Miinchen 13: 114-141.

LARSEN, K. 1961. — New species of Veratrum and Orchidantha
from Thailand and Laos. Bot. Tidsskr. 56: 345-350.

NaKAlI, T. 1941. — Notae ad Plantas Asiae Orientalis (XVI). J.
Jap. Bot. 17: 189-210 (Lowiaceae on p. 201).

RIp.ey, H. N. 1893. — On the Flora of the eastern coast of the
Malay Peninsula. Trans. Linn. Soc. Il Bot. 3: 267-408
(Lowiaceae p. 383).

1924. — Lowiaceae, in Flora of the Malay Peninsula
4: 291-293.

SCHUMANN, K. 1900. — Musaceae, in Engler, Das Pflanzenreich
IV, 45: 1-42.

SCORTECHINI, B. 1886 (25 October). — Descrizione di nuove
Scitamineae trovate nella Peninsula Malese. Nuov. Giorn.
Bot. Ital. 18: 308-311.

TOMLINSON, P. B. 1959. — An anatomical approach to the classi-
fication of the Musaceae. Journ. Linn. Soc. Bot. 55:

779-809.

TOMLINSON, P. B. 1962. — Phylogeny of the Scitamineae —
Morphological and Anatomical considerations. Evolution
16: 192-213.

——— 1969. — Anatomy of the Monocotyledons (ed. C. R.
Metcalfe) vol. If11 Commelinales —- Zingiberales. Claren-
don Press, Oxford.

2. Description of the genus

Leaf-bearing stems are short, erect or suberect, often bearing
lateral branches so that the plants are tufted; each branch bears
first a short 2-keeled prophyll backing the axis from which it
arises. Roots are relatively thick; air-canals are present, but not
conspicuous as they are in Musa. Leaves are 2-ranked, their
petioles rather long with sheathing bases which are very close
together on the stem; leaf-blades are long-elliptic, smooth, often
with crisped edges, the lateral veins at a very small angle to the
midrib so that they appear to run longitudinally, joined across by
close rather regular fine cross-veins which are conspicuous in
dried leaves but not in living ones.

242 Gardens’ Bulletin, Singapore — X XV _ (1970)

Inflorescences are terminal on short leafy branches of the stem:
they persist for a long time, sometimes after the leaves below them
have died. There is no main inflorescence-axis bearing primary
bracts, as in all other Scitamineae; the whole inflorescence is a
series of monochasial cymes. The basic unit of branching has at
its base a 2-keeled prophyll and then three bracts and a terminal
flower-bud; the distal bract enfolds the flower-bud and has no bud
in its axil; the next lower bract encloses a bud which will produce
the next branch of the monochasial cyme; the basal bract encloses
a bud which does not immediately develop but may later form
the beginning of a new branch-system of the inflorescence. An
old inflorescence shows a succession of scars where the flowers
have fallen, with two bract-scars between each (the scar of the
basal 2-keeled prophyll is not very distinct). The flower-scars are
in two longitudinal rows not far apart; thus the cyme is near to
being a cincinnus. The bracts are all tubular at the base, and
their exact orientation is not easy to establish.

Flower-structure. Flowers last one or two days after opening. The
inferior ovary has an apical prolongation which bears the floral
parts. This prolongation was called a calyx-tube by Ridley but is
solid; in its centre is a small strand of delicate tissue connecting
the base of the style with the axis of the ovary. The three sepals
are narrow, with almost parallel sides and short tips; they are
slightly fleshy and become limp on the second day. The pezvals
are arranged as in an orchid, two small lateral ones and a much
larger labellum at the base of the flower. The lateral petals are
little longer than the stamens and form with the base of the
labellum (which overlaps them but is not joined to them) a short
tube around the stamens. The labellum is usually about as long
as the sepals, the basal apart (claw) being narrow and the apical
part (blade) being much wider than the sepals; it is at first firm
and slightly fleshy with a slimy surface, on the second day putres-
cent except for the base. The five stamens are of simple structure,
opening inwards, in a close semicircle around the style and facing
the labellum. (Larsen mentions a staminode replacing the sixth
stamen, but I did not note this, nor does it appear in Ridley’s or
Larsen’s drawings). The style is slender, about as long as the
stamens; the concave 3-lobed stigma faces the labellum, its lobes
variously toothed or fringed and joined at the base to form a
shallow cup with raised tumid edges, open towards the labellum:
the backs of the stigmatic lobes (which show in a front view of
the flower) are polished, not receptive.

The ovary, sheathed by the floral bract, is trilocular with axile
placentation and many small ovules. The fruit is a loculicidal
capsule which bears at its tip the shrivelled basal part of the pro-
longation of the ovary. Lane states that there is a joint between
the sterile tip of the ovary proper and the extension bearing the
floral parts, but I did not notice this. The seeds are hairy, much
as in the genus Globba (Zingiberaceae) and have a lacerate aril
consisting of a few long lobes (not mentioned nor figured by Ridley
though present in his specimen now at Kew).

Holttum — Orchidantha 243

3. Species, Key and Descriptions

Style with stigmas 23-44 cm long; stigmas
fimbriate, fimbriae 5-6 mm long
Labellum 10-12 cm long, its blade 5 cm wide;
sepals 11-14 cm long
1. O. fimbriata

Labellum 6 cm long, its blade 2.2 cm wide;
sepals 6-64 cm long
2. O. siamensis
Style with stigmas not over 1.6 cm long; stigmas
with short teeth or trilobed

Ovary with extension 8-12 cm long;
sepals 6-10 cm long
3. O. longiflora

Ovary with extension 2-54 cm long;
sepals 24-3 cm long
Stigma-lobes bilobed with a small tooth
in the sinus; ovary with extension
54 cm long
4. O. maxillarioides
Stigma-lobes finely toothed; ovary with
extension 2—24 cm long
Sepals greenish white; petals 10 x 4 mm;
labellum brown with yellow lines
5. O. laotica

Sepals pale yellow at base to dull
purple at apex; petals 84 x 14 mm;
labellum blackish violet
6. O. borneensis

1. Orchidantha fimbriata Holttum, sp. nov.

O. longiflora sensu Ridley, Fl. Mal. Penin. 4 (1924) 292; M.T.M.
in Gard. Chron. Ser. III, 20 (1896) 658, with fig; not Lowia
longiflora Scort.

A Lowia longiflora Scort. differt: stylo cum stigmatibus triplo
longioribus; stigmatibus longe fimbriatis; lebello duplo longiori et
latiori.

Leaf-blade light green, to 100 x 15 cm, base narrowly decurrent,
edges undulate; petiole with sheath to 30 cm long. Inflorescence
little branched; 2-keeled bracts 18 mm long; intermediate bracts
to 7 cm long, green; floral bract to 15 cm long, dull purple. Ovary
with extension 12—20 cm long, purple, distal part 4 mm diameter.
Sepals reflexed, 11-14 cm long, 2 cm wide when flattened, narrowed
very gradually to the mucronate apex, edges strongly recurved
when flower first expands, flattening later, dark dull purple
throughout or green with a dark purple base. Petals at first rolled
back, cream with dark purple tip and base, 27 mm long, 6 mm
wide near base, narrowly triangular, apex filiform. Labellum 10-
12 cm long, 5 cm wide when flattened, base dark purple-brown,

244 Gardens’ Bulletin, Singapore — X XV (1970)

narrow, with inflexed sides, widening abruptly beyond the stigma
to a creamy white irregularly grooved and folded blade, towards
apex usually 3-lobed, lateral lobes reflexed, middle lobe smaller
and straight, texture rather fleshy, surface smooth and somewhat
mucilaginous. Stamens 24 mm long in all, filament 3 mm, flushed
with purple, anther cream. Style 2 cm long, purplish, wiry, stigma
2.4 cm long, white at base beneath (receptive surface) divided
almost to the base into 3 narrow dark purple shining lobes arranged
near together almost in a horizontal plane like a trident, their
distal edges bearing many curved fimbriae 5-6 mm long. Fruit
84 cm long including neck; seeds as in O. maxillarioides, 1 cm
long, aril-lobes to 15 mm long.

TYPE: Haniff 3957 (Ridley’s no. 16313), Perak, G. Kerbau,
S. Batu (SING). Other specimens in SING. PERAK. Ridley s.n.
1898, Tambun. Curtis s.n. Aug. 1898, Ipoh. SELANGOR. Ridley
8169, Gua Batu. Goodenough s.n. 1896, Ulu Langat. PAHANG.
Burkill & Haniff 16445, Benus valley, Bentong. TRENGGANU.
Corner s.n, 20.11.1935, Sungei Nipa, Kemaman ‘“‘abundant on the
hillsides in the stream valleys”. CULT. Hort. Bot. Singap. Sappan
s.n. 4.11.1926 (fig).

In addition to the above, there is in Kew Herbarium a specimen
collected by Scortechini in Perak (the name not in his handwriting)
presumably distributed after his death from Calcutta. There are
also flowers from a plant cultivated in the Botanic Garden at
Calcutta.

The flowers have at first an unpleasant odour, described by
Corner (on the label of his Kemaman specimen) as “smelling of
bugs and coconut oil’. On the second day the lip is putrescent,
the sepals become flat, and the odour is more like that of a
fungus. Such odours usually attract flies, but I did not observe
any flies visiting the flowers on cultivated plants at Singapore, nor
any fruits produced by those plants.

In addition to the Kemaman specimen recorded above, Corner
also collected another Orchidantha in Ulu Ayam swamp (S.F.N.
30353). He reports this as smaller than O. fimbriata as found at
S. Nipa, the flowers apparently similar but without scent (unfor-
tunately no complete flower was preserved). He records the Malay
name Lobor and that the leaves are used for wrapping bras pulut
in cooking. He also noted a smell of bananas when the plant was
crushed.

After the manuscript was submitted for publication, Dr. Hsuan
Keng of the Botany Department (University of Singapore) informed
me that Professor A. N. Rao collected a clump of this plant on
Gunong Panti in South Johore in 1969 and grew it in the Depart-
ment. Late September 1970, the plant produced flowers which
were photographed (see Plate 1).

2. Orchidantha siamensis Larsen, Bot. Tidsskr. 56 (1961) 347, 348,
igt2}¢3:

Leaf-blade 35 x 6~-7 cm, cuspidate; petiole with sheath 28 cm.
Ovary with extension 10-14 cm long; sepals 6-63 cm x 8-10 mm,
dull purple; petals 15-20 x 2 mm; labellum, narrow basal part 3
cm long, purple, blade 3 x 2.2 cm, white, emarginate; stamens

Holttum — Orchidantha 245

10 mm long; style 10-15 mm long, stigma 15 mm, lobes laciniate,
laciniae to 6 mm long; fruit 24-3 cm long, seeds 7 mm long, arill-
lobes 10 mm.

TYPE: Kerr 7148, Bachaw, Pattani, S. Thailand (K). Also
collected at some locality by Kiah, S.F.N. 24292 (SING).

3. Orchidantha longiflora (Scort.) Ridl., Flora Mal. Penin. 4 (1924)
292, nomen tantum. H. Keng, Gard. Bull. Sing. 24 (1969) 347-
349. — Lowia longiflora Scort., Nuov. Giorn. Bot. Ital. 18 (1886)
308, t. 11. K. Schum. in Engler Pflanzenreich IV, 45 (1900) 40,
fig. 1OA. — Orchidantha calcarea Hend., Gard. Bull. Str. Settl.
T (1935) to.

Leaf-blade to 55 x 6 cm, somewhat recurved with raised un-
dulate edges, base long-decurrent and not sharply distinct from
the broadly grooved petiole; petiole with sheath to 20 cm long.
Ovary with extension 8-12 cm long, tinged with pink. Sepals
6-10 cm long, to 15 mm wide, oblong, shortly narrowed to apiculate
tip, edges more or less revolute, dark brown-purple towards apex,
olive green with brown veins towards base. Petals 2.8 cm long,
3 mm wide at the base, with long slender tip, nearly white.
Labellum 5-6 cm long, 2.2-2.5 cm wide, base dark purple with
overlapping edges quite enclosing sepals and petals, blade ovate,
creamy white, strongly rugose with raised veins, tip (always?)
with a triangular sinus 14 mm deep, the raised midrib ending in
a point in the sinus, a little longer than the lobes. Stamens in all
12 mm long, filaments short, widened and purplish brown at the
base. Style 1 cm long; stigma-lobes purple, with truncate apices
finely and irregularly toothed, midlobe 6 mm long, laterals shorter.

TYPE. The type of this species has not been found, but the
agreement of Scortechini’s descriptions and figure with Henderson’s
specimen, especially in dimensions and shape of stigmas and
labellum, leave no doubt that Henderson had re-discovered Scor-
techini’s species (the discrepancies are that Scortechini gave 10
cm as length of sepals, Henderson 6 cm, and Scortechini 12 cm
for length of “‘calyx-tube’’, Henderson 8-10 cm). In the Singapore
Herbarium is a specimen collected by Wray, no 3366, from Upper
Perak, labelled Lowia longiflora Scort. in an unknown hand, which
is clearly this species; Henderson evidently did not notice it. The
type of O. calcarea Hend. is Henderson 26023, Lenggong, Upper
Perak. No other specimens are known.

4. Orchidantha maxillarioides (Ridl.) K. Schum. in Engler, Pflan-
zenreich, Musaceae (1900) 42. Rid]. Flora Mal. Penin. 4 (1924)
293. — Protamomum maxillarioides Ridl., Trans. Linn. Soc. II
Bot. 3 (1893) 383, t. 66. — Lowia maxillarioides (Ridl.) Baker,
Bot. Mag. (1894) t. 7351.

Leaf-blade to 33 x 8.5 cm, concave with broadly waved edge,
base rather broadly cuneate and sharply distinct from the petiole;
petiole narrowly grooved, 20-40 cm long, slender; sheath 10-20
cm long. Inflorescence often with several branches close together,

246 Gardens’ Bulletin, Singapore — XXV (1970)

branches sometimes branched again; floral bract 3.5 cm long, dark
purple. Ovary with extension 20 mm longer than bract, purplish.
Dorsal sepal hardly 3 cm long, 44 mm wide, edges a little reflexed;
lateral sepals 3.2 cm x 54 mm, slightly concave, edges not reflexed,
all sepals translucent purplish, greenish towards apex, oblong, the
apical quarter narrowed to acute tip. Petals 7 mm long, less than
2 mm wide, oblong with short abrupt tip, straight, dull purplish
with pale veins. Labellum 21 x 10 mm, elliptic, apex rounded,
basal 3-4 mm with sides upcurved, meeting but not overlapping
the petals, cream with faint dull purple mottling especially towards
the apex, base entirely flushed with dark purple, surface with about
3 longitudinal folds on either side of a broad plane median band.
Stamens hardly 5 mm long, filaments shorter than anther, whitish.
Style white, 4 mm long; stigma dull dark violet, base hollow, each
of the three lobes 2 mm long and wide, bilobed, with a small
tooth in the sinus. Fruit purple, 3 cm long, 1.7 cm wide, 3-angled,
narrowed at apex to a stiff appendage 15 mm long; seeds 7 mm
long, 5 mm wide, short-hairy, spherical above a narrow base; aril
of c. 6 stiff narrow lobes longer than seed.

TYPE. Ridley 2399, Pulau Tawar, Pahang. Other specimens:
Henderson 21864, Tembeling, Pahang. Cult. Bot. Gard. Singapore,
Md Nur s.n. 4.10.1926.

The origin of the plants cultivated at Singapore is not recorded.
They do not usually bear fruits, but there is a ripe capsule on
Ridley’s original collection. The flowers have no perceptible odour.

5. Orchidantha laotica Larsen, Bot. Tidsskr. 56 (1961) 349, fig.

es

Leaf-blade 35-50 x 10-15 cm; petiole with sheath 45 cm.
Ovary with extension 2.5 cm long. Sepals 3 cm x 5 mm, greenish
white; petals 10 x 4 mm; labellum 2-2.5 cm long, brown with
yellow lines; stamens 5-6 mm long; style 5 mm, stigma 3 mm,
lobes shortly laciniate.

TYPE. Kerr 21284, Muang Baw near Wieng Chan, Laos (K).

6. Orchidantha borneensis N. E. Brown, Gard. Chron. N.S. 26

(1886) 519.

Leaf-blade 15-25 x 6-9 cm; petiole 10-25 cm. Ovary with
extension 1.8 cm. Sepals 2.5—2.8 cm long, base pale yellowish, apex
dull purple; petals 8 x 14 mm, truncate with a fine bristle at apex;
labellum 2 cm long, linear-lanceolate, acuminate, blackish violet
(as petals); stamens 5 mm long; style “as long as the stamens,
slender, terete, whitish, terminating in a blackish violet 3-parted
fimbriate crest with a V-shaped stigma below it facing the
labellum’’.

TYPE. Cult. Compagnie Continentale d’Horticulture, Gand, 23
June 1885, origin Borneo (K). A letter from Florence attached
to the type sheet at Kew confirms that this species was published
two days before Lowia longiflora Scort.

Plate 1. Orchidantha fimbriata Holttum.

Photographs of plant from Gunong Panti. Divisions in cm.
Left: Habit of flowers. Right: Close-up of one to show fimbriate stigmas.
(Photographs by Mr. D. Teow),

The Arborescent Naucleae of New Guinea and
Solomon Islands (Rubiaceae)

By C. E. RIDSDALE*

Department of Botany,
The University, Bristol, U.K.

Rubiaceae of the New Guinea region were the principal
interest of Th. Valeton in the early part of this century. The
Naucleae were considered in his treatise on the Rubiaceae (Bot.
Jahr. 60, 1925). More recent collections have revealed the
presence of additional species in the area which are referable
to the genus Neonauclea Merrill. The present paper considers
this group in detail in an attempt to integrate the earlier
information with a study of the more numerous later collections,
and reassesses the taxonomy of the genus Neonauclea and its
relationship to other arborescent Naucleae. The complex generic
synonymy is not considered in detail as this is the subject of
a detailed review being prepared by Bakhuizen van den Brink
(personal communication).

The Linnean genus Nauclea was based on _ heterogenous
material; that which remains has been located by Bakhuizen
van den Brink and found to be congeneric with Adina Salisb.
Merrill’s homonym excludes Nauclea L. and is a circumscription
based on Sarcocephalus Afz. ex Sabine. Nauclea L. is not in
current usage and is not conserved. Bakhuizen van den Brink
suggests conservation of Nauclea Merrill against Nauclea L. and
Sarcocephalus Afz. ex Sabine and conservation of Adina Salisb.
against heterotypic Nauclea L. Whilst this would avoid emenda-
tion of binomials to some extent and conserve the current usage
of the genera, it can hardly be said that reversion from Adina
Salisb. to Nauclea L. and from Nauclea Merrill to Sarcocephalus
Afz. ex Sabine is a less satisfactory solution. Sarcocephalus Afz.
ex Sabine was in use up to 1915 and persisted in some areas
until 1963. Adina Salisb., on the other hand, has some 170 years
of use but most of its thirty or so members were originally
ascribed to Nauclea L. Furthermore, the Malesian representatives
of Adina are not congeneric with Nauclea L. and Bakhuizen
van den Brink suggests that they should be accommodated in
a new genus, Metadina. Thus the alternative to the proposal,
to conserve Nauclea Merrill and Adina Salisb., would, in most
cases, require only reversion to earlier names. Later taxa would
require recombination.

* Formerly of Division of Botany, Lae, New Guinea.

248 Gardens’ Bulletin, Singapore — XXV (1970)

Valeton (Bot. Jahr. 60, 1925) severely criticised Merrill's
concept of Nauclea and Neonauclea but preferred the Nauclea
concept of Korthals. There is a clear case, however, for the
retention of Neonauclea and the erection of Metadina as
morphologically and geographically well defined genera.

The description is based on material from the region under
discussion and field notes are included, being compiled from
personal observation or from data supplied by reliable foresters.
Frequently the material preserved on herbarium sheets is not
sufficient to identify the collection as the plants are relatively
rarely collected in the vegetative condition where the characters
of the stipules may be observed. Combined keys are included
so that it should be possible to identify the material in any
condition.

Key to Genera of Naucleae

Climbers. Inflorescence modified into hooks; corolla imbricate;
calyx lobes short, inappendiculate; seeds winged Uncaria
(not considered further)

Trees.

Lobes of corolla valvate; calyx short, truncate; stigma mitriform;
fruit dehiscing from the apex and remaining attached at the
base only; leaves close veined with 10-12 pairs of veins. Plants
of riverine swamp forests Mitragyna

Lobes of the corolla imbricate; calyx distinctly lobed; stigma
and fruit otherwise;
Flowers axillary; stigma subglobose; on dehiscing fruit axis
crowned by the calyx remains; stipules small, less than 5 mm
Metadina
Flowers terminal; stigma various; fruit not as above; stipules
greater than 5 mm;
Fruit a fused syncarp or a concrescent aggregate of capsules;
Calyces fused; fruit a fused syncarp; stigma lanceolate;
stipules obtuse Sarcoce phalus

Calyces concrescent: fruit a loose aggregate of capsules;
stigma lanceolate; stipules long sepuhinate. Trees of wet,

poor, swamp forest Anthocephalus
Fruit at maturity composed of numerous free cocci, initially
a pseudosyncarp: calyces completely free Neonauclea

Mitragyna Korthals

Mitragyna speciosa Korthals Verh. Nat. Gesch. Bot. tome 35 (1840).
M. speciosa Korth. Obs. de Naucl. Ind. 19 (1939), nomen nudum.
Stephygyne speciosa Korth. Verh. Nat. Gesch. Bot. 160 (1840).

S. parvifolia K. Schum. Fl. Kaiser-Wilhelmsland 127 (1889).
Nauclea speciosa Miq. FI. Ind. Bat. ii: 140 (1856).

Ridsdale — Arborescent Naucleae 249

Tree attaining 30 m (190 ft) commonly 11-25 m (60-81 ft),
bole 12-15 m (40-50 ft), sometimes flanged. Other bark greyish
green, shallowly sculptured, and with pustular lenticels; under
bark green; inner bark straw turning brown on exposure. Wood
straw, sapwood ill defined. Young branches dark brown,
internodes 2.5-6 cm, nodes pubescent. Stipules elliptic 7-15 mm
long x 4-5 mm wide, sparsely pubescent. Leaves ovate (7-)
10-14 x 3-5 cm, apex obtuse, abruptly acuminate; base obtuse,
slightly decurrent in the petiole; glabrous above, pubescent on
the veins below or sub-glabrous with tufts of hairs in the axils
of the veins. Lateral veins (9—) 11-15. Petiole 2.4 cm. /nflorescence
solitary; peduncle 6-10 mm to bract scar. Capitulum including
styles c. 3cm diameter, subtended by a pair of reduced leaves
(c. 4.5 x 1.5-2 cm) immediately below a pair of ovate pubescent
bracts. Calyx cupuliform c. 2 mm, shallowly lobed. Bracteoles
4 mm with slightly expanded apex. Corolla tube c. 5 mm, lobes
filiform 2.5—3 mm, valvate in bud. Style c. 5 mm, mitrate portion
c. 2 mm. Fruiting capitulum c. 20 mm diameter, individual
cocci 7 mm dehiscing from the apex but remaining attached
at the base.

Widespread tree of swamp forest expecially on riverine flood
plains.

TERRITORY OF NEW GUINEA. Sepik: NGF 3758;
Hollrung 674; Ledermann 6880. PAPUA. NGF. 4586, 33409;
Brass 8689; Hoogland 4217; Saunders 98. IRIAN BARAT.
Valeton 195; bb 25670 (Ned. Ind. Forest. Service).

?-MOLUCCAS; PHILIPPINES, BORNEO.

Metadina Bakkuizen van den Brink

Ab Adina Salisb (typus A. globiflora Salisb.) differt ovulis in
loculis numerosissimis, alabastris terminalibus conicis, habitu
arboris.

Type Nauclea trichotoma Z. & M. in Moritzi, Syst. Verz. 61, 1864.

Syn. Adina trichotoma (Z. & M.) Benth. & Hook. f. ex Hook. f.
& Jacks., Ind. Kew. 1: 43, 1895.

The genus as erected above was proposed and delimited by
Bakhuizen van den Brink (Taxon 19: 472 (1970) ) to contain the
Malesian taxa previously referred to Adina Salisb. It is
characterized by the small acuminate stipules, presence of
bracteoles in the capitulum, and small obtuse-truncate calyx lobes.
The calyx and ovary easily detach from the basal portion which
is immersed in the tissue of the capitulum. The spathulate
bracteoles arise from this basal portion and are interspersed
between dense hairs. The genus occurs throughout the Phillipine
and Indonesian Islands but only just reaches the extreme western
part of New Guinea.

250 Gardens’ Bulletin, Singapore —X XV _ (1970)

Metadina multifolia (Hav.) Ridsdale comb. nov.
Adina multifolia Haviland Journ. Linn. Soc. 33: 45 (1897).

Tree attaining 25 m (81 ft), bole c. 13 m (42 ft). Outer bark
greyish purple, slightly flaky; inner bark creamish white. Sapwood
white, heartwood straw yellow-orange. Young twigs rusty brown
with scattered lenticels; internodes 1-2.5 cm. Stipules small
acuminate c. 4.5 x 2 mm, glabrous. Leaves elliptic (5—) 7-15 x
(2-) 3-6 cm, base cuneate, slightly decurrent into the petiole,
glabrous. Lateral veins 7-8 prominent; petiole 5-15 mm.
Inflorescence axillary, solitary; peduncle from the first leaf to
the bract scar 2.5-4.5 cm, bract scar to capitulum c. 1 cm;
bract small, coronate-dentate, usually persistent. Capitulum
including styles 12-20 mm diameter. Calyx 1.5 mm overall,
upper portion 0.5 mm, slightly papillose, lobes small, elliptic,
obtuse, margin ciliate; columnar base densely pubescent and
surrounded by bracteolar portion, apex of ovary slightly raised
at margin bearing a ring of hairs and 5 bracteolae; bracteolae
spathulate, ciliate hairy at margin. Corolla 4-5 mm papillose,
lobes 1 mm. Style exserted c. 4 mm. Fruiting capitulum c. 10 mm
diameter, remains of the calyx (bracteolae bearing portion)
persistent on the axis after dehiscence. Fig. 1 B-—E.

IRIAN BARAT. bd 32711 (Ned. Ind. Forest. Service).
Kostermans 109 (=bb 33341); BW. 956, 3705, 3887, 5767;
P. v. Royen 3511. MOLUCCAS. Ambon: Buwalda 6140; bb
25933 Morotoai. Kostermans 827, 1650; Tangkilisam 213
(=bb 33886).

Sarcocephalus Afz. ex Sabine

Mature leaves generally ovate to orbicular, greater than 12 cm;
stipules greater than 1 cm; flowering capitulum greater than 35 mm
S. coadunata

Mature leaves elliptic, less than 12 cm long; stipules less than 1
cm; flowering capitulum less than 35 mm
S tenuiflorus

Sarcocephalus coadunata (Roxb. ex Smith) Druce (as Sarcocephalis)
Rep .Bot. Exch. Club Brit. Isles, 4: 644, 1916 (1917).
Nauclea coadunata Roxb. ex Smith in Rees, Cycl. xxiv n. 6 (1813).

Medium tree attaining 25 m (81 ft.), usually smaller, branches
more or less horizontal, not buttressed, bole stout. Outer bark
dark brown, thick, deeply and flaky fissured; inner bark straw.
Wood yellow. Young twigs rugose; internodes 2-5 cm. Stipules
broadly ovate to orbicular, 3-4 cm long. Leaves broadly ovate
15-30 x 7.5-15 cm, sometimes larger, apex obtuse, base obtuse
or cordate sometimes decurrent in the petiole; glabrous to pubes-
cent on the undersurface. /nflorescence usually a solitary capitulum;
peduncle 2-5 cm. Bracts, large ovate, deciduous. Capitulum in-
cluding styles 4 cm diameter, without flowers 1 cm. Calyces fused

Ridsdale — Arborescent Naucleae 251

E

Fig. 1. Sarcocephalus tenuiflorus & Metadina multifolia: A, S. tenuiflorus:
Corolla and stigma; B-E, M. multifolia: B, Complete flower; C, Calyx;
D-E, Basal bracteoles.

252 Gardens’ Bulletin, Singapore — XXV (1970)

lobes deciduous, clavate c. 3 mm, vestigial lobes triangular, slightly
pubescent. Corolla cream to yellow, tube 7 mm, lobes 4-5 mm,
glabrous. Style exserted c. 7 mm, stigma spindle-shaped. Syncarp
c. 2.5-3 cm; seeds ovoid, not winged.

Widespread tree of secondary regrowth, grasslands and savannah.
Throughout S.E. Asia.

Sarcocephalus tenuiflorus Hav., Journ. Linn. Soc. 33: 32 (1897).
Nauclea tenuiflora (Hav.) Merrill, J. Wash. Acad. Sci. 5: 337
(1915).

Medium tree attaining 25 m (81 ft.). Outer bark reddish brown;
inner bark straw. Branchlets smooth, internodes 2-4 cm. Stipules
ovate to orbicular, 6-8 mm, slightly keeled. Leaves elliptic to
narrowly obovate 7-12 x 3-6 cm; apex acuminate, base cuneate;
glabrous. Lateral veins 6-8. Inflorescence solitary. Bracts ovate,
caducous. Capitulum including styles 25-30 mm diameter. Calyces
fused, lobes spathulate 1 mm. Corolla white, tube 16 mm, lobes
4-5 mm. Style 8-10 mm, stigma lanceolate 3 mm. Syncarp 10-20
mm diameter. Seeds ovoid, not winged. Fig. 1A.

Scattered in wet swamp forest frequently associated with Metro-
xylon swamps.

TERRITORY OF NEW GUINEA. NGF 3829, 9963,
14192, 19144, 22886, 35132; Hollrung 829 (Type). PAPUA.
Saunders 68; Schodde 2385. IRIAN BARAT. BW /31/1.

Further investigation will probably show that this species is only
part of a variable complex of N. subditus (Miq.) Merrill to which
the type was referred by K. Schumann.

| Anthocephalus A. Rich.
Anthocephalus chinensis (Lamk.) A. Rich. ex. Walp.

Tree attaining 30 m (100 ft.), bole straight, often buttressed.
Outer bark grey, pustular, finely furrowed; inner bark straw. Wood
white, soft, grading to orange heartwood. Branchlets glabrous;
internodes 1-4 cm. Stipules lanceolate-acuminate 1.5—2 cm, glab-
rous. Leaves elliptic (rarely elliptic-orbicular) 15-50 x 8-25 cm;
apex slightly acuminate, base obtuse to rotundate; glabrous above,
pubescent on the undersurface. Lateral veins (7—) 10-15, prominent
below. Petiole c. 2.5 cm. /nflorescence solitary, peduncle 2.54
cm. Bracts small c. 2 mm. Capitulum including styles 3-5 cm
diameter. Calyx 5-8 mm, linear-elliptic, sub-glabrous. Corolla tube
8 mm, lobes 2-3 mm. Style exserted c. 10 mm. Fruiting capitulum
20-45 mm diameter, individual cocci c. 6 mm.

Widespread in regrowth communities and wet lowland swamp
forest.

Ridsdale — Arborescent Naucleae 253

Neonauclea Merrill

This is the most diverse and widespread genus of the Naucleae;
in the New Guinea region it is represented by some thirteen species
which are to be found from the lowlands to mid-montane altitudes.
It is frequently difficult to separate the species both in the her-
barium and in the field using the available literature. The characters
which have been found to be useful are considered below.

VEGETATIVE CHARACTERS

It is possible to identify the species by means of vegetative
characters which are frequently ignored in keys constructed from
herbarium material. Thus two keys are included which should
serve the field botanist and also for the identification of herbarium
material. The most distinctive feature of the genus is the stipules
(fig. 2); characters of shape and pubescence are distinctive to the
level of species groups. Bakhuizen van den Brink has independ-
ently drawn attention to the stipular characters and has used them
in the delimitation of the genus. The New Guinea species do not
agree with this proposed delimitation as those with acuminate
stipules would be excluded from Neonauclea and would appear
nearer to Metadina, reference to the calyx immediately confirms
their position in Neonauclea where they are placed in the present
work. Experience has shown that in the majority of species these
characters are constant. Some variation has been noted during
the period of shoot expansion, especially in N. obversifolia and
N. hagenii. During this period some degree of elongation of the
stipules has been observed but this is readily recognisable and the
general shape of the stipule is still maintained. Pubescence is
usually characteristic to the species level but may vary to a small
degree with maturity and locality. The keel found in some species
occasionally shows some variation in the degree of development,
e.g. N. gordoniana generally is without a keel but sometimes has
a slight keel developed which somewhat alters the shape. This
situation does not arise in species which have a pronounced keel,
such species are generally acuminate. It is probable that this
variation is a reflexion of the structure and shape of the shoot apex.

Collectors often ignore the vegetative parts as flowering is
terminal on lateral shoots and hence the vegetative phase is
frequently not represented in herbarium collections.

FLORAL CHARACTERS

The lateral shoots in the reproductive phase have several charac-
ters which vary with the degree of maturation. In the early stages
the capitulum is completely surrounded by a pair of bracts, typi-
cally broadly ovate; these are early caducous. The first pair of
leaves below the capitulum is usually highly reduced and caducous
in some species (fig. 3).

Gardens’ Bulletin, Singapore —X XV (1970)

254

Fig. 2. Stipules of Neonauclea: A, N. acuminata; B, N. perspicuinervia;
C, N. hagenii; D, N. gordoniana; E, N. lanceolata; F, N. chalmersii;
G, N. obversifolia; H, N. brassii; 1, N. solomonensis; J, N. maluensis;

K, N. forsteri; L, N. versteeghii; M, N. clemensii.

Ridsdale — Arborescent Naucleae 255

Fig. 3. Neonauclea forsteri: showing relationships between foliar leaves,
reduced leaves below the capitulum, stipules and the bracts.

256 Gardens’ Bulletin, Singapore — X XV (1970)

The appearance of the calyx changes with the degree of maturity.
There are five lobes which surround and overtop the immature
corolla, these being attached to the basal part by a thin shaft; as
the corolla expands the calyx lobes gradually break across the
narrow base of the shaft. It is apparent that in some species there
is a change in the general morphology of the lobes at this time
due to shrinkage and contortion of the different areas (fig. 6). At
maturity the calyx consists of a cylindrical base with the remains
of the five shafts appearing as vestigial lobes. These appendiculate
lobes are characterisic of the genus. In the fruiting stage the calyx
becomes further contorted and caps the individual cocci. The size,
morphological form, and type of pubescence of the lobes are
characteristic to the species groups.

Within the genus the corolla is of limited variability which is
expressed in the shape of the tube and in the size of the lobes. In
the fruiting stage the size of the whole capitulum and of the indi-
vidual cocci is the only general character of taxonomic value.

ECOLOGY

The genus reflects a family characteristic of being commonly
found in transition zones and seral communities. The trees are
occasional as components of the canopy layer under primary
forest conditions but more frequently occur in the subcanopy.
Many species are found in advanced regrowth communities.
The genus appears to have undergone a degree of ecological
evolution from forest types, showing relic pachycaul construction,
to types found in transient communities which may be small
leptocaul shrubs. This change has resulted in some degree of
parallelism with species outside New Guinea.

HERBARIUM KEY TO SPECIES OF NEONAUCLEA

Leaves pubescent, at least on the veins below
Leaves sub-auriculate, sessile or slightly

petiolate N. clemensii
Leaves not sub-auriculate, distinctly
petiolate
Leaves cuneate at the base or sub-
orbicular, obtuse N. versteeghii
Leaves truncate, obtuse, or cordate at
the base N. forsteri

Leaves glabrous

Calyx lobes with persistent acicular pro-
cessess and leaves linear to narrowly
elliptic, c. 5x as long as wide N. chalmersii

Ridsdale — Arborescent Naucleae

Calyx and leaves otherwise
Corolla pubescent

Corolla 5-6 mm; capitulum 20-35
mm diameter; fruit cocci 4 mm
Corolla 10-12 mm; capitulum 55-70
mm diameter; fruit cocci 10 mm

Corolla glabrous
Corolla < 10 mm; capitulum

35 mm (without fls. <12 mm);

fr. capitulum < 20 mm

Stipules narrowly linear, obtuse,
generally more than 3 x as
wide
Stipules glabrous
Stipules finely pubescent

Stipules obovate-spathulate, less
than 3x as long as wide

Corolla > 10 mm; capitulum
40 mm (without fls.> 15 mm);
fr. capitulum > 20 mm

Corolla less than 13 mm

Stipules and petioles glabrous;
stipules spathulate-ovate,
keeled; corolla narrowly
campanulate, lobes c. 1 mm;
style exserted c. 8 mm
Calyx lobes c. 5.5 mm,

apiculate

Calyx lobes c. 3 mm, obtuse

Stipules and petioles finely
pubescent; stipules acute, not
keeled; corolla broadly cam-
panulate, lobes 2-2.5 mm;
style exserted c. 12-15 mm

Corolla greater than 13 mm
Stipules acute

Stipules ovate, elliptic or
obovate

Peduncles and nodes pubes-
cent; stipules broadly
ovate to obovate; pubes-
cent; corolla lobes c. 6 mm;
stamens exserted

Peduncles and nodes glab-
rous; stipules elliptic to
elliptic ovate, glabrous

257

N. lanceolata

N. hagenii
ssp. papuana

N. gordoniana
N. obversifolia

N. solomonensis

N. brassii
N. maluensis

N. hagenii

N. acuminata

N. forsteri

N. perspicuinervia

258

Gardens’ Bulletin, Singapore — XXV (1970)

FIELD KEY TO SPECIES OF NEONAUCLEA

Leaves linear to narrowly elliptic, c. 5x as
long as wide; calyx lobes with long acicular
appendages at apex

Leaves and calyx appendages otherwise
Stipules acute at apex

Stipules pubescent
Stipules linear oblong; capitulum 25—
35 mm; style exserted c. 5 mm;
leaves elliptic, generally apiculate,
usually less than 11 cm
Stipules trigonal lanceolate; capitulum
exceeding 50 mm; style exserted
c. 12 mm; leaves broadly elliptic to
ovate, not apiculate, usually exceed-
ing 11 cm
Stipules glabrous
Stipules strictly acute, spathulate,
keeled (lower 2/3 with + parallel
sides)
Stipules acuminate, not keeled
Stipules long acuminate, c. 15 mm;
mature leaves large, usually ex-
ceeding 15 cm
Stipules trigonal to trigonal lanceo-
late, usually less than 12 mm;
mature leaves less than 15 cm.
(Riverine trees)
Corolla glabrous
Corolla papillose

Stipules obtuse at the apex
Leaves pubescent
Leaves sub-auriculate,
slightly petiolate
Leaves not sub-auriculate, distinctly
petiolate
Leaves cuneate at the base, or sub-
orbicular obtuse

Leaves truncate, obtuse or cordate
at the base, never suborbicular

sessile or

Leaves more or less glabrous
Corolla densely pubescent; stipules
ovate; leaves usually with a pro-
nounced acumen

N. chalmersii

N. obversifolia

N. hagenii

N. brassii

N. acuminata

N. hagenii
N. hagenii
ssp. papuana

N. clemensii

N. versteeghii

N. forsteri

N. lanceolata

Ridsdale — Arborescent Naucleae 259

Corolla glabrous; stipules not ovate(some-
times ovate-spathulate); leaves without
pronounced acumen
Stipules narrowly linear obtuse; |
less than 4 mm wide N. gordoniana

Stipules ovate-spathulate, broadly
elliptic or obovate; greater than
4 mm wide
Mature leaves usually exceeding
18 cm; stipules exceeding
IS x 8 mm
Peduncles and nodes pubescent N. forsteri

Peduncles and nodes glabrous N. perspicuinervia

Mature leaves usually less than
18 cm; stipules smaller than
1S x18 mm
Stipules spathulate ovate, base
distinctly narrow N. solomonensis

Stipules not spathulate, base
not distinctly narrow
Leaves generally exceeding
10 cm N. brassii

Leaves smaller than 10 cm WN. maluensis

DESCRIPTION OF SPECIES

a
Neonauclea clemensif Merrill & Perry, J. Arnold Arb. 29: 164

(1948).

