Flora Malesiana

Myristicaceae

Series I — Seed Plants

Cover: Myristica cinnamomea King

Photograph W.J.J.O. de Wilde

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H.P. Nooteboom W.J.J.O. de Wilde
L.G. Saw

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Flora Malesiana

Series I — Seed Plants

Volume 14 — 2000

Myristicaceae

(W. J. J.O. de Wilde)

THE LuESTHER T. MERTZ LIBRARY

THE NEW YORK BOTANICAL GARDEN

ISBN 90-71236-47-1
All rights reserved

© 2000 Foundation Flora Malesiana

No part of the material protected by this copyright notice may be reproduced or utilized in any
form or by any means, electronic or mechanical, including photocopying, recording, or by any
information storage and retrieval system, without written permission from the copyright owner.

ABSTRACT

Flora Malesiana. Series I, Volume 14 (2000) iv + 1-634, by W.J.J.O. de Wilde
(edited by P.F. Stevens), published by the Nationaal Herbarium Nederland, Universiteit
Leiden branch, The Netherlands, under the auspices of Foundation Flora Malesiana.

ISBN 90-71236-47-1

Contains the taxonomic revision of one family, Myristicaceae, for Malesia, i.e. the area
covering the countries Indonesia, Malaysia, Brunei Darussalam, Singapore, the Philip-
pines, and Papua New Guinea.

W.J.J.O. de Wilde, Myristicaceae, pp. 1-622.

A pantropical family of trees, in Malesia represented by six genera: Endocomia
(4 species), Gymnacranthera (6), Horsfieldia (97), Knema (75, only one species in New
Guinea), Myristica (152, of which the majority endemic to New Guinea), and Para-
myristica (1, Papua New Guinea). Altogether there are 335 species of the family in
the Malesian area. Some species are of economic importance, for instance Myristica
fragrans, nutmeg.

The general part consists of 28 pages and also includes paragraphs on vegetative
anatomy by P. Baas & J. Koster, on palynology by R.W.J.M. van der Ham, and on
phytochemistry and chemotaxonomy by R. Hegnauer.

Myristicaceae are dioecious. In addition to the general keys, mainly based on male
specimens, also regional keys are given for the larger genera Horsfieldia, Knema, and
Myristica, based on female (fruiting) specimens.

For each species full references, synonymy, keys to infraspecific taxa, diagnostic
descriptions, field-notes, distribution, and annotations regarding relationships or dif-
ferences with resembling species are presented. Genera and species are arranged alpha-
betically.

This treatment is illustrated with 94 line drawings (many full-page), 6 maps, and
4 pages with colour photographs* (inserted after p. 8).

Index to scientific plant names of taxa treated in this volume (accepted names and
synonyms) on pp. 623-632.

Lists of revised families in Flora Malesiana on pp. 633-634.

*) The grant of the Dr. Hendrik Muller’s Vaderlandsch Fonds, The Hague, for the repro-
duction and inclusion of the colour photographs, is gratefully acknowledged.

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Flora Malesiana, Series I, Volume 14 (2000) 1— 632

MYRISTICACEAE
(W.J.J.O. de Wilde, Leiden, The Netherlands)!

Myristicaceae R.Br., Prodr. (1810) 339, ‘Myristiceae’; Warb., Monographie der Myr-
isticaceae, Nova Acta Acad. Caes. Leop.-Carol. 68 (1897) 1—680; J. Sinclair, Gard. Bull.
Sing. 16 (1958) 205—470, pl. I-XIV; Whitmore, Tree Fl. Malaya | (1972) 315-345; W.J.
de Wilde, Blumea 30 (1984) 173-196; 39 (1994) 341—350: Beitr. Biol. Pflanzen 66 (“1991’,
1992) 95-125; in Soepadmo & Saw (eds.), Tree Fl. Sabah & Sarawak 3 (2000) 335-473;
Kiihn & Kubitzki in Kubitzki et al., Fam. & Genera Vascular Plants 2 (1993) 457-467
(with extensive literature references). — Type genus: Myristica Gronov.

Dioecious or monoecious (Endocomia) monopodial shrubs or trees with aromatic oil-
cells, (2—)5—15(—40) m high (Virola sessilis, Brazil, 30 cm high); stilt-roots sometimes
present; branching whorled, the crown often pyramidal. Bark and wood exuding a watery
pink or red sap on cutting. Wood white or soon brownish on cutting, soft; rays narrow, tan-
niniferous tubes present; growth rings usually distinct and visible to the naked eye. Twigs
tomentose, glabrescent. Hairs uniseriate, often with armed cells, appearing as stellate or
dendroid. Leaves simple, entire, alternate, usually distichous in plagiotropic shoots, exstipu-
late, often with indumentum on both surfaces when young, glabrescent, or remaining on
the lower surface; lateral nerves distinct, interarching at the margins; reticulations some-
times faint; vernation conduplicate (Asian genera); dark dots (corky hair bases) on lower
leaf surface present or absent; glands absent. /nflorescences from foliar axils to ramiflorous
(cauline), pedunculate, compound or not, with or without bracts at base, the flowers vari-
ably arranged at the ends of the branches, or a short, woody (sub)sessile simple or 2—4-fid
scar-covered wart- or worm-like brachyblast with the flowers in (sub)umbels at the end;
bracts caducous. Female inflorescences similar to male, usually less branched. Flowers
unisexual, mostly pedicellate; bracteole present or absent. Perianth gamophyllous, glo-
bose to rotate, buds 2- (Horsfieldia, p.p.) to 5-lobed (Endocomia, p.p.), cleft to variable
depths, lobes little or much recurved; the female usually urceolate, larger, more swollen
than the male; disc rarely present. Androecium with synandrium sessile or stalked (andro-
phore), in Myristica mostly with a short sterile apex; anthers dorsally adnate to the col-
umn and laterally to each other, or free, in Knema stellately arranged around a disc,
2—45 in number, ellipsoid to linear, extrorse, opening by longitudinal slits, 2-loculed,
tetrasporangiate; immature pollen sacs septate. Ovary monocarpellate, sessile, superior,
ovoid; style absent or short, stigma 2-lobed or rarely peltate with few to many laciniations;
ovule 1, mostly anatropous, + basal. Fruits (sub)globose to oblong, rarely transversely
elongate, dehiscing by a longitudinal circumferential suture into 2 valves, rarely indehiscent;
pericarp leathery, carnose or subligneous, glabrous or tomentose. Avril (orange-)red (or
yellow?), completely covering the seed, laciniate to various depths or sometimes (sub)entire,
attached to the base of the seed between the hilum and the micropyle. Seeds with testa of
three layers; albumen hard, mostly ruminate, containing oil, sometimes starch. Embryo
small, near the base of the seed or a little above; cotyledons connate at base (peltate or
cup-shaped) or free; radicle basal, cylindric.

1) With contributions by R.W.J.M. van der Ham (palynology), R. Hegnauer (phytochemistry &
chemotaxonomy), J. Koster and P. Baas (vegetative anatomy). Most of the original drawings are
by J.H. van Os and some by R. van Crevel.

ho

Flora Malesiana, Ser. I, Vol. 14 (2000)

DISTRIBUTION

Pantropical with c. 500 species in 20 genera, more or less equally distributed over and
restricted to the three main continental areas: 8 genera (with few species) in Africa, of
which 5 in Africa proper and 3 in Madagascar, 6 genera in America, and 6 in Southeast
Asia, mainly Malesia.

The largest genera are Virola (c. 50 species) in America, and Horsfieldia (c. 100 spe-
cies), Knema (c. 90 species), and Myristica (c. 170 species) in Asia. The latter three, to-
gether with Endocomia and Gymnacranthera, have widespread distributions; Endocomia
occurs from S China to New Guinea (Map 1), Gymnacranthera from S Peninsular Thai-
land to New Guinea (Map 2); Horsfieldia occurs from Sri Lanka and S China east to the
Solomon Islands and N Australia, with centres of species richness in W Malesia, main-
ly Borneo and New Guinea (Map 3); Knema has a distribution from India to W New
Guinea. with a centre of diversity in W Malesia, mainly Borneo (Map 4);the genus
Myristica has the largest distribution, from India to N Australia and far into the Pacific,
with centres of speciation in W Malesia and, particularly, New Guinea (100 species)
(Map 5). Paramyristica (1 species) is endemic to Papua New Guinea (Map 6).

In the present treatment the now official name Papua Barat has been used instead of
Irian Jaya.

HABITAT

Low or medium (rarely canopy) trees in various types of primary lowland rain forest,
including kerangas and marshy forest. Some species of Horsfieldia (in New Guinea) and
several of Knema (e.g., Mt Kinabalu area) and Myristica (in New Guinea) occur in
montane forest. Occasionally species are ‘sciophilous nomads’ (fast growing, shade toler-
ant), notably some Horsfieldias in New Guinea, and few are found in secondary forest.
Sometimes Myristicaceae constitute a considerable component of the forest, especially
of the middle storey of the lowland rain forest, but they are not gregarious.

According to Koster & Baas (1981) leaf anatomical characters are xeromorphic, which
is unexpected in view of the mesic ecology of recent Myristicaceae (see Vegetative
Anatomy’ ).

ECOLOGY

Pollination & flower biology — Flowering and fruiting generally occurs throughout the
year. The usually + carnose, yellow or brown, inside creamy, pink, or red flowers of
several genera have repeatedly been reported as being fragrant, e.g. Horsfieldia irya and
Myristica fragrans. Anthesis presumably is mainly nocturnal, and small beetles may
effect pollination; nectar is not reported for any species. Male plants of Myristica pro-
duce over 50 times as many flowers as do females (Armstrong & Drummond 1986;
Armstrong & Tucker 1986; Armstrong & Irvine 1989). Myristica subalulata and some
related species are myrmecophilous, the ants possibly involved in pollination (De Wilde
1998). Besides the coloured inside of the perianth, in Knema also the staminal disc may
be brightly (purple-red) coloured, the contrast with the creamy-white pollen possibly
heightening the attraction of pollinators.

Dispersal — The brown-black seeds contrasting with the orange or red aril and the inner
surface of the (red, pink, or white) opened pericarp attract birds and suggest bird disper-

De Wilde — Myristicaceae 3

SS Pe

Se

Map 1. Distribution of the genus Endocomia W.J. de Wilde
(see also p. 29).

Map 2. Distribution of the genus Gymnacranthera (A.DC.) Warb.
(see also p. 40).

Flora Malesiana, Ser. I, Vol. 14 (2000)

Horsfieldia

I sect. Horsfieldia
II sect. /rya
III sect. Pyrrhosa

Q

Hl . 2 =

al Wk: Mite a

as Bie

e ** opener ee : ;
o * : o

xxxxx H. crassifolia

Map 3. Distribution of the genus Horsfieldia Willd.
I = sect. Horsfieldia; Il = sect. Irya, east of Wallace’s Line (broken line); III = sect. Pyrrhosa, west
of Wallace’s Line. Distribution of H. irya is indicated by a dotted line, that of H. crassifolia by a line
of asterisks. Crosses indicate the approximate areas of species with a number of perianth lobes
deviating from that of section areas II and III (see p. 55 for further information).

Map 4. Distribution of the genus Knema Lour.
(see also p. 222).

De Wilde — Myristicaceae 5

Map 5. Distribution of the genus Myristica Gronov.
The inlet indicates the eastern part of the total distribution area, with Pacific islands and Fiji
(see also p. 359).

3 fang
SS

Map 6. Distribution of the genus Paramyristica W.J. de Wilde
(see also p. 619).

6 Flora Malesiana, Ser. I, Vol. 14 (2000)

sal (Howe & Vande Kerckhove 1980), in Asia by fruit pigeons and doves, hornbills, and
birds of paradise (Sinclair 1958). Monkeys and rodents may disperse fruits or seeds as
well. Dissemination by water may occur in Horsfieldia wallichii, of which the seeds
float because of air trapped between aril and seed, and in Horsfieldia irya, a widespread
riverine species with cavities in the endosperm. The seeds of the marsh nutmeg, Myristica
elliptica, are reported to float, also when the aril is missing.

Germination & seedling — Seeds remain viable for a restricted period only, a few weeks,
and germinate only in a damp, shady environment and therefore the natural reintegration
of Myristicaceae in secondary forest is impossible. Germination is (mostly) hypogeal.
The cotyledons remain within the testa, the taproot and hypocotyl emerge, the shoot is
erect, initially with reduced leaves (cataphylls), mostly borne spirally: the Horsfieldia
type (subtype) of seedling (De Vogel 1979), a common type in tropical woody dicotyle-
dons.

References: Armstrong, J.E. & B.A. Drummond, Biotropica 18 (1986) 32-38. — Armstrong,
J.E. & A.K. Irvine, Amer. J. Bot. 76 (1989) 74-85, 86-94. — Armstrong, J.E. & S.C. Tucker,
Floral development in Myristica (Myristicaceae). Amer. J. Bot. 73 (1986) 1131-1143. — De Vogel,
E. F.,, Seedlings of Dicotyledons (1979) 1-203. Pudoc, Wageningen. — De Wilde, W. J.J.O., Blumea
43 (1998) 165-182. — Howe, H.F. & G.A. Vande Kerckhove, Science 210 (1980) 925-926. —
Sinclair, J., A revision of the Malayan Myristicaceae. Gard. Bull. Sing. 16 (1958) 205—472.

TAXONOMY

The family Myristicaceae is homogeneous and clearly belongs (also phytochemically)
within the Magnoliales. Phylogenetic analysis of the genera revealed that the family is
monophyletic, of African origin (Sauquet 1998), and seemingly most related to the An-
nonaceae, mainly through the ruminate endosperm. Canellaceae have been suggested
to be allied through the monadelphous androecium (Wilson & Maculans 1967), but ac-
cording to a recent cladistic analysis of all families (APG 1998) this family is as yet not
properly placed as it falls beyond the recognized basal orders. The foliage is generally
strongly reminiscent of Annonaceae. The 3-lobed perianth reminds of Lauraceae and
Annonaceae. Myristicaceae are distinguished, however, by their unisexual flowers with
uniseriate perianth, and monocarpellate 1!-ovuled female flowers.

Within the family the relationship of the genera is unclear, and initially one single
genus, Myristica (divided into sections), was recognized until Warburg (1897) divided it
into several genera. The genera of Madagascar possibly retain the most primitive char-
acters, viz. pollen morphological, not or less consolidated stamens, and a little devel-
oped aril.

Within the larger Malesian genera, Horsfieldia, Knema, and Myristica, subgenera or
series have been recognized (Warburg 1897; Uphof 1959; Sinclair 1968; De Wilde 1979),
but as explained under these genera, the distinctions are not sharp and only allow for
informal grouping of species, largely reflected in the keys to the species.

Practical taxonomic notes — All members of the family Myristicaceae can be recog-
nized in the field by their general habit, i.e., a slender bole with monopodial crown, the
branches more or less verticillate and tiered, and the rather long exstipulate distichous

De Wilde — Myristicaceae 7

leaves like those of the Annonaceae. The latter family differs in its flower structure,
fruits, and lack of the red exudate of the cut bark. Further differences are obvious in the
transversely cut twig where the bark has radiating parenchyma in the Annonaceae, and
also the pollen is different. The genera of Myristicaceae can be told apart on vegetative
characters only with experience, and flower, inflorescence, and fruit characters are nec-
essary for a definite identification. Useful characters are the non-striate, usually finely
lenticellate and granulate bark of the twigs in Gymnacranthera, or the dry leaves not
readily breaking into pieces in most species of Knema and Gymnacranthera (both have
reticulate sclerenchyma in the mesophyll). Certain leaf and wood anatomical characters
can be used in genera diagnoses. All six Malesian genera may reach timber size.

Since myristicaceous specimens either have male flowers, or female flowers and/or
fruits, keys have been constructed for both sexes, using also vegetative characters. How-
ever, specimens with only female flowers may be difficult to assign to a particular ge-
nus, and one should use both types of keys, paying particular attention to the inflores-
cence type.

References: APG (Angiosperm Phylogeny Group), An ordinal classification for the families of
flowering plants. Ann. Missouri Bot. Gard. 85 (1998) 531-553. — De Wilde, W. J. J.O., New ac-
count of the genus Knema (Myristicaceae). Blumea 25 (1979) 321—478; in E. Soepadmo & L.G.
Saw (eds.), Tree Flora of Sabah & Sarawak 3 (2000) 335-473. — Sauquet, H, Phylogénie des
Myristicaceae. Rapport de stage. Institut Agronomique, Paris-Grignon (1998). — Sinclair, J., The
genus Myristica in Malesia and outside Malesia. Gard. Bull. Sing. 18 (1968) 1-540. —
Uphof, J.C.Th., in: A. Engler & K. Prantl, Die natiirlichen Pflanzenfamilien ed. 2, 17a II (1959)
177-199. — Warburg, O., Monographie der Myristicaceae, Nova Acta Acad. Caes. Leop.-Carol. 68
(1897) 1-680. — Wilson, T. K. & L.M. Maculans, The morphology of the Myristicaceae. I. Flow-
ers of Myristica fragrans and M. malabarica. Amer. J. Bot. 54 (1967) 214—220.

VEGETATIVE MORPHOLOGY

Notes — 1) Sizes given in the descriptions always relate to (measurements in) the dry
state. When single measurements are given they indicate length. 2) Comprehensive dis-
cussions on the morphology of Asian Myristicaceae have been published by Sinclair
(i958. 1961, 1968).

Growth form — Asian Myristicaceae are always trees, though sometimes only a few
metres high. On germination the erect shoot, which initially develops into the orthotropic
main stem, carries a number of spirally arranged cataphylls, soon passing into normal
leaves. Growth is normally monopodial in flushes, and each season at the end of the new
flush the leaves are produced + crowded into a several-leaved pseudowhorl. The
plagiotropic lateral shoots hence are + whorled, + horizontal, and so are the main branches
on the stem. This growth form of the trees is according to the model of Massart (Hallé et
al. 1978; De Wilde 1992). In the plagiotropic shoots, usually in the herbarium speci-
mens, the leaves are generally (sub)distichous (in Malesia in a few species of Horsfieldia
phyllotaxis is spiral). In some species (e.g. Gymnacranthera ocellata, Paramyristica
sepicana), an apical bud with bud scales is formed, the latter leaving ring-shaped scars
at the base of the innovations. Buttresses are frequent, stilt-roots occasional. The mono-
podial crown often is + pyramidal in outline. Presence of stilt-roots and other character-
istics have been summarized in the field-notes of most species.

8 Flora Malesiana, Ser. I, Vol. 14 (2000)

Bark — The bark of the trunk is smooth or fissured (furrowed), scaly, or dippled, in
certain species of Myristica and Knema blackish and gritty (the penarahan arang group
of foresters). The inner bark is fibrous, red-brown. When cut the inner bark (and wood)
exude a clear, pale to intense dark red sap (kino), usually free flowing, and typical for the
family. The generally soft sapwood is pale , darkening brown-red on exposure; the heart-
wood often is dark-coloured; the core of old trees is commonly brittle, or reported as
being rotten. Field-notes on bark and wood characteristics have been given under most
species. Some photographs of bark have been published by Sinclair (1958).

Indumentum — Almost all Asian Myristicaceae have some sort of indumentum (of sparse
minute hairs to thickly woolly), composed of uniseriate hairs (see ‘Anatomy’) which
may be scale-like, stellate, or dendroid. Very often the hairs are rust-coloured and early
shed, but an indumentum is usually present on the sterile apical leaf bud, the very apex
of the twigs, inflorescences, and the flower buds; it may persist on the lower leaf sur-
face, dependent on the species. The length of the hairs, viz. short (0.1 mm) versus longer
(0.2 mm or more) is diagnostic. The indumentum of the fruits, if present, is always
diagnostic.

The indumentum in Myristicaceae is also used for the distinction of genera, as ex-
plained for Malesia below in ‘Vegetative anatomy’ by Koster and Baas. The hairs are
essentially uniseriate, but the cells may be branched to one or two sides, forming sessile
(scale-like) to long-dendroid hairs.

Twigs — The thickness (diameter) of the twigs, measured at the apex in the distal 10 cm,
and whether they are terete (as usual), (blunt) triangular, ridged (mostly at both sides in
between the insertion of the petiole), or + flattened, can be used as diagnostic characters.
The bark of the twigs may be longitudinally grooved (striate) to various degrees, and in
older twigs may become characteristically longitudinally cracked and later on flaking.
Colour of the bark of the twigs is usually some shade of brown, straw, or greyish (pale)
and contrasting with the dark drying colour of the petiole. Only when colours are con-
trasting it is mentioned in the descriptions, being characteristic for certain species, espe-
cially in Horsfieldia, and for the whole genus Endocomia. The bark of the twigs may be
lenticellate to various degrees, according to the species. In Gymnacranthera the twigs
are always + flattened and strongly lenticellate; in Knema and Endocomia lenticels are
(almost) absent. Some New Guinean Myristicas are characteristically myrmecophilous
with part of the twigs thickened and ant-inhabited (De Wilde 1998).

Leaves — The leaves are simple, exstipulate, pinninerved, and spirally inserted (dis-
persed) on orthotropic axes. However, in the plagiotropic fertile twigs, i.e. in herbarium
specimens, the leaves are usually distichous; rarely the phyllotaxis is spiral, as in some
species of Horsfieldia. The blade varies between elliptic to lanceolate, often being broadest
at or slightly above, sometimes below the middle. The margin is occasionally revolute
on drying, and only then it is mentioned in the descriptions. The lower leaf surface
usually is pale and may be papillose or not, or covered with alveolar material (Koster &
Baas 1981, 1982), characteristic for Knema or, e.g., Horsfieldia iryaghedhi and certain
species of Myristica. The indumentum may be persistent on the lower leaf surface, but in
most species of all genera it is early falling. Very characteristic for certain species in

(For legends, see page 622)

Photo 1 — Gymnacranthera bancana (Miq.) J. Sinclair

am ’ abies...
Photo 2 — Gymnacranthera forbesii (King) Warb. var. forbesii

(For legends, see page 622)

Photo 3 — Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. farquhariana

Photo 4 — Myristica fatua Houtt. subsp. fatua

(For legends, see page 622)

Photo 5 — Knema glauca (Blume) Petermann var. glauca

Photo 6 — Knema kinabaluensis J. Sinclair

(For legends, see page 622)

Photo 7 — Knema sumatrana (Blume) W.J. de Wilde

Photo 8 — Horsfieldia wallichii Warb. Photo 9 — Myristica iners Blume

De Wilde — Myristicaceae 9

Horsfieldia, Knema, and Myristica is the presence or absence of dark-coloured, red-
brown or blackish dots and/or dashes, especially on the lower leaf surface, i.e., corky
warts developed from the bases of fallen hairs, visible with a strong lens. Much finer
dark spots representing tannin-conglomerations are often present. The presence or ab-
sence of dots (and dashes) is important for species distinction and for this purpose, to a
lesser extent, the presence or absence of microscopic papillation on the lower leaf sur-
face is used (to be seen with a magnification of x 60). In general, one should always
inspect the lower leaf surface when determining a myristicaceous specimen. For the dis-
tinction of the many similar species of Myristica in New Guinea the size of the leaf
blades is used; as arbitrary demarcation smaller or larger than about 15 cm is chosen for
the leaf length. Whether the midrib and lateral nerves are raised or sunken above (they
are always raised beneath) are useful taxonomic characteristics, as is the number of
lateral nerves. The distinctness of the veins connecting the laterals (in the descriptions
‘lines of interarching’) near the blade margin, as well as the nature of the tertiary vena-
tion (in the descriptions simply ‘venation’) are of taxonomic importance at the species
level. The size of the ultimate areoles formed by the veinlets is important for the distinc-
tion of some Knema species. In some species of all genera the colour of the blade on
drying is used for species delimitation, i.e. green in Knema viridis. The angle of the lat-
eral nerves to the midrib (in the descriptions indicated by degrees) may be diagnostic.

The sterile apical leaf bud, of a typical elongate-conical shape, has a characteristic
indumentum (hairs always appressed in Myristica) and more or less characteristic shape
and size. It consists of a single leaf only, and is present and visible as soon as the previ-
ous leaf in the flush has developed and expanded. In Asian species the vernation is
conduplicate. In a few species, especially those from higher elevations, some bud scales
may be present on the apical bud which ends the flush, and this is also rather character-
istic for lowland taxa such as Paramyristica and some Myristica and Gymnacranthera
species, e.g. G. ocellata, where these bud scales leave two distichous rows of closely set
scars at the transition between innovations. In species of a (presumably) more or less
seasonal environment (drought, or cold in the mountains), + ellipsoid or ovoid, sterile or
fertile (inflorescence) buds, normally 10 mm long or much less, composed of several
cataphylls, can be found axillary to leaves; in these buds the first two scales are minute
and essentially placed transversely and opposite (De Wilde 1992), as is common in di-
cotyledons.

References: De Wilde, W.J.J.O., The genera of Myristicaceae as distinguished by their inflores-
cences, and the description of a new genus, Bicuiba. Beitr. Biol. der Pflanzen 66 (*1991°, 1992) 95—
125; Census of Myristica (Myristicaceae) in New Guinea anno 1994. Blumea 40 (1995) 237-344.
— Hallé, F, et al., Tropical trees and forests. An architectural analysis. Springer, Berlin etc. (1978).
— Koster, J. & P. Baas, Comparative leaf anatomy of the Asiatic Myristicaceae. Blumea 27 (1981)
115-173; Alveolar material in the Myristicaceae. Linn. Soc. Symp. Series, No. 10, Suppl. (1982)
131-138. — Sinclair, J., Gard. Bull. Sing. 16 (1958) 205; 18 (1961) 102-327; 33 (1968) 1-540.

REPRODUCTIVE MORPHOLOGY

Inflorescences — Myristicaceae are practically always dioecious, except Endocomia,
which is monoecious, female flowers being mixed within the more numerous male flowers
in each paniculate inflorescence. The inflorescences are useful for the recognition of the

10 Flora Malesiana, Ser. I, Vol. 14 (2000)

genera; in detail, inflorescences are also characteristic of species (see De Wilde 1992).
They are always axillary (rarely somewhat supra-axillary) amongst or below the leaves,
and provided with bracts. They are polythetic, which means that their branches are never
terminated by a flower. The male inflorescences are often larger and with many more
flowers than the female, and show more interspecific differences. Endocomia, Gymna-
cranthera, Horsfieldia, Paramyristica, and part of Myristica have non-woody, panicle-
like inflorescences of short duration, while Knema and Myristica, partly, have woody,
condensed, knob-like, scar-covered brachyblasts producing at the apex flowers over
several seasons. These two types of inflorescence belong to basically different types, a
single and a plural (multiple) type (De Wilde 1992):

The architecture of the basic single type of inflorescence corresponds with the typical
mode of vegetative branching in the family. This type is axillary, compound, provided
with a smooth, non scar-covered, common peduncle; the first lateral branches are op-
posite, but with branches higher up essentially dispersed. The knob-like inflorescences
of Knema, and those of Myristica, partly, can be regarded as derived from these by re-
duction of branching and clustering of flowers. The superficially similar paniculate in-
florescences of the remaining genera appear to be derived from a number of the basic
type inflorescences, arranged in a way again comparable to the mode of vegetative branch-
ing.

In the multiple-type inflorescence the common peduncle is always provided with the
scars of basal prophylls. Clustering of the flowers into flower heads or (sub)umbels adds
to the variation in appearance of the inflorescences, but the presence or absence of scars
of prophylls at and towards the base of the main peduncle is an essential and easily seen
criterion. Knema and Myristica (both those with knob-like as well as panicle-like inflo-
rescences) have the single type, Endocomia (partly), Horsfieldia, and Gymnacranthera
have the plural type. In Paramyristica the inflorescences are essentially as in Myristica,
but they are panicle-like and arranged in a short-shoot, ending in a vegetative bud.

The two highly distinctive forms of inflorescences in the genus Myristica, discrimi-
nating the two sections Myristica and Fatua, both belong to the single type; that of sect.
Fatua, as that of Knema, being a strongly condensed form of the panicle-like inflores-
cences of sect. Myristica. As could be expected, there are quite a number of intermediate
forms in Myristica.

Fig. 1. Schematic representation of male inflorescences in Myristicaceae. — a: single type, of
Endocomia rufirachis — b: plural type, distally branched, of Endocomia macrocoma subsp. longipes
— ¢: ditto of Gymnacranthera forbesii var. forbesii — d-g: plural type, generally distally branched,
of Horsfieldia, d: H. polyspherula, e: H. clavata, f: H. spicata, g: H. iryaghedhi (the latter with a
strongly aberrant inflorescence within the genus Horsfieldia, the position of the first basal
ramifications of the single-type partial inflorescences is often not clear) — h-k: a choice of forms
of single-type inflorescences in Knema, h: K. pseudolaurina, i: K. furfuracea, J: K. celebica, k: K.
tridactyla — \-o: a choice of forms of single-type inflorescences as found in Myristica:: sect.
Myristica: |: M. iners, m: M. schleinitzit, n: M. fragrans; sect. Fatua: 0: M. fatua — p: inflores-
cence of Paramyristica sepicana; note that here the single-type inflorescences, which are similar in
Myristica, are distichously grouped into a short-shoot ending in a vegetative bud.

De Wilde — Myristicaceae

12 Flora Malesiana, Ser. I, Vol. 14 (2000)

Survey of Malesian genera with description of their inflorescences (Fig. 1)

1) Endocomia — Inflorescences predominantly of the plural type, panicle-like, resem-
bling those of most Horsfieldias; only in one species, E. rufirachis, a few single-type
inflorescences have been found. Flowers either faintly clustered, or clustered into
loose umbels or umbel-like racemes, often with the apical flowers the youngest. —
Fig. la, b.

2) Gymnacranthera — Inflorescences of the plural type, panicle-like, resembling those
of Horsfieldia but branching also from the axils of the lower cataphylls, so that the
common peduncle is short or absent. Flowers solitary, or grouped in small few-flow-
ered racemes or small clusters. — Fig. Ic.

Similar inflorescences with basal branching are prevalent in the Horsfieldia clavata-
group (New Guinea).

3) Horsfieldia (with 3 sections) — Inflorescences of the plural type, panicle-like, vari-
able in size and shape. Main peduncle with scars of cataphylls (bracts) and basal
(cataphyll-like) prophylls present; ramification into partial inflorescences predomi-
nantly higher up in the inflorescence, but mainly basally in the H. clavata-group (with
deviating androecium, New Guinea); see also under Gymnacranthera. Flowers (in
most species, sect. Jrya and Pyrrosa) on the ultimate branches, solitary or in few-
flowered loose clusters, or in H. iryaghedhi (sect. Horsfieldia, Sri Lanka) flowers
numerous, clustered into dense capitula. — Fig. 1d-g.

4) Knema — Inflorescences of the single type. They consist of persistent scar-covered
brachyblasts (of long duration), simple or + digitately 2—4-branched, producing flowers
at the apex each flowering season. Flowers each season few to many, closely set,
often appearing as an umbel, spirally arranged, axillary to minute caducous bracts.
Common peduncle absent, or up to 10 mm long, smooth. The Knema-type of inflores-
cence also occurs in Myristica sect. Fatua and is thought as having originated by
suppression of most of the common peduncle and by contraction of the flower-bear-
ing raceme. — Fig. 1h—k.

5) Myristica — Inflorescences of the single type. The inflorescences of the two sections
as recognized by Sinclair (1968: 30, 41), sect. Myristica and sect. Fatua, have differ-
ent designs based on the same plan, taking into account the necessary reductions and
corresponding mode of clustering of the flowers. — Fig. 1 l-o.

Sect. Myristica: inflorescences branched, panicle-like, generally lasting only one flow-
ering season; peduncle (hypopodium) distinct, often flattened; the proximal pair of
main branches and the central branch may be developed variously; sometimes the
central branch is lacking, or all three hardly may be developed. Within sect. Myristica
all kinds of intermediates between the situations of Fig. 4 1-n can be found, some-
times within one species.

Sect. Fatua: inflorescences as in Knema, i.e., simple or digitately few-branched, with
woody scar-covered axes, producing flowers for several seasons; peduncle absent or
short, not flattened (Fig. 1o). Also in the Fatua-type inflorescences the two opposite
basal side branches are generally clearly visible, with the central branch either well
developed or represented by only a single flower or the scar of its pedicel, e.g. as in
M. fatua.

De Wilde — Myristicaceae 13

Although Sinclair’s sections differ in inflorescence type, it is also true that for many
species their assignment to a section is arbitrary. For instance, in sect. Myristica, the
type species M. fragrans has male inflorescences which may be judged as short-lived
with a few flowers, but generally they grow quite old and last for many flowering
seasons, each season producing one or a few flowers at the ends of one or two slender
scar-covered raceme-like partial inflorescences, usually with a single middle-flower;
this is the dichasial inflorescence as described by Warburg (1897: 42) and later au-
thors. Recently these inflorescences have been described accurately by Armstrong &
Tucker (1986). The Knema-like inflorescences of sect. Fatua can be related easily to
a variant of the single-type inflorescence occurring in sect. Myristica.

6) Paramyristica — Here the inflorescences are truly of a mixed nature. The true (par-
tial) inflorescences are similar to the single-type, condensed inflorescences of
Myristica, but these are distichously arranged on a lateral short-shoot which ends,
significantly, in a small vegetative bud. This arrangement seems also predominant in
a few New Guinean species of Myristica, for instance M. markgraviana (see De Wilde
1995):

Flowers — The unisexual flowers are campanulate or urceolate, waxy-creamy or yellow
outside, greenish, creamy, yellow, red, or white (Knema galeata) inside. They can be
glabrous or brownish hairy on both surfaces. Flowers are frequently fragrant (e.g.,
Horsfieldia iryaghedhi, Myristica fragrans). The uniseriate perianth is (hard) carnose,
and cleaves at anthesis along previously developed lines of suture into 2—5 lobes, to
various depths, + according to the genus. The perianth splits deepest, nearly to the base,
in Endocomia and part of Knema; in some species of Horsfieldia it opens only incon-
spicuously at the very apex. The perianth lobes usually curve back at anthesis (espe-
cially or only in female flowers), except for Horsfieldia and possibly Paramyristica. In
the descriptions the size of the dry mature buds is given, and the length of the lobes by
stating the depth of the cleft by fractions. In Knema the full-sized flower buds remain
closed for a long time before opening. According to the genus the short or long pedicels
may or may not have one bracteole (rather large in Myristica, small in Knema). In some
species of Horsfieldia the pedicel (best to be seen in male flowers) is more or less dis-
tinctly jointed at the base; this feature can be used in species delimitation. The female
flowers (generally somewhat larger than the male flowers) have a single monocarpellate
ovary (hairy or glabrous), with sessile or short-stipitate, usually bilobed stigma, the lobes
being simple or variously lobulate again; they are conspicuously many-lobulate espe-
cially in species of Knema, style and stigma are conspicuously small in Myristica. The
androecium of the male flowers is most distinctive for the genera, as explained below
(Fig. 2).

Note on the construction of the androecium, and its naming in the six genera

The male organ in the unisexual flowers is formed by the fusion of a variable number
of stamens. Together they form the androecium, situated in the centre of the flowers.
Each anther consists of a pair of bisporangiate lobes or thecae (which are usually septate
when immature). When fully fused, the number of anthers may be difficult to count, but
it is half that of the often closely appressed thecae. Genera in which the filaments of the
individual stamens have remained partly free (e.g. in the Madagascan genus Maloutchia),

14 Flora Malesiana, Ser. I, Vol. 14 (2000)

:
0

b Cc d e f
Fig. 2. Schematic drawings of the androecium of Endocomia (a), Gymnacranthera (b), Horsfieldia
(c), Knema (d), Myristica (e), and Paramyristica (f).

can be regarded as comparatively primitive in this respect. The Malesian genera all have
(almost) completely fused stamens, forming an androecium of which the shape is char-
acteristic for the genus. For convenience’s sake, schematic drawings with a brief expla-
nation of the construction of the androecium and the terms used in the taxonomic text
are given for each genus. The (partly) connate or free anthers are shown in solid black.
In the case of connate anthers these form the synandrium and the fused filaments below
form a stalk called androphore. When the androphore is absent or inconspicuous, the
synandrium coincides with the androecium. In Knema the anthers are mostly free, and
the androecium is described in a different way. In the descriptions the number of thecae
is given, but in Knema only the number of anthers. In Figure 2 the letters correspond
with the explanation below.

a: Endocomia — The synandrium consists of short anthers adnate by their backs to a
short slender column, carried on a stalk or androphore. In the descriptions the follow-
ing terms are used: synandrium, androphore.

b: Gymnacranthera — The androecium practically entirely consists of elongate anthers
which are tightly connate into an elongate synandrium by most of their dorsal and
lateral sides. The stalk or androphore is so short that the synandrium appears to be
sessile. Therefore solely the all-comprising term androecium is used in the descrip-
tions.

c: Horsfieldia — Here the synandrium is formed by elongate anthers which are com-
pletely or for a lower part attached by their backs to a swollen, solid or usually hollow
column. The synandrium is either subsessile (as in Gymnacranthera), or carried on a
comparatively short stalk or androphore. In the description either only the general
term androecium is used when an androphore is practically absent, or the terms
androecium and androphore when the latter is obvious.

d: Knema — The androecium is formed by an apical disc-like structure to which the ses-
sile or free anthers are attached radially. Below the disc there is a cylindrical or taper-
ing stalk. In the descriptions the terms staminal disc and staminal column are used.

De Wilde — Myristicaceae 15

e: Myristica — Tightly set elongate anthers are completely adnate by their backs to a
rather thick central column, together forming the elongate synandrium. Above this,
the column is usually produced into a sterile portion, called the sterile apex, whereas
the synandrium is carried on a distinct cylindrical stalk or androphore. The terms
used in the descriptions are sterile apex, synandrium, and androphore.

f: Paramyristica — The subglobose synandrium is formed by (tightly set) elongate an-
thers which are completely fused by their backs to a swollen but largely hollow col-
umn, as in most species of Horsfieldia, and is carried on a comparatively long stalk
or androphore. Terms used in the taxonomic descriptions are synandrium and andro-
phore.

Fruits — The fruits are ellipsoid or oblong, more rarely (sub)globose, and they vary
strongly in size (I-12 cm long); only in Gymnacranthera all species have rather small
fruits. Fruits are essentially similar in construction in all genera, when fully ripe a firmly
fleshy or + coriaceous unicarpellate capsule, circumferentially opening at ventral and
dorsal side, at the latter, though not completely to the base. Usually the fruits are vari-
ously rusty pubescent or glabrous, pale yellow or creamy, pinkish, or salmon, or in
Endocomia canarioides glossy dark purple. The colourful unit of brown or black seed
with bright orange or red aril remains attached to the inner base of the pericarp (which
often is brightly coloured inside). In Endocomia the colour of the aril possibly is not
always red, probably in some species yellow, but this needs further observation. The
showy open fruits with contrasting colours supposedly attract frugivorous birds.

The fruit (pericarp) usually shrinks considerably on drying, and shape and size infor-
mation given in the descriptions concerns dried specimens.

Seeds — The single seed is generally similar in shape to that of the fruit. The endosperm
is ruminate, the embryo small, the seed coat ligneous, (blackish) brown, or grey, and
covered by the firm-fleshy aril. According to Corner (1976) the construction of the seed
coat is anatomically characteristic for the various Asian genera. The tegmen is massive
(Corner 1976; Van Heel 1982) and causes the characteristic rumination of the seed
(rumination in Annonaceae and some other families is of both testal and tegmic origin).
The testa in Endocomia is (mostly) variegated, in general an infrequent feature of
seeds.

The rather thin (sometimes thick) hard-fleshy aril is a true aril, originating from fu-
nicular as well as exostomal tissue. It is always well developed and completely covering
the seed in Asian Myristicaceae (reduced in Myristica ingens from New Guinea) and
either entire or shallowly to deeply laciniate, according to the genus. The aril is entire or
only shallowly lobed in Knema and Horsfieldia, in the latter sometimes + elongate above
the seed into a short folded tube; the aril is incised to about halfway in Endocomia and
deeply cleft (nearly) to the base in Gymnacranthera, Myristica, and Paramyristica. The
embryo is small and shows variation in the position of the cotyledons and whether or not
they are partially connate, according to the genera (Warburg 1897; Sinclair 1958). The
endosperm (albumen) is hard and contains fat and/or fixed (not volatile) oil, and some
essential (volatile) oil (3—8% in seeds of M. fragrans, which contains a narcotic); starch
may be present, is abundant in Myristica, and absent in Gymnacranthera and Horsfieldia.

16 Flora Malesiana, Ser. I, Vol. 14 (2000)

References: Armstrong, J.E. & S.C. Tucker, Floral development in Myristica (Myristicaceae).
Amer. J. Bot. 73 (1986) 1131-1143. — Corner, E.J.H., The seeds of Dicotyledons. Cambridge
Univ. Press (1976). — De Wilde, W.J.J.O., The genera of Myristicaceae as distinguished by their
inflorescences, and the description of a new genus, Bicuiba. Beitr. Biol. der Pflanzen 66 (‘1991’,
1992) 95-125; Census of Myristicca (Myristicaceae) in New Guinea anno 1994. Blumea 40 (1995)
237-344. — Sinclair, J., Gard. Bull. Sing. 16 (1958) 205; 33 (1968) 1-540. — Van Heel, W.A.,
Notes on the structure of developing seeds of Knema and Horsfieldia (Myristicaceae). Blumea 28
(1982) 53-60. — Warburg, O., Monographie der Myristicaceae, Nova Acta Acad. Caes. Leop.-
Carol. 68 (1897) 1-680.

FOSSILS

Little is known about fossil Myristicaceae. A leaf fragment, Myristicophyllum, is de-
scribed from E Borneo (Geyler 1887); fossil wood, Myristicoxylon princeps, has been
described from the Cretaceous in the Sahara (Boureau 1950).

References: Geyler, H.T., Uber fossile Pflanzen von Labuan aus Vega. Exped. Vetensk.,
Jakttagelser, Stockholm 4 (1887) 499, t. 33, f. 3-6. — Boureau, E., Etude paléoxylologique du
Sahara (IX). Sur un Myristicoxylon princeps, n. gen, n. sp., du Danien d’ Asselar. Bull. Mus. Hist.
Nat. Paris II, 22 (1950) 523-528.

CHROMOSOMES

According to the summary presented by Kithn & Kubitzki (1993), based on Marawetz
(1986), and Plant Resources of South-East Asia 5 (2, 1995 & 3, 1998)], chromosome
numbers are high and interpreted as (paleo)polyploid. Known are for Knema: 2n = 42
(K. intermedia: n = 21), Gymnacranthera: 2n = 44 (G. farquhariana vat. zippeliana: n =
21), Horsfieldia: 2n = 50 (H. iryaghedhi: n= 25), Myristica: 2n = 42 or 44 (M. elliptica:
n=21, M. fragrans 2n = 42). In the New World much higher numbers have been found
(Osteophloeum: 2n = c. 280).

References: Kiihn, U. & K. Kubitzki, in: K. Kubitzki, F.G. Rohwer & V. Bittrich (eds.), The fam-
ilies and genera of vascular plants, vol. 2 (1993): Myristicaceae: 461. — Marawetz, W., Pl. Syst.
Evol. 152 (1986) 49-100.

USES

In most myristicaceous species the heartwood is poorly differentiated from the sapwood,
and the wood is of minor commercial importance. The timber is suitable for temporary
light constructions. The wood, mainly from the blackish-stemmed group (including
Myristica lowiana), is mostly soft or moderately hard or heavy; perishable, but easily
treated with preservatives; it is easy to work, but sometimes splits soon. The sapwood is
pale, sometimes not well defined, but often the heartwood is dark reddish brown.

The seeds of some species may be used for their fat content or their fragrance, also as
medicine. Myristica fragrans is most important, yielding nutmeg (seeds), mace (aril),
and the spicy pericarp can be candied. The seeds of M. argentea (W New Guinea) is of
minor importance.

Myristicaceae are rarely used in silviculture. Some data are given in PROSEA 5 (2,
1995 & 3, 1998). Propagation is by seed, and shade should be provided for germination
and growth. A few species are ornamental (e.g., Horsfieldia iryaghedhi), or may be
introduced as such (e.g., H. sylvestris)

De Wilde — Myristicaceae 17

Kino — This substance, in the field-notes called exudate or sap, oozes from freshly cut
bark in larger or lesser quantities according to individual species. It is also present in the
wood, twigs, and to a lesser extent in petioles and inflorescence axis. Its presence is an
excellent field test when one suspects a tree to belong to the Myristicaceae. The colour
varies from dark red to pink; less often it has an orange tint. Kino is not so obvious in
very young trees. The amount probably varies within a species with time. It contains
tannin and gum and has left many an indelible stain on the clothes of plant collectors.
Warburg (1897) stated that the kino of one species has been used in America as a styptic.
Its function is not known, but it may help the wounds of a damaged tree to heal. It has
been described as bloody and gruesome and Malays have aptly given Myristicaceae
names including darah, the Malay name for blood.

References: De Wilde, W.J.J.O., in: M.S.M. Sosef et al. (eds.), PROSEA: Plant Resources of
South-East Asia 5 (3) (1998) 214-216 (Endocomia W.J. de Wilde), 292—296 (Horsfieldia Willd.).
— De Wilde, W.J.J.O., et al., Gymnacranthera (A. DC.) Warb.; in: R.H.M.J. Lemmens et al. (eds.),
PROSEA: Plant Resources of South-East Asia 5 (2) (1995) 255-260. — Sambas, E.N., Knema
Lour.; in: M.S.M. Sosef et al. (eds.), PROSEA: Plant Resources of South-East Asia 5 (3) (1998)
317-320. — Sangat-Roemantyo, H., et al., Myristica Gronov.; in: R.H.M.J. Lemmens et al. (eds.),
PROSEA: Plant Resources of South-East Asia 5 (2) (1995) 346—356. — Warburg, O., Monographie
der Myristicaceae, Nova Acta Acad. Caes. Leop.-Carol. 68 (1897) 1-680.

VEGETATIVE ANATOMY

(J. Koster, leaf anatomy & P. Baas, wood anatomy)

Leaf anatomy — A detailed description of the leaf anatomy of the Asian Myristicaceae
was given by Koster & Baas (1981). For short leaf anatomical accounts see also Schouten
(1986) and De Wilde (1994). Metcalfe (1987) summarized the vegetative anatomy of
the whole family. For the present survey more specimens were examined, including the
two new genera Endocomia and Paramyristica.

About 65 of the species belonging to the six Asian genera have been examined leaf
anatomically. Individual species will not be mentioned in this synopsis, although many
of the species examined can be distinguished by their leaf anatomical characters. Genera
will be mentioned when a character has diagnostic value on the genus level.

Hairs are present, at least in young leaves, on both surfaces or only on the abaxial
surface (in Gymnacranthera and some Myristica species). A hair is composed of one
row of short to tall cells, having one or two arms each, one or two (rarely more) cells
nearest the epidermis (the so-called stalk cells) excepted. In Gymnacranthera, Myristica,
and Paramyristica the cells have two arms, of unequal length in Gymnacranthera; the
cells in Endocomia, Horsfieldia, and Knema have one arm. In older leaves the hairs have
often been shed, but the upright walls of the most proximal parts remain as cutinized
rings on the epidermis. These rings are subtended by one to numerous small cells, ar-
ranged in a circle or oval.

Alveolar material, as an irregularly structured cutinaceous layer overlying the cuticle
proper, is present on the abaxial side in many species. The thickness of the cuticle proper
measures up to 18 um adaxially and to 11 um abaxially. The cuticular flanges on the ad-
axial surface are usually more or less sinuous at high focus and more or less straight at
lower focus; thin areas of cuticle are present in the loops of the undulations. The cuticular
flanges usually show inconspicuous pitting.

18 Flora Malesiana, Ser. I, Vol. 14 (2000)

Abaxial papillae sometimes occur. Large empty idioblasts (partially) with a thin cuti-
cle or without a cuticle, probably secretory cells, are often present. Some species abound
in regular cork warts; groups of basal cells of hairs are probably the origin of some of
these structures.

Stomata are usually confined to the abaxial epidermis; the stomatal type is paracytic.
The dimensions of the guard cell pairs range from 8 to 21 um for the width and from 15
to 39 um for the length. The guard cells are often embedded in the subsidiary cells,
which are dome-shaped in Gymnacranthera. In Knema, Myristica, and Paramyristica
the stomatal complex is (strongly) sunken; the bordering epidermal cells show papillae.
In Knema and some species of Myristica these papillae are more or less horizontally
directed, leaving a star-shaped opening above the stomatal complex. In most species of
Myristica and in Paramyristica the more or less upright papillae form a ring above the
stomatal complex.

An adaxial hypodermis is sometimes present, either as a continuous layer or only
locally. An inconspicuous abaxial hypodermis has been recorded for a few species.

The mesophyll is dorsiventral (rarely isobilateral), with mostly two or three, some-
times up to four adaxial layers of palisade parenchyma. In the leaf margin the cells
adjacent to the epidermis often have sclerified walls.

The midrib is abaxially prominently raised, and adaxially raised in most Horsfieldia
species, in Endocomia, Knema, Myristica, and Paramyristica. There is a more or less
straight adaxial vascular bundle (sometimes strongly interrupted) and an arc-shaped abax-
ial bundle, sometimes joined together. The phloem is arranged in separate strands, often
in two layers. One to numerous phloem bundles are interspersed in the ground tissue
between the main bundles, often accompanied by xylem elements; in some Knema spe-
cies there is a complete collateral bundle in the pith. In Gymnacranthera the adaxial
bundle is absent. The whole system is surrounded by groups of sclerenchyma fibres,
which also occur in the pith, often even in the centre of the phloem bundles. The ground
tissue is from centre to periphery parenchymatous to collenchymatous, often interspersed
with cells with sclerified walls; in Gymnacranthera and Knema there are often several
layers of these cells at the periphery of the midrib. Sometimes adaxial chlorenchyma 1s
continuous in the midrib.

The veins are supplied with collateral bundles; the major veins may have a more
complex vascular system, not unlike that of the midrib. Sclerenchyma caps are present
at the abaxial and adaxial sides; in Knema a sclerenchymatous bundle sheath is found. A
usually poorly differentiated parenchymatous bundle sheath, in Knema sometimes con-
tinuous to the epidermides, surrounds the bundle and the sclerenchyma. In Knema strands
are present, consisting of sclerenchyma fibres only, in position and distribution not un-
like the vein bundles.

The petiole at its basal end has a vascular system consisting of three more or less arc-
shaped collateral bundles with free phloem bundles adaxially. The sclerenchyma is usu-
ally confined for the greater part to the abaxial sides. The vascular system of the distal
end is intermediate between that of the basal end and the midrib.

Crystals may be present in various types. Large druses in enlarged mesophyll idioblasts
frequently occur, often adjacent to epidermal cells, which may be extremely flattened
and have a thin cuticle and thin and short cuticular flanges (in Myristica adaxially and in
Endocomia, Gymnacranthera, and Horsfieldia adaxially and abaxially). Small druses

De Wilde — Myristicaceae 19

have also been found, but not in Gymnacranthera. Small spindle-shaped particles often
occur, usually grouped in cells of the mesophyll and the ground tissue of the midrib.
Other crystal types have been found in one or a few species only.

Large, more or less spherical cells frequently occur in the mesophyll and the ground
tissue of the midrib. Usually they contain oil, in some species probably tannin- or muci-
lage-like substances. The large empty idioblasts in the epidermis have been mentioned
above. Fairly thick-walled tubule-shaped cells have sometimes been found, adaxially
and abaxially of the sclerenchyma caps of the vein bundles. The content of these cells is
probably tannin.

Sclereids are often present as brachy- to astrosclereids in the ground tissue of the
midrib. Filiform, rarely branched sclereids have been recorded for Gymnacranthera.
Astrosclereids and thick filiform, branched sclereids are present in a few species only.

The genera Gymnacranthera and Knema can be distinguished by a combination of
leaf anatomical characters. Leaf anatomy provides no means to discriminate between
Endocomia and Horsfieldia and between Myristica and Paramyristica.

Wood anatomy — The wood anatomy of the Myristicaceae is fairly uniform. For a de-
tailed family description and literature survey see Metcalfe (1987). Wood anatomy of
the main Malesian genera is summarized in the PRosEA Handbooks 5: 2 & 3 (Lemmens
et al. 1995; Sosef et al. 1998) and pictured by Ilic (1991). The wood is diffuse-porous
with vessels medium-sized and in low density (usually 3—12/sq.mm), solitary and in
radial multiples. Perforations mixed simple and scalariform, but one of the types domi-
nant or (virtually) exclusive in some species. Intervessel pits ranging from scalariform
to opposite and alternate. Vessel-ray pits often coarse and with reduced borders to sim-
ple. Fibres thin- to medium thick-walled, with minutely bordered to simple pits, often
septate around the vessels. Parenchyma scanty paratracheal to narrowly vasicentric and
often also in zonate bands. Rays typically 1—2(—3)-seriate, heterocellular and composed
of fairly large cells. Crystals present in ray cells or absent. Tanniniferous tubes, usually
in very low frequency, present in all species studied, and virtually unique to the family
Myristicaceae (except sporadic occurrence in some members of the U/maceae). Oil and/
or mucilage cells present among the axial and ray parenchyma in some species.

References: De Wilde, W.J.J.O., Paramyristica, anew genus of Myristicaceae. Blumea 39 (1994)
341-350. — Ilic, J., CSIRO Atlas of Hardwoods (1991) 331-334. — Koster, J. & P. Baas, Com-
parative leaf anatomy of the Asiatic Myristicaceae. Blumea 27 (1981) 115-173. — Lemmens,
R.H.M.J., I. Soerianegara & W.C. Wong (eds.), Plant Resources of South-East Asia 5 (2). Timber
Trees: Minor commercial timbers (1995) 255-260. — Metcalfe, C.R., Anatomy of the Dicotyle-
dons, Ed. 2, Vol. 3 (1987) 56-71. — Schouten, R.T.A., Revision of the genus Gymnacranthera
(Myristicaceae). Blumea 31 (1986) 451—486.— Sosef, M.S.M., L.T. Hong & S. Prawirohatmodjo
(eds.), Plant Resources of South-East Asia 5 (3). Timber Trees: Lesser-known timbers (1998) 292—
296, 317-320, 603-662.

PALYNOLOGY
(R.W. J. M. van der Ham)

The pollen morphology of the Myristicaceae has been poorly known for a long time.
The earliest more extensive account is that by Wodehouse (1937), who dealt with 36
species of the American genera, providing detailed descriptions and drawings. A more

20 Flora Malesiana, Ser. I, Vol. 14 (2000)

inclusive treatment is the light microscopic study by Agababian (1970) of 10 genera
from America, Africa and Asia. Further, pollen of a limited number of species is de-
scribed in pollen floras, of which Tissot et al. (1994) stands out by informative light and
scanning electron micrographs (Gymnacranthera, Knema, Myristica). Generic accounts
are those by Siddiqi & Wilson (1975; 8 spp. of Knema) and Medeiros Carreira (1985; 36
spp. of Virola, incl. Bicuiba). A preliminary paper by Walker (1976) contains a short
family description based on light and scanning electron microscopic data. Comprehen-
sive descriptions of all American, African and Madagascan genera, including scanning
and transmission electron micrographs, are in a series of papers by Walker & Walker
(1979, 1980, 1981, 1983). The Asian genera, among which the largest in the family
(Horsfieldia, Knema, Myristica), are still in need of elaborate palynological study. To
date the pollen of the Asian Endocomia (4 spp.) and Paramyristica (1 sp.) and the Afri-
can Staudtia (2 spp.) is entirely unknown.

Pollen grains of Myristicaceae are usually subspherical to slightly boat-shaped mon-
ads with a single, probably always distal aperture. Occasional chance tetrads, observed,
for instance, in /ryanthera, are tetragonal. The outline in polar view is subspherical to
elliptic, or sometimes obtusely rectangular. Outline in equatorial view is subspherical to
elliptic, or often obtusely triangular with a straight to slightly convex apertural side and
a more or less strongly convex nonapertural side. Pollen grain size (largest equatorial
diameter) is mostly between 20 and 40 um. Pollen of Brochoneura 1s smaller (14-21
uum). Some other genera have larger pollen grains: Gymnacranthera (up to 49 um), Knema
(up to 57 um), Myristica (up to 59 um), and Mauloutchia (up to 69 um).

Aperture morphology is relatively simple and not much diverse. It ranges from dis-
tinctly sulcate via indistinctly sulcate (sulcoidate) to more or less ulcerate or ulceroidate.
The aperture margins are often not clearly defined, which in ulceroidate groups may lead
to a superficially inaperturate condition (cryptoaperturate). Wodehouse (1937) observed
that an apertural area, even in pollen with a hardly recognizable aperture in the exine,
shows a distinctly thickened intine. Sometimes such intine parts seem to be acetolysis-
resistant (Walker & Walker 1983).

Exine thickness is from 0.5 to 5 um. Rather thin exines are found in Brochoneura
(0.5 um), Pycnanthus (0.5 pum, exclusive echinae) and Scyphocephalum (0.8 \um). Fairly
thick exines (3—5 um) occur in the coarsely reticulate Myristica pollen. Exine stratification
is usually distinct, with a thin to thick infratectal layer, which is mostly columellate. In
Brochoneura the infratectum is thin, so that the granulae observed by Walker & Walker
(1979) might actually represent short columellae. In Mauloutchia pollen the verrucate/
scabrate sexine elements seem to stand directly on the nexine, although irregular colu-
mella-like structures occur as well. The allegedly primitive granular infratectum reported
by Walker & Walker (1979) seems to be part of a granular sexine structure, which is
rather a derived feature. Distinct infratectal granules were found so far only in Otoba,
more or less adhering to the inner tectum surface and mixed with columellae. In view of
the other pollen characters (see below) this is probably also a derived condition. The
tectum as well as the nexine can be relatively thin to rather thick. In a few genera most or
only the inner part of the nexine may be lamellate. Because of the absence of any con-
trast in the nexine in transmission electron micrographs, the whole exine is considered
to be ectexinous.

De Wilde — Myristicaceae 21

Exine ornamentation 1s the most diverse character in Myristicaceae pollen: from psilate/
perforate via finely fossulate to coarsely reticulate, with several derivations. Pollen of
Brochoneura (Madagascar) has a simple massive psilate/perforate tectum. Pollen of the
American genus Otoba is psilate/imperforate with a proximal, + protruding (coarsely)
reticulate area. In Compsoneura and Virola (both American) the ornamentation is finely
fossulate to coarsely reticulate. In both genera reticulate pollen with scabrate (+ banded)
muri is found. Such ornamentation occurs also in /ryanthera (America) and Coelocaryon
(Africa), while the finely fossulate type with vaguely banded muri of Haematodendron
(Madagascar) and the crotonoid type of Scyphocephalum (Africa) can be easily joined.
The finely fossulate type of Compsoneura and Virola is also known from Gymnacranthera
(Asia), and the (more) coarsely reticulate type from the American Bicuiba and Osteo-
phloem, and the Asian Horsfieldia, Knema and Myristica. In Horsfieldia the reticulum is
sometimes interrupted, so that an intectate condition remains. Ornamentation in the
Madagascan genus Mauloutchia is diverse: scabrate, verrucate (verrucae scabrate or
smooth) or scabrate/echinate. Scabrate verrucae occur also in Cephalosphaera (Africa).
The pollen of Pycnanthus (Africa) is finely reticulate/echinate.

Concluding, the pollen of the Myristicaceae is diverse. Virtually every genus in the
family is palynologically distinct. A rigid subdivision based on pollen morphology, how-
ever, is difficult, but may be attempted after a more extensive study of the Asian genera.

The family has a modest fossil pollen record (Muller 1981). Pycnanthus type pollen
is known from the Upper Eocene and Lower Miocene of Africa, and pollen grains of the
Virola type from Pliocene and Quaternary sediments in Guyana.

References: Agababian, V.S., Biol. Zh. Armenii 23, 5 (1970) 58—69. — Medeiros Carreira, L.M.,
Bol. Mus. Paraense E. Goeldi, Bot. 2 (1985) 29-76. — Muller, J., Bot. Rev. 47 (1981) 1-142. —
Siddiqi, M.R. & T.K. Wilson, Pak. J. Bot. 7 (1975) 197-200. — Tissot, C., H. Chikhi & T.S. Nayar,
Publ. Dép. Ecol. Inst. Fr. Pondichéry 35 (1994). — Walker, J.W., Linn. Soc. Symp. Ser. 1 (1976)
251-308. — Walker, J.W. & A.G. Walker, Ann. Missouri Bot. Gard. 66 (1979) 731-755; Amer. J.
Bot. 67 (1980) 603-611; Grana 20 (1981) 1-17; Amer. J. Bot. 70 (1983) 315-326. — Wodehouse,
R.P., Brittonia 2 (1937) 397—402.

PHYTOCHEMISTRY AND CHEMOTAXONOMY
(R. Hegnauer)

General remarks — Chemical characteristics of the family were discussed in a number
of reviews in recent time (Hegnauer 1969, 1989, 1990; Gottlieb 1979). Many references
and structural formulae are given in these surveys. Therefore a compact résumé of pres-
ently known facts and some references to most recent chemical investigations of myris-
ticaceous plants, with emphasis on Asian taxa, seem to be appropriate here. Most mem-
bers of the family are locally used in traditional medicine. This is one of the reasons why
we are relatively well informed about its secondary metabolites. Moreover, Myristica
fragrans yields the famous spices nutmeg and mace. There is plenty of literature about
this plant, its cultivation and its products (e.g., Briicher 1977, Purseglove et al. 1981,
Delaveau 1987, Flach & Tjeenk Willink 1989). Because nutmeg, if taken in large amounts,
is toxic and causes among other symptoms hallucinations, pharmacologists and ethno-
botanists interested in psychotropic plants became also involved in nutmeg research
(Efron 1967).

22 Flora Malesiana, Ser. I, Vol. 14 (2000)

Chemistry of the family — Essential oils, lignans and neolignans, flavonoids in the
widest sense and biogenetically related phenolic compounds, peculiar acetogenins based
on long-chain fatty acids, and large amounts of a special type of triglycerides in seeds
are outstanding myristicaceous features. Moreover, tryptamine-derived alkaloids were
reported in several genera, and diterpenoids and triterpenoids were detected only errati-
cally hitherto.

Essential oils — Belonging to woody polycarps (i.e. Magnoliidae—Magnolianae
sensu Takhtajan 1980) Myristicaceae have oil cells in most of their parts and usually are
aromatic plants. So far only essential oils of nutmeg and mace were investigated thor-
oughly (Purseglove et al. 1981). There are scarcely qualitative differences between the
oils of nutmeg and mace, but rather marked quantitative differences. The same is true
between nutmeg oils of different production centres (West India [Grenada], Indonesia
and East India) and even between individual trees of the island Grenada. Roughly nut-
meg yields 7-16% and mace 4—17% essential oil. Nutmeg oil contains 61-66% mono-
terpene hydrocarbons (85-93% individual trees of Grenada), mainly pinenes and sabi-
nene, 5—15% (6.6-12%) monoterpene alcohols and their esters, such as geraniol and
linalool, and their acetates, and 2-18% (0—3.5%) phenylpropanoid ethers, among which
myristicin, elemicin and safrole predominate. These aromatic allylphenolic ethers are
assumed to represent the main toxic and psychotropic constituents of nutmeg and mace.
The essential oil of leaves of M. fragrans also contains mainly pinene and myristicin.

Lignans and neolignans — These plant constituents are dimers of phenylpropanoids
(C, ... Co + Cy ... Cor). According to Gottlieb and Yoshida (1989) lignans and neo-
lignans should be defined biogenetically not purely on chemical arguments. They con-
sider lignans as Cg—Cg--linked dimers of phenolic derivatives of cinnamy! alcohol
(CgHs-7CH = 8CH-2CH OH) or phenolic derivatives of cinnamic acid (CgHs-’CH =
8CH-°COOH). On the other hand neolignans are dimeric derivatives of allylbenzenes
(CgHs-7CH>-8CH = 9CH); e.g. eugenol) or propenylbenzenes (CgHs-7CH = 8CH-°CH3;
e.g. isoeugenol), and different types of linkage between the two monomers OCCUT, €.8.
RuguaGalees) C-Sta8- 7 aSeoee =o), O- Ona ; 4-Q-5’ etc. In both, lignans and neolig-
nans, one or two additional linkages between the two units are often present. In lignans
Cy and Co: carry oxygen and in neolignans they do not. A botanical argument which
favours such a lignan-neolignan distinction is their distribution in seed-plants. Lignans
occur everywhere in gymnosperms and angiosperms, and neolignans are mainly (not
wholly!) restricted to Magnoliidae—Magnolianae (Takhtajan 1980) which correspond
with woody Polycarpicae + Piperales of Wettstein (1935). In Myristicaceae both true
lignans and neolignans occur frequently. Conserva et al. (1990) call neolignans “the
most conspicuous constituents of Myristicaceae.” Examples of neolignans occurring
in the family are dehydroguaiaretic acid and | ,2-dihydrodehydroguaiaretic acid of the
stem bark of Knema furfuracea (Pinto et al. 1990), and lignans are represented, e. g. by
asarinin. horsfieldin and dihydrocubebin, from leaves, bark, wood and seeds of Hors-
fieldia iryaghedhi (Gunatilaka et al. 1982; Tillekeratne et al. 1982). By condensations
with chalcones or dihydrochalcones neolignans can give rise to still more complex
phenolic compounds such as the lignoflavonoids iryantherin A to J of South American
Iryanthera taxa (Conserva et al. 1990; Silva et al. 1995). Finally it should be mentioned
that not all phytochemists follow the lignan-neolignan-definition of Gottlieb and Yosh-
ida. Many chemists prefer the older definition which considers all 8-8’-linked dimeric

De Wilde — Myristicaceae 23

phenylpropanoids as lignans, and dimers with other linkages, e.g. the 5-5’-linked dehy-
drodieugenol and the 8-5’-linked carinatone of Virola carinata, as neolignans.

Flavonoids and biogenetically related phenolic compounds — If flavonoids are defined
as phytoconstituents derived from a cinnamic acid and three acetates (malonates) which
yield the phloroglucinol- or resorcinol-type A-ring, this class of natural products com-
prises many chemical subclasses. Myristicaceae are outstanding producers of flavonoids
sensu lato. At present the following types of flavonoid phenolics are known from the
family: Flavonols (e.g. kaempferol, quercetin), flavones (e.g. apigenin, luteolin, 7,4’-
dimethoxyflavone, a 5-desoxyflavonoid), flavanones (pinocembrin), several chalcones
and dihydrochalcones, several diarylpropanes, isoflavones (e.g. 2’-hydroxyformonon-
etin [also a 5-desoxyflavonoid]), pterocarpans (demethylhomopterocarpin, maackiain),
flavan-3-ols (catechins, e.g. fisetinidol) and flavans. Moreover, the family is rich in con-
densed tannins which are based on catechins and leucoanthocyanidins (flavan-3,4-
diols). Leucoanthocyanidins have not yet and catechins only rarely been isolated from
Myristicaceae. Nevertheless the production of adstringent kino-like exudates by most
species and the demonstration of the presence of procyanidins in a few species render
possible the conclusion that myristicaceous tannins are of flavanoid nature. Still another
type of natural products is represented by stilbenes. In contrast to flavonoids which are
C6-C3-Cg compounds, stilbenes are Cg-C>-C,, products, because during the biosynthesis
of usual stilbenes one CO; is lost. Stilbenes occur in Myristicaceae, e.g., as mono-, di-
and trimethylethers of resveratrol (= 3,5,4’-trihydroxystilbene).

Peculiar acetogenins based on long chain fatty acids — Two main types of such
acetogenins or polyketides occur in the family. (a) The anacardic acid-cardol-type of
alkyl- or alkenylphenols which is based on ordinary polyketides. If polyketide synthesis
starts with a cinnamic acid molecule this pathway yields w-phenylalkanyl]- and -alkenyl-
phenols. The phenolic part of such acetogenins originates from cyclization of the last
three or four acetyl units of the polyketide chain and bears one, two or three phenolic
hydroxyls (= phenol-, resorcinol- and phloroglucinol-type compounds). Additionally this
aromatic ring may carry a carboxyl group (anacardic acids sensu stricto) or an acetyl
group (acetophenone derivatives). Phenolic alkanones, such as the malabaricones, have
an Oxo group in the side chain next to the aromatic ring. In some species 3-alkyl- or
3-w-phenylalkylisocoumarins occur; these metabolites are lactones of an anacardic acid
carboxyl with a 2’-hydroxyl in the side chain. Thus this type of biogenetically related
phenolic polyketides is extremely diverse in Myristicaceae. (b) The second class of
acetogenins bears a methyl- or methylene-butanolide or -butenolide structure which is
probably formed by a reaction of the carboxyl group terminating the polyketide chain
with a pyruvate unit or its enolic form. Examples of this type of acetogenins are iryellip-
tin, grandinolide and the juruenolides of /ryanthera elliptica, grandis, jururensis and
ulei and Virola surinamensis (Lopes et al. 1994, 1996).

Most recent phytochemical investigations treat mainly lignanoids, flavonoids and
acetogenins. Examples are: Horsfieldia iryaghedhi (Gunatilaka et al. 1982; Tillekeratne
et al. 1982). Knema austrosiamensis (Gonzales et al. 1993, 1996), K. elegans (Spencer
et al. 1980), K. furfuracea (Pinto et al. 1990; Zahir et al. 1993), K. glomerata (Lu
Zeng et al. 1994), K. laurina (Kijjoa et al. 1991; Gonzalez et al. 1996), K. tenuinervia
subsp. setosa (Kijjoa et al. 1991). Myristica dactyloides (Herath & Priyadarshini 1996,
1997). Pycnanthus angolensis (Omobuwajo et al. 1992). For 1,3-diarylpropanes and

24 Flora Malesiana, Ser. I, Vol. 14 (2000)

| 3-diarylpropan-2-ols and catechins Virola elongata and minutiflora are noteworthy
(Kijjoa et al. 1981). Virola venosa (Kato et al. 1992) and Virola aff. pavonis (Martinez &
Torres 1997) represent a notable example for the vicarious occurrence of lignans and
neolignans in the family.

Seed fats (oils) — Myristicaceae store large amounts of triglycerides in seeds; they
are accompanied by proteins and in some species by starch. The triglycerides of the
family contain saturated fatty acids as main acids, usually 14:0 (myristic) and 12:0
(lauric) and sometimes 16:0 (palmitic) or 18:0 (stearic). The presence of large amounts
of 14:0 in several seed fats explains the fact that trimyristin could be isolated from the
seeds of a number of species. Often the seed lipids contain a large portion of unsa-
ponifiable matter, i.e. non-triglycerides. The non-triglyceride part consists of essential
oils, lignans, acetogenins and other resinous matters.

Alkaloids — As already mentioned (Efron 1967) tryptophan-derived protoalkaloids
and B-carboline alkaloids occur in several species of Virola. Recently 5-methoxy-
N,N-dimethyltryptamine, 6-methoxy-2-methyl-1 .2,3,4-tetrahydro-B-carboline and hors-
filine, C;3H6N20>, a new oxindole alkaloid, were isolated from leaves of Horsfieldia
superba (Jossang et al. 1991). Leaves of Osteophloeum platyspermum contain the
methylether of N-methyltryptophan. Thus a special metabolism of tryptophan yielding
psychotropic tryptamines and simple indolic alkaloids seems to be present in New World
(Virola and Osteophloeum species) and Old World (Horsfieldia species) members of the
family. Bennett and Alarcon (1994) published recently a remarkable ethnobotanical
paper about Amazonian Myristicaceae and about hallucinogenic uses of Osteophloeum
platyspermum and Virola duckei in Ecuador.

Meroterpenoids (= compounds of partly terpenoid origins) — Fruits (seed kernels,
arilli, pericarps) yield lipid fractions which often contain besides triglycerides appreci-
able amounts of ‘resinous matter’ of varying composition (essential oils, lignanoids and
flavonoids, acetogenins and, in some instances, meroterpenoids). Such a meroterpenoid
is komboic acid of seeds of Pycnanthus kombo. It amounts to ca. 23% of total seed ‘fat’
and was characterized as 16(2’,5’-dihydroxy-3’-methylphenyl)-2,6,10,14-tetramethyl-
2.6,10,14-hexadecatetraenoic acid, i.e. a 2-geranylgeranyl-substituted 6-methylhydro-
quinone with one of the last methyl groups in the geranylgerany] side chain oxidated to
COOH. Biogenetically related tocotrienols (vitamin E group), of which 2,8-dimethyl-
2-(4,8,12-trimethyl-3,7,1 |-tridecatrienyl)-6-chromanol was the main product, were iso-
lated from fruits of Iryanthera grandis, and seeds of Otoba parvifolia yielded a series of
farnesylated aromatic to semiaromatic and ring-constricted compounds which probably
all derive from the same biogenetic pathway, i.e. farnesylation of gentisic acid and con-
sequent modifications of the resulting farnesylgentisic acid (Ferreira et al. 1989, 1995).
Gentisic acid may also be involved in the biosynthesis of komboic acid and the toco-
trienols in which an aliphatic diterpene in place of the sesquiterpene farnesol is com-
bined with an aromatic ring.

Diterpenes, triterpenes and phytosterols — Small amounts of phytosterols and tetra-
and pentacyclic triterpenes are ubiquists in unsaponifiable matters of plant lipids. Accu-
mulation of diterpenes and triterpenes seem to be much more restricted and rather rare in
the family. The tetracyclic triterpenes cycloeucalenol and 24-methylenecycloartanol were
isolated from wood of Cephalosphaera usambarensis, and nutmegs yielded a saponin
with oleanolic acid as sapogenin. Diterpenoids were isolated from leaves and twigs of

De Wilde — Myristicaceae 25

Osteophloeum platyspermum (a kaurane and three eperuane derivatives), and recently
stem bark of Staudtia kamerunensis yielded staudtienic acid, Cy9H2,602 with a rearranged
abietane structure (Noumbissie et al. 1992).

Summary and chemotaxonomic remarks — Myristicaceae are chemically characterized
by a number of metabolic features.

1) They store large amounts of triglycerides of buttery or fatty consistency because
12:0 and 14:0 (16:0, 18:0) are their main fatty acids.

2) They deposit variable amounts of essential oil in oil cells which occur in practi-
cally all plant parts.

3) They have an extremely diverse flavonoid metabolism which results in the produc-
tion of rather characteristic compounds in a number of their taxa, e.g. isoflavones, ptero-
carpans, | ,3-diarylpropanes and their 2-ols, chalcones and dihydrochalcones, flavanones,
flavones and catechins.

4) Lignans and/or neolignans occur in large numbers and often in large amounts in
all investigated plant parts of practically every member of the family.

5) Two types of polyketides (acetogenins) occur widely in the family: the anacardic
acid-type with several variants and the methylbutanolide-type with a smaller number of
variants.

6) Myristicaceae are relatively tannin-rich plants, but the precise nature of their tan-
nins is not yet known. All available facts indicate a universal presence of condensed
tannins, i.e. oligo- and polymeric proanthocyanidins.

Idioblasts storing essential oils characterize woody polycarps or Magnolianae (Takh-
tajan 1980). Within this taxon Myristicaceae resemble Lauraceae mostly in their seed
fats and in their secondary metabolism. Both families are virtuous producers of neolig-
nans and lignans and of polyketides of the butanolide-type. The typical alkaloids of
Magnolianae are benzyltetrahydroisoquinoline-type bases. They are widespread in
Lauraceae, but seem to be totally absent in Myristicaceae, but also in Winteraceae,
Calycanthaceae and some other, mostly small relictic families. Myristicaceae, however,
are not an alkaloid-free taxon. Some of its members produce tryptamine derivatives
including B-carboline and the oxindole horsfiline. The tendency to replace benzylisoqui-
nolines by tryptamine derivatives does also occur in some members of Lauraceae (Aniba
p.p., Nectandra p.p. and Umbellularia p.p.) and is characteristic of Calycanthaceae.
Myristicaceae could represent a member of Magnolianae in which accumulation of ben-
zylisoquinolines was totally replaced by intensifying the production of lignanoids and
flavonoids s.1. Finally it should be mentioned that Myristicaceae resemble strikingly
Papilionoideae (many 5-desoxyflavonoids, isoflavones, pterocarpans, | ,3-diarylpropanes,
flavans and fisetinidol-type catechins).

References: Bennett, B.C. & R. Alarcén, Osteophloeum platyspermum and Virola duckei (Myt-
isticaceae): Newly reported as hallucinogens from Amazonian Ecuador, Econ. Bot. 48 (1994) 152—
158. — Briicher, H., Tropische Nutzpflanzen (Ursprung, Evolution und Domestikation) (1977) 426-
427: Myristica fragrans; Springer-Verlag Berlin. — Conserva, Lucia M., et al., Phytochemistry 29
(1990) 3911-3918: Dihydrochalcones and the dihydrochalcone-based flavonolignans iryantherin
A to E from fruits and barks of Iryanthera laevis, paraensis and ulei. — Delaveau, P., Les épices;
histoire, description et usage des différents épices, aromates et condiments (1987), Ed. Albin Michel
S.A., Paris: Part IV. Passez muscade (Myristica fragrans): 154-164. — Efron, D.H. (Editor-in-
chief), Ethnopharmacologic search for psychoactive drugs, Public Health Publ. No. 1645 (1967),

26 Flora Malesiana, Ser. I, Vol. 14 (2000)

U.S. Government Printing Office, Washington, D.C.: Part HI: 185-229: Myristica fragrans, with
3 contributions treating mainly psychoactivity, toxicology, pharmacology and chemistry of nutmeg.
Part IV. South American snuffs with 6 contributions of which one treats botanical origins of snuffs
known as Yakee and Parica (prepared by some tribes with barks of Virola calophylla, calophylloi-
dea and elongata), one treats the snuff Epéna prepared from the bark of Virola calophylloidea, and
one treats the identification of N-methyltryptamines and B-carbolines in Epéna and Parica-snuff
preparations and in the bark of Virola calophylla, and an other one treats the psychoactive action of
tryptamine derivatives. The harmala alkaloids (= 8-carbolines) are treated in part V which is de-
voted to Malpighiaceae which yield the snuffs Ayahuasca, Caapi and Yagé. — Ferreira, A.G., et al.,
Phytochemistry 28 (1989) 579-583; 40 ( 1995) 1723-1728. — Flach, M. & M. Tjeenk Willink, in E.
Westphal & P.C.M. Jansen (Eds.) PROSEA, A selection (1989) 192-196: Myristica fragrans. Pudoc
Wageningen. — Gonzalez, M.J.T.G., et al., Phytochemistry 32 (1993) 433-438: Two resveratrol
methyl ethers, a diarylpropane, a flavan, two resorcinol- and two phloroglucincol-type alkanones
and two corresponding alkenones and the two lignans episesamin and xanthoxylol from wood of
Knema austrosiamensis. — Gonzalez, M.J.T.G., et al., Phytochemistry 43 (1996) 1333-1337: Sat-
urated and monounsaturated anacardic acids, alkylphenols and alkanoylphenols from wood of
Knema austrosiamensis and bark of K. laurina and 5-hydroxy-3’ ,4’-methylene-dioxyflavan from
bark of K. laurina. — Gottlieb, O.R., Chemical studies on medicinal Myristicaceae of Amazonia,
J. Ethnopharmacol. 1 (1979) 309-323. — Gottlieb, O.R. & M. Yoshida, Lignans, in: J.W. Rowe
(Ed.), Natural products of woody plants (1989) 439—S11. Springer-Verlag, Berlin. — Gunatilaka,
A.A.L., et al., Phytochemistry 21 (1982) 2719-2723: Lignans asarinin, dihydrocubebin and hors-
fieldin and trimyristin from seeds of Horsfieldia iryaghedhi. — Hegnauer, R., Chemotaxonomie
der Pflanzen, Vol. V (1969) 144-153, 435-437, 457. — Hegnauer, R., Chemotaxonomie der Pflanzen,
Vol. VIII (1989) 108-118: Comments on the chemistry of Polycarpicae (= Magnoliidae sensu Cron-
quist), including fatty acid-derived acetogenins of Myristicaceae. — Hegnauer, R., Chemotaxon-
omie der Pflanzen, Vol. IX (1996) 101-111. — Herath, H.M.T.B. & A.M.A. Priyadarshini, Phyto-
chemistry 42 (1996) 1439-1442; 44 (1997) 699-703: Neolignans and arylalkanones (malabaricones)
from bark. — Jossang, A., et al., J. Org. Chem. 56 (1991) 6527-6530. — Kato, M.J., et al., Phyto-
chemistry 31 (1992) 283-287: Two 5-desoxyflavones from flowers, pericarp and leaf and ten
lignans from pericarp, aril, seed and root and a special type of w-phenylalkanoylphenol (1-[11-
phenylundecanoyl]-3-hydroxycyclohexan-2,6-dione) which occurs in all parts and was already known
from Virola elongata and sebifera. — Kijjoa, A., et al., Phytochemistry 20 (1981) 1385-1388:
Virolane and virolaflorine, two 1,3-diarylpropanes from wood of Virola minutiflora and virolanol,
virolanol-B and -C, three 1,3-diarylpropan-2-ols and (—)-fisetinidol from wood of V. elongata. —
Kijjoa, A., et al., Planta Medica 57 (1991) 575-577: Bark of both species yielded only several
acetogenins of type (a). — Lopes, N.P., et al., Phytochemistry 35 (1994) 1469-1470; 43 (1996)
1089-1092: 17 Neolignans from leaves and seeds and juruenolide C and D and 3 propiophenones
from seeds and a new juruenolide from seedlings. — Lu Zeng et al., J. Natural Products 57 (1994)
376-381: Stem bark yielded 10 (a)-type polyketides among which kneglomeratanol and the two
acetophenones kneglomeratanone A and B and the isoflavones fromononetin, biochanin-A and 8-
O-methylretusin. — Martinez V., J.C. & R. Torres Ch., Phytochemistry 44 (1997) 1179-1182: Leaves
yielded nine neolignans among which otobaphenol. — Motter Magri, Fatima M.. et al., Phytochem-
istry 43 (1996) 669-671: Unripe pericarps yielded an 1,4-dioxane-type neolignan (linkages 7-O-7’
and 8-0-4’) and 4 butanolide-type polyketides. — Noumbissie, B.E., et al., J. Natural Products 55
(1992) 137-139. — Omobuwajo, O.R., et al., Phytochemistry 31 (1992) 1013-1014: 2’-
Hydroxyformononetin and its 7-methy] ether from wood. — Pinto, Magdalena M.M., et al., Phyto-
chemistry 29 (1990) 1985-1988: (a)-Type acetogenins among which an isocoumarin and two
neolignans (dehydroguaiaretic acid and its 1,2-dihydro derivative) from bark. — Purseglove, J.W..,
et al., Spices, Vol. 1 (1981), Logman Group Ltd., London: Nutmeg and mace p. 174-228: History -
Botany - Ecology - Cultivation - Diseases - Pests - Improvement - Products and end-uses - Process-
ing and manufacture - Chemistry - Standard specifications - Production, trade and markets - 3 pp.