Large tree attaining 30 m (100 ft); bole to 15 m (50 ft),
frequently 9-12 m (30-40 ft), rarely buttressed. Crown large,
4-4 total height. Outer bark pale grey-brown, vertically
fissured, flaking vertically into pieces c. 2.5-10 cm (1 x 4 in);
under bark brown; inner bark straw to very pale brown, fibrous.
Sapwood straw to pale brown; heartwood deep salmon pink.
Young twigs somewhat rugose; internodes c. 5 cm, stem broadening
at the nodes which are rugose to hairy. Stipules obovate,
obtuse, 20-25 mm x 10-12 mm, keeled; densely hairy on the
keels, hairs ferruginous; other portions of the stipule finely
pubescent. Leaves obovate 18-26 cm x 12-18 cm; apex obtuse
or slightly sub-apiculate; glabrous above, densely hairy on the
under surface, especially on the veins. Main vein broad at the
base, c. 5 mm, canaliculate above; lateral veins 10-12, obliquely
ascending and anastomosing at the margin. Petiole short (to 6 mm)

260 Gardens’ Bulletin, Singapore — XXV (1970)

or absent. Inflorescence with 3 capitula; peduncle from the
bract scar to the first leaf scar c. 4 cm. Bracts early caducous,
ovate, 8 mm x 7 mm, outside finely pubescent, inside glabrous.
Capitulum including styles 30-45 mm diameter, without flowers
attached 10 mm. Calyx lobes deciduous, 5 mm, spathulate
rotund-ovoid, sub-apiculate when young, densely hairy; swollen
apex c. 1.5 mm at maturity; part of calyx persisting with ovary
2 mm, exceedingly hairy on the vestigial lobes and with a ring
of hairs on the cylindrical base. Corolla 11 mm, glabrous, dirty
white; lobes 3-4 mm, reflexed at maturity. Style 16-20 mm
long, exserted for c. 7 mm. Fruiting capitulum without fruit
attached 25 mm diameter, individual cocci c. 8 mm long. Figs. 4
and 2 M.

TERRITORY OF NEW GUINEA. Markham Valley:
NFG 1607, 2513, 3169, 3271, 3309, 7373, 7539, 9402, 9831;
Macintosh s.n. Bulolo. Hartley 10921. Kui: NGF 22500.
PAPUA. Iawarere: Brass 687.

Neonauclea versteeghii Merrill & Perry, J. Arnold Arb. 25: 188

(1944).

Tree attaining 23 m (75 ft); bole 12-19 m (40-60 ft), buttresses
unrecorded. Outer bark brown, sometimes with shallow vertical
fissures, peeling into small flakes; inner bark brownish, turning
yellow near the cambium. Sapwood yellow; heartwood pink to
salmon red. Young twigs with thin pale brown bark, slightly
furrowed; internodes c. 2 cm on mature stems, nodes glabrous.
Stipules obtuse, ovate to obovate 20-25 x 7-15 mm, pubescent
especially at base; faintly keeled. Leaves sub-orbicular to broadly
obovate 24-34 x 18-25 cm; apex obtuse; base cuneate (rarely
truncate and decurrent); chartaceous to subcoriaceous, glabrous
above, pubescent below on the costa. Lateral veins 8-10, distinct,
hairy, anastomosing at the margin. Petiole 15—-SO mm, faintly
pubescent when young, glabrous at maturity. Inflorescence with
one or three capitula, peduncle 35-50 mm to bract scar. Bracts
caduous (not seen). Capitulum including styles c. 50 mm diameter.
Calyx lobes deciduous 4-4.5 mm, golden sericeous, swollen apex
of the lobe obtrigonal, densely hairy on the edges of the faces;
part persisting with the ovary 2.5 mm with short hairs on the
vestigial lobes and a ring of long hairs on the cylindrical base.
Corolla 14 mm, glabrous, pale yellow; lobes 2-3 mm. Style
exserted for c. 9 mm. Fruiting capitulum without fruit attached
25 mm diameter, individual cocci 8 mm. Figs. 5 and 2 N.

IRIAN BARAT. Brass & Versteegh 13509 (TYPE):
BW 2972, 10769, 11780, 12196; van Royen & Sleumer 7538.

The type has large suborbicular leaves slightly decurrent into
the petiole; the other material has distinct cuneate leaf bases
which seem more typical of species.

Ridsdale — Arborescent Naucleae 261

Fig. 4. Neonauclea clemensii: A, Mature corolla; B, Calyx surrounding
| immature corolla; C, Calyx at maturity; D, Individual fruit coccus.

262 Gardens’ Bulletin, Singapore — XXV (1970)

Fig. 5. Neonauclea versteeghii: A, Mature corolla; B, Calyx surrounding
immature corolla; C, Calyx from mature corolla; D, Individual calyx
appendage at maturity; E, Individual fruit coccus.

Ridsdale — Arborescent Naucleae 263

Neonauclea forsteri (Seem. ex Hav). Merr. J. Wash. Acad. Sci.
5: 540 (1915).

N. cardiophylla Merr. & Perry, J. Arnold Arb. 25: 189 (1944).

Nauclea forsteri Seem. Fl. Vit. 121 (1866). Nomen nudum.

N. Forsterii Haviland J. Linn. Soc. 33: 56 (1897).

N. orientalis Forst. f. Prodr. n. 85; p. 15 (1786).

N. rotundifolia Guill. Ann. Nat. Sci. 7: (1837).

Tree attaining 12 m (40 ft): bole frequently crooked; buttresses
thick, spreading, c. 61 cm (2 ft) high. Outer bark light brown
to grey, fissured: inner bark yellow. Wood pale yellow. Young
twigs somewhat rugose, internodes 2-4 cm, stems broadening
at the nodes to c. 8 mm, nodes ferruginously hairy. Stipules
ovate, c. 25 x 15 mm, pubescent, slightly keeled at the base.
Leaves broadly obovate to ovate (14-) 18-22 x 10-15 cm,
apex obtusely acuminate, base truncate to obtuse, glabrous or
pubescent. Main vein broad c. 2.5-3.5 mm at base, lateral veins
5-9, oblique, prominent, faintly anastomosing at the margin.
Inflorescence a single capitulum. Peduncle c. 2 cm, ferruginously
pubescent. Bracts broadly ovate, densely ferruginously hairy at
the base and lower 4. First pair of leaves below the bracts
much reduced, c. 5 x 3 cm, caducous. Capitulum including
style 50-70 mm diameter. Calyx lobes caducous, in the bud
c. 7 mm long, oblong, densely hairy, hairs pale brown, apex very
dark brown black, in the young flowers calyx lobes 10 mm long,
rotund-ovoid, densely hairy on the angles of the faces, shaft thin:
part persisting with the ovary 4 mm, pubescent. Corolla 8-12 mm
to the base of the lobes; lobes 2-4 (—6) mm long, narrow,
irregular, white, scented. Stamens exserted from corolla, style
exserted 10-12 mm. Fruiting capitulum 25-35 mm _ diameter,
individual cocci c. 10 mm. Figs. 3 and 6.

FIJI, SOCIETY IS., BRITISH SOLOMON ISLANDS.
BSIP 1442, 2919, 3119, 4229, 4750, 5392, 5665, 5803, 5089,
7584, 8433, 8465, 8651, 8690, 8822, 8830, 9020, 9306.
BOUGAINVILLE. Craven (& Schodde) 115; Waterhouse
25 (TYPE N. cardiophylla).

This species is based on a specimen in the Forster herbarium.
Seeman’s name occurs only in observation and is invalid;
Haviland clearly considered Forster’s specimen as the basis of
the name and so I designate as lectotype Forster’s specimen
from Society Is. (K.). The species shows varying degree of
pubescence of the leaves from more or less glabrous, to hairs
on the veins, to densely pubescent. Haviland’s conception of
the species is very broad and the material from the Philippines
needs to be re-examined as to specific status; also the material
from the Pacific Islands with large foliaceous stipules may be
separable from the species as typified above.

264 Gardens’ Bulletin, Singapore — X XV (1970)

heer
, (i v4
A

t-

Fig. 6. Neonauclea forsteri: A, Mature corolla; B, Calyx surrounding
immature corolla; C, Calyx at maturity; D, Calyx from mature
corolla; E, Individual fruit coccus.

Ridsdale — Arborescent Naucleae 265

Neonauclea chalmersii (F. v. Muel.) Merrill, J. Wash. Acad. Sci.

a, S29 41915).

Nauclea chalmersii F. v. Mueller, Notes on Papuan Plants 8: 44

(1886).

Shrub or small horizontally spreading tree, attaining 8-11 m
(25-35 ft), usually less than 4 m (13 ft). Outer bark pale grey
brown, thin, 1-2 mm, smooth or flaky, longitudinally cracked;
under bark greenish to pale straw; inner bark straw to pinkish
brown, fibrous. Wood straw coloured. Young twigs smooth, old
twigs with bark longitudinally ridged; internodes on mature
stems 1 cm or less; stems expanded at the nodes, glabrous.
Stipules oblong to oblong-ovate up to 22 mm. finely pubescent.
Leaves linear to narrowly elliptic, 8-16 x 2-4 cm; young
leaves sparsely pubescent soon becoming glabrous; more or
less sessile (occasionally with a petiole to 6 mm). Lateral veins
7-9 (-11) arising at an acute angle to the main vein, anas-
tomosing at the margin, somewhat prominent above. Inflorescence
generally a solitary capitulum, peduncle to bract scar 15-40 mm.
Bracts caducous, ovate c. 22 x 15 mm. First leaf below the
capitulum reduced, c. 4 x 2 cm. Capitulum including styles 45-55
mm diameter, without flowers attached 10 mm. Calyx lobe deci-
duous, 2.5 mm, with a setaceous appendage 5-6 mm, shaft 2.5-3
mm: part persisting with ovary 1.7-2 mm with aristate vestigial
lobes. Corolla 11 mm, glabrous, pale purple; lobes 4 mm. Stamens
half exserted. Style exserted 9 mm. Fruiting capitulum c. 30-35
mm diameter, individual cocci c. 10 mm long. Figs. 7 and 2 F.

Leptocaul tree on river banks and flood bars.

TERRITORY OF NEW GUINEA. NGF 3127, 22681.

PAPUA. NGF 5830, 5858, 7147, 14265, 17000, 31727;
Brass 553, 1381, 5584, 28892; Carr 16533; Crutwell 597, 692;
Forbes 191; Hartley 10630; Hoogland 48684; van Royen 4255
Saunders 137; Schodde 2580, 2970; White 357.

Neonauclea lanceolata (BI.) Merrill, J. Wash. Acad. Sci. 5: 504
(1915).

N. schlechteri (Val.) Merrill & Perry, J. Arnold Arb. 25: 187
(1944).

Nauclea lanceolata Blume, Bijdr. 1010 (1827).

N. affinis Miq., Fl. Ind. Bat. ii: 137 (1856).

. gracilis Vidal, Phan. Cuming, Philip., 176 (1885).

. philippinensis Haviland, J. Linn. Soc. 33: 52 (1897).
. tenuis Haviland, l.c. 33: 55 (1897).

. schlechteri Valeton, Bot. Jahr. 60: 50 (1925)

ae a oe

266 Gardens’ Bulletin, Singapore — XXV (1970)

Fig. 7. Neonauclea chalmersii: A, Mature corolla; B, Calyx surrounding
immature corolla; C, Calyx from mature corolla; D, Individual fruit
coccus; E, Seed.

Ridsdale — Arborescent Naucleae 267

Tree attaining 25 m (81 ft), frequently 12-19 m (40-60 fb),
crown large, about one third of total height. Outer bark grey
brown, flaky with vertical cracking; under bark pink; inner bark
pink to pinkish white. Sapwood cream, turning pink to orange
at the heartwood. Young twigs smooth, but sometimes with ridges
on the bark; internodes on the mature parts of the stem c. 10
mm, stems broadening at the nodes. Stipules ovate 6 (-9) x 4-5
mm, faintly keeled, minutely pubescent. Leaves elliptic 12 (—-15)
x 6-8 cm, apex caudate-acuminate (10 mm); base cuneate,
attenuate into petiole; glabrous. Lateral veins 5-7, oblique,
anastomosing at the margin, not prominent above. Inflorescence
generally a single capitulum; peduncle to bract scar 12-15 mm;
first foliar leaves sometimes reduced to 4-4 normal leaf size.
Bracts caducous. Capitulum including styles 20-35 mm diameter,
without flowers attached 10 mm. Calyx lobes deciduous, 4 mm,
spathulate oblong, obtuse, hairy; part persisting with the ovary
1.5 mm, exceedingly hairy on the vestigial lobes, scattered hairs
on the cylindrical basal part. Corolla 5-6 mm broadly campanu-
late, cream, densely hairy, expecially at the base of the corolla
_ lobes. Style exserted 4 mm. Fruiting capitulum (8-) 10-15 mm
diameter, individual cocci c. 4 mm long. Figs. 8 and 2 E.

TERRITORY OF NEW GUINEA. Schlechter 16925 (TYPE
N. schlechteri); NGF 3029, 3829, 5952, 7307, 11200, 12796,
13611, 21196, 21696, 27457, 29926; Brass 31903, 32440;
Darbishire 404; Hartley 10007710420, 10512, 11404; McAdam
270. PAPUA. Forbes 535 (TYPE N. tenuis); Brass 23671; Carr
15445, 15461, 15565, 15785, 15885; Pullen 5439; Schodde
2184. IRIAN BARAT. Brass 12595, 13534; Drs. van Leeuwen
10652. PHILIPPINES, CELEBES, BORNEO.

| Neonanclea hagenii (Laut. & K. Schum.) Merrill, J. Wash. Acad.
Sci. 6: 540 (1915).

N. dahlii (Val.) Merrill & Perry, J. Arnold Arb. 25: 188 (1944).
Nauclea hagenii Laut. & K. Schum. FI. Deutsch. Siidsee, 577 (1901).
N. dahlii Valeton, Bot. Jahr. 60: 51 (1925).

Small tree attaining 15-19 m (50-60 ft), bole to 8 m (25 ft).
Outer bark blackish grey with small slightly raised lenticels; under
bark pale silver brown; inner bark yellow, fibrous. Wood cream.
Young twigs smooth, or with some longitudinal ridges; internodes
on the mature stem 20-25 mm. Stipules trigonal lanceolate, 10-15
. mm, densely fine pubescent (rarely glabrous). Leaves broadly
elliptic to ovate elliptic, 8-16 (-18) x 4-9 cm, apex acuminate,
base shortly angustate, petiole faintly pubescent. Lateral veins
6-10, slightly oblique, faintly anastomosing at the margin; main
vein broad at base (2-3 mm), prominent; lateral veins prominent

268 Gardens’ Bulletin, Singapore — XXV_ (1970)

Fig. 8. Neonauclea lanceolata: A, Mature corolla; B, Calyx at maturity:
C, Calyx from mature corolla; D, Individual fruit coccus; E, Seed.

Ridsdale — Arborescent Naucleae 269

Fig. 9. Neonauclea hagenii: A-D, N. hagenii subsp. hagenii: A, Mature
corolla; B, Individual calyx appendage at maturity; C, Calyx sur-
rounding immature corolla; D, Individual fruit coccus. E-F, N. hagenii
subsp. papuana: E, Mature corolla; F, Individual calyx appendage at

maturity.

270 Gardens’ Bulletin, Singapore — X XV (1970)

for # of length. Inflorescence generally a single capitulum, rarely
with three capitula. Peduncle to bract scar (2—) 3-5 cm. First leaf
sometimes reduced to 4 normal leaf size. Bracts caducous, ovate,
acuminate. Capitulum including styles 50-65 (—70) mm diameter,
without flowers attached 15-20 mm. Calyx lobes deciduous, 6-7
mm, densely sericeous, shaft 2—2.5 mm, lobes swollen, cylindrical
elongate, apex acuminate; part persisting with the ovary 2-2.5
mm, pubescent. Corolla 10-12 mm, broadly campanulate, pale
green; lobes 2-2.5 mm. Stamens exserted. Style exserted 12-15
mm, cream. Fruiting capitulum 25-30 mm diameter, individual
cocci c. 10 mm. Figs. 9 A-D and 2 C.

Small tree always associated with riverine vegetation or seral
forest on flood plains.

TERRITORY OF NEW GUINEA. Lauterbach 175 (TYPE
N. hagenti); NGF 944, 3136, 5602, 9552, 13893, 14344, 14717,
18772, 18831, 22500, 24654; Brass 32519; Darbishire 8163;
Hartley 10165, 10226, 10948; Hoogland 4977, 5065. BIS-
MARCK ARCHIPELAGO. NGF 24148; Waterhouse 955-B.
IRIAN BARAT.

Neonauclea hagenii (Laut. & K. Schum.) Merr. subsp. papuana
(Val.) Ridsdale stat. nov.

N. papuana (Val.) Merrill & Perry, J. Arnold Arb. 25: 187 (1944).
Nauclea papuana Valeton, Lorentz’s Nova Guinea 8: 449 (1911).

It does not seem possible to separate N. papuana (Val.) M. & P.
from N. hagenii (Laut. & K. Schum.) Merr. by characters of
specific nature. The plants here included are separated by the
papillose pubescence of the corolla and by the glabrous nature of
the stipules. The subspecies occupies a similar ecological situation
to N. hagenii being found in seral flood plain forest along rivers.
It seems to replace N. hagenii in the drier monsoon areas of New
Guinea and the Moluccas. Fig. 9 E-F.

MOLUCCAS; Morotai: Kostermans 866, 1350. Aru
Islands: Buwalda 2592; bb 25423 (Ned. Ind. Forest Service).
IRIAN BARAT. Versteegh 1032 (Type); BW 1115, 1194,
7467, PAPUA. Brass 6955.

Neonauclea moluccana (Mig.) Merr. (?Nauclea glabra Roxb.)
appears to be closely related. It was described from, and is
represented by, fruiting material; the size of the fruiting capitulum
(12-15 mm diameter) and of the individual cocci (3-4 mm) is
smaller than in the N. hagenii group. The shape of the leaves
and of the acuminate stipules are similar to subsp. papuana.

Ridsdale — Arborescent Naucleae 271

Fig. 10. Neonauclea gordoniana: A, Mature corolla: B, Calyx surrounding
immature corolla; C, Calyx from mature corolla; D, Individual fruit
coccus.

272 Gardens’ Bulletin, Singapore — XXV (1970)

Neonauclea gordoniana (F. M. Bailey) Rids., comb. nov.

Nauclea gordoniana F. M. Bailey. Bot. Bull. Dept. Aegric.,
Brisbane 10: 22 (1895).

Large tree attaining 30-37 m (100-120 ft), crown dense. Outer
bark pale grey brown, flaky: inner bark cream to pink, fibrous,
turning grey brown on exposure. Sapwood straw to creamy brown;
heartwood red-brown. Young twigs pale grey brown with numerous
lenticels; internodes on the mature stems 2-4 cm, stems broadening
at the nodes. Stipules linear obtuse, 10-18 x 2-3 (4) mm, rarely
keeled, glabrous. Leaves elliptic to broadly elliptic, 10-17 (—22)
x 4-9 (-11) cm, apex slightly acuminate, base obtuse; glabrous.
Lateral veins 6-9, prominent above, raised below; main vein broad
(2-3.5 mm), canaliculate; lateral veins obliquely ascending, anasto-
mosing at the margin. /nflorescence with 1-3 capitula; peduncle
10-15 mm. Bracts early caducous (not seen). Capitulum including
styles 20-35 mm diamenter, without flowers attached 10 mm. Calyx
lobes deciduous 2.5 mm. shaft 1.5 mm, swollen lobes 1 mm, apex
clavate to clavate-turbinate with hairs strongly developed on one
side at maturity; part persisting with the ovary 1.5-2 mm, pubes-
cent on vestigial lobes and with scattered hairs on cylindrical base.
Corolla 5-8 mm, lobes 2.5 mm. Style exserted c. 5 mm. Fruiting
capitulum 12-15 mm diameter, individual fruit cocci c. 4 mm.
Figs. 10 and 2 D.

AUSTRALIA. Cowley, Baron River (Type). Fletcher
13431; Brass 19066. TERRITORY OF NEW GUINEA.
NGF 8463. PAPUA. NGF &608, 10436, 33405; Brass 22061,
25477; Schodde 2501, 2592; Carr 15381.

Neonauclea obversifolia (Val.) Merrill & Perry, J. Arnold Arb.
25: 188 (1944).

Nauclea obversifolia Valeton, Bot. Jahr. 60: 52 (1926).

Large tree attaining 46 m (150 ft), bole to 21 m (70 ft), usually
buttressed to 1-2 m (3-6 ft). Outer bark grey brown, flaky, with
a few scattered pustular lenticels and longitudinal fissures, fibrous;
inner bark yellow brown to straw. Sapwood pale straw brown;
heartwood orange brown. Young twigs smooth; internodes on the
mature stems 1-2 cm, stems expanded at the nodes. Stipules linear
or rarely linear ovate (ensiform) 6-10 x 1.5 mm, finely pubescent
(usually greyish on dried specimens, sometimes ochre). Leaves
elliptic 7-11 (-16) x 3-6 cm, apex apiculate, base cuneate;
lateral veins 6-8, slightly oblique, faintly anastomosing at the
margin. Inflorescence with 1-3 capitula; peduncle to bract scar
15-40 mm, first foliar leaves often reduced to 4 normal leaf size.
Bracts caducous. Capitulum including styles 25-35 mm diameter,
without flowers attached 6-9 mm. Calyx lobes deciduous 2.5 mm,
turbinate to deltoid, sometimes clavate when young, hairy. Corolla
5-7 mm, glabrous; cream, lobes small c. 1 mm (in dry specimens
corolla abruptly funnel-shaped). Style exserted 4-6 mm. Fruiting
capitulum 12-15 mm diameter, individual cocci 3-4 mm. Figs.
11 and 2 G.

Ridsdale — Arborescent Naucleae 273

’ Fig. 11. Neonauclea obversifolia: A, Mature corolla; B, Calyx surround-
| ing immature corolla; C, Calyx from mature corolla.

274 Gardens’ Bulletin, Singapore — XXV (1970)

IRIAN BARAT. BW 28990; TERRITORY OF NEW
GUINEA. Schlechter 16785 (tTyPE); NGF 510, 1175, 1201,
3412, 12373, 24764, 27484; Hartley 10834= 11048, 12196
Hoogland 4979, Hoogland & Craven 10207. PAPUA? Forbes
368. NEW BRITAIN. NGF 3412, 32644.

Neonauclea solomonensis Ridsdale sp. nov.

Arbor magna ad 30 m, ramulis laevibus, internodiis ramulorum
maturis 10-15 (-20) mm. Stipulae spathulatae-obovatae 10-14
mm longae, 5-7 mm latae (ad basim 3 mm latae), carinatae,
glabrae. Folia obovata vel obovata-elliptica 10-20 cm longa, 4-7
cm lata, apice acuminata, basi cuneata vel obtusa, glabra; nervorum
Jateralium primatiorum paria 7-9 oblique ascendentium, ante
Marginem anastomosantium (in siccitate atro-brunneis per duos
longitudinis trientes), subtus prominentium. Pedunculi 20-45 mm
longi, pedunculi laterales pedunculos medios superantes; foliorum
primum par infra capitulum ad tertiam partem foliorum normalium .
amplitudinis. Capitula (stylis inclusis) 30-40 mm diametro. Calyx
44.5 mm, lobis deciduis 2.5 mm, clavatis-ellipsoideis, apice-
brunneo, dense pubescentibus; parte persistente cum ovario 1.5
mm. Corolla 8 mm longa. Stylus 4 mm corollam superans. Capitula
fructifera c. 20 mm diametro, coccis individuis 6 mm longis.

TYPE BSIP 8739 (K)

Large tree attaining 30 m (100 ft); bole straight, buttresses thick,
plank-like, spreading, to height of 2.5 m (8 ft.); crown deep and
large. Outer bark grey brown, finely scaly fissured; under bark
dull yellow; inner bark sulphurous yellow turning khaki on expo-
sure, fibrous. Wood yellow brown, hard. Young twigs smooth,
slightly lenticellate; internodes on mature stem 10-15 (—20) mm.
Stipules spathulate-obovate, c. 10-14 mm X5-7 mm (3 mm at
base), keeled glabrous. Leaves obovate elliptic, 10-20 x 4-7 cm
(incl. petiole), apex acuminate, base cuneate to obtuse; glabrous.
Lateral veins 7—9, obliquely ascending, anastomosing at the margin
(drying dark brown for % their length), prominent beneath. /nflore-
scence with 1-3 capitula; pedunucle 20-45 mm, lateral peduncles
often long than the median. First pair of leaves persistent, some-
time reduced to c. 4+ normal size. Bracts early caducous, broadly
ovate. Capitulum including styles 3-4 cm diameter. Calyx lobes
deciduous c. 2.5 mm, swollen expanded portion clavate-ellipsoid,
apex dark, pubescent; part persisting with the ovary 1.5 mm.
Corolla c. 8 mm. Style exserted 4 mm. Fruiting capitulum c. 20
mm diameter, individual cocci c. 6 mm. Figs. 12 D-F and 2 I.

BRITISH SOLOMON ISLANDS. BSIP 96, 1505, 3740,
3888, 4100, 5864, 6347, 7526, 7623, 8739, 8985, 9161, 9373.
BOUGAINVILLE ISLAND, NGF 593, 815, 2857.

Ridsdale — Arborescent Naucleae PF ie

Fig. 12. Neonauclea acuminata & N. solomonensis: A-C, N. acuminata:
A, Mature corolla; B, Calyx surrounding immature corolla; C, Calyx
from mature corolla. D-F, N. solomonensis: D, Mature corolla; E,

Individual calyx appendage at maturity; F, Calyx surrounding imma-
ture corolla.

276 Gardens’ Bulletin, Singapore — XXV (1970)

Neonauclea brassii S. Moore, J. Bot. 65: 241 (1927).

Slender tree, bole straight, crown narrow. Outer bark pale
brown, finely scaly. Young twigs with lenticels; internodes 15—25
mm, stem broadening at the nodes. Stipules strictly acute, lower
2 with ‘+ parallel sides, sometimes elliptic — spathulate 8-10 x
3-4 mm, keeled, glabrous. Leaves broadly elliptic, 10-17 x 5-7 cm
(including petiole 10-14 mm) apex obtuse slightly apiculate, base
cuneate; glabrous. Lateral veins 8-10, obliguely ascending. Inflores-
cence generally a single capitulum; peduncle to bract scar 40 mm.
First leaf often reduced to % normal leaf size. Bracts caducous
(not seen), Capitulum including styles 40 mm diameter. Calyx
lobes deciduous, 5.5 mm, swollen lobes clavate, apex slightly
acicular, pubescent; part persisting with the ‘ovary 3.5 mm,
densely hirsute on the vestigial lobes and with scattered hairs
on the cylindrical base. Corolla 10 mm, narrowly campanulate;
lobes c. 2 mm. Style exserted c. 8 mm. Individual fruit cocci
c. 6 mm. Fig. 13 A-C.

TERRITORY OF NEW GUINEA. NGF 10579; Cult. Lae
Bot. Gdn. PAPUA. Brass 658 (TyPE); Carr 16486; Hoogland
3446.

Neonauclea maluensis (Val.) S. Moore, J. Bot. 65: 242 (1927).

Tree c. 25 m (81 ft). Stems smooth with scattered lenticels,
slightly broadening at the nodes; internodes 6-15 mm. Stipules
spathulate, apex acute to slightly obtuse (4-) 6-10 x 24 mm,
slightly keeled. Leaves obovate 7-8.5 x 3-4 cm, apex obtuse
abruptly acuminate, base cuneate slightly decurrent in the petiole;
Lateral veins 7-9, slightly oblique arching and anastomosing
at the margin. Inflorescence a solitary capitulum; peduncle to
bract scar 2—3.5 cm. Bracts caducous (not seen). First leaf some-
times reduced to c. 2-3.5 x 1.5-2 cm. Capitulum including styles
30-35 mm diameter. Calyx in the bud stage 4.5 mm overall,
lobes 2.5 mm (elongating in the flowering stage to 3.5 mm),
ellipsoidal, silvery pubescent; part persisting with the ovary
1.5—2 mm. Corolla 6-7 mm, cream; lobes short 1.5 mm. Style
exserted for 7-8 mm. Fruiting capitulum 20-25 mm diameter,
individual cocci 7.5 mm long. Figs. 13 D-F and 2 I.

IRIAN BARAT. BW 2147, 3688, Drs. van Leeuwen 11092.
Brass 11922. TERRITORY OF NEW GUINEA. Robbins
2503.

There is a great deal of variation in these specimens upon
which the description is based. The species was described from
Ledermann 7872 and 10570, collected in the Sepik basin and
the original description suggests that the leaves are larger. The
species probably occurs throughout the coastal and foothill forest
from Irian Barat to the Sepik and Ramu drainage areas as I
have seen sterile material growing in this area which could
probably be referred to this species. Further exploration and
collections will help to clarify this rather ambiguous species.

Ridsdale — Arborescent Naucleae 277

2mm

Fig. 13 Neonauclea brassii & N. maluensis: A-C, N. brassii: A, Mature
corolla; B, Individual calyx appendage at maturity; C, Calyx from
mature corolla. D-F, N. maluensis: D, Mature corolla; E, Individual
calyx appendage at maturity; F, Calyx from mature corolla.

278 Gardens’ Bulletin, Singapore — XXV (1970)

Fig. 14. Neonauclea acuminata

Ridsdale — Arborescent Naucleae 279

Neonauclea acuminata Ridsdale sp. nov.

Abor magna ad 30 m alta, ramulis rugosis vel fissuratis;
internodiis ramulorum maturorum 25-40 mm, nodis_glabris..,.
Stipulae lanceolatae-acuminatae, 16-25 mm longae, glabrae. Folia
late elliptica vel aliquantum obovata 8-18 cm longa, 6-12 cm
lata, apice abrupte acuminato (vel cuspidato), basi truncata,
glabra; nervorum lateralium primariorum paria 8-10, oblique:
ascendentium, ante marginem anastomosantium. Pedunculi robusti.
15-25 mm longi, ad apicem bracteati, bracteis caducis non visis;
foliorum primum par infra capitulum saepe minus amplum.
Capitula (stylis inclusis) 50-70 mm diametro. Calyx c. 9 mm,.
lobis deciduis 6-7 mm longis, spathulatis vel trigonis-spathulatis,.
dense puberulis; parte persistente cum ovario 4 mm; _lobis.
vestigialibus truncatis, pubescentibus. Corolla campanulata 13-16.
mm longa, sordida-albida vel purpuralbida; lobis 3 mm. Stylus.
8-12 mm corollam superans. Capitula fructifera 40 mm diametro,,
coccis individuis 12 mm longis.

TYPE NGF 24952 (K)

Large tree attaining 30 m (100 ft). Young branchlets rugose:
to fissured, internodes of mature branchlets 25-40 mm, nodes
glabrous. Stipules lanceolate-acuminate 16-25 mm, glabrous.
Leaves broadly elliptic to slightly obovate 8-18 x 6-12 cm,.
apex abruptly acuminate, base truncate. Peduncles robust 15—25
mm. Capitulum (including styles) 50-70 mm diameter, Calyx
c. 9 mm, lobes deciduous 6-7 mm, densely puberulous, part
persisting with ovary 4 mm. Corolla 13-16 mm, dirty white to:
purple-white. Style exerted 8-12 mm. Fruiting capitulum 40 mm
diameter.

This species is immediately distinguished by the large leaves,
which have a similar appearance to N. perspicuinervia Merrill
& Perry, the long acuminate glabrous stipules and by the large
size of the flowering capitulum and its various parts. The higher
montane, over 2,000 m (6,500 ft), trees have a tendency towards.
smaller more obovate leaves. Figs. 12 A-C, 14 and 2 A.

TERRITORY OF NEW GUINEA. NGF 3218, 3351],.
6212, 9104, 10208, 10470, 12790, 21139, 24952, 42130.
Hartley 12330, 12477.

Neonauclea perspicuinervia Merrill & Perry, J. Arnold Arb.
25: 188 (1944). |

Large tree c. 30 m (100 ft), bole c. 12 m (40 ft), crown broad..
Outer bark grey thinly flaky; under bark orange-yellow; inner
bark straw sapwood straw; heartwood yellow-brown. Young
' branches with deeply ridged pale scaly bark; internodes 1.5-2.5
cm. Stipules ovate-elliptic to broadly spathulate 1.5-2 x 1 cm,
slight obscure keel, glabrous. Leaves elliptic (9-) 15-25 (-36) x
8-15 (—25) cm (including petiole 2.5-3 cm), apex obtuse abruptly
acuminate, base obtuse to rotundate, slightly decurrent in the:

280 Gardens’ Bulletin, Singapore — X XV (1970)

4mm

Fig. 15. Neonauclea perspicuinervia: A, Mature corolla; B, Calyx from
mature corolla with single appendage attached; C, Individual fruit
coccus.

Ridsdale — Arborescent Naucleae 281

petiole. Lateral veins 9-11, prominent beneath arching and
anastomosing at the margin, leaf margin slightly crenulate.
Inflorescence a single capitulum, peduncle to bract scar 3-4 cm.
Bracts ovate c. 2 cm x 2 cm caducous. Capitulum including
styles c. 7 cm diameter. Calyx c. 15 mm overall, shaft c. 3.5 mm,
swollen apex c. 5 mm, densely white sericeous; part persistent
with the ovary 6 mm pubescent. Corolla 15 mm creamy white;
lobes 2—3.5 mm. Style exserted for 10 mm. Individual fruit cocci
c. 35 mm. Figs. 15 and 2 B.

IRIAN BARAT. Kanehira & Hatusima 12387 (TYPE);
Brass 13557; BW 569/; Drs. van Leeuwen 10769. TERRI-
TORY OF NEW GUINEA. NGF 24737.

Acknowledgments

I acknowledge with thanks the Directors of the following
institutes for placing their facilities at my disposal during my
visits: Herbarium Bogoriense, The British Museum and the
Royal Botanic Gardens Kew. I also thank Professor A. J. Willis,
Dr. D. Gledhill and Mr. D. A. Campbell, who read the manu-
script and offered valuable criticism, and Dr. Bakhuizen van
den Brink for making his paper on the nomenclature of the
Naucleae freely available to me before publication. I am indebted
to Mr. J. S. Womersley, Chief, Division of Botany, Lae for
the use of facilities during my employment with Department of
Forests, New Guinea and for freedom to pursue studies in the
herbarium and on field expeditions. Finally I thank my colleagues
Mr. M. J. E. Coode and Mr. E. Henty for their constant
encouragement.

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The Identity of Hyphaene natalensis Kuntze

By C. X. FURTADO*

Botanic Gardens, Singapore

SYNOPSIS

Hyphaene natalensis Kuntze (1847) was misidentified with
H. crinita Gaertn. (1791) by Martius (1849) apparently on the
supposition that the types of both these species had come from
South East Africa, where both the Dutch and the British had
settlements. This view of Martius was widely accepted in the
Floras of the regions, even though, contrary to Martius’ view,
Kirk (1863) had stated that H. crinita is an inland species. This.
was apparently due to Kirk’s misidentification of the taxon named
by Kuntze as H. natalensis and by Martius as H. crinita. According
to Kirk’s specimens in Kew, it is obvious that he had confused the
species partly with a form of H. coriacea and partly with H.
turbinata (mistaking the latter and H. schatan as forms of true
H. coriacea). Since the species (H. natalensis) from Kaffraria and
Natal was left unnamed by Kirk, Drude (1895) referred this
species to a form of H. coriacea (on vegetative characters given
by Kirk).

Since then more than one species of East Africa have been
referred to H. crinita. In 1924 Beccari showed that no fruit from
Fast Africa would agree with H. crinita as depicted by its author,
Gaertner. However, South African botanists themselves were not
certain where Hyphaene species are found in the region; and
relying on the information of a South African botanist who had
worked on the palms for the flora of the region, Beccari was
convinced that the type of H. natalensis was not from Natal and
so he typified the species on a specimen from Mozambique.

However, it is shown here that H. natalensis is found in
coastal regions of East Africa from Kaffraria to Mozambique and
the species has been retypified on Natal plant.

The fact that Kuntze (1847) had given a detailed description
of H. natalensis has generally been overlooked, probably because
this description is separated from its diagnosis and placed in the
discussion where Kuntze gives the reasons for keeping separate
H. thebaica and H. crinita (united by Martius, 1838). This original
description, given by Kuntze, is reprinted here and a new description
based on new material is given.

In the summer of 1967, while on vacation in Portugal, I under-
took to determine, among other things, the unnamed specimens
of Hyphaene in the Herbarium of the Overseas Botanical Institute
(Centro de Botanica) in Lisbon. Being then wholly unfamiliar
with the systematics of the genus, I did not foresee the great
difficulties with which I would be confronted in this enterprise.

* Retired botanist.

284 Gardens’ Bulletin, Singapore — XXV _ (1970)

Fig. 1. Translation of the label of Gaertner’s type:

Hyphaene crinita of Mr. Gaertner [Synonym] H. natalensis Kuntze
[in] Linn[aea]. (Amsterdam Garden probably brought from Kaffra-
ria and sent to Gaertner). H. thebaica is different. I have united to
this Kuntze’s species. (sg.) Martius.

Furtado — Hyphaene natalensis 285

The two binomials widely employed in the accounts of Hyphaene
species of East Africa are Hyphaene coriacea Gaertner (1788) —
the type species of the genus — and H. crinita Gaertner (1791) —
the second species instated under the genus. Yet the fruits of
the species identified as such do not agree with Gaertner’s drawings
of the types.

In his monograph Beccari (Borasseae: 29 and 30 (1924))
undertook to clarify this problem. He showed that there was no
species of Hyphaene in East Africa which would match with
Gaertner’s drawing of H. crinita and that the only specimen that
would agree in a great measure with Gaertner’s drawing of H.
coriacea was from the regions around the Red Sea. Beccari had
examined a large number of specimens from Central, North and
West Africa but he made no attempt to see if any of these could
fit with the type drawing of H. crinita. Further, Beccari’s identifi-
cation of H. coriacea represented a species quite different from
that shown by Gaertner’s drawing and he also ignored the geogra-
phical particulars which Brown (Tuckey, Exp. River Zaire,
Appendix, 5: 457 (1818)) had apparently himself obtained from
the collector of the type specimen. But, what is most important for
our purpose here is Beccari’s contention that the binomial H.
natalensis Kuntze has priority for the species from South East
Africa, the same that is commonly called H. crinita (op. cit.: 39
(1924) ). Either because of the defects indicated above or because
the mongraph was wholly in Italian, Beccari’s views have generally
been ignored by botanists so that even in recent literature these
two binomials have been employed in the erroneous sense given
- them by Martius (Hist. Palm, 3, ed 2: 226 (1849)) and by Kirk
(Journ. Linn. Soc. Bot. 9: 234 and 235 (1867) ).