De Wilde — Myristicaceae Di

of References. — Silva, Dulce H.S., et al., Phytochemistry 38 (1995) 1013-1016: Iryantherin G
to J from fruits of Iryanthera grandis. — Spencer, G.F., et al., J. Natural Products 43 (1980) 724—
730: Three neolignans and several anacardic acid- and resorcinol-type acetogenins from seeds. —
Takhtajan, A.J., Outline of the classification of flowering plants (Magnoliophyta), Bot. Rev. 46
(1980) 225-359. — Tillekeratne, L.M.V., et al., Phytochemistry 21 (1982) 467—476: Bark, wood
and leaves yielded the lignans asarinin and dihydrocubebin and dodecanoylphloroglucinol. —
Wettstein, R., Handbuch der systematischen Botanik, 4. Aufl. (1935), Franz Deuticke, Leipzig-
Wien. — Zahir, A., et al., J. Natural Products 56 (1993) 1634-1637: Leaves yielded two new
phenyl-acylphenols, knerachelin A and B.

VERNACULAR NAMES

The preferred trade name applicable for Myristicaceae in general is penarahan. Penin-
sular Malaysian names are: chendarah, chendarahan, darahan, penarah, penarahan,
pendarah, and pendarahan; in Sabah: darah-darah (preferred name); in Sarawak: binarah
(Murut), bindara (Kelabit), jela bala (Kenyah), kayo bela (Kayan), kayo raha (Berawan),
kumpang (preferred name), pang (Bidayuh), pendarahan, pumpu (Bidayuh Sadong),
raha meban (Punan Tutoh); the names balun ijok, darah-darah, penaharan, and pianggu
are also used in Sarawak; in the Philippines: duguan (Filipino language).

The group of Myristicas with black, gritty bark is called penarahan arang in Malaya
and parts of Indonesia. Some Knema species in Indonesia are called ki-mokla (Sundanese).
Specific names are, e.g., swamp nutmeg for Myristica elliptica (W Malesia) or man-
grove nutmeg for M. hollrungii (New Guinea); the papua nutmeg is M. argentea, a spe-
cies locally cultivated in SW Papua Barat. Some other Malay names for Myristicaceae
may be mentioned here: the nutmeg, Myristica fragrans, is called buah pala, while wild
nutmegs are called pala bukit or pala hutan. Horsfieldia irya is pianggu and H. crassifolia
is jangkang paya. Jangkang means stilt-roots and paya is swamp. In Sarawak and Bru-
nei, Myristicaceae are generally called kumpang. In New Guinea the family has many
different names in the local languages. This information about the vernacular names
comes largely from Sinclair (1958).

Reference: Sinclair, J., A revision of the Malayan Myristicaceae. Gard. Bull. Sing. 16 (1958)
205-472.

KEY TO THE GENERA
(based on male flowering specimens)

la. Inflorescence a sessile or to 3(—5) mm peduncled, short, tubercle- or worm-like pro-
tuberance, usually woody, with scars of fallen pedicels and bracts. Bracteole pres-

BCS aetitceciceSyatte sta diccah io fs Wie Ge Beka ic = Bhoee bt Got ele ites cee tee age eee 2
b. Inflorescence branched, panicle-like, short- or long-peduncled, the distal parts of
the branches woody and with scars, or not. Bracteole present or absent ....... 3

2a. Androecium a stalked disc with the anthers sessile, contiguous or largely free,
stellately attached by their bases. Bracteole mostly small, at the base of the perianth
Onlower to median on the pedicel: %4..00. (55... oe See oe Knema (p. 222)

b. Androecium a stalked elongate column with the anthers completely fused dorsally,
apex of the column often a sterile protuberance, or + flat, very rarely shallowly
hollowed (M. markgraviana, M. hooglandii). Bracteole embracing the perianth, at
dipncamiic apex ol the pedicel ..........t:.nGckoe. eae Myristica (p. 359)

Flora Malesiana, Ser. I, Vol. 14 (2000)

Sa.

6a.

la.
. Aril entire or laciniate up to halfway or less ..........-- 000s eres eee eees +
2a.

Bay

_ Bracteole present, at or towards the apex of the pedicel. [Inflorescences with scar-

covered perennial distal parts present or not.] .......------- Myristica (p. 359)

“Bracteole absent . Ste acne Scr os eee es ae) wie ean Tr cee ge «eles aero 4
. Synandrium variable, with the central column solid or excavated, usually consider-

ably broader than the androphore. Perianth inside glabrous, lobes not reflexed at
a thEST Senda? are El vi ae ls) acts ets ene: els ete ich 5

. Synandrium elongate or + globose, central column at apex not excavated, narrow,

about as wide as or narrower than the androphore. Perianth inside hairy or papillary
hairy, lobes erect or reflexed at anthesis. [Inflorescences panicle-like, usually branched
from near the base and with some basal cataphyll scars present.] ............ 6
Male perianth small, less than 4 mm long (6—7 mm in H. superba); androphore much
shorter than synandrium, glabrous. Inflorescence with peduncle with basal cataphyll
scars, not branched from base; apical vegetative bud absent ..............-----
we Hal axtpaireanventacuaed. orn doayla «7 \Paee Ut RMPS tell. Swe: Sai Ae A eee Horsfieldia (p. 54)

. Male perianth 5—6 mm long; androphore nearly as long as synandrium, with minute

hairs at base. Inflorescence a brachyblast composed of partial inflorescences of the
Myristica sect. Myristica-type (see also p. 360), ending in a vegetative bud.......
seh nd ps ah ATONE, SN OREN OP NP 2 ROR yr ee Paramyristica (p. 619)
Synandrium elongate, anthers free in the apical part. Perianth lobes erect at anthesis
ee ee PT oo oo: eee Gymnacranthera (p. 40)

. Synandrium short, (depressed) globose, anthers completely sessile, without apically

free part. Perianth lobes spreading or reflexed at anihesise eek Endocomia (p. 29)

KEY TO THE GENERA
(based on female flowering and fruiting specimens, also using vegetative characters)

Aril divided into segments to or almost to the base ......-- +... seer reese )

Inflorescences or infructescences paniculate, with scars of basal cataphylls, without
apical vegetative bud. Bark of twigs smooth or very finely striate. Fruits 1.5—-3 cm
long. [Bracteole absent. Leaves not conspicuously brittle when dry.) =" «eee
aa ea Sl e SI ed RE oN8) book Fd A |S) dey ge 2 a Gymnacranthera (p. 40)

_ Inflorescences either 1) simple or furcate, short, (sub)sessile, or panicle-like with

peduncle, without basal cataphyll scars and without terminal bud, or 2) compound,
consisting of those of the foregoing type distichously arranged on short-shoots, with
apical vegetative bud. Bark of twigs usually striate, longitudinally cracked, or flak-
ing. Fruits 15-8 con long) oi.aciae days 92 it «ce eye eo Se 5
Inflorescence a compound synflorescence, with apical vegetative bud. Bracteole (fe-
male flowers not seen) absent. Fruits 5 cm long, conspicuously pubescent. Crowded
linear basal cataphyll scars usually present on innovations. [Leaves brittle when dry. ]
Se eda cat core ces ps fsck os cS eg Se oe oh ee Rat Se Paramyristica (p. 619)

_ Inflorescences without apical vegetative bud, rarely present (M. carrii, M. hooglandii,

M. markgraviana). Bracteole scar on fruiting pedicel present. Fruits of variable size,
pubescent or glabrescent. Crowded basal cataphyll scars on twigs usually not present.
[Leaves brittle when dry.] .........252 22sec sees ene ees Myristica (p. 359)

De Wilde — Myristicaceae (Endocomia) 29

4a. Aril at apex convoluted or shallowly laciniate. Seed not variegated, not pointed at
Macecncunetat esis tes. valle. Wade ne emis. 2) Ze sh Bee SE Ue ee x

b. Aril coarsely incised for about the upper 1/3, or + entire in the Philippines. Seeds
usually variegated, often bluntly pointed at one end. [Bracteole absent. Stigma nar-
rowly 2- or few-lobed. Inflorescence paniculate. Monoecious. Leaves brittle when
POE VRMISINOCIOW.) oi. 20.00. OA, ee tee wee Endocomia (p. 29)

5a. Bracteole absent. Inflorescence paniculate. Stigma minutely 2-lobed (or few-lobed
in H. iryaghedhi, introduced). Leaves brittle when dry, lower surface usually not
whitish (papillose and whitish below only in H. iryaghedhi) .. Horsfieldia (p. 54)

b. Bracteole present. Inflorescences short, wart- or worm-like, simple or forked, scar-
covered, (sub)sessile. Stigma few- to many-lobed. Leaves generally not brittle when
dry; lower surface usually + whitish [venation on upper surface finely reticulate and
usually clearly visible, unlike most other genera.] ............ Knema (p. 222)

ENDOCOMIA

Endocomia W.J. de Wilde, Blumea 30 (1984) 179; Beitr. Biol. Pflanzen 66 (*1991", 1992) 95: Tree
Fl. Sabah & Sarawak 3 (2000) 338. — Type species: Endocomia macrocoma (Miq.) W.J. de
Wilde.

Horsfieldia sect. Pyrrhosa subsect. Papillosae Warb., Mon. Myrist. (1897) 265. — Based on Hors-
fieldia papillosa Warb., H. prainii (King) Warb., H. canarioides (King) Warb.

Trees, monoecious (always?). Twigs never angular or ridged, without lenticels, some-
times flaky. Leaves membranous or chartaceous, slightly brittle when dry, lower surface
not pale, not papillose, dots absent; reticulation lax, distinct or not. Inflorescences with
male and female flowers mixed (always?), sometimes pseudoterminal (terminal bud of
twig abortive), paniculate, branched from near the base; basal cataphylls caducous; bracts
caducous; flowers in small clusters, buds either in the same or in different stages of
development. Flowers short or long pedicellate, at base not articulated; bracteole absent.
Male flowers: perianth rotate, thinly leathery, inside pubescent, yellowish or (in New
Guinea) bright red; buds small, ellipsoid, ovoid, or subglobose, cleft to c. 3/4 to nearly
the base, lobes 3-5, spread or recurved in anthesis; androecium with androphore narrow,
glabrous, synandrium small, (depressed) globose or short-ellipsoid, with 2-6 completely
fused, short (0.5 mm or less) elliptic anthers. Female flowers: similar to male; ovary
ovoid, glabrous, stigma sessile, minute, 2-lobed, lobes narrow or broad and 2—5-lobu-
late. Infructescences small or large panicles, to 30 cm long. Fruits ellipsoid or (ob)ovoid,
glabrous, yellow or purplish; pericarp thin or thick, fleshy-leathery; aril yellowish (?) or
red, subentire or to halfway laciniate; seeds usually pointed at apex, variegated; albu-
men ruminate, without (?) starch; embryo incompletely known. — Fig. 3, 4.

Distribution — The genus has 4 species, ranging from South China through South-
east Asia and Malesia east to New Guinea; not in Central & East Java, Lesser Sunda
Islands, Solomon Islands or Australia. — Map | (see p. 3).

Note — In most species female flowers have not been seen; they should be found in
the large, paniculate, predominantly male-flowerd inflorescences, or in purely female
inflorescences.

30 Flora Malesiana, Ser. I, Vol. 14 (2000)

KEY TO THE SPECIES

la. Flowers in one cluster or semi-cluster all in about the same stage of development.
Synandrium depressed-globose, slightly broader than long. Androphore short, about

as long as or shorter than the synandrium. Anthers 4—6. Fruits 4.5-7 cmlong .. 2

b. Flowers in one cluster usually in different stages of development. Synandrium glo-
bose or short-ellipsoid, about as broad as long or longer than broad. Androphore
about as long as or longer than the synandrium or, in Java, sometimes shorter than

the synandrium. Anthers mostly 3-6. Fruits of Vatlable SIZES « ...6)se:sn- een 3

2a. All male perianths on a plant (3- or) 4-lobed. Anthers 4-6. Leaf buds, twig apex and
inflorescences with grey-brown hairs 0.1—0.2 mm long, sometimes glabrescent.
Nerves above flat or but little raised. Pericarp 2-10 mm thick . . 1. E. canarioides

b. Male perianths more or less evenly mixed 4- and 5-lobed. Anthers 4. Leaf buds, twig
apex and inflorescences with rusty hairs 0.5 mm long, sometimes late glabrescent.
Nerves raised above. Pericarp 1.5—3(-4) mm thick ..........- 3. E. rufirachis

3a. Leaves 8-20 cm long, drying greenish; nerves 7—12 pairs. Inflorescences weak and
slender, rather poorly flowered. Fruits 4.5—7 cm long, pericarp 5-8 mm thick, drying
BrOWies . Abe 2 Pa nenaatd «bei oath) SORE Iaee et aaa A oie 4. E. virella

b. Leaves larger, 15—35 cm long, usually drying dark brown, nerves 1 1—24 pairs. Inflo-
rescences variable. Fruits up to 4.5(—5.5) cm long, pericarp less than 5 mm thick,
Giyinetblackisht= 52 RETR ROI, nic wrt eee 2. E. macrocoma

1. Endocomia canarioides (King) W. J. de Wilde

Endocomia canarioides (King) W.J. de Wilde, Blumea 30 (1984) 190, f. 3e—h. — Horsfieldia
canarioides (King) Warb., Mon. Myrist. (1897) 294, t. 21; Gamble, Mat. Fl. Malay Penins. 5
(1913) 208; Ridl., Fl. Malay Penins. 3 (1924) 55. — Myristica canarioides King, Ann. Roy. Bot.
Gard. Calc. 3 (1891) 304, pl. 134. — Horsfieldia macrocoma (Mig.) Warb. var. canarioides
(King) J. Sinclair, Gard. Bull. Sing. 16 (1958) 389, f. 55. — Lectotype: King’s coll. 10064, Pe-
ninsular Malaysia.

Myristica racemosa King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 328, pl. 173. — Horsfieldia racemosa
(King) Warb., Mon. Myrist. (1897) 347; Gamble, Mat. Fl. Malay Penins. 5 (1912) 222; Ridl., FI.
Malay Penins. 3 (1924) 60. — Type: Curtis 934, Peninsular Malaysia.

2Embelia ridleyi King & Gamble, Mat. Fl. Malay Penins. 4 (1905) 112; J. Sinclair, Gard. Bull. Sing.
15 (1956) 31. — Type: Ridley 6324, Peninsular Malaysia.

Tree 10-35 m. Twigs (grey-)brown, 1.5—4(—10) mm diameter, with minute grey-brown
hairs 0.1 mm long, early glabrescent, bark striate, on older twigs sometimes longitudi-
nally cracking, lenticels very inconspicuous. Leaves membranous or chartaceous, ellip-
tic to oblong, 8-30 by 4-13.5 cm, base subcordate or rounded to short-cuneate, apex
acute-acuminate; upper surface dark brown, glabrous, lower surface early glabrescent;
midrib on both surfaces early glabrescent, above flat or slightly raised; nerves 11-19
pairs, lines of interarching distinct or not; venation lax, distinct or not on both surfaces;
petiole 8-30 by 1.5—4 mm; leaf bud 6-25 by 1-3 mm, with dense (grey-)brown hairs
0.1—0.2 mm long or less. Inflorescences predominantly with male flowers, generally
behind the leaves, (sub)glabrescent or with sparse greyish hairs 0.1 mm or less, variable
in shape, loose or condensed, many-flowered, 6-25 by 3-15 cm, peduncle 0.5-6 cm
long, bracts caducous, not seen. Flowers in loose or dense clusters of 3-8, all in about

De Wilde — Myristicaceae (Endocomia) 3]

the same stage of development; perianth outside with + sparse greyish hairs 0.1
(—0.2) mm, or glabrous, lobes inside with pale hairs 0.2—0.3 mm, often in longitudinal
rows. Male flowers: pedicel 1.5—2.5 mm; bud broadly ellipsoid-obovoid, 1.5—1.6 by
1.3-1.4 mm, apex broadly rounded or somewhat depressed, base + tapering, subcircular
in cross section, cleft 3/4—4/5, lobes 3 or 4 (or 5), at sutures 0.2(—0.3) mm thick;
synandrium depressed globose, 0.3—0.5 by 0.5—0.6 mm, anthers 4—6, androphore 0.3—
0.4 by 0.3 mm. Female flowers not seen; ovary in very immature fruits glabrous. Fruits
(solitary or) 2—6 together in panicles of 15—25 cm long, ellipsoid-oblong, (4.5—)5—7 by
2.5-3.5 cm, apex (narrowly) rounded, base rounded or narrowed for 3—5 mm, glabrous,
drying dark brown or blackish, finely granulate; pericarp 2-10 mm thick; fruiting pedi-
cel 15-20 mm long; aril laciniate for c. 1/3 to nearly halfway; seeds ellipsoid-oblong,
4—5 cm long, apex subacute or bluntly beaked for up to 3 mm long, testa purplish brown-
ish variegated. — Fig. 4e-h.

Field-notes — Bark smooth, grey- or dark-brown, flaking in small thin pieces, or fi-
nely fissured or cracked; slash inner bark brownish white, yellowish brown, or red, with
watery reddish exudate; slash wood white or pale yellow. Leaves glossy on both sur-
faces. Flowers (pale) green. Fruits green turning yellow (Peninsular Malaysia) or purple
brown (N Sumatra), ellipsoid-oblong, very large, to 10-12 cm long, mature aril yellow.

Distribution — Peninsular Thailand; Malesia: Sumatra, Peninsular Malaysia, Singa-
pore.

Habitat & Ecology — Evergreen, open bamboo forest, lowland rain forest, on flat
land or hillsides; 0-300 m altitude; fl. Jan.—June; fr. June—Sept.

Note — The fruits are very variable in the thickness of the pericarp. When more
material becomes available, two varieties may be distinguished through this character.
Fruiting specimens with thin pericarp may be difficult to distinguish from large-fruited
specimens of Endocomia macrocoma subsp. prainit.

2. Endocomia macrocoma (Miq.) W.J. de Wilde

Endocomia macrocoma (Miq.) W.J. de Wilde, Blumea 30 (1984) 182; Tree Fl. Sabah & Sarawak 3
(2000) 339. — Myristica macrocoma Migq., Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 207; 2 (1865)
49, p.p. (excl. specimens from Sulawesi = Horsfieldia irya). — Horsfieldia macrocoma (Miq.)
Warb., Mon. Myrist. (1897) 299, t. 21; J. Sinclair, Gard. Bull. Sing. 16 (1958) 392; 23 (1975) 75,
p.p. — Lectotype (here designated): Teijsmann 5553, Moluccas, Halmahera.

For more references and synonyms see the subspecies.

Trees 5-50 m. Twigs (pale) grey brown to dark brown, 2—6(—10) mm diameter, with
grey-brown to rusty hairs 0.1—0.5 mm, early to late glabrescent, bark (coarsely) striate
or longitudinally cracking, sometimes flaking, lenticels absent or inconspicuous. Leaves
membranous or chartaceous, elliptic or obovate to oblong-oblanceolate, (12—)15—35
(—40) by (4—)5—12 cm, base (narrowly) rounded to attenuate, apex acute-acuminate;
upper surface olivaceous to brown, glabrous, lower surface glabrous or (early) glabrescent;
midrib above slender, flat or slightly raised; nerves 11—24 pairs, lines of interarching
generally indistinct; venation lax or fine, sometimes slightly trabeculate, on both sur-
faces distinct or not; petiole 10-25 by 1.5—3.5 mm, glabrous to late glabrescent; leaf
bud 8-25 by 1.5-3.5 mm, with dense grey brown to rusty hairs 0.1—0.5 mm. Jnflores-
cences (see note 1) slender to stout, much or little branched, glabrescent or with persist-

32 Flora Malesiana, Ser. I, Vol. 14 (2000)

ent greyish or rusty hairs 0.1—0.5 mm long, in male 6-30 by 2-25 cm, moderately to
many-flowered, peduncle 0-5 cm long, bracts + elliptic, 1.5—3 mm, + thinly pubescent,
caducous. Flowers 4-10 together, buds usually in different stages of development, in
+ umbelliform clusters (3—)5—30 mm spaced along the main branches of the inflores-
cence; perianth outside early glabrescent or thinly with hairs 0.1-0.5 mm long, lobes
inside towards the apex with few to many (pale) brownish red hairs 0.2—0.4 mm long,
usually in rows in spaces between the anthers, sometimes only near the lobe sutures.
Male flowers: pedicel 1.5-7 mm long; buds broadly ellipsoid or broadly ovoid, 1.5—2.3
by 1.3-2 mm, apex rounded to subacute, base rounded to attenuate, in cross section sub-
circular or 3- or 4-angular, cleft 2/3—4/5, lobes 3 or 4 (or 5), at sutures 0.2—0.3(—0.5)
mm thick; synandrium (depressed) globose to ellipsoid, 0.2—0.5 by 0.3—0.5 mm, anthers
3—6, androphore slender, longer to shorter than the androecium, 0.3—1 mm long. Female
flowers: pedicel 2-6 mm; buds ovoid-ellipsoid, 2.2-2.6 by 1.7—2 mm, cleft 2/3—5/6,
lobes 0.2—0.3 mm thick; ovary (narrowly) ovoid, 1.3-1.8 by 0.8—1.2 mm, glabrous, stigma
0.2-0.3 mm high, narrowly to broadly 2-lipped, in the latter case the lips minutely 2—5-
lobulate. Fruits 3-12 in a pendent loose panicle (10—)15—30 cm long, ovoid, (narrowly)
ellipsoid, or obovoid, 1.5—4.5(—5.5) cm long, glabrous; pericarp 1-3 mm thick, black-
ish, finely granulate, not or sparingly warted; the aril almost entire to laciniate to about
halfway; seeds usually with pointed apex, testa variegated (not always in New Guinea).

Distribution — Widespread, continental Southeast Asia and Malesia; see under the
subspecies.

Notes — 1. The inflorescences often have mixed male and female flowers, the latter
possibly most frequent towards the end of the branches; other species may have separate
male and female inflorescences on the same twig.

2. The flowers in a subumbel or cluster are generally in different stages of develop-
ment, as in Endocomia virella, hence open flowers are found together with closed ones
and usually with much smaller, still growing flower buds.

KEY TO THE SUBSPECIES

la. Synandrium (0.3—)0.4—0.5 mm long, with (3 or) 4—6 anthers, androphore 0.3—0.6
mm long. Stigma broadly 2-lipped and minutely lobulate .........-...----- 2

b. Synandrium 0.2-0.3 mm long, with 2 or 3 anthers, androphore 0.7—1 mm. Stigma
narrowly 2-lipped. [Perianth inside greenish to pale yellowish. Fruits 34.5 cm long. ]

byae fStOS ots = Eieryel atte eTAS as book os bee DORNER ots Seer b. subsp. longipes

2a. Plants either early glabrescent or leaf buds, inflorescences and flowers with greyish
or pale brown hairs 0.1—0.2 mm long or less. Perianth inside greenish to yellowish
(continental Southeast Asia, W Malesia, Philippines) or red (New Guinea). Fruits
2:54 emulongascraiets er die vitro: Be re ee c. subsp. prainii

b. Leaf buds, inflorescences and flowers with + conspicuous rusty hairs 0.2-0.5 mm
long. Perianth inside greenish yellow. Fruits 1.7—3.5 cm long a. subsp. macrocoma

a. subsp. macrocoma

Endocomia macrocoma (Migq.) W.J. de Wilde subsp. macrocoma; W.J. de Wilde, Blumea 30 (1984)
184, f. 2i: 3d: 41 (1996) 375. — Horsfieldia macrocoma (Miq.) Warb. var. macrocoma; J. Sinclair,
Gard. Bull. Sing. 16 (1958) 393.

1oS)
io)

De Wilde — Myristicaceae (Endocomia)

Myristica nesophila Migq., Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 206, p.p. — For the syntype de
Vriese s.n. (L), Bacan I., not the lectotype = Horsfieldia.

Horsfieldia leptocarpa Warb., Mon. Myrist. (1897) 346, t. 21 (excl. Forster s.n., Sulawesi = Hors-
fieldia irya). — Horsfieldia leptosperma, nomen, in obs. sub Horsfieldia olivaeformis Warb.,
Mon. Myrist. (1897) 352. — Type: de Vriese s.n., fr., Sulawesi or Buru.

Gymnacranthera ibutii Holthuis, Blumea 5 (1942) 183, f. 4. — Type: Lam 2976, Talaud I.

Leaf bud and twigs rusty pubescent, hairs 0.2-0.5 mm long, twig apex sometimes
early glabrescent. Inflorescences rather condensed, much branched, 6—15 cm long, the
flowers densely rusty pubescent, hairs 0.2—-0.5 mm long. Pedicels of male and female
flowers 3-4 mm long. Male flowers: buds 3-lobed, cleft to c. 4/5, inside greenish yel-
low; synandrium subglobose to ellipsoid, 0.5 by 0.4—-0.5 mm, anthers 4, androphore
0.4—0.6 mm long. Female flowers: ovary + narrowly ovoid, stigma rather broadly
2-lipped and very finely lobulate. Infructescences 10—16 cm long. Fruits fusiform, ellip-
soid, + obovoid, or pear-shaped, 1.7—3.5 by 1.2—1.9 cm, apex acute to broadly rounded,
base rounded or narrowed; pericarp 1—3 mm thick; fruiting pedicel 6—12 mm long: aril
at apex laciniate 1/3—1/2; seeds + ellipsoid, 1.5—2 cm long, apex + acute, testa blotched
and variegated. — Fig. 3i, 4d.

Field-notes — No buttresses. Outer bark 0.2—0.7 mm thick, slightly fissured or not,
little peeling; inner bark 9-18 mm, pale red to pink-ochre, with some pale reddish wa-
tery exudate; sapwood whitish to yellowish tinged red, gradually passing into the darker
heartwood. Perianth (inside) greenish yellow. Fruits yellow or orange, aril bright red, at
apex incised to 1/3—1/2; seeds mottled brown.

Distribution — Malesia: Moluccas (Talaud, Morotai, Halmahera, Bacan, Obi, Buru,
Seram, Ambon); possibly Sulawesi (sterile coll.) and Philippines (NE Luzon); see notes.

Habitat & Ecology — Forest with little undergrowth, on alluvial flats locally with
stagnant water, or hill slopes; on porous volcanic soil, or loam soil with stones; also in
disturbed forest; 0—400 m altitude; fl. & fr. throughout the year.

Notes — |. Subsp. macrocoma is arbitrarily separated from subsp. prainii, the latter
being very variable in many features, and mainly differing by a much less developed
indumentum on leaf buds and inflorescences, and by its generally larger fruits. Subsp.
macrocoma has a remarkable variation in fruit shapes. Its fruits are generally + ellipsoid,
but fruits from Bacan and Obi are small, 1.7—2.2 cm long and either broad-ellipsoid or
obovoid; in Buru fusiform fruits 28 mm long are found.

2. Ridsdale c. s. ISU 317, from Palanan, NE Luzon, strongly deviates in its very hairy
appearance, and may represent a separate taxon.

b. subsp. longipes W. J. de Wilde

Endocomia macrocoma (Miq.) W.J. de Wilde subsp. longipes W.J. de Wilde, Blumea 30 (1984)
185, f. 2f-h; Tree Fl. Sabah & Sarawak 3 (2000) 339. — Type: de Vogel 888, SE Kalimantan.

Leaf bud with hairs dull brown, 0.1 mm long or less, twig apex early glabrescent.
Inflorescences lax, little to much branched, 6—30 cm long, with the flowers glabrous or
sparingly pubescent, hairs grey-brown, 0.1 mm long; male and female flowers with slender
pedicels (2-)4—7 mm long. Male flowers: buds 3- (or 4-)valved, cleft to 4/5—5/6, inside
greenish to yellowish; synandrium (depressed) globose, 0.2—0.3 by 0.2—0.4 mm, anthers
2 or 3, androphore slender, (0.6—)0.7—1 mm long. Female flowers: ovary narrowly ovoid,

Flora Malesiana, Ser. I, Vol. 14 (2000)

yh felties
Mae)
Mt Lo!

ages MAN
Ta

Na hhep \
St Sey
uy
\ Mages

+7}

Be 4 Ay

4 177
4 AUN We (faite are
% AEE AAI ea An
+e \ wR ue att! 2
veg a
ee
es
woe

v.

De Wilde — Myristicaceae (Endocomia) 35

stigma shortly and narrowly 2-lipped. Infructescences 15-30 cm long. Fruits narrowly
ovoid-ellipsoid, 3.5—4(—5.5) by 1.4—2 cm, apex blunt or subacute, base rounded or nar-
rowed; pericarp 1—2 mm thick; fruiting pedicel 15 mm long; aril laciniate to 1/5; seeds
3 cm long, the apex slightly acute or to 2 mm beaked, testa variegated with longitudinal
blotches. — Fig. 3f-h.

Field-notes — Bark of trunk greyish or chocolate brown, somewhat fissured or not
and little to profusely scaling in small thin pieces; outer bark 2 mm thick, brown; inner
bark 7-17 mm, cream, light brown(-red), or yellowish, with pale orange or reddish wa-
tery exudate; sapwood pale yellow or pale brown. Perianth inside yellowish or pale
green. Fruits hanging from the branches, green.

Distribution — Malesia: E Sumatra and Borneo (Sarawak, Sabah, E and S Kalimantan).

Habitat & Ecology — Primary forest, mixed dipterocarp forest, riverside forest, on
stream banks or alluvial flats, on deep clay soil, low ridges with sandy or sedimentary
soil; O-1000 m altitude; fl. & fr. throughout the year.

Note — Related to Endocomia virella, with similar flowers, but the latter has leaves
drying distinctly greenish, and much larger fruits.

c. subsp. prainii (King) W.J. de Wilde

Endocomia macrocoma (Miq.) W.J. de Wilde subsp. prainii (King) W. J. de Wilde, Blumea 30 (1984)
187, f. 3b, c. — Myristica prainii King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 299, pl. 126. —
Horsfieldia prainii (King) Warb., Mon. Myrist. (1897) 292, t. 21. — Types: King’s coll. 417,
431; Carter s.n., Andaman.

Horsfieldia papillosa Warb., Mon. Myrist. (1897) 291, t. 21. — Myristica papillosa (Warb.) Boerl.,
Handl. 3 (1900) 85. — Type: male specimen, cultivated in Bogor Botanical Garden,, origin
unknown.

Horsfieldia merrillii Warb. in Perkins, Fragm. Fl. Philipp. (1904) 49; Merr., Philipp. J. Sci., Bot. 2
(1907) 274; Enum. Philipp. Flow. Pl. 2 (1923) 182. — Types: Merrill 2233, 2370, Mindoro.
Horsfieldia oblongata Merr., Philipp. J. Sci., Bot. 13 (1918) 286; Enum. Philipp. Flow. Pl. 2 (1923)
182; Markegr., Bot. Jahrb. Syst. 67 (1935) 148 (for the New Guinean specimens only). — Type:

Ramos Philip. pl. 1393, Luzon.

Horsfieldia trifida A.C.Sm., J. Arnold Arbor. 22 (1941) 60. — Type: Brass & Versteegh 14017,

Irian Jaya.

Leaf bud pubescent, hairs greyish to dull brown, 0.1—0.2 mm long or less, twig apex
early glabrescent. Inflorescences condensed to lax, usually much-branched, 8-25 cm
long, with the pedicels and perianths sparingly to densely short-pubescent, hairs grey

Fig. 3. Inflorescences and flowers of Endocomia. — E. rufirachis (J. Sinclair) W.J. de Wilde. a. Flow-
ering twig; flowers all in about the same stage of development and rachis of inflorescence at base
without cataphylls; b. male flower in mature bud stage; c. ditto, at anthesis, note papillose-hairy
inner surface of perianth lobes; d. section of male perianth showing stalked androecium; e. longi-
tudinal section of androecium, schematic, androphore and central column drawn solid black. —
E. macrocoma (Miq.) W.J. de Wilde subsp. longipes W.J. de Wilde. f. Inflorescence with female
and male flowers, all in different stages of development; g. male flower at anthesis; note reflexed
perianth lobes and slender androphore; h. female flower at anthesis. — E. macrocoma subsp. macro-
coma. i. Female flower in section [a—e: BNB For. Dept. 1716; f-h: SAN 66753, 1: Atasrip 103]. —
Scale bar for a, f = 2 cm; for b—e, g—-i = 0.8 mm.

36 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Endocomia) 37

(-brown), 0.1—0.2 mm long; pedicels of male and female flowers slender, 1.5—2.5 mm
long. Male flowers: buds 3- or 4- (or 5-)lobed, cleft to (2/3—)3/4—5/6, inside greenish to
yellowish, or red; synandrium (depressed) globose or ellipsoid, (0.3—)0.4—0.5 by 0.3—
0.5 mm, anthers (3 or) 4, or 5 or 6 (New Guinea, see note 1), androphore slender, (0.2—
)0.3-0.6 mm long. Female flowers: ovary ovoid, with broad 2-lipped stigma.
Infructescences 10-30 cm long. Fruits narrowly to broadly ellipsoid to ovoid, (2.2—)3—
4.5(-5.5) by (1—)1.2—2.5 cm, apex obtuse or often subacute, base rounded, or with an up
to 5 mm long narrowed part; pericarp 1—2.5 mm thick; fruiting pedicel S—15 mm long;
aril almost entire (Philippines, New Guinea) or laciniate c. 1/5; seeds ellipsoid, 2—3.2
cm long, apex acute or shortly beaked, testa variegated by longitudinal markings, in
New Guinea not or only indistinctly so. — Fig. 4b, c.

Field-notes — Without or with short buttresses up to 200 x 30 x 4 cm, branches often
horizontally spreading, or drooping. Bark grey to blackish brown, smooth, without or
with shallow fissures, shallowly irregularly peeling or not; exudate watery, colourless or
pale red to brownish, once recorded as slightly milky; blaze pale brown to salmon; wood
white or straw or salmon-cream. Perianth inside greenish to yellow (W Malesia), once
purple (Thailand) or dark red to deep maroon (New Guinea), anthers creamy to pale
yellow, ovary green with brown or blackish stigma; flowers with sweet scent. Fruits
glossy green, turning yellow, in the Philippines and New Guinea orange, aril bright red.

Distribution — South China (Yunnan), India (Assam, Andaman I.), Bangladesh,
Burma, Indochina; Malesia: W Sumatra, W Java, Philippines, New Guinea.

Habitat & Ecology — Lowland and hillside forest, riverine or swamp forest; by streams,
on alluvial or clayey soil, in limestone country, or on copper-rich soil; 0-500 m altitude;
fl. & fr. throughout the year, but fl. in W Malesia mainly July, Aug., in the Philippines
mainly Mar.—May, in New Guinea throughout the year.

Uses — The wood is used for house-building (Sepik area).

Notes — 1. A variable subspecies, mainly in the indumentum, some features of the
male flowers, the shape and size of the fruits, and the colour of the seeds. This variation
is more or less correlated with the geography. Specimens from the Philippines differ in
inflorescences and indumentum, especially that of the flowers, composed of grey-white
rather long hairs 0.1—0.2 mm. Specimens from New Guinea stand apart. They are char-
acterized by generally shorter inflorescences and infructescences, androecium with 5 or
6 anthers, perianth dark red inside, fruits orange and small, sometimes only 2.2—2.5 cm
long, testa non-variegated or only faintly so. Specimens from outside New Guinea have
usually only 4 anthers, perianth greenish or yellowish inside, fruits yellow (once orange
in the Philippines) and the testa always variegated. In other features the New Guinea
specimens agree with those from elsewhere in the range of the species.

Fig. 4. Infructescences, fruits, and seeds of Endocomia. — E. rufirachis (J. Sinclair) W.J. de Wilde.
a. Infructescence. — E. macrocoma (Miq.) W.J. de Wilde subsp. prainii (King) W.J. de Wilde.
b. Infructescence; note complete arils and pointed seeds; c. ditto; note arils faintly laciniated, in-
completely covering the seeds pointed at apex. — E. macrocoma subsp. macrocoma. d. Infruc-
tescence. — E. canarioides (King) W.J. de Wilde. e. Fruit in spirit; f. ditto, opened; note deeply
laciniated aril; g & h. seed; note variegated testa [a: Kostermans 9579; b: PNH 7281 (Edafo);
c: Kostermans & Soegeng 478; d: Atasrip 103; e—g: de Wilde/Duyfjes 18877 (spirit); h: King’s coll.
10064). — Scale bar for all = 2 cm.

38 Flora Malesiana, Ser. I, Vol. 14 (2000)

2. In Endocomia the seeds are almost always variegated, except in E. macrocoma
subsp. prainii from New Guinea, where the seeds are dull greyish brown, not variegated
or only faintly so, possibly because the aril is not laciniate.