Fortunately my studies in Lisbon enabled me to show that
H. crinita is identical with H. togoensis Becc. (1924) from North
Tropical West Africa and that H. coriacea is a species probably
common in North Mozambique and Kenya, with no resemblance
to any species from South East Africa and the regions along the
Red Sea. More recently, when my paper was in the advanced stage
of print, the Biological Institute of the University of Tuebingen,
Germany, loaned me the holotypes of Gaertner’s two species and
enabled me to confirm my identifications (Furtado in Rev. Garcia
da Orta 15: 459 t.19 (1970) ).

The identification of Gaertner’s species raised the problem of
identifying the species from East Africa erroneously referred to
Gaertner’s binomials. Already H. coriacea sensu Becc. (1924)
has named as H. beccariana Furtado (op. cit.: 44 Ic. 5 A and B
(1970) ); but the South African species misidentified as H. crinita
by Martius (1849) and Kirk (1867) and as H. coriacea var. minor
by Drude (Engl. Bot. Jahrb. 21: 126 (1895) ) and correctly referred
to as H. natalensis by Beccari (op. cit.: 39 (1924)) has not been
properly typified. Through Mr. Hein Wicht, F.R.G.S., of Durban,
fresh specimens of fruits and leaf-hastulas from South East Africa
were obtained. These enabled me to re-typify the species and clear
its nomenclature. The results of this inquiry are given here.

286 Gardens’ Bulletin, Singapore — XXV (1970)

YMMI”-;UYqqvq

Uy

HYP HAENE CRINITA GAERTN.
Kon done p €-TROP- AFRICA

Leite YH YY Yui

Plate 1 Hyphaene crinita sec Kirk in Kew

Apparently these specimens collected by Kirk at Kongoni, Nyasaland
(= Malawi) are the basis of Kirk’s statement that H. crinita sec
Mart. (= H. natalensis) is an inland species, and not a coastal one.
Three obovate fruits are forms of H. coriacea, and the third with a
campanulate outline is H. schatan. The date given on the specimen
is 1859, which might indicate the one when the specimen was studied
and incorporated in the Kew herbarium. (Photo published by
permission of the Director, Royal Botanic Gardens, Kew).

Furtado — Hyphaene natalensis 287

History of H. natalensis

In 1847, Kuntze (Linnaea 20: 15) described Hyphaene natalensis
based on fruits obtained from the Port of Natal by one Geinzius
in 1843. Since previously Martius (Hist. Palm, 3, ed 1: 227 (1839) )
had accepted only two species of the genus, H. thebaica with H.
crinita as a synonym, and H. coriacea, Kuntze gave reasons for
recognising H. crinita as quite distinct from H. thebaica stating
that the fruits of the former resembled more those of his new
species (H. natalensis) though in no way identical. (Fig. 1)

Kuntze’s remarks on the shape of the fruit of H. crinita appearing
more like that of H. natalensis, and the fact that the Dutch had
long established settlements in South East Africa have apparently
misled Martius (Hist. Palm. 3, ed. 2: 225 (1849) ) into making an
assumption that the type fruit of H. crinita had probably come
from Kaffraria.

It is on this geographical assumption alone that Martius reduced
to H. crinita not only H. natalensis but also the specimens sub-
mitted to him from Kaffraria by one Drege. Martius described
the fruits of the species being subobliquely obovate-pyriform,
convex at the vertex, a description that is applicable to the fruit
of H. crinita but not to that of H. natalensis which was described
as “‘obovate depressed at both the ends”’.

In 1867 Kirk (Journ. Linn. Soc. 9: 235) having studied the
Hyphaene species from the Zambesi region and its neighbourhood
changed completely the sense of H. crinita, perhaps basing his
conclusions on vegetative characters of the stem and the ligule. He
described the fruit as oblong and stem as “cylindrical’’, meaning
perhaps that the stem is solitary and neither ventricose (as in
H. ventricosa) nor branched (as in H. coriacea as_ interpreted by
him). Now, H. natalensis was said to be from the seacoast, but
Kirk stated that it is an inland species from the Zambesi and the
Nyassa regions and stressed that it is “‘not found on the seacoast’.
Kirk’s view was adopted by Wendland (Bot. Zeit. 39: 92 (1881) ).*

In 1895, Drude (Bot. Jahrb. 21: 176) modified somewhat the
sense by excluding a synonym given by Kirk but preserved Kirk’s
general interpretation. Drude at once saw that Kirk had left
nameless the Kaffraria species represented in the Berlin Herbarium.
On particulars given by Kirk of the ligule and the ecology of H.
coriacea, Drude identified Bachmann’s specimens from Kaffraria
in the Herbarium of Berlin as a dwarf form of H. coriacea from
Mozambique and called it doubtfully H. coriacea var. minor Kirk,
although Kirk had indicated his two varieties without naming
them. However, contrary to the views of Kirk, Drude applied
H. crinita also to the maritime specimens from Tanzania and
Madagascar.

Now, Kirk characterized H. coriacea as a species having a
dichotomous branched stem, and perhaps Drude was unaware that
the species from Kaffraria was normally non-branching and
cespitose. Nevertheless, it is surprising that neither Kirk nor

* Plate 1.

288 Gardens’ Bulletin, Singapore —X XV _ (1970).

Wendland nor even Drude followed Martius, and that all adopted’
H. crinita to the plant from Kaffraria whence the type of the.
species was supposed to have come, giving instead entirely a new
interpretation to the taxon.

Ignoring all these differences, Wright (Fl. Cap. 7: 30 (1897) &
Fl. Trop. Afr. 8: 121 (1901)) combined the views of Martius,
Kirk and Drude, and included under H. crinita both H. natalensis
and H. petersiana, thereby regarding the species as widely distri-.
buted both inland and along the seacoast from Kaffraria in the-
south to Tanzania and Madagascar in the north.

Marloth (Fl. S. Africa 4: 47 & 50 (1915) ) credited two species
to South East Africa, namely, H. crinita and H. ventricosa. The
former, according to him, remained dwarf in the southern coast-.
district of Natal, though it reached 30 ft. in height on the Limpopo
and the Zambesi. The other, Marloth stated, occurred in the north.
He gave a photograph on plate 12 of the same work with the
caption: ‘“‘Hyphaene crinita Gaertn. in the Northern Transvaal
near Zoutpansberg.”’

From the facts that Marloth could not loan to Beccari any
fruiting specimens from South East Africa and that he considered
highly doubtful the records of the occurrence of Hyphaene species.
anywhere in Natal and its southern district or even in its immediate
north (cf. Beccari: 40 (1924)), it seems obvious that Marloth
had not seen any specimens of Hyphaene either in a herbarium
or live excepting the one from the Northern Transvaal the habit
of which he illustrated. It is not easy to identify the species as
more than one species have similar habit. Whatever information
he had given in the Flora on the habit and distribution of the
plant was evidently reproduced from the previous books. For the
same reason it is not easy to determine what Marloth meant by
H. ventricosa, since this species was then known only from the
Zambesi region, and not from South East Africa.

In all these systematic treatments no attempt was made on
taxonomic grounds to discover a species that would agree either -
with the types of H. crinita and H. natalensis or with their original
descriptions. As a result, these binomials have been equivocally
used, although H. natalensis has been regarded often as a synonym
H. crinita. As said above, Beccari was the first to make a systematic
survey to identify these two species. In his monograph on Borasseae
(1924) he indicated that true H. crinita Gaertn. could not be found
anywhere in East Africa and that its description varied so much
from that of H. natalensis that these two species could not be
identical.

In order to determine the exact identity of H. natalensis,
Beccari approached Marloth who had elaborated palms for Flora
of South Africa (1915) for fruiting specimens of the two palms,
Hf. crinita and H. ventricosa mentioned by him in the Flora. In
reply to this, Marloth maintained that the occurrence of a
Hyphaene species in Natal was highly doubtful, admitting that it
occurred more to the North (Beccari op. cit.: 40 observ. (1924) )..

Furtado — Hyphaene natalensis 289

From this information Beccari concluded that the type fruits of
H. natalensis were probably brought to Natal from elsewhere so
that Geinzius could acquire them in Natal from where he could
post them to Loddiges in England. Hence Beccari typified the
species on a specimen collected by Schinz at Pisini near Delagoa
Bay. He could not however ignore the facts that Martius had
received specimens from one Drege from two localities in Kaffraria
and that Drude had Bachmann’s collections from Pondoland also
in Kaffraria. Without seeing any specimen from these collections
Beccari included them in H. natalensis. Even in studying the fruit
collected by Schinz, Beccari saw only Kuntze’s diagnosis. He
complained that this was too brief for the correct identification of
the species. He had overlooked an extensive description of the
fruit and its internal structures provided by Kuntze on the next
page following the diagnosis of H. crinita.

As said above, Beccari’s conclusion regarding the non-occurrence
of H. crinita in East Africa and the priority of H. natalensis has
been overlooked or ignored by botanists even today.

Phillips (Genera S. Afr. Flow. Pl.: 126 (1926) ) credited only
one species of Hyphaene for South East Africa, found in Natal
and the northern Transvaal. He did not name the species but it
is obvious that he meant the species to be H. crinita, a binomial
widely adopted then as it is now by South African botanists for
naming the species from the region. Evidently, like Marloth, he
was unaware of the Hyphaene specimens that grew in the Kaffraria
region.

Recent travellers in East Africa have had some of their specimens
from Mozambique and Tanzania named as H. crinita. In ‘“‘Prin-
cipes’’, journal of the Palm Society of America, this binomial is
frequently applied equivocally to specimens that in no way
represent H. natalensis. Brenan (Check List of Forest Trees &
Shrubs, Tanganyika: 399 (1949) ), who utilized Beccari’s mono-
graph on Borasseae, stated that the record of H. crinita in the
region is doubtful and suggested, for some unspecified reasons,
that the taxon from the region referred to H. crinita might be a
form of H. ventricosa. Brenan mentioned a specimen (Stuhlman 24)
which Drude had identified as H. crinita, a name used by Drude
to a species akin to H. schatan.

Provenance of the Type of H. natalensis

Since Beccari was misled to believe that no Hyphaene species
occurs in Natal, it is imperative that H. natalensis be typified on
the specimens from the region.

An inquiry into the specimens preserved in different herbaria
was not of much help in the identification of this species, and I
could not locate Kuntze’s types. A note published in ‘‘Principes,”’
a journal already mentioned above, put me in touch with Mr.
Hein Wicht of Durban, a keen observer of palms in South Africa,
and whose familiarity with the Hyphaene species of South East
Africa has enabled me not only to dissipate many false conceptions

290 Gardens’ Bulletin, Singapore — X XV _ (1970)

made by current botanical workers on the Hyphaene species of
the region, but also to typify H. natalensis.

Now, Durban is a new name for the former Port of Natal. Fruits
of Hyphaene species occurring not only along the coast but also
in the reserve of Kruger National Park were sent to me. It was
obvious that there were at least three distinct forms or species in
South East Africa, and Wicht agreed that they appeared distinct,
but botanists in South East Africa insist that only one species
exists in the region. The most important contribution for my
purpose was Wicht’s familiarity with the exact places where the
coastal species occurred in the north and the south of Durban,
i.e. from Kaffraria in the south to Mozambique a little beyond
the Tropic of Capricorn in the north. Unfortunately, the scope of
this inquiry had to be limited to South East Africa only.

Now, from the specimens received from Wicht it was evident
that Marloth had no proper basis for belittling the record of
the provenance of the type of H. natalensis, nor did Phillips (1926)
for ignoring the Hyphaene palms that were stated to exist in
Kaffraria by Martius (1849) and Drude (1895). The specimens can
be collected even now. Wicht assures me that he had seen a view
of “the Port of Natal in 1840” with a Hyphaene species in the
background. The type fruit of H. natalensis was collected only in
1843! Wicht recalled that a little south of Durban, at Amazintoti,
there used to be cespitose Hyphaene palms growing wild in a
property that belonged to him, though now they have disappeared,
owing to the impact of cultivating more economic plants.

In 1890, Wood, Curator of Botanic Gardens, Durban (Gard.
‘Chron. 8: 381) stated that in the Gardens there were male plants
which he suspected were sown by a former Curator Mcken; he
also noted that solitary plants of this palm are found on the sandy
plains in close proximity to the coast throughout the whole length
of the colony.

There is also evidence in the Kew Herbarium to show that
Marloth was wholly mistaken as to the occurrence of a Hyphaene
species in the Port of Natal, for, in 1859, only 15 years after
the collection of the type specimen of H. natalensis, one Sanderson
sent a leaf and male spadix of a Hyphaene species from Natal to
Kew where it was labelled H. crinita. It has to be recalled that
unlike Hyphaene nuts which German industrialists then sought
from any part of Africa in order to test the suitabilty of their
kernel as a substitute for vegetable ivory, leaves and male flowers
were not in such a demand. Therefore Sandersons’ specimens must
be accepted as having been collected from plants growing in or
near the Port of Natal then generally known as Natal. Kew has
also fruits collected in Natal by the Curator of the Durban
Gardens( Mcken) in 1869. These came apparently from the Port.
Wood (1890) reported that a collection of palms growing at the
neglected part of the Gardens was from seeds sown apparently by
Mcken. This would suggest that the orignal palms were becoming
a nuisance and had to be removed to a “neglected part of the
Gardens”, and it might not have been the first occasion for
eliminating the palms from important places in the city.

at

Furtado — Hyphaene natalensis

Hy Pn AEME CRIMITA
]. pata eae

Plate 2 Hyphaene natalensis Kuntze (Neotopotype in Kew).
Collected by Mcken in Natal in 1869. Top: Twin fruits. Bottom left:
entire fruit. Right: Longitudinal section. (Photo published by per-

mission of the Director, Royal Botanic Gardens, Kew).

292 Gardens’ Bulletin, Singapore — XXV (1970)

Now, by the phrase ‘“‘fructibus ex portu Natalensi misit
Geinzius’’ found in the protologue of H. natalensis, Kuntze
obviously wished to say that Geinzius had collected the fruits from
the plants growing in the port city of Natal and not that Geinzius
had acquired them there or posted them from that city. Geinzius
usually gave details of fruits, seeds, bulbs, etc. sent by him from
Africa to be grown in the gardens in England and Germany. Thus
many new species were described on material (or its progeny)
sent by Geinzius and some were even named after him. If he
did not know the provenance of the fruits, he would have informed
the recipient, and so Kuntze would probably have noted the fact
with such words as “‘patria ignota’’ (or incerta). Walpers (Annales
1: 751 (1848) ), who was in contact with Kuntze, also understood
that the palm grew (“‘crescit’’) at the Port of Natal.

The type could not have been derived from a palm introduced
and cultivated in Natal; for, in the very early stages of the
development of the Port, the authorities would have been con- ©
centrating on improving the aesthetic aspects of the surroundings.
This would mean ridding the place of as many native trees as
feasible and planting instead exotic trees chosen for their beauty
or shade; and, when there was a native Hyphaene, one would not
take the trouble to bring in a similar plant from another place of
the same country. Probably this work of eliminating native trees
led to the existence of male adult plants of the species in 1859
and 1890.

In view of this, it is evident that the type fruits of H. natalensis
must have been taken from a palm growing spontaneously in or
near the Port of Natal. As no species of Hyphaene is found growing
now in this locality, one has to explore its vicinity to see if any
is found to agree with the original description of the species.

The Protologue of H. natalensis

As said above, the protologue of H. natalensis is divided into
two parts. The first contains: (1) a brief diagnosis of the fruit;
(2) a brief description of a juvenile leaf since the nuts sown in
England in 1843 had not produced normal leaves till December
1846; and (3) the information that (a) the palm was listed by
Loddiges in his list of palms issued in 1845 and (5) the fruit was
from the Port of Natal received from Geinzius in 1843.

The second part of the protologue contains Kuntze’s reasons
for excluding H. crinita from the synonymy of H. thebaica, and
for recognizing these two species as distinct, in addition to H.
natalensis (newly described) and H. coriacea. It is in the latter
part that Kuntze gave a comprehensive description of the fruit
of H. natalensis. As this description has generally been overlooked
by botanists including Walpers (Annales 1: 751 (1848)) who

Furtado — Hyphaene natalensis 293

usually reproduced full descriptions of the species listed, and also
by Beccari (1924), both the diagnosis and the detailed description
are reprinted here as found in Linnaea 20: 15 (1847):

Hyphaene natalensis Kze

‘“Fructibus obovatis, utrinque depressis, laevibus, putamine sub-
globoso, breviter pedicellatis.

Loddiges palmae 1845 p. 7 No. 64.
Fructus ex portu Natalensi misit Geinzius a. 1843.”
(The descrpition of the juvenile leaf is not transcribed here.)

The detailed description of H. natalensis given under H. crinita
is as follows:

‘“Hujus fructum paucis describamus. Insidet pedicello 4-5 lin.
longo, 3-34 lin. crasso. Est 24 pollices longus, basi 14, ante apicem
rotundato-depressum 21 poll. diametro, lateribus levissime
depressus, subangulatus, epidermide laevi, castaneo-fusca, tenui
et fragili obductus. Sarcocarpium 14-2 lin. crassum, rubellum;
mesocarpium 2-3 lin. crassum, lignosum, ratione ad pedicellum
habita, obliquum (in H. thebaica sarcocarp. 3 lin. crass. pallidum;
mesocarpium | lin. crass.). Putamen 12 lin. longum, 15 latum, basi
attenuatum, obtusum, apice late truncatum s. depressum, laeve,
extus fibris rigidis fuscis mesocarpii cavitati partim, imprimis basi
et apice adnatum. (In H. thebaica putamen oviforme a basi latiore
obliqua apicem versus attenuatum, in apice ipso truncato-
umbonatum). Cavitas albumins (ut in H. thebaica) putamini con-
formis. Cavitas embryonis in specimine examinato non
reperiebatur.”’

Apparently Kuntze did not have a very normal fruit, for not
only did it lack the embryo but its putamen was much broader than
long, attenuate and obtuse at the base, and broadly truncate or
depressed a the apex. In the specimens (from the district of Natal)
recently examined, the putamen is nearly as long as it is broad,
somewhat truncately rounded at the base, and curvedly attenuate
from the upper third, forming almost an angle at the apex above
the embryo in the albumen. The absence of the embryo might
have been the cause of some disorganization in the structure of the

fruit such that the latter became somewhat flat or depressed at
the vertex.

Re-typification of H. natalensis

Now, there are four considerations which make the re-typification
of the species an easy task. In the first place the original description
is extensive giving the size and the shape of the fruit ;secondly, there
is Only one species of Hyphaene extensively distributed in the sandy
regions between the coast and the mountains of South East Asia
from Pondoland in Kaffraria in the South to the north in
Zululand; thirdly, there is no other species of Hyphaene found

294 Gardens’ Bulletin, Singapore —XXV (1970)

growing in the region having even a limited distribution; and
fourthly, the description agrees with the fruit of this Hyphaene
even though there are some minor variations not known to Kuntze
who probably based his description on one fruit or two of the
same plant. Therefore the coastal species of South East Africa
must be Kuntze’s species of which the type fruit had been
collected by Geinzius in 1843. A new description of the species
is therefore given here with its synonymy:

Hyphaene natalensis Kuntze in Linnaea 20: 15 (1847); Walpers,
Ann. 1: 751 (1848); Becc., Borass: 39 (1924) p.p. Figures 1-2,
Plate 2.

H. coriacea sec. Wendl. in Bot. Zeit. 39: 93 (1881).

H. coriacea var. minor Kirk ex Drude in Engl. Jahrb. 21: 126
(1899) 126 quoad specimina pondolandensia.

H. crinita sec. Mart., Hist. Palm. 3, ed 2: 227 (1849); Wendl.
in op. cit. 39: 92 (1881); Salom., Palm. Gatt. in Arten: 43 (1887);
Drude in op. cit. 21: 123 (1895); Wright, Fl. Cap. 7: 30 (1897)
et Fl. Trop. Afr. 5: 121 (1901); Marloth, Fl. S. Afr. 4: 50 (1915):
Omnino pro parte.

Caulis caespitosus, simplex vel rarissime bifurcatus, socialis,
1-4 m, interdum usque ad 6 m altus. Folia costapalmata, in lamina
petioloque nigropunctulata vel lepidota, deciduo pruinosa; ligula
in folio juvenili in petiolum oblique transversa, folii adulti
bilateralis, dimensione variabilis, in latere altiore ad usque 18 cm
longa, in altero tertio-brevior, margine plus minusve aculeata,
aculeis inaequalibus robustis praedita, apice sensim in costam
exeuns, interdum oblique transversa ut in folio juvenili. Pedicellus
12-15 mm longus, supra medium constrictulus, inde basin versus
incrassatus. Fructus 5.5-6.5 cm altus, apice circa 5.2 cm in diam.,
ad stigma depressum 3-3.5 cm crassus, oblique obovatus, basin
versus attenuatus, imo latere dorsali truncato rotundatus, ventrali
supra stigma curvato-ventricosus; apice oblique convexiusculus,
latere dorsali altior quam ventrali, vertice obsolete trigonus et in
perimetro stigmatico-pedicellari saepisse longitudinaliter promi-
nulus; infra apicem obscure angulatus; epidermide fusco brunneo,
nitido, iregulariter depresso-punctulato, tenui et fragili, obductus.
Sarcocarpium in parte ossea vertice oblique truncatum, basi rotun-
dato-truncatum. Putamen circa 3 x 3 cm, ovoideum, apicem versus
paulo attenuatum, basi late truncatum, extus fibris fuscis endocarpii
cavitati, imprimis basi et apice, adnatum. Albumen durum, album,
cum embryone apicali.

AFRICA AUSTRO-ORIENTALIS: in campis sabulosis siccis
inter litus et montes, ad usque 150 in altus. Provincia Natal in
Horto vel in Portu natalensis (Mcken — Neotopotype, K); in
subprov. Zululand, Mtunzini (Wicht 1); in vico Hibberdone
prope urbem Skottsberg (Wicht 3); Margate (Wicht 4).
Provincia Kaffraria, Portus Edward (Wicht 5).

A male spadix, collected in Natal in 1859 by Sanderson, is
preserved in Kew.

295

Furtado — Hyphaene natalensis

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A: Entire fruit (lateral view). B: Idem, dorsal view. C: Top view. D
Fruit, longitudinal section. E: Hastula. F: Hastula of an undeveloped

leaf.

296 Gardens’ Bulletin, Singapore —XXV (1970)

Plate 3 Hyphaene natalensis
Photographs by H. Wicht ;
A: Shows the cespitose character of the palm.

B: Cespitose character of the palm is often suppressed by
who utilize the leaves for making ropes and baskets.

C: Male plant cut off to extract Gemaan (toddy).

D: The palm occasionally branches in some places more often than in
others. Injury caused by fires is suspected to induce branching.

Africans

a ee CROCS NZ RY OCS ER PENNE ON FOO oa’ &

Acknowledgements

My thanks are due to Mr. Hein Wicht from Durban for
obtaining specimens from different parts of South East Africa; to
the Chief, Botanical Research Institute, Pretoria, Republic of
South Africa, for obtaining a photocopy of Kuntze’s original des-
cription of H. natalensis; to the Keeper of Herbarium, University
of Tuebingen, Germany, for loaning the holotype of H. crinita
Gaertner; to the Director, Botanic Gardens, Kew, for obtaining
for me photographs of some East African specimens preserved in
the Herbarium, and to the Director, Botanic Gardens, Singapore,
for putting the library and the herbarium facilities at my disposal
during the course of this study.

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Asian Species of Hyphaene

By C. X. Furtapo*

Botanic Gardens, Singapore

ABSTRACT

In this study two species of Hyphaene are described as new:
one from Israel (H. sinaitica) and the other from Ceylon (H.
taprobanica); both these are referred in previous literature as
H. thebaica. The correct name of the third species, H. indica, is
shown to be H. dichotoma (White) Furtado: this is common in
the north west coast of India. A fourth species (H. reptans) has
been recorded from the mountains of Arabia and has a flat,
trailing stem which branches but the fruits are unknown.

It was generally believed that the genus Hyphaene was exclu-
sively African so that any species found in Arabia and in other
parts of Asia were regarded as introduced from Africa and
acclimatized in Asia. Since all the adult species from the regions
outside Africa had branched stems, they were all referred to as
H. thebaica, a name that is still being widely but erroneously
used to denote also any African species that has a branched stem.
Thus about the year 1883 Haeckel (A Visit to Ceylon, ed. Engl.:
180 (1883)) noticed a species of Hyphaene growing in the Botanic
Gardens, Peradeniya, Ceylon and found that both its vegetative
characters and the flowers and fruits were so remarkably different
that it should not be called H. thebaica. The palm however pro-
duced a branched stem and information obtained at the Gardens
showed that attempts had been made to introduce H. thebaica
from Egypt, though apparently there was nothing to show that
the palm growing at Peradeniya was actually from the seed
obtained from Egypt. The extant records are not of any value to
determine this now. From an analogy, I might cite a case from
the Botanic Gardens, Singapore. Here, too, attempts were made
to acclimatize Hyphaene nuts from Egypt and India; but the
branched Hyphaene growing in the Gardens, though labelled
“H. thebaica — Egypt’, is actually H. dichotoma (H. indica).

To return to the Peradeniya plant, Haeckel wrote such a glowing
account on the ability of H. thebaica to alter its materially
important characteristics in response to changes in the environ-
ments or climate so as to discourage any systematic inquiry. No
doubt Beccari (Engl. Jahrb. 38: 8 (1906) & Agric. Colon. 2 (3):
15 et seg (1908) & Borasseae 23 et seq (1924)) had shown that,
besides H. thebaica, there are several other species which have
dichotomously branched stems, and that true H. thebaica is a
species restricted in its distribution from the mountains of the
Sudan where the Nile originates to the lower regions along the

* Retired botanist.

300 Gardens’ Bulletin, Singapore — X XV _ (1970)

Nile up to 27° Lat. N. and not to the seashore of the Mediterra-
nean or the Gulf of Suez or the Red Sea, where there are many
other branched species which cannot be said to be in any way
closely related to H. thebaica.* Similarly Beccari reported other
branched-stemmed species from the regions of North West Africa
and the Gulf of Guinea. Nevertheless, Haeckel’s view on the
fantastic ability of H. thebaica to change fundamental characters
in response to the changes in environments has become current
even in modern books (e.g. Taeckholm & Drar, Fl. Egypt 2: 279
(1950) & McCurrah, Palms of the World: 104 (1963)). To get a
precise idea of Haeckel’s views, the following particulars are taken
of “‘H. thebaica’” as given by Blatter (Palms Brit. Ind. & Ceyl.:
157 (1926) ), with a proviso that so far I have not found any true
H. thebaica grown in India or Ceylon.

CULTIVATION IN INDIA: — Old specimens of the Egyptian Doum
Palms may be seen in many a garden in India and Ceylon, and, as a
rule, they are much better developed than the tree growing in Egypt.
The climate seems, indeed, to exercise a great influence upon the
development of this palm. When Haeckel (1883) saw the Doum Palm
in Ceylon he was surprised to find it there under an aspect so altered
that he could scarcely recognize.

“Adaptation,” he says, ‘‘to perfectly different conditions of existence
have made the Doum Palm of Egypt quite another tree in Ceylon.
The trunk is developed to at least double the thickness, much larger
than in its native land; the forked branches are more numerous but
shorter and more closely grown; the enormous fan-leaves are much
larger, more abundant and more solid; and even the flowers and
fruit, so far as my memory served me, seemed to be finer and more
abundant. At any rate, the whole habit of the tree had so greatly
changed in the hot-house climate of Ceylon that the inherited physiog-
nomy of the tree had lost many of its most characteristic features.
And all this was the result of a change of external conditions and
consequent adaptation, more particularly of the greater supply of
moisture which had been brought to bear, from its earliest youth, on
a plant accustomed to the dry desert climate of North Africa. These
splendid trees had been raised from Egyptian seed, and in twenty
years had grown to a height of thirty feet.”

Since H. thebaica has a very limited distribution in nature even
in Egypt itself, the identity of the species reported from Arabia
and elsewhere outside Africa as referred to the Egyptian species
becomes a problem of first importance. Beccari (1908 & 1924) made
an attempt to identify them but was handicapped for lack of fruit-
ing specimens. However, he was able to determine one species
growing in the island of Diu on the north west coast of India;
he named it H. indica. The situation has not improved very much,
except that here two more new species from Asia are added. The
three species recorded here appear to be allied to a group of
African species or subspecies classified by Beccari under H. multi-
formis, but generally the epidermis in the fruits of the latter taxa
seem to be wrinkled and not discoloured into a network of lines
nor partly rimulose as in the Asian ones.

*Mahmood Ahmad Abu-Alula reports recently the occurrence of
individual trees (parts of Saudi Arabia) and pure stands (Sudan and The
United Arab Republic).

Furtado — Asian Species of Hyphaene 301

Enumeration of the Species

1. Hyphaene dichotoma (White) Furtado comb. nov.

Borassus dichotoma (dichotomus) White in Graham, Cat. PI.
Bombay: 226 (1839); Mart., Hist. Palm 3 ed 2: 318 (1850):
basinym. Figure 1.

B. flabellifer sec. in Hk. f., Fl. Brit. Ind. 6: 482 (1893) quoad
syn B. dichotoma.

H. coriacea sec. Dalziel & Gibs., Bomb. Fl. Suppl.: 95 p.p.
(1861).

H. indica Becc. in Agric. Colon. 2, fasc. 3: 173 t.3 fig. (1908)
et Borass.: 30 t.20 figs. 3-3’ & t.28 (1924): Blatter, Palms.
Brit. Ind. & Ceyl.: 187 tt. 33 & 34 & fig. 24: (1926):
syn. nov.

1. thebaica sec. Cooke, Fl. Bomb. 2: 811 p.p. (1907).

Beccari (1908) gave details of this species under H. indica
describing the fruit as obovate-pyriform and giving an illustration
of its longitudinal section; it could not therefore be allied to H.
thebaica though it was long regarded to be that species introduced
in Cutch in Gujarat. If the species is not really an indigenous
but an introduced palm in India, Beccari suggested searching for
its original parent in Arabia especially in Oman where, according
to reports noticed by Martius, similar palms were seen growing by
ancient travellers. Later Beccari (1924) showed that the Indian
species is allied to H. macrocarpa (Becc.) Furtado (=H. multiformis
subsp. H. macrocarpa Becc.) from infra-equatorial east Africa.

INDIA: Island of Diu and Ahmedabad in Gujerat.

The protologue of H. dichotoma given under Borassus dicho-
tomus is as follows:

“Oka-mundel, covers the whole of Diu Island and is also
found in various parts of Goozerat. Mr. Vaupel.

A solitary tree grows on a Hill at Mazagon [Bombay], and it
is branched like the Doum Palm. The fructifications have not
been examined. See a sketch of the tree, Madras Journal of
Science. N.”’

Now, Hooker (1893) and others regarded this species to have
been based on an abnormal branching of Borassus flabellifer;
he therefore reduced the former to a synonym of the latter. How-
ever, the phrase “it is branched like the Doum Palm” should be
taken to mean that the palm produces (perhaps repeatedly) dicho-
tomous branches as in the Doum and one should not expect ab-
normal branching (if such were really the case) to occur uniformly
in all the palms from the Diu Island and from the various parts
of Gujarat. Martius (1850) who accepted the species as new
remarked that the palm in Bombay was brought by Parsis from
‘Gujarat and could easily be distinguished from B. flabellifer.
Besides, monstrous branching in the latter species, he stressed, was
irregular and is rare even on the Coromandel of India, where the

302 Gardens’ Bulletin, Singapore —X XV (1970)

species is common, Martius added a new locality, Admedabad
(also in Gujarat) where Baron von Hugel (Car. = a misprint for
Bar.) had collected the specimens.

Gammie (fide Beccari) suggested that this species occurred as
far south as Goa and Blatter 1926 published a plate of a palm
growing at Bassein north of Bombay.

2. Hyphaene taprobanica Furtado spec. nov.

H. thebacia sec. Haeckel, A Visit to Ceylon, ed Engel.: 180
(1883); Becc., Borasseae: 24 t 13 (1924); Macmillan, Trop. Plant. &
Gardening: 161 t. (1935); Blatter, Palms Brit. Ind. & Ceyl.: 165
(1926); Taeckholm & Drar, Fl. Egypt 2: 279 (1950); McCurrah,
Palms of the World: 104 (1960): omnino pro parte zeylanica.
Figure 2, Plate 1.

Ab H. dichotoma cui affinissima, fructibus obovato-oblongis,
dorso paulo ventricosis cum sarcocarpio osseoso multo tenuiore
quam putamen paulo majore, ovoideo-oblongum vel ovatum, in
uno latere magis convexum apice rotundatum, basi fere truncatum
Sat distincta.

Caulis iterum iterumque dichotomus, cum ramis primis a basi
ejusdem modice remotis. Fructus obovate oblongus, dorso magis
convexus, vertice in perimetro stigmatico-pedunculari depressius-
culus, ventre obscure carinatus ad basin carpellis duobus steri-
libus plerumque conspicue tuberculatus, circa7 cm. altus, 5.5
cm. latus, 4-4.5 cm. crassus, epidermide nitidus, obscure tessel-
latus, partim irregulariter rimulosus. Sarcocarpium in parte fibrosa
4.5-7 mm. crassum, in parte osseosa 2-3 mm. crassum. Putamen
ovoideo-oblongum vel ovatum, basi rotundato-truncatum uno latere
magis convexum, vertice late rotundatum. Albumen conforme,
pariete 7-8 mm. crassum, embroyone carens.

ZEYLONA: probabiliter indigena, in locis supralittoreis. Cult.
in Horto Botanico, Peradeniya, feminea tantum (Tissera: June
1969, K, holotypus).

All the fruits, even the very young ones, have the same shape,
being much ventricose on the dorsal side and slightly carinate on
the ventral side which is taller at the apex and bears two knobs
of the vestigial carpels at the base. The epidermis is at first nitid
and partly dark brown, but later it is almost dark and opaque,
tessellately marbled all over and cracked in parts. Like H. sinaitica
and H. dichotoma, this appears to be a coastal species and so the
external characters regarding the colour of the fruit might be
abnormal and the effects of the cold mountainous climate of
Peradeniya.

Beccari’s t.11 attributed to H. thebaica shows a very variable
habit as in the members of the “H. multiformis’ group. Some
plants are branched at the ground level, others higher up. There is
one that has not branched at the base but instead has a very thick
stem and a few short and stout branches higher up—a phenomenon
also noticed in the Ceylon plants, though this is totally different

303

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1

Furtado — Asian Species of Hyphaene

phaene dichotoma (White) Furtado (H. indica Becc.)

(Ex Ic. Beccariana in Borass. t. 20 figs. 3 & 3%).

Hy

Fig. 1.

304 Gardens’ Bulletin, Singapore — XXV (1970)

from the Egyptian species as illustrated by Beccari. In the absence
of fruits I have assumed that all Ceylon plants are specifically
identical. The solitary plant from the Peradeniya Gardens has
not branched at the base but has bifurcated several times higher up.

When Dr. J. C. Willis, former director of the Botanic Gardens,
Peradeniya, provided in 1908 a male specimen from Jaffna to
Beccari in Italy with the information that the species was intro-
duced by the Peradeniya Gardens from Egypt but that it had then
become diffused on the island; he was obviously acting on the
long current belief of the staff of the Gardens that the species
is H. thebaica introduced into the Gardens from Egypt (cf. also
Haeckel op. cit. 1883), but the species is in no way related to the
Egyptian plant. I am inclined to believe that the species is a
native of the sea-coast of Ceylon, and is introduced into Pera-
deniya. The fact that Peradeniya in all these years has only one
female palm which, being dioecious, has not produced nuts with
an embryo hence are not viable, and that formerly the species
seems to have been widely distributed along the sea-coast of
Ceylon (e.g. Jaffna, Galle, Colombo, Batticaloa, etc.) favour my
view. As noted in the introduction there might have been unsuc-
cessful atempts to acclimatize the Egyptian species in Ceylon.

Due to the mistaken identity of the Ceylon species as H. thebaica,
Haeckel (1883) wrote, as said above, a glowing account to show
that this species readily evokes numerous changes in the habit,
shape and size of flowers and fruits. Though as early as 1908 &
1924 Beccari had shown that H. thebaica is a misnomer applied
to several species from Egypt and elsewhere and had described a
new Hyphaene from India, the fantastic adaptability of H. thebaica
has, as shown above, been quoted even in modern books.

Beccari ( Agric. Colon. 2, (3), 177 (1908) ), who had examined
the leaves of both H. dichotoma and H. taprobanica (the latter
from a male specimen from Jaffna) noted that the leaf of the
Ceylon plant had a greater number of segments which were also
much shorter, the lamina being less deeply divided and very much
lepidote on both the surfaces. Further, the male spadix was more
robust and longer than 1 m., but the spathes were glabrous in
the lower parts and fugaciously furfuraceous above (not densely
tomentose as in H. dichotoma), their tips broadly triangular and
longer than the respective axillary branch, the spikelets being
inserted at the mouth of the spathe, and each branch bearing
generally five spikelets (in H. dichotoma, the spikelet-bearing
branch is very much longer than its axillant spathe). Apparently
the same spadix was later mistaken by Beccari as having come
from the Botanic Gardens in Peradeniya and was described and
illustrated as of H. thebaica in Borasseae 24 t.13 (1924).

Since fruits were lacking in the specimen, Beccari could not
decide whether the Ceylon Hyphaene represented a new species
quite distinct from the region of Cutch or another i.e. one already
known.

Furtado — Asian Species of Hyphaene 305

——

Fig. 2. Hyphaene taprobanica Furtado (ex collectione fructuum holo-
typica)
A: Fructus latere visus. B: Idem ventre visus. C & D: Sectiones
verticales duorum fructuum.

306 Gardens’ Bulletin, Singapore — XXV (1970)

3. Hyphaene sinaitica Furtado spec. nov.

H. thebaica sec. Mart. Hist, Palm. 3: 22 (1849); Hart. Fauna &
Flora of Sina, Petra & Wady ‘Arabah’: 9, 28 & 111 (1891);
Zohary, Arb. Flor. Israel & Transj. in Imp. For. Inst., Oxford,
Paper 26: 12 (1951); Tomlinson, Anat. Mon. 2 Palmae: 158
(1961) omino pro parte. Figure 3.

H. coriacea ? sec. Busse in Karst. et Schenck, Veget. Bildr. 5
ser. fasc 7 t.44 (1907) quoad palmas austro arabicas.

Ab H. dichotoma, cui affinissima, fructibus dimidio minoribus,
ambitu irregulariter obovato-oblongis et in sectione transversa
tetragonoideis, vel apicem versus compressis fere bifacialibus am-
bitu obovatis, cum epidermide fragili partim discolore lineata,
paulo rimulosa, et sarcocarpio osseoso multo tenuiore differt.