3. Endocomia rufirachis (J. Sinclair) W.J. de Wilde

Endocomia rufirachis (J. Sinclair) W.J. de Wilde, Blumea 30 ( 1984) 192, f. 2 a-e, 3a; Tree Fl. Sabah
& Sarawak 3 (2000) 340. — Horsfieldia macrocoma (Miq.) Warb. vat. rufirachis J. Sinclair,
Gard. Bull. Sing. 16 (1958) 393. — Type: Wood SAN A 4770, Sabah.

Tree 10-40 m. Twigs brown, 3.5—5(—14) mm diameter, with rusty hairs 0.5 mm long,
early to late glabrescent, bark coarsely striate, tending to flake, lenticels absent or incon-
spicuous. Leaves membranous to thinly chartaceous, elliptic-oblong to oblong-lanceo-
late, 18-38 by 6-11 cm, base subcordate, broadly rounded, or short-cuneate, apex acute-
acuminate; upper surface drying brown, glabrous, lower surface early glabrescent; midrib
on both surfaces often late glabrescent, above rather flat to moderately raised; nerves
17-25(—30) pairs, above (partly) raised, sometimes late glabrescent, lines of interarching
often distinct and regular; venation lax, distinct or faint above; petioles 8-25 by 1.5—3
mm, + late glabrescent; leaf bud 8-18 by 2-3.5 mm, with dense rusty hairs 0.3—0.5 mm.
Inflorescences (with male or mixed male and female (?) flowers) between or behind the
leaves, sometimes apical, 3-5 times branched, many-flowered, 8-30 by 5—25 cm, pe-
duncle 1.5—5 cm, with dense rusty hairs 0.5—0.7 mm long, bracts elliptic-oblong, 2-6
mm long, caducous. Flowers usually in rather dense clusters of 5-10, sometimes flow-
ers more dispersed and only 2 or 3 together, all + in the same stage of development,
perianth outside with hairs 0.2-0.5 mm long, lobes inside with dense rusty hairs 0.2—0.3
mm. Male flowers: pedicel slender, 2—3.5 mm; buds obovoid to + obconical, 1.5—1.8 by
1.2-1.5 mm, apex rounded to subtruncate, base + tapering, in cross section faintly an-
gular or subcircular, pubescent or glabrescent towards the apex, cleft 3/4—4/5, the lobes
(3 or) 4 or 5, oblong, 0.2 mm thick; synandrium depressed-globose, (0.2—)0.3 by 0.5—0.6
mm, anthers 4, sessile, androphore 0.3 by 0.2—0.3 mm. Female flowers not seen. Fruits
6-18 in large panicles up to 30 cm long, ellipsoid-oblong, 4.5—6.5 by 1.8-2.5 cm, apex
narrowly rounded, base subacute or often tapering into a 4-10 mm long narrowed part,
glabrous, drying blackish, finely granulate, without lenticels, not warted; pericarp 1.5—3
(—4) mm thick; fruiting pedicel 8-16 mm long; aril laciniate to 1/5—1/3; seeds ellipsoid-
oblong, 3.5—4.5 cm long, apex beaked for 0.2-4 mm, testa with elongate purplish and
brownish dapples. — Fig. 3a-e, 4a.

Field-notes — Buttresses sometimes present, short and rounded. Bark usually black-
ish, brittle, smooth with superficially longitudinal cracks or with paper-thin flakes; inner
bark 10-15 cm thick, pale yellow, (reddish) brown, or orange, cambium yellowish to
red; sapwood pale, whitish to brown-yellow, soft; exudate slight, pale red from inner
sapwood. Flowers yellow with reddish or brown-red indumentum. Fruits green to yel-
low, aril bright red.

Distribution — Malesia: Borneo (Sarawak, Sabah, E Kalimantan).

Habitat & Ecology — Primary and logged-over lowland rain forest on flat land and
hill slopes or on periodically inundated ground; on leached clays, loam soils, black soil,
also on sandstone and limestone; 0-400 m altitude; fl.& fr. throughout the year.

De Wilde — Myristicaceae (Endocomia) 39

Notes — 1. Characterized by the inflorescences in which all the flowers are in about
the same stage of development, by the conspicuous rusty indumentum on leaf buds and
inflorescences, and by the large fruits.

2. Endocomia rufirachis resembles Horsfieldia motleyi, which has similarly pubes-
cent flowers. Endocomia macrocoma subsp. macrocoma (Moluccas) may have a simi-
lar, rather conspicuous yellowish red indumentum, but differs in the one-clustered flow-
ers 1n various stages of development, the shape of the androecium, and the smaller fruits.

3. In spite of a fairly large number of flowering specimens, only male flowers have
been found. Some of the collections have male inflorescences with fruits separately
attached to the herbarium sheet; both are apparently collected from one single tree. In
the related Endocomia macrocoma, the female flowers are usually scattered in between
the more numerous male ones in one inflorescence, or can be found on separate female
inflorescences of the same tree.

4. Endocomia virella W. J. de Wilde

Endocomia virella W.J. de Wilde, Blumea 30 (1984) 194; Tree Fl. Sabah & Sarawak 3 (2000) 342.
— Type: SAN 49546 (Sadau), Sabah.

Tree 8—20 m. Twigs pale grey-brown, |.5—4 mm diameter, with grey hairs 0.1 mm or
less, early glabrescent, bark finely irregularly striate, slightly longitudinally cracking,
not flaking, lenticels absent. Leaves membranous to thinly chartaceous, elliptic(-oblong),
8-20 by 3.5—-8 cm, base cuneate, apex acute-acuminate; upper surface dark olivace-
ous to dull green, glabrous, lower surface olivaceous to dull pale green, glabrous; midrib
above slender, flat; nerves 7—12 pairs, above flat or but little raised, lines of interarching
distinct or not; venation lax, generally indistinct on both surfaces; petiole 10-18 by 1.5—
2 mm; leaf bud 6-10 by 0.6—1 mm, with rather sparse greyish hairs 0.1 mm or less.
Inflorescences slender and lax, 5—15 by 1.5—8(—10) cm, few-flowered, peduncle 0.5—2.5
cm, glabrous or glabrescent, hairs sparse, greyish or pale brown, less than 0.1 mm, early
caducous, bracts not seen. Flowers (male) in lax umbel-like clusters of 2—6 flowers,
5—20 mm spaced along the main branches of the inflorescence, the buds in one cluster
usually in different stages of development; perianth outside with sparse hairs less than
0.1 mm, soon glabrescent, lobes inside towards the apex with pale hairs 0.2—0.4 mm
long in between the anthers. Male flowers: pedicel slender, 4-6 mm long; buds broadly
ellipsoid or broadly ovoid, 1.8—2 by 1.5—1.6 mm, + 3- (or 4-)angular, apex rounded to
subacute, base (broadly) rounded, cleft c. 5/6, lobes 3 or 4, widely spreading or + re-
curved, 0.2 mm thick; synandrium globose or depressed globose, 0.2—0.3 by 0.3—0.4
mm, anthers 3 or 4, androphore slender, 0.8—1 by 0.2 mm. Female flowers not seen.
Fruits 1—4 in little-branched panicles 12—25 cm long, ellipsoid-obovoid to ellipsoid-
oblong, (4.5—)5—7 by 2.5—3.5 cm, apex rounded, base subattenuate or with an up to 7
mm long narrowed part, glabrous, drying bright to dark brown, finely granulate; pericarp
+ woody, 5-8 mm thick; fruiting pedicel 25-30 mm; aril laciniate to 1/4—1/3; seeds
ellipsoid-oblong, 4 cm long, apex subacute, testa elongately purple-brown variegated.

Field-notes — Once recorded with buttresses. Bark greenish, brown-yellow, yellow-
ish green, or black, smooth or scaly; inner bark either reddish, orange-yellow, with clear
red sap, smelling; sapwood white, soft or medium hard. Perianth greenish to yellowish,
anthers yellow. Fruits green.

40 Flora Malesiana, Ser. I, Vol. 14 (2000)

Distribution — Malesia: Borneo (Sarawak: 4th Div.; Brunei; Sabah: Beaufort Hill).

Habitat & Ecology — Primary forest on hillsides, ridges; brown or blackish soil;
0—400 m altitude; fl. Jan., May; fr. Jan., Aug., Oct.

Note — Characterized by the minute indumentum, the rather small leaves drying
green, the slender and lax inflorescences, the flowers which are in each cluster in differ-
ent stages of development and greenish when mature, the long-stalked synandrium of
only 3 or 4 anthers, and the large fruits drying brown. Specimens may resemble those of
the Bornean subspecies of Endocomia macrocoma, but they dry brown and have smaller
fruits.

GYMNACRANTHERA

Gymnacranthera (A. DC.) Warb., Ber. Pharm. Ges. 2 (1892) 227; Ber. Deutsch. Bot. Ges. 13 (1895)
82: Mon. Myrist. (1897) 131; Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 222; Ridl., Fl. Malay
Penins. 3 (1924) 61; J. Sinclair, Gard. Bull. Sing. 16 (1958) 434; 17 (1958) 96; R.T.A. Schouten,
Blumea 31 (1986) 451; W.J. de Wilde, Tree Fl. Sabah & Sarawak 3 (2000) 343. — Myristica
sect. Gymnacranthera A.DC., Ann. Sc. Nat. 4, 4 (1855) 31; Prodr. 14, 1 (1856) 200; Migq., Fl.
Ind. Bat. 1, 2 (1858) 63; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 304. — Type species: Myristica
paniculata A. DC. = Gymnacranthera paniculata (A. DC.) Warb.

Shrubs or trees, dioecious. Twigs often somewhat compressed, lenticellate, not flaky.
Leaves rarely brittle when dry, lower surface pale, not papillose, dots absent; reticula-
tion lax, faint. Inflorescences paniculate, branched from near the base (female more con-
densed); flowers in loose clusters, all in the same stage of development; basal cataphylls
caducous: bracts caducous. Flowers short-pedicellate, bracteole absent. Male flowers:
perianth urceolate, membranous or leathery, inside pubescent, yellow; buds ellipsoid,
cleft to about 1/2 or less, lobes (2 or) 3 or 4, slightly outcurved (in female + reflexed);
androecium sessile or with short and narrow androphore, synandrium (long-)ellipsoid,
with 5-12 linear anthers, dorsally connate to the central column, laterally and at apices
free, the connective sometimes slenderly protruding. F emale flowers: shorter and wider
than male, + ovoid; ovary broadly ovoid, pubescent, stigma small, sessile, 2-lobed, each
lobe entire or 3—6-lobulate. Infructescences paniculate, several-fruited. Fruits ellipsoid
(globose in G. canarica, S India), 2-3.5 cm long, hairy or glabrescent; pericarp thick-
leathery; aril laciniate to near the base (as in Myristica); seeds not variegated; albumen

ruminate, containing a fixed oil, starch absent; cotyledons divaricate, connate at base. —
Fig. 5-7.

Distribution — The genus has 7 species, of which one, G. canarica (King) Warb., in
S India, and 6 in S Thailand and Malesia east to E New Guinea; not known from Java,
the Lesser Sunda Islands, and Palawan (Philippines). — Map 2 (see p. 2):

KEY TO THE SPECIES

la. Male perianth 4-6 mm long; androecium shorter than the perianth tube. Young twigs
and inflorescences with woolly hairs 1(—2) mm long; lower leaf surface with persist-
ent woolly hairs 0.5 mm long. Leaves 18-42 cm long, petiole 3-5 mm thick. Fruits
2.5-3.5 cm long, with hairs 0.3-0.5 mm. — Peninsular Malaysia, Sumatra, Borneo
eayh eaatealeteaety mad. 2 Lane ate. LA RE ik Gad ie i Menor 1. G. bancana

De Wilde — Myristicaceae (Gymnacranthera) 4]

b. Male perianth 2—5 mm; androecium as long as the perianth tube. Young twigs and
inflorescences glabrescent, or with hairs less than 0.5 mm; lower leaf surface gla-
brescent or with appressed hairs, indumentum less than 0.5 mm high. Leaf size vari-
able, petiole 1-3 mm diameter. Fruits 2-3 cm long, pubescent or glabrescent .. 2

2a. Young twigs with + woolly hairs to 0.5 mm long. Leaves densely pubescent below.
Fruits 2.3-3 by 2—2.2 cm, indumentum conspicuous, pericarp 3—5 mm thick. —
AIRES AWCSI Hy: 5 0 OU tok Sa ee eee eae oe Bee: 5. G. maliliensis

b. Young twigs glabrescent or variously pubescent by appressed hairs, less than 0.2
mm long. Leaves below glabrescent or with + scattered hairs. Fruits generally smaller,
Pldorescent OF pubescent, pericarp up to 2 mm thick 2:2". 722 heer eee. - 3

3a. Twigs, including the apical part, conspicuously densely set with lenticels. Lower
leaf surface with brownish hairs. Midrib flat above; nerves at c. 45° to the midrib in
the middle of the leaf. Fruits ellipsoid-ovate with truncate base, 2 cm long, short-
Bobecccitte | DOMEeONE 2.208 tnt. os RT ares Sab ee: coe 6. G. ocellata

b. Twigs towards the apex without or with but a few lenticels. Lower leaf surface
glabrescent or with scattered, inconspicuous, greyish or pale brown hairs. Nerves at
more than 45° to the midrib. Fruits globose to ellipsoid-oblong, base not truncate,
Peo cin lone. elabrescent or pubescent, 2244 nes eee eer eee ee 4

4a. Twigs at apex 1-2 mm, about 10 cm lower down 2—3.5 mm diameter. Leaves 5—17
(—27) by 1.5—5.5(-8.5) cm; midrib flat or sunken above. — W & E Malesia (in
E Malesia not rarely twigs thicker and leaves larger) ....... 3. G. farquhariana
b. Twigs at apex (2—)2.5—4 mm, about 10 cm lower down (3—)3.5—5.5 mm diameter.
Leaves larger, 14-33 by (5.5—)6—13 cm; midrib usually sunken ............. 5
5a. Lateral nerves on lower leaf surface distinct but little prominent, at 60—70° to the
midrib in the middle of the leaf; blade drying flat. Anthers 6(—8), straight. — Borneo
5c do ie SRR ae gaa eM Oe: PAR ee Rate Alaa 2 2. G. contracta

b. Lateral nerves on lower leaf surface very dinstinct and prominent, at (35—)40—50
(—55)° to the midrib in the middle of the leaf; blade generally drying irregularly un-
dulate. Anthers 6-10, sometimes twisted. — S Thailand, W Malesia . 4. G. forbesii

1. Gymnacranthera bancana (Miq.) J. Sinclair

Gymnacranthera bancana (Miq.) J. Sinclair, Gard. Bull. Sing. 16 (1958) 436, f. 53, pl. XH A; 17
(1958) 99: R.T.A. Schouten, Blumea 31 (1986) 463, f. 3a—f. 4; W.J. de Wilde, Tree Fl. Sabah &
Sarawak 3 (2000) 345. — Myristica bancana Miq., Fl. Ind. Bat., Suppl. (1861) 383; Warb.,
Mon. Myrist. (1897) 518. — Type: Teijsmann 3279, Sumatra, Bangka.

Myristica murtonii Hook. f., Fl. Brit. India 5 (1886) 105; King, Ann. Roy. Bot. Gard. Calc. 3 (1891)
297, pl. 124 ter. — Gymnacranthera murtonii (Hook. f.) Warb., Mon. Myrist. (1897) 357, t. 20;
Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 223; Ridl., Fl. Malay Penins. 3 (1924) 61. — Type:
Murton 13, Singapore.

Myristica ferruginea Wall. ex King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 298, pl. 125. — Type:
Wallich Cat. n. 6803 (lecto), Singapore.

Myristica amplifolia Warb., Mon. Myrist. (1897) 517. — Type: Anonymous n. 16 ‘Medang Simpai’,
Palembang, Sumatra.

Gymnacranthera murtonii (Hook. f.) Warb. var. borneensis Warb., Mon. Myrist. (1897) 359. —
Myristica murtonii Hook.f. var. borneensis (Warb.) Boerl., Handl. 3 (1900) 88, nom. alt. —
Gymnacranthera bancana (Migq.) J. Sinclair var. borneensis (Warb.) J. Sinclair, Gard. Bull. Sing.
16 (1958) 439; 17 (1958) 100. — Syntypes: Beccari 1211, 3977, Kuching, Sarawak.

14 (2000)

Flora Malesiana, Ser. I, Vol.

“ ‘
Wo
f

ay

De Wilde — Myristicaceae (Gymnacranthera) 43

Tree 15—40 m. Twigs 3—6(—8) mm diameter, rusty woolly-tomentose with hairs 0.5—1
mm long, glabrescent, greyish, bark smooth or finely cracked, densely set with lenticels.
Leaves coriaceous, elliptic to lanceolate, 18—42 by 7.5—-19 cm, base short-attenuate to
rounded or subcordate, apex acute(-acuminate), margin often revolute; upper surface
olivaceous, often glossy, lower surface grey-brown, hairs dense, rusty, 0.5 mm long,
glabrescent; midrib above flat, 1-2 mm wide, nerves (13—)15—23 pairs, at 60—70° to the
midrib, prominent below, venation indistinct on both surfaces; petiole late glabrescent,
10-20 by 3-5 mm; leaf buds 10—20 by 4-6 mm, with dense brown hairs 0.5—1 mm.
Inflorescences paniculate, with hairs 1-2 mm long; in male 6—10 cm long, up to 6 cm
wide, many-flowered, in female 2—5 cm long, fewer-flowered; bracts triangular, 2—4
mm long, pubescent, caducous. Flowers rusty pubescent inside and outside, hairs 0.2—
0.3 mm long. Male flowers: pedicel 1-3 mm long; buds ellipsoid-oblong, 4-6 by 2-3
mm, cleft 1/4 to nearly 1/2, lobes (long) triangular, slightly spreading, tube 2.5—3.5 mm
long; androecium truncately ellipsoid, (sub)sessile, 1.5—2 by 0.8—1.2 mm; anthers (7—)9
or 10, subsessile, free apices + erect, 0.2-0.3 mm long. Female flowers (immature):
pedicel 1 mm long; buds coriaceous, ovoid, 4-6 mm long, lobes 3; ovary subglobose,
densely pubescent, stigma sessile. Fruits 2-8 per infructescence, ellipsoid(-oblong), 2.2—
3.5 by 1.5—-2.2 cm, with rusty hairs 0.3(—0.5) mm long; pericarp 3 mm thick; fruiting
pedicel 3-5 mm long. — Fig. 5a-f; Photo 1.

Field-notes — A handsome tree, especially when in flower; crown dense or spread-
ing; bole smooth, no buttresses. Bark brown to grey, slightly fissured, finely or thickly
flaky, or scaly; slash wood white to yellow. Flowers golden yellow with a brownish
tinge, with a spicy odour when crushed.

Distribution — Malaysia: Sumatra (Aceh, Indragiri, Jambi, Palembang, Bangka, Riau),
Peninsular Malaysia (Johore), Singapore, Borneo (Sarawak, Brunei).

Habitat & Ecology — Primary and degraded dryland forest, also swamp forest and on
hillsides and ridges, on granite, sand and sandy loam soil; up to 250 m altitude; fl. Sept.,
Oct.; fr. throughout the year.

2. Gymnacranthera contracta Warb.

Gymnacranthera contracta Warb., Mon. Myrist. (1897) 360, t. 20 (excl. Motley 1284 = Gymna-
cranthera forbesii); J. Sinclair, Gard. Bull. Sing. 16 (1958) 439, f. 54, p.p.; 17 (1958) 100, p.p.;
R.T.A. Schouten, Blumea 31 (1986) 471, f. 4; W.J. de Wilde, Tree Fl. Sabah & Sarawak 3 (2000)
347. — Myristica contracta (Warb.) Boerl., Handl. 3 (1900) 88. — Type: Beccari 32] (male,
female), Sarawak.

Tree 5—26 m. Twigs subterete to angular or + ridged, 3-4(—5.5) mm diameter, hairs
minute, early glabrescent, bark chocolate to greyish brown, finely longitudinally cracked

Fig. 5. Gymnacranthera bancana (Miq.) J. Sinclair. a. Habit of leafy twig; b. male inflorescence;
c. male flower; d. ditto, opened, showing porportionally small androecium; e. fruits; f. fruit, opened,
showing thin pericarp and deeply laciniated aril of seed. — G. ocellata R.T. A. Schouten. g. Female
flower, opened, showing pubescent ovary with obliquely 2-lipped sessile stigma, each lip shallow-
ly lobulate; h. fruits, note truncate bases [a: SF 39502 (Sinclair), b—d: SF 40045 (Sinclair); e, f:
Sinclair & Kadim 10436; g: Endert 4835; h: S 39255 (Ilias Paie)|. — Scale bar for a, b, e, h = 2 cm;
for cdf o/=— 1865 min for f= 1icm:

44 Flora Malesiana, Ser. I, Vol. 14 (2000)

with lenticels when older. Leaves chartaceous or subcoriaceous, elliptic-oblong to lan-
ceolate, sometimes + parallel-sided, (16—)20-29 by 6-9.5 cm, base short-attenuate to
broadly rounded, apex (acute-)acuminate, margin flat; upper surface + brown, often glossy,
lower surface greyish to violaceous, with scattered appressed hairs, glabrescent; mid-
rib above slightly sunken (grooved), narrow, | mm wide, nerves (11-)13-18 pairs, at
60—70° to the midrib, below slender, distinct but not much prominent, venation indis-
tinct at both surfaces; petiole 10-20(-25) by 2-2.5 mm; leaf buds 4-8 by 2-3 mm, some-
times with scale-like cataphylls, densely minutely appressed-pubescent. Inflorescences
in male (broadly) paniculate, 3.5—4.5 cm long, up to 4 cm wide, many-flowered, with
rusty hairs 0.3-0.5 mm long, in female 1.5—2 by | cm, fewer flowers but more dense;
bracts broadly triangular, 2.5 by 3 mm, pubescent, caducous. F’ lowers rusty pubescent
inside and outside, hairs 0.10.2 mm long. Male flowers: pedicel 2—2.5 mm long; buds
ellipsoid-oblong, 2.5-2.8 by 1.5—2 mm, cleft 1/3 to nearly 2/3, lobes 3 or 4, triangular,
slightly spreading, tube 1-1.8 mm long; androecium truncately ellipsoid-oblong, 1.7 by
0.6-0.7 mm, up to 0.2 mm stiped; anthers 6(—8), subsessile, free apices 0.3 mm long,
erect. Female flowers: pedicel 1-2 mm long; buds narrowly obovoid, 3—3.5 by 2 mm,
cleft 2/3—3/4, lobes 3 or 4, long-triangular, curved outward or reflexed, tube 0.7—1 mm
long; ovary subglobose, minutely pubescent, 1.2 mm diameter, stigma sessile, minutely
2-lobed. Fruits c. 7 per infructescence, ellipsoid, 2—2.2 by 1—-1.4 cm, with hairs 0.1 mm,
or early glabrescent; pericarp 1.2—1.4 mm thick; fruiting pedicel 3—7 mm long.

Field-notes — Bark reddish brown, nearly smooth, very fine scaly; inside hard, pale
reddish brown. Flowers yellow. Fruits dark red.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah; possibly also W Kali-
mantan: Suzuki 9993).

Habitat & Ecology — Primary lowland forest, 0-580 m altitude; fl. July—Sept.; fr.
Oct.—Dec.

3. Gymnacranthera farquhariana (Hook. f. & Thomson) Warb.

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb., Mon. Myrist. (1897) 365, t. 20; R.T.A.
Schouten, Blumea 31 (1986) 476, f. 7; W.J. de Wilde, Tree FI. Sabah & Sarawak 3 (2000) 348.
— Myristica farquhariana Wall. ex Hook. f. & Thomson, Fl. Ind. (1855) 161, p.p. — Lectotype:
Wallich Cat. no 6795 (K), Singapore.

For more references and synonyms see the varieties.

Tree 3-30(—45) m. Twigs faintly angular, + ridged or lined when young, 1—2(-3.5)
mm diameter, with hairs 0.1 mm, early glabrescent, bark chocolate or grey, later on grey-
brown, smooth, lenticellate. Leaves thinly chartaceous to coriaceous, elliptic to lanceo-
late, 5-17 by 1.5—5.5(—6) cm, in E Malesia up to 27 by 8.5 cm, apex acute(-acuminate),
base attenuate, margin conspicuously revolute or not, olivaceous or brown, sometimes
glossy above; lower surface with sparse appressed hairs 0.1 mm, glabrescent, pale brown
to grey-purplish; midrib above flat or sunken (grooved), to | mm wide; nerves 7-11
(—15) pairs, flat or raised below, discolorous or not, venation indistinct; petiole 6—15
(-18) by 1—-2.5 mm; leaf buds slender, 5-10 by 1-2 mm, with dense appressed greyish
hairs 0.1 mm. Inflorescences paniculate, with rusty hairs 0.2 mm long; in male 2.5—12
cm long, up to 8 cm wide, many-flowered, in female up to 4 by 2 cm, fewer flowered;
bracts broadly triangular, 1 by 2 mm, pubescent, caducous. Flowers rusty pubescent
inside and outside, hairs 0.1 mm long. Male flowers: pedicel 1.5—4 mm long; buds ellip-

De Wilde — Myristicaceae (Gymnacranthera) 45

soid(-oblong), 2.54 by 2-3 mm, cleft 1/3—2/3, lobes 3 or 4, (long-)triangular, slightly
spreading, tube 1—2.3 mm long; androecium truncately ellipsoid-oblong, 1—2.5 by
0.8—1 mm, stiped up to 0.3 mm; anthers (6 or) 7—11(—13), subsessile, free apices up to
0.5 mm long, erect or somewhat incurved. Female flowers: pedicel 1.5—3 mm long, buds
ovoid or obpyriform, 2-3 by 1.5-2 mm, cleft 1/2—3/4, lobes 3 or 4, long-triangular,
strongly spreading or recurved, tube short-urceolate, 0.7—1.5 mm long; ovary subglobose,
1 mm diameter, densely minutely pubescent, stigma minute, subsessile. Fruits up to 13
per infructescence, subglobose to ellipsoid-oblong, 1.8—2.8 by 1.1—1.9 cm, glabrescent
(glabrous) or inconspicuously pubescent, hairs 0.1 mm long; pericarp | mm thick; fruit-
ing pedicel 4-15 mm long.

Distribution — A widespread species ranging from Peninsular Malaysia to the Bis-
marck Archipelago. Four varieties of Gymnacranthera farquhariana are recognized to
accommodate its particularly complex variation. The varieties are very close to each
other, and several specimens are not easily placed within a variety.

KEY TO THE VARIETIES

la. Leaves oblong-lanceolate, small, 5—13.5 by 1.54.5 cm; nerves below not or hardly
raised, i.e. usually they cannot be felt with the finger. Fruits short-ellipsoid to glo-
bose. — Sumatra, Peninsular Malaysia, Borneo ............ b. var. eugeniifolia

b. Leaves small or large, 6-27 by 2-8.5 cm; nerves below usually clearly visible and
contrasting, usually distinctly raised and to be felt with the finger. Fruits various 2

2a. Leaves coriaceous, elliptic(-oblong), 6—15(—17) by 3—5.5(—6) cm, usually with con-
spicuously revolute edge. [Fruits globose to short-ellipsoid.] — Sumatra, Peninsular

PNA SEAS OLE ON peers, as tore aye Sips os Ny os eahnl os anche ah 2 clea a. var. farquhariana
b. Leaves membranous, chartaceous, or sometimes coriaceous, (elliptic-oblong to) lan-
ceoltcs 827 by2—8.5 cm, marein flat, rarely revolute... pan eee ie eee 3

3a. Fruits globose to short-ellipsoid, fruiting pedicel 8-15 mm long. — Philippines . . .
rn lites. CL OE. fe, Hoth oe. Qt ee See c. var. paniculata

b. Fruits ellipsoid to oblong, rarely subglobose (Moluccas), fruiting pedicel 4-8 mm
long. — W Malesia (nerves on lower surface of blade usually pale yellowish), E Ma-
lesia (nerves usually reddish brown and contrasting), rare in the Philippines......
nae 2 RRR Be ieee asm irae Seti pe 8s) 99 d. var. zippeliana

a. var. farquhariana

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. farquhariana; R.T.A. Schouten,
Blumea 31 (1986) 477; W.J. de Wilde, Tree Fl. Sabah & Sarawak 3 (2000) 349. — Gymna-
cranthera farquhariana (Hook. f. & Thomson) Warb., Mon. Myrist. (1897) 365, t. 20, p.p. —
Myristica farquhariana Wall. ex Hook. f. & Thomson, FI. Ind. (1855) 161, p.p.; A.DC., Prodr.
14, 1 (1856) 200, p.p.; Mig., Fl. Ind. Bat. 1, 2 (1858) 63, p.p.; Hook. f., Fl. Brit. India 5 (1886)
108, p.p.; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 305, pl. 138.

Myristica griffithii Hook. f., Fl. Brit. India 5 (1886) 109 (excl. Maingay 1306A & B = var. eugenii-
folia); King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 304, pl. 135 (excl. Curtis 2406, 2458 = Hors-
fieldia penangiana). — Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. griffithii
(Hook. f.) Warb., Mon. Myrist. (1897) 368; Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 226,
p.p.; Ridl., Fl. Malay Penins. 3 (1924) 62. — Gymnacranthera eugeniifolia (A. DC.) J. Sinclair
var. griffithii (Hook. f.) J. Sinclair, Gard. Bull. Sing. 16 (1958) 447, f. 57; 17 (1958) 113. —
Lectotype: Griffith 4356 (K), Malacca.

46 Flora Malesiana, Ser. I, Vol. 14 (2000)

Myristica farquhariana Wall. ex Hook. f. & Thomson var. major King, Ann. Roy. Bot. Gard. Calc.
3 (1891) 306, pl. 136, f. 4. — Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var.
major (King) Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 226; Ridl., Fl. Malay Penins. 3
(1924) 62. — Lectotype: Griffith 4355 (K), Malacca.

Tree 3-30 m. Leaves coriaceous; blade elliptic(-oblong), widest at or above the mid-
dle, 6—15(—17) by 3-5.5(—6) cm, usually with a conspicuously revolute margin; nerves
7-11 pairs, on the lower leaf surface distinct and discolorous or not,:but always clearly
raised and to be felt with the finger; petiole 8-18 by 1.5—2 mm. Fruits 1—6 per infruc-
tescence, globose to short-ellipsoid; fruiting pedicel 6-15 mm long. — Photo 3.

Field-notes — Crown dense or spreading; bole smooth, in peat swamps sometimes
with buttresses or with a few stilt-roots. Bark dark brown, brittle; slash wood whitish.
Flowers yellow. Fruits yellow to orange, very spicy.

Distribution — Peninsular Thailand; Malesia: S Sumatra (Lampung), Peninsular
Malaysia, Singapore, Borneo.

Habitat & Ecology — Primary and degraded forest; mostly in peat swamp forest,
occasionally on hillsides; 0-1000 m altitude; fl. mainly Apr.—July; fr. Sept.—Mar.

b. var. eugeniifolia (A. DC.) R.T.A. Schouten

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. eugeniifolia (A.DC.) R.T.A.
Schouten, Blumea 31 (1986) 480; W.J. de Wilde, Tree Fl. Sabah & Sarawak 3 (2000) 349. —
Myristica eugeniifolia A.DC., Ann. Sc. Nat. 4, 4 (1855) 29; Prodr. 14, 1 (1856) 190; Miq., Fl.
Ind. Bat. 1, 2 (1858) 58; Hook. f., Fl. Brit. India 5 (1886) 113. — Gymnacranthera eugeniifolia
(A.DC.) J. Sinclair, Gard. Bull. Sing. 16 (1958) 444, p. p. — Gymnacranthera eugeniifolia
(A. DC.) J. Sinclair var. eugeniifolia; J. Sinclair, Gard. Bull. Sing. 16 (1958) 444, f. 56, pl. XIV,
p.p.; 17 (1958) 112, p.p. — Type: Gaudichaud 116, Penang.

Myristica farquhariana auct. non Hook. f. & Thomson, p.P.: Hook. f., Fl. Brit. India 5 (1886) 108;
King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 305. — Gymnacranthera farquhariana auct. non
(Hook. f. & Thomson) Warb., p.p.: Warb., Mon. Myrist. (1897) 365; Gamble, Mat. Fl. Malay
Penins. 5, 23 (1912) 225; Ridl., Fl. Malay Penins. 3 (1924) 62.

Myristica griffithii auct. non Hook. f.: Hook. f., FI. Brit. India 5 (1886) 109 (as for syntype Maingay
1306 only).

Gymnacranthera apiculata Warb., Mon. Myrist. (1897) 359, t. 20. — Myristica apiculata (Warb.)
Boerl., Handl. 3 (1900) 88. — Type: Beccari 2246, Borneo, Sarawak.

Tree 3-30 m. Leaves chartaceous or coriaceous; blade oblong-lanceolate, widest at or
below the middle, 5—13.5 by 1.5—4.5 cm, margin little or conspicuously revolute or not;
nerves 6-10 pairs, on lower leaf surface not or hardly raised, usually concolorous and
not to be felt with the finger; petiole 7-14 by 1-1.5 mm. Fruits 1—3(—5) per infructescence,
subglobose, 1.8—2.2 by 1.3-1.8 cm; fruiting pedicel 4-13 mm long.

Field-notes — Crown small, narrow, dense or not; bole smooth, no buttresses. Bark
brown(-grey), finely fissured, with small scales; wood white to pale brown. Flowers bright
yellow. Fruits green turning golden yellow to orange, very spicy.

Distribution — Malesia: throughout Sumatra, Peninsular Malaysia, Singapore, and
Borneo.

Habitat & Ecology — Primary and degraded forest, on dry land (hillsides and ridges)
as well as in (periodically) wet places, near streams and rivers, and in kerangas; found
on limestone and on sandy soils; 0-1300 m altitude; fl. mainly Mar.—Oct.; fr. July—Feb.

De Wilde — Myristicaceae (Gymnacranthera) 47

Note — The distribution of var. eugeniifolia largely coincides with that of var.
farquhariana. Specimens from Borneo here referred to var. eugeniifolia were formerly
sometimes determined as Gymnacranthera contracta, a species now accepted in a much
more restricted sense. Also specimens intermediate with var. zippeliana occur, espe-
cially in Sabah and Sarawak.

c. var. paniculata (A.DC.) R.T.A. Schouten

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. paniculata (A.DC.) R.T.A.
Schouten, Blumea 31 (1986) 481. —Myristica paniculata A.DC., Ann. Sc. Nat. 4, 4 (1855) 31;
Prodr. 14, 1 (1856) 200; Mig., Fl. Ind. Bat. 1, 2 (1858) 63; Fern.-Vill., Nov. App. (1880) 177;
Vidal, Phan. Cuming. (1885) 139; Rev. Pl. Vasc. Filip. (1886) 221.— Gymnacranthera paniculata
(A. DC.) Warb., Mon. Myrist. (1897) 370, t. 20; Merr., Philipp. J. Sci., Suppl. 1 (1906) 55; Enum.
Philipp. Flow. Pl. 2 (1923) 181; Elmer, Leaf]. Philipp. Bot. 3 (1911) 1059; J. Sinclair, Gard. Bull.
Sing. 17 (1958) 104. — Gymnacranthera paniculata (A. DC.) Warb. var. paniculata; J. Sinclair,
Gard. Bull. Sing. 17 (1958) 104, f. 2. — Type: Cuming 901, Luzon.

Myristica farquhariana auct. non Hook. f. & Thomson, p.p.: Hook. f., Fl. Brit. India 5 (1886) 108
(for the Philippine specimen).

Gymnacranthera laxa Elmer, Leafl. Philipp. Bot. 8 (1915) 2772. — Type: Elmer 13715, Philip-
pines, Mindanao.

Gymnacranthera acuminata Merr., Philipp. J. Sci., Bot. 12 (1917) 265; Enum. Philipp. Flow. Pl. 2
(1923) 181. — Type: FB 21074 (Cenabre & Cortes), Philippines, Samar.

Gymnacranthera macrobotrys Merr., Philipp. J. Sci., Bot. 13 (1918) 284; Enum. Philipp. Flow. PI.
2 (1923) 181. — Type: BS 1171 (Ramos), Philippines, Leyte.

Tree 8-14 m. Leaves chartaceous; blade (oblong-)lanceolate, 9-21 by 3-6 cm, mar-
gin not revolute; nerves (8—)9-11 pairs, on lower leaf surface distinct, sometimes
discolorous, little raised; petiole 8-15 by 1-2 mm. Fruits 1 or 2(-4) per infructescence,
subglobose or short-ellipsoid, 1.8-2.3 by 1.5—1.9 cm (when large with marked ridge on
the suture); fruiting pedicel 8-15 mm long.

Field-notes — Bark smooth, brittle, pale grey-brown, | cm thick. Fruits orange; seeds
banded brown and black, with but little spicy taste.

Distribution — Malesia: Philippines.

Habitat & Ecology — Forest on ridges as well as by rivers and lakes; up to 1400 m
altitude; fl. mainly Apr—June, Aug.—Oct.; fr. Jan.—July.

Note — This variety is restricted to the Philippines. It is very close to var. zippeliana
from which it differs only in the fruits; in var. zippeliana the fruits are usually ellipsoid-
oblong, not subglobose, and have a shorter fruiting pedicel. FB 2/074 (the type of G.
acuminata) and Sulit 14603, from Samar, deviate in rather small leaves with glossy
upper surface.

d. var. zippeliana (Miq.) R.T.A. Schouten

Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. zippeliana (Miq.) R.T.A. Schouten,
Blumea 31 (1986) 482: W.J. de Wilde, Tree Fl. Sabah & Sarawak 3 (2000) 349. — Myristica
zippeliana Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 50; Scheff., Ann. Jard. Bot. Buitenzorg |
(1876) 45. — Gymnacranthera zippeliana (Miq.) Warb., Mon. Myrist. (1897) 373. — Gymna-
cranthera paniculata (A. DC.) Warb. var. zippeliana (Migq.) J. Sinclair, Gard. Bull. Sing. 17 (1958)
108, f. 3. — Type: Zippelius s.n., Irian Jaya, Bird’s Head.

48 Flora Malesiana, Ser. I, Vol. 14 (2000)

Gymnacranthera suluensis Warb., Mon. Myrist. (1897) 373; Elmer, Leafl. Philipp. Bot. 3 (1911)
1058: Merr., Enum. Philipp. Flow. Pl. 2 (1923) 181. — Syntypes: Vidal 3546, 3561, Philippines,
Sulu I., Basilan.

Tree 3—-30(—45) m. Leaves thickly membranous or chartaceous, sometimes sub-
coriaceous; blade (oblong-)lanceolate, widest at or above the middle, 7.5—27 by 3.5-8.5
cm, margin not revolute; nerves 8-1 1(—15) pairs, on lower leaf surface distinct or not,
but always clearly raised, either pale yellowish, concolorous (W Malesia), or reddish or
purplish brown, discolorous (generally in E Malesia); petiole 7-15 by 1—2.5(-3) mm.
Fruits (1 or) 2-13 per infructescence, ellipsoid to oblong, rarely subglobose, (1.5-—)
1.8-2.5 by 1.1-1.5 cm; fruiting pedicel 4-8 mm long.

Field-notes — Crown dense, narrow; bole smooth, no buttresses. Bark brown or grey,
slightly fissured and finely flaky or scaly; inner bark dark brown, 5-14 mm thick, slash
wood white to yellow; heartwood yellow to brown, hard. Flowers golden yellow to brown,
odourless or faintly sweet; androecium brownish; pollen whitish. Seeds dark brown.

Distribution — Malesia: Peninsular Malaysia, S Philippines, and New Guinea (incl.
the Bismarck Archipelago), not in Java or the Lesser Sunda Islands. The most wide-
spread taxon of Gymnacranthera.

Habitat & Ecology — Variable; found in primary or in degraded and secondary for-
est, mostly on hillsides and ridges, in New Guinea usually on foothills and riverbanks,
also near the coast: on sandstone, clay, loam, and granite rock; 0-900 m altitude, in Bor-
neo 400-1200 m; fl. mainly Jan.June, Aug.—Oct. (Borneo mainly July—Nov.); fr. Jan.—
June, Aug.—Sept. (Borneo Mar.—May, Oct.—Nov.). Locally abundant, but frequently
mentioned as growing in regenerating forest.

Uses — In Papua Barat (Bird’s Head) the bark, together with lime, is used to prepare
the skins of birds.

Notes — 1. Gymnacranthera farquhariana var. zippeliana is variable in leaf size. In
Sulawesi, Moluccas, and New Guinea in particular, specimens with large leaves up to 27
cm long are found. The fruits also are variable in shape and size. Particularly distinctive
fruits are found in de Vogel 3789, 3795, 3955, from Bacan I., which have small, almost
globose fruits 1.2 cm diameter; the trees are 40—45 m high (other specimens have never
been recorded as being taller than 33 m). In all other characters these specimens agree
with var. zippeliana, but more material may show them to represent a separate taxon.

2. The fruits of some specimens from New Guinea are + ellipsoid(-oblong), and are
intermediate in shape with those of var. paniculata; they are placed in var. zippeliana
because of their short fruiting pedicels.

3. Occasionally specimens are found with particularly brittle leaves when dry, a char-
acter not often met with in other Gymnacranthera species.

4. Some collections from New Britain have leaves drying dull with the nerves sunken
above, and pale, often purplish-whitish below, but intermediate forms exist with the re-
mainder of var. zippeliana.

4. Gymnacranthera forbesii (King) Warb.

Gymnacranthera forbesii (King) Warb., Mon. Myrist. (1897) 363, t. 20; Gamble, Mat. Fl. Malay
Penins. 5, 23 (1912) 224: Ridl., Fl. Malay Penins. 3 (1924) 61; J. Sinclair, Gard. Bull. Sing. 16
(1958) 441, f. 55, pl. XIII B; 17 (1958) 101, f. 1A, C; R.T.A. Schouten, Blumea 31 (1986) 472,

De Wilde — Myristicaceae (Gymnacranthera) 49

474; W.J. de Wilde, Tree Fl. Sabah & Sarawak 3 (2000) 350. — Myristica forbesii King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 306, t. 137. — Lectotype: Forbes 2976 (female fl.) (K), Sumatra,
Benkulu.

Tree 5—35 m. Twigs subterete to angular or ridged, (2—)2.4—4(—5.5) mm diameter,
hairs 0.1 mm or less, early glabrescent, bark chocolate, later on brown or grey, finely
cracked, with many lenticels. Leaves chartaceous or coriaceous, elliptic to oblong-lanceo-
late, widest at or below the middle, 14—33 by (5.5—)6—13 cm, base (short-)attenuate to
broadly rounded, apex (acute-)acuminate, blade coarsely undulate on drying, margin not
revolute; upper surface olivaceous to brown, sometimes glossy, lower surface grey
(-purple) to brownish, with scattered appressed minute hairs, glabrescent; midrib above
somewhat sunken (grooved) above, 1.5 mm wide, nerves 10-18 pairs, at (35—)40—50
(—55)° to the midrib, yellowish, distinct and (very) prominent below, venation some-
times distinct at the lower surface; petiole 8—20 by 1.5—3 mm; leaf buds 5—10 by 1.54
mm, with dense greyish hairs 0.1 mm. /nflorescences paniculate, with (grey-)rusty
+ woolly hairs 0.2—0.4 mm long; in male 4-12 cm long, up to 8 cm wide, many-flow-
ered, in female 1-4 cm long, 1-3 cm wide, generally fewer flowered; bracts triangular,
3 by 3.5 mm, pubescent, caducous. Flowers grey-rusty pubescent inside and outside,
hairs 0.1-0.2 mm long. Male flowers: pedicel 2—3 mm long; buds ellipsoid-oblong,
2.54 by 1.5—2 mm, cleft 1/3—nearly 1/2, lobes 3 (or 4), (long-)triangular, only slightly
spreading, tube 1.5—2.3 mm long; androecium + truncately ellipsoid-oblong, stiped to
0.2 mm long, 1.5—2.3 by | mm; anthers 6—10, subsessile, often somewhat twisted, free
apices 0.3-0.5 mm long, + erect. Female flowers: pedicel 2 mm long; buds ovoid or
pyriform, 2—2.5 by 1.7—2 mm, cleft to c. 3/4, lobes (2 or) 3, strongly spreading or recurved,
tube short-urceolate, 0.5—0.7 mm long; ovary subglobose, 1—1.3 mm diameter, densely
minutely pubescent; stigma sessile, minutely 2-lobed. Fruits 4-25 per infructescence,
ellipsoid-oblong, 1.8—2.4 by 1—1.4 cm, (early) glabrescent or short-pubescent, hairs 0. 1
mm; pericarp | mm thick; fruiting pedicel 4-10 mm long. — Photo 2.

Distribution — S Thailand; Malesia: Sumatra, Peninsular Malaysia, Singapore, Bor-
neo. Somewhat arbitrarily two varieties can be distinguished, of which one confined to
Borneo.

Note — The leaves of G. forbesii usually dry coarsely undulate, not flat as in other
species.

KEY TO THE VARIETIES

la. Leaves chartaceous to coriaceous; nerves on lower surface moderately prominent,
0.3—0.5 mm wide. Infructescences usually not branched at the base, few-fruited. —
Benmsular Malaysia:Sumatha,Bormeo: ...2.95. 62252022. 42s a. var. forbesii

b. Leaves (very) coriaceous; nerves on lower surface more prominent, 0.5—0.7 mm wide.
Infructescences conspicuously branched from the base, many-fruited. — Borneo
ES eos 2 ice Beet n © Oe Ss Secs Seles ee b. var. crassinervis

a. var. forbesii

Gymnacranthera forbesii (King) Warb. var. forbesii; J. Sinclair, Gard. Bull. Sing. 17 (1958) 101,
f. 1A, C, p.p.; R.T.A. Schouten, Blumea 31 (1986) 474; W. J. de Wilde, Tree Fl. Sabah & Sarawak
3 (2000) 350.

50 Flora Malesiana, Ser. I, Vol. 14 (2000)

Twigs (2—)2.5-3(-4) mm diameter. Leaves chartaceous or subcoriaceous; blade ellip-
tic to oblong, 14-28 by (5.5—)6-13 cm; nerves on lower surface moderately prominent,
0.3-0.5 mm wide. Infructescences not or little branched at base, with 4—10 fruits.

Field-notes — Crown irregular, dense; bole usually straight, not buttressed. Bark soft,
grey to brown, smooth, finely fissured, or thinly flaky; the inner bark pink to red-brown,
laminated, sometimes fibrous; slash wood white to pale yellow. Flowers brown-green in
buds, bright yellow at anthesis; pollen whitish. Fruits brown-green turning orange.

Distribution — S Thailand (Pattani); Malesia: Sumatra, Peninsular Malaysia, Singa-
pore, Borneo.

Habitat & Ecology — Primary and degraded forest; hillsides and riverbanks, alluvial
forest; on sandy and limestone-derived soils; up to 600 m altitude; fl. Feb.—Apr., Aug.—
Sept.; fr. May—July(—Aug.), Dec.—Jan.

b. var. crassinervis (Warb.) J. Sinclair

Gymnacranthera forbesii (King) Warb. var. crassinervis (Warb.) J. Sinclair, Gard. Bull. Sing. 17
(1958) 102, f. 1B; R.T.A. Schouten, Blumea 31 (1986) 475; W.J. de Wilde, Tree Fl. Sabah &
Sarawak 3 (2000) 350. — Gymnacranthera crassinervis Warb., Mon. Myrist. (1897) 362, t. 20.
— Myristica crassinervis (Warb.) Boerl., Handl. 3 (1900) 88. — Lectotype: Beccari 1119 (K),
Sarawak.

Twigs (2.5-)3—4(-5.5) mm diameter, lower down 3.5-5.5 mm. Leaves coriaceous;
blade elliptic-oblong to oblong-lanceolate, 16-33 by 6-12 cm; nerves on lower surface
strongly prominent, conspicuous, 0.5—0.7 mm wide. Infructescences often conspicu-
ously branched from the base, with 8-25 fruits.

Field-notes — Bole without buttresses. Bark grey to brown, smooth or sometimes
slightly flaky, slash wood white-orange-brown. Flowers bright yellow. Fruits brown green,
orange-red when ripe.

Distribution — Malesia: most of Borneo (according to Sinclair W Kalimantan, but
no specimens seen).

Habitat & Ecology — Primary and degraded dryland as well as wet forest, alluvial
forest; on sandy(-clay) and loamy soils; up to 1250 m altitude; fl. mainly Apr., Aug.—
Oct.; fr. Apr., Aug.—Dec.

Note — Gymnacranthera forbesii vat. crassinervis usually is easily recognized and
distinguished from var. forbesii and other species of Gymnacranthera by its stout twigs
and leaves and usually strong orange-yellowish lateral nerves, which are very distinctly
raised on the lower leaf surface. Gymnacranthera bancana also is a stout species, but
leaves and young twigs are always rusty tomentose, whereas the present variety is al-
most glabrous.

5. Gymnacranthera maliliensis R.T. A. Schouten

Gymnacranthera maliliensis R.T.A. Schouten, Blumea 31 (1986) 467, f. 5. — Type: van Balgooy
3960, eastern Central Sulawesi.

Tree 6-20 m. Twigs terete to slightly angular, 1.5—2.5(—4) mm diameter, lower down
3-4 mm, when young rusty pubescent by woolly hairs, not very appressed, hairs woolly,
to 0.5 mm long, glabrescent, greyish to brown with the bark smooth or finely longitu-

De Wilde — Myristicaceae (Gymnacranthera) 51

dinally cracked, densely set with lenti-
cels. Leaves chartaceous or coriaceous,
(oblong-)lanceolate, 10—28(—30) by 3—7
(—9) cm, base attenuate, apex acute
(-acuminate), margin not or but slightly
revolute; upper surface olivaceous to
brown and often glossy, lower surface
brown to purplish grey, with dense ap-
pressed hairs 0.2—0.3 mm long, late gla-
brescent; midrib above flat or slightly
sunken, | mm wide, nerves 8-17 pairs,
at 45—60° to the midrib, distinct but

Fig. 6. Schematic drawing, in longitudinal and
transverse section, of the androecium of Gym-
nacranthera maliliensis R.T.A. Schouten; note

the minute appendix of the connectivum; the not prominently raised below, venation
broken line indicates the depth of the incision forming a coarse network + indistinct
between the anthers. — Scale bar = 1 mm. at both surfaces; petiole 7-14 by 1.5-2

(—2.5) mm; the leaf buds 10 by 2 mm,

densely appressed-pubescent. /nflores-
cences (broadly) paniculate, with rusty hairs 0.5 mm long; in male 4.5—7 cm long, up to
6 cm wide, many-flowered, in female (from infructescences) little or much branched,
rather few- to many-flowered, 3-5 cm long; bracts broadly triangular, 2.5 mm, pubes-
cent, caducous. Flowers rusty pubescent inside and outside, hairs 0.1—0.2 mm long.
Male flowers: pedicel 2.5—-4 mm long; buds ellipsoid-oblong, 3.5—4 by 2 mm, cleft to
1/4—1/2, lobes 3 or 4, (long-)triangular, somewhat spreading, tube 2—2.7 mm long;
androecium + truncately ellipsoid, 1.8—2.3 by 1.7 mm, stipe 0.2 mm; anthers 8-10,
subsessile, the free apices 0.5 mm long, erect or curved slightly inward. Female flowers
not seen. Fruits up to 25 in immature infructescences, 3-6 when mature, ellipsoid, 2.3—
3 by 2-2.2 cm, hairs persistent, rusty, 0.2 mm; pericarp 3—5 mm thick; fruiting pedicel
stout, 4-8 mm long. — Fig. 6, 7.

Field-notes — Tree to 20 m, dbh to 25 cm, with red sap. Flowers dark yellow. Unripe
fruits brown.

Distribution — Malesia: E Central Sulawesi, east of Malili.

Habitat & Ecology — Primary and degraded forest on ultrabasic (nickel-containing )
soils; 200-500 m altitude; fl. Feb., July; fr. Oct.

Note — Endemic to Central Sulawesi; restricted to soils derived from ultrabasic rock.
Distinguished from the only other species occurring in Sulawesi, G. farquhariana var.
zippeliana, by the more conspicuous indumentum of woolly hairs on the young twigs
and the large fruits with thick pericarp.

6. Gymnacranthera ocellata R.T.A. Schouten

Gymnacranthera ocellata R.T. A. Schouten, Blumea 31 (1986) 469, 3g, h; W.J. de Wilde, Tree Fl.
Sabah & Sarawak 3 (2000) 351. — Type: SAN 72177 (Saikeh), Sabah.

Tree 10-25 m. Twigs somewhat angular or + compressed, 2—3.5(—6.5) mm diameter,
with appressed hairs 0.2 mm high, glabrescent, the bark brown or grey, smooth or finely
cracked, with dense conspicuous lenticels of mixed size, even when young; at the base

Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Gymnacranthera) 55

of each innovation a group of scars occurs (see note). Leaves chartaceous or thinly
coriaceous, ovate-elliptic to oblong-lanceolate, widest at or below the middle, 10—25 by
4—9.5 cm, base short-attenuate to broadly rounded, apex acute, margin not revolute;
upper surface olivaceous or brown, often glossy, lower surface grey-brown, with dis-
tinct, but not very dense appressed rusty hairs, late glabrescent; midrib above flat, nar-
row, | mm wide; nerves 11—18 pairs, at c. 45° to the midrib, distinct but not prominent
below, venation on both surfaces a coarse, rather indistinct network; petiole 9-18 by
2-2.5 mm: leaf buds + stout, 10 by 3 mm, composed of the unexpanded leaf often with
in addition several scale-like cataphylls, densely appressed-pubescent. Inflorescences
(broadly) paniculate, with rusty hairs 0.3—0.5 mm long; in male 3—8.5 cm long, up to
7 cm wide, many-flowered, in female to 2 by 1.5 cm, few-flowered; bracts broadly tri-
angular, 2.5 by 3 mm, pubescent, caducous. Flowers rusty pubescent inside and outside,
hairs 0.1(—0.2) mm long. Male flowers: pedicel 1.5—3.5 mm long; buds ellipsoid-
oblong, 3—4(—5) by 2—3(—3.5) mm, cleft 1/3—1/2, lobes 3 or 4, (long-)triangular, at an-
thesis somewhat recurved, tube 2—2.5 mm long; androecium truncately ellipsoid, 1.5—
2.3 by 0.7—1 mm, stipe to 0.2 mm; anthers 7-10, subsessile, the free apices 0.4—0.5 mm
long, + erect. Female flowers: pedicel 1.5—2.5 mm long; buds narrowly ovoid, 2.7—3 by
2 mm, cleft 1/2—3/4, lobes (2 or) 3, long-triangular, + recurved, tube 0.5—1.3 mm long;
ovary subglobose, 1.2-1.5 mm diameter, densely minutely pubescent, stigma sessile,
shallowly 2-lobed. Fruits 4-10 per infructescence, ovoid-ellipsoid, usually with trun-
cate base, 1.8—2.2 by 1.1—1.3 cm, hairs 0.1-0.2 mm long; pericarp 0.7—1.3 mm thick;
fruiting pedicel 3-7 mm long. — Fig. 5g, h.

Field-notes — Bole smooth, 10—17 m; no buttresses. Bark smooth, fissured, regularly
cracked, or flaky, grey or dark red-brown; inner bark 10-15 mm thick, brown; sapwood
4 cm, whitish streaked with pale red; heartwood light brown to blackish brown. Flowers
yellow; anthers (pollen) whitish yellow. Fruits green turning orange(-brown).

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, Kalimantan, incl. Nunukan).

Habitat & Ecology — Primary dryland forest, kerangas, forest on ridges and low
hills; on tuff, sand, and sandy loam; up to c. 1300 m altitude; fl. mainly June—Novy.; fr.
July—Dec.

Note — The specific epithet refers to the numerous small eye-like pale lenticels on
the twigs. Gymnacranthera ocellata may also be recognized by the conspicuous and
numerous scars of cataphylls at the base of each seasonal shoot, possibly related with a
marked seasonal growth.

Fig. 7. Gymnacranthera maliliensis R.T. A. Schouten. a. Habit of leafy twig with immature fruits;
b. twig with male inflorescence; c. male flower; d. ditto, opened, showing androecium; e. infruc-
tescence; f. fruit, opened, showing thick pericarp and deeply laciniated aril of seed [a: van Balgooy
3884; b—d: van Balgooy 3960; e, f: bb Cel./V 255]. — Scale bar for a, b, e = 2 cm; for c, d =
1.65 mm; for f = 1 cm.

54 Flora Malesiana, Ser. I, Vol. 14 (2000)

HORSFIELDIA

Horsfieldia Willd., Sp. Pl. 4 (1806) 872 [non Blume = Harmsiopanax Warb.(Araliaceae)], Pers.,
Symb.2 (1807) 635; Warb., Mon. Myrist. (1897) 130, 262: J. Sinclair, Gard. Bull. Sing. 16 (1958)
368: 27 (1974) 133-141; 28 (1975) 1-181; W.J. de Wilde, Gard. Bull. Sing. 37, 2 ((1984’, 1985)
115-179: 38. 1 (1985) 55-144; 38, 2 (‘1985", 1986) 185-225; 39, 1 (1986) 1-65; Blumea 32
(1987) 459-472; 41 (1996) 375-381; Tree Fl. Sabah & Sarawak 3 (2000) 352. — Pyrrhosa
Endl.. Gen. Pl. (1839) 830, nom. illeg. — Type species: Horsfieldia odorata Willd. [= Horsfieldia
iryaghedhi (Gaertn.) Warb.].

Myristica sect. Pyrrhosa Blume, Rumphia | (1837) 190-192, t. 62-64. — Subsequent authors treat-
ed the genus Horsfieldia as defined at present partly under Myristica sect. Pyrrhosa as well as
under several other sections of Myristica, e.g., sections Caloneura p.p., Eumyristica p.p.,
Horsfieldia, Irya (see Sinclair, 1958: 368), and under the here accepted and discussed sections.
— Lectotype species: Myristica glabra Blume [= Horsfieldia glabra (Blume) Warb.].

Shrubs or usually trees, dioecious. Twigs sometimes angular or with two raised lines,
lenticellate, not flaky. Leaves sometimes dispersed, brittle when dry, lower surface not
pale, papillose only in H. iryaghedhi, dots present or absent; reticulation lax (never forming
a close raised network as in Knema). Inflorescences (on older wood in H. sabulosa)
paniculate, usually branched several times; flowers numerous, all in about the same
stage of development, solitary, or clustered (in H. iryaghedhi the male with flowers
sessile in dense flower heads); basal cataphylls caducous; bracts small or large, caducous.
Flowers small, mostly short-pedicelled, at base articulated or not, bracteole absent. Male
flowers: perianth + globose or cup-shaped, leathery, inside glabrous, yellowish (never
red): buds (depressed-)globose, (transversely) ellipsoid, reniform, or clavate, laterally
compressed or not, cleft to variable depths, lobes 2 or 3 (or 4), not spread at anthesis;
androecium sessile or with short narrow androphore, glabrous, synandrium variously
shaped (cup-shaped, globose, ellipsoid, cylindrical, or trigonous), laterally compressed
or not: anthers 2—c. 25, largely or entirely connate into a central column hollowed to
different depths at apex; anthers straight, curved, or inflexed into the apical cavity to
variable depths. Female flowers: slightly larger than male, globose to ovoid-ellipsoid;
ovary glabrous or pubescent, style absent, stigma small, deeply 2-lobed (more-lobed in
H. iryaghedhi). Infructescences branched, few- to many-fruited. Fruits globose or ellip-
soid, 1-8 cm long, pericarp leathery or somewhat fleshy, with or without lenticel-like
tubercles, glabrous or pubescent, perianth sometimes persistent; aril entire or only shal-
lowly lobed; seeds rarely globose, not variegated; albumen ruminate, with fatty oil but
no starch; cotyledons connate at base. — Fig. 8-33.

Distribution — More than 100 species, ranging from Sri Lanka through NE India to
S China (Kwangsi, Hainan) and through Malesia and the Caroline Islands east to the
Solomon Islands and N Australia. The genus is absent in the Lesser Sunda Islands and
8 species occur exclusively outside the Malesian area. Except for a few widely distrib-
uted species, e.g., H. amygdalina (extra-Malesian), H. glabra, H. irya, H. laevigata, H.
tuberculata, most species have a limited distribution. — Map 3 (see p. 4).

Distinct centres of species development are New Guinea and Borneo, and to a lesser
extent Sumatra and Peninsular Malaysia. Three sections have been recognized here, and
they occupy largely exclusive areas. Section Horsfieldia, with only H. iryaghedhi, is
confined to Sri Lanka. Section /rya (with c. 40 species) is, except for the widespread

De Wilde — Myristicaceae (Horsfieldia) 55

H. irya, confined to East Malesia, the Solomons and N Australia. Section Pyrrhosa oc-
curs west of Wallace’s Line. In distribution, sections /rya and Pyrrhosa overlap for a
narrow area in the Philippines and Sulawesi. They are morphologically segregated main-
ly by a different number of perianth lobes. Some species with the aberrant number of
3 lobes occur in section /rya: H. angularis, H. olens, and H. sepikensis. In section Pyrrho-
sa the deviating number of 2 lobes is found in H. longiflora, H. thorelii, and H. amygdali-
na, partly (these three species are extra-Malesian), and H. crassifolia and H. sterilis. For
a further explanation, see De Wilde (*1984’, 1985).

Habitat & Ecology — Trees of primary rain forest, persisting in degraded forest or
sometimes in old secondary growths; sometimes in marshy forest (H. irya); stilt-roots
are present in some species. Some species reach or occur in montane areas, and the wide
altitudinal range contrasts with those of the other Malesian genera of the Myristicaceae.

According to Sinclair (1958) the bark of species of Peninsular Malaysia is usually
reddish brown, smooth or more often striate, or rough with circular or irregular dents,
sometimes flaking but mostly not. The flowers are usually waxy yellow, and often sweet
scented; those of H. iryaghedhi have a particularly strong smell.

Taxonomy — There is a large morphological diversity in the flowers of the genus
Horsfieldia. Schematic drawings of the androecia of most species have been depicted
here on pages 58—61 as Plates 1—3 [after W. J. de Wilde, Gard. Bull. Sing. 37, 2 (1984,
1985) 129-137].

Three sections can be recognized and are supposedly of unequal taxonomic weight
but with significant different ranges of distribution (see above): 1) sect. Horsfieldia,
containing one single species, the type species of the genus, rather deviating from all
other species, 2) sect. Jrya, containing most species with predominantly a 2-lobed peri-
anth, and 3) sect. Pyrrhosa, most of its species with predominantly a 3- or 4-lobed
perianth. The descriptions of the three sections have been given here separately and are
not included in the treatment of the species, which are all listed alphabetically. [For
an extensive discussion of the subdivision of Horsfieldia into the three sections, see
W.J. de Wilde (°1984’, 1985) 125-136.]

Section Horsfieldia

Myristica Gronov. sect. Horsfieldia A. DC., Prodr. 14, 1 (1856) 200; Migq., FI. Ind. Bat. 1, 2 (1858)
63. — Myristica Gronov. sect. /rya auct. non Hook. f. & Thomson: Benth. & Hook. f., Gen. Pl. 3
(1880) 137, for Horsfieldia only. — Myristica Gronov. sect. Eumyristica Hook. f. & Thomson
subsect. Horsfieldia (A. DC.) King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 282. — Type species:
Horsfieldia odorata Willd.

Myristica Gronov. sect. Pyrrhosa Blume, Rumphia | (1835) 190, p.p., for Myristica horsfieldii
only, not the lectotype species. — Myristica Gronov. sect. Eumyristica Hook. f. & Thomson, FI.
Ind. (1855) 162, p. p., for Myristica horsfieldii only.

Horsfieldia Willd. sect. Orthanthera Warb., Mon. Myrist. (1897) 268, p.p., for the lectotype species
only. — Horsfieldia Willd. sect. Trivalves J. Sinclair subsect. Orthanthera (Warb.) J. Sinclair,
Gard. Bull. Sing. 16 (1958) 371, p.p., nom inval., provisional name only. — Lectotype species:
Horsfieldia iryaghedhi (Gaertn.) Warb.

Phyllotaxis of plagiotropic shoots distichous. Lower leaf surface with alveolar tissue,
epidermis papillose, stomatal complex sunken; without larger dark-coloured dots (cork

56 Flora Malesiana, Ser. I, Vol. 14 (2000)

warts). Flowers in male inflorescences sessile, at base not articulated, arranged in many-
flowered subglobose dense heads; perianth in buds elongate-obconical, + angled, cleft
1/2-3/4, 3- or 4-(in female: 2- or 3-)lobed. Androecium elongate, subcylindrical, andro-
phore distinct; column narrow, narrowly hollowed for over halfway; anthers 3—5, erect.
Stigma sessile, many-lobulate. — One species, Sri Lanka. — Map 3 (p. 4).

Section Horsfieldia is monotypic, the species H. iryaghedhi deviating from all other
Horsfieldias by some anatomical characters of the leaf, male flowers sessile and ar-
ranged in dense heads with a thick receptacle, angular buds, anthers largely connate, but
not back to back so that a narrowly hollowed central column is formed; the stigma in the
female flowers is many-lobed, not 2-lobed as in the other Horsfieldia species.

Section Irya

Horsfieldia Willd. sect. Irya (Hook. f. & Thomson) Warb., Mon. Myrist. (1897) 123, 267, p.p.;
W.J. de Wilde, Gard. Bull. Sing. 37, 2 ((1984’, 1985) 127. — Myristica Gronov. sect. /rya
Hook. f. & Thomson, Fl. Ind. (1855) 159; A. DC., Prodr. 14, 1 (1856) 202; Miq., FI. Ind. Bat. 1,
2 (1858) 64; Benth. & Hook. f., Gen. PI. 3 (1880) 137, p. p., excl. sect. Horsfieldia, King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 284, p.p., for the smaller part only. — Horsfieldia Willd. sect.
Irya (Hook. f. & Thomson) Warb. subsect. Euirya Warb., Mon. Myrist. (1897) 123, 267, p.p., for
the type species only. — Type species: Myristica irya Gaertn. [= Horsfieldia irya (Gaertn.) Warb.].

Myristica Gronov. sect. Pyrrhosa Blume, Rumphia | (1837) 190, p.p., for Myristica javanica and a
few other species only, excl. lectotype species Myristica glabra (= sect. Pyrrhosa) and Myristica
horsfieldii (= Horsfieldia iryaghedhi, sect. Horsfieldia), A.DC., Prodr. 14, 1 (1856) 202, p.p.;
Mic., Fl. Ind. Bat. 1, 2 (1858) 64, p.p., excl. Myristica glabra (= sect. Pyrrhosa).

Horsfieldia Willd. sect. Pyrrhosa (Blume) Warb. subsect. Bivalves Warb., Mon. Myrist. (1897) 262
(incl. series Smithii and series Globularia). — Horsfieldia Willd. sect. Bivalves J. Sinclair, Gard.
Bull. Sing. 16 (1958) 370, 371, comb. inval., provisional name only. — Type species not indi-
cated, several species listed.

Phyllotaxis in plagiotropic shoots distichous. Lower leaf surface without alveolar
tissue, epidermis not papillose, stomatal complex not sunken; rarely (H. inflexa) with
larger dark-coloured dots. Flowers pedicelled, not articulated at base, solitary or in loose
clusters; buds rarely globose or obovoid, usually laterally compressed, in outline cir-
cular, elliptic, or pear-shaped, not angular (in young stages of H. sylvestris faintly so),
cleft to variable depths, perianth mostly predominantly 2-lobed, rarely 3-lobed.
Androecium various, often + zygomorphic, laterally compressed, sometimes anthers at
apex bi-laterally incurved, or obconical and + actinomorphic; androphore distinct or
not, central column broad or narrow, little to much hollowed: anthers few to many, (sub)-
erect, or at apex incurved into apical hollow of the column. Stigma minutely 2-lobed. —
Mainly E Malesia (including the Philippines), and one species, H. irya, distributed over
almost the whole area of the genus. — Map 3 (p. 4).

Almost all of the 40 species of this section have predominantly 2-lobed perianths,
with a more or less zygomorphic androecium, because it is laterally compressed or with
the anthers at apex incurved from two sides into an apical hollow of the column.

Aberrant are H. olens and H. sepikensis with 3- or 4-lobed perianths, but with the
androecium tending to be zygomorphic. Horsfieldia an gularis has 2—4-lobed perianths.

Also aberrant are the species of the group of H. clavata, with a 2-lobed perianth but a
club-shaped non-zygomorphic androecium.

De Wilde — Myristicaceae (Horsfieldia) 57

A few mutually related species from continental SE Asia, placed in section Pyrrhosa,
viz. H. longiflora, H. thorelii and H. amygdalina, have (partly) a 2-lobed perianth, and a
zygomorphic androecium, especially H. longiflora. They blur the distinction between
sections Irya and Pyrrhosa. Section Irya occurs mainly in East Malesia, with only
H. irya extending far beyond the main range of distribution of the section. Within sec-
tion /rya, eight groups of species can be distinguished, a survey of which is given by
De Wilde (“1984’, 1985: 128).

Section Pyrrhosa

Horsfieldia Willd. sect. Pyrrhosa (Blume) Warb., Mon. Myrist. (1897) 262, p.p.; W.J. de Wilde,
Gard. Bull. Sing. 37, 2 (1984’, 1985) 130. — Myristica Gronov. sect. Pyrrhosa Blume, Rumphia
1 (1837) 190, t. 62—64, p. p., for the smallest part incl. the lectotype t. 64, f. LA, B; Hook. f. &
Thomson, Fl. Ind. (1855) 160; A. DC., Prodr. 14, 1 (1856) 202, p.p.; Miq., Fl. Ind. Bat. 1, 2
(1859) 64, p.p.; Benth. & Hook. f., Gen. Pl. 3 (1880) 136; King, Ann. Roy. Bot. Gard. Calc. 3
(1891) 282. — Horsfieldia Willd. sect. Pyrrhosa (Blume) Warb. subsect. Eupyrrhosa Warb.,
Mon. Myrist. (1897) 265 (excl. Horsfieldia macrocoma = Endocomia). — Lectotype species:
Myristica glabra Blume [= Horsfieldia glabra (Blume) Warb.].

Myristica Gronov. sect. Eumyristica Hook. f. & Thomson, FI. Ind. (1855) 162, p.p., for Myristica
superba = Horsfieldia superba (Hook.f. & Thomson) Warb. only. — Myristica Gronov. sect.
Caloneura A.DC., Prodr. 14, 1 (1856) 192, p.p., for Myristica superba Hook.f. & Thomson
only [= Horsfieldia superba (Hook. f. & Thomson) Warb.].

Myristica Gronov. sect. Irya auct. non Hook. f. & Thomson: King, Ann. Roy. Bot. Gard. Calc. 3
(1891) 284, p.p.

Horsfieldia Willd. sect. Irya (Hook. f. & Thomson) Warb. subsect. Euirya Warb., Mon. Myrist. (1897)
267, p.p., excl. Horsfieldia irya (type species of sect. /rya).

Horsfieldia Willd. sect. Irya (Hook. f. & Thomson) Warb. subsect. Trivalves Warb., Mon. Myrist.
(1897) 267. — Horsfieldia Willd. sect. Trivalves J. Sinclair subsect. Trivalves, J. Sinclair, Gard.
Bull. Sing. 16 (1958) 370, 371, comb.inval., provisional name only. — Type species not indi-
cated.

Horsfieldia Willd. sect. Orthanthera Warb., Mon. Myrist. (1897) 268, p.p., for Horsfieldia ralunen-
sis and Horsfieldia sylvestris only, excl. the lectotype species Horsfieldia iryaghedhi.

Phyllotaxis of plagiotropic shoots dispersed or distichous. Lower leaf surface without
alveolar tissue, epidermis not papillose, stomatal complex not sunken; with or without
larger dark-coloured dots (or cork warts). Flowers with a pedicel (short in H. wallichii),
at base with or without articulation, solitary or in loose clusters; buds (depressed) glo-
bose, obovoid, or ellipsoid, not laterally compressed, not angular, cleft c. 2/3 or less;
perianths predominantly 3- (or 4-)lobed (rarely 2-lobed, H. longiflora, H. sterilis). Androe-
cium (sub)circular or more or less triquetrous in cross section, never laterally compressed,
depressed-globose, ellipsoid, or obovoid, usually with a broad central column with an
apical hollow of variable shape and depth; androphore usually narrow at base; anthers
few to many, + straight or curved, almost entirely connate or free for about the upper half
(H. polyspherula-group). Stigma minutely 2-lobed. — Continental SE Asia, West Malesia
(including the Philippines). — Map 3 (p. 4).

Section Pyrrhosa contains c. 60 species, mainly with exclusively or predominantly
a 3- (or 4-)lobed perianth; species with a 2-lobed perianth are H. longiflora (Vietnam),
H. crassifolia (Sumatra, Peninsular Malaysia, Borneo) and H. sterilis (Sabah). For a
brief discussion see De Wilde (“1984’, 1985: 132).

58 Flora Malesiana, Ser. I, Vol. 14 (2000)

Legends to Plates 1-3:

Semi-schematic drawings of the androecia of most species of Horsfieldia, except
H. ampla (1), H. discolor [21], H. disticha [22], H. perangusta [59], and H. ses-
silifolia [75], of which no male flowers are available.

Between square brackets the number has been given of the alphabetically arranged
species of the present revision.

Lateral view (left), longitudinal section (right), apical view (top), white: anthers;
black: sterile tissue (i.e., androphore and central column).

Plate 1 — /: H. iryaghedhi [36] — 2: H. irya [35] — 3: H. spicata [79] — 4: H. inflexa [33]
— 5: H. moluccana var. moluccana [44a] — 6: H. parviflora [56] — 7: H. ob-
scurinervia [49] — 8: H. ardisiifolia [6] —9: H. talaudensis [88] — 10: H. samarensis
[72] — 11: H. smithii (77) — 12: H. olens [51] — 13: H. sepikensis [74] — 14: H. syl-
vestris [87] — 15: H. coryandra [16] — 16: H. urceolata [94] — 17: H. crux-meliten-
sis [19] — 18: H. clavata [13] — 19: H. squamulosa [81] — 20: H. ampliformis [2] —
21: H. angularis [5] — 22: H. iriana [34] — 23: H. aruana {7| — 24: H. subtilis var.
subtilis [84a] — 25: H. schlechteri [73] — 26: H. basifissa [9] — 27: H. sinclairii
[76] —28: H. psilantha [62] — 29: H. laevigata var. laevigata [37a] — 30: H. pilifera
[60] — 31: H. lancifolia [38].

Plate 2 — 32: H. decalvata [20] - 33: H. tuberculata [93] — 34: H. corrugata [15] — 35:
H. pachycarpa [53] — 36: H. pulverulenta |64] — 37: H. leptantha [40] — 38: H.
hellwigii var. hellwigii [31a] — 39: H. ralunensis [67] — 40: H. sabulosa [71] —41:
H. atjehensis [8] — 42: H. sucosa subsp. sucosa [85a] — 43: H. pallidicaula vat.
pallidicaula [55a] — 44: H. sparsa [78] —45: H. triandra [91] — 46: H tristis [92]
— 47: H. fulva [27] — 48: H. superba [86] — 49: H. grandis [30] — 50: H. wallichii
[96] — 51: H. pulcherrima [63] — 52: H. flocculosa 25] — 53: H. motleyi [46] —
54: H. tomentosa [90] — 55: H. cf. gracilis [29] — 56: H. paucinervis [57] — 57: H.
splendida [80] —58: H. rufo-lanata [70] — 59: H. reticulata [68] — 60: H. crassifolia
[18] —6/: H. carnosa [12].

Plate 3 — 62: H. sterilis [82] — 63: H. hirtiflora [32] — 64: H. brachiata [11] — 65: H.
pachyrachis [54] — 66: H. ridleyana [69] — 67: H. obtusa [50] — 68: H. tenuifolia
[89] — 69: H. macilenta [41] — 70: H. laticostata [39] — 71: H. nervosa [47] — 72:
H. polyspherula var. polyspherula [61a] — 73: H. oligocarpa [52] — 74: H. endertit
[24] — 75: H. valida [95] — 76: H. borneensis [10] — 77: H. fragillima [26] — 78: jak
androphora [4] — 79: H. amplomontana [3] —80: H. montana [45] — 81: H. elongata
[23] — 82: H. punctata [65] — 83: H. costulata [17] — 84: H. subalpina subsp. sub-
alpina [83a] — 85: H. obscura [48] — 86: H. xanthina subsp. xanthina [97a] — 87:
H. majuscula [43] — 88: H. coriacea [14] — 89: H. penangiana [58] — 90: H., punc-
tatifolia [66] — 91: H. macrothyrsa [42] — 92a: H. glabra var. glabra [28a] — 92b:
H. glabra var. javanica [28b].

Magnification for 3-11, 15-19, 21-27, 29-40, 45—48)50) 52, (ApPap Ole:
Magnification for /, 2, 12-14, 20, 28, 41-44, 49, 51, 53-73, 76-80, 82—92a, b
= x 10.

» 0
Oe. YU O88
ey | a] a)

ae

O

7 le TD 0. VO.

FO ji -_ os ag

os O) oP ce
308 rho.

= a" U) 4
we 2

=> = oS | ‘ :
2 2) Og OF O8

o
0. @.

fs

(+)
0.0

Ge, WO,

Plate 2 — For legends, see page 58

De Wilde — Myristicaceae (Horsfieldia)

&) a am. den a Mer,
@. YQ, @O. 08. GO.
é«) \ @)
00.24.00.0000, 09,
® ® C*)
Od. ¢. oe, @.

(+) @)
EB S0 aaa:

(+)
epee
16.8. 6.08.08.
er in enh ig
Sey Oboe

(\@ 1@-6- 6.
Plate 3 — For legends, see page 58.

62 Flora Malesiana, Ser. I, Vol. 14 (2000)

Notes to the Keys:

Besides a general key to the species (1), based on male flowering specimens, five sep-
arate regional keys (2—6) are given, based on female flowering and fruiting specimens
and with emphasis on vegetative characters and partly on distribution.

In species with a laterally compressed androecium (generally in flowers with a
2-lobed perianth) the shape and size of the androecium, as given in the keys and descrip-
tions, always concern the outline as seen laterally.