Caulis dichotomus. Fructus circa 5 cm. altus, 4.5 cm. latus,
4-4.5 cm. crassus, nitidus, atro-bodius, obovato-oblongus, ventre
obscure carinatus et plus minusve porcatus, interdum apicem versus
sensim angustatus, sectione transversa tetragonoideus vel tetra-
gonoideo-bifacialis, cum epidermide partim fracta et retiforme
marmorata aut delapsa, etiam profunde rimulosa. Sarcocarpium
in parte fibroso 6-8 mm. crassum; in parte Osseosa circa 2 mm.
orassum. Putamen ovatum, basi oblique truncatum, circa 3 cm.
altum, 2.5 cm. in diam. cum pariete 1.5—2 mm. crasso. Albumen
conforme, cum pariete 5-7 mm. crasso; embryo apicalis.

PALESTINA: Wadi Tabba, ad versus meridiem a Eileth
(Goldring — sn. 15-5.69, holotypus, K).

This palm was first reported by Hart (1891) from Aqaba, but
Zohary (1951) found the species near the road between Aqaba
and Um Rashrash. Later he noted several trees of different ages
at 8 km north of Aqaba and remarked: “This is the northernmost
station of a tree so widely distributed in tropical regions. Here
it grows at approximately sea-level, on alluvial, somewhat saline
soil and ripens normal fruits.”

Here again there is a faint echo of Haeckel’s view regarding
mercurial changeability of the characters of H. thebaica in response
to climatic conditions, although the climate in Aqaba could not
have been very much different from that in some parts of Lower
Egypt where the species has a restricted distribution. The fruit of
H. thebaica is always conspicuously carinate, broadly truncate
at the base and much narrowed towards the apex. Its epidermis is
light brown or spadiceous and remains intact. In H. sinaitica, on
the other hand, the fruit is obovate and obscurely carinate, with
its epidermis dark brown and partly discoloured in lines so as to
give it a net-like appearance. In addition it cracks in parts rather
deeply in dry sarcocarp.

Martius, relying on the reports of several travellers, remarked
that H. thebaica occurred in several parts of Arabia in Sinai, along
the West coast Arabia (Yambu, Medina, etc.) and also in Oman.
Blatter recorded the species from Shaik Othman in Aden, and
Busse suspected the species from South Arabia to be H. coriacea.

Furtado — Asian Species of Hyphaene 307

scm

Fig. 3. Hyphaene sinaitica Furtado (ex collectione fructuum holotypica)

Fructus diversi, forma variabiles, omnes obovoidei, basin versus
tetragonoidei, apicem versus plus minusve plerumque compressi,
parte compressa nunc dorso, nunc ventre, nunc latere, vel rarius
dorso ventrique obscure sita.

308 Gardens’ Bulletin, Singapore — X XV (1970) :

Plate 1 Hyphaene taprobanica Furtado
Plants growing at Batticaloa, Ceylon.

(Photographed by Mr. I. Balachandran of St. Michael’s College,
Batticaloa).

Furtado — Asian Species of Hyphaene 309

Though from geographical considerations these records may be for
H. sinaitica, the possibility of the existence of more than one
species* should not be excluded; for unless the species is widely
distributed through the ocean currents, isolation in habitat tends
to produce local forms in Hyphaene species. There is a need there-
fore to study the specimens from different places on the coast of
Arabia.

4. Hyphaene reptans Becc. in Agric. Col. 2 (3): 151 (1908) &
Borasseae: 49 t.45 fig 6 (1924).

SOUTH ARABIA: Wadi Hadiea, Hadramaut, alt. 400 m.
(Lundt — K); Abrail, alt. 650 m. (Lundt — K).

The palm is said to have a trailing stem which is flat on two
sides and branches. From the ligule (hastula) the illustrated leaf
which is thought to be of an adult plant appears to be juvenile.
Fruit is not known.

However. H. dankaliensis var haycockensis Becc. (Borass: 29
(1924) ) is from the Haycock island in the Red Sea on the side of
South Arabia. Its stem is reported to be 4—5 cm. high. H. reptans
might be allied to this, but the nature of the stem of the latter is
not known.

Acknowledgements

It is my pleasant duty here to thank the Director, Botanic
Gardens, Kew, England, for the loan of the specimens from the
herbarium and for the photocopy of the original description of
Borassus dichotomus — the basinym of H. dichotoma (White)
Furtado; to Dr. C. C. Heyn, Botanist of the Jerusalem University,
Israel, for obtaining Hyphaene fruits from Wadi Tabba in the
Gulf of Aqaba, the type of H. sinaitica; to Mr. Maxwell Tissera,
a student at the University of Ceylon, for obtaining nuts of the
Peradeniya palm, the type of H. taprobanica, and also photographs
and information of the same and other palms recorded here; to
Mr. Mahmood Ahmad Abu-Alula, Director of The Technical
Office of The Agricultural Affairs in Jeddah (Saudi Arabia) for
fruit specimens, photographs and notes of H. thebaica: and to the
Director, Botanic Gardens, Singapore. for providing me library
and herbarium facilities to make these studies.

_ * A supply of H. thebaica fruits arrived lately. indicating the occurrence
in Saudi Arabia.

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A New Search for Hyphaene guineensis Thonn.

By C. X. FuRTADO*

Botanic Gardens, Singapore

Synopsis

Hyphaene guineensis Thonn. was described by Thonning him-
self who had visited the lower regions of Ghana towards the end
of the 18th century, but the species was published posthumously
in 1829, over twenty years after the destruction of the type in the
Copenhagen Herbarium during a bombardment of the city by a
gunboat. Since then no duplicates have been located and a great
deal of confusion had arisen over the precise identity of the
species. Sometimes it has been regarded only an infra-equatorial
species of West Africa, the fact being overlooked that the type
was from the supra-equatorial region. However, those who attri-
bute it to the latter region, have identified it with different montane
species, as if none is found in the lower regions. |

The account of a German expedition effected along the west
coast of Africa in 1873-76 shows that a single, unbranched
stemmed species, identified by them as H. guineensis, was quite
common on the coastal lands from the Cape Palmas eastward to
Ghana and Nigeria and southwards as far as Angola. Attempts
were therefore made to locate specimens in the herbaria of the
coastal states of supra-equatorial regions and to obtain nuts from
living trees. No specimens were available of any coastal species
and botanists did not seem to have seen any species in the coastal
region. But the Keeper of the Herbarium of the University of
Ghana and his associates visited the places on the coast near
Accra and on the banks of the Lower Volta (regions apparently
studied by Thonning) and obtained specimens which represent
some forms of H. guineensis.

This induced me to study the specimens in the Kew Herbarium
collected in Upper Guinea. They could not be located previously
when a search was made during my earlier attempt to identify
the species in Lisbon. The result of my study shows that this
species varies a great deal in Ghana, probably because of the
ocean winds (referred to also by the German explorers of 1873-
76). Some forms agree precisely with what I had described earlier
as H. doreyi from Angola but others are quite different. However,
a specimen from the Kew Herbarium, collected in Ghana in 1886
and labelled as H. thebaica, has been chosen as the neotopotype
of H. guineensis because it agrees most closely with the original
description of the species. The wide distribution of the species is
explained on the assumption that the species has originated in
Angola and subsequently spread to the north through the Benguela
Ocean currents.

* Retired hotanict

312 Gardens’ Bulletin, Singapore —XXV (1970)

2 t soaps
MOMS Hay\

Ly Lp 4
ye) me,
f/\ Mig? pf

Fig. 1. Hyphaene baikieana Furtado (Holotypus — K).

A: Fructus latere visus. B: Aspectus ejusdem vertice visus. C: Sectio
ejusdem verticalis.

s

Furtado — Hyphaene guineensis 313

The Kew specimens also revealed the misleading nature of
figures of H. macrosperma in Beccari’s Borasseae (1924). As a
result, the three forms which I had referred to H. crinita have
been separated as H. crinita Gaertn., H. baikieana Furtado sp.
nov. and H. togoensis Becc. Other species confused with H.
guineensis are H. macrosperma Wendl. (H. dahomeensis Becc.)
and H. tuleyana Furtado sp. nov. The last is apparently the same
taxon which Drude had identified as H. thebaica and which
Tomlinson had referred together with H. togoensis to H. guineensis.
Since H. thebaica has been widely confused, three forms of this
species have been illustrated.

Probable Origin of Confusion

One of the most misapplied binomials of the genus Hyphaene
is H. thebaica (.) Mart.; it seems to be a handy name to indicate
any branched species of Hyphaene that one is not able to identify
with certainty, be it a native of northern Egypt, the western coast
of India, or from anywhere in Arabia, Madagascar, Kenya,
Mozambique, Angola, Upper Guinea or found cultivated in any
of the Gardens of Asia or America. A recent case of such mis-
application is by Menninger (Fantastic Trees: 72 (1967) ) who has
given under that name photographs of a palm that obviously
belongs to the group of H. hbaronii Becc. if not identical with it.

When this practice is adopted in authoritative Floras without
attempts being made to distinguish any peculiar systematic forms
found in the region dealt with from those growing outside that
region, students of the genus get generally discouraged. This may
be one of the main reasons for the confusion that exists in the
nomenclature of the species of the genus.

In my own case, though I had pertinent Floras, Check- Lists,
many journals and popular books, I soon found myself confronted
with so many contradictions and equivocal use of binomials that
I had to give up my intention of making myself familiar with the
genus Hyphaene and of determining its few specimens that were
available in the herbaria in Lisbon. It was only when a copy of
Beccari’s revision of Hyphaenes (Borasseae, 1924) was obtained
that I was able to carry out my undertaking.

Thus H. guineensis, to quote an example that concerns this
paper, was credited to Upper Guinea in the region of the Niger
by Drude, (Engl. Jahrb. 21: 110 & 123 footnote 3, (1895)) who
based his conclusion on Pechuel-Loesche’s report on Die Loango
Expedition, 1882, (quoted here below) and on the specimen from
Dr. Baikie’s Niger Expedition which has been described here as
H. baikieana, Drude, who wished to adopt H. congensis Kirk
nom. nud. in Salomon’s list of palms (1887) for the taxon in
Lower Guinea, overlooked the fact that H. guineensis identified
by Pechuel-Loesche was an unbranched coastal palm. Rendle
(Cat. Welw. 2: 82 (1899) ) re-adopted H. guineensis for an Angolan
palm, while Wright (FI. Trop. Africa 8: 120 (1901)) recorded the

314 Gardens’ Bulletin, Singapore — XXV _ (1970)

species as though from Lower Guinea only. But Hutchinson
(Hutch. & Dalziel, Fl. W. Trop. Afr. 2: 132 (1936)) treated the
species as if of Upper Guinea only (i.e. Ghana, Togoland and
the neighbouring regions) and made no remarks about other
senses given to the same binomial, nor did he state whether there
was any reason to accept or reject the species as interpreted by
Rendle, Wright and others, or even by Beccari. Irvine, who had
worked for a number of years as a professor of biology in the
Gold Coast Colony (now the independent state of Ghana), credited
only one species to the country, the one that produced a branched
stem; he called it H. thebaica and remarked that it was not an
uncommon species in the drier regions of Ghana and Togoland
(Plants from Gold Coast: 235 (1930) ). Later he recorded the same
species also from Nigeria and Sierra Leone (West African Botany
1931: 176 & 188 (1941, reprint)). In 1961 (Woody Plants of
Ghana: 780-781 pl. 34a) Irvine followed the practice of Wright,
Hutchinson and others, and so extended the distribution of H.
thebaica also to Tanganyika, Somaliland and Arabia. Obviously
he did not agree with or consult Beccari (1924). The latter had
restricted the use of the binomial to a taxon native only of the
Sudan and Egypt, had adopted H. togoensis, H. dahomeensis and
H. macrosperma (with H. guineensis as yet doubtful) for the
species from the supra-equatorial region of tropical West Africa,
and other names for the species of other regions of North and
East Africa and India.

In an authoritative Flora like that of Hutchinson and Dalziel
(1936) one expects that the authors would accept or reject with
justification the previous interpretations by Irvine and others of
H. thebaica as found in Ghana and the neighbouring countries
and from which Beccari had excluded the three or four species
mentioned above. Hutchinson, however, followed an unusual
procedure. He did not credit H. thebaica to Ghana and the neigh-
bouring regions; instead of identifying the species discussed by
Beccari and Irvine, he exonerated himself on the plea that he had
not seen from the region any authentic specimens of these taxa.
So he merely listed H. guineensis, H. togoensis and H. dahomeensis
and, despite greater systematic differences recorded by Beccari
between the last two, he expressed his suspicion that the last two
were probably only synonyms of the first! For some unknown
reason Hutchinson ignored H. macrosperma Wendl. founded on
a specimen collected by Baikie apparently during his Niger
Expedition. He referred the reader to the Appendix of the Flora
which had probably been already prepared although issued only
in 1937 (and reprinted also in 1955). In this Appendix, Dalziel,
its compiler, disregarded Hutchinson’s opinion and quoted the
vernacular names of all Hyphaene species including that of H.
guineensis from the region under H. thebaica as if he regarded
merely as synonyms the different species recorded by Beccari
from Upper Guinea and North Africa. Dalziel’s statement that
Hyphaene is a monoecious palm is probably a misprint for
dioecious. (As to the word Sokuti auoted by Dalziel, it can

Fig. 2. Hyphaene crinita Gaertnet

Three views of the type fruit preserved in Gaertner’s Herbarium
at the University of Tuebingen, Germany.

316 Gardens’ Bulletin, Singapore — X XV _ (1970)

apparently be applied to any palm tree in the Ewe language of
Ghana, for according to information obtained by Dr. Hossain,
Soku means a “‘palm fruit’ and Sokuti a ‘‘palm tree’’).

Yet Beccari, Irvine and even Hutchinson and Dalziel were con-
temporaries who were dealing with the objects seen growing in
the regions. Excepting the types of H. guineensis which are lost,
specimens in herbaria could have been consulted. Beccari not
only had described the different species but also given illustrations
of the fruits (mostly types) for the guidance of systematists. Under
these circumstances failure of making a systematic survey of the
species recorded from Ghana and the neighbouring countries on
the plea that no authentic specimens were seen constitutes a
lacuna that is hardly excusable in a Flora that claims to be
authoritative. Besides, Kew Herbarium had some specimens from
Ghana and the neighbouring regions.

Tomlinson, interested in the anatomy of palms, and while
attached at one stage to the University of Ghana, noticed only
one Hyphaene species indigenous in Ghana and Nigeria, the same
that for fifty years or more previously was mistaken for H.
thebaica. As the types of H. guineensis were collected in Ghana
more than 150 years earlier and only one species had definitely
been recorded since that time from the territory, Tomlinson felt
justified in adopting the binomial H. guineensis for the only
Ghanaian species known to him, and, according to him, this taxon
was wholly different from the true H. thebaica from Egypt (Anat.
Mon. II. Palmae, 158 (1961))*. Besides. in the absence of the
types and any other specimen from the type region, Thonning’s
data of H. guineensis were, until recently, considered too inade-
quate to identify the species precisely. The procedure adopted by
Tomlinson seemed then to be the most logical. for it did not leave
the Ghanaian species unnamed, nor did it permit either of the
two binomials i.e. H. guineensis and H. thebaica to be used
equivocally for the same species of West Tropical Africa, nor H.
guineensis to be excluded from Ghana where the type of the
species was collected.

However, in the second edition of the Flora of W. Trop. Africa
3: 169 (1968), Russell stressed that lack of adequate herbarium
material was the reason for disregarding both Beccari’s and
Hutchinson’s treatment of the different species from West Tropical
Africa and for following Dalziel in adopting for all of them the
name H. thebaica. Russell seems to have some doubt as to H.
guineensis Thonn. being from Upper Guinea. He was probably
unaware that Thonning himself had collected the type of his
species in a former Danish settlement in Ghana, and not in Lower
Guinea as Wright (F/. Trop. Afr. 8: 121 (1901)) had erroneously
recorded.

Now, Beccari had received from several parts of the Gulf of
Guinea fruits which were not identical with H. togoensis. Further,

*In comparing the anatomy of the leaves Tomlinson mistook H
sinaitica from Israel for H. thebaica—q.v.

Furtado — Hyphaene guineensis 317

H. togoensis was a palm that normally branched dichotomously
while H. guineensis is taken to be a palm that normally does not
branch, a reason why Beccari placed it in his Section Ventricosae.

Beccari (1924) who had vast experience with the genus Hyphaene
was of the opinion that its species produced very localized forms.
This led him to believe that the taxon identified as H. guineensis
from Lower Guinea by Rendle, Wright and others could not be
the species so named from Upper Guinea. This was also the
opinion of Drude (1895) who wished to distinguish the species
from Lower Guinea as H. congensis. As the Angolan species did
not produce a ventricose stem and had moreover thickly lepidote
leaves, it could not belong to the Ventricosae of Beccari. In addi-
tion, the Angolan fruits, being capitate and somewhat constricted
below the apex, could not be described as sub-rounded turbinate
(turbinato-subrotunda) as noted in the protologue of H. guineensis.
H. congensis was a nomen nudum. The species was therefore
named H. doreyi by me in honour of the collector who had pro-
cured for me a most complete specimen of the species and also
many details of the plant’s early and later growth. Besides this.
there were from the same region two more species which obviously
did not belong to the Ventricosae.

I was also fortunate enough to receive from Tomlinson speci-
mens from Ghana and so was able to verify that what he had
named as H. guineensis was identical with H. togoensis. In view
of this I included H. guineensis sec. Tomlinson and H. togoensis
Becc. in the synonymy of H. crinita Gaertn. as newly typified by
me but excluded from it H. dahomeensis and H. macrosperma
(Furtado in Rev. Garcia de Orta 15, 1967: 449 & 459 (1970)).

A Search for Literature

When I was trying to determine the origin of a specimen in a
Lisbon herbarium, a British systematist, who had widely explored
the regions along the Niger and the neighbouring countries,
informed me that Hyphaene spp. are never found in low, humid
regions, and that in Ghana, Nigeria, etc. they are found only in
the upper reaches of the river Niger where it passes through arid
country. He therefore rejected my suggestion that the specimen in
question could have been from any of the collectors who had
visited only the lower regions of the Congo, Nigeria, Ghana or
any of the islands in the Gulf of Guinea i.e. the collectors who
could have sent the specimen to Lisbon.

This surprised me indeed, for most Hyphaene specimens with
which I was acquainted in the Lisbon Herbaria were from the
lower regions, often from the sea-coast of Mozambique, Angola
and the Congo, and Drude (Engl. Jahrb. 21: 123 (1895)) had
recorded at least one species from the coastal region of Loango.
Consulting again Schumacher’s account (Kong. Danske Vidensk.
Selsk. Nat. Math. 3: 25-26 (1828)) of Thonning’s exploration,
it became obvious that the latter had botanized generally in the
low regions around Accra and other places as far east as the

318 Gardens’ Bulletin, Singapore — XXV (1970)

Lower Volta i.e. in the territory that constituted the then Danish
Guinea. Now, Tomlinson’s field notes on the specimens he pre-
sented to me state that the species he had recorded for Ghana
was from the higher regions only and was common in North
Nigeria. This was also true according to Irvine (1930 & 1961)
who had identified it as H. thebaica. It was evident therefore that
H. guineensis sec. ‘Tomlinson could not be Thonning’s species,
though it was the same as Irvine’s H. thebaica.

Chipp’s A List of Trees, Shrubs and Climbers of the Gold Coast
(1913) and The Gold Coast Forest: A Study in Synecology (1927),
did not list any Hyphaene species from Ghana though the latter
work recorded the results of careful sample surveys of a large
area Of Lower Ghana. However, an inquiry into the account of
Die Loango Expedition 1873-1876 made under the patronage of
Die Deutschen Gesellschaft Zur Erforschung Aequatorial-Africas,
published in three parts 1879-1882 & 1907, furnished very interest-
ing data. Guessfeldt (Abth 1: 46 (1879)) stated that.H. guineensis
occurred either isolated or more frequently singly in open groves
in narrow strips of sandy ground parallel to the sea-coast. The
frontispiece plate, drawn by one of the artists of the Expedition,
shows palms with separate, non-cespitose and unbranched stems,
few with two stems growing together. Apparently this is the same
species which Smith had identified from its outward appearance
as a Hyphaene species, but Brown thought that this was a species
of Corypha (= Borassus) because the branching of the stem was
not mentioned, overlooking the fact that Smith had already iden-
tified another fan-leafed species as Corypha (Tuckey, Exped. Zaire
App. 5, 456-457 (1818)). If all Hyphaene species were taken to
produce branched stems as Brown and others then considered,
it need not be stressed when naming the genus, even though a
species would fail to bifurcate occasionally.

Unfortunately Wright (1901) consulted only a drawing of the
palm on the frontispiece of the first part of Guessfeldt (1879) who
had observed the palm in Cabinda on the Loango Coast, north of
the River Congo (Zaire). But in the first half of the third part
of the Report (1882), Pechuel-Loesche mentions the occurrence
of ‘‘this erect (unbranched), sea-loving, fan-leafed, forest-shy”
non-cespitose palm, either alone or in open rows, or in extensive
groves on sandy places along the sea-coast from the eastern part
of Liberia near Cape Palmas eastwards to the Ivory and the Gold
Coasts, and the Niger delta and southwards to Angola a little
south of 7° L.S., near Ambrizette. Further south a repeatedly
divided species identified as H. coriacea* and H. benguellensis are
reported to occur.

The maritime species of the north as well as of the south was
idéntified as H. guineensis because it bore an undivided, solitary
stem, and because it grew along the sea-coast. According to
Pechuel-Loesche, humid winds seem to make the Ivory and the
Gold Coasts less favourable to vegetation and apparently also to
H. guineensis which was less common here than on the western

* H. coriacea is native of East Africa.

Furtado — Hyphaene guineensis 319

14

ffalhn ei

Fig. 3. Hyphaene guineensis Thonn. (Neotopotypus — K)

A and D: duo fructus similiformes sed magnitudine dispares. B: Apex
fructus vertice visus. C: Aspectus ejusdem ab imo magis versus
carinam.

320 Gardens’ Bulletin, Singapore-—X XV (1970)

side of Cape Palmas. However, he could not determine whether
this sparseness of the palm was due to natural influence or to
human interference as the places wherein this Hyphaene should
have thrived were utilized by the local villagers to grow coconuts.
Elsewhere in the east both coconuts and H. guineensis were found
erowing together.

A New Search for the Palms

From the observations made by Mr. J. D. Sampayo d’Orey in
whose honour H. doreyi was named, the palm remains stem-
less even for a number of years, and the same fact has been
recorded by Welwitsch (on herbarium labels of specimens not
definitely associated with any fruits) who stated that vast tracts
might be noticed along the sea-coast covered with stemless palms
which from time to time grow tall individually and produce flowers
and fruits. If such a phenomenon occurred also in Ghana, botanical
collectors would generally have ignored the plants. Further, modern
botanical explorers in Upper Guinea seemed to hold the view that
the species of Hyphaene grow only in the higher regions. But, as
we have seen from Schumacher’s information quoted above, the
holotype of H. guineensis must have come from the lowlands.
This and the remarks of Pechuel-Loesche that an unbranched form
of H. guineensis was widely distributed along the sea-coast from
Cape Palmas eastwards to the Gulf of Guinea and southwards to
Angola suggested the need of exploring the low regions of Upper
Guinea for an unbranched species of Hyphaene.

Accordingly, I wrote to several institutions in Liberia, the Ivory
Coast, Ghana and the neighbouring regions, to see if any species
of Hyphaene could be located in the lowlands especially along
the sea-coast. Unfortunately, the institutions that replied had no
specimens of such a palm in their herbaria, nor were they aware
of the existence of any living specimens in the region. The Her-
barium of the University of Ghana had no specimens, but Mr.
LL. M. Quartey, Curator of the University Gardens, Legon, Ghana,
and Dr. M. Hossain, Botanist at the University and Curator of the
University Herbarium, together obtained for me specimens of
‘“‘Hyphaene guineensis sec Tomlinson” cultivated at the University
Gardens. They informed me that these plants were probably a
progeny of the plants so labelled at the Achimota School Arbore-
tum. Obviously these were the plants which Irvine had identified
as H. thebaica in his books, but the change in nomenclature was
made by Tomlinson* long after Irvine’s retirement from the
Achimota School.

Since this plant did not represent a species of the lowlands, both
Dr. Hossain and Mr. Quartey were encouraged to visit some of
the possible sites where Thonning might have found his H.
euineensis. They were successful in locating unbranched Hyphaene

*On a specimen he sent me from the Achimota School Gardens,
Tomlinson remarked: “Apparently introduced but wild in North Nigeria.”

Furtado — Hyphaene guineensis

id
rages
ih in

i th
Hate
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Higy: fs

Cie
yd
dc Vr

' PTE : ein?
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* in Ca Tan

aap:

ie
<n AN \
aw

JURAIM! DEL |

Fig. 4. Hyphaene guineensis Thonn. (Ayittey GC. 37,579) (forma H. doreyi)
A: Fructus. B: Fructus geminatus. C: Sectio fructus verticalis. D:

Aspectus ejusdem vertice visus.

322 Gardens’ Bulletin, Singapore — XXV (1970)

palms along the sea-coast near Accra and along the Lower Volta
region. The fruits varied a great deal but there is no doubt that
the specimens represented H. doreyi. Geographically this could
be readily accepted as Thonning’s H. guineensis, but as there were
important differences in the fruit, a further inquiry was necessary.
Elderly people of the locality informed Dr. Hossain that the palm
generally grows along the sea-coast though it has now disappeared
from many places.

Thonning’s Description of the Fruit

Thonning gave an extensive description of the leaves and male
flowers. He did not see the female flowers, and gave a meagre
description of the fruit. From the fact that he did not mention
that the stem was branched or cespitose, it can be inferred that
the palm was solitary and had an unbranched stem. The most
important part of the long description is ‘‘drupa turbinato-
subrotunda, obtusissima, obsolete trigona’’ which may be trans-
lated as follows: ‘‘fruit turbinately sub-rounded, very obtuse at
the top, obsoletely trigonous.”’

Wendland (Bot. Zeit. 39: 92 (1881)) described the fruit as
“‘obliquely depressed, obovate, flattened at the vertex, rounded
or obtusely trigonous, 6 cm tall, 60 x 65 mm in diam. The seed
obliquely roundish, hardly 3 cm high, 37-40 mm in diam. From
West Coast of Central Africa.”

The fact that Wendland was a keen collector of palm fruits and
that his description of the fruit of H. guineensis agrees well with
that given by Thonning suggest that he might have had access to
one of the isotypes distributed by Thonning to several botanists
in Europe. Moreover Wendland gave the dimensions of the fruit
and the seed and stated that the fruit came from ‘‘West Coast of
Central Africa’’, particulars he could not have known except from
herbarium specimens because no one had mentioned them before.
In fact subsequent workers on the species have ignored these
particulars. Neither Thonning nor Wendland described the fruits
as being abruptly capitate and somewhat constricted below the
apex or in the middle, characteristics of H. doreyi both in Angola
and in Ghana. However it has to be recalled that Pechuel-Loesche
mentioned that the sea winds seemed to affect unfavourably the
growth of vegetation in the Ivory and the Gold Coasts in general,
including also H. guineensis. Variations in the fruit could have
been one of the responses, or else there could be another lowland
species in Ghana. Hence Dr. Hossain and his associate were
requested to make a further survey of the areas, while all the
specimens preserved in the Kew Herbarium from Upper Guinea
were obtained on loan for study.

The subsequent explorations in Ghana were restricted to the
previous localities in former Danish Guinea where Thonning was
a Danish official; the newer specimens were again more akin to
those from Angola. However the Kew Herbarium has from an
indefinite place in Ghana fruits which, though representing H.

Furtado — Hyphaene guineensis

Wh,

323

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Cy \\(\ NT;
Ay
DW) Hie SN

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al
7 ho

———

ms,
\\\Nas
Tees an SS SA
SS
ASS
¢ } i
¢ MD Lis fit 9
f Wh -
“fii;
) ‘|

Fig. 5. Hyphaene guineensis Thonn. (leg. Hossain GC. 37,578).
A: Fructus latere visus. B: Sectio ejusdem verticalis. C: Aspectus

ejusdem vertice visus.

324 Gardens’ Bulletin, Singapore — XXV _ (1970)

doreyi, agree generally with Thonning’s and Wendland’s descrip-
tions of H. guineensis. Since the fruits have a dark brown epidermis
which cracked in the parts exposed to the sun (“‘cortice fragili’’
according to Thonning), there is no doubt that the specimen is
from the lowland, and the name H. thebaica given in the Her-
barium is a misnomer, for the latter species grows on highlands
and produces fruits with yellowish brown epidermis which does
not crack even on drying. Hence the specimen in the Kew Her-
barium is chosen as the neotype of H. guineensis with H. doreyi
as a synonym.

As said above, Beccari, relying on his vast experience with the
genus which shows a great tendency to differentiate into localized
forms, discounted the idea that the distribution of one and the
same species or variety could occur in such wide areas as the
sea-coast of Angola south of the equator near the River Quanza
and in Upper Guinea above the equator. However H. guineensis
seems to have originated in Angola and is distributed to the north
by the Benguela Ocean currents — a factor which Beccari had
not taken into account. Another peculiarity generally observed in
Angola but so far not in Ghana is that the species form stemless
colonies as mentioned above, sometimes occupying vast tracts of
land. But, after some months or even years, the specimens grow
stems individually and produce flowers and fruits.

Hyphaene crinita Gaertn.

In trying to identify H. guineensis Thonn. I stumbled on the
problem of the identity of H. crinita rather unexpectedly, because
H. crinita was generally regarded as a species from South East
Africa and H. guineensis was described as from North West Africa.
When I was working in Lisbon, my first conclusions as to the
identity of H. crinita were drawn and later published in the Review
Garcia de Orta (15 (4), 1967: 449 (1970)). These were based
entirely on the study of the figures published by Gaertner (1891)
and Beccari (1924). Subsequently I have had opportunities to
examine the actual type fruits of H. crinita and H. baikieana
Furtado (= H. macrosperma sec Becc.), and good fruits of H.
togoensis from the palms cultivated in Ghana. As a result I found
that the figures of H. macrosperma sec Becc. are partly misleading
and that these taxa had better be kept separate.

The probable reasons why Martius was misled to confuse H.
crinita with H. natalensis are given in a paper on the identity of
the latter species, published in this Bulletin (pp 283-287).
However, it is often overlooked that the Dutch had a “factory”
(comptoir) in Ghana and also some plantations, so that nuts from
this region could have easily found a place in the seed collection
of the Botanic Garden at Amsterdam where the type fruit of H.
crinita was found. It is true that Lawson and Price (Bot. Journ.
Linn. Soc. Lond. 62 (3): 321 (1969)) have ignored this fact and
quoted Junghans to show that the former Danish Guinea is the

Furtado — Hyphaene guineensis 325

ey
sy,

Ra
iT WN
il

an »

Fig. 6. Hyphaene macrosperma Wendl. (ex Beccari t. 18 figs. 7 and 71).
A: Entire fruit; B: Longitudinal section.

326 Gardens’ Bulletin, Singapore —XXV (1970)

area represented by Ghana. But Jeppesen (Danish Plantations on
the Gold Coast in Geogr. Tidskrift 65: 73-88 (1966) shows that
the Danes had only small plantations and defence posts dotted
in the eastern parts of Ghana especially towards the Lower Delta,
and Lasegue (Musee Bot. Delessert: 70 (1845)) notes that Palisot
de Beauvais from Lisbon had reached Chamah, a Dutch “‘factory”’
(comptoir) on the Gold Coast lying between the Cape of Three
Points and Cape Corse (modern Cape Coast) and thence to Koto,
a Danish ‘“‘comptoir’ on the same coast on the River Volta.
Therefore the type of H. crinita in Amsterdam could have been
sent from Ghana by a Dutch planter.

The fact that H. crinita is closely allied to H. togoensis and
H. baikieana, both of which are based on the types collected in
Upper Guinea supports the view that the type of H. crinita had
also come from that region. All three species have fruits which
are broad at the apex and narrowed at the base and which are
slightly longer than broad. The fruits of H. crinita and H. baikieana
are on the dorsal side almost vertically straight though the base
is slightly rotundate in the former, broad and cordulate in the
latter. In H. togoensis the fruit is dorsally convex. The broadest
diameter is below the middle in this species, in the middle or
slightly above in H. crinita, and near the apex in the third. In
both H. crinita and H. baikieana fruits are laterally compressed
so that it appears somewhat triangular in cross-section. Though
the available fruit specimens of H. baikieana have only embryonic
seed, that of H. crinita is half-developed (known only from one
fruit), and those of H. togoensis are fully developed. The distinc-
tions shown above seem to justify recognition of the three taxa as
distinct species. On the other hand, none of these three species
are allied either to H. thebaica or to H. natalensis.

The Species form Upper Guinea

Since H. guineensis has been definitely identified, advantage is
taken here to enumerate the species that have been found in the
region and to give their synonyms.

1. Hyphaene baikieana Furtado spec. nov

H. crinita sec. Furtado in Rev. Garcia Hort. 15, 1967: 449 Ic. 2
fig. A tantum (1970) quoad synonymum H. macrosperma sec.
Becc. p.p. Figure 1.

H. guineensis sec. Drude in Engl. Jahrb. 21: 124 (1895) in nota.

H. macrosperma sec. Drude op. cit.: 123; Becc., Borass: 27 t. 19
fig. 6 (1924) p.p.

H. crinita valde affinis, sed fructibus bilateraliter compressis,
in sectione transversa trigonoideis, supra stigma abrupte ventricosis,
dorso gibbosis, apice oblique truncatis haec species sat distincta.

Furtado — Hyphaene guineensis 327

Fig. 7. Hyphaene tuleyana Furtado (Holotypus — K).

A: Fructus latere visus. B: Aspectus ejusdem ab imo. C: Aspectus
ejusdem dorsalis. D: Fructus verticaliter sectus ut partis interiores
et earum dispositio appareant.

328 Gardens’ Bulletin, Singapore — XXV (1970)

Fructus omnes immaturi visi, unusquisque cum semine vix
evoluto, ambitu obovato pyriformes, ex dorso latiores ventrem
versus angustati, autem lateraliter compressi, dorso ad basin gib-
bosi vel sub-cordulati et ad apicem rotundati, ventre supra stigma
abrupte carinati, vertice oblique truncati, centro depressiusculi, ad
apicem carinae altiores, epidermide non fragili, flavescente minute
punctato, 7-8 cm alti, 5—6 cm lati, 44.5 cm crassi. Infructescentiae
rami duo visi, usque ad 15 cm longi, uterque amenta tres, 8-11
cm longa, 1 cm diam. ferens.

AFRICA BOREO-OCCIDENTALIS: Regio ad Fluvium Niger
(Baikie leg. circa 1859 — K, holotypus).

Since this specimen belonged to ‘‘Dr. Baikie’s Niger Expedition’,
both Drude and Beccari separately mistook it to be the type of
H. macrosperma, a species described to have an “ovate” fruit as
in A. thebaica but rounder in cross-section, and not obovate-
pyriform and triangular. Geographical considerations misled Drude
to identify this species as H. guineensis.

The peculiar facets shown in the entire fruits photographed by
Martelli for illustrating Beccari’s monograph are somewhat mis-
leading and the fruit represented in Furtado figure 1, B, (copied
from Beccari) is not found in the collection at Kew, though it is
the one that had made me identify Beccari’s H. macrosperma
with H. crinita.

Because the fruit is yellow and ventricose and shows no cracks
in the epidermis, I surmise that, like H. crinita, this species is an
inland one and of higher and drier regions and that it has a stem
that branches dichotomously.

The specimen was originally named as ‘“‘Hyphaene macrocarpa
Wendl.?”” but later named by Martelli as ““H. macrosperma
Wendl.”’ on grounds that it is so named by Beccari in his manu-
script on “‘Borasseae’”’. Wendland did not describe any species
called H. macrocarpa and so this might have been an error for
“H. macrosperma Wendl.?’’.

The holotype is indicated as if it was collected by Barter in
Jan. 1860. However, Barter had died in July 1859 (Hutchinson
& Dalziel, Fl. W. Trop. Afr. 1: 71 (1927)). The specimen is also
indicated as being from “Dr. Baikie’s Niger Expedition”, with
which Barter seems to have been associated only in 1857, though
all the collections from Dr. Baikie’s Niger Expedition (1854-59)
are attributed erroneously to Barter. I have therefore not cited
the name of Barter. Further, the date “Jan. 1860” seems to be
the date when the specimens were received at Kew or when they
were sorted out and incorporated into the Kew Herbarium. A
duplicate of this specimen was donated to Drude in Berlin and
was similarly labelled (op. cit. p. 124 footnote).

2. Hyphaene crinita Gaertn., Fruct. 2: 13 t. 82 fig. 4 (1791);
Mart. Palm. 3 ed 2: 225 (1949) pro parte typica; Becc. in
Agric. Col. 2: 157 (1908) 157 p.p. Borasseae: 29 fig. 13 (1924):
Furtado in Rev. Garcia de Orta 15 (4), 1967: 449 t. Ic. 2 excl.
A & B, excl. syn. et addenda: 459 t, 19 fig. infra (holotypus)
(1970). Figure 2,

Furtado — Hyphaene guineensis 329

Fig. 8. Hyphaene thebaica (Ethiopia — K).

A: Lateral view of the fruit. B: Its dorsal views. C: The basal view.
D: Nut vertically dissected.

330 Gardens’ Bulletin, Singapore — XXV (1970)

ws Wt
hy i\,
"0 |

Heh uu i, Yi
i) Wu f i Mi

E

Fig. 9. Hyphaene thebaica (cult. University Garden, Cairo, Egypt, leg.
Dr. Vivi Tackholm).
A: Fruiting spikelet with the topmost fruit as seen dorsally. B: Fruit
as seen laterally. C: The basal view of the fruit. D: Vertical section
of the fruit. EZ: Hastula.

Furtado — Hyphaene guineensis 331

H. thebaica sec Mart., Palm. 3: 227 (1839) quoad synon H.
crinita Hutch. & Dalz., Fl. W. Trop. Afr. 2: Appendix: 507
(1955) p.p.

AFRICA: Holotypus in Herb. University of Tiibingen, Germany.

3. Hyphaene guineensis Thonn. apud Schum. in Kong. Dansk.
Vid. Natur. Math. Afhand. 4: 219 (1829); Guessfeldt, Die
Loango Exped. 1873-76, Abt. 1:46 (1879) & Titelbild; Wendl.
in Bot. Zeit. 39: 92 (1881); Pechuel-Loesche, Die Loango Exped.
Abt. 3 (1): 76 & 121 (1882); Drude in Engl. Jahrb. 21: 110 &
123 (1895) p.p.; Rendle, Cat. Afr. Pl. Welw. 2: 83 (1899); Wright
in Fl. Trop. Afr. 8: 121 (1901); Becc., Borasseae: 47 (1924);
Faber in Schimp., Pflanzengeogr. ed. 2, 1: 533 fig. 166 (1935);
non Hutchinson in Hutch. & Dalz., Fl. West Trop. Afr. 2:
392 (1936) & Tomlinson, Anat. Mon. II. Palmae: 158 (1961).

Figures 3-5.

H. congensis Kirk kin Salom.,Die Palm. Gewachse: 43 (1887);
Drude op. cit.: 124 nota; Becc. op. cit.; 49: nomen nudum.

H. doreyi Furtado in Rev. Garcia de Orta 15, 1967: 450, tt.
5 & 6 (1970) syn. nov. |

H. thebaica sensu Dalziel in Appendix: 508: (1955) p.p.; Russell
in Fl. W. Trop. Afr. 3 (1), ed. 2: 169 (1968) p.p.

WEST AFRICA: Ghana loc. incert. (Colonial & Indian Exhi-
bition of 1886 No. 92 — K, neotype); Tema Paradise Beach
(Hossain GC 37,578 & GC 38,675); Lower Volta Agbozume
District, at Sonuto near Keta Lagoon (Quartey & Ayittey; GC
37,579, & Hossain & Enti GC 38,668. :

Several specimens of this species in the Herbaria of Kew, BM
& Lisbon have been named by me as H. doreyi.

This species seems to have originated in Angola in the region
of Quanza and become widely distributed further to the northern
regions of Upper and Lower Guinea by the Benguela Ocean
currents.

In view of the great many variations noted in the collections
made by Dr. M. Hossain of Ghana and his assistants, H.
gossweileri Furtado (1970) might, on further studies, prove to be
a form of H. guineensis.

4. Hyphaene macrosperma Wendl. in Bot. Zeit. 31: 92 (1881);
Drude in Engl. Jahrb. 21: 123 (1895) non in nota; Becc.,
Borass.: 27 (1924) parte typica; Furtado in Rev. Garcia de Orta
15 (4), 1967: 453 Ic. 4 figs. A & B (1970). Figure 6.

H. dahomeensis Becc., Borass.: 48 tt. 18 fig. 7 & 42 figs. 14
(1924).

AFRICA BOREO-OCCIDENTALIS: Dahomey (Paisson-
Herb. Becc. holotypus).

Beccari & Drude identified this with H. haikieana.

332 Gardens’ Bulletin, Singapore — XXV_ (1970)

a: i 7
TN 9 ey den? 0
\ Ah) A a | - : /
( \ in “Bhi pie? Yip 4;
‘

\

B 4 nif

jucaimi
del.

Fig. 10. Hyphaene thebaica (Egypt, leg. Walsingham — K).

A: Lateral view of the fruit. B: Fruit as seen from below. C: The
dorsal view of the fruit. D: Fruit as seen from the top. E: Vertical
section of the fruit.

Furtado — Hyphaene guineensis 333

5. Hyphaene tuleyana Furtado spec. nov. Figure 7.

H. thebaicae arcte affinis sed fructibus fere tertia parte minori-
bus, ambitu laterali quadratis vel obovoideo quadratis, apice
curvatis, ventre carinatis, carina etiam apicem versus valde dilatata
haec species admodum diversa. Ab H. occidentali, quacum propter
magnitudinem fructus facile confusa, fructibus conspicue carinatis
in sectione transversa ovoideo-trullatis, ambitu laterali obovato
quadratis et dorsali rotundato-quadratis haec species sat distincta.

Caulis basi solitarius, circa ad 3 m. altitudinem semel bifurcatus,
ramis 7-9 m. longis. Amenta duo visa, parte fertili 18-20 cm.
longa, 2 cm. in diam., parte basali sterilia, 4-5 cm. longa. Fructus
circa 5 cm. altus, 6—6.25 cm. latus, 5 cm. in diam., in perimetro
stigmatico pedicellari ambitu obovoideo-apice arcuatus, dorso
curvatus, supra stigma carinatus, carina valde dilatata, basi inflexa,
imo sulcata, utrinsecus paulo depressa, apice altiore quam dorsum
vel eo aequa, ambitu dorsali quadrangulari-oblongus, in angulis
rotundatus, basin versus paululo dilatatus vel non, medio vel
supra cinctulus, apice recurvus; vertice in linea dorso-carinali
sulcatus, sulco ante apicem carinae terminato; sectione transversa
late ovoideo-trullatus, epidermide non friabili, luteo brunnescente
vel cinnamomeo.

AFRICA BOREO-OCCIDENTALIS: Nigeria borealis (Tuley
in 1964 — K, holotypus).

A very close ally of H. thebaica, from which this species is
readily distinguished by its much smaller fruits which are obovate-
quadrangular on the broadest side with arcuate apex, and almost
quadrangular on the dorsal side, sometimes a little broader at the
base and often constricted above the middle. The carina is dilated
above and its tip is often the highest point in the fruit. The dent
at the apex is more towards the carina than in the centre of the
vertex which is the case in H. thebaica.

The fruit is more or less the size of that of H. occidentalis but
differing from it in being obovate in the broadest side and having
a well-developed carina making the species ovoid-trullate in cross-
section.

A photograph taken by Tuley (apparently in 1964) shows that
the palm does not divide underground but only when the stem is
about 3 metres high. The tree was branched only once.

The specimen and the photographs were apparently made in
1964 and probably Russell (Fl. W. Trop. Afr. 3 (1) ed. 2: 169
(1968)), included also this species under H. thebaica.

Tomlinson identified the specimen of H. togoensis growing in
the Achimota Gardens as H. guineensis and referred in the note
at the herbarium to the wild North Nigerian species which may
be H. tuleyana. This may also be partly the species from Bornu
in North Nigeria which Drude had referred to as H. thebaica
(Engl. Jahrb, 21: 110 (1896)).

334 Gardens’ Bulletin, Singapore — X XV (1970)

Hyphaene thebaica (L.) Mart. Figures 8-10.

Since this binomial has often been misapplied to several un-
related species, advantage is taken here to illustrate three of its
forms. The species has to be typified on t. 133 of Martius (Palm.
1838-49), while Beccari’s t. 20 (Borasseae, 1924) has to be ex-
cluded, being apparently a member of the group referred by
Beccari as “Hyphaene multiformis’” (Borasseae: 32 (1924)).

The fruit of H. thebaica varies a great deal in size, depending
mostly upon ecological conditions. It is always ovate or ovoid
in shape, being almost truncate at the base where it is more or
less grooved on the sides of the keel. This last is broadly ventricose
in the lower half and gradually narrowed towards the obtuse or
slightly retuse apex. The dorsal side is arcuate, broad towards the
base, slightly narrowed towards the apex.

Acknowledgements

This study has been made possible through the generous co-
operation of several persons and institutions to whom I am very
grateful. In particular I should like to thank the following: The
Director, British Museum (Science), London, for the photocopies
of the pertinent parts of the Die Loango Expedition, 1873-1882,
by Guessfeldt & Pechuel-Loesche; the Director, Royal Botanic
Gardens, Kew, for the loan of the herbarium specimens from
Upper Guinea and Egypt and for the photocopy of the treatment
of Hyphaene by Russell in Flora of West Tropical Africa, 1968;
the Curator, Botanical Museum, Copenhagen, Denmark, for the
photocopy of Jeppesen’s account of Danish Plantations in the
Gold Coast (Geogr. Tidskr. 65 (1966)); the Curator, the Rijksher-
barium, Leiden, the Netherlands, for photocopies of Wendland’s
papers on Hyphaene; the Keeper of Herbarium, University of
Tuebingen, Germany, for the loan of Gaertner’s type of H. crinita;
the Centro de Botanica, Lisbon, Portugal, for the copies of the
pertinent parts of Schumacher’s paper on Thonning’s journey in
the Gold Coast; Dr. Vivi Tackholm, Faculty of Science, Cairo
University, Giza, Egypt, for the specimens of H. thebaica grown
in the University Garden in Giza; Dr. M. Hossain of the University
of Ghana and his assistants for obtaining good specimens of H.
togoensis and for visiting some localities where Thonning had
worked, and collecting from there specimens of H. guineensis:
and the Director of Botanic Gardens, Singapore, for providing
me the library and herbarium facilities to carry out my work.

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“Vol XXV, Part Tl 15th October, 1971

CONTENTS

= Bo. PAGES
_Tixter, P.: A Contribution to Bryological smnowledge of
Z a Fraser’s Hill Station (Malaysia) - - 335-353
Be _ CORNER, E, J. H.: Meruloid Fungi i in Malaysia - - - 355 — 381.
Be 2 REVIEW: Grasses of Malaya (H. Keng) - - - - 383 — 384
P< - Oprruary: Juraimi bin Samsuri 3 apiwe os - 385

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————— ee —

A Contribution to the Bryological Knowledge of
Fraser’s Hill Station (Malaysia)

By P. TIrxier*

SUMMARY

The author gives the list of his own collections from Fraser’s Hill
(Pahang). It indexes 78 species of Mosses and 67 of Liverworts and four
are new to Science: Syrrhopodon fraserianus, Trichosteleum fruticolum,
Cololejeunea pahangiana and Lejeunea pilifera.

In the course of my monographic studies of the Liverwort genus
Cololejeuna, I was struck by the paucity of herbarium collections
and the small number of localities visited in Malaya. In the work
of Benedix (1947) only 14 species were listed while the localities
were confined to the Southern part of the Malayan Peninsula such
as Mt. Ophir and to Singapore. His publication drew attention at
once to Fraser’s Hill as a hill station for the collection of
Liverworts. As for the Malayan Mosses, the last publication is by
Dixon and dates as far back as 1926. Subsequently few data has
been published.

Fraser’s Hill is a station situated 112 km from Kuala Lumpur,
at an altitude of about 1,400 m on a ridge (of the Main Range)
near the water parting between Pahang and Selangor, and has an
annual rainfall between 3,000 and 4,000 mm. Discovered in 1916,
the site was surveyed in 1919 and was soon developed by the
Governments of Pahang and the Federated Malay States. The
location and the vegetation resemble those observed in Bach Ma
(Central Vietnam) in that Fraser’s Hill also lies on top of a ridge
and the vegetation is similarly an oak mountain forest. As is also
the case in Bach Ma, Cyathea reaches 5-6 m high and also many
palms such as Caryota, Licuala, and Pinanga occur. The Northern
side may be wetter than the Southern, but nowhere was the elfin-
woodland forest seen. The vegetation remains relatively disturbed
and bushes of ferns represent the secondary vegetation. It was
observed that practically all the collections in windy and sunny
localities were made on the very various coniferous trees planted
along the roads of the station and in the garden of Fraser’s Hill
Hotel. For the Mosses in my list, new species for Fraser’s Hill are
marked ‘“*” and those for Malaya marked “**”,

The author wishes to thank Dr. C. X. Furtado, Botanist in
Singapore, for kindly correcting the Latin diagnosis in this paper.

*Laboratoire de Cryptogamie, Muséum National d’Histoire Naturelle
(Paris); and, Faculté des Sciences, Phnom Penh (République Khmére)

336

Gardens’ Bulletin, Singapore — XXV (1971)

Mosses

*

*

*

Dicranoloma braunii (C. Muell.) Par.

On coniferous barks, 25/XII/1969, nos 4611, 4763, 4762.

Indochina, Malesia, New Guinea, New Caledonia, Pacific
Islands.

Leucoloma molle (C. Muell.) Mitt.

On trunks in evergreen forest, 26/XII/1969, no 4756.

Indochina, Malesia.

Campylopus caudatus (C. Muell.) Mont.

On trunks, 28/XII/1969, no 4858.

Himalaya, Ceylon, Sumatra, Borneo, Moluccas, Luzon,
Mindanao.

Campylopus hemitrichus (C. Muell.) Jaeg.

On trunks and rocks, 26/XII/1969, no 4764.

Luzon, Borneo, Sumatra.

Leucobryum aduncum Doz. & Molk.

On rotten logs in evergreen forest, 26/XII/1969, no 4750.

From Himalaya to New Caledonia.

Leucobryum bowringii Mitt.

On bark in evergreen forest, 28/XII/1969, no 4838.

Ceylon, Malesia, Indochina, China, Formosa, Japan.

* Leucobryum javense (Brid.) Mitt.

*

*

Very common on trunks, grounds and banks, 25/XII/1969,
no 4610.

India, Ceylon, Indochina, Malesia, China, New Guinea.

Octoblepharum albidum Hedw.

On trunks of coniferous trees, 25/XII/1969, no 4608.

Pantropical.

Exodictyon blumii (C. Muell.) Fleisch.

On trunks, in evergreen forest, 26/XII/1969, no 4739.

Java, Borneo, Luzon, Negros, Thailand, Cambodia, Vietnam.

Syrrhopodon albidus Thw. & Mitt.

On trunks, in evergreen forest, 26/XII/1969, no 4760.

Ceylon, Malaya.

Syrrhopodon fraserianus sp. nov.

On bark, in sunny position, 26/XII/1969, no 4769.

Syrrhopodon tristichus Nees

On bark in evergreen forest, 26/XII/1969, no 4740.

Ceylon, Sumatra, Java, Ambon, Borneo, Thailand, Luzon,
Negros.

Thyridium fasciculatum (Hook. & Grev.) Mitt.

On twigs in sunny position, 25/XII/1969, nos 4719, 4775.

Nepal, Ceylon, Malaya, Thailand, Cambodia, Java, Sumatra,
Moluccas, Mauritius, New Caledonia, Louisiade, Palau,
Carolines, Samoa, Pitcairn.

Thyridium undulatum (Broth. & Ged.) Fleisch.

On trunks, in sunny station, 26/XII/1969, no 4757.

Sumatra, Java, Sulawesi, Borneo, New Guinea, Singapore.

Tixier — Bryophytes, Fraser's Hill 337

**

*

*

*

*k *

**

Calymperopsis langbianensis P. Tixier
On twigs of coniferous trees, 25/XI1I/1969, no 4721.
Thailand, Cambodia, Vietnam.

Calymperes nietnerii C. Muell.
On trunks in evergreen forest, 26/XI1I/1969, no 4792.
Java, Ceylon, Thailand.

Calymperes serratum A. Braun

On rotten wood in evergreen forest, 26/XII/1969, no 4789.
Indochina, China, Malesia, New Caledonia, Micronesia.
Hyophila involuta (Hook.) Jaeg.

On wet banks, 26/XII/1969, no 4780.

Ceylon, Malesia, Indochina.

Hydrogonium consanguineum (Thw. & Mitt.) Hilp.

On wet banks, 28/XII/1969, no 4843.

Ceylon, Luzon, Singapore, Java.

Brachymenium exile (Doz. & Molk.) Bosch. & Lac.

On wet banks, 26/ XII/ 1969, no 4793.

India, Japan, Formosa, Korea, Ceylon, Java, Sumatra, Hawaii,
Vietnam.

Brachymenium nepalense Hook.

Very common, on trunks, roofs etc., 28/XII/1969, no 4853.

Indochina, India, Malesia.

Rhodobryum giganteum (Schwaegr.) Par.

On banks, 25/XII/ 1969, no 4606.

Mascarhenas, India, Ceylon, Malesia, Indochina, China,
Japan.

Rhizogonium spiniforme (Hedw.) Bruch.

Very common, but weakly developed, 26/ XII / 1969, no 4773.

Pantropical.

Mniodendron divaricatum (Reinw. & Hornsch.) Lindb.

On ground in wet evergreen forest, 26/XII/1969, no 4796.

Sumatra, Java, Borneo, Sulawesi, Luzon, Negros, Mindanao,
Mindoro, Malaya. | |

Philonotis hastata (Duby) Wijk & Marg.

On wet banks, 27/ XII/1969, nos 4821, 4823.

Madagascar, Thailand, Sumatra, Java, Malaya, Borneo, Luzon,
Laos, Hawaii.

Macromitrium cumingii C. Muell.

On coniferous bark, 26/XII/1969, no 4771.

Thailand, Java, Philippines, New Guinea, Carolines, Marshall,
Vietnam.

Macromitrium reinwardtii Schwaegr.
On coniferous barks, in sunny position, 28/XII/1969, no 4836.
Java, Borneo, Sulawesi, Tahiti, Tasmania, Cambodia.

Groutiella goniorrhyncha (Doz. & Molk.) Wijk. & Marg.

On bark and branches of coniferous trees, 25/XII/ 1969,
no 4735.

India, Indochina, Malesia, New Guinea.

338

*

%

*

*

*

*

**

*

Gardens’ Bulletin, Singapore — XXV (1971)

Desmotheca apiculata (Doz. & Molk.) Lindb.

On branches in sunny positions, 27/XII/1969, nos 4817,
4833.

Singapore, Sumatra, Java, Borneo, Ambon, Vietnam.

Racopilum schmidii (C. Muell.) Mitt.

On trunks in evergreen forest, 27/XII/1969, no 4816.

Nilghiris, Philippines, Indochina.

Endotrichella elegans (Doz. & Molk.) Fleisch.

On banks mixed with Aerobryopsis longissima, 28 / XIT/ 1969,
no 4851.

Sumatra, Java, Malesia, Vietnam, Formosa, Philippines,
Carolines.

Garovaglia plicata (Brid.) Bosch & Lac.

On coniferous bark, 28/XII/1969, no 4837.

Sikkim, Sumatra, Java, Ceram, Luzon, Mindanao.

Pterobryopsis crassicaulis (C. Muell.) Fleisch.

On trunk of Podocarpus sp., 25/XII/1969, 4603, 4728.

Ceylon, Sumatra, Java, Philippines, Vietnam.

Pterobryopsis gedehensis Fleisch.

On trunk in evergreen forest mixed with Homaliodendron
flabellatum, 27/XII/1969, no 4801.

Java, Cambodia, Mindanao.

Symphysodontella cylindracea (Mont.) Fleisch.

On trunk in evergreen forest, 27/XII/1969, no 4824.

Sumatra, Java, Mindanao, Ambon, New Caledonia, Samoa,
Tahiti, Rarotonga, Cook Islands, Thailand, Fiji, Vietnam.

Papillaria fuscescens (Hook.) Jaeg.

On trunks and twigs in sunny stations, 25/ XII/ 1969, no 4732.

Himalaya, India, Ceylon, Indochina, China, Malesia.

Aerobryidium filamentosum (Hook.) Fleisch.

Pendulous on twigs in sunny stations, 26/ XII/ 1969, no 4786.

India, Himalaya, China, Ceylon, Indochina, Malesia.

Aerobryopsis longissima (Doz. & Molk.) Fleisch.

On banks and on twigs in sunny stations, 25/ XII/1969, nos
4605, 4718, 4737.

From Himalaya till Pacific Islands.

Floribundaria floribunda (Doz. & Molk.) Fleisch.

Common -on twigs in sunny stations, 25/XII/ 1969, no 4720.

Madagascar, India, Himalaya, Indochina, Ceylon, Japan, New
Guinea, Pacific Islands.

Floribundaria pseudofloribunda Fleisch.

On branches in evergreen forest, 27/XII/1969, no 4815.

Northern Vietnam, Java, Luzon, New Guinea, New Hebrides,
Queensland.

Trachyloma indicum Mitt.

On coniferous trunks in sunny station, 27/ XII/ 1969, no 4803.

Ceylon, Indochina, Malesia, New Guinea.

Tixier — Bryophytes, Fraser’s Hill 339:

*

*

*

*

**

*

**

*%

*

Homaliodendron flabellatum (Smith) Fleisch.
On trunks in evergreen forest, 26/XII/1969, no 4752.
From India to Pacific Islands.

Pinnatella kuehliana (Bosch & Lac.) Fleisch.
On trunks in evergreen forest, 27/XII/1969, no 4814.
Sulawesi, Sumatra, Java, Malaya, Ceram, New Guinea.

Pinnatella microptera Fleisch.

On coniferous trunks in sunny stations, 28/XII/1969, no
4839. ;

Luzon, Borneo, Thailand, Singapore.

Ephemeropsis tjibodensis Goeb.

Epiphyllous in evergreen forest, 28/XII/1969, no 4666.

South East Asia from Laos to New Guinea.

Distichophyllum mittenii Bosch & Lac.

On rotten logs in evergreen forest, 26/XII/1969, no 4785.

Ceylon, Malaya, Java, Palawan, Cambodia, Formosa,.
Aneityum, New Caledonia.

Chaetomitrium torquescens Bosch & Lac.
On twigs in sunny positions, 27/ XII/1969, no 4809.
From Java to New Guinea.

Lopidium trichocladon (Bosch & Lac.) Fleisch.

On trunks in forest and sunny positions, 27/XII/1969, no
4825.

Java, Moluccas, Thailand, Laos, Vietnam, Cambodia.

Cyathophorella tenera (Bosch & Lac.) Fleisch.

On trunks in evergreen forest and in sunny positions, 27/XII/
1969, no 4827.

Sumatra, Sumbawa.

Thuidium meyenianum (Hpe.) Doz. & Molk.
On ground mixed with Pogonatum, 26/ XII/1969, no 4784.
Himalaya, Ceylon, Malesia, New Guinea, Melanesia.

Rhynchostegium celebicum (Lac.) Jaeg.
On trunks in evergreen forest, 25/XII/1969, no 4615.
Luzon, Thailand, Vietnam, Sumatra, Java, Sulawesi, Hawaii.

Campylodontium flavescens (Hook.) Bosch & Lac.

(Syn.: Cribrodontium wernerii Herz.)

At the base of tree in sunny positions, c. fr., 27/XII/1969, no
4826.

From Himalaya to Luzon.

Clastobryella asperrima Dix.

On twigs, 25/XII/1969, nos 4722, 4800; 27/XII/1969, no
4835.

Malaya, New Caledonia, Aneityum.

Acanthocladium alboalare P. Tixier.

On trunk in evergreen forest, 26/XII/1969, no 4772.

Vietnam.

340

*

**

*

**

**

ex

Gardens’ Bulletin, Singapore — X XV _ (1971)

Trismegistia calderensis (Sull.) Broth.

On rotten wood in evergreen forest, 26/XII/1969, nos 4738,
4768.

Sumatra, Java, Malaya.

Trismegistia rigida (Mitt.) Broth.

Same station, 26/XII/1969, nos 4739, 4741.

Indochina, Malesia, New Guinea, New Caledonia.

Meiothecium hamatum (C. Muell.) Broth.

On twigs of coniferous trees, 25/XII/1969, no 4731. (cf)

Ceylon, Sumatra, Java, Fiji, Panay, Samoa, Vietnam,
Cambodia.

Acroporium aciphyllum Dix.

On banks, 28/XIT/1969, no 4848.

Malaya.

Acroporium albidissimum : Dix.

On rotten wood in evergreen forest, 26/ XII/ 1969, no 4751.

Thailand, Malaya.

Acroporium diminutum (Brid.)) Fleisch.

On rotten wood in evergreen forest, 28/XII/1969, no 4855.

Java, Philippines, Moluccas, Malaya, Vietnam.

Acroporium falcifolium (Fleisch.) Fleisch.

On rotten wood in evergreen forest, 27/XII/1969, no 4799.

Sumatra, Java, Malaya, Sulawesi, Philippines.

Acroportum hamulatum (Fleisch.) Fleisch.

On trunks in evergreen forest, 26/XII/1969, nos 4732. 4770,
4781, 4782: 28/XII/1969, nos 4846, 4847.

Java, Luzon, Vietnam, Cambodia, Malaya.

Acroporium longicuspis Dix.

On rotten wood in evergreen forest, 27/ XII/ 1969, no 4842.

Malaya.

Acroporium secundum (Reinw. & Hornsch.) Fleisch.

‘On banks, 28/XII/ 1969, nos 4837, 4847, 4776.

Sumatra, Java, Borneo, Sulawesi, Ceram.

Acroporium cf vincensianum (Thér.) Broth.

Common on rotten logs in evergreen forest, c.fr., 26/XII/ 1969,
nos 4738, 4758, 4798.

Vietnam, Cambodia.

Trichosteleum boschii (Doz. & Molk.) Jaeg.

On trunk in evergreen forest, 26/XII/1969. no 4769.

Indochina, Malesia.

Trichosteleum fruticolum sp. nov.

Small collections on twigs, 28/XII/1969, no 4852.

Known from Bokor in Cambodia.

Trichosteleam hamatum (Doz. & Molk.) Jaeg.

On rotten wood in evergreen forest, 26/XII/1969, no 4787.

From South East Asia to the Pacific Islands.

Acanthorrhynchium. papillatum (Harv.) Fleisch.

On rotten wood in evergreen forest, 26/XII/1969, no 4742.

From South East Asia and Nepal to the Pacific Islands.

Tixier — Bryophytes, Fraser's Hill 341

* Taxithelium kerianum (Broth.) Broth.
On trunks in evergreen forest, 26/XII/1969, no 4765.
Australia, Malesia, Indochina, New Guinea.
* Taxithelium vernierii (Duby) Besch.
On twigs in evergreen forest, 27/XII/1969, no 4819.
Indochina, Malesia.
** Glossadelphus zollingerii (C. Muell.) Fleisch.
On twigs, 27/XII/1969, no 4834.
Java, Sulawesi, Luzon, Fiji, Hawaii.
Ectropothecium buitenzorgii (Bel.) Mitt.
Very common, banks, base of trunks, twigs, 25/XII/1969,.
nos 4725, 4774.
Indochina, Malesia.

* Etcropothecium ichnotocladum (C. Muell.) Jaeg.
On rotten wood in evergreen forest, 27/ XII/ 1969, no 4813.
Luzon, Sumatra, Java, Borneo, Sulawesi, Ambon, Vietnam.

Isopterygium albescens (Hook) Jaeg.

On wood near the ground in evergreen forest, 25/XII/ 1969,
no 4736; 26/XII/1969, no 4745; 27/ XII/ 1969, no 4831.

From Nepal to Pacific Islands.

Isopterygium bancanum (Lac.) Jaeg.

On rotten wood in evergreen forest, 28/ XII/ 1969, no 4839.

Java, Malaya, Banka, Luzon.

Pogonatum junghuhnianum (Doz. & Molk.) Doz. & Molk.

On banks in forest, 27/XI1/1969, no 4830.

Java, Nilghiri, Indochina, Philippines.

Pogonatum macrophyllum Doz. & Molk.

On banks in forest, 26/XII/1969, nos 4790, 4791; 25/XIIL/
1969, no 4604; 27/XII/1969, no 4829; 28/XII/1969, no:
4857.

Malaya, Sumatra, Java, Moluccas, Sulawesi, Luzon, Negros,.
Mindanao, Cambodia.

*

Liverworts

Anthoceros glandulosus Lem. & Lindb.

On stones in the garden of the hotel, 27/ XII/ 1969, no 4820:
Common in South East Asia.

Marchantia acaulis St.

On wet bank, 26/XII/1969, no 4794.

Perak.

Metzgeria furcata (L.) Dum.

Epiphyllous in evergreen forest, 27/ XII/ 1969, no 4703.
Cosmopolitan.

Chiloscyphus argutus (Reinw. Blum Nees) Nees

On rotten wood in evergreen forest, 27/XII/1969, no 4849.
From Tropical Asia to Australia.

* Species new for Fraser's Hill. Die Ww lied 1 bat) Lo
** Species new for Malaya.

342

Gardens’ Bulletin, Singapore— XXV (1971)

Chiloscyphus communis St.

On rotten wood in wet ground, 26/XII/1969, no 4776;
28/XI11/1969, no 4849.

Himalaya, Indochina, Malesia, Japan, Queensland.

Bazzania calcarata comb. nov. ( = Mastigobryum calcaratum
Sande Lacoste, Ann. Mus. Lugd. Bat. 1863 in Stephani
Spec. Hep. III, 1908, 498).

On rotten wood in evergreen forest, 26/ XII/ 1969, no 4766.

Java, Malaya, Borneo.

Bazzania caudata comb. nov. ( = Mastigobryum caudatum
St., Spec. Hep. III, 1908, p. 947).

On rotten wood in evergreen forest, 26/ XII/ 1969.

Pulau Penang.

Bazzania commutata comb. nov. ( = Mastigobryum commuta-
tum L. & G., Spec. Hep. 1839-1851, p. 97 et in Stephani,
Spec. Hep. III], 1908, p. 509).

On bark, 25/ XII/1969, nos 4609, 4612.

Java.

Bazzania intermedia (G.) Lellorg.

On rotten wood in evergreen forest, 26/XII/1969, no 4783.

Ambon, Banda, Samoa, Philippines, Tonkin, Nilghiri, Ryu
Kiu.

Bazzania malaccensis comb. nov. ( = Mastigobryum malac-
cense St., Species Hepaticarum, III, 1908, p. 494).

On rotten wood in evergreen forest, 26/XII/1969. no 4768.

Perak.

Bazzania pectinata comb. nov. ( = Mastigobryum pectinatum
L. & G., Spec. Hep., p. 84 et Stephani, Spec. Hep., IIL,
1908, p. 445).

On bark in forest, 26/XII/1969, nos 4743, 4759.

Bazzania spiralis (Reinw. Bl. Nees) Meijer.
On rotten wood in evergreen forest, 26/XII/1969, no 4755.
Thailand, Malaya, Banka, Sumatra, Borneo, Cambodia.

Lepidozia cf quadridens (N. & E.) Lindb.
On rotten log in forest, 26/XII/1969, nos 4749, 4779.
Java, Indo Malesia, Brazil.

Herberta cf parisii (St.) Miller.

In low situation on trunk, 28/XII/1969, no 4859.

Plant between H. parisii and H. ceylanica. This genus seems
new for Malaya.

Mastigophora diclados (Brid.) Nees.

In clusters along the banks of the roads, 26/XII/1969, no
4767. On wood, in clear forest, 26/XII/1969, no 4754.

Pantropical.

Radula nymanij St.

On trunks, in sunny situations, 27/XI1/1969, no 4810.

Epiphyllous on Araucaria leaves, 26/XI1I/ 1969, nos 4718, 4680.

Java, Samoa, Philippines, Vietnam, New Guinea.

Tixier — Bryophytes, Fraser's Hill 343

Caudalejeunea stephanii (Spr.) St.
Epiphyllous in evergreen forest, 26/XII/1969, no 4718.
From Mascarenhas to Tropical Oceania.

Cheilolejeunea (X enolejeunea) meyeniana Kachroo & Schuster
Epiphyllous in evergreen forest, 26/XII/1969, no 4664.
South East Asia.

Cheilolejeunea (Xenolejeunea) imbricata Schuster & Kachroo.

Bark of coniferous trees, 25/XII/1969, no 4729. Epiphyllous
in evergreen forest, very common, 26/XII/1969, nos 4620,
4671; 27/XII/1969, nos 4685, 4716, 4718.

Tropical Asia and Oceania.

Cololejeunea (Cryptolejeunea) ciliatilobula Schifin.
Epiphyllous in evergreen forest, 26/ XII/1969, nos 4647, 4670.
Vietnam, Cambodia, Malaya, Java, Sumatra, Moluccas.

Cololejeunea (Taeniolejeunea) fioccosa (L. & L.) St.

Epiphyllous in evergreen forest, 27/ XII/1968, nos 4688. 4704.

Formosa, Japan, China, Philippines, Borneo, Java, Sumatra,
Chittagong, Thailand, Cambodia, Laos, Vietnam, Sierra
Leone.

Cololejeunea (Taeniolejeunea) faicatoides Benedix.

Epiphyllous in evergreen forest, 27/XII/1969, nos 4692,
4699, 4703, 4686; 26/XII/1969, nos 4673, 4668.

Sumatra, Java, Borneo, Thailand, Cambodia, Sulawesi,
Philippines.

Cololejeunea (Taeniolejeunea) gynophthalma Benedix.

Epiphyllous on Araucaria leaves, 26/XII/1969, nos 4618,
4635, 4636, 4670.

Sumatra, Java, Vietnam, Thailand, Luzon, Cambodia.

Cololejeunea (Lasiolejeunea) jelinekii (St.) Benedix.

Epiphyllous in evergreen forest, 26/XII/1969, nos 4632,
4635, 4636, 4657: 27/XI11/1969, no 4705.

Sumatra, Java, Nicobar.

Cololejeunea (Lasiolejeunea) karstenii (St.) Benedix.

Epiphyllous in evergreen forest, 26/XI1/1969, no 4646;
27/XI1/ 1969, no 4708.

Java, Vietnam (?).

Cololejeunea (Taeniolejeunea) leonidens Benedix.

Very common, epiphyllous on palms, 26/XII/1969, nos
4620, 4621, 4622, 4623, 4624, 4625, 4626, 4627, 4628. 4679,
4678; 27/XII/1969, nos 4680, 4692, 4698, 4700, 4718.

Sabah, Malaya, Java.

Cololejeunea pusilla P. Tixier.

Epiphyllous in evergreen forest, 27/XII/1969, no 4683.

Thailand, Cambodia.

Cololejeunea (Pedinolejeunea) pahangiana sp. nov.

Epiphyllous in evergreen forest, 27/XII/1969, no 4703.

344

Gardens’ Bulletin, Singapore — XXV (1971}

Cololejeunea (Taeniolejeunea) oshimensis (Horik.) Benedix.

Epiphyllous in evergreen forest, 26/XII/1969, nos 4653,
4654, 4656, 4670, 4671, 4688, 4699; 27/XII/1969, nos
4703, 4714, 4715, 4716, 4718.

Japan, Formosa, Foukkien, Vietnam, Cambodia, Thailand,
Malaya, Sumatra, Java, Sulawesi, Laos.

Cololejeunea (Pedinolejeunea) triapiculata comb. nov.

(= Leptolejeunea triapiculata Herz. Ann. Bryol. 1932, 5, 95)

Epiphyllous in evergreen forest, 26/XII/1969, no 4646,
27/XI1/1969, no 4703.

Java, Sumatra.

Cololejeunea (Chlorocolea) sigmoidea S. Jovet-Ast & P. Tixier.

Epiphyllous in evergreen forest, 26/ XII/ 1969, no 4620.

Java, Sabah, Thailand, Cambodia, Vietnam. -

Cololejeunea wightii St.

Epiphyllous in evergreen forest, 27/XII/1969, no 4708.

Malaya, Thailand, Cambodia, Micronesia.

Colura acroloba (Mont.) S. Jovet-Ast .

Epiphyllous in evergreen forest, 26/XII/1969, nos 4648,
4655, 4658, 4663; 27/XII/1969, nos 4689, 4715, 4716,
4718.

Ceylon, Cambodia, Sumatra, Java, Vietnam, Borneo,
Philippines, New Caledonia.

Diplasiolejeunea brachyclada Ev.

Epiphyllous in evergreen forest, 26/XII/1969, no 4665;
27/XII/ 1969, no 4689.

Caribbean area and South East Asia.

Drepanolejeunea laevicornua St. .

Epiphyllous in evergreen forest, 26/ XII/ 1969, nos 4653, 4660.

Java.

Drepanolejeunea tenera Goeb., forma goebelij Herz.

Epiphyllous in evergreen forest, 26/XII/1969, no 4674;
27/XI1/1969, nos 4699, 4705.

Sumatra, Java, Borneo.

Drepanolejeunea tenuis (Nees) Schiffn.

On twigs of coniferous trees, 25/XII/1969, no 4612.

Sumatra, Java, Philippines, Vietnam.

Drepanolejeunea thwaitesiana (Mitt.) St.

Epiphyllous in evergreen forest, very common, 26/ XII/ 1969,
nos 4658, 4688; 27/XII/1969, nos 4706, 4718.

Ceylon, Sumatra, Java, Borneo, Philippines, New Guinea,
Vietnam, Thailand, Cambodia.

Lejeunea pilifera sp. nov.

On trunks and twigs in sunny positions, 27 / XII/ 1969, no 4811.

Lopholejeunea subfusca St.

On wood in evergreen forest, 27/XII/1969, nos 4802, 4828.

Ceylon, Cambodia, Thailand, Vietnam, Malaya, Sumatra,
Krakatau, Java, Borneo, Philippines, Formosa, Ryu Kiu,
New Guinea, New Caledonia, Samoa, Tahiti.

Tixier — Bryophytes, Fraser's Hill 345

Leptolejeunea massartiana Herz. ex. Schiffn.
Epiphyllous in evergreen forest, 27/X1I/1969, no 4714.
Java, Sumatra.

Leptolejeunea foliicola St.
Epiphyllous in evergreen forest, 27/XII/1969, no 4716.
Malaya, Sumatra, Java, Cambodia, Vietnam, Philippines.

Leptolejeunea subacuta St.

Epiphyllous in evergreen forest, 27/XII/1969, no 4716.

Ryu Kiu, Vietnam, Cambodia, Bonins, Japan, Malaya,
Sumatra, Borneo, Philippines, Botel Tobago.

Leptolejeunea vitrea (Nees) L.

Epiphyllous in evergreen forest, 26/ XII/1969, no 4718.

Sumatra, Java, Borneo, Philippines, Malaya, Moluccas, Laos,
Cambodia, Vietnam.

Leucolejeunea xanthocarpa (L. & L.) Ev.
Epiphyllous in evergreen forest, 26/XII/1969, no 4665.
Pantropical.

Microlejeunea cucullata (Nees) St.

Bark of coniferous trees, 25/XII/1969, no 4724; epiphyllous
in evergreen forest, very common, 27/XII/1969, nos 4699,
4718.

Tropical Asia and Oceania.

Pycnolejeunea excisula St.
Epiphyllous in evergreen forest, 26/XII/1969, no 4665.
Java.

Pycnolejeunea cf gigantea St. .

Epiphyllous in evergreen forest, 26/XII/1969, no 4620.

Species between Pycnolejeunea gigantea St. (with smaller
lobuli) and P. grandistipulata (Gott.) St., South American
taxon with leaves slightly acuminate at the apex.

Pycnolejeunea cf. punctata St.
On bark of coniferous trees, 25/XII/1969, nos 4726, 4846.
Bismarck Archipelago.

Ptychantus striatus (L. & L.) St.

On bark, 28/XII/1969, no 4850.

Japan, Ryu Kiu, Formosa, Ambon, Malesia, Burma, Assam
India, Madagascar, Africa.

Stenolejeunea apiculata (Gott.) Schuster.

Epiphyllous in evergreen forest, 26/XII/1969, nos 4646,
4665, 4682: 27/XII/1969, nos 4702, 4705.

Java, Philippines, Formosa, Malaya, Cambodia, Vietnam,
New Caledonia.

Taxilejeunea compressiuscula (St.) Eifrig
On sunny bark, 28/XII/1969, no 4845.
Johore, Sumatra, Java, Philippines.

Taxilejeunea luteola (St.) Eifrig
On twigs of Podocarpus, 25/X11/1969, no 4607.
Java.

346 Gardens’ Bulletin, Singapore —X XV (1971)

Thysananthus spathulistipus (Reinw. Bl. Nees) St.

On twigs, 26/XII/1969, no 4753.

Southern Africa, Madagascar, India, Ceylon, Malesia,
Vietnam, Cambodia, Laos.

Tuyamaella angulistipa (St.) Hattori

Corticolous on twigs of coniferous trees, 25/XII/1969, no
4612, 4723

Epiphyllous 26/XII/1969, nos 4628, 4678: 27/XII/1969,
no 4716.

Java, Perak, Borneo, Vietnam, Cambodia.

Frullania apiculata Auct. var. goebelii Schiffn.

Epiphyllous in evergreen forest, 27/XII/1969, nos 4685,
4718; corticolous 28/XII/1969, no 4841. -

Ceylon, Vietnam, Malesia, Moluccas.

Frullania cf. cochleata St.
On trunk, 25/XII/1969, no 4734.
New Guinea.

Frullania grandistipula Lindenb.
On twigs, 28/XII/1969, no 4854.
Ceylon, Sumatra, Java.

Frullania meyeniana Lindenb.

On trunk in sunny station, 25/XII/1969, no 4733.

Java, Philippines, Vietnam, Thailand, Cambodia, Laos,
Hawai.

Frullania ternatensis Gott.

Pendulous, on trunks, 25/XII/1969, no 4730, 28/ XIT/ 1969,

no 4818.