(1) GENERAL KEY TO THE SPECIES
(based on male flowering specimens)

la. Leaves papillose beneath. Male flowers sessile, packed into dense subglobose capi-
tula: buds + obconical, mutually appressed, angular. Perianth 3-lobed, cleft 1/5—2/3.
Androecium stalked, anthers* 3-5. — Sri Lanka, elsewhere cultivated........-..
er ts ss Se i oe 5 ss se eR ore arate 36. H. iryaghedhi

b. Leaves not papillose beneath. Male flowers subsessile or usually pedicelled, mutu-
ally free or at least not densely clustered; buds variable, not or only somewhat angu-

lar. Perianth densely clustered before anthesis in H. sylvestris from E Malesia . 2
2a. Leaves in fertile twigs distichous, membranous, usually with whitish marks of ir-
regular shape and size. Perianths globose, 2-lobed, 1—1.5(—2) mm diam. Androecium
not or hardly laterally compressed; anthers 6—10, for the larger part connate, form-
ing a shallow or deep saucer-shaped column, free apices 0.3 mm, + incurved; andro-
phore distinct, tapering. — Sri Lanka to Solomon Is. (throughout Malesia), gener-
ally Goastal-MVene «2c. sno: soa i 35. H. irya

b. Leaves in fertile twigs distichous (alternate), or dispersed (spirally), or mixed in the
same specimen. Leaf of variable consistency, usually without whitish marks. Peri-
anths 2—4-lobed, variable in shape and size. Androecium various; anthers few to
many, connate or mutually for some distance free, column variable; if column deeply
cup- or saucer-shaped, then anthers at apex free for at least halfway, or deeply inflexed
into the cup; androphore Various... 2..- se. ees ole Seg Ses SI aoe 3
3a. Perianth 2-lobed, or sometimes a few flowers in one inflorescence 3- or 4-lobed. —
Species mainly from E of Wallace’s Line, or the following from W Malesia: H. cras-
sifolia (Sumatra, Peninsular Malaysia, Singapore, Borneo), H. penangiana, p.Pp.
(Sumatra), H. sterilis (Borneo), H. sucosa subsp. bifissa (Borneo) .......---- 4

b. Perianth 3- or 4-lobed, or perianth with both 3 and 4 lobes present, sometimes the
odd flower in an inflorescence with 2-lobed perianth. — Species from W Malesia,

and the following from New Guinea: H. angularis, H. olens, H. sepikensis ... 43
4a. Androecium not laterally compressed (in cross section + circular), not longer than
wide. — W Malesia ..)...2.2 sss cule Solder = sande = oy ees oe 5

b. Androecium laterally compressed or not; if not or only slightly so then the androecium
(including androphore) longer than wide; androecium not or little laterally compres-

sed in odd 3- or 4-lobed flowers. — E Malesia ........---- +++ essere rere: 8

Sa. Leaves coriaceous, lower surface with inconspicuous, subpersistent, dense indumen-
tum beneath (in Borneo the hairs often deciduous) [dark dots and dashes present. ] —
Swamp forest on peat or sand .....--.-- +. eee ee eee eee 18. H. crassifolia

*) In the species descriptions the number of thecae, twice the number of anthers, is given.

De Wilde — Myristicaceae (Horsfieldia) 63

b.

6a.

b.

Ta.

b.

8a.

9a.

14a.

Leaves + membranous, glabrous or early glabrescent beneath. — Not from peat
BE Mp OKeStiad aactwersnreee bays «. aeaercmieanpien «wh hordes Oe apes 6
Twigs 2 mm diam.; bark not pale on drying. Leaves 7-12 cm long, with dots be-
neath. [Lateral nerves flat and inconspicuous above.] ...... 58. H. penangiana
Twigs 2—5(—10) mm diam.; bark pale on drying, often contrasting with the blackish
dried petioles. Leaves 14 cm long or more, without dots beneath ........... 7
Androecium depressed-globose, largely consisting of anthers; apical hollow flat
SATE ENA) MINA SOMES TES. AELO, GUNS. PSR te IO 85b. H. sucosa subsp. bifissa
Androecium broadly obovoid, consisting of a large sterile basal part at apex with
omsmiall anthers (26s. 6 thecae) 22 PUP MIR ee 82. H. sterilis
Inflorescences spike-like, unbranched or short-branched, lateral branches to 5
(-10) mm long. Inflorescences, flowers and petioles blackish on drying, usually
contrasting with the paler grey-brown twigs. Anthers inflexed. — Moluccas .... .
ee De SHE LIN VME AOS Pal SR oe sect Seance ecerts 79. H. spicata

_ Inflorescences branched, the side branches at least 5 mm long. Inflorescences, flow-

ers, and petioles brown, not contrasting with the colour of the twigs ........ 9
Androecium cup- or saucer-shaped, moderately laterally compressed; anthers at
one or both sides of the androecium distally distinctly incurved into the central
hollow. — E Malesia: Philippines, Sulawesi, Moluccas, NW New Guinea ... 10

_ Androecium laterally flattened or not, the column either 1) solid, 2) broadly but

shallowly hollowed only up to c. 1/3, or 3) narrowly slit-like channelled to vari-
able depths; anthers straight or only slightly incurved, never inflexed into the cen-

tral hollow. — E Malesia: Moluccas, whole of New Guinea .............. Wh
_ Leaves with dots beneath (lens!). Twigs angular or ridged ...... 33. H. inflexa
. Leaves without dots. Twigs terete, angular, or winged .............--.--- 1]

. Perianth together with the pedicel + pear-shaped, i.e., pedicel tapering. Petiole com-

paratively long, 1—2.6 cm. Twigs terete, not angular. [Leaf blade (6—)8—22(—25)
PC oie fs eee ee oes ness tye oe eee 44. H. moluccana

_ Perianth (in lateral view) short-cuneate, rounded, or subtruncate at base; pedicel

+ abruptly passing into the perianth, not tapered. Petioles comparatively shorter.
lmestctcte OF AMOWIAL occ. : gy 6 eke ce ea 12

. Perianth cleft about halfway. Anthers connate; androphore short or absent .. 13
_ Perianth cleft to + 2/3 or more. Anthers mutually free at least in the incurved or

‘i L225 bo aa ae eee Rm resent Sent OOTY ersten torts op ee 14

_ Anthers 18—25, inflexed at both sides into thin-walled laterally compressed an-

droecium cup. Perianth 2.5— 4 mm wide. Leaves membranous, matt on drying. —
RRC CAS aos yee tegen Mike Ore AED eke cb Sega pe ee 56. H. parviflora

. Anthers 11-12, inflexed at only one side of the androecium cup; cup thick- and

firm-walled. Perianth 2—2.2 mm wide. Leaves chartaceous, + glossy above. — Philip-
(ESCA EZCT) ee ee nena artemis See 49. H. obscurinervia
Anthers free only in the inflexed distal parts, the basal parts connate into a cup-
shaped column. Androphore minute, only c. 1/10 of the androecium length .. 15

. Anthers free for at least 2/3. Androphore longer, c. 1/3 of the androecium. [Twigs

generally angular or ridged. Perianth 2-3 mm wide; pedicel glabrous. Anthers all
inflexed into the centre of the androecium.] — Moluccas ....... 77. H. smithii

Flora Malesiana, Ser. I, Vol. 14 (2000)

Lia:

2la.

23a.

24a.

. Twigs angular or winged. Perianth 4 mm wide, glabrous. — Philippines ........

4 Socata peeps bee Roi @ is Sklar inant (i waa Se 6. H. ardisiifolia

. Twigs terete. Perianth 2.5—3 mm wide .............. 5s see ee eect eee 16
a. Pedicel pubescent, shorter than the perianth. Inflorescences densely finely pubes-

cent. Anthers inflexed at both sides of the laterally compressed androecium. —
Moluccas, (Talaudd.) nebiaieder. cso ce. eee leet 88. H. talaudensis

. Pedicel glabrous, longer than the perianth. Inflorescences with sparse hairs less

than 0.1 mm long. Anthers inflexed at one side only of the androecium. — Philip-
pines,(Samar)» ic lovieSemelin te ase teertt bho etre lien: 72. H. samarensis
Flower buds + angular, arranged into dense semi-globose clusters. Androecium much
longer than wide, not or slightly laterally compressed. Leaves lanceolate(-linear),

usually parallelsided BAA .ks sxe whet eee a ee 87. H. sylvestris
. Buds not angular, not densely clustered ..........- 062 - see eee eee eee 18
Twigs angular, ridged, or winged, at the apex as well as lower down ....... 19
. Twigs not winged, i.e., terete or only somewhat angular or lined at apex .... 74

_ Perianth 2-, 3-, or 4-lobed, subspherical, hardly or not laterally compressed, + glossy,

not collapsing on drying. — Papua Barat (Bird’s Head) ....... 5. H. angularis

. Male perianth predominantly 2-lobed, little or much laterally compressed, matt on

drying, slightly or strongly collapsing on drying ...........--++++++s55- 20

_ Leaves thinly coriaceous. Pedicel about as long as or longer than the perianth.

Buds cleft almost to the base. Hairs of inflorescences and pedicel 0.2—0.3 mm.
Anthers 10—14; the column hollow for c. 1/4 ...........-.....- 34. H. iriana

_ Leaves membranous. Pedicel shorter than the perianth. Buds cleft to 2/3—3/4. In-

florescences and pedicel almost glabrous, hairs 0.1 mm or less. Anthers (12—)14—
($= columm SOliG Or AIMOSU SO) eo caso oho ee oa 7. H. aruana
Inflorescences 25-35 cm long. Buds + pear-shaped. Androecium longer than broad;
androphore 0.5 mm long or more, about half as long as the anthers or longer 22

_ Inflorescences 20 cm long or less. Buds of variable shapes. Androecium longer or
shorter than broad; androphore short or long .......-.-.- +++ esse eee eeee 23
. Buds glabrous (?), 4 by 2 mm. Anthers 10, androphore nearly as long as the an-
thers. Inflorescences to 25 cm long, glabrescent .............-. 1. H. ampla
. Buds pubescent, 3 by 3 mm. Anthers 7, androphore about half as long as the an-
thers. Inflorescences 25-35 cm long, pubescent ........... 2. H. ampliformis
Inflorescences delicate, 2—5(—8) cm long, | or 2 (or 3) times branched. — New
Guinea, including Amn Is..........0/. ssc os gate os ++ i aslonie ete ne 24
_ Inflorescences stouter, 5-20 cm long, not or 1—4 times branched (inflorescences of

H. sinclairii and H. basifissa from New Guinea sometimes small). — Whole of
E Malesia, including Sulawesi; not in the Philippines .......-.....---++: 30
Buds pubescent or late glabrescent, distinctly or only somewhat longer than broad.
Anther-bearing part of the androecium much shorter than the elongate club-shaped
ANTICO PMOLE far. sieved a WEG eee 8 ala leek oc ca 2D

. Buds glabrous or early glabrescent, about as long as or shorter than broad. Andro-

phore much shorter than the anthers ............--- ee eeee eee eeees 29

. Buds together with tapering pedicel long pear-shaped. Androphore glabrous . 26

b. Buds globose or ellipsoid, distinctly marked-off from the slender pedicel. Andro-

phore glabrous or pubescent... . 2.22. eee eee er ene 27

De Wilde — Myristicaceae (Horsfieldia) 65

30a.

52d.

36a.

. Pedicel and bud together 10-12 mm long ............. 19. H. crux-melitensis
meeoicel. and budtogetherS mm long sn... cee ee eee: 13. H. clavata
. Buds subglobose or ellipsoid in outline, lobes 0.5—0.8 mm thick. Twig apex, leaf
Dbucsanduntlorescences: withhairs'0/1—0.2 mney neha Albee... 28
. Buds subglobose in outline, lobes 1—1.5 mm thick. Hairs 0.3(—0.4) mm. [Androphore
eiaroust-Pedicel)3immilong)) . Jon wo0\ele ae yee eee 94. H. urceolata
. Androphore (at least in the lower half) densely pubescent. Pedicel 3—3.5 mm long.
moossaienic. l/(Sronlyids esa nee Mee, oes $1. H. squamulosa
. Androphore glabrous or with a few scattered hairs. Pedicel (3—)4—6.5 mm long.
PosmemciinliG—)lWarnyeo Rishon. .\. als. Oey. Behe 16. H. coryandra
. Buds + laterally compressed, + obtriangular in lateral view, 1.8—3 mm wide, usual-
ly + collapsing on drying. Androphore 0.2—0.5 mm long, shorter than the an-
Line omer vey eg ited #0 le orhomrarod, Jh.\hn De ee Ee eed: 84. H. subtilis
. Buds subglobose, 1-2 mm wide, not or but slightly compressed, wrinkled on dry-

ing but not collapsing. Androphore 0.4—0.5 mm long, about half the length of the
PRMMCE We Nake ocy BE gee sande ay Qe eee 73. H. schlechteri
Bud and pedicel together + pear-shaped; apex broadly rounded in lateral view, the
lower (1/4—)1/3—1/2 tapered and gradually passing into the + tapered pedicel (these
characters not always clear in certain specimens of H. tuberculata). Buds glabrous,
fueescenor Slabreseent| ty) sart.. ¥aGn oF) Ve ee eR eee. Si

. Buds in lateral view circular, ovate, obovate, elliptic, transversely elliptic, or reni-

form; at base short-attenuate, rounded, or truncate, not tapered; pedicel + slender.
Buds hairy (at least at base) or in H. basifissa, H. psilantha, and H. sinclairii gla-
Prensiomelabresceity .au 2. et eA MO ee, Re ey 35

. Buds glabrous. — A variable species ................... 93. H. tuberculata
. Buds minutely pubescent, or in H. corrugata (at 1200—1900 m altitude) early gla-

bane CHE e SD (100) bree toy ethers Eeelsy schegh tee". S12, Ree 32
Leaves + lanceolate, 5—16 cm long. Buds cleft only c. 1/6. — Sulawesi.........
Se. th Ar antld htelin 2oudlal wieks alee 38. H. lancifolia

. Leaves elliptic to lanceolate, 12-30 cm long. Buds cleft about halfway ..... 33
. Pedicel 1.5—2 mm long; buds 2.3 mm long, thinly pubescent. Anthers 6. — Moluccas;

MITE TBE IMIG ah we cork SG ici lias & ges enki es beens Oe 20. H. decalvata

. Pedicel generally longer; buds 2.5—3.5 mm long. [Vegetatively and sometimes in

fruit much resembling H. laevigata.] — New Guinea; at 450—2000m ...... 34

. Pedicel stoutish, 2-4 mm long. Buds + membranous, glabrescent, with or without

a few scattered blackish warts. Anthers 8-12 .............. 15. H. corrugata

. Pedicel stout or slender, 2-5 mm long. Buds + fleshy, pubescent, blackish warts

AUSents Anite ES: Sahi4s i) eet A eR ESN ee ee ae 53. H. pachycarpa

. Inflorescences pubescent to nearly glabrous. Leaf bud and twig at apex with rusty

or greyish hairs, 0.1—0.4(—0.5) mm long. Leaves + glabrescent, or with scattered
stellate hairs, beneath: when younger ¢ wa eee ee ee 36

. Inflorescences generally thick-woolly tomentose. Leaf bud and twig at apex with

conspicuous, coarse, rusty hairs, (0.3—)0.5—1.5 mm. Leaves with (sub)persistent
indumentum, at least on and near the midrib beneath .................... 40
Buds + glabrous, subglobose, 2-3 mm diam., cleft to the base, not collapsing on
ign ee eres: 2 REIMER We os. cule od ws yee Lee ems ae 9. H. basifissa

Flora Malesiana, Ser. I, Vol. 14 (2000)

40a.

_ Buds glabrous or hairy, size and shape variable, cleft c. 1/2 to near the base; col-

lapsingtan drying OF NOs case. 4 6 4 «'</QANARER © Ta Fhe a 3M

a. Buds (almost) wholly with + persistent indumentum, hairs may be very minute and

scattered. [Leaves olivaceous-brown, without a reddish tinge. Hairs of leaf bud
0.1—0.2 mm, usually greyish.] — Moluccas, New Guinea, incl. New Britain . 38

_ Buds (and pedicel) glabrous or glabrescent, at least the upper 4/5. — E Papua New

Guined ose Re POSED. CARA. St Ss OS Sloe OS 39

_ Buds 1.2—1.9 mm diam. — A variable species ..........----- 60. H. pilifera
_ Buds 2—3.3 mm diam. — A variable species .......-..--+--- 37. H. laevigata
_ Buds (2—)2.5—3.5(—4) mm diam., cleft 1/2-2/3. Leaves 20-40 cm long, olivace-

ous or brown, not reddish tinged. — Papua New Guinea (Bagabag I., Long I., New
Brito: Newslceland) tres. £onee A) beers... Ate cee. Sie. Se 62. H. psilantha

_ Buds (1-)1.5—2 mm diam., cleft c. 1/2. Leaves 6—20 cm long, generally with a

reddish tinge on drying, especially the midrib and nerves. [Hairs of leaf bud to 0.1
mm long, greyish brown. Buds larger and pedicel pubescent in deviating speci-
mens.] — E Papua New Guinea ..........--.0+-sseeteee 76. H. sinclairii
Buds largely pubescent, towards the base thick-walled and coriaceous, the remain-
der collapsing on drying; both male and female buds opening at apex by small
pore-like slit less than | mm long. Androecium subellipsoid, mainly consisting of
the column with 2 minute anthers at the apex, just below the pore. Leaves coriaceous,
+ bullate: with harsh hairs leaving rough thickened bases .. 64. H. pulverulenta

. Buds glabrous or pubescent, membranous or chartaceous, + not collapsing on dry-

ing, cleft at least c. 1/3. Androecium mainly consisting of 10-16 sessile anthers.
Hairs not harsh, not leaving rough thickened bases ......--.+++-+++++++: 4]

. Buds pubescent, cleft 3/4-5/6. Anthers 10-14. Leaves membranous or charta-

GEGliss BAe oO RR a ee te eee 40. H. leptantha

. Buds glabrous, except at the very base; cleft c. 1/2 or less. Leaves generally mem-

rans ete fe a 42

. Buds subglobose. Anthers 12-16. Leaves oblong(-lanceolate), at apex (acute-)acu-

minate, not caudate (always?) .....-.--- sees eee ee eee ees 31. H. hellwigii

_ Buds obovoid or ellipsoid. Anthers 10(-12). Leaves oblong-lanceolate, at apex

andater ee... Yk OR FE Sa ee OR Lee chee 67. H. ralunensis

_ Leaves in fertile shoots dispersed, i.e. in 3 or more rows along the twigs. Leaf bud

proportionally short and broad (about 4 times longer than broad, or less) .... 44

_ Leaves in fertile shoots distichous (in rare cases a few on the same plant in 3 rows).

heat bud ‘generally moreslenden Ute ee rhe APs aa See 52

- Leaves + clustered towards the end of the twigs. Leaf bud and inflorescences with

hairs 0.5—1 mm long. Bark of older twigs often blackish, flaking. — Borneo
(Sarawak, Brunei); sandy soils .......--2+- see sere errr ees 71. H. sabulosa

_ Leaves clustered or not. Leaf bud and inflorescences with hairs up to 0.2 mm long.

Bark of older twigs not or slightly flaking ......-- +--+ sees errr etree 45

a Leaves with dotsibeneath (lens!)\eaw. .. ee ee es a, Se aia 46
Sieeavesiwithout dots | eee hs kbs eel. 9 Sis Se 47
a. Bark of twigs (grey-)brown, not contrasting with the dark colour of the petioles;

older bark not flaking. Leaves in 2 or 3 rows. —W, C&S Sumatra, Java.......
Ge SAE OE) OF PAI 5 Pre ded sm ace et tee acer ener 28. H. glabra

De Wilde — Myristicaceae (Horsfieldia) 67

b. Bark pale, grey or yellowish brown, rather contrasting with the petioles; older bark

Sia.

more or less flaking. Leaves in 3—5 rows. — Sumatra (N Aceh); atc. 1300m....

ELIT Tey AiUC ted Stee Sen is ol BR A 8. H. atjehensis
. Bark of twigs brown, not contrasting with the colour of the petioles ....... 48
. Bark of twigs pale, greyish or straw, contrasting with the blackish brown colour of

Mewes! LU UTA AEs Soke he OR Ac, SON a VES I 50
. Twigs ridged or short-winged. — Sumatra ................. 32. H. hirtiflora
Pier iesucrete, neitherridped nor Wiliged ee CRE eey Moe ee Seeeiere. 49
. Leaf pubescent beneath. — Sumatra, Peninsular Malaysia ..... 86. H. superba
Mitealvelabrous beneath. — Borneo». . ..02.. Nae i: 26. H. fragillima

. Pedicel articulated. Androecium with depressed apex, apical hollow broad, either

shallow or rather deep, reaching up to nearly halfway the column; androphore largely
hidden by the anthers. Leaves mosly distichous, sometimes tristichous. — Sumatra,

Eemarsular IWidlaysia;! 21.9.) A2.. a0 ae 85a. H. sucosa subsp. sucosa
. Pedicel mostly not articulated. Androecium not or but slightly depressed, the api-
cal hollow narrow and inconspicuous. Leaves generally in 3-5 rows ....... Sil
. Androphore + absent. Leaves blackish brown. — Borneo .. 55. H. pallidicaula
. Androphore 0.3—0.4 mm long. Leaves bright brown. — Peninsular Thailand,
Mimrdina ieenimsular Malaysia. 4. ..220«600%.06.01 cee ee ees 78. H. sparsa
. Buds + obconical-obovoid, very leathery, cleft 1/6—1/5(—1/4) only. Androecium
fmoinateranthers 3 /Sumatra'. 2s sow es 2 ee I 91. H. triandra
. Buds of variable shapes, cleft 1/5(—1/4) or more. Androecium various, anthers 4 or
more Guale flowers not known in Hi perangusta): . 2028. 088.5 2S. Se. 53

. Buds ellipsoid or obovoid, 3-8 mm long, cleft 1/5—1/4(—1/3). Leaves + parchment-

like, matt above with finely wrinkled surface (in H. sessilifolia male flowers not
hnowimandileaves notso distinctly matt)». 4.50 Pee SS 1 54

. Buds either 1) ellipsoid or short pear-shaped, 3.5 mm long or less (though some-

times rather large in H. endertii, H. flocculosa, H. majuscula, and H. wallichit), or
2) (depressed) globose; cleft (1/4—)1/3—1/2 or more. Texture of leaves variable in-

cluding coriaceous, never parchment-like and not typically matt above ..... 58
. Twigs 5—7(—10) mm diam. Leaves 25—70 cm long, indumentum persistent beneath
as Hansa lant OP. OD sti.) eRe aL ee ee ee 311
. Twigs 3-5 mm diam. Leaves up to 30 cm long, (largely) glabrous beneath, or with
some hairs persistent on the midrib. [Male perianth 3-5 mm long.] ........ a5
. Apical leaf bud with hairs 0.1—0.3 mm. Leaf midrib broad, margin flat ..... 56

. Leaf bud with hairs | mm long. Leaf midrib above line-shaped, margin (dry) rolled-

DEM ti a taal sg elie en ee Ee See 59. H. perangusta

. Leaves to 32 cm long, dark olivaceous. Twigs (yellowish) brown, coarsely striate

and tending to crack longitudinally. Inflorescences (almost) glabrous. Pedicel not
nerculatedeAmthersi 2-08 hii 2a oO, Pe eee 92. H. tristis

. Leaves up to 24 cm long, fulvous-brown. Twigs brown, finely striate, not cracking.

Inflorescences pubescent. Pedicel articulated. Anthers 10-12 ..... 27. H. fulva
Buds 7-8 mm long, perianth and pedicel glabrous. Petiole 6-15 mm long. — Su-
matras Peninsular Malaysia? 2tie eta eae, 86. H. superba

. Male buds not known; female perianth and pedicel pubescent. Petiole almost ab-

Seni Domeod Sasawakjyen ios. ete. Ae See: 75. H. sessilifolia

Flora Malesiana, Ser. I, Vol. 14 (2000)

_ Leaves with (sub)persistent indumentum beneath; in H. gracilis and H. wallichii
sometimes hairs vestigial on and near midrib and nerves .....-....+++++-- 59
_ Leaves glabrous or early glabrescent beneath; in some species often with vestigial
hairs'on the lower midnilyls i. wi a OR es 69
leaves: notiscabrousabove! jailsew ou noo. «20 S.C) a aaa: 60
Hveavestscabrous above tema .cst irene. ctsche oe seein eins 30. H. grandis
Leaves with dots and/or dashes beneath, obscured by hairs or not (lens!) .... 61
_ Leaves without dots or dashes beneath ...........-- 0 eee eee ee eee eeee 63
_ Buds short pear-shaped, 2—2.5 mm long; pedicel indistinct, (.3—1 mm long. Leaves
thinly pubescent beneath, often + glabrescent ..........---- 96. H. wallichii
. Buds subglobose, 1—1.5 mm diam., pedicel slender, 0.5—1.5 mm long. Leaves with
conspicuous indumentum beneath .......... 6... eee renee eee eee ees 62
. Twigs 2-3 mm diam.; leaf blades 7-15 cm long, [lateral nerves 5—9 per side]. —
Borneo (Sarawak); sandy soils .......6.--. eee eee eee 57. H. paucinervis
. Twigs 5—8(—14) mm diam.; leaf blades (18—)24—36 cm long. — Sumatra, Penin-
Sulan Malaysian list) G0 erie nay ae. ASPEN Ata eet se 63. H. pulcherrima
_ Buds broadly ellipsoid or obovoid, 2-3 mm long; androecium longer than broad.
Pedicel (1.5—)3—4 mm long. Twigs and lower leaf surface with hairs 1.5—2 mm
lorie RT 1. LS hE Panes Oe 25. H. flocculosa
_ Buds (depressed) globose, 2.5 mm diameter or less; androecium as broad as or
broader than long. Pedicel 3 mm long or less. Twigs and lower leaf surface with
nas tose S ema som es rs a SE, Sse, ie ee 64
_ Buds 1 mm diam., with persistent indumentum or late glabrescent. .........---
denice ER, ad ee Cs. «ee 46. H. motleyi
_ Buds 1—2.5 mm diam., glabrous or glabrescent ........--+- +++ sess eeee 65

a. Pedicel not or indistinctly articulated (this character not quite clear in H. gracilis

And ivr ufolanata)™. ean. A eet, Tee) CSRS or ol: Se 66
uPedicel articulatediaa.t00n. Ye eee ae eee eee 68. H. reticulata
. Upper leaf surface dark brown on drying, with venation + indistinct. — S Penin-

sular Thailand, Peninsular Malaysia .......-...---5-5005- 90. H. tomentosa
. Upper leaf surface olivaceous on drying, venation distinct or not. — Borneo 67

a. Twigs 3.5—7(-13) mm diam. Leaves 10— 45 cm long; venation distinct above; lat-

eral nerves 11 pairs or more. Buds 1.5—2.3 mm diam.; anthers 8 ormore .... 68

b. Twigs 3 mm diam. Leaves 12-21 cm long; venation indistinct above; lateral nerves

68a.

14-17 pairs; [lower leaf surface with sparse subpersistent hairs, vestigial mainly

on nerves and midrib]. Buds 0.6—1 mm diam.; anthers 16-18 ... 29. H. gracilis
Leaves 18—45 cm long; lateral nerves 18—25 pairs, sunken above. Buds 1.5—2 mm
diam.: anthers 8-10. — Borneo; lowland ...........-..--- 80. H. splendida

b. Leaves smaller, 10-23 cm long; lateral nerves 1 1—16 pairs, raised above. Buds 2—

2.3 mm diam.; anthers c. 15. — Borneo (Sarawak, Sabah); montane, 900-1400 m
Seer is we vite igs peace CE IS 8, ET SS 70. H. rufo-lanata

69a. Twigs at apex pale, greyish or straw, contrasting with the dark brown petiole 70
b. Twigs at apex (dark) brown, not contrasting with the petiole ..........--. 75
70a. Androecium angular in cross section. Anthers free for the upper half or more. —

Borneo: heath forest on sand or peat soil .........-..-55-- 52. H. oligocarpa

b. Androecium (sub)circular in cross section. Anthers largely connate ......-- 71

De Wilde — Myristicaceae (Horsfieldia) 69

(eltar

b.

Poa.

80a.

8la.

Leaves chartaceous, bright brown. [Pedicel not articulated. Androecium with small
apical hollow.] — Borneo; mostly in heath forest on sandy soils .. 12. H. carnosa
Leaves membranous, (blackish) brown. — Usually growing in forest on richer soils,
ppeleradimen Sarid) were. ereme Ah s> |; oes A. A Get AO at. G2:
Pedicel articulated. Androecium strongly depressed-globose; apical hollow broad
with flattish bottom, to nearly halfway into the androecium. — Sumatra, Peninsu-
Pentel Stat? aise tend chee IOS 48. H. sucosa subsp. sucosa

. Pedicel not articulated. Androecium ellipsoid to slightly depressed-globose; with

the apical hollow small and narrow (male flowers not known in H. discolor). —

Remnsular Malaysia: Romeo ly ital. cee oe eR eS PEL. 73
. Buds (sub)globose, 1.5—2(—2.2) mm long, androecium subglobose or short-ellip-
soid, | mm long. Leaves distichous or dispersed. — Borneo; up to 700m ... 74
. Buds ellipsoid, 2-2.4 mm long, androecium ellipsoid, 1.8—2 mm long. Leaves dis-
tichous. — Peninsular Malaysia; atc. 1300m .............. 23. H. elongata
eerste 0): cmilongvor less is. wn bhava Bae REG x toate 55. H. pallidicaula
BiaRncsHS Ci lone OrmMorenecs ad WL el ee 21. H. discolor
. Twigs ridged or nearly winged, also in the older wood .................. 76
. Twigs not ridged; sometimes twigs faintly ridged, lined, or angular in the apical
PeeAnIVIMTALED. street hon hid Se eee ence? Laeaten) 79
Piodsiclett nearly to the base:— New Guinea. ........ cee. Aue hee. . Wh
einosielett /2=2/3»—— West: Malesia w:2.¥). swt). bee Eee 78

. Buds slightly broader than long, short-pubescent in the lower half. Androecium

slightly broader than long; anthers erect, not incurved ........ 5. H. angularis

. Buds subglobose to broadly ellipsoid, glabrous. Androecium longer than broad,

+ obovoid, the anthers with apex free and incurved, those of one side of the androe-

cmnmclaspmetheothers iin Longe wee eee eee 51. H. olens
. Buds 2.5 mm diam., pubescent. — N Sumatra ............. 32. H. hirtiflora
. Buds 1-1.5 mm diam., glabrous. — S Peninsular Thailand, Sumatra, Peninsular
Malay siayBOmeoiwcwi! d an tshhuewe Seek he eee 11. H. brachiata

. Buds short pear-shaped, 2(—2.5) mm long, subsessile with the pedicel much shorter

than the perianth, 0.3-1 mm long, thickish. Leaves coriaceous, pubescent or
glabrescent, with dots and dashes beneath (lens!); lateral nerves flat or sunken above.
Br SOIT NOW 22. psntidiab naive sas oe dersineion bisa gs artnnysibese sige wove OREM toe Oe 96. H. wallichii

. Buds variable in shape and size, the pedicel proportionally longer and more slender

(buds obovoid with pedicel short in H. glabra var. oviflora). Leaves variable, nerves
raised or sunken, dots present or absent. Twigs solid or faintly hollow ...... 80
Inflorescences stout, the rachis towards the base 5—8 mm diam. Androecium about
as broad as long, triquetrous in cross section, [anthers entirely connate]. — W Bor-
eceeies his ulin ewok 5 duke jess epee 54. H. pachyrachis

. Inflorescences large or small, the rachis towards base 4(—4.5) mm thick or less.

Androecium triquetrous or circular in cross section. — Whole of W Malesia . 81
Androecium 3- or 4-angular in cross section. Anthers + erect, free for about half-
way or more. Buds 1.5(—2) mm diameter or less. Pedicel articulated. Leaves with
the lateral nerves raised above; dots absent (H. ridleyana with leaves small, nerves
sunken, male buds 1 mm diam.). — Most of W Malesia, not in Sulawesi, rare in the
PAROLES) Pesce ene ee elise eee arate ee iceeeaene Rec ected orate eee 82

70

Flora Malesiana, Ser. I, Vol. 14 (2000)

- Androecium in cross section circular, ellipsoid, or subtriangular with rounded an-

gles. Anthers + curved, almost entirely connate, free apices c. 1/3 or less. Buds
(1.3—)1.5 mm diameter or more. Pedicel articulated or not. Leaves with the lateral
nerves raised, level, or sunken above; dots present or absent (lens!) ........ 89

_ Leaves 5—16 cm long; midrib and lateral nerves level or sunken abovegwer =. 83
_ Leaves small or large, 5—28 cm long; midrib and lateral nerves raised above . 84

33a. Twigs and inflorescences rather glabrescent. Leaf apex acute or acute-acuminate;

84a.

86a.

87a.

88a.

89a.

nerves faint above. Anthers 4—6. — Peninsular Malaysia, Borneo..........----
PAT ee i er a ook ee ec Cee Er aaa 69. H. ridleyana

. Twigs late glabrescent, inflorescences with persistent indumentum. Leaf apex blunt;

nerves flat or but little raised above, clearly visible. Anthers 9 or 10. — Borneo
(Sarawak) sulunetin-e betel eid. ko de eRe, 50. H. obtusa
Leaves early glabrescent beneath, also on the midrib; leaf apex long acute-acumi-
nate. Twigs rather smooth, lower down cracking longitudinally. — Borneo (Bru-
nena .h 22 i cox deus lesb Heekus os Ho AE ee 22. H. disticha

_ Leaves early glabrescent beneath, but midrib sometimes late glabrescent; leaf apex

acute-acuminate. Twigs striate, lower down coarsely striate or finely cracking 85

. Twigs 1—2(—4) mm diam. Leaves 7-18 cm long, thinly membranous to subcharta-

ceous; petiole slender, 1—1.5(—2) mm diam. Inflorescences delicate, up to 9 cm
long; buds dumm diamin;.. 4.) BER Ae ese: CEE ea eee 86

. Twigs 1-5(—8) mm diam. Leaves of variable sizes, chartaceous or coriaceous; the

petiole 1.5-4(—8) mm diam. Inflorescences up to 15(—20) cm long; buds 1-2 mm
distin. UA ohscie isn ecto RR A SE CGS 22 O RR ee 87
Leaf bud with hairs (0.1—)0.2 mm long; twigs at apex and leaves glabrous; inflores-
cences with sparse stellate hairs 0.2 mm long, glabrescent. Leaves drying to a grey-
ish tinge. Male buds short pear-shaped, tapering into:the pedicels j-n eee re
bral Be GI IO Bes re 0h ate ee etal 89. H. tenuifolia

_ Leaf bud, apical part of twig, petiole, midrib beneath and inflorescences with woolly

stellate-dendroid hairs (0.2—)0.5 mm long; leaves olivaceous on drying. Male buds

globose or depressed-globose ........-- +--+ ss eeeee reese 41. H. macilenta
Twigs and leaves stout, the midrib broad above, at the transition to the petiole at
least 3 mm wide. Inflorescences 10-20 cm long. — Borneo; forests on poor soil,
including sand and peat ......-----s see e eset e treet eee 39. H. laticostata

. Twigs and leaves less robust; midrib above towards the insertion of the petiole less

than 3 mm wide. Inflorescences up to 15-20 cm long. — On poor or rich soil 88
Leaves 16-28 cm long, leaf base + rounded or short-attenuate, nerves 16-19 pairs,
very prominent above. — Borneo (Sarawak) ......-+.+---+->- 47. H. nervosa

_ Leaves 7-28 cm long, base short- to long-attenuate, nerves 6-16 pairs, raised to

variable degrees above. [On drying, colour of leaves above and beneath usually
much contrasting, generally more so than in the related species. ] — Variable, with
3 varieties (based on fruit size). Sumatra, Peninsular Malaysia, whole of Borneo,

Philippines (Mindanao, Palawan) ......------+--+--+--> 61. H. polyspherula
Leaf bud, apical part of twig, and inflorescences with hairs 0.2 mm long or more
(hairs 0.1—0.4 mm long in H. punctata) ........+ 00sec cree eens 90

. Leaf bud, apical part of twig, and inflorescences with hairs (0.2-)0.1 mm long or

le8Seccodle me fives deetlien oe eee ee. hr ee = Laeeni aia 97

De Wilde — Myristicaceae (Horsfieldia) 7\

90a.

94a.

95a.

96a.

100a.

Buds ellipsoid, 2.5—3.5 mm long; androecium longer than broad. [Leaves thick
and brittle, apex bluntish; lower surface usually with conspicuous pale golden

aerrnesears: (Leis!) Jil. Qasr sah: SP BA ee ee ee 24. H. endertii
. Buds (sub)globose; androecium not longer than broad ................- 9]
. Buds 2.5—3 mm diam. (or 1.5 mm in Hallier 624 from W Borneo, see the notes),
lemme: = Sumatran Nk newn’ eee eye GE ..2 95. H. valida
. Buds (1—)1.2—2.5 mm diam., cleft 1/3—2/3(—3/4). — Peninsular Malaysia, Bor-
en eee ive ti Teed hone ish a ea es RS SI. 2 92
MberVcsen ithidetsybencathir@ens!) 0s fy ek ee eS 93
. Leaves without dots beneath. [Pedicel not articulated.] ................. 94

. Pedicel articulated. Lateral nerves flat or sunken above. Lower leaf surface cin-

namon or chocolate, contrasting with upper surface. — Borneo..............
MMP USCA A HELEALG, S21 )00' Ghee a REUADE BUDE Soc te, RM a ewe 10. H. borneensis

. Pedicel not articulated. Lateral nerves largely raised above. Upper and lower leaf

surface not much contrasting. — Peninsular Malaysia ...... 65. H. punctata
Buds 2—2.5 mm diam.; androecium sessile, broadly saucer-shaped. Leaves 20—45
ecmalone..== Lowland forest 3) (is Joh Alt A Se 26. H. fragillima

. Buds (1—)1.4-2.2 mm diam.; androecium (depressed-)globose, with the apical

hollow small, concealed by the apices of the anthers. Leaves 4-35 cm long. —
Monrancworest at S0O—2000 My ». ane. ncanctene ee ape beeoee See eI aers a5
Androecium with slender androphore 0.3—0.8 mm long, not hidden by the an-
thers. Leaves membranous, 9-18 cm long, dark brown, [apex acute-acuminate]
oe po ee ee eee en eS ce nc 4. H. androphora

. Androecium (sub)sessile, androphore absent or up to 0.5 mm, largely hidden by

BP MAMT OEE S ge lic ere ecy cud), a Ry, abe wast) Suaialaci oe. Sic bate gt ge 96
Leaves chartaceous or membranous, to c. 35 cm long, olivaceous-brown; apex
acute-acuminate. Inflorescences to 20 cm long. — Borneo (Mt Kinabalu)......
«oo ot Oe ROE REN ote aie oa RM a pene aeRemmnntet Ss | 3. H. amplomontana

. Leaves coriaceous, 4-14 cm long, blackish; apex obtuse to subacute. Inflores-

eences + —l6'cnmllone AGlt. ASN tC, Es Ct 2 Ee Ie ae 45. H. montana
mibcaveswwithdots beneatix: Je yiles 0) 1ath JL, ft Fi SRS B® 104
. Leaves without dots beneath (dots should not be confused with smaller, blackish
POOWMUS)) 6 He te gam aye SK bibl Je Bae. Ayla wht eene SMe ae RAR Sane neg eee 98
. Buds + ellipsoid; androecium + obovoid, the apical part of the anthers deeply
inflexed into the apical hollow. — New Guinea ........... 74. H. sepikensis

. Buds and androecium of variable shapes; the anthers + straight or curved, at apex

not inflexed. — W and E Malesia, not in New Guinea ................. 99

. Perianth coriaceous; lobes thick, towards the base (0.3—)0.4—1 mm thick.