Malaya, Ceylon, Sumatra, Java, Borneo, Sulawesi. Ternate,
Philippines, Vietnam, Laos, Cambodia.

New Species

Syrrhopodon fraserianum sp. nov. Fig. 1

Planta parva, corticola, brevis, pallide viridis. Folia usque
3.5-4.0 mm longa, 0.4 mm lata ad partem vaginalem. Cancellinae
1.4 mm longae cum 5 ordinibus cellularum. Cellulis parietum
crassis, 7-8 u in diametro, unipapillosis. Apex folii cum margine
denticulato, innervoque spinosus. Alia desunt.

Small corticolous plant, light green, stem short. Leaves 3.54.0
mm long, 0.4 mm wide near the sheath. Lattices 1.4 mm long,
with 5 rows of cells. Cells of the lamina 7-8 » in diameter, with
thick walls and a papilla on both sides of the leaf. Apex with
some teeth on the margin and spiniferous on the back of the
nerve.

This little species is between Syrrhopodon larminatii Broth &
Par. by the ornament of the cells, and, S. leucophanoides Card.
& P. Vard. and S. calymperoides Card. & P. Vard. by the form
of the leaf.

Tixier — Bryophytes, Fraser’s Hill 347

4mm

4769

Fig. 1. Syrrhopodon fraserianum:
a, Leaves; b, Cancellinae; c, Top of the leaf; d, Cells.

348 Gardens’ Bulletin, Singapore — X XV _ (1971)

Trichosteleum fruticolum sp. nov. Fig. 2

Similis T. subcucullifolio; haec planta ramicola, breviter ramosa.
Folia obtusa, in apice, marginibus recurvis, 1.8-2.0 mm _ longa,
1.2 mm lata. Cellulis alaribus, inflatis, pariete incrassatis, cellulis
laminae 70 » longis, 7 » latis cum papillis crassis, 5 » in diametro.
Apex leviter crenulatus, brevis cum cellulis scutulatis. Cetera
desunt.

In habit similar to T. subcucullifolium, on twigs. Leaves obtuse,
apex tubulate, 1.8-2.0 mm long, 1.2 mm wide. Cells of lamina
thick walled, elongated, 70 » long, 7 » wide. Papillae 5 » in
diameter. Apex lightly crenulated. Margin cells 70 » long and
De wide.

This species is near the following: 7. chaetomitriopsis Dix.
from Thailand, 7. montanum P. Tix. and T. subcucullifolium
Par. & Broth from Vietnam, 7. everettii Dix. from Borneo and,
finally T. brongsermae Zant. of New Guinea.

It is also known from Mount Bokor in Cambodia.

Cololejeunea pahangiana sp. nov. Fig. 3

Planta magna, pallide viridis, affixa substrato. Caules 1-2 cm
longi, 0.06 mm crassi, leviter sinuosi, cum foliis 2.2 mm lati.
Rhizoidea hyalina multa. Folia inter se 0.7 mm dissita. Lobus
‘margine hyalinus cauli non continguus, uno ordine cellularum
lato, adnatus ad caulem, 1.2 mm longus, 0.9 mm latus. Cuticula
cellularum papillosa, parietibus earum incrassatis intermedo.
‘Cellulae hyalinae rectangulares 25-40 x 15-20 uw: marginales
15-20 x 15-20 yw; basales 40-60 x 15-25 yw. Lobulus planus
erectus, margine semi hyalinus, divergens, lobo parallelus, 0.4 mm
Jongus, 0.15 mm latus. Papilla hyalina ad partem proximam dentis
medii. Androecia axiales vel lateralia, 0.6 mm alta, bracteis
2-jugis. Capitula feminea lateralis. Bracteae florales marginatae.
cum lobo 0.8 mm longo, 0.8 mm lato, et lobulo 0.4 mm longo,
0.15 mm lato. apice denticulatus. Perianthium plano-obrotundatum,
0.9 mm altum, 0.8 mm latum, valide rostratum.

Large plant, light green, autoicous applied to the substratum.
‘Stem up to 1-2 cm long, 0.06 mm thick, lightly sinuous, width
including leaves 2.2 mm. Numerous hyaline rhizoids. Leaves 0.7
mm apart. Lobe oval, with a hyaline margin, adnate to the stem,
1.2 mm long, 0.9 mm wide. Leaf cells with papillous cuticle and
intermediary thickenings. Hyaline cells: 25-40 x 15-20 u; cells of
the edge: 15-20 x 15-20 wp; basal cells 40-60 x 15 x 25 u.
Lobule 0.4 mm long, 0.15 mm wide, divergent, erect, plane, parallel
to the lobe; its distal edge has a row of semi hyaline cells.
Male inflorescence lateral or axial, with 2 tiers of fertile bracts.
Female inflorescence lateral, female bracts with a lobe of 0.8
mm long and 0.8 mm wide, lobule 0.4 mm high and 0.15 mm
wide. Perianth plane, almost globoid, 0.9 mm high and 0.8 mm
wide, with a strong beak.

Tixier — Bryophytes, Fraser's Hili 349

Fig. 2. Trichosteleum fruticolum:
I, Leaves; 2, Apex of the leaf; 3, Auricle: 4, Cells of
the margin; 5, Cells of the lamina.

350 Gardens’ Bulletin, Singapore —XXV (1971)

EF olejeunea pal lana:

I, Stem; 2, Leaf; 3, Base of the lobe; 4, Cells of the apex;
5, End of the hyal margin; 6, Lobule; 7, Perianth; 8, Lobule
of bract leaf; 9, Male inflorescence.

Tixier — Bryophytes, Fraser's Hill . 351

Lejeunea (?) pilifera sp. nov. Fig. 4

Planta parva, luteo virens, corticola foliicolaque. Caules usque
ad 5 cm longi, 0.06 mm crassi cum foliis 0.8 mm lati. Folia
inter se 0.4 mm dissita. Lobus rotundatus 0.4 mm longus, 0.3
mm latus; lobulus 100 » x 100 ». Amphigastrium bifidum usque
ad mediam partem. Cellulae lobi lobulique, pariete tenuae,
isodiametricae 30 x 40 wu. Cellulae amphigastrii similes. Rhizoides
ex cellulis marginalibus lobi.

Small plant, yellowish, corticolous or epiphyllous. Stems to
5 cm long 0.06 mm thick, width with leaves 0.8 mm. Leaves
spaced at 0.4 mm; lobe rounded, 0.4 mm long, 0.3 mm wide.
Under-leaves divided to the middle of the organ. Cells of lobe,
lobule, and underleaf isodiametric with thin walls. Rhizoids
Springing up of the cells in the leaf margin. The prescence of
such organs is not reported for the genus Lejeunea in the paper
of W. Degenkolbe (Britorgane bei beblatterten Lebermoose. —
Ann .Bryol., 1937, X, pp. 43-96).

Conclusions

1. Interest of the collections

The object is to find out if these gatherings bring an interesting
contribution to the knowledge of Bryology in Malaya. In my
lists most of the Liverworts are practically new for the region
and for Mosses 31 species are now for Fraser’s Hill, 25 for
Malaya and 2 for Science.

2. Biogeographical connections

For the mosses, the community floristic coefficients between
Fraser’s Hill and the different countries are as follows:

Malesia (*) cre oo
Indochina es 10
Ceylon . 2 3U
New Guinea n 20
Northern India ss 20
Pacific Islands - 14
Southern India, New Caledonia = 10
China, Micronesia, Formose, Japan PA under 10

We can see that the affinities of our bryological population go
to the rest of Malesia, Indochina and Ceylon (between 90 and 30).
From 20 to 10 we have India, New Guinea and New Caledonia.
Few connections appear with the Northern Bryofloras. Fraser’s
Hill is found to have a good many Javanese species (Tjibodas)
hitherto unknown for Malaya. Thus these collections establish a
bridge between the Southern Indochina bryoflora and the Javanese.
Yet, the Malayan Peninsula remains a “center of distribution”
for species of the Sematophyllaceae and particularly of the genus

(*) Malesia as in Flora Malesiana.

— XXV (1971)

332

Tixier — Bryophytes, Fraser’s Hill 353

Acroporium, Dixon, in 1926, gave 35 species of Acroporium for
Malaya, let us remind ourselves that one can find only 12 species
in Thailand and 9 in New Caledonia, for example.

Literature Cited

Benedix, E. H. 1953. Indomalayische Cololejeuneen. Feddes
Rep. 134: 1-88.

Dixon, H. N. 1924. New Species of Mosses from Malay Peninsula.
Bull. Torrey Bot. Club: 51, p. 225-229,

— 1926. A List of Mosses of the Malay Peninsula.
Gard, Bull. Str. Settle. 4: 1-46.

Froelich J. 1955. Die von Prof. Dr. V. Schiffner i, den Jahren 1893-
94 in Ceylon, Penang, Singapore, Sumatra und Java gesammelten
Laub und Torfmoose. Ann. Nat. Mus. Wien, 59; pp. 66-116.

Johnson A, 1959. The Genus Sphagnum in Malaysia. Gard. Bull.
Singapore, 17, 2: p 312-324.

Merulioid Fungi in Malaysia

by E. J. H. CORNER
Botany School, Cambridge, U.K.

Meiorganum (1 sp.), Merulius (8 sp.) and Phaeophlebia (1 sp.) are
recognised. New species are Merulius agathidis, M. irpicoides, M.
luridochraceus, M. purpurascens and M. rubrotremellosus. Serpula
and Gyrodontium are placed under Merulius. A list of specific
names in Merulius is supplied.

In preparing an account of Malaysian boleti I met the problem
of Meiorganum. Heim considered it to represent a new family
Meiorganaceae intermediate between Boletaceae and Polyporaceae.
I have known the fungus for many years, for it is not uncommon
in Southeast Asia, and I have always regarded it as a species of
Merulius. Hence I have been lead to investigate the problem.

In recent years Merulius has been whittled down to an insigni-
ficant group of species about M. tremellosus with or without the
small alliance of M. corium (Donk 1964, Parmasto 1967). The
brown-spored species are separated as Serpula in Coniophoraceae
(Cooke 1957, Donk 1964), and the remainder has been submerged
in Corticiaceae. The evidence is not convincing. There are inter-
mediates between Serpula and Merulius and, besides Meiorganum,
there are other merulioid species in Southeast Asia with very much
more complicated construction than occurs in other Coniophora-
ceae and Corticiaceae. The merulioid character is the continuous
unthickened hymenium with gelatinous subhymenium, raised on
folds, pores or spines the trama of which is also gelatinous or
muciaginous. That merulioid fungi may degenerate into corticioid
states in the same way as clavarioid, stereoid and hydnoid fungi
is clear. This state is generally assumed to be primitive because
it is simple but to me the evidence points to the derivation of the
corticioid by simplification (Corner 1968, 1970). The problem,
therefore, is from what ancestry the more complicated merulioid
fungi may have been derived. It is likely that yet others of more
demonstrative nature may be found in the tropics, and the points
needs development.

Merulius and Serpula

The main distinction is white spores for Merulius s. str. (Corti-
ciaceae) and brown spores for Serpula (Coniophoraceae). Other
details lie in the construction of the hymenium and subhymenium
and in the cyanophilous character of the spores in cotton blue.
The distinction in spore-colour is so obvious in many species that
the generic separation has become customary. Yet, some species
have spores that are described as white or yellowish, e.g. M.
byssoideus, MM. crassus, M. fuscescens, M. molluscus, M. mon-
tanus and M. pseudomolluscus. Of these M. byssoideus has been

356 Gardens’ Bulletin, Singapore — XXV (1971)

considered to be very near to M. ftignicolus with ferruginous
spores, if not identical (Ginns 1968). Allied with M. byssoideus
is the species described here as M. luridochraceus and its spores,
under the microscope, vary from very pale yellowish to pale
brownish ochraceous, and they are cyanophilous as in Serpula.
M. molluscus has been transferred to Serpula independently by
Karsten and Donk (1964 p. 209), though the spores have been
described as white in the mass (Bourdot et Galzin 1927). Leucogy-
rophana Pouz. has been made for this species because of its
white or yellowish cyanophilous spores, turning brown in Melzer’s
iodine; four other species have been added by Parmasto (1967).
Donk reduces Leucogyrophana to Serpula and Ginns finds the
amyloid reaction uncertain (as I find for M. agathidis p. 365).
The hymenium and subhymenium in all three genera are exactly
alike and in strong contrast with the dry and often loose hymenium
with large projecting basidia of Coniophora (Lentz 1957). The
distinction in spore-colour between Merulius s. str. and Serpula
breaks down; they have the same general construction; only the
uncertain significance of cyanophily remains. I find that the white
spores of M. rufus, close to M. tremellosus, are slightly cyanophi-
tous. Hence I prefer to use Merulius in the wide Friesian sense
to include Serpula, and this fulfils the cogent need, often expressed
by plant-pathologists, to preserve the name Merulius lacrimans as
one of the most widely employed in practice (Harmsen 1956,
i960). There is an excessive tendency now to split up the genera
of basidiomycetes on characters the universal applicability of which
has not been established from world monographs. From the point
of view of the general structure of the fruit-body I can see no
difference between Merulius and Serpula; that they are related to
Coniophora is conjecture. Specialists, of course, employ subgeneric
categories as if they were genera, but botany requires the main
ideas. It seems to me that, as with clavarioid, thelephoroid, boletoid
and polyporoid fungi, the white spore is derived from the coloured
which in the case of Mertilius is not necessarily ferruginous; it
may be olivaceous or pinkish (Meiorganum).

Gyrodontium

This genus was made for the common brown-spored Merulius
with hydnoid spines, previously called Hydnum henningsii Bres.,
but with H. versicolor B. et Br. as an earlier name, as shown by
Maas-Geesteranus. Instead of folds or pores covered by the
hymenium, this species has long spines which, if occasionally con-
nate, are never poroid. It certainly proves that Serpula, at least,
is much more complicated than Coniophoraceae. The hyphae in
the spines are longitudinal with mucilaginous walls, as in the gills
of Paxillus but not inflated, and this direction indicates a positively
geotropic growth of the spine as a special emergence of the meru-
lioid hymenium. However, the brown-spored M. irpicoides (p. 367)
has spines or irpiciform plates which are free or shortly connected
at the base, and M. (Serpula) pinastri has pores which may
become strongly irpiciform (Cooke 1957). Thus this hydnoid dis-
tinction fails and I treat Gyrodontium as part of Merulius s.l.

Corner — Merulioid F ungi in Malaysia 357

Meiorganum and Boletaceae

Meiorganum has the appearance of an upside-down boletus.
The fruit-bodies grow from tree-trunks or branches, living or
dead, and develop the pileus at the end of a shortly descending,
but expanding stem; the pileus then flattens horizontally into a
circular or spathulate form and develops tubes on the underside,
The hyphae inflate and the tubes have a phylloporoid trama of
Strictly longitudinal, mucilaginous and scarcely inflated hyphae.
Heim compared the one known species M. neocaledonicum with
Gyrodon merulioides which, though erect, may be lignicolous
with lateral pilei, but it has a bilateral boletoid trama, cyanes-
cent flesh, ochraceous brown spores and, evidently, a very differ-
ent struct to the surface of the pileus and stem (Singer 1945); to
this one may add the African G. intermedius (Singer and Grinling
1967). On the other hand, the spores, hymenium, minutely lacunose
flesh with short branching fascicles of hyphae, and the colouration
of Meiorganum are typical of brown-spored Merulius. If Serpula is
recognised, Meiorganum differs merely in the shape of the fruit-
body, which is bridged by the pleuropodal M. agathidis, and in
the colour of the spore-print which is dull vinaceous pink to fawn
brown.

The pores of Meiorganum develop from the centre of the young
discoid hymenium as fine reticulate wrinkles. Dried specimens
collected in this state, which is fertile, pass exactly for blackened
Merulius. Heim emphasized the fact that the tubes of Meiorganum
are not separable from the flesh; they have a distinct mucilaginous
construction, as in phylloporoid boleti, but the finely lacunose
flesh breaks away with them and is the cause of the apparent
difference.

I retain Meiorganum distinct from Merulius chiefly for con-
venience because it appears to be exactly intermediate between
Gyrodon or Paxillus (as P. panuoides) and Meruliaceae and shows
that the ancestry of this family must be sought in the origin of
Boletaceae, not in Coniophoraceae or Corticiaceae. This is borne
out by the spore-basidium relation d = 0.48 w, as in many boleti.
This relation reveals the pink-brown spore as an endospore in a
mucilage sheath, developed through compression of the spore-
circles (Corner 1971).

Meiorganum is the most elaborate merulioid fungus yet found.
Its boletaceous affinity is clear. It shows how the resupinate meru-
lioid fungi could have been derived by loss of form-factors in
the development of the tubes and in loss of colour of the spore;
if a young dorsifixed Meiorganum became resupinate with rudi-
mentary tubes it would be Serpula. Thus it shows how the boletoid
fruit-body, primitively terricolous, has become adapted to the
lignicolous habitat and, in using the lignin as support, economised
in construction. So many advanced merulioid fungi thrive all over

358 Gardens’ Bulletin, Singapore —XXV (1971)

the world. On the upgrade theory from resupinate forms, Meior-
ganum must be the peak of evolution in Coniophoraceae, yet it is
about the most uncommon. The upgrade theory has no biological
support and, lacking this, will die of its own sterility.

The problem shifts to what may be the difference between
Meruliaceae and Boletaceae. There is no difference in basidia and
spores because Boletus xylophilus Petch, for instance, has as small
structures (basidia 18-24 x 6~-7.5u, spores 4.5-6.7 x 3.3-4.3y). It
does not lie in the fertile edge to the pore, for many boleti have
this character as well as Paxillus. It does not lie in the tramal
structure of the tubes, the inflation of the hyphae or in the
incurved margin of the pileus, for all these boletoid points occur
in Meiorganum. It does not lie in the structure of the surface of
the pileus because this is most variable in Boletus and Paxi!lus.
It lies simply in the loss of the stem and the resupinate tendency
which substitutes this; it lies, that is, in those major characters of
the fruit-body over which the present attitude is to gloss.

Merulius agathidis and Hygrophoropsis

M. agathidis has sessile imbricate pilei with dichotomous meru-
lioid gills the sides of which become strongly ribbed, making the
gills appear phylloporoid, though they are not poroid-reticulate.
It resembles Paxillus panuoides Fr. but the gill-trama of M.
agathidis is composed of uninflated and laxly interwoven hyphae.
In this respect it resembles Hygrophoropsis flabelliformis (B. et C.)
Corner, which has white and slightly cyanophilous spores and
dichotomous gills without ribbing. It is difficult to exclude H.
flabelliformis from Hygrophoropsis and it is as difficult to exclude
it from Merulius. I have discovered a second collection of mine
from Singapore with pilei up to 6 cm wide, otherwise identical
with H. flabellijormis, and I had placed it under Merulius. lf
tubes, spines and branching folds are admitted into Merulius
because there are intermediates, then radial direction of the
branching folds will give the dichotomous merulioid gill with its
mucilaginous hymenium and subhymenium, as in Hygrophoropsis,
the construction of which is as close to that of Merulius as to
Paxillus with which it is placed in classification; and none of these
has close affinity with Coniophora. Indeed, they have not the
hymenium of Coniophora which comes like a red herring across
the trail of Merulius. Loss of gills would result in Podoserpula
Reid (Kew Bull. 1963, 437), which offers merely a stem as a
distinction from Merulius, though it seemed to me that it had a
thickening hymenium (Corner 1966, 6. 79, as Cantharellus pusio
in error). Again there is little affinity with Coniophora though
Podoserpula is placed in Coniophoraceae by Reid and Donk. If
Hygrophoropsis and Podoserpula belong with Merulius s. str. with
white spores, then there is evidence for the ancestral diversifica-
tion of Merulius s.1. into species with coloured and colourless

Corner -— Merulioid Fungi in Malaysia 359

spores, and this supposition supports the idea of the family
Meruliaceae, divergent from Boletaceae and convergent in sim-
plification with Coniophoraceae and Corticiaceae.

Phaeophlebia

This genus differs from white-spored Merulius in the absence of
a compact, exposed layer of basidia. It is based on the single,
almost cosmopolitan species first described as Merulius strigoso-
zonatus. In other respects it is typical Merulius with the rufous
orange, rugulose and mucilaginous hymenium, but the basidia are
separated by and immersed among very narrow lobulate hyphal
ends which build an epihymenium through which the maturing
basidia project; it is the kind of hymenium that occurs in Acan-
thophysium (Cunningham 1963) and has been called a catahy-
menium (Lemke 1964). Cooke described the spores as yellowish
to pale brown but, as observed by Cunningham and myself, the
mature spores are hyaline and white in the mass; possibly the
action of potash on dried specimens, which give then a brown
solution, colours the spores. Cooke described, also, gloeocystidia
which neither Cunningham nor I have observed; I suspect that
they are the immature and immersed basidia.

Cunningham placed the species in Stereum where it is even
more out of place than in Merulius. He seems not to have appre-
ciated the mucilaginous structure of the flesh and hymenium; his
illustration is rather diagrammatic and shows closely septate,
branched but not lobulate, hyphal ends in the hymenium.

Donk has followed Talbot in considering Phaeophlebia as
synonymous with Punctularia. They appear to have the same
general construction but the very small and discrete fructifications
of Punctularia seem different from the effuso-reflexed masses of
Phaeophlebia with imbricate and tomentose pilei. Moreover,
neither Cunningham nor I have seen the dark brown swollen
dendrophyses or the fissured hymenium which Talbot described
for Punctularia. Unless a fungus is discovered to bridge these
differences, I prefer to distinguish the two genera. Nevertheless,
if Acanthophysium can be reduced to Aleurodiscus and retained
in Corticiaceae (Donk 1964), then Phaeophlebia goes back to
Merulius.

Byssomerulius Parm. and Meruliopsis Bond

The white-spored species of Merulius without clamps in the
hyphae of the fruit-body have been separated as Byssomerulius,
based on M. corium Fr. (Parmasto 1967). It is difficult to see how
it differs from Meruliopsis which has been reduced to Merulius
s. str. (Donk 1964, p. 262), with the name M. taxicolus (Pers.)
Duby for the type of Meruliopsis. I agree with Donk, and cite M.

360 Gardens’ Bulletin, Singapore —XXV (1971)

purpurascens (p. 370) as a species intermediate between Bysso-
merulius and Meruliopsis. 1 have also collected in Venezuela a
pileate ally of M. corium with short spines or papillae, instead of
pores, and slightly cyanophilous spores; it has probably been
described as a Hydnum, but it shows how much there is still to
learn about merulioid fungi. Donk has shown, too, the uncertainty
that may attend the definition on the absence of clamps. From a
tropical viewpoint I regard these genera as merely series in the
complex of Merulius.

Byssomerulius raises, also, the problems of Caloporus and
Gloeoporus as transfers from Polyporaceae (Donk 1960, p. 192;
1962, p. 227); many other tropical species assigned to Poria
become involved and I am unable to comment on them at present.

The merulioid hymenium

Thin resupinate species of Merulius, similar to Corticium, have
such slight hymenial folds or reticulations that they suggest mere
physical buckling under tangential stress. This effect hardly suits
Phaeophlebia the excessive tangential growth of which throws the
margin of the pileus into undulations with little or no effect on
the underlying hymenium. It does not explain the long and narrow
tubes (up to 6 x 0.5 mm) in M. eurocephalus with longitudinal
hyphae in the dissepiments, or the phylloporoid trama of Meior-
ganum developing beneath merulioid wrinkles, or the long and
discrete spines of M. versicolor with their longitudinal hyphae.
These outgrowths are organised in a marginal pore-field (Corner
1932). They differ from the polyporoid in being covered from the
first by the fertile hymenium. It seems that they develop by con-
tinual sympodial outgrowth of hyphae into the hymenium along
the edges of the pores or at the tips of the spines; thus they have
longitudinal tramal hyphae. Generalised intercalation of basidia
may stretch the internal hyphae but not direct them because inter-
calation alone merely pulls hyphae apart as in the cantharelloid
gill-fold (Corner 1966). Lines of feeble outgrowth, dichotomising
as they radiate with marginal growth, produce the gill-folds of
M. agathidis, but massive intercalary growth on the sides of the
gills disaligns the internal hyphae. The exact method of contruc-
tion remains to be discovered but there must be some lamellate,
poroid or hydnoid motivation that raises the merulioid fungi far
above the corticioid level. Apart from Meiorganum, Merulius
agathidis, M. eurocephalus and M. versicolor appear to me as
descendants of the stock of Paxillus and Boletus which, in losing
agaricoid form-factors, have become aphyllophoralean with unin-
flating hyphae.

This view of the descent of Meruliaceae from the boletoid stock
differs widely from the current. I trust that mycologists who have
so kindly sent me their papers on resupinate fungi will pardon
the disagreement which arises from the effort to assimilate the
merulioid fungi of the tropics.

Corner — Merulioid Fungi in Malaysia 361

Key to the merulioid fungi in Malaysia

1. Spores coloured, pinkish, olivaceous or yellow brown.
2. Hymenium with spines or irpicoid plates -7mm long.

3. Forming large thick brackets with or without a resupinate part;
flesh 3-14mm thick, spongy-floccose. Sp. 3.5-4.7 x 2.5-7, fuscous
olivaceous. Cystidia none. Hyphae without clamps .....................

M. versicolor p. 373

3. Resupinate on dead Poria; flesh 0.3mm thick. Sp. 44.7 x 3-3.5 y.
Cheilocystidia fusiform, thin-walled. Hyphae with clamps .........
M. irpicoides p. 367

2. Hymenium lamellate or poroid.

4. Hymenium lamellate with ridges on the sides of the dichotomous,
yellow to orange gills —3mm wide, spuriously poroid. Pilei —7cm
radius, imbricate, tomentose. Flesh 3—6mm thick. Sp. 3.3-4.2 x
\.J-2Zy. Hyphae clamped. On wood of Agathis ..............:.secneses

M. agathidis p. 365

4. Hymenium poroid or gyroso-plicate.

5. Pendent, subungulate, then applanate and dorsifixed, —20cm
wide. All parts turning pink or vinaceous, then fuliginous on
bruising. Tubes —9mm long; pores 1-3mm wide, dull orange
yellow, then fuscous or cinnamon. Flesh 3—5cm thick, minutely
lacunose. Smell strong. Sp. 3.7-4.3. x 2.7-3yu, pinkish to
vinaceous brown. Hyphae clamped, inflated ........................

Meiorganum p. 362

5. Not so. Pores yellow to ferruginous, -0.S5mm wide.

6. Effuso-reflexed with pilei —18cm radius. Flesh 5—20mm thick.
Tubes —6mm long. Sp. 4--5.5 x 3-4y. ferruginous ochraceous.
Hyphae clamped; with many fine fibre-hyphae ..................

M., eurocephalus p. 366

6. Resupinate, very thin. Tubes -2mm long. Sp. 3.5-4.5 x
2-2.7u, pale ochraceous s.m. Hyphae clamped; without fibre-
Se. i ee ns a ne M. luridochraceus p. 369

1. Spores white.

7. Effuso-reflexed with imbricate tomentose pilei -4cm radius, rufous
sepia to purplish brown, thin, pliant. Hymenium rugulose, rufous
orange, then umber to blackish. Sp. 5-7 x 3-4y. Basidia immersed,
then protruding singly through a layer of 1-2 lobulate hyphae ......

Phaeophlebia p. 374

7. Not so. Hymenium usually poroid. Basidia in a compact superficial
layer.

8. Hyphae without clamps in the fruit-body. Resupinate with narrow
flange-like pilei. Sp. 3-3.5 x 1.2-1.5y ....... M. purpurascens p. 370

8. Hyphae with clamps.

9. Floccoso-subcoriaceous, thin, resupinate. Hymenium lurid yellow
to cinnamon ochre. Sp 3.3-4.2 x 2.3-2.74, pale ochraceous ......
M_ luridochraceaus p. 369

9. Waxy subcoriaceous, subgelatinous. Sp. 3.5-5 x 2-2.5u.

10. Effuso-reflexed, wholly rose-red, -15mm_ radius. Without
Shee MA beie 8.532. aiveegei.-. M. rubrotremellosus P. 371
10. Resupinate. Hymenium ochraceous drab then brownish,

pinkish or pale orange. Gloeocystidia frequent, clavate ......
M. rufus p. 372

362 Gardens’ Bu_letin, Singapore — XXV (1971)

Meiorganum

Heim, Rev. mycol. Paris 31 (1966) 157 (diagnosis); id. 30 (1965)
307 (illustrated).

Fruit-bodies lignicolous, substipitate, dorsifixed, descending,
conical then ungulate to applanate, spathulate to flabelliform or
eccentric, fleshy, putrescent; margin incurved. Tubes more or less
inseparable, becoming compound with wide angular merulioid
pores, waxy mucilaginous with phylloporoid trama, developing
centrifugally from a small smooth hymenial disc at the apex of
the primordium; pore-edges fertile; hymenium merulioid, not
thickening. Flesh finely lacunose. Spores dull vinaceous pink to
fawn brown, small, smooth. Cystidia present. Hyphae monomitic,
thin-walled, inflating, clamped, minutely fasciculate-reticulate in
the flesh. Sp. 1.

M. neocaledonicum Heim l.c.

Fig. 1, Plate I

Sessile or substipiate, descending, subungulate, usually eccentric,
then applanate with the horizontal limb -13 cm radius and
21 cm wide, with incurved margin and poroid hymenium on the
underside, finally undulate or lobed, solitary to subcaespitose and
imbricate, putrescent, all parts turning dingy pink or vinaceous,
then fuliginous, on cutting or bruising, drying blackish. Upperside
sterile, uniformly pallid fawn bistre to light fulvous fawn or rufous
fawn, smooth or matt, somewhat appressedly inoderm-fibrillose
towards the margin, somewhat ruguiose when old, not zoned or
sulcate, ferruginous fawn villous at the basal attachment; margin
incurved, entire, pallid white to pallid yellowish, pruinoso-veluti-
nate. Tubes -9 mm long, waxy subgelatinous, dingy fawn orange
or fuscous orange; pores becoming 1-3 mm wide, at first small
and developing in more or less radial rows at the margin, angular,
compound, with uneven entire edge, dull orange yellow, becoming
fuscous orange or cinnamon fawn. Flesh 3—5 cm thick in the
centre, spongy floccose, more or less finely lacunose but often
minutely (especia‘ly near the surface and in the subhymenium over
the tubes), pallid dingy white to dingy buff or pallid vinaceous.
Smell strong of gas-tar and fenugreek, resembling Hymenogaster.

On stumps and fallen rotting trunks, also on living trees, in the
forest. Singapore, Sumatra, Borneo, New Caledonia.

Spores 3.7-4.3. x 2.7-3 p, dingy pinkish to dull vinaceous drab
or fawn brown in the mass, smooth, broadly ellipsoid, with a thin
mucilage sheath, with a small gutta 1.5 » wide, cinnamon brown
under the microscope, darkening to fuscous ferruginous in potash,
the apiculus 0.2 »; possibly slightly cyanophilous in cotton blue.
Basidia 18-28 x 5.5-6.5 », clamped; sterigmata 4, 2.5-3.5 » long.
Cystidia 35-75 x 5-7 u, narrowly fusoid, the obtuse to subacute
apex 2-4 » wide, thin-walled, colourless, projecting —-40 pu, gen-
erally rather thickly granular encrusted in the immersed part,

Corner — Merulioid Fungi in Malaysia 363

Fig. 1. Meiorganum neocaledonicum. Developing fruit-bodies in Ls.,
showing the lacunar flesh, x 1; spores, x 1000; part of the
hymenium, x 500.

364 Gardens’ Bulletin, Singapore — X XV _ (1971)

internally somewhat oleaginous, abundant, not forming a sterile
edge. Hyphae 2.5-15 (—23) » wide, clamped, the cells 60-220 »
long in the flesh, the walls thin or slightly thickened (subdiffluent
in potash), more or less concrescent in short anastomosing micro-
fibrils, branching at a wide angle, often with oleiferous hyphae
—-11 » wide; in the tube-trama 2-6 » wide, with thickened mucila-
ginous walls, strictly longitudinal, very compact, clamped; sub-
hymenium 15-20 » thick, compact, with hyphae 2-3 » wide. Surface
of the pileus with more or less appressed or slightly divergent
hyphae 3-5 p» wide, with obtuse ends, thin-walled, the cells 13-38 »
long, not organised into a palisade. Colour caused by orange yellow
granules on the outside of the hyphae, especially in the hymenium
and subhymenium, the tissue turning dark brown in potash and
giving a yellow solution; in alcohol-formalin, giving out a dingy
purplish colour.

Coll. Singapore, Bukit Timah, Corner s.n. 24 Jan. 1936, 19 Nov.
1939.—Sumatra, Brastagi, 1500 m alt., Corner s.n. Sept. 1931.—
Borneo, Kinabalu, east ridge, 1300 m alt., RSNB 592, 14 June
1961; RSNB 8228, Mesilau, 1600 m alt., 10 April 1964.

Development gymnocarpic, beginning with conical primordia
growing directly or obliquely downwards, becoming cylindric or
cigar-shaped, the obtuse apex then forming a minute hymenial
disc in its centre and widening by marginal growth into the centric
or eccentric pileus; hymenium fertile almost from the first, smooth,
then developing fine reticulate wrinkles from the centre outwards,
the wrinkles gradually deepening by marginal. and intercalary
growth and widening into the tubes.

This description is taken from Malaysian material. Except for
the presence of clamps, there are merely slight differences from
that given by Heim. The New Caledonian fungus grew on wood
of Agathis, the Malaysian on dicotyledonous wood, even from the
trunks (? dead heartwood) of living trees. The New Caledonian
specimens were slightly blackish scaly, but I think this was only
the consequence of bruising as all the tissue is rufescent-nigrescent.
There are slight differences in colour which is, in any case, hard
to describe, and I found that the Bornean specimens were more
richly coloured than those from Singapore. The cystidia in the
Malaysian specimens were encrusted in their immersed parts, but
the incrustation dissolves in potash. The smell of the New Cale-
donian fungus was said to be ‘fine odeur suave’, whereas that of
the Malaysian fungi was to me exceptionally noiseme. With
regard to the clamps, present in all the Malaysian material, Heim
reported their absence from the New Caledonian, but I think that
an error has crept in because in the centre of his. Fig. 11 (Rev.
mycol. 30, p. 316) a clamp is shown on an uninflated hypha.
and the oblique septa that he emphasized are the septa of inflated
clamps.

It is possible that this striking fungus reached Lloyd and might
have been described by him, though I have not detected it in his
publications.

aii gill if
coeteurd

mane im errs

Plate 1. Meiorganum neocaledonicum. Fruit-bodies, north Borneo, mt. Kinabalu.

Plate 2. Merulius agathidis. Fruit-bodies, north Borneo, mt. Kinabalu.

Corner — Merulioid Fungi in Malaysia 365

Merulius

Fr., Syst. Myc. 1 (1821) 326.—Boninohydnum Ito et Imai, Trans.
Sapporo nat. Hist. Soc. 16 (1940) 127—Byssomerulius Parmasto,
Eesti NSV Tead. Akad. Toim. Biol. 16 (1967) 383.—Gyrodontium
Pat., Ess. Tax. Hym. (1900) 117; Cooke, Gen. Homobas. (1953)
42; Donk, Taxon 5 (1956) 79.—Gyrophora Pat.. Hym. Eur. (1887)
143.—Gyrophana Pat., Cat. rais. Pl. cell. Tunis (1897) 53.—
Leucogyrophana Pouz., Ceska Mykol. 12 (1958) 32; Parmasto,
Consp. Syst. Corticiacearum (1968) 77.—Serpula S. F. Gray, Nat.
Arr. Brit. Pl. 1 (1821) 637; Cooke, Mycologia 49 (1957) 197.—
Trabecularia Bonord., Bot. Ztg. 15 (1857) 211.—Xylomizon Pers.,
Mycol. europ. 2 (1825) 26.—Xylophagus Murr., Torreya 3
(1903) 7.

M. agathidis sp. nov. Plate 2.

Pilei —7cm applanati sessiles horizontales caespitosi imbricati, spathulati
dein suborbiculares, aetate lobato-sinuati, villoso-tomentosi, flavidi dein
cinnamomeo-olivacei, basim versus aurantio-brunnei; margine incurvato
villosulo. Lamellae -3.5mm_ latae, confertae dichotomae rugosae (ut
videtur poroideae sed non reticulatae), ceraceo-mucilaginosae obtusae
integrae, luride flavae dein aurantiaceae, aetate fusco-cinnamomeae. Caro
3-6mm crassa, floccosa, mollis aquosa, fusco-brunneo-ochracea, tomento
—lmm crasso. Odor fortis acidus. Sporae 3.3-4.2 x 1.7—2,,, pallide ochraceae
leves. Hyphae fibulatae. Ad lignum Agathidis in silva. Borneo, Malaya
(typus RSNB 8123, CGE).

Pilei -7 cm radius, sessile, applanate, horizontal, imbricate,
forming masses -12 x 10 cm from a common trunk, semicircular,
finally lobed and undulate, villous-tomentose, pale chrome yellow
then cinnamon olivaceous, at the base dull orange brown; margin
incurved, finely vilious. Gills -3.5 mm wide, radiating, rather
crowded, repeatedly dichotomous, developing vertical ridges on
the sides and appearing subporoid (but not reticulate), rather waxy
mucilaginous, the edge obtuse and entire, bright chrome yellow
then bright orange, finally clouded fuscous cinnamon. Flesh 3-6
mm thick, softly floccose, watery, not gelatinous, fuscous brownish
ochraceous, with a thick tomentum —1 mm. Smell sour, strong,
rather fruity on drying.

On dead trunks of Agathis in the forest. Malaya, Borneo.

Spores 3.3-4.2 x 1.7—2 », pale ochraceous under the microscope,
slightly darker in potash, smooth, subcylindric, straight with obtuse
ends or suballantoid, the slightly thickened wall cyanophilous in
cotton blue and often, but not in all cases, red-brown in Melzer’s
iodine. Basidia 15-17 x 4 wu; sterigmata 4. Cystidia none; gill-edge
fertile. Hyphae 2-7 (-10) » wide, clamped, scarcely inflated,
radiating and much interwoven, branched at a wide angle, the
walls thin or slightly thickened (0.5-1.5 » and subdiffluent in
potash); 2-3.5 « wide in the gill-trama, laxly interwoven; 2-3.5 yu
wide in the tomentum on the pileus, projecting irregularly. Colour
evidently caused by a fine granular excretion in scattered patches
on the hyphal walls, more or less soluble in potash.

Coll. Malaya, Negri Sembilan, Gunong Angsi, 700 m_ alt.
Corner s.n. 3 July 1930.—Borneo. Kinabalu, Mesilau, 1600 m
alt., 6 April 1964, RSNB 8123.

366 Gardens’ Bulletin, Singapore — X XV (1971)

This is one of the few basidiomycetes that I have found on
dead wood of Agathis. It recalls the north temperate Paxillus
panuoides of coniferous wood and the tropical Hygrophoropsis
flabelliformis. It is probably not uncommon and opportunity should
be taken to obtain a spore-print. Old specimens of the Malayan
collection had the older parts of the gill puberulous with excurrent
cystidium-like hyphae -80 x 3-5 un, aseptate, slightly tapering,
with thin brownish walls; it seemed that they belonged to some
mould but it was not possible to trace them.

M. eurocephalus (B. et Br.) Petch, Ann. r. bot. Gdn Peradeniya 4
(1910) 408.

Polyporus eurocephalus B. et Br., J. Linn Soc. Bot. 14 (1875)
48.—Serpula eurocephala (B. et Br.) W. B. Cooke, Mycologia 49
(1957) 212 (with full synonymy); Cunningham, Thelephoraceae
of Australia and New Zealand (1963) 329.—M. pseudolacrymans
P. Henn. (1901).—Gyrophana pseudolacrymans (P. Henn.) S. C.
Teng, Fungi of China (1964) 761.—M. insignis Wakefield, Kew
Bull. (1917) 107—M. consimilis Lloyd, Myc. Writ. 7 (1922)
1122; Ginns, Mycologia 61 (1969) 363.

Sessile, applanate, often imbricate, effuso-reflexed varying resu-
pinate, spreading laterally - 40 cm in extent, the pileate brackets
-18 cm radius and 32 cm wide, slightly convex; upperside
appressedly cottony fibrillose, nearly smooth, dry, soft, white or
pale yellowish white, becoming golden ferruginous from the spores;
margin entire, thick. obtuse, becoming subacute, white, the resu-
pinate part with white villous-tomentose border; attached with
white or yellowish mycelial strands. Hymenium polyporoid; tubes
— 6 mm long, often of unequal length, soft, waxy, subdeliquescent,
separable from the flesh. yellow then lurid ferruginous yellow,
often tinged greenish; pores 0.2-0.5 mm wide, round or elongate,
often 2-3 confluent, uneven with obtuse waxy edges, often toothed
in the ascending parts of the fruit-body, concolorous. Flesh 5—20
mim thick at the base of the pilei, 2-10 mm at 5 mm from the
margin, spongy, soft floccose, dry, white or pale yellowish. Smell
slight, sour.

On dead fallen and standing trunks of trees, bamboos and
palms, also on dead roots, in the forest. Pantropical; frequent in
Malaya and Singapore, lowland.

Spores 4-5.5 x3-3.7 (-4.2) pw, bright ferruginous ochraceous
in the mass, broadly ellipsoid to ovoid or even subglobose, smooth,
with slightly thickened walls and thin mucilaginous sheath (0.3 1),
aguttate, contents oleaginous. Basidia 30-40 x 6-8 », clamped:
sterigmata 4, 3-6 » long. Cystidia none. Hymenium not thickening,
continuous over the pore-edges; subhymenium with mucilaginous
hyphae 1.5-2.5 » wide. Hyphae 2—6.5 » wide, thin-walled, clamped,
often branched from the clamp, rather loosely interwoven without
conspicuous radiating hyphae (except at the margin), colourless,
not encrusted, accompanied by copious slightly thick-walled olea-
ginous hyphae 1-3 mm long or more, —6 mu wide, occasionally

Corner — Merulioid Fungi in Malaysia 367

branched, apparently aseptate, tapering to long filiform tips. 0.5—-1
wide, permeating the flesh in all directions as elastic fibrils, also in
the tube-trama, and very abundant in the mycelium and rhizo-
morphs; hyphae in the tube-trama 2-4 » wide, more or less longi-
tudinal with mucilaginous walls.

Coll. Singapore, Gardens Jungle, Reservoir Jungle.—Malaya,
Pahang, Tembeling, Kuala Tekai.—Borneo, Brunei.

This is a rather common, striking and omnivorous fungus. The
Malayan fungus was described as M. insignis and it may be a
variety. Petch said for the Ceylon fungus that the pileus turned
red or purple when bruised or cut, which I have not noticed in
the Malayan; he also gave the spores as 5-8 x 3-4 p», which is
larger than the Malayan.Cooke’s description appears to have been
drawn from dried specimens and he cites collections from Malaya,
but he seems to have overlooked with others the very abundant
oleiferous fibre-hyphae which are readily seen as long fibrils on
teasing out the tissue of the pileus. These hyphae are the basis of
the dimitic character assigned to the tissue by Ginns. The exact
nature of these hyphae, similar to the fibre-hyphae of M. lacry-
mans, needs enquiry from young living primordia.

M. irpicoides sp. nov. Fig. 2

Resupinatus separabilis, subtus tomentosus, margine flavido-albo villoso;
contextu 0.2-0.3mm crasso, flavido-albo. Hymenium irpicoideum, haud
poroideum, ceraceo-subgelatinosum, luride ochraceo-flavum dein brunneo-
aurantiacum; spinulis -7 x 0.5—1.5mm, subcylindricis obtusis liberis v.
plus minus connexis, saepe applanatis, longitudinaliter rugosis. Sporae
4-47 x 3-3.5y, brunneo-ochraceae s.m., leves, Hyphae fibulatae. Ad
receptaculum Poriae emortuum. Borneo (typus RSNB 84/8, Kinabalu,
Mesilau, 1650m alt., 22 April 1964, CGE).

Resupinate with thin flesh 0.2-0.3 mm thick, tomentose on
the underside, separable, yellowish white, margin villous. Spines
—~7 x 0.5-1.5 mm, odontioid-irpicoid, subcylindric or joined into
flattened, often longitudinally ridged, plates, not poroid-anasto-
mosing, obtuse, waxy subgelatinous, lurid ochre-yellow then brow-

nish orange.

On the dead fructifications of Poria in the forest. Borneo,
Kinabalu.

Spores 4-4.7 x 3-3.5 », brownish ochraceous under the micro-
scope, ellipsoid, smooth, obtuse, with a thin mucilage sheath (0.3 »).
Basidia 26-32 x 5.5-6.5 », clamped; sterigmata 4, 5 « long. Cheilo-
cystidia -90 x 5-8 pn, projecting —40 », sometimes immersed, scat-
tered along the fertile edge of the plates, fusiform, subacute or
obtuse, thin-walled, often with a fine thin granular incrustation on
the projecting part: pleurocystidia none. Hymenium not thickening,
not encrusted, continuous over the spine-tips; subhymenium c. 30
thick, composed of mucilaginous hyphae 1-2.5 » wide. Hyphae
clamped, but some secondary septa without clamps; in the flesh
and spines 3-12 (—18) » wide with the cells —1000 » long or more,
with mucilaginous walls, not encrusted, fairly compact, regularly
longitudinal in the spines; in the tomentum (0.5-0.6 mm thick)
2-3.5 » wide, much interwoven, clamped, thin-walled, with a dark
brown incrustation, often in annuli, soluble in potash.

Gardens’ Bulletin, Singapore — X XV (1971)

368

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Fig. 2. Merulius irpicoides. Diagram of the structure of the fruit-body

Corner — Merulioid Fungi in Malaysia 369

This may be near to the New Caledonian M. chlorina Pat.,
which also grew on a polypore, but its hymenium was olivaceous
and shortly poroid; no cystidia were recorded for it; there is no
mention of a tomentose undersurface; the spores, 3-4 x 2.5-3 p,
were described as white by Patouillard and yellow brown by Cooke.

M. irpicoides may also be related to M. castaneus Lloyd on
Chamaecyparis and Pinus in Japan, but that species has thick-
walled encrusted cystidia (Ginns 1969). For M. castaneus the genus
Cystidiophorus Bond. et Ljub. has been proposed (Imazeki and
Hongo, Fungi of Japan II, 1965, 125 n. 244); such generic distinc-
tion hardly applies to M. irpicoides.

M. luridochraceus sp. nov.

Resupinatus —12cm latus, floccoso-firmus separabilis; contextu 0.2-0.5mm
crasso, albido; margine angusto albo-villoso. Hymenium mox gyroso-
plicatum, demum irregulariter poroideum, tubulis -2mm_ longis, poris
0.2-0.5mm latis, ceraceo-gelatinosum, laete flavum, dein luride ochraceum,
aetate fulvoferruginosum, sicco fusco-olivaceum nigricans. Sporae 4-4.5 x
2.2-2.7 v. 3.3-4.2 x 2.3-2.7u, pallide ochraceae s.m., leves. Hyphae
fibulatae. Ad lignum putridum. Borneo (typus RSNB 5837, CGE).

Resupinate —12 cm in extent, floccose-firm, separable; context
0.2-0.5 mm thick, whitish; margin narrow, white villous. Hymen-
ium soon gyrose-plicate, then irregularly poroid with tubes -2 mm
long, pores 0.2—-0.5 mm wide, waxy gelatinous, bright yellow then
lurid ochraceous, in age fulvous ferruginous, drying fuscous oliva-
ceous and blackening.

On rotten wood in the forest. Borneo, Kinabalu.

Spores 44.5 x 2.2-2.7 » or 3.3-4.2 x 2.3-2.7 u (RSNB 5837),
pale ochraceous or brownish under the microscope, slightly darker
in potash, smooth, ellipsoid, the wall slightly thickened, with a
thin mucilage sheath, not amyloid, or merely slightly brownish,
cyanophilous in cotton blue. Basidia 20-30 x 5-7 m; sterigmata
4, 4-6 » long. Cystidia none. Hymenium not thickening, continuous.
Hyphae 1.5—5 p. wide, clamped, thin-walled or slightly thick-walled,
scarcely encrusted, often branched from the clamp, more or less
interwoven, subparallel in the basal part of the flesh, compact; in
the tube-trama 1.5-3 » wide, mucilaginous.

Coll. RSNB 5837, Pinosuk Plateau 1700 m alt., 17 March 1964;
RSNB 8489, Mesilau 24 April 1964; Corner s.n., Mesilau, 1600
m alt., 5 Feb. 1964.

This appears to be frequent on Mt. Kinabalu and always resu-
pinate. It seems to be near to M. byssoideus Burt with darker
brown, wider spores 4.5-6 x 3.5-4.5 ». The spores in the type,
RSNB 5837, were so pale that I took them at first to be colourless,
but re-examination and comparison with the other collections
showed that they were slightly ochraceous. A spore-print is needed,
for the species seems exactly intermediate between Serpula and
Merulius s. str.

370 Gardens’ Bulletin, Singapore —X XV (1971)

M.. purpurascens sp. nov.

Resupinatus, 0.4mm -crassus, subcoriaceus, margine saepe anguste reflexo
pileiformi -5mm lato; superne flavidus, ochraceus v. subaurantiacus,
anguste sulcatus, tomentosus v. subvillosus. Hymenium mucilaginosum
rugulosum, poroideum v. gyroso-irpicoideum, poris 0.2-0.5mm latis, 0.5mm
altis, aurantiacum dein cinnamomeum, vinaceum v. fuscum. Sporae 3-3.7
x 1.2-1.5y, albae subarcuatae inamyloideae. Hyphae sine fibulis. Ad
ramulos emortuos in silva. Borneo, Malaya, Solomon Isl. (typus Corner
sn. 29 Nov. 1930, Pahang, CGE).

v. flavido-albus var. nov.

Differt hymenio pallide flavido-albo, sicco incarnato, gloeocystidiis
sparsis 20-35 x 6.5-9.5,. Ad ramulos in silva. Malaya (typus Corner s.n.
8 Nov. 1930, Pahang, CGE).

Resupinate, rather closely adnate but separable, widely effused.
often with narrow flange-like reflexed margin or pileus —-5 mm
wide; upperside pale yellow, ochraceous or orange buff, narrowly
sulcate, tomentose or subvillous; margin yellow to pale buff. dis-
tinct, narrow. Hymenium mucilaginous, rugulose, poroid or gyrose-
irpicoid with projections —0.5 mm high, pores 0.2—0.5 mm wide,
Orange then more or less cinnamon, vinaceous or fawn. Flesh
—0.4 mm thick, felted, subcoriaceous, dry, pallid buff. Dried mate-
rial with pale pinkish buff to vinaceous rose or fuscous purple
hymenium; material in alcohol-formalin turning deep purple
magenta.

On dead branches and twigs in the forest, chiefly montane.
Borneo, Malaya, Solomon Isl. (? Australia, New Zealand).

Spores 3-3.7 x 1.2-1.5 p, white, smooth, subcylindric or sub-
allantoid, not amyloid, 2-guttulate. Basidia 15-18 x 354 u,
without clamps; sterigmata 4, 34 long. Hymenium continuous;
subhymenium 20-25 yp thick, mucilaginous, with hyphae 1.5-2.5 »
wide. Cystidia none, but ? small subfusiform basidioles 4-6 uu
wide. Hyphae 2-5.5 » wide, without clamps, long-celled, the
branches often constricted at origin, cells —200 p» long, closely
interwoven, the walls becoming 0.5—1.5 » thick, smooth or thinly
encrusted (RSS 585), often brownish; with abundant irregular
small crystals and amorphous purplish brown matter in the
hymenium and subhymenium, insoluble in potash but more or less
soluble in Melzer’s iodine.

Coll. Malaya, Pahang, Fraser’s Hill, 1100 m alt., Corner s.n.
29 Nov. 1930.—Borneo, Kinabalu, 1650 m alt., 22 April 1964,
RSNB 8422.—Solomon Islands, Guadalcanal, Gallego, 1000 m alt.,
7 July 1965, RSS 585; Popomanasiu, 2200 m alt., 26 Oct. 1965,
RSS 1623.

v. flavido-albus var. nov.

Differs in the pallid yellowish white hymenium, drying pinkish,
and in the presence of very sparse gloeocystidia 20-35 x 6.5-9.5 nu,
clavate, immersed or in the hymenium.

On sticks in the forest. Malaya, Pahang, Tembeling, Corner s.n.
8 Nov. 1930.

This species without clamps in the fruit-body is close to the
almost cosmopolitan M. corium (Pers.) Fr., from which it differs
in the more richly coloured hymenium, the smaller spores and

Corner — Merulioid Fungi in Malaysia 371

basidia (spores 5-8 x 2.5-4 py, basidia 16-24 x 46 » in M.
corium), and perhaps in the way in which the whole tissue becomes
deep purple in alcohol-formalin. The Australasian species des-
cribed by Cunningham (1963) as M. ravenelii Berk. seems identical,
though it has entirely resupinate fruit-bodies (as may happen in
the Malaysian) and slightly longer and narrower spores 4-5. x 0.8-1
u. Other exotic species without clamps, placed in Byssomerulius
Parmasto, which seem very similar are these: —

M. glaucinus Bourd. et Galz., on pine-wood in France, pellicular
with sparse cystidium-like organs.

M. hirtellus Burt, on dicotyledonous wood in North America,
with cylindric cystidia (Ginns 1968).

M. rubicundus Litsch., on wood of Alnus in Siberia, with cylin-
dric cystidia; described as M. glaucinus. v. lutescens Pilat (Bull.
Soc. mycol. Fr. 49, 1933, 293).

Whether any of these show the purple colour in alcohol-formalin
is not known. The very sparse gloeocystidia of M. purpurascens
v. flavio-albus resemble those of M. rufus and are not comparable
with the hymenial cystidia of the other species.

M. ravenelii Berk. is now interpreted as a synonym of Poria
taxicola (Pers.) Bres., which has been given its own genus Meru-
liopsis. M. purpurascens differs in the less regularly poroid under-
surface, the yellow to orange hymenium (pink to brownish in P.
taxicola), and the narrower hyphae with long cells. I note also in
specimens of P. taxicola that there are sterile, fusiform, obtuse
basidioles frequently in the hymenium, which I have not seen for
certain in M. purpurascens. Dried specimens, however, are very
similar.

M. rubrotremellosus sp. nov.

Effuso-reflexus, pileis -18mm in radio flabelliformibus 1—2mm crassis,
roseo-ruber, ceraceo-gelatinosus. Hymenium minute poroideo-reticulatum,
alveolis 0.3-0.6mm latis, etiam 1—2mm radiatim elongatis, concolore. Sporae
4-5 x 2-2.3y, albae suballantoideae inamyloideae. Hyphae fibulatae. Ad
lignum putridum in silva. Borneo (typus RSNB 2834, CGE).

Effuso-reflexed; pilei -18 mm radius, flabelliform, 1-2 mm thick,
wholly rose-red, waxy subgelatinous. Hymenium minutely poroid-
reticulate with meshes 0.3-0.6 mm wide, sometimes elongate
radially 1-2 mm, concolorous.

On a rotten branch in the forest. Borneo, Kinabalu, Tenompok,
1650 m alt., 6 Sept. 1961, RSNB 2834.

Spores 4-5 x 2-2.3 u, white. smooth, ellipsoid-cylindric to sub-
allantoid, not amyloid. Basidia 25-30 4.5-5 mw; sterigmata 4,
4-5 u« long. Cystidia none, the pore-edges becoming fertile but
apparently with narrow, closely set, excrescent hyphae at first.
Hymenium not thickening, ? not mucilaginous; subhymenium ?
not mucilaginous, the hyphae 1.5-2.5 » wide. Hyphae of the flesh
2-4 » wide, clamped, thin-walled, rather long-celled, radiating and

372 Gardens’ Bulletin, Singapore — X XV (1971)

interwoven, on the surface of the pileus compact and appressed;
with pinkish amorphous granular matter among the hyphae, es-
pecially towards the upperside of the fruit-body; the tissue slightly
pinkish in potash with the cytoplasm of the hyphae turning pinkish,
scarcely changing colour in alcohol-formalin.

This seems at first to be an ally of M. tremellosus Fr., the
tissue Of which turns bright pink in alcohol-formalin, but neither
hymenium nor subhymenium appear to be mucilaginous, and, as
the edges of the hymenial outgrowths may be sterile at first, it is
possible that this is not a Merulius but a poroid ally of the rose-red
Hydnum roseomaculatum P. Henn. et Nym., which is not un-
common on Mt. Kinabalu; yet the hyphae of this Hydnum may
lack clamps (Maas-Geesteranus, Proc. Kon. Nederl. Ak. Wet.
Amsterdam, ser. C, 70, 1967, 63.).

M. rufus Fr., Syst. Myc. 1 (1821) 327: Bourd. et Galz., Hym. Fr.
(1927) 349.

M. pilosus Burt, Mycologia 14 (1922) 184; Ginns, Mycologia 60
(1968) 1229, f.11.— Phlebia merulioides Lloyd, Myc. Writ. 4 (1915)
537; Ginns, Mycologia 61 (1969) 369.—P. castanea Lloyd, Myc.
Writ. 7 (1922) 1108; Ginns, Mycologia 61 (1969) 361.— P. rufa
(Fr.) Christiansen, Dansk Bot. Ark. 19 (1959-60) 164.—? M.
nothofagi G.H. Cunningham, N. Z. Plant Diseases Div. Bull. 83
(1950) 7; Thel. Austral. N. Z. (1963) 324, f. 199.

Resupinate, separable, waxy coriaceous; flesh 0.3-0.7mm thick,
tough, pallid white; margin determinate, white, minutely fimbriate
or byssoid. Hymenium irregularly irpicoid-poroid, the irregular
tubes —1 x 0.2-0.5 mm, with blunt fertile edges, white then pale
ochraceous drab tinged brownish or pinkish, often pale pinkish
orange when young. Smell rather fragrant.

On fallen trunks, branches and twigs, mainly of Fagaceae.
North temperate, Borneo, (? Australia, New Zealand).

Spores 3.5-4.5 (-5) x 2-2.5 uw, white, subcylindric, obtuse,
straight, smooth, not amyloid, slightly cyanophilous in cotton blue.
Basidia 20-28 x 3.7-4.5 yw; sterigmata 4, 3 » long. Hymenium not
thickening; subhymenium —50 u thick, the hyphae 1.5—-2.5 » wide.
Gloeocystidia 28-45 x 6-8 » in the hymenium, 28-75 x 6.5-15 »
when immersed, clavate, thin-walled, smooth, with dense oleaginous
guttulate contents, sparse or frequent, immersed or slightly pro-
jecting, abundant at the margin before the development of the
tubes. Hyphae clamped, often branching from the clamp; in the
flesh 3-6 » wide, radiating, longitudinal, compact, the walls some-
what thickened (toughly subgelatinous in potash), the tissue rather
toughly waxy gelatinous; in the tube-trama 2-4 » wide, with sub-
gelatinous walls and oily contents, interwoven and longitudinal,
fairly compact; basal hyphae 4-8 » wide, the walls 1-2 » thick. All
the tissue, except the middle part of the flesh, turning bright red
pink in alcohol-formalin, only slightly pinkish in potash, the pink
colour caused by the slight reddening of the hyphal cytoplasm.

Coll. Borneo, Kinabalu, Mesilau, Pinosuk Plateau, Bembangan
River, 1300-1800 m alt.; RSNB 1763, 1831, 2635, 2643, 5786.

Corner — Merulioid Fungi in Malaysia 373

The presence of gloeocystidia in this species has been noted
by Ginns, but they seem to have escaped others. The Bornean
material is identical with that which I have collected in Great
Britain. I cannot detect any distinction in M. nothofagi. M. rufus
is near to M. tremellosus but the fruit-bodies are much thinner,
entirely resupinate, and with smaller spores.

M. versicolor (B. et Br.) comb. nov.

Hydnum versicolor B. et Br., J. Linn. Soc. Bot. 14 (1873)
59.— Gyrodontium versicolor (B. et Br.) Maas G., Persoonia 3
(1964) 190; Proc. Kon. Nederl. Ak. Wet. Amsterdam ser. C, 69
(1966) 323.— Hydnum henningsii Bres., Bull. Soc. mycol. Fr. 6
(1890) xlviii, pl. 9, f.2— G. henningsii (Bres.) Pat., Ess. Tax.
(1900) 117.— Hydnum flavidum Lloyd, Myc. Writ. 6 (1920) 957.—
G. flavidum (Lloyd) Reid, Kew Bull. 17 (1963) 268.— G. ser-
puloides (P. Henn.) Reid, 1.c. 268.—Boninohydnum pini Ito et
Imai, Trans. Sapporo nat. Hist. Soc. 16 (1940) 127.—? Hydnum
boveanum Mont., Ann. Sci. nat. Bot. Ser. 2, 4 (1835) 194: Maas
Geesteranus, Persoonia 3 (1964) 157, 191 ut G. boveanum (Mont.)
Maas G.

Sessile or effuso-reflexed, commonly with imbricate brackets,
sometimes solitary or entirely resupinate. Pileus —12cm radius,
2lcm wide, often much smaller, cottony subfloccose to nearly
smooth, pale yellowish white to pale clear primrose yellow; margin
obtuse, then subacute, entire, white. Spines 3—7 x 0.5-Imm, crow-
ded, conical, subacute or obtuse, even subtruncate, discrete, often
angular or somewhat longitudinally rugulose, waxy, brittle, clear
primrose yellow then mustard yellow to fuscous olive or olivaceous
cinnamon. Flesh 3—14mm thick at the base of the pileus, 0.5-6mm
at 5mm from the margin, soft, spongy, floccose, not gelatinous,
pale yellowish to bright primrose yellow. Smell aromatic, some-
what of caramel, slightly pungent, not sour.

On dead standing or fallen trunks, bamboos and palms in
primary and secondary forest, frequent -—1300m alt. Malaya,
Singapore, Africa, Asia.

Spores 3.5—4.7(-S) x 2.5—2.7(-3)u fuscous olivaceous in the
mass (not ferruginous), ellipsoid, obtuse, smooth, with a fine
mucilage sheath. Basidia 15-20 x 4-5; sterigmata 4, 2.5—3, long.
Cystidia none. Hymenium not thickening, continuous over the tips
of the spines from the outset. Hyphae monomitic, without clamps,
not or scarcely inflating long-celled; in the flesh 4-84 wide, thin-
walled, flexuous, longitudinal, without oleiferous or fibre-hyphae;
in the spines 3—Sy wide, mucilaginous, longitudinal; in the sub-
hymenium 2-3.5y, mucilaginous.

This is the common bracket hydnum with yellow to olive brown
spines that at once reveals itself as a Merulius. It is as omnivorous
as M. eurocephalus and the two often cannot be distinguished
from the upperside. Gyrodontium capense Reid seems to differ
only in the slightly larger spores 4.8-5.8 x 3-3.8p.

374 Gardens’ Bulletin, Singapore —X XV (1971)

Phaeophlebia

W.B. Cooke, Mycologia 48 (1956) 401 (with full synonymy of
28 names); Donk, Taxon 6 (1957) 107; Lentz, Sydowia 14 (1960)
127.— Auricula Lloyd, Myc. Writ. 7 (1922) 1152 (non O.K. 1891);
Donk l.c. 21.—? Punctularia Pat., Bull.. Herb. Boiss: 3 (1895) 57:
Talbot, Bothalia 7 (1958) 140, 164; Donk, Persoonia 3 (1964) 287.

P. strigosozonata (Schw.) W.B. Cooke L.c.

Merulius strigosozonatus Schw., Trans. Am. phil. Soc..n.s. 4 (1832)
460. Fie aire ae |

At first conchate or pezizaeform and dorsifixed, then through
marginal growth widely effuso-reflexed with the horizontal pileate
part 14cm radius, spathulate-semicircular or joining in flanges,
sometimes imbricate, becoming undulate and plicate from excessive
tangential growth; upperside closely tomentose, zoned, subsulcate,
dark rufous sepia, madder brown, purplish. brown, bay brown
or umber, with darker fuscous brown to blackish zones, finally
wholly blackish umber and fuscous drab; margin yellowish or
white, tomentose. Hymenium subgelatinous, closely and narrowly
radiating rugulose with shallow obtuse folds, becoming very
crowded in the central part of the hymenium but not poroid,
bright orange near the margin, becoming rufous orange, then
vinaceous umber, somewhat rufous on bruising, drying black.
Flesh pliant coriaceous, in two layers; upper tomentose layer
0.8-1.2mm thick, dark brown; lower subgelatinous layer 0.5—-0.8mm
thick, paler. Smell sour, rank.

On fallen logs in the forest. Singapore, Malaya, New Guinea,
Solomon Isl. Cosmopolitan (? Africa).

Spores 5-7 x 3-4 (6-84 long, Cunningham), white, smooth,
ellipsoid to subcylindric, not amyloid, slightly cyanophilous in
cotton blue. Basidia 28-34 x 5-5.5u (30-S0u long, Cunningham),
clavate; sterigmata 4, 4-5y long. Cystidia none. Hymenium with
the basidia in a more or less discontinuous layer separated by
narrow, 1—2u wide, profusely and irregularly branched, often
lobulate, mucilage hyphae, overtopping the immature basidia as
a mucilage layer —20u thick, the mature basidia protruding through
this layer; subhymenium —30y thick, mucilaginous. Hyphae mono-
mitic, clamped, not inflating; in the lower layer of the flesh 24u
wide, more or less radiating, rather compact, with mucilaginous
walls, colourless, long-celled; in the tomentum 3-5, wide, at first
perpendicular to the surface then interwoven, with smooth brown
walls —1.5, thick, sparsely branched, the growing tips colourless
and thin-walled, very dense and almost agglutinated in a thin layer
c. 20u thick at the junction with the gelatinous layer of the flesh.
Hymenium and tomentum turning dark brown in potash, scarcely
changing colour in alcohol- formalin.

Corner — Merulioid Fungi in Malaysia 375

mf
a eae — ~ dl
- Va no forte) De J-f-
BO TA Ly TALES GD EIGEN LIL IDG,
4 4 ea Aa VAS a 8 ee Paes SP Get Taw owen we MY ee 4 =
Sx fe ~- + ~‘w=3 a te se ee wate eo &@, © Ves . . =< ae! oe
eke Be ean Coe ns Seo. ROI Berto GY} SSS Se ie ee le a ge
Bes aS ae Se ee a a
NE See A ecg ee ner Oa
eo — —_ * sa. % es ae i Sa SEE Bee | —
e———— es — ( —_—_—, at 5 ES \(—_,
—s — —— ——_——_ — 1
— a = \ \ —— <—— \ Je -—_— | \ \ \—
—_— ——<——_— = —__ = <a ——
ee ee ee a
=- > oe TT <a —— ——— © —— ) '~————- . — , ——= “3
Be ny es ie ie = "4 _—™~ ee Set Pree > hae ay os
—_— —_— 6... = = “ eee EF s
—— ‘ et —_ “~SSNN “~ =~ aN -___ - a a es ey ees
Wit asa wa;n ena “5 Mall sion Sea No eavellal i ian Aye
DIO DPMP POD ro HIATT MS SIT ON TAR ALES
+ ? POM RCANN LU apd ,)e> or I) s on\ii\t Wy, i
TEPER ISRO Cian NTN Mii"

=

: ROAR CN ELT
MIMIC THe yet itWS

Fig. 3. Phaeophlebia strigosozonata. Diagram of the structure of the
pileus in Ls., x 25; hymenium, x 1000.

376 Gardens’ Bulletin, Singapore — XXV _ (1971)

Coll. Singapore, Bukit Timah, Corner s.n. April 1929, 29 March
1931.— Malaya, Johore, Corner s.n. Mawai-Jemaluang Rd., 30
March 1941; Pahang, Tembeling, Corner s.n. 6 June 1931; Kelan-
tan, Sungei Ketil, Corner s.n. 1 June 1931.—New Guinea, Woitapi,
2000m_ alt., Corner s.n. 23 Aug. 1960.— Solomon Isl., Small
Malaita, RSS 18/0, 27 Nov. 1965.

This fungus resembles superficially Stereum spadiceum but is
easily distinguished by the irregularly rugulose, miucilaginous
hymenium. Though I obtained spore-prints from the Malaysian
material, I failed to find any fertile basidia until I examined my
collection from the Solomon Islands; even then I saw few among
the many that were immature. The hymenium does not form a
discrete layer. The young fruit-bodies are dorsifixed as in
Meiorganum. '

In October 1931, I grew six fruit-bodies on a dead branch taken
from the forest on Bukit Timah in Singapore. The largest took
19 days to reach its full size (semicircular pileus 27mm in radius,
44mm wide) and then persisted and spored for twenty days, when
it became mouldy. It had a rate of marginal growth of 1—-1.5mm
in 24 hours, but this increased to 1.5—-2mm in 24 hours when the
fruit-bodies were c. 7mm in radius; this rate declined to 1-1.5mm
after a few days and then increased to 2mm on the 14th day before
declining again. The other fruit-bodies grew in much the same
manner but stopped at various sizes from 7—22mm in radius.
However, when radial growth stopped, vigorous tangential growth
began and caused the margin to become undulate and folded.
The zonation of the pileus corresponded with the 24 hour in-
crements in growth, which were mainly, if not entirely, nocturnal.
Dry weather impeded marginal growth but did not stop sporing
which continued throughout the life of the fruit-body after the
first 3-4 days. The growing margin was white but the new tissue
on either side quickly coloured.

Corner — Merulioid Fungi in Malaysia 377

References

Bourbor, H. et A. Galzin (1927) Hymenomycetes de France.

Cooke, W. B. (1957) The genera Serpula and Meruliporia. My-
cologia 49, 197-225.
Corner, E. J. H. (1932) The fruit-body of Polystictus xanthopus.
Ann. Bot. 46, 71-111.
(1966) A monograph of cantharelloid fungi. Ann. Bot.
Mem. 2.
(1968) A monograph of Thelephora (Basidiomycetes).
Nova Hedwigia, Beiheft. 27.
(1971) Studies in the basidium II. (in press).

CUNNINGHAM, G. H. (1963) The Thelephoraceae of Australia and
New Zealand. New Zeal. Dept. Sci. Ind. Res. Bull. 145.

Donk, M. A. (1958) The generic names proposed for Hymenomy-
cetes. LX, ‘Meruliaceae’ and Cantharellus s. str. Fungus
28, 7-15.
(1960) The generic names proposed for Polyporaceae.
Persoonia 1, 173-302.
——- (1962) Notes on _ resupinate Hymenomycetes VI.
Persoonia 2, 217-238.
(1964) A conspectus of the families of Aphyllophorales.
Persoonia 3, 199-324.
GINNS, J. H. (1968) The genus Merulius I. Species proposed by
Burt. Mycologia 60, 1211-1231.
(1969) The genus Merulius II. Species of Merulius and.
Phlebia proposed by Lloyd. Mycologia 61, 357-372.
———— (1970) The genus Merulius III. Species of Merulius and
Phlebia proposed by Schweinitz and Peck. Mycologia 62,
238-255.
HARMSEN, L. (1956) On Merulius silvester Falck and M. himan-
tioides Fr. Friesia 5, 257-263.
——— (1960) Taxonomic and cultural studies on brown spored
species of the genus Merulius. Friesia 6, 233-277.
LEMKE, P. A. (1964) The genus Aleurodiscus s. str. in North
America. Can. J. Bot. 42, 213-282.
LENTZ, P. L. (1957) Studies in Coniophora. I, the basidium.
Mycologia 49, 534-544.
PARMASTO, E. (1967) Corticeaceae U.R.S.S. IV. Eesti NSV Tead.
Akad. Toim. Biol. 16, 377-394.
-———— (1968) Conspectus systematis Corticiacearum. Inst.
Zool. Bot. Ac. Sci. R.P.S.S. Estonicae, Tartu.
SINGER, R. (1945) The Boletaceae of Florida II. Farlowia 2, 223-
303.
SINGER, R. and K. GRINLING (1967) Some Agaricales from the
Congo. Persoonia 5, 353-377.

378 Gardens’ Bulletin, Singapore — XXV (1971)

Index of species attributed to Merulius

Merulius Fr. (1821)

Merulius St.-Am. (1821) =
Cantharellus Fr.

Merulius S. F. Gray (1821) =
Hygrophoropsis Maire

affinis Jungh. (1838)

affinis Vel. (1922)

agathidis Corner

albidus Pers. (1825)

albostramineus Torrend (1913) = ?
M. porinoides Fr.

albus Burt (1917) = Poria cocos
(Schw.) Wolf

alveolaris Pers. (1925) = Favolus
ambiguus Berk. (1872)

americanus Burt (1917) = M.
himantioides Fr.

armeniacus Berk. (1872)

atropurpureus W. B. Cooke (1943) =
?. M. taxicolus (Pers.) Duby

atrovirens Burt (1917) = M. pinastri
(Fr.) Burt

aurantiacus §. F. Gray (1821) =
Hygrophoropsis aurantiacus (Fr.)
Maire

aurantiacus Kl. (1836) = ? M.
molluscus Fr.

aurantiacus Pers. (1822)

aurantius Lloyd (1922) = M.corium
(Pers.) Fr.

aureus Fr. (1828)

qauroreus Pers. (1825) = Cantharellus
lutescens Fr. f. Donk (1969)

baileyi B. et Br. (1883) = an agaric
f. Cunningham (1963)

bellus B. et C. (1872)

binominatus Mass. (1913) = M.
eurocephalus (B. et Br.) Petch

borealis Rom. (1911) = Athelia f.
Parmasto

brassicaefolius Schw. (1822) = M.
lacrimans Fr.

bryophilus Pollini (1824) =
Leptoglossum bryophilum (Fr.)
Ricken

byssoideus Burt (1917)

byssinus Pers. (1825)

canaliculatus Pers. (1825)

candicans Pers. (1825) =
Leptoglossum muscigenum (Fr.)
Karst.

candicans Roum. (1886) = M.
pinastri (Fr.) Burt

candidus Lloyd (1923) = Poria
calcicolor Sacc. et Syd.

cantharelloides Pers. (1825) =
Cantharellus tubaeformis Fr. f.
Donk (1969)

cantharellus Pers. (1825) =
Cantharellus cibarius Fr.

carbonarius Lloyd (1920) = M.
lacrimans Fr. v. carbonarius
(Lloyd) W. B. Cooke

carbonarius Pers. (1825) =
Geopetalum carbonarium (Fr.) Pat.

carmichaelianus Berk. (1860)

castaneus Lloyd (1916) =
Cystidiophora castanea (Lloyd)
Imazeki

ceracellus B. et C. 1872)

cervinus Pers. (1825) = Cantharellus
lutescens Fr. f. Donk (1969)

chilensis Speg. (1920)
chlorina Pat. (1908)

cinereus Pers. (1825) = Cantharellus
cinereus Fr.

cladopus Pers. (1825)

clavatus Secr. (1833) = Gomphus
clavatus S. F. Gray

collariatus Pers. (1825)

conchoides Lloyd (1922) = M.
corium (Pers.) Fr.

confluens Schw. (1822) = M. corium
(Pers.) Fr.

consimilis Lloyd (1922) = M.
eurocephalus (B. et Br.) Petch

corium (Pers.) Fr. (1828)
cornucopioides Pers. (1825)
crassus Lloyd (1925)

crispatus Fr. (1821)

crispus Pers. (1825) = Plicatura

croceus Duby (1830) = M. aureus
Fr.

crocicreas Ces. (1878)
crucibulum Pers. (1825)

cubensis Burt (1917) = M. corium
(Pers.) Fr.

cupressi Schw. (1822) = a gall.

cupularis Pers. (1825)

cuticularis Lév. (1855)

cyatheus Vel. (1922)

debriscola Lloyd (1924) = not a
fungus (Ginns 1969)

deliciosus Brig. jun. (1878) =
Cantherellus deliciosus (Brig. jun.)
Sacc.

demissus (Berk.) Pat. (1908)

densus Rick (1960)

destruens Pers. (1825) = M.
lacrimans Fr.

Corner — Merulioid Fungi in Malaysia

dilatatus Pers. (1825) = Cantharellus
tubaeformis Fr. f. Donk (1969)

diobensis Beeli (1929)

domesticus (Pers.) Falck (1907) =
M. lacrimans Fr.

dubiosus Bres. (?)

dubius Burt (1917) = Poria
sanguinolenta (Fr.) Cke.

elegans Pers. (1825)

elliottii Mass. (1892)
epiphyllus Pers. (1825)

erectus Lloyd (1921) = ? a
clavarioid fungus (Ginns 1969)

eurocephalus (B. et Br.) Petch (1910)

fagineus Pers. (1825) = Plicatura

farlowii Burt (1917) = M. serpens
Fr. (Byssomerulius Parmasto)

fissus Pers. (1825)

fiavescens Bres. (1920)

foetidus Pers. (1825)

foliaceus Pers. (1825)

fugax Fr. (1821) = M. molluscus Fr.

fuligineus Pers. (1825) =
Cantharellus infundibuliformis Fr.

fulvus Lasch (1829) = Phlebia
radiata Fr.

fuscescens Bres. (1920)

fuscus Lloyd (1925)

fuscus Pers. (1828)

fusisporus Rom. (1911) =
Parmasto

gelatinosus Lloyd (1922) =
himantioides Fr.

gelatinosus Petch (1925)

giganteus Saut. (1877) =
lacrimans Fr.

_ glaucinus Bourd. et Galz. (1923)
gloeoporus Rick (1960)
— gossypinus Rick (1960)

_ guillemotii Boud. (1894) =
lacrimans Fr.

_gyrosus Burt (1917) =
Rom.
_haedinus B. et C. (1872) =
corium (Pers.) Fr.
helvelloides Pers. (1825)
_hexagonoides Burt (1917)
himantioides Fr. (1821)

_hirsutus Burt (1917) = M. corium
(Pers.) Fr.

hirtellus Burt (1917) = ? M. bellus
= Byssomerulius

Athelia f.

M. borealis

T ee.