Androecium ellipsoid-obovoid, longer than broad .................... 102

. Perianth thinner, lobes at base 0.2—0.3 mm thick. Androecium subglobose, broad-

ly ellipsoid, or obovoid, not or but little longer than broad. [Androphore narrow,

only 0.2 mm long. Leaves membranous to chartaceous.]............... 100
Pedicel (1—)1.5—2 mm long, about as long as the perianth. — Peninsular Malay-
SIA MBORMIEO LEE SR eRe 6 22 ek ok 25 8 8 2 ee eee RRP, Roe) eae 101

. Pedicel shorter than the perianth, 0.5 mm long. — Sulawesi, Philippines.......

a eee ORE PUREED otal BAK Cid Th WT PELE Ee ee ke Oe 17. H. costulata

2, Flora Malesiana, Ser. I, Vol. 14 (2000)

10a. ABudsiclefecal/Qese eres EE Ls aoe 83. H. subalpina
elds. clett 2/34 ee ae A II Crore een 48. H. obscura
102a. Bark of twigs not flaking. Leaves + membranous .......---+++++++++: 103

b. Bark of twigs flaking or not. Leaves coriaceous. [Pedicel not articulated. Anthers
(3—)4-8; androphore rather broad, tapering, (0.1—)0.2—0.3 mm long.] — Borneo
(Sarawak, Sabah); kerangas, montane forest; 800-1550 m ... 97. H. xanthina

103a. Pedicel articulated. Anthers 7-9; androphore rather broad and tapering, 0.2—0.5

mm long. — Sumatra, Peninsular Malaysia; 0-1000m ..... 43. H. majuscula

b. Pedicel not articulated. Anthers 5 or 6; androphore narrow, 0.1—0.2 mm long,
hidden by the anthers. — C Sulawesi; 100-450 m ........--. 14. H. coriacea

104a. Twigs 2.5—3(—4) mm diam. Leaves (8—)12 cm long or more. Male buds (sub)-
elobose, 1)5=4:2immn diamnts vases £1). 2). a lees 2h) Se 105

b. Twigs 1.5-2 mm. Leaves 5-12 cm long. Male buds + ellipsoid or globose, 1.2—
1G tm dongth eae cee se eee SE 58. H. penangiana
105a-eBudsclett 3/4—4/5; anthers 7 —llionies: |. . « 1a: oe eee 106
b. Buds cleft 1/3—2/3; anthers 9-20. — Sumatra, Java ........-.++++++55 107
106a. Anthers 7-9. Dry fruits 4-5 cm long, pericarp 10-20 mm thick. — N Sumatra,
Peninsular Malaysia, Borneo .....--.--6- +e sete eee 66. H. punctatifolia

b. Anthers c. 11. Dry fruits 2 cm long, pericarp 1.5 mm thick. — Peninsular Malay-

Saude Lien didies s caine se SOA ote we ae 65. H. punctata

107a. Buds 3—4.2 mm diam.; anthers 15—20.— N & C Sumatra 42. H. macrothyrsa
b. Buds 1.5—2.5 mm diam.; anthers 9-15. — Variable, with 3 varieties. Java, W,C &
Ka GinivAataale < <o:.cchcut) 2s CEE a eens hae s.4 ee Core 28. H. glabra

(2) REGIONAL KEY TO THE SPECIES __ PENINSULAR MALAYSIA, SINGAPORE
(based on female flowering and fruiting specimens)

1a. Flower buds pubescent at base. [Ovary pubescent. Fruits pubescent, ellipsoid, 2.5—4
cm long. Flowers in dense clusters and strongly fragrant in male specimens.] —
Originating from Sri Lanka, cultivated in Penang, Singapore and elswhere.......

sn ibys Semepwen ved rides 4 bend de pave ces ates Aen Cee athe FoI Manet ge 36. H. iryaghedhi

b. Buds glabrous or early glabrescent .....--- +22 +--+ +++ 20s see Teens 2
Jay Perianth 2-lobed: Ovary glabrous 2. 22.5. 2.2) sec are + io bo Ra 3
b. Perianth 3- (or 4-)lobed. Ovary glabrous or pubescent .......--..+-++++000 5

3a. Twigs ridged or lined. Leaves membranous, often with irregular whitish blotches.
Fruits and seeds globose. — Plant usually riverine, in coastal areas .. 35. H. irya

b. Twigs not lined. Leaves without pale blotches. Fruits and seeds ellipsoid ..... -
da. Leaves membranous, glabrous beneath, dots absent. — Gardens’ Jungle, Singapore
(originating from E Malesia) .........---- +++ sss errr eres 56. H. parviflora

b. Leaves coriaceous, beneath pubescent and with dots (lens!) — Kerangas, peat swamp
fateSb moines teeta ee eee Beare oe eel 18. H. crassifolia

5a. Leaves with dots beneath (dots not to be confused with smaller dark punctation) 20
by. Leaves without dots beneath’. s.g.7che..- 226s v5 6 ae occ > AR ee 6
6a. Leaves with persistent indumentum beneath ........--- +2205 ee errr 7

b. Leaves glabrous or glabrescent beneath ......--- ++ +++ seer eter eres: JU

De Wilde — Myristicaceae (Horsfieldia) 73

7a.

b.

8a.

b.
9a.

Ovary pubescent. Fruits pubescent or at least with vestigial indumentum near the
Base spenanth Not persistent. Las |i. Lele 90. H. tomentosa
Ovary glabrous or with some incidental minute hairs. Fruits glabrous, perianth (at
ie eee ESE) METSISLENE cis A ies Hts minke My Sa IAT 2 a te 8
Hairs on lower (and upper) leaf surface harsh, with hardened hair bases, in older
leaves rendering the surface scabrous. Fruits |-1.4 cm long .... 30. H. grandis
leaves not. Scabrous..Fruits 2icm lone or more aera IE eee ee eee a 9
Twigs 3-5 mm diam. Leaf bud and twig apex with hairs 0.2—0.3 mm long. Leaf
blade 13-21 cm long, [matt above on drying. Fruits 2.2-2.4(-3) cm long] .......
Pe TSE Vn.) 2 Nd SU DA lads REE De 27. H. fulva

. Twigs 5-10 mm diam. Leaf bud and twig apex with hairs 0.5 mm long or more.

wecanlade 2 0=40(=70) cia long 4. 10.1, Re, BES ae Penance 10

. Leaf bud, twig apex and lower leaf surface with rather stiff, rust-coloured hairs

USS mm lone. Fruits 3:8=5.5 em long \ 2). ee wee eer 86. H. superba

. Leaf bud, twig apex and lower leaf surface with yellow-brown or pale brown woolly

iagse 2, mint lone Fruits\s:corlone'”.. 2th ecw ee 25. H. flocculosa

. Twigs pale brown or straw, contrasting with the blackish petiole. Leaves either

distiehous/or in 5=5) rows along the twigS” 2 at gees teem ales cease ae ar 12

. Twigs brown, not contrasting with the petiole. Leaves distichous .......... 14
Mibcavcs UIStiChoOus OF iI) SHOWS! As... ).) 08 ae SR ae eesti el Oe 13
. Leaves in 3—5 rows along the twigs. [Pedicel not articulated. Fruits 3-5.5 cm long,

BeRAMPheMOl PELSISLETIL, |! 2a". OURS Le. i tereleuclete Steere one neem 78. H. sparsa
. Leaves distichous. Pedicel not articulated. Male perianth + ellipsoid, 2—2.4 mm
Ma PaO SPN OC IKMOW HAG. i608. i 30k iia ates Age ene 23. H. elongata
. Leaves distichous or in 3 rows. Pedicel articulated (this character best seen in male
flowers). Male perianth globose, smaller. Fruits 2.5—3.5 cm long, perianth persist-
Bite oo <adgsie foley SOIR RE. Es SPOPORREEDL OMEN Pr BeptonenCrIAC cent 85a. H. sucosa subsp. sucosa
. Leaf upper surface matt on drying caused by fine wrinkles; nerves flat or sunken.
Biutsmsually with persistent, pertanthy 2). Sele cpctpe eee 27. H. fulva
. Leaves above not particularly matt, not finely wrinkled; nerves flat or raised. Peri-
Fimo lpersistent underthe fruits Ay. sa). > ey eloeyactaee terior ree it
mleateral nerves fat or buttamtly raised above .¢).\0.192..0. ange here © 16
. Lateral nerves distinctly raised above. [Pedicel articulated; this character best seen
GMA SEL O Wer Se]! Oey LO LADS. SOURS ae A Sn ED heres oui 18
. Leaf bud with hairs | mm long. [Midrib narrow, line-shaped, sunken above; blade
KONE NA Eyes oer irs elerelt. amrresd cna Bod Gace geter 59. H. perangusta
Ricah bud with hairsimuUchsShorteny.a4s< she Saas See eee eee = 7.
. Leaf bud with hairs 0.1 mm long. Twigs 2.5—5 mm diam. Leaves 15—27 cm long.
pedicel motarticulatedtt tie0 )..2 See ee 83a. H. subalpina subsp. subalpina
. Leaf bud with hairs 0.2-0.4 mm long. Twigs 1.5—3.5 mm diam. Leaves 5—15 cm
loneyRediccl-articulatedag AAvwe? (a) 2 Oe Bay eee re 69. H. ridleyana
. Leaf bud with hairs 0.1 mm long. Midrib early glabrescent beneath. Fruits 4.5—6.5
emt lone, swith thick pericarp. Sh. 00+. See ee ewe eee 43. H. majuscula
. Leaf bud with hairs 0.1 mm long or usually much longer. Midrib often late glabres-

cent beneath. Fruits 2-4 cm long, pericarp 2—5(—7) mm thick ............ 2)

Flora Malesiana, Ser. I, Vol. 14 (2000)

3a.

Sa.

. Buds glabrous (pubescent at base in H. hirtiflora and H. triandrd) eee
- Leaves membranous, often with irregular whitish blotches. Fruits globose, 1.5—2

. Twigs + angular, lined or low-ridged. [Fruits 2-3(—4) em long.| 11. H. brachiata
. Twigs terete, neither lined nor ridged .........- 6. esse ence eee eee eens 20
. Twigs 2-5 mm diam. Leaves usually chartaceous. Fruits 1.9-3.5 cmlong.......

Rael of stern fab bee em ee aes, «carte reenter 61. H. polyspherula

. Twigs 1-3 mm diam. Leaves membranous. Fruits 2.3=2.4 cm long icetgeiaeer an -

i EE DE. Mn bin bid he eat ce cede Meigen ie 41. H. macilenta

a. Leaves usually with persistent indumentum beneath ..............+++-5: 22
. Leaves glabrous or glabrescent beneath ...........+ +++ tees eee eeeeeees 23
a. Leaves + pubescent, sometimes late glabrescent, with dots and dashes beneath.

Ovary glabrous. Fruits 4-6 cm long, glabrous, perianth usually persistent .......
Eb gerer Phe oea att yee SEMerseeetect sith taeid. eae dere 96. H. wallichii

_ Leaves always pubescent beneath, with dots, not with dashes. Ovary pubescent.

Fruits 1.6—1.8 cm long, shaggy-hairy, perianth not persistent 63. H. pulcherrima

. Twigs 1.5—2 mm diam. Fruits 1.1-2 cm long ......-.----- 58. H. penangiana
. Twigs somewhat stouter, 2.5—5 mm diam. Fruits 2 cm long. or More see 24
_ Leaves coriaceous, apex blunt or subacute. Fruits 2-2.3 cm long, pericarp thin. —

Montane species of C Peninsular Malaysia .......------+--- 65. H. punctata

_ Leaves membranous, apex acute-acuminate. Fruits 4.5—8 cm long, pericarp 10—20

mm thick. — Widespread in W Malesia; forests up toc. 1100m .......--..--:
PS eae ae er enters. oo Re ee rans e seta. c 66. H. punctatifolia

(3) REGIONAL KEY TO THE SPECIES — SUMATRA, JAVA

(based on female flowering and fruiting specimens)

_ Flower buds pubescent at base. [Ovary pubescent. Fruits ellipsoid, 2.5—4 cm long,

pubescent. In male the flowers in dense clusters, strongly fragrant.] — Cultivated,
originating from Sri Lanka ..............5++++++--eee 36. H. iryaghedhi

i)

cm diam., glabrous; pericarp 1-2 mm thick; seeds globose. [Perianth 2-lobed; ovary
glabrous.] — Riverine or marshy, usually near the coast .........- 35. H. irya

_ Leaves of different consistency, usually not blotched. Fruits and seeds ellipsoid. —

Plantnot. coastal .24 82 teeeee ee ee oe een coe ee 3
Perianth 2-lobed. Leaves with dots beneath (dots not to be confused with smaller
punctation of different origin, lens!) ............-+++++++++ sess eeeaeees 4

_ Perianth 3- (or 4-)lobed. Leaves with or without dots beneath: ».\. ees Mae 5
4a.

Leaves coriaceous, 10-20(—28) cm long, finely pubescent beneath. Twigs 2-6 mm
diam. Fruits 1.5—2.2 cm long, with persistent perianth. — Peat swamp or padang
POLLEY Meee eee OM MES CEPT EAR a FOP Pot ky ohne RR Ott bs Lt 18. H. crassifolia

_ Leaves membranous or thinly chartaceous, 5—12 cm long, glabrous beneath. Twigs

1.52 mm diam. Fruits 1-2 cm long; perianth not persistent. — Mixed forest ....
ER OM ob 5 ahi tae eet Bendis wok 58a. H. penangiana subsp. penangiana
Ovary and fruits (at least at base) pubescent. Leaves with persistent indumentum
beneath Sa 2c dewiaatlialec mages Geos nao i eRe ASE) Et ee 6

. Ovary and fruits glabrous (fruits almost glabrous in H. triandra). Leaves glabrous

or pubescent beneatlinas Sot ingen Bee Hee EEE EEA 7

De Wilde — Myristicaceae (Horsfieldia) 75

6a.

14a.

Twigs 2—5 mm diam. Leaves 9-27 cm long, lower surface without dots. Fruits with
hairs 0.5 mm long or less. — Thailand, Peninsular Malaysia; specimens from Su-
MmiEmanmorseen liranehet<l den ddan. eRe ere 90. H. tomentosa

. Twigs 5—8 mm diam. Leaves 20-36 cm long, with dots beneath (lens!). Fruits with

Eeriipsee SENT OFS G22. GOR Ed Ahad a,c c/o vee Slee me eerie 63. H. pulcherrima

. Leaves with dots beneath (lens!). Lateral nerves generally flat or sunken above

ere itd. canolwiuuy b. 1: 2 Shek) Deere ee 21

. Leaves without dots beneath. Nerves either raised or flat to sunken above .... 8
. Leaves with persistent indumentum beneath. Fruits with persistent perianth .. 9
. Leaves glabrous or glabrescent beneath. Fruits with persistent perianth or not 11
. Hairs harsh, older leaves scabrous beneath. [Fruits 1-1.4 cm long.]............

RC hi CTA Ua bb ih BS Ee aa ee ae 30. H. grandis

Haims sotter: older leaves not scabrous beneath?) 22022 0 2a Pera: 10
. Twigs 5-8 mm diam.; leaves 20-40(—70) cm long. Leaf bud with hairs 0.5-1 mm.

erick seo. Svein Lome AU at) Pahl aed Re Sl Sih eet 86. H. superba

. Twigs 3—5 mm diam.; leaves 13-21 cm long. Leaf bud with hairs 0.2—0.3 mm long.

[Fil iG P30 dak (Cy Oe pee ea eco i APRN CLP ach, Pacurig Naik cole 27. H. fulva

. Lateral nerves flat or sunken or but faintly raised above. Colour of lower leaf sur-

face generally greyish brown, not much contrasting with upper surface ..... 192

. Lateral nerves distinctly raised above. Colour of the lower leaf surface bright brown

Omchocolate, contrasting withthe upper suriace 5220-2 ee oe eee 16

. Twigs 1.5—3 mm diam. Leaf bud, twig apex, and young inflorescences with woolly

Hans 307 min: eaves:s=9icnr long | Ae eee een 91. H. triandra

. Twigs (2—)3-10 mm diam. Leaf bud, twig apex, and inflorescences 0.1—0.3 mm

imnemibeaves morethan lOicmilone: |...) 25.504: igs we es en pg 113

. Leaves distichous, matt on drying caused by minutely wrinkled upper surface. Bark

Hen IOS straWwiOn DLOWACh 25 HLA! Vee RS MN ee eS Peers = ie: 14

. Leaves distichous or in 3—5 rows; not particularly matt, upper surface not finely

wrinkled. Bark of twigs pale, grey-brown or straw, contrasting with the blackish
emo lcrrmnyer ils sir atts. i belire a ds ech: Winistds Mas ity gaete Bad eae Ie Pa ito 15
Leaves elliptic-oblong to oblong, olivaceous-brown above. Stem grey-brown, not
much contrasting with petiole. Fruits brown on drying, 2.2-3 cm long, perianth
BSUSISLER Ged). th acl) e kotoneaksldarr ke SN geet eos Kine eee 27. H. fulva

. Leaves elliptic-oblong to lanceolate, dark olivaceous above. Stem pale, + yellow-

ish brown, rather contrasting with the petiole. Fruits blackish on drying, 1.5 cm

ae eMetANt NOW PELSISLEME y joins, </41 orn) weaqugel found eer oe et 92. H. tristis
. Leaves distichous or in 3 rows. Fruits 2.5—3.5 cm long, with the perianth persistent.
Pedicel articulated (best seen in male flowers) ... 85a. H. sucosa subsp. sucosa
. Leaves in 3-5 rows. Fruits 3—5.5 cm long, perianth not persistent. Pedicel not ar-
melated pauiiwa janis. Peete. wiki 208. GROW bee Lene 78. H. sparsa
. Perianth 4-lobed. Pedicel not articulated. Fruits 8-9 cm long. [Flowers known only
from the remnants persistent under the young fruits.] .......... 95. H. valida
. Perianth generally 3-lobed. Pedicel articulated. Fruits up to 6.5 cmlong .... 17
moiwiessancularwathdinesionmidges: |p. tae cleans. PORE So eee 18

L Twigs tesete,or faintly angular.neither lined normdgedV. aes heise - 19

Flora Malesiana, Ser. I, Vol. 14 (2000)

2a.
. Perianth predominantly 3- (or 4-)lobed .....------ ++ eee eee eee eee i
3a.

. Buds pubescent (known only in male flowers). Fruits 5—6 cm long. — N Sumatra

Perl correct re EE RS. DOSS 6 ets. APIO. BAN! 32. H. hirtiflora

. Buds glabrous. Fruits 2-4 cm long ......----- ++ s0+ sree eee 11. H. brachiata
. Twigs 1-3 mm diam. Leaves membranous, midrib beneath late glabrescent. Fruits

BAD A crn lGns rei ee ee eee OSE OCR NE cartes Ce heaaee 41. H. macilenta

. Twigs 2-5 mm diam. Leaves chartaceous. Fruits 1:9—65.cmiong aAeaeeee 20
_ Leaf bud and inflorescences with hairs 0.1—0.2 mm long. Midrib beneath early gla-

brescent; Fruits4:5=615 ‘emlong se. Sco" kes ee 43. H. majuscula

Leaf bud and inflorescences with hairs 0.1—0.6 mm long. Midrib often late gla-

brescent. Fruits 1.9-3.5 cm long. — Variable; 3 varieties based on fruit size .....
tedexd ch oHobree tb eee eRe? «rere eerie * 61. H. polyspherula

_ Leaves usually with persistent indumentum beneath (sometimes glabrescent), and

with both dots and dashes (lens!). Twigs conspicuously hollow. Fruits 4-7 cm long,
the perianth generally persistent .........-----++++++eeee> 96. H. wallichii

. Leaves glabrous or glabrescent beneath, with dots, not with dashes. Twigs not con-

spicuously hollow. Fruits variable ........--. 0600s terest es 22,

_ Bark of twigs pale, greyish to straw-coloured, contrasting with blackish petiole.

Leaves dispersed in 3—5 rows. [Female flowers and fruits not known.| — Sumatra
(NPA GEh): at: cal S00. ae rae tel jn eee arises + 8. H. atjehensis

_ Bark of twigs brown, not contrasting with petiole. Leaves distichous (or in 3 rows

in A. glabra, p.p.). -=.. +» - agakayoomeuey- 12) oe Se 23

_ Fruits (4.5—-)5—8 cm long, pericarp 10-20 mm thick ...... 66. H. punctatifolia
. Fruits 1-2.5 cm long, pericarp much thinner .........-- +--+ esse seers 24
. Twigs 1.5—2 mm diam. Leaves 5—12 cm long. Fruits |.1—2 cmlong eae

ee att iirtabate der Aah wth aero ference amas oie" 58. H. penangiana

. Twigs 2.5—4(—6) mm diam. Leaves (8—)12 cm long or more. Fruits 1.8—2.5 cm long

MIRA UCL OF) LAME os 8 ce BE. 6 $s Spee ee ee yi)

_ Leaves distichous. — C & N Sumatra .................; 42. H. macrothyrsa
_ Leaves distichous or in 3 rows. — S Sumatra, Mentawai Is., north to Simeulué [.,

eave tee ye Me eh tt PRIMM alee s Soe ccs Dac ene eeeme 28. H. glabra

(4) REGIONAL KEY TO THE SPECIES — BORNEO
(based on female flowering and fruiting specimens)

_ Leaves membranous, usually with irregularly shaped whitish blotches. Fruits glo-

bose, 1.5—2 cm diam., glabrous; pericarp 1-2 mm thick; seeds globose. [Perianth
2-lobed; ovary glabrous.] — Plants usually growing not too far from the coast ...
AB IOG CA ILOG SILA 2 ON AL Spa Some Pntemta saree cctier e eer ceNe oid tee a 35. H. irya

_ Leaves variable, usually not white-blotched. Fruits subglobose or ellipsoid; seeds

ellipsoid. — Plants coastal or not ... 2.2.6. ... ee eee eee eee eee eee nn aes 2
Périanth: 2-lobed sits 6 2 O08 Re EE? Se 6 Se ee RE se Ee 3

Leaves coriaceous, densely short-pubescent and with dots beneath (lens!). Twigs
grey-brown, not contrasting with petioles ........-....---- 18. H. crassifolia

_ Leaves membranous, glabrous and without dots beneath. Twigs greyish or straw-

coloured, contrasting with blackish petioles ........-..-- +++ eee seers +

De Wilde — Myristicaceae (Horsfieldia) aa

Aa.

Inflorescences + spike-like, 5-10 cm long. Perianth persistent under the fruits (al-
mas) 1, pormeo!(SE Sabah) 2... .+.00e shes eaek oe ee ose ae 82. H. sterilis

. Inflorescences branched, 1—2 cm long. Perianth not persistent under the fruits. —

Borneo (Sarawak, Sabah, E, C & S Kalimantan) . 85b. H. sucosa subsp. bifissa

Mieiwes imo or more rows along the twigs 25.05 2)78-) okie oe alee? => 6
Beare aishichous bias «dtodd isn ilg paw hel ee sae mie bate 8
. Leaves + clustered at the apex of the twigs. Petiole proportionally long and slender,

25-50 mm long. — Northern Borneo; sandy soils ........... 71. H. sabulosa
Mieaves not clustered, Petiole proportionally shorter -°-1.2 23ee1-2 7. eee ee - i}

. Leaves 10-30 cm long. Twigs pale, grey or straw-coloured, contrasting with the

blackish petioles. Buds 2.5-3 mm long. Fruits 1.5-6 cm long 55. H. pallidicaula

. Leaves 20—45 cm. Twigs brown, not contrasting with the petioles. Buds 4-5 mm

lonesEroits 6S cmilongy sit). Axe Gentn eee 2e8 oe Iga ae 26. H. fragillima
eiwiersincd or ridged whit. SNe oes Sees eee 11. H. brachiata
. Twigs terete or faintly angular, neither distinctly lined nor ridged .......... 2

. Leaves with (sub)persistent indumentum beneath (sometimes largely glabrescent

opty Bi UCD LATS 112) ee ERIE ERE At nl UENO oo ce ten) All “So cb tc ae 10

. Leaves glabrous or early glabrescent beneath (midrib sometimes late glabrescent)

ee Ree. orc Pie SRC: Zee ae AC eae eee ade 18

. Plant stout; leaves 50 cm long, petiole 3 mm long only. — Borneo (lowland Sarawak)

OIE ALLY is tH ourn Ou. Gel ae Ok. t. Poo eee 75. H. sessilifolia

. Plants variable in habit; leaves large, but petiole proportionally much longer .. 11
. Older leaves with scabrous hair scars above and beneath. Fruits 1-1.4 cm long ..

BPC Polis Wi. algae eC MOL b eS Rl. cthvoie bk Aeoh Se ere & 30. H. grandis
Bee Aves NOt SCADEOUS) PTUitS 1anger «.45)50,05 gens © ys eee ee ee eee 12
. Leaves with dots and/or dashes beneath (lens!) ............. 96. H. wallichii
. Leaves with or without dots beneath, never with dashes ................. 13
. Flower buds with persistent indumentum; fruits without persistent perianth .....

IE Doo ea regs url pauiecs sole IAL AEE AEE Oa) CO So: AT 46. H. motleyi
. Buds glabrous or early glabrescent; fruits with persistent perianth or not .... 14
. Ovary pubescent; fruits sometimes pubescent only towards the base ....... 15
mOvcrvrand fruits glabrous: past sive whcieoeeh ht wel. SURIMeeeee aes Sele ada k 16

. Twigs 3.5—5 mm diam. Leaves 10-23 cm long; nerves 11-16 pairs. Fruits largely

elabrescent: perianth not persistent |4).sy-sc). Gee ee eee ee 70. H. rufo-lanata

. Twigs 4-7 mm diam. Leaves 18—45 cm long, nerves 18—25 pairs. Fruits pubescent,

MPCESISteNt PEHianth: 2 2.).2 «00+ oka eee eee 80. H. splendida

. Twigs 3-6 mm diam. Leaves 18-35 cm long; upper surface sometimes bullate;

Imerves 17—20 pairs.) Fruits 2.3—2.J1em lone e eee esse. 68. H. reticulata

. Twigs 1.5-3 mm diam. Leaves often smaller, not bullate. Fruits 1-1.5 cm long 17
. Leaves membranous, beneath without dots; nerves 14-17 pairs . 29. H. gracilis
. Leaves thinly chartaceous, beneath with dots; nerves 5—9(—11) pairs ...........

LE «SL: nee eee eee nee rE Re ee ee 57. H. paucinervis

. Leaves with dots or with dots and dashes beneath (lens!) (dots not to be confused

with smaller, irregularly spaced points, which are usually present) ......... 19

. Leaves without dots beneath (enlarged hair scars sometimes present) ...... Ze

Flora Malesiana, Ser. I, Vol. 14 (2000)

19a

25ab

28a.

oa

_ Leaf bud, twig apex, and young inflorescences with hairs 0.2 mm long or more
ER ER. ed wre oes thee sears whento nee AIR Red SEN 20
Leaf bud, twig apex, and young inflorescences with hairs 0.1 mm long or less 2Al
. Twigs hollow. Leaves often with persistent indumentum beneath. Pedicel not ar-

ticulated. Fruits with persistent perianth ................+.- 96. H. wallichii
. Twigs (almost) solid. Leaves glabrescent beneath. Pedicel articulated. Fruits with-

Olt persistent periamthiey as LS EEE 10. H. borneensis
. Twigs 1.5-2 mm diam. Leaves 5—12(—17) cm long; nerves 8-11 pairs. Fruits 1.1—2
cnlone; pericarp thin ees 7a errs ee ere 58. H. penangiana
Twigs 2.5—4 mm diam. Leaves 9-21 cm long, nerves 11-16 pairs. Fruits 4.5—-8 cm
longewath thickipenicanp? 214 20 2.522. eS oe 66. H. punctatifolia
Twigs pale, grey-brown or yellowish, contrasting with the dark brown petioles 23
. Twigs brown on drying, + not contrasting with the petioles ...........+.-- |
_ Leaves membranous, (blackish) brown above, somewhat paler beneath. Perianth

persistent under the fruits. — Mixed forest ..........---- 55. H. pallidicaula
_ Leaves usually chartaceous, bright brown or olivaceous above. Perianth not per-

Sistent wider the fruits” 17. ice eo ee scare ererenenere ciate 5 eR 24

. Twigs 2-3 mm diam. Leaves 7—16 cm long, bright brown or chocolate beneath,

contrasting with the grey-olivaceous upper surface. Fruits 1.8-2.7 cm long. —
Kerancast peatilorest”s Vie pameestee: Pete c casa rem tne 52. H. oligocarpa
. Twigs 3-10 mm diam. Leaves 13-35 cm long, the lower surface not conspicuously
contrasting with upper surface... 2... 1.2 ee eee eee 25
Fruits 5 cm long or more. Leaves not conspicuously matt above . 21. H. discolor
_ Fruits 2 cm long or less. Leaves matt, caused by finely wrinkled upper SULLAce =. .
we bela ce te bute Saye toe ta vane Yosnite Gp todo eamememibags taimamabe, Sete been Pann Aoi Ren Noi 26
_ Leaves (elliptic-)oblong. Fruits 1.6—2 cm long. — Heath forest, peat swamp forest
Be Sa te nce te tates + i RL Tae ee os uote). sae eRe Re 12. H. carnosa
. Leaves elliptic-oblong to lanceolate. Fruits 1.5 cm long. — Mixed forest.) 4+-
WOT EN OR 5 che oo rate a AIR eRe ER Vero gee re a arerar ce Ne 92. H. tristis
_ Leaf bud and immature inflorescences with hairs 0.1 mm long or less. Lateral nerves
flat or sunken, or but little raised above ............---- 2025 ese e scans 28
_ Leaf bud and immature inflorescences with hairs 0.1 mm long or more; lateral
nerves above raised or not; if hairs only 0.1 mm long, then the lateral nerves above
distinctly raised, at least in the lower half ...........--. 0s eee eee ee eees 30
Species from lowland limestone, up to c. 700 m. Leaves membranous. Fruits not
known — Boreo:(NE. Kalumantan).7-)3-2 22 se: 22 48. H. obscura
. Montane species; 800-1800 m. Leaves membranous or coriaceous. Fruits 5 cm
long or less. — Borneo (Sarawak, Sabah) ........--- +++ e esse eres 29
_ Leaves without distinct large hair scars beneath (lens!). Fruits 3-5 cm long .....
ae hy fe EG Roth A eee 83b. H. subalpina subsp. kinabaluensis
_ Leaves with (usually) distinct yellowish enlarged hair scars beneath. Fruits 3.5—5
Shh Als Socks Gales Pa PAHs LS: a 97. H. xanthina
_ Pedicel not articulated (best seen in male flowers). Fruits with perianth persistent
Oe AOS <p, AEA Ai ts AO IG RES LAE oe rte ee 31
. Pedicel articulated. Perianth not persistent under the fruits ..........-..-- 36

De Wilde — Myristicaceae (Horsfieldia) 719

31a.

Leaves 15—45 cm long. Fruits 6 cm long or more; perianth + persistent. — Low-
PO mommontanemorest hy ses ck) b A oe eee 19 EE 32
. Leaves 5—20 cm long. Fruits 2-4 cm long; perianth not persistent. — Montane for-
Eeesias O02 OOO Tad ateligotea is ates les op delde te Meera owls Dheke See ele 34
. Nerves 11-22 pairs. — Borneo (Sabah: Mt Kinabalu); 1000-1500m ..........
Blac isei te sos atheist: oo coheed ae 3. H. amplomontana
. Nerves 20-30 pairs. — Forests up toc. 1000m ............-----++-55-- 33

_ Female flowers and fruits not known. — Hallier 624, Mt Damoes, W Kalimantan;

probably an undescribed species close to H. valida, see there . 95. aff. H. valida

. Perianth 4-5 mm long. Fruits 6-8 cm long, pericarp 10-20 mm thick ..........

8s cris (I deere Gdk aient + remit. vceeions SERIE 26. H. fragillima

. Leaves membranous, apex acute-acuminate. Fruits 2.4-3 cm long .............

ae le es See ae i cro o 4. H. androphora

. Leaves chartaceous or coriaceous, apex rounded to (sub)acute, not acute-acumi-

opis . eee er ene ee ee ee rises GI: ce ee. 35

. Leaves chartaceous or coriaceous, without large hair scars beneath (lens!). Peri-

ne monn lone. Fruits 227i LONG bo. 9. ty non og eeaetee on) ee 45. H. montana

. Leaves usually strongly coriaceous, usually with large hair scars beneath. Perianth

Poss in longesF ruts 34. Crm NOM orga oot icon oemecpenenenee eee 24. H. endertii

. Male inflorescences very stout, the rachis towards the base 5-8 mm diam. [Female

inflorescences, flowers, and fruits not known.] — Borneo (W Kalimantan) ......

i rere ree ere ee 54. H. pachyrachis
iialeimlorescences JESS SOUL oe ak strc oO ocean Ree ea ene ei ieee 3]
. Midrib on upper leaf surface towards the transition to the petiole 3 mm broad or

fos, . ER eee ae ererenas Meiers 39. H. laticostata
Mevtrartiar ASE MATMOWE! < 6c cule. osc ee se ae ee aie ae © + We ener eee 38
. Leaves 16-28 cm long, base rounded or short-attenuate; nerves 16-19 pairs. —

RITE ORCS AGA WAKe)! gioco: pete at i text cor yok Se ches 2 oie eae 47. H. nervosa
. Leaves 5-28 cm long, base rounded, short-, or long-attenuate; nerves 5—15(—20)

DLIEES « see ee RSet eR me INE NSIS ak ks apatgare™ 39
. Lateral nerves sunken, flattish, or but slightly raised above ............... 40
ME MAN MEIVE SALSCGADOW Es 6 2 lac ried 0 give o 25 epicenter eee oe 4]
mieaieape erOunded=s PHUIES MOUSESI oes es eeu oe en 50. H. obtusa
mitcat apex acute-acuminate. Fruits 5-2 em long .-..-.>2an- 69. H. ridleyana
. Leaf bud and young inflorescences with hairs 0.1—0.2 mm long. Leaves on drying

dull, greyish brown, colour of upper and lower surface not much contrasting. Fruits
PP POMUOMO a. cate ae ces cocks 2 Senn sre 2 a ayer ere gee eee 89. H. tenuifolia

. Leaf bud and young inflorescences with hairs 0.2 mm long or more; if hairs 0.1

mm long, then the olivaceous to dark-brown upper leaf surface much contrasting
Witte Ciinamom COLOUR DCMEAUM ... ce a's «<2 che a sens amie everest cee 42

. Twigs 1-3 mm diam. Leaves membranous, 10—18(—27) cm long. Fruits 2.3-2.4 cm

Ol Sim cpeeneee stewie ae abiesas, - saa MOM iret See odiods 2s 41. H. macilenta

. Twigs 2-5 mm diam. Leaves chartaceous; leaves and fruits of variable sizes . 43
. Twigs early glabrescent; bark + longitudinally cracking. Leaf apex long acute-

acuminate. Fruits 2.8—3.2 cm long; pericarp hard-woody, 8—10 mm thick. — Bor-
MEGUMI IOAN ere etek Sota Acts a eee Sie eee re eemete ees 22. H. disticha

80

Flora Malesiana, Ser. I, Vol. 14 (2000)

b.

Twigs late glabrescent; bark striate, not cracking. Leaf apex acute-acuminate, the
acumen not conspicuously long. Fruits 1.9-6 cm long. — Variable, with 3 varieties
(based on fruit size). Whole of Borneo ............-.--- 61. H. polyspherula

(5) REGIONAL KEY TO THE SPECIES — PHILIPPINES, SULAWESI, MOLUCCAS

la.

Da:

3a.

4a.

Sa.

6a.

Ta.

8a.

(based on female flowering and fruiting specimens)

Leaves membranous, usually irregularly whitish blotched. Perianth 2-lobed; ovary
glabrous. Fruits globose, 1.5—-2 cm diam., glabrous; pericarp 1-2 mm thick; seeds

globose. — Riverine or marshy, mostly near the coast ...........-- 35. H. irya
_ Leaves variable, usually not whitish blotched. Fruits subglobose or ellipsoid; seeds
ellipsord.— Coastal or mOtwen acme ines + <7 te SR + ho 2
Perianth 3-lobed. [Ovary and fruits glabrous.] ...........-.2 00 eee eee eees 3
. Petianth2-lobed) due. Get Rees ee ES eh 5

Leaf bud and inflorescences with hairs 0.2-0.6 mm long. Pedicel articulated. [Fruits
2(-2.5) cm long.] — Philippines (Mindanao, Palawan) ........---+++++++++++-
SA Poche Ota SLA at, Eo eae ee Oe ee 61a. H. polyspherula var. polyspherula

Fos es A bods Cao ae De aes lk RM MURR TIS 6S 4
Fruits 3.5-7 cm long; (dry) pericarp (4-)8—-15 mm thick. Leaves + membranous,
olivaceous-brown, midrib glabrous above; leaves sometimes with whitish blotches
as in H. irya. — Philippines, Sulawesi ...............-++--- 17. H. costulata

_ Fruits 4 cm long, (dry) pericarp 3.5—8 mm thick. Leaves membranous to thinly cori-

aceous, brown: midrib towards the base pubescent above in younger leaves. —
G@ Siilawesiiieat. hloae . ee Ts tec os eee 14. H. coriacea
Twigs 4-14(—20) mm diam. Leaves 20-45 cm long, petiole 2—7 mm long. Leaf bud
and inflorescences with hairs 0.3—1(—1.5) mm long. Buds 3.5—5 mm long, glabrous;
ovary glabrous. Fruits 3.5-5.5 cm long, glabrous ............. 87. H. sylvestris

. Twigs more slender. Leaves smaller, petiole comparatively longer. Leaf bud and in-

florescences with hairs up to 0.2 mm long. Buds 3(—3.5) mm long or less. Fruits up
to 3 cm long (in H. lancifolia to 3.5 cm long) .........--+ ++ +eeeee eee eeee: 6
Leaves + chartaceous, oblong-lanceolate to lanceolate. Buds cleft c. 1/4; ovary pu-
bescent. Fruits often + pear-shaped, 2.5-3.5 cm long, early glabrescent; (dry) pericarp
A= < nam thick. ——iSUlaWeSia ahem mrtn ten foe ise atk lars re 38. H. lancifolia

_ Leaves of different consistency, generally broader, oblong to oblong-lanceolate. Buds

cleft.c. 1/3 or more. Fruits 1=3 cml long. ..naq6 cs. Bgl eee a
Ovary and fruits pubescent; hairs on the fruits may be inconspicuous and only re-
maining at the very base near the insertion of the pedicel (lens!); pericarp thick or

ih ee er ee eens OM EOTIC TE coice Ao op o 8
. Ovary and fruits glabrous; pericarp 1-2 mm thick. [Female flowers and fruits not
known in H. aruana and H. samarensis.]. 2... 6. 0066 6 ec i De Be ee 11
Flower buds 2.5—3 mm long, cleft 1/3—1/2. Fruits (1.6—)1.8—3 cm long; pericarp
2 raniinat ti nee oes cee a ke a ee iim Goo ena 37. H. laevigata

_ Female flowers not known. Fruits smaller, pericarp thinner .........--.+-+-- 2)

De Wilde — Myristicaceae (Horsfieldia) 81

9a.

2a.
. Twigs terete, sometimes lined in-between the bases of petioles but neither angled

Fruits 1.5—1.6 cm long, short-ellipsoid. [Male buds transversely ellipsoid, cleft
2/3—4/5. Leaves 8-30 cm long, membranous or chartaceous; nerves flat, inconspic-
uous. Twigs terete, not ridged.] — Moluccas (Talaud Is.), possibly Sulawesi ....
Bers test: Aanueee bad Bas. 0 tu. Soe 88. H. talaudensis

menmtecismmallen. L113; Ca VOME, vcscisa.0sGre cor ensieie the te eRe PP Ree 10
10a.