Parmasto)
hispidulus O. K. (1898)
hydnoides P. Henn. (1904)

379

hydrolips Merat (1821) =
Cantharellus cinereus Fr.

imbricatus Balfour-Browne (1955)

incarnatus Schw. (1822)

incrassatus B. et C. (1849) =
Meruliporia incrassata (B. et C.)
Murr.

infundibuliformis Cke. et Mass.
(1888) = Rodwaya
infundibuliformis (Cke. et Mass.)
Syd.

insignis Wakef. (1917) = M.
eurocephalus (B. et Br.) Petch

interruptus Bres. (1925)

irpicinus Pk. (1894) =
(Fr.) Burt

irpicoides Corner

lacrimans Fr. (1821)
lacrymans Pers. (1825)

laeticolor B. et Br. (1878) =
molluscus Fr.

lamellosus Pers. (1825)
lepidus Rom. (1911)

leucophaeus Pers. (1825) =
Geopetalum carbonarium (Fr.) Pat.

lichenicola Burt (1917) =
mollusca (Fr.) Cke.

lignosus Berk. (1854)

lividus Bourd. et Galz. (1923) =
Phlebia f. Parmasto

lobatus Pers. (1825) = Leptoglossum
lobatum (Fr.) Ricken

luridochraceus Corner
luteolus O. K. (1891)

macedonicus Pilat (1938)
melanoceras Mont. (1859)
miniatus Wakef. (1931)

minor Falck (1912) =
(Harmsen 1952)

moelleri Bres. et P. Henn. (1896)

molluscus Fr. (1821) =
Leucogyrophana Pouz.

molluscus Pers. (1825)
montanus Burt (1917)

muscigenus Pers. (1822) =
Leptoglossum

muscigenus St.-Am. (1821) = =
Leptoglossum muscigenum (Fr.)
Karst.

muscorum Pers. (1825)

M. pinastri

Poria

? nom. nud:

nigripes Pers. (1825) =
Hygrophoropsis aurantiacus (Fr.)
Maire

niveus Fr. (1828) = ? Plicatura (f.
Pilat, Bull. Soc. mycol. Fr. 49,
1933, 292)

nothofagi G. H. Cunningham (1950)

380

ochraceus Lloyd (1921) =
(Pers.). PF;

odoratus Schw. (1822) =
Cantharellus odoratus (Schw.) Fr.

ottonis Pers. (1825)

pallens Berk. (1841) = M. rufus Fr.

pallens Schw. (1832) = M. corium
(Pers.) Fr.

papyraceus Fr. (1828) = M.
himantioides Fr.

M. corium

papyrinus Quel. (1888) = M. corium
(Pers.) Fr.

patellaeformis B. et C. (1872)

pelliculosus Cke (1891) = M. corium

(Pers.) Fr.

petropolitanus Fr. (1838) =
niveus Fr. f. Pilat (1933)

pezizoideus Speg. (1881)
phlebioides Bourd. et Galz. (1923)
pilosus Burt (1922) = M. rufus Fr.
pinastri (Fr.) Burt (1917)

pinorum Britz. (1909) = M. pinastri
(Fr.) Burt

polychromus Petch (1917) =
himantioides Fr. (2? M.
eurocephalus)

porinoides Fr. (1821)

pruinatus Secr. (1833) Cantharellus
tubaeformis Fr.

pruni Pk. (1906) = M. rufus Fr.

pseudocantharellus Pers. (1825)
pseudolacrymans P. Henn. (1901) =
M. eurocephalus (B. et Br.) Petch
pseudomolluscus Parmasto (1962) =
Leucogyrophana t. Parmasto
puiggarii Speg. (1919) = ? M.
eurocephalus (B. et Br.) Petch
pulverulentus Fr. (1828) = M.
pinastri (Fr.) Burt v. M. umbrinus
Fr. f. Parmasto
purpurascens Corner
purpureus Pers. (1825) =
violaceus Fr.
purpureus Rick (1938)

quéletii Schulz. (1885)

radiatus Pers. (1822)

ravenelii Berk. (1872) =
(Pers.) Duby

retirugis Pers. (1825) =
Leptoglossum retirugum (Fr.)
Ricken

rimosus Berk. (1891)

roseus Schw. (1822) = Cantharellus
roseus (Schw.) Fr.

rubellus Pk. (1882) = M. incarnatus

Schw.

Craterellus

M. taxicolus

Gardens’ Bulletin, Singapore — X XV (1971)

rubicundus Litsch. (1934) =
Byssomerulius Parmasto

rubrotremellosus Corner
rufus Fr. (1821)
rugulosus B. et C. (1867)

sartilaginosus Wettst. (1885)
sclerotiorum A. Moell. (1907)

serotinus Pers. (1825) =
Leptoglossum muscigenum (Fr.)
Karst.

serpens Fr. (1821 = Byssomerulius
Parmasto

sesia Pers. (1828)

sessilis B. et Br. (?)

silvester Falck (1907) = ? M.
himantioides Fr.

similis B. et Br. (1875) = M.
eurocephalus (B. et Br.) Petch

sinuosus Pers. (1825) = :
Pseudocraterellus

sodiroi (Pat.) Rick (1960)

sordidus B. et C. (1891) = M. :
pinastri (Fr.) Burt

sororius Burt (1917)

spadiceus B. et C. (1867)

spathularius Schw. (1822) =
Dacryopinax Martin

spissus Berk. (1872) = Meruliporia
incrassata (B. et C.) Murr.

squalidus Fr. (1828) =
himantioides Fr.

squamula Pers. (1825)

stereoides P. Henn. (1900)

stratosus Pilat (1936)

strigosozonatus Schw. (1832) =
Phaeophlebia

subambiguus P. Henn. (1897) =
eurocephalus (B. et Br.) Petch

subaurantiacus Pk. (1885) =
molluscus Fr.

succineus Lloyd (1924) =
(Pers) FT.

sulcatus Pk. (1878) = nomen
dubium (Ginns 1970)

sulphureus Burt (1917) not Merulius
(Ginns 1968)

sulphureus Vel. (1922)

taxicolus (Pers.) Duby (1830)

tenellus Pers. (1825)

tenuis Pk. (1894) =
Fr.

tenuissimus B. et Br. (1883), ?

“identity (Cunningham 1963)

terrestris (Pk.) Burt (1917) =
lacrimans Fr.

tesselatus Bres. (1890) =
eurocephalus (B. et Br.) . Pot

M. corium

M. himantioides

Corner — Merulioid Fungi in Malaysia

tignicola Harmsen (1952) = M.
pinastri (Fr.) Burt f. W. B. Cooke

tomentosus Burt (1917) = ? Poria

tortilis Pers. (1825)

torulosus Pers. (1825)

tremellosus Fr. (1821)

tubaeformis Secr. (1833) =
Cantharellus tubaeformis Fr.

tubiformis Pers. (1825) =
Cantharellus infundibuliformis Fr.

_ turfosus Pers. (1825)

uliginosus Pers. (1825) =
Leptoglossum lobatum (Fr.) Ricken

ulmi Pk. (1906) = M. corium (Pers.)
Fr.

umbonatus Pers. (1825) =
Cantharellula umbonata (Fr.)
Singer

381
umbrinus Fr. (1828) = ? M.
himantioides Fr., ? M.
pulverulentus Fr.
vastator Pers. (1825) = M. lacrimans

Fr.
versiformis B. et Br. (1875)

villosus Pers. (1825) = Cantharellus
tubaeformis Fr.

violaceus (Fr.) Pat. (1900)
violaceus Pers. (1825)
wrightii Berk. (1872)

xanthopus Pers. (1825) =
Cantharellus lutescens Fr.
xylostromaticus (Berk.) Rick =

Porotheleum leucobombycinum W.
B. Cooke

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iy ian Pas 9 Al) wept 2a ’ i 194) Mure ! Pek axg
= wi (dro) eee iy
ff 1 say rAprclianiwe! wraiy dts ; x= iS.
4 ™ } ei)? Bs ta gl ipa bs wot
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pases

REVIEW
Grasses of Malaya

GILLILAND, H. B., Grasses of Malaya, V1+319 pages, with 67
figures, 22 partially coloured plates, and 36 photo plates. Singapore
Government Printer (P.O. Box 485, Singapore); 1971. S$$30.

At last a highly useful and beautifully bound botanical treatise
on Malayan grasses is available.

H. N. Ridley’s monumental work, Flora of the Malay Peninsula,
was published in 5 volumes nearly half a century ago. Since then
much has been added to the knowledge of the Malayan flora. A
project of writing a revised Flora of Malaya initiated in early
1950’s, of which only the first two volumes, namely, Orchids of
Malaya (1953) and Ferns of Malaya (1954), both by R. E. Holttum,
came out many years ago and has gone through several editions.
The present book, Grasses of Malaya, is listed as the third volume
of this distinguished series.

Before the late Professor Gilliland left Singapore in 1965, a
manuscript “composed of Keys and species descriptions...” was
handed over to Mr. H. M. Burkill, former director of the
Singapore Botanic Gardens. Professor Gilliland then returned to
S. Africa where he passed away shortly afterwards. For several
years, Mr. Burkill, besides performing his official duties as Director
of the Gardens, took up the painstaking job of editing and revising
the manuscript and brought it into the present form.

In the original Malayan flora, the grasses occurred only on the
fringe of the tropical rain forests, and naturally they were rather
poorly represented. After the opening and destruction of the forests,
there followed the introduction (both intentionally and unintention-
ally) of new grasses into this part of the world. Thus it is justifiable
and necessary to include a good many naturalized and cultivated
grasses in the account.

Besides an informative introduction, this book consists of two
parts. The first part, dealing with the bamboos, is mainly based on
Dr. Holttum’s work on Malayan bamboos; the second part, or the
main part, dealing with grasses, follows the pattern set up by
Dr. Bor’s work on grasses of Burma, Ceylon, India and Pakistan.
Since economically the grasses are probably of greater importance
than any other family of plants, a brief summation of economic
uses Of many grass species contained in the work, mainly taken
from I. H. Burkill’s well-known Dictionary, adds much to the
practical value of this book. All these are clearly acknowledged by
the editor.

The main body of the book — The generic and species descrip-
tions and the keys — is admirable. The descriptions are, in general.
brief but to the point. And because of the careful selection of the
most obvious characters, the keys are specially easy to work with.

The present writer, however, would like to suggest a few minor
points for consideration of improvement in future editions.

384

1. Professor J. L. Audus’ excellent drawings, 22 in all, partially
coloured, certainly add much beauty and value to the book.
It would, however, bring harmony and uniformity to the
book if the scales of measurement of the drawings could
be converted into the metric system.

2. Most of the text-figures are very good; only a few of them
e.g. Fig. 31, on p. 161 and Fig. 51, on p. 233, probably
should have been improved.

3. It should be better to indicate clearly the fresh material or
herbarium specimens of which photographs were taken
as reproduced in the plates at the end of the book, especially
with regard to the collector and the collecting numbers where
herbarium specimens were used.

4. In some cases (e.g. on pp. 18, 19 etc.), vernacular names
are given immediately below the basinyms and synonyms
and also in the same lettering. This could be very confusing.
It would probably be better to transpose all vernacular
names to the end of the description as is the practice of
most authors.

5. Preferably, the reference on the grasses of Java by Backer
(on p. 10) should be replaced by the excellent chapter on
grasses (as “‘Poaceae’’) in the 3rd volume of Flora of Java
(Pp. 495-641, 1968) by Backer and Bakhuizen. This was
based on Backer’s original manuscript and elaborated by
Ch. Monod de Froideville.

This book will be a welcome and useful addition to any botanist’s |
or agriculturist’s working library.

(HSUAN KENG; Assoc. Professor of Botany,
Department of Botany, University of Singapore.)-

Obituary

Juraimi bin Samsuri

It is with great regret that we have to record the sad demise
of Juraimi bin Samsuri, who was the artist of the Singapore Botanic
Gardens. Juraimi, as usual, reported for duty on 30.6.71 but just
before noon asked to be excused from work for the rest of the day,
as he was not feeling well. He returned home, and at about
2.00 p.m. collapsed suddenly and passed away. He leaves behind
a widow and five children.

Of simple taste and quiet disposition, Juraimi was born in
Singapore on 14th April, 1923. He was educated at the Tanglin
Besar Malay School and Victoria School, where he excelled in art.
He had no specialised training, but through perseverance and
hardwork mastered the technique in line drawing and painting.

He joined the Botanic Gardens service in 1942 as a Label
Printer and because of his talents in the field of art he was subse-
quently appointed as the Gardens’ artist. He served under many
prominent botanists and horticulturists including Holttum, Corner,
Henderson, Furtado, Ewart, Addison, Purseglove, the late Sinclair,
H. M. Burkill, Chew and others. His drawings and paintings in
their publications bear testimony to his skill as an artist. In the
last three years before his death he showed a great interest in
photography and was responsible for taking many of the pictures
which were required, especially for horticultural work in the
Department; but it is as an artist of botanical drawings that he
will be well remembered. He did not confine his interest only to
his work as an artist, but also assisted wholeheartedly in the
diverse extension work of the Department.

He is sadly missed by his many colleagues and friends. To his
widow and children, we extend our deepest sympathy. A.G.A.

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A sions ant
Lane a i :
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INDEX, VOL. XXV

Basionyms and synonyms appear in italics, new taxa, namcs
and status in boldface. Page numbers in boldface refer to taxonomic
treatments, in italics to illustrative matter on the same, preceding

or following page(s).

Acanthocladium alboalare, 339.

Acanthophysium, 359.

Acanthorrynchium papillatum, 340.

Acroporium, spp. listed, 340.
Adicea mooreana, 157.

Adina, 247, 249.
globiflora, 249.
multifolia, 250.
trichotoma, 249.

Aerobryidium filamentosum, 338.
Aerobryopsis longissima, 338.
Aleurodiscus, 359.

Anabaena sp., 184.

Ancistrocladus, 235, 237.
tectorius, 235, 236.

Ankistrodesmus falcatus, 182.

Anthocephalus, 248.
chinensis, 252.

Anthoceros glandulosus, 341.

Artabotrys, 235.
suaveolens, 235.

Arthrodesmus, spp. listed, 184.

Ascocentrum, 105.
micranthum, 106.

Auricula, 374.

Bazzania, spp. listed, 342.
Boletaceae, 357.

Boletus. 358, 360.
xylophilus, 358.

Boninohydnum, 365.
pini, 373.

Borassus dichotoma, 301.
flabellifer, 301.

Brachymenium, spp. listed, 337.
Byssomerulius, 359, 365, 360,
371.

Calopus, 360.

Calostoma, 109.
berkeleyi, 109.
cinnabarinum, 109.
fusca, 109.
insignis, 109.
junghuhnil, 109.
lurida, 109.
lutescens, 109.
orirubra, 109.
pachystelis, 109.
ravenelti, 109.
retisporum, 109.
sarsinil, 109, 7/0, //1.
viridis, 109.

Calymperes, spp. listed, 337.
Campylodontium flavescens, 339.
Campylopus, spp. listed, 336.
Caudalejeunea stephanii, 343.
Chaetomitrium torquescens, 339.
Cheilolejeunea, spp. listed, 343.

Chiloscyphus, 342.
argutus, 341.

Clastobryella asperrima, 339.

Cleisostoma micranthum, 106.
poilanei, 106.
tixieri, 106.

Closteriopsis longissima var.
tropica, 182.

Closterium, spp. listed, 183.
Coelastrum microporum, 182.

Cololejeunea, spp. listed, 343-344.
pahangiana, 343, 348, 350.

Colura acroloba, 344.
Coniophora, 356, 358.

Coniophoraceae, 355, 356, 357,
358,..359.

Corticaceae, 355, 357, 359.
Corticium, 360.

Corypha, 318.

Cosmarium, spp. listed, 184.
Cyathophorella tenera, 339.
Cystidiophorus, 369.

388

Dactylococcopsis sp., 184.

Dendrocnide, 1, 7, 112, 113, 141.

index to collections studied, 97—
104.

index to taxonomic descriptions,
104.

species excluded, 96.

species unidentified, 95.

sect. DENDROCNIDE, 3, 4, 7,

8, 34.
SARCOPUS, 3, 4, 7, 8,

sect.
34.

amplissima, 4, 9, 81, 82, 84,
85, 86, 88.

anacardioides, 42.

basirotunda, 4, 10, 34, 35.

batanensis, 22.

carriana, 4, 10, 45, 46.

celebica, 9, 92, 93.

chingiana, 32.

coerulea, 96.

contracta, 53, 95.

corallodesme, 10, 51. 53, 54.

cordata, 10, 56, 58, 59.

costata, 7, 11.

crassifolia, 8, 18, 20.

densiflora, 8, 24, 25.

diffusa, 21.

elliptica, 2, 4, 8, 16, 17.

excelsa, 2, 7, 10, 64, 65.

gigantea, 95.

harveyi, 3, 9, 88, 89, 90.

kajewskii, 2, 9, 94.

kjelbergii, 2, 8, 86, 87.

kusaiana, 71.

latifolia, 10, 71, 73, 76, 95S.

longifolia, 3, 9, 78, 79, 80, 86,
94.

luzonensis, 2, 9, 40.
var. anacardiodes, 42.
var. luzonensis, 40, 4/.

meyeniana, 4, 8, 21, 22, 24.
forma meyeniana, 21, 23, 24.
forma subglabra, 22, 24.

microstigma, 4, 9, 53, 82, 83,
86, 95.

milnei, 88, 90.

mirabilis, 2, 8, 90, 9/.

morobensis, 10, 62, 67, 68.

moroides, 2, 4, 9, 44, 55, 56, 57.

nervosa, 10, 69, 70.

nitida, 47, 50.

oblanceolata, 2, 4, 8, 14, 75, 16.

peltata, 2, 4, 9, 18, 42, 45, 47.
var. murrayana, 45.
var. peltata, 42, 43.

photinophylla, 3, 10, 60, 6/,
62, 64, 66, 67.

platyphylla, 22.

pulus, 36.

rechingeri, 3, 9, 74, 75, 76, 78.

rigidifolia, 8, 26, 27, 92, 94.

saipanensis, 71.

salomonensis, 79.

schlechteri, 9, 71, 72, 76, 76,
77, 88

sessiliflora, 95.

sinuata, 3, 4, 9, 36, 37, 38, 40,
i, 2

Gardens’ Bulletin, Singapore — XXV (1969-1971)

stenophylla, 11, 12.

stimulans, 4, 7, 8, 11, 13, 14,
32.

subclausa, 3, 8, 26, 28, 29.

subglabra, 22.

subpeltata, 24.

ternatensis, 3, 10, 47, 48, 49,
$1;1\53.

torricellensis, 3, 10, 51, 52.

urentissima, 3, 4, 8, 32, 33, 34.

venosa, 8, 30, 3/.

vitiensis, 3, 10, 62, 62, 63.

warburghii, 79.

Desmogonium rabenhorstianum,
180.

Desmotheca apiculata, 338.

Dicerostylis, 108.
lanceolata, 107.

Dicranoloma braunui, 336.
Dictyosphaerium pulchellum, 183.
Dimorphococcus lunatus, 183.
Diplasiolejeunea brachyclada, 344.
Distichophyllum mittenii, 339.
Drepanolejeunea, spp. listed, 344.

Ectropothecium, spp. listed, 341.
Endotrichella elegans, 338.
Ephemeropsis tjibodensis, 339.
Euastrum, spp. listed, 183-184.
Euglenia, spp. listed, 182.

Eunotia maior var. emarginata, 180.

Exodictyon blumii, 336.

Ficus, 2.

Fleurya, 111, 115.
aestuans, 164.
alatipes, 124.
bulbifera, 121.
canadensis, 118.
capensis, 152.
caravellana, 164.
cordata, 128, 164.
cuneata, 170.
disepala, 126, 129.
funigera, 157.
glandulosa, 164.
glomerata Gaud., 145, 149.
glomerata Griseb., 170, 172.
grossa, 159.
ingrata, 164.
interrupta, 145.
lanceolata, 161, 163.
lurida, 164.
mitis, 152.
mooreana, 157.
ovalifolia, 149.
paniculata, 167, 170.
peduncularis, 152, 154.

Index

Fleurya (cont.)

perrieri, 164.
petiolata, 164.
photinophylla, 60.
podocarpa, 149, 152.
ruderalis, 170.
spicata, 115, 145, 149.
umbellata, 170, 172.
urophylla, 157.
urticoides, 124, 126.
var. glabrata, 124.

Fleuryopsis, 115.
Floribundaria, spp. listed, 338.

Freycinetia, 189, 209.
enumeration of species, 195.

keys to Malayan species, 193-195.
species of uncertain position, 222.

sect. AURICULIFOLIAE, 217.

sect. FILIFORMICARPAE, 220.

sect. HEMSLEYELLA, 230.
sect. POLYSTACHY, 219.

sect. RACEMOSIFLORAE, 214.

sect. SARAWAKENSES, 215.
acuminata, 190, 204.

angustifolia, 189, 190, 193, 194,
196, 197, 199, 210, 212, 214.

arborea, 191.
bennettii, 204.
biloba, 209, 211, 224.
borneensis, 211, 225.
brunoniana, 197.
confusa, 189, 190, 191, 192,
193, 194, 203, 232.
var. minima, 209, 213, 232.
concinna, 212, 225.
corneri, 189, 190, 193, 195,
198, 198, 212, 225.
ciliaris, 211, 225.
creaghii, 213, 229.
debregeasiana, 197.

discoidea, 209, 213, 226, 227.

distigmata, 189, 203
gitingiana, 209, 211, 213, 230.
hemsleyi, 211, 226.
imbricata, 189-195, 203, 210,
Zils, 21 ole
var. hispidula, 204, 215.
var. kuchinensis, 213, 215.
javanica, 189-191, 792, 193,

194, 195, 196, 207, 204, 206,

210, 205.
var. expansa, 209, 228.
var. javanica, 228.
kalimantanica, 209, 213, 230,
aad.
kamiana, 189-194, 198, 201],
205.
kinabaluana, 209, 211, 219.
kingiana, 203.
lucens, 190, 195, 196.
lucida, 211, 217, 2/8.
malaccensis, 190, 197.
montalbanica, 210.
montana, 190, 195, 196.
palawanensis, 209.
var. andersoniana, 209, 211,
222.

parviaculeata 209, 212, 220,
221.

rigidifolia, 189-191, 192, 193,
194, 203, 204, 210, 212,
220, 223.

robinsonii, 228.
var. meijeri, 209, 213, 228.

sarawakensis, 212, 213, 215.

schefferi, 203, 204.

sumatrana, 189, 190, 192, 193,
194, 198, 199, 201, 210, 211,
ZiT.
var. penangiana, 192, 199,
200, 201, 202.

tenuis, 211, 214.

valida, 190, 199.

villarii, 203.

winkleriana, 213, 230.

Frullania, spp. listed, 346.
Frustulia rhomboides, 180.

Garovaglia plicata 338.
Girardinia marginata, 124.
Globba, 242.

Gloeoporus, 360.
Glossadelphus zollingerii, 341.
Gomphonema subtile, 182.
Groutiella goniorrhyncha, 337.

Gyrodon, 357.
intermedius, 357.
merulioides, 357.

Gyrondontium 365, 356.
boveanum, 373.
capense, 373.
flavidum, 373.
henningsii, 373.
serpuloides, 373.
versicolor, 373.

Gyrophana, 365.
pseudolacrymans, 366.

Gyrophora, 365.

Haynea ovalifolia, 149, 152.
Herbertia cf. parisii 342.

389

Homaliodendron flabellatum, 339.

Hormidium sp., 183.

Hydnum, 360.
boveanum, 373.
flavidum, 373.
henningsii, 356, 373.
roseomaculatum, 372.
versicolor, 356, 373.

Hydrogonium consanguineum, 337.

Hygrophoropsis, 358.
flabelliformis, 358, 366,

390 Gardens’ Bulletin, Singapore — XXV (1969-1971)

Hylophila, 106, 108.
cheangii 106.
lanceolata, 106.
mollis, 108.

Hyophila involuta, 337.

Hyphaene, species from upper

Guinea, 326.

baikieana, 3/2, 313, 324, 326,
331.

baronii, 313.

beccariana, 285.

benguellensis, 318.

congensis, 313, 317, 331.

coriacea, 283, 285, 287, 292,
301, 306,, 318.
var. minor, 285, 287.

crinita, 283, 284, 285, 286,
287-294, 313, 315, 317, 324,
326,328; 331,

dahomeensis, 313, 314, 317, 331.

dankaliensis var. haycockensis,
309.

dichotoma, 299, 301, 302, 303,
304, 306.

doreyi, 311, 317, 320, 322, 324,
331

gossweileri, 331.

guineensis, 311-318 passim,
319, 320. 324. 322,323, 324,
326, 328, 331, 333.

indica, 300, 301, 303.

macrocarpa 301, 328.

macrosperma, 313, 314, 317,
324,..925,;0 3205 e826, oa.

multiformis, 300, 301, 302, 334.

natalensis, 283-290 passim, 291,
292, 293, 294, 295, 296, 324,
326.

occidentalis, 333.

petersiana, 288.

reptans. 299, 309.

schatan, 283, 289.

sinaitica, 299, 302, 306, 307,
309, 316.

taprobanica, 299, 302, 304, 305,
0

308.
thebaica, 283-328 passim, 329,
330, 331, 332, 3a 324.
togoensis, 285, 313, 314, 316, 317,
324, 326, 333.
tuleyana, 313, 327, 333.
turbinata, 283.
ventricosa, 287, 288, 289.

Isopterygium, spp. listed, 341.

Laportea, by 2o93n4 185 115.
species, excluded, 175, 176.
TRIBUS Urereae, 114.
sect. FLEURYA, 111, 113, 114,
1150126. Wa5, fot aaa:

sect. LAPORTEA, 111, 114,
115, 145.

sect. SCLEPSION, 115.

aestuans, 114, 117, 128, 163, 164,
165.

agusanensis, 40, 42.

alatipes, 113, 114, 117; 124, 725,
526, °T33:

amberana, 11340814, 113, 431,
132, 135,137.

amplissima, 84,

anacardioides, 42.

annamica, 11, 12.

ardens, 36, 39.

armata, 139, 141.

basirotunda, 34.

batanensis, 22.

bathiei, 164.

brunnea, 11, 12.

bulbifera, 113, 114, 117, 121,
122, 145.

caffra, 112, 113, 117, 155, 156.

canadensis, 113, 114, 115, 117,
118, 719, 120, 121, 123.

_chingiana, 32, 34.

conduplicata, 30, 32.
contracta, 95.
corallodesme, 53.
cordata, 58, 60.
cordatifolia, 58, 60.
costata, 11.
var. contracta, 95,
crassifolia, 18.
crenulata, 36, 71.
var. luzonensis, 40, 42.
var. nervosa, 69.
var. nitida, 47, 50, 51.
cuneata, 112, 113, 114, 116,
170, /71.
decumana, 112, 113, 114, 116,
139, 1/40.
densiflora, 22, 24.
dielsii, 121, 123.
diffusa, .21,. 22.
disepala, 114, 117, 126, 127.
divaricata, 118, 120.
elliptica, 16.
evitata, 121.
floribunda, 112, 114, 116, 173,
174.
forrestri, 142, 145.
gaudichaudiana auct., 58.
gaudichaudiana Wedd., 21, 22.
gigantea, 39.
gigas, 7, 64, 66.
giraldiana, 142, 145.
glabra, 47, 50, 51.
gracilipes, 11, 12.
grossa, 112, 117, 159, 159, 160.
grossedentata, 142, 145.
hainanensis, 11, 12.
harveyi, 88, 90.
humbertii, 131, 133, 135.
humilis, 139, 141.
integrifolia, 36, 39.
interrupta, 113-116, 145, 145,
147, 154, 166.
kotoensis, 22.
kusaiana, 71.
lanaensis, 11, 12.
lanceolata, 117, 161, /62.
latifolia, 39, 71, 72.
latilanceolata, 40, 42.
laxiflora, 42, 45.
leytensis, 21, 22.
longifolia, 79, 81.
longispica, 142, 145.
luzonensis, 40, 42.
macrostachya, 112, 113, 114, 116,
142, 143, 144.
mammosisetosa, 47, 50, 51.

Index

Laportea (cont.)

manombensis, 131, 133.

meyeniana, 21.

microstigma, 82, 84.

milnei, 88, 90.

mindanaensis, 22.

mirabilis, 90.

monticola, 30, 32.

mooreana, 112, 113, 117, 157,
158,.159, 161.

moroides, 55, 56.

murrayana, 45.

oblanceolata, 14.

oleracea, 121, 123.

ovalifolia, 112, 113, 114,
148, 149, 150, 154, 163.

peduncularis, 112, 117, 152,
E53, 155, 159.

pedunculata, 139.

peltata auct., 42.

peltata Gaud. ex Decne., 44,
55, q

pendula, 28, 30.

Hermernt, 12, dia: 114. 116, 135,
136.

photinophylla, 60.

platyphylla, 22.

pterostigma, 21, 22.
forma subglabra, 22.

pustulosa, 36, 39, 40.

rechingeri, 74.

rigidifolia, 26.

ruderalis, 112, 113, 114, 116.
166, 167, 168, 172.

saipanensis, 71, 74.

salomonensis, 79, 81.

schlechteri, 76, 79.

schomburgkii var. vesicolor,
Ae

110,

septentrionalis, 116, 133, 134.
sessiliflora, 95.
sinensis, 121, 123.
sinuata, 36.
stimulans, 11.
var. contracta, 95.
subclausa, 28.
subglabra, 22.
subpeltata, 24.
sumatrana, 11, 12.
terminalis, 121, 124.
ternatensis, 47, 50, 51.
thorelii, 11, 12.
torricellensis, 51.
triplinervia, 30, 32.
urentissima, 32, 34.
venosa, 30, 32.
violacea. 112, 114, 117, 128, 129,
130, 131.
vitiensis, 62, 64.
vitifolia, 129, 131.
vriesiana, 42, 45.
warburghii, 79, 81.
weddellii, 117, 133, 137, 138.

Lejeunea (?) pilifera, 344, 351, 352.
Lepidozia cf. quadridens, 342.
Leptolejeunea, spp. listed, 345.

Leucobryum, spp. listed, 336.
Leucogyrophana, 356, 365.

391

Leucolejeunea xanthocarpa, 345.
Leucoloma molle, 336.
Liverworts, new spp., 346-353.
Lopholejeunea subfusca, 344.
Lopidium trichocladon, 339.

Lowia, 239.
longiflora, 239, 243, 245, 246.
maxillarioides, 245.

Lowiaceae, 239, 240.

Lowioideae, 239.

Macromitrium cumingii, 337.
reinwardtii, 337.

Marchantia acaulis, 341.
Mastigophora diclados, 342.
Meiorganaceae, 355.

Meiorganum, 355, 356, 357, 358,
360, 362, 376.
neocaledonicum, 357, 362, 363,
364.

Meiothecium hamatum, 340.
Melosira, spp. listed, 180.
Merismopedia sp., 184.
Meruliaceae, 359.

Merulioid fungi in Malaysia, key
to, 355-361.

Meruliopsis, 359, 360, 371.
Merulius, 355, 356, 357, 358, 359,
360.

Merulius, 355-360, 365.
index of species attributed to,
378-381.
agathidis, 356, 357, 358, 360,
364, 365.
byssoideus, 355, 356, 369.
castaneus, 369.
chlorina, 369.
consimilis, 366.
corium, 355, 359, 360, 370.
crassus, 355.
eurocephalus, 360, 366, 373.
fuscescens, 355.
glaucinus, 371.
var. lutescens, 371.
hirtellus, 371.
insignis, 366, 367.
irpicoides, 356, 367, 368, 369.
lacrymans, 356, 367.
luridochraceus, 356, 369.
molluscus, 355, 356.
montanus, 355.
nothofagi, 372, 373.
pilosus, 372.
pinastri, 356.
pseudolacrymans, 366.
pseudomolluscus, 355.

392 Gardens’ Bulletin, Singapore — XXV_ (1969-1971)

Merulius (cont.)

purpurascens, 360, 370, 371.
var. flavido-albus, 360, 370.

ravenelii, 371.

rubicundus, 371.

rubrotremellosus, 371.

rufus, 356, 371, 372, 373.

strigosozonatus, 359, 374.

taxicolus, 359.

tignicolus, 356.

tremellosus, 355, 356, 372, 373.

versicolor, 360, 373.

Metadina, 247, 248, 249, 253.
multifolia, 250, 25/.

Metzgeria furcata, 341.
Micrasterias, spp. listed, 184.
Microlejeunea culcullata, 345.

Mitragyna, 248.
speciosa, 248.

Mniodendron divaricatum, 337.
Mosses, Fraser’s Hill, 336-341.
Musa, 239, 240, 241, 242.
Musaceae, 239, 240.

Nanocnide, 114.

Nauclea, 247.
affinis, 265.
chalmersii, 265.
coadunata, 250.
dahlii, 267.
forsteri Seem, 263.
Forsterii Hav., 263.
glabra, 270.
gordoniana, 272.
gracilis, 265.
hagenii, 267, 270.
lanceolata, 265.
obversifolia, 272.
orientalis, 263.
papuana, 270.
philippinensis, 265.
rotundifolia, 263.
schlechteri, 265.
speciosa, 248.
subditus, 252.
tenuiflora, 252.
tenuis, 265.
trichotoma, 2A9.

Naucleae, 247.
key to genera, 248.

Navicula radiosa var. minutis-
sima, 180,

Neonauclea, 247, 248, 253.
species, key to, 256-259.
acuminata, 254, 257, 258, 275,

278, 279.

brassii, 254, 257, 258, 259, 276,
277.

cardiophylla, 263.

chalmersii, 254, 256, 257, 258,
265, 266, 267.

clemensii, 254, 256, 258, 259,
261.

dahlii, 267.

forsteri, 254, 255, 256-259,
263, 264.

gordoniana, 253, 254, 257, 259,
271, 272.
hagenii, 253, 254, 257, 258, 267,
269, 270.
spp. hagenii, 269.
spp. papuana, 257, 258, 269,
270.

lanceolata, 254, 257, 258, 265,
268.

maluensis, 254, 257, 259, 276,
277.

moluccana, 270.

obversifolia, 253, 254, 257, 258,
Biz, 2273.

papuana, 270.

perspicuinervia, 254, 257, 259,
279, 280.

schlechteri, 265.

solomonensis, 254, 257, 259,
274, 275. |

versteeghii, 254, 256, 258, 260,
262.

Nephrocytium agardhianum, 182.
Netrium digitus, 183.

Nitzschia palea, 182.

Oblixilis, 115.
Octoblepharum albidum, 336.
Oedogonium sp., 183.

Orchidantha, 239, 240, 241.
species, key to, 243.
borneensis, 239, 243, 246.
calcarea, 239, 245.
fimbriata, 240, 243, 244, 246.
laotica, 240, 243, 246.
longiflora, 239, 240, 243, 245.

maxillarioides, 240, 243, 244,
245.

siamensis, 243, 244.

Oscillatoria sp., 184.

Index

Papillaria fuscescens, 338.

Paxillus, 356, 357, 358, 360.
panuoides, 357, 358, 366.

Pediastrum, spp. listed, 182.
Phacus, spp. listed, 182.

Phaeophlebia, 359, 360, 374.
strigosozonata, 374, 375.

Philonotis hastata, 337.

Phlebia castanea, 372.
merulioides, 372.
rufa, 372.

Pinnatella, spp. listed, 339.
Pinnularia, spp. listed, 180.

Pleurotaenium, spp. listed, 183.

Protamomum, 239.
maxillarioides, 239, 24S.

Pilea mooreana, 157.
Podoserpula, 358.
Poikilospermum, 3, 78.
Pogonatum, spp. listed, 341.
Polyporus eurocephalus, 366.

Poria, 360.
taxicoloa, 371.

Pterobryopsis, spp. listed, 338.
Ptychanthus striatus, 345.
Punctularia, 359, 374.

Pycnolejeunea, spp. listed, 345.

Racopilum schmidii, 338.
Radula nymanii, 342.
Rhizogonium spiniforme, 337.

Rhodobryum giganteum, 337.

Rynchostegium celebicum, 339.

Saccolabium micranthum, 10S,
106.
fissum, 106.
humile, 105, 106.

Sarcanthus, 105.

Sarcocephalus, 247, 248, 250.
coadunata, 250.
tenuiflorus, 250, 25/, 252.

Scenedesmus, spp. listed, 183.
Sceptrocnide, 111, 114, 115.
Schychowskia, 115.

393

Scitamineae, 239, 240, 242.
Selenastrum bibraianum, 182.

Serpula, 355, 356, 357, 365, 369.
eurocephala, 366.

Smitinanda, 105.

micrantha, 106.

humilis, 106.
Sphaerocystis schroeteri, 182.
Spirogyra sp., 183.
Spirulina gomontii, 184.
Spondylosium, spp. listed, 184.
Staurastrum, spp. listed, 184.
Stenolejeunea apiculata, 345.

Stephygyne parvifolia, 248.
speciosa, 248.

Stereum spadiceum, 376.

Symphysodontella cylindracea,
338.

Syrrhopodon, spp. listed, 336.
fraserianum, 346, 347.

Taxilejeunea, spp. listed, 345.
Taxithelium, spp. listed, 341.
Tetraedron sp. 182.

Tetrallantos lagerheimii, 183.
Thrixspermum, 105.

Thuidium meyenianum, 339.
Thyridium, spp. listed, 336.
Thysananthus spathulistipus, 346.
Trabecularia, 365.
Trachelomonas, spp. listed, 182.
Trachyloma indicum, 338.
Trichoglottis appendiculifera, 108.

Trichostelium, spp. listed, 340.
fruticolum, 348, 349.

Trismegistia, spp. listed, 340.
Tuyamaella angulistipa, 346.
Uncaria, 248.

Urera, 1, 6.
amberana, 131.
amplissima, 84, 86.
commersoniana, 95.
excelsa, 64, 66.
leichardiana, 60, 62.
rotundifolia, 64, 66.

394 Gardens’ Bulletin, Singapore — X XV (1969-1971)

Urtica, 112.

aestuans, 164.
affinis, 145, 149.
ardens, 36, 40.
bulbifera, 121, 123.
canadensis, 118.
capitata, 145.
caravellana, 164.
coerulea, 96.
contracta, 95.
cordata, 164.
crenulata, 36.
cuneata, 170, 172.
decumana, 139.

divaricata, 118, 164.

divergens, 164.
evitata, 121.
ferox? * 2.
fruticosa, 11, 12.
gaudichaudii, 145.
gigantia, 95.
hirsuta, 164.

interrupta, 145, 148.

javanica, 145, 148.
latifolia, 164.
lomatocarpa, 145.
mitis, 152.
nemorosa, 164.
obnoxia, 36.
ovalifolia, 84, 86.
paniculata, 167.
peduncularis, 152.
peltata, 42, 45, 56.
petiolata, 164.
pulus, 36.
pustulata, 118.
ruderalis, 167.
rumphii, 139.
schimperiana, 164.

sessiliflora, 145.
sinuata, 36, 40.
spicigera, 145.
stimulans, 11, 12.
var. obtusa, 11.
tuberculata, 164.
umbellata, 21.

Urticaceae, 1, 111.

TRIBUS Urereae, 6, 114.

Urticastrum, 7, 115.

alatipes, 124.
amplissimum, 84.
bulbiferum, 121.
costatum, 11.
decumanum, 139.
divaricatum, 118.
evitatum, 121.
gaudichaudianum, 21.
gigas, 64.
harveyi, 88.
laxiflorum, 42.
microstigma, 82.
milnei, 88.
moroides, 55.
oleraceum, 121.
peltatum, 42.
Photinophyllum, 60.
pterostigma, 21.
sinuatum, 36.
stimulans, 11.
vriesianum, 42.

Xanthidium urniforme, 184.
Xylomizon, 365. —
Xylophagus, 365.

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