Fruits subglobose. Twigs + flattened, usually lined or low-ridged. Leaves 12—25
cm long, membranous; nerves flat, inconspicuous. Male buds + pear-shaped, cleft
fomemes == Moluecasius sn llioiias gs ose oR ee 20. H. decalvata

. Fruits short-ellipsoid. Twigs terete, not lined. Leaves 5—14 cm long, chartaceous,

nerves inconspicuous on both surfaces. Male buds + obtriangular, cleft about half-
Meee atlippines: (Luzon) ie. 4 jo. es LEE. b 49. H. obscurinervia

. Twigs angular or ridged. [Species distinctive only in male flowering specimens. |

ree es ileal CIARA UM REIS LS OE dao, EA ae ENE Pr Se Loti LS 2

. Twigs (sub)terete or sometimes faintly angular, or shallowly lined ......... 13
. SW New Guinea, possibly Aru and Tanimbar Is. [Female flowers and fruits not

| SIRT pkgs eA RES RAPA rae ote oe Se etna as ARR Das Cee ks a 7. H. aruana

. Moluccas (Seram, Banda, Dammar I., possibly Ternate) ........ 77. H. smithii
Philippines: (Ovary glabrous oralmost so.) ............--:- 6. H. ardisiifolia
. Bark of twigs pale, grey-brown, contrasting with the blackish petioles. [Fruits black-

ismondrying; I.5—2,.cm long.|=— Moluccas ....2.25. 05.206. ee. 79. H. spicata

. Twigs brown, in colour not contrasting with the petioles ................. 14
. Fruits + globose to subellipsoid, 0.9—1.2 cm long (to 2 cm in New Guinea); black-

ish on drying. — Aru Is., New Guinea .......... 84a. H. subtilis var. subtilis

. Fruits 1.1-1.6 cm long (fruits not known in H. samarensis) .............. 5
MMU PIMESM SAMAR)! 5.2 pues : Seuot SM S F ok) steele ays b ESE 72. H. samarensis
BP OMICCASOUIAWESL ceo uc onic hae oes os eee rath ese sce ee nee ee 16
. Fruits ellipsoid, 1.5 cm long; blackish on drying. — Moluccas (Morotai, Obi Is.)

sco 5 SiO AE eae Pe 44a. H. moluccana var. moluccana

. Fruits subglobose or ellipsoid, 1.1—1.6(—2) cm long, brown on drying. — Sulawesi

akeabaena: Ts.)s. MoOluCCAaS (Sram): secs cuoe, as2ya <reneen or @ Shee oe 56. H. parviflora

(6) REGIONAL KEY TO THE SPECIES — NEW GUINEA
(based on female flowering and fruiting specimens)

. Leaves membranous, often with irregular whitish blotches. Perianth 2-lobed; ovary

glabrous. Fruits glabrous, globose, 1.5—2 cm diam.; pericarp 1-2 mm thick; seeds
globose. — Riverine or marshy, usually not far from the coast ..... 35. H. irya

. Leaves of different consistency, generally without whitish blotches. Fruits glabrous

or pubescent, globose or ellipsoid; if globose either only 1 cm diam. (H. subtilis),
or the pericarp more than 2 mm thick, at least at one side; seeds mostly ellipsoid.
SE MASEL OT MOL: sedan ss (Ree VA chee che, cP erties ch eset, one era Sie (ca EER 2
ihviss.ancled or ndged.— Aru Is., New Guinedy...¢ 45 ..5. so thi 0a. eee ae 3

nor nmdged. —— New, Guinea, to Solomomls., 3 0: 6 cy tncye ees es aeietes deta eeutt =e 8

Flora Malesiana, Ser. I, Vol. 14 (2000)

4a.

Sa.

6a.

10a.

4. Leaves with dots beneath (lens!). Perianth 2-lobed. — New Guinea (Bird’s Head to

WiSepiProw Psa Oe Ss LR se he TER 33. H. inflexa
‘Leavesuwithoutdots beneath” Stiri ions 5s as Sad I, cr 4
Pesianth 6-@Or 42 Nlobed! Scc adits a. 6 be weal elecrces ooo eo om Sa 3)
. Perianth 2-lobed) ... . ccs cbs ccs oc ces GHEN Seo 4.) 11. 6
Ovary glabrous (?). Fruits 10-16 mm long, glabrous. Leaves chartaceous, 7-14 cm
long; petiole comparatively long and slender, 11-20 mm long. — SW & S New
Guineai(Dieul, Westem Prov) ok on ee ee es OE 51. H. olens
. Ovary pubescent. Fruits 17-20 mm long, pubescent at base. Leaves membranous
or thinly chartaceous, 10-27 cm long; petiole 7-15 mm. — Papua Barat (Bird’s
Head) ates PEvature 6 ee Pee Hews Bois omens | )-nteiggegs 5. H. angularis
Flower buds depressed-globose, lobes nearly | mm thick; ovary pubescent. Fruits
17-20 mm long, pubescent. Leaves membranous or thinly chartaceous. — Papua
Barat(Birdis.Head) es see ei” soe etotnes ao-sare 5. H. angularis
. Female flowers and fruits not known ............ 00 e eee eee eee eee eee 7

a. Leaves membranous. — SW New Guinea; possibly Aru and Tanimbar Is........

je We fer lise dnc Didier eee tote. heretic tebe prea 7. H. aruana
_ Leaves thinly coriaceous. — SW New Guinea (a species close to H.. G7UANA) caso
eA deed: eaei ord beter ctabest aei deem ef) eer Sora: «Rie 34. H. iriana
_ Perianth 3- (or 4-)lobed, [cleft almost to the base.] — Papua New Guinea (East
S Epike PLOV. )s es -feyeysy ssi Cece a hg ce recat ect kchoaae aie See 74. H. sepikensis
aPerianth.2-lobed | vais levis atectieeeeneet tone ho ae se Riche ee 2
Ovary and frmitslabrous, «22. 2 ee ee 4 1: ee 10
. Ovary and fruits pubescent. [Hairs on fruits either distinct or small and inconspicu-
ous and usually only to be seen at the base of the fruits near the insertion of the
stalk (lens!). Fruits ellipsoid, sometimes globose in H. sinclairii.] ........- 16
Leaf bud, twig apex, and inflorescences with hairs 0.3—1.5 mm long. Leaves 17—45
cm long, often + parallel-sided; nerves 30—40 pairs. [Fruits ellipsoid, 3.4—-5.5 cm
long, glabrous.] — Moluccas and W & C New Guinea. 2 = 87. H. sylvestris
_ Leaf bud, twig apex, and inflorescences with hairs 0.2 mm long or less; hairs in H.
moluccana and H. tuberculata 0.1—0.3 mm long. Leaves generally smaller; nerves
FEWEL 4c. o0 bt 6 4 Sikora ete SE ee ee he eS Ce aecs ee 11
. Fruits globose or subglobose, [not beaked and without pseudostalk], 1.4 cm diam.
OE LESS fe re ee coe ONE neste cee ese ee oes a o.oo 12

b. Fruits ellipsoid, 1.3 cm long or more .........-+ ++ eee e eter eects 13

_ Fruits brown on drying; pericarp 1.5-3 mm thick. Buds cleft nearly to the base. —
Northern parts of Papua Barat and Papua New Guinea ........ 9. H. basifissa

b. Fruits blackish on drying; pericarp | mm thick. Buds cleft c. 1/3. — Aru Is., whole

GE New GUINEA: ao 6 cs <, cue oe are epee id aa 84a. H. subtilis var. subtilis
. Fruits to 2 cm long, blackish on drying; apex pointed/beaked or not, base without
or with long or short pseudostalk .........----+++-+-+ 0+ +++ 200s cues 14

b. Fruits 1.3—3.7 cm long, (dark) brown on drying; apex rounded, base mostly with-

Out pseudostalk "Sep sees ee es oe ihe)
- Pseudostalk of fruit (1.5—)2-6 mm long. — Papua Barat (Jayapura), Papua New
Guinier (W Sepik PIOV)) .. once oe eee a 73. H. schlechteri

b. Pseudostalk absent or up to 3 mm long. —Whole of New Guinea 84. H. subtilis

De Wilde — Myristicaceae (Horsfieldia) 83

15a.

BD:

23a:

24a.

Buds 2 mm long, cleft 1/2—4/5. Fruits 1.3—2.8 cm long, pericarp 1—2 mm thick. —
PieluccaseWw. New (Guinea! 2..20 VNerekh. Pee. 44. H. moluccana
Buds 2—3 mm long, cleft 1/2—2/3. Fruits 1.5—3.7 cm long, pericarp 1-8 mm thick.
— Papua New Guinea (Milne Bay Prov., Bismarck Archipelago, Papuan Islands)
ee er RE UA BELL TSE So a), oe 2G Se & 93. H. tuberculata

. Leaf bud, twig apex, and inflorescences with hairs 0.2 mm long or less (0.1—0.3

MaMa ALI EPP SELANING) FSA LL Sek SES Ne AOI. 17

. Leaf bud, twig apex, and inflorescences with hairs 0.5—1 mm long (0.2—0.5 mm

long in H. ampliformis; indumentum not known in H. ampla) ............ 21

. Hairs 0.1-0.3 mm long. Infructescences and female inflorescences large, much

branched, 10-16 cm long. Fruits 1.7—2.2 cm long, pericarp 1-2 mm thick. — Papua

New Guinea (Bismarck Archipelago, Bagabag I., Long I.) ....62. H. psilantha
. Hairs 0.1—0.2 mm long, or less. Infructescences and female inflorescences 10 cm
OMMIAGOfMESS-sFTULES VanLADIC ..2)..0. cote oveteedsrs cs sareo-s>snscoics SM Meee: © Reece ee 18
. Fruits 1.6 cm long or less; pericarp 1-3 mm thick. Buds pubescent ........ 12)
. Fruits 1.5 cm long or more; pericarp 2 mm thick or more; if fruit 1.5 cm long, then
MMOS GODOSE ANG. DUS G1abCOUS ait accu: cue pines aloe murat mele ae Guede © eyes 24
. Fruits at apex rounded, not apiculate; pseudostalk absent ...... 60. H. pilifera
DEaicsrapiculate, pseudostalk to'S mim Tomees. 6). 42) 0es eee oe ae ela, « 20

. Leaves broadly obovate to oblong, 12—20 by 5—11 cm. Pedicel widening to above

and gradually passing into the bud. Fruits (including | mm long pseudostalk and
2 mm long apiculum) 1.4 by 0.8—0.9 cm, fruiting pedicel 9-14 mm long, distinctly
Mae RLU MME wo ca hassle stele Sih a's “afiaseratahate olen Mowat eeente 19. H. crux-melitensis

. Leaves elliptic to lanceolate, 4.5—20 by 0.7—6 cm. Pedicel clearly marked off from

the bud. Pseudostalk of fruits 1.5—5 mm; fruiting pedicel up to 10 mm long, not or

PUMIMeMADCTING: <.y.cceccone ME A BODINE Ue ae MARS Sees AEE te DA|
. Pseudostalk of fruits 5 mm long. [Fruits excluding pseudostalk but including 2—3
mm long apiculum 1.6—1.7 by 0.9-I1cm.] .............. $1. H. squamulosa
MEScUaOstalk Of fruits l).5—S main LONG se. hp certo ena Beene aa eat hE 22

. Leaf bud, twig apex, and inflorescences with hairs 0.3 mm. Tertiary venation of

leaves below coarse and distinct. [Female flowers not known; fruits not known
with certainty; male buds subglobose, with thick perianth.] .. 94. H. urceolata

. Leaf bud, twig apex, and inflorescences with hairs 0.1—0.2 mm. Tertiary venation

Penetally less distinct. ([Malejpertanth: ditieremts)) 2 soa ae ss el hereto ae 23
Fruits excluding the 1.5—2.5 mm long pseudostalk, but including the 0.5—2 mm
long apiculum, 1.2—1.5 by 0.8—1 cm. Male buds clearly marked off from the pedi-
ell Aaah ALE pa CURR a eR eR oat ue 16. H. coryandra

. Fruits excluding the 1.5—2 mm long pseudostalk, but including the 2 mm long apic-

ulum, 1.3 by | cm long. Male pedicel broadening to above and gradually passing
HGNC MNT se Pah Ts eal, LOLS PACT EY, SNL ae 13. H. clavata
Buds 2—2.4 mm long, glabrous. Leaves 6—14 cm long. [Fruits (sub)globose, short-
ellipsoid, or obovoid, 1.5—2.5 by 1.5—2 cm; pericarp 4—6 mm thick.] — E Papua
RE WMG fi ity, WAMU Ee O8 Tine, beets Cae See kes, Engel 76. H. sinclairii

. Buds 2.5 mm long or more, pubescent or glabrescent. Leaves 10 cm long or more

OE VMN Ss SONI OEE ehaiy ae dob ee ale academe aman ott SENS shd SABI ce, 25

84 Flora Malesiana, Ser. I, Vol. 14 (2000)

25a. Fruits 1.6-3 cm long, usually with coarse pale wart-like lenticels; (dry) pericarp
2-6 mm thick (sometimes much resembling small-fruited H. pachycarpa). —
Moluccas, whole of New Guinea including Bismarck Archipelago; 0-1000 m....
Pebarsgetes B) icc he ee ee NS, nh eh) ee 37. H. laevigata

b. Fruits (3—)3.5—7.5 cm long; pericarp (4—)5 mm thick or more. — New Guinea;
(450) 1000=2000 mat eee Miiete See os a ee ee PGS Se 26

26a. Buds pubescent. Fruits 3—4.5 cm long; pericarp 4-10 mm thick a: Seekers
ey apaad leet 22 Re sities eroereeenie lind Lecae stogae 53. H. pachycarpa

b. Buds glabrescent. Fruits 6—7.5 cm long; pericarp 10-20 mm thick 2...
with al oleae ita: Wine seein es . eee Sees 15. H. corrugata

27a. Leaf bud and inflorescences with hairs 0.2—0.5 mm long. Fruits not known. [Leaf

bud. female flowers, and fruits not known in H. ampla.| .........+.+++++: 28

b. Leaf bud and inflorescences with hairs 0.5—1(-1.5) mm long. Fruits usually con-
spicuously pubescent ........c.. 62. aoe. SE el EN 29

28a. Inflorescences glabrescent. — Papua New Guinea (Sepik Prov.) .. 1. H. ampla

b. Inflorescences pubescent. — Papua New Guinea (Sepik and Morobe Prov.)......
otal ban Goethe he PRE eee eee) CUO italia ot See 2. H. ampliformis

29a. Leaves coriaceous, beneath with harsh hairs, when shed leaving thickened scars.
Buds 4 mm long, opening with narrow pore-like slit. [Fruits 3-5 cm long, pericarp

A Temimuticke| News Guincaye teem: aris ele cee re 64. H. pulverulenta

b. Leaves membranous or chartaceous. Buds cleft 1/4-1/2 ............++.-: 30
30a. Flowers [only the male known] entirely pubescent. [Fruits 2—2.4 cm long, pericarp
Ae momtnicke | — New Guinea Wit. sats +l.) 2 «oye sre oe 40. H. leptantha

b: Flowers largely glabrescent)... 3.2.2 0265-1. 5.6 ee ee ak
31a. Leaves generally oblong-lanceolate, at apex caudate. Buds 4 mm long. Fruits 2.5—
3 cm long. — Papua New Guinea (New Britain) ........... 67. H. ralunensis

b. Leaves oblong(-lanceolate), at apex not caudate (always?). Buds 3 mm long. Fruits
|.2-2.8 cm long. — Most of Papua New Guinea (incl. New Britain and New Ire-
(cl) eae een Emer teOMEni ye UPMREEE sours a ome mro Lo 2 em» 31. H. hellwigii

1. Horsfieldia ampla Markgr.

Horsfieldia ampla Markgr., Bot. Jahrb. Syst. 67 (1935) 148; W.J. de Wilde, Gard. Bull. Sing. 38, 1
(1985) 95. — Type: Ledermann 9639 (B, lost), Papua New Guinea, Sepik Prov.

Small tree 4—5 m. Twigs terete. Leaves cuneate-obovate, up to 40 by 16 cm, base
+ attenuate, apex short-acuminate; nerves 16-18 pairs, straight, sharply raised beneath
and connected towards the margin; petiole 1 cm long. Inflorescences on the older wood,
to 25 by 10 cm, glabrescent, loosely flowered. Male flowers yellow, clavate, 4 by 2 mm
(excluding pedicel?), the perianth 2-lobed, cleft to hardly 1/4. Staminal column thick;
thecae to c. 20, the androphore about as long as the anthers or slightly shorter.

Distribution — Malesia: NE Papua New Guinea (Sepik Prov., ‘Aprilfluss’; mountain
slope near camp 18).

Habitat & Ecology — Dense, very humid forest, on mountain slope at 200—400 m
altitude; male fl. Nov. 1912.

De Wilde — Myristicaceae (Horsfieldia) 85

Notes — 1. Known only from the type, now destroyed. Keyed out by Markgraf with
H. crux-melitensis, both having clavate flowers. He mentions in the key that the perianth
(not the androecium) is largely hollow, and in a note that the species is peculiar amongst
the New Guinea Horsfieldias because of its large male flowers, which in other species
are smaller and almost always broader than long, and that it is without close relatives.

2. The species is possibly related to or may be identical with H. ampliformis.

2. Horsfieldia ampliformis W. J. de Wilde

Horsfieldia ampliformis W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 95, f. 14. — Type: Hoogland
& Craven 11085 (male fl.), New Guinea, Sepik Prov.

Tree 5—8 m. 7wigs when young narrowly ridged, 4~7(—10) mm diameter, early or late
glabrescent, with + woolly hairs 0.2—0.5 mm long; bark coarsely striate, lenticellate, not
flaking. Leaves thickly membranous, (elliptic-)oblong, (18—)25—38 by (6—)7—13 cm, base
short to long-attenuate, apex attenuate-acuminate; upper surface drying dark brown, lower
surface early or late glabrescent or with rather scattered stellate hairs 0.2—0.5 mm long;
dots absent; midrib above + narrow, flattish; nerves 18—22 pairs, above thin, flat or sunken;
lines of interarching beneath not very prominent; venation lax, indistinct on both sur-
faces; petiole 4-6 by 3—4 mm; leaf bud 25—40 mm long, with hairs 0.2—0.5 mm. Jnflo-
rescences below the leaves; in male: many-flowered, 4 to 5 times branched, 25—35 by
20-30 cm, peduncle 4—5 cm; in female: 9-10 by 6—8 cm; all branches with rather loose
hairs 0.2—0.5 mm long; bracts (seen only in female) 5 mm long, caducous. Flowers 2—5
together in male and female, with loose hairs (0.1—)0.2—0.3 mm long, in female glabrescent
towards apex; perianth 2-lobed; pedicel not articulated. Male flowers: pedicel 2-4 mm
long, buds largely hollow, broadly obovoid, laterally + flattened, 3—3.3 by 3—3.2 mm,
apex obtuse to broadly rounded, base shortly tapering, cleft c. 1/3, lobes 0.3 mm thick;
androecium small, + flattened, 2.5 by 1—1.2 mm, apex broadly rounded, synandrium
1.5-1.8 by 1-1.2 mm, narrowly hollowed for 1/5—1/3 at apex (Plate 1: 20); thecae 14,
free apices 0.1—0.2 mm; androphore 0.8—1 by 0.5—0.6 mm. Female flowers: pedicel 1—2
mm long, buds broadly ovoid, 3 by 2.6—2.8 mm, cleft c. 2/3; ovary broadly ovoid, 2—2.2
by 1.8—2 mm, densely pubescent with hairs 0.1 mm or less, stigma short, not or hardly
lobed, 0.1 by 0.4 mm. Fruits not seen. — Fig. 8.

Field-notes — Small tree, 8 m high. Flowers medium green, yellow at anthesis.

Distribution — Malesia: northern Papua New Guinea (Sepik, Morobe Prov.).

Habitat & Ecology — Lower montane rain forest; 1200-1300 m altitude; fl. Apr.,
Aug.

Notes — 1. Horsfieldia ampliformis is close to H. ampla, of which no material has
been seen. According to the description the latter differs by the more elongate, possibly
glabrous perianths, 4 by 2 mm, the androecium with 20 thecae, and the glabrescent in-
florescences. The two species have a peculiar long-stalked androecium and larg, male
inflorescences. Horsfieldia ampla was collected at only 200—400 m altitude.

2. Known only from a male and a female flowering specimen. The perianths of the
female specimen, Craven & Schodde 1463 (Morobe Prov.), are glabrescent in the upper
half; it could be H. ampla. Moreover, as the hairs on the leaf buds are slightly shorter
than those of the male specimen, this specimen is difficult to distinguish from the vari-
able and widespread H. laevigata.

Flora Malesiana, Ser. I, Vol. 14 (2000)

86

De Wilde — Myristicaceae (Horsfieldia) 87

3. Horsfieldia amplomontana W.J. de Wilde

Horsfieldia amplomontana W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 34; Tree Fl. Sabah &
Sarawak 3 (2000) 361. — Type: Clemens 30536, Sabah.

Tree 10-20 m. Twigs 3.5—6(—10) mm diameter, early to rather late glabrescent, hairs
0.3—1(—1.5) mm long; bark coarsely striate, not flaking, lenticels small, not contrasting
in colour and inconspicuous. Leaves membranous to chartaceous, elliptic-oblong to ob-
long-lanceolate, 15—35 by 5—11 cm, base short-attenuate to narrowly rounded, apex acute-
acuminate; upper surface glabrescent, except towards the base of the midrib in young
leaves, olivaceous to brown, lower surface glabrous (glabrescent), without dots or hair
scars, Sometimes pale and contrasting with upper surface; midrib slender, + raised above,
nerves 11 or 12 pairs, above flattish or raised, lines of interarching not distinct; venation
lax, + distinct or not; petiole 8—15 by 2.5—3.5 mm, glabrescent; leaf bud 15—22 by 3-4
mm, with dense rusty hairs 0.5—1(—1.5) mm long. /nflorescences behind the leaves, with
shaggy rusty hairs 0.5 mm; in male: 3 or 4 times branched, many-flowered, 10-21 by
10-16 cm, peduncle 1.5—4 cm long, in male in loose clusters of 5—10; in female (in
fruit): 7-8 cm long; bracts broadly ovate-ellipsoid, 2-7 mm long, finely pubescent, cadu-
cous; flowers glabrous, perianth 3- (or 4-)lobed, pedicel sometimes + articulated (see
note 2). Male flowers: pedicel 0.8—1.5(—2) mm long, buds (depressed-)globose, 1.5—2 by
2—2.3 mm, apex and base (broadly) rounded, glabrous, cleft 1/2—2/3, lobes 0.2 mm thick,
not or only slightly collapsing on drying; androecium depressed-globose, 0.6—1 by 1.1-
1.8 mm, apex broadly rounded, base rounded or sagged (Plate 3: 79); thecae 20-26, al-
most completely sessile, 0.8—1.2 mm long, free apices up to 0.1 mm, incurved, conceal-
ing a+ 3-radiate apical slit or cavity 0.2—0.5 mm deep; column broad, solid; androphore
rather narrow, (0.1—)0.2—0.4 mm long, completely or partly hidden by the anthers. Fe-
male flowers (from fruit): 3-lobed, 3 mm long. Fruits 1-3 per infructescence, ellipsoid,
apex and base rounded, 7-8 by 4.5—5 cm, glabrous, drying dark brown, finely to coarsely
tubercled, pericarp 15 mm thick; fruiting pedicel 3 mm long; perianth persistent.

Field-notes — Large tree. Bark grey, fissured; outer bark soft, 5 mm thick; inner bark
white, soft, 5 mm; cambium pale; sapwood white; exudate from bark sticky. Flowers
golden. Ripe fruits orange.

Distribution — Malesia: Borneo (Sabah: Mt Kinabalu).

Habitat & Ecology — Primary and degraded forest, ridge forest; on sandstone; 1000—
1500 m altitude; fl. Nov., Dec., Feb.; fr. Nov.

Notes — |. Close to H. montana, both having very similar male flowers, but H. am-
plomontana differs considerably by its stouter twigs, larger leaves, larger male inflores-
cences, and very much larger fruits with a thick pericarp; in H. montana the fruit is only
2—2.7 cm long, and the perianth is not persistent.

2. The pedicels are generally not articulated, although some flowers of SAN 18843
seem to have an articulation, but this may be an artefact caused by drying.

Fig. 8. Horsfieldia ampliformis W.J. de Wilde. a. Twig apex with leaves; b. twig with male inflores-
cence axillary to fallen leaf; c. mature male flower, perianth opened, showing androecium; d. twig
with female inflorescence; e. female flower, opened, showing finely pubescent ovary and minute
2-lobed stigma [a—c: Hoogland & Craven 11085; d, e: Craven & Schodde 1463). — Scale bar for
fb —=2.cm: for cae= 1-7 mim;

88

Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 9. Horsfie

b. mature male flower; c. ditto, longitudina
tudinal section, schematic; e. twig with infru

Idia androphora W.J. de Wilde. a. Branch with leafy twig and male inflorescence,
lly opened, showing androecium; d. androecium, longi-

ctescence, fruits mature, aril complete [a—d: Noote-
boom & Chai 01710, type; e: Sinclair, Kadim & Kapis 8977]. — Scale bar for a, e = 2 cm; for b—d
= 0.85 mm.

De Wilde — Myristicaceae (Horsfieldia) 89

4. Horsfieldia androphora W. J. de Wilde

Horsfieldia androphora W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 32, f. 30; Tree Fl. Sabah &
Sarawak 3 (2000) 362. — Type: Nooteboom & Chai 01710, Sarawak.

Tree 7-20 m. Twigs 2-4(—5) mm diameter, with rusty hairs 0.3-0.6 mm, rather late
glabrescent; bark blackish brown, finely striate, not flaking; lenticels small, inconspicu-
ous. Leaves membranous, elliptic to oblong, 9-18 by 3.5—6.5 cm, base attenuate, apex
acute-acuminate; upper surface drying dark brown or blackish brown, glabrous, lower
surface glabrescent (except midrib); dots absent; midrib above raised, beneath with some
vestigial indumentum or late glabrescent; nerves 9—13 pairs, raised above, lines of inter-
arching on the lower surface irregular and not very conspicuous; venation lax, distinct or
not; petiole 10-12 by 1.5—2 mm, glabrescent; leaf bud 8—12 by 2—3 mm, with rusty hairs
0.3-0.6 mm. Inflorescences with + dense hairs 0.2—0.6 mm; in male: rather many-flow-
ered, 3 (or 4) times branched, 6-14 by 3.5—9 cm, peduncle 1—2 cm long; in female (from
infructescences): 3—4 cm long; bracts densely short-pubescent, ovate-elliptic,
+ acute, 3 mm long, caducous; flowers (male) in clusters of 2—6 each, perianth 3-lobed,
glabrous, pedicel glabrous or with a few minute hairs 0.1 mm at the very base, not ar-
ticulated. Male flowers: pedicel 0.5—2 mm, slender; buds globose, 1.4—2(—2.2) mm
diameter, cleft 1/3 to nearly 1/2, lobes 0.2 mm thick; synandrium depressed-globose,
somewhat flattened or impressed at apex and/or base, in cross section rounded, (0.6—)
0.8—1 by (0.8—-)1-1.3 mm, androphore slender, (0.3—)0.4—0.8 mm long (Plate 3: 78);
thecae 14-22, almost completely connate, incurved, concealing the apical hollow,
0.2-0.3 mm deep. Female flowers not seen. Fruits 2-5 per infructescence, ellipsoid,
2.4-3 by 1.4-2 cm, apex subacute to rounded, base rounded or shortly narrowed, gla-
brous, drying dark brown, finely tuberculate, without lenticels, pericarp 1.5—2 mm thick;
fruiting pedicel 2 mm long; perianth not persistent. — Fig. 9.

Field-notes — Bark chocolate to reddish brown, narrowly cracked, longitudinally
furrowed, or cut into rectangular blocks; sap watery, more or less colourless (tree in
flower), or blood red (tree in fruit). Twigs chocolate, with rusty hairs. Flowers yellow.
Fruits smooth, orange, testa whitish grey.

Distribution — Malesia: Borneo (Sarawak, Sabah).

Habitat & Ecology — Montane forest, mossy forest, wooded sandstone ridges, 800—
1200 m altitude; fl. Mar., Oct.; fr. Mar., June.

Notes — 1. Regarding the general morphology of the androecium, the present species
belongs to the H. grandis group. It seems closest to H. tomentosa from S Thailand and
Peninsular Malaysia, the two having a long-stalked synandrium in common. Horsfieldia
tomentosa has generally larger flowers, a pubescent lower leaf surface, and smaller fruits
which are pubescent or glabrescent. Horsfieldia androphora grows in mountains; H. to-
mentosa is restricted to lowlands.

2. Horsfieldia androphora keys out beside H. fragillima (also with non-articulated
pedicels), but the latter species differs in many characters including general habit and
fruit size; its saucer-shaped androecium is quite different.

5. Horsfieldia angularis W. J. de Wilde

Horsfieldia angularis W. J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 97. — Type: BW 5828 (male
fl.), New Guinea, Bird’s Head.

90 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 15-30 m. Twigs 2-angular, lower down subterete, with two ridges, 3—7(—10) mm
diameter, early glabrescent, hairs grey-brown, 0.1 mm or less; bark striate, distinctly
coarsely lenticellate, not flaking. Leaves membranous to thinly chartaceous, oblong
(-lanceolate), 10-27 by 3-7.5 cm, base attenuate, apex acute-acuminate; upper surface
drying pale to dark brown, often finely paler pustulate, lower surface glabrescent, hairs
very minute, grey, stellate, less than 0.1 mm; dots absent; midrib slightly raised above;
nerves 12-15 pairs, above thin and flattish or slightly sunken, lines of interarching not
distinct; venation lax, faint; petiole 7-15 by 2-3 mm; leaf bud 10-15 by 2—2.5 mm, with
hairs 0.1 mm. Inflorescences with rather dense hairs 0.2—0.3 mm; in male and female:
2 or 3 times branched, rather few-flowered, 3—4 by 2—2.5 cm, peduncle 0.3—0.6 cm long;
bracts not seen, caducous; flowers (male) generally 2—4 together; perianth 2—4-lobed in
male, 2(—3)-lobed in female, in the lower half with hairs 0.1(—0.2) mm long; pedicel
pubescent, not articulated. Male flowers: pedicel not tapering, 1-2 mm long; buds in
lateral view circular or slightly transversely ellipsoid, not or only slightly laterally com-
pressed, not collapsing on drying, 1.7—2.3 by 2.2-3.2 mm, cleft c. 9/10, lobes (0.2—)0.3
mm thick; androecium slightly laterally flattened (in 3- or 4-lobed flowers about 3- or 4-
angular in cross section), above broadly rounded, 1.2—1.5 by 1.5—2.2 mm (Plate 1: 2/);
thecae 24 to c. 40 (in 4-lobed flowers), + erect, free parts at apex to 0.1 mm, central
column at apex narrowly hollowed for (1/3—)1/2; androphore absent, the androecium
+ broadly attached. Female flowers: pedicel 1—-1.5 mm long; buds depressed globose,
2.5 by 3-3.2 mm, cleft c. 3/4; ovary + depressed globose-ovoid, 1.2 by 1.5 mm, densely
short-pubescent, style and stigma minutely 2-lobed, 0.1 by 0.3 mm. Fruits 5—10 per
infructescence, short-ellipsoid, 1.7—2 by 1.4-1.7 cm, pubescent at very base, with coarse
paler-coloured lenticel-like tubercles; pericarp thick-woody, 3—5 mm thick; fruiting pedi-
cel 3-5 mm long; perianth not persistent.

Field-notes — Sometimes buttressed to 1 by 0.5 m; bark sometimes fissured, or peel-
ing off in small scales; with red exudate; sapwood pale brown or white; heartwood not
discernible or pinkish. Flowers greenish. Fruits yellow(-brown), sour and edible.

Distribution — Malesia: Papua Barat (Bird’s Head, subprov. Manokwari).

Habitat & Ecology — Primary forest; on clayey soils; locally common on the coastal
plain up to 600 m in Kebar Valley; 0-600 m altitude; fl. Feb., Aug.; fr. Feb., Oct.

Note — Much related to H. basifissa, of which sterile specimens are difficult to iden-
tify since their twigs too are rather ridged. Horsfieldia angularis is distinguished from
H. basifissa by 1) the more strongly ridged and somewhat stouter twigs, 2) the more
hairy and 2—4-lobed flowers with thicker lobes, 3) the hairy ovary and the thinly pubes-
cent ellipsoid fruits. Both species have thickish, subglobose male buds, which hardly
collapse on drying, and which at anthesis are cleft to the base.

6. Horsfieldia ardisiifolia (A. DC.) Warb.

Horsfieldia ardisiifolia (A. DC.) Warb., Mon. Myrist. (1897) 274; J. Sinclair, Gard. Bull. Sing. 28
(1975) 3: WJ. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 72, f. 9. — Myristica ardisiifolia A. DC.,
Ann. Sc. Nat. Bot. 4, 4 (1855) 31, t. 4; Prodr. 14, 1 (1856) 203, ‘ardisiaefolia’. — Type: Cuming
1702, Philippines.

Horsfieldia warburgiana Elmer, Leafl. Philipp. Bot. 3 (1911) 1061; Merr., Enum. Philipp. Flow. PI.
2 (1923) 183. — Type: Elmer 12297, Philippines.

Horsfieldia gigantifolia Elmer, Leafl. Philipp. Bot. 9 ( 1925) 3120, 3129; 10 (1939) 3763; nom. nud.

De Wilde — Myristicaceae (Horsfieldia) 9]

Tree 5—10 m. Twigs flattened 2-angular, lower down terete with two ridges, 3—6(—13)
mm diameter, at first with bright rusty hairs 0.3—0.5(—0.8) mm long, early glabrescent;
bark smooth to striate, distinctly lenticellate, not flaking. Leaves membranous, (ellip-
tic-)oblong, 20—40 by 5.5—15 cm, base nearly rounded to attenuate, apex acute-acumi-
nate; upper surface drying olivaceous to blackish brown, finely minutely paler pustulate
or not, lower surface early glabrescent except for some indumentum remaining on the
midrib, hairs coarse 0.3—0.5 mm; dots absent; midrib fairly broad, flattish above; nerves
18-28 pairs, slender above, flattish, lines of interarching regular and distinct beneath;
venation lax, inconspicuous; petiole 13-16 by 3—4.5 mm; leaf bud 10-20 by 3-4 mm,
with hairs 0.3—0.8 mm long. Inflorescences thinly with stellate-dendroid hairs 0.3 mm;
in male: 3 or 4 times branched, rather many-flowered, broadly pyramidal, 7-16 by 6—14
cm, peduncle 0.5—1(—2) cm long; in female: 4-8 cm long; bracts broadly ovate, pubes-
cent, 3-4 mm long, caducous; flowers (male) solitary or 2-4 together, perianth 2-lobed,
glabrous, pedicel sometimes at first with sparse hairs, slender, not articulated. Male flow-
ers: pedicel 1—2(—4) mm; buds transversely ellipsoid or reniform, moderately laterally
compressed, drying dull, more or less collapsed on drying or not, 2.5-3 by 4—4.5 mm,
below sometimes with a basal sinus, cleft 4/5—5/6, the lobes 0.2(—0.3) mm thick;
androecium broadly transversely ellipsoid, slightly laterally flattened, hollow, 1.5 by 3—
3.5 mm (Plate 1: 8); thecae (36—)40—48, connate for about halfway, forming a cup with
the anthers from one side deeply inflexed, those from the other side for a large part over-
arching the former; anthers sometimes slightly sagged at base, hiding the narrow andro-
phore, 0.2—0.3 mm long. Female flowers: pedicel 2(—2.5) mm long; buds subglobose-
ovoid, 2.5 mm diameter, cleft c. 1/2; the ovary broadly ovoid-subglobose, 1.5—1.7 mm
diameter, glabrous, stigma consisting of 2 minute sessile lobes 0.1—0.2 mm. Fruits 2—6
per infructescence, broadly ellipsoid to subglobose, 20—25 by 17-20 mm, glabrous (or
possibly with few minute hairs at base), finely rugulose, without marked tubercles, dry-
ing (reddish) brown; pericarp 1.5-2 mm thick; fruiting pedicel 3-6 mm long; perianth
not persistent. — Fig. 10.

Field-notes — Flowers yellow, fragrant. Fruits orange-red.

Distribution — Malesia: Philippines (Luzon, Mindoro, Sibuyan, Samar, Leyte).

Habitat & Ecology — Lowland forest in moist valleys; O-400 m altitude; fl. & fr.
throughout the year.

Note — Horsfieldia ardisiifolia is close to species like H. parviflora and H. smithii,
both from the Moluccas, all of which have anthers strongly incurved or inflexed into the
androecium cup. Horsfieldia ardisiifolia is distinguished by thick winged or ridged twigs,
large leaves, coarse hairs on the leaf buds, male buds 4—4.5 mm wide, and a broad an-
droecium with the anthers deeply incurved and clasping each other.

7. Horsfieldia aruana (Blume) W. J. de Wilde

Horsfieldia aruana (Blume) W. J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 100. — [Palala aruana
Rumph., Herb. Amb. 7 (1755) t. 24.] — Myristica aruana Blume, Rumphia | (1837) 191;
J. Sinclair, Gard. Bull. Sing. 28 (1975) 112, 118, 119, 122-124, in the synonymy of Horsfieldia
spicata. — Horsfieldia novo-guineensis Warb., Mon. Myrist. (1897) 271, t. 23, nom. nov., p.p.,
for the lectotype only. — Lectotype: those specimens of Zippelius s. n. at L, annotated by Blume,
W New Guinea.

Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Horsfieldia) 93

Tree c. 15 m. Twigs 2-angled, becoming subterete with two ridges, 3—5 mm diameter,
early glabrescent, hairs c. 0.1 mm long; bark striate, not flaking; lenticels small, incon-
spicuous. Leaves membranous, elliptic-oblong, 15—29 by 5—9.5 cm, base attenuate, apex
acute-acuminate; upper surface drying olivaceous to brown, lower surface early
glabrescent; dots absent; midrib slightly raised above; nerves 13—15 pairs, slender, flat-
tish; venation lax, indistinct; petiole 10-15 by 1.5—2.5 mm; leaf bud c. 10 by 1.5 mm,
hairs c. 0.1 mm long. Inflorescences among or below the leaves, with sparse hairs c. 0.1
mm or less; in male: 3 or 4 times branched, 5—8 by 4—5 cm, rather many-flowered,
peduncle 0.5—1.5 cm long; bracts not seen, caducous; flowers (male) in loose clusters of
2-5 each, perianth 2-lobed, glabrous; pedicel slender, sparsely pubescent, not articu-
lated. Male flowers: pedicel 1—1.5 mm long; buds in lateral view circular to somewhat
transversely elliptic, laterally compressed, blackish and collapsing (always?) on drying,
1.5—2 by 2—2.5 mm, cleft 2/3—3/4, lobes c. 0.2 mm thick; androecium much compressed
laterally, c. 1.5 by 2 mm, above broadly truncate-rounded (Plate 1: 23); thecae 28-36,
distal free parts 0-0.1 mm, column almost completely solid; androphore to 0.1(—0.2)
mm. Female flowers and fruits not seen.

Distribution — Malesia: SW Papua Barat; possibly also Moluccas (Aru and Tanimbar
Islands, see note).

Habitat & Ecology — Not known.

Note — Specimens perhaps to be included in H. aruana are Buwalda 4969 trom the
Aru Is. and bb 24414 from the Tanimbar Is.; the male flowers of both are immature. The
synandrium of Buwalda 4969 is cleft to c. 1/10 only; however, in bb 2441/4 it appears
cleft nearly 1/4 or 1/5; the irregular whitish blotches on the leaves are similar to those
usually found in H. irya and H. smithii.

8. Horsfieldia atjehensis W.J. de Wilde

Horsfieldia atjehensis W.J. de Wilde, Gard. Bull. Sing. 38, 2 (‘1985’, 1986) 186. — Type: Bangham
882, Sumatra, N Aceh.
Horsfieldia amygdalina auct. non (Wall.) Warb.: Merr., J. Arnold Arbor. 8 (1934) 61.

Tree c. 10 m. Twigs terete, 3.5—5(—8) mm diameter, pale grey to yellowish brown,
early glabrescent, with grey-brown hairs less than 0.1 mm; bark coarsely striate and
tending to flake; lenticels rather conspicuous towards the apex of the twig. Leaves in 3—
5 rows, thinly chartaceous, obovate-oblong to oblong-lanceolate, 13-25 by 4.5—9 cm,
base long-attenuate, apex acute-acuminate; upper surface drying dark brown, lower sur-
face early glabrescent, with scattered dots (lens!); midrib flat above; nerves 10—12 pairs,
flat above, lines of interarching indistinct; venation lax, indistinct or invisible on both
surfaces; petiole 12-15 by 2.5-3.5 mm; leaf bud 15 by 3.5—4 mm, with dense grey-brown
hairs less than 0.1 mm. Inflorescences (female not seen) behind the leaves, glabrescent,

Fig. 10. Horsfieldia ardisiifolia (A.DC.) Warb. a. Leafy twig apex, note ridged twig; b. twig with
male inflorescence in axil of fallen leaf; c. mature male flower, lateral view; d. ditto, opened, show-
ing androecium; e. androecium, longitudinal section, schematic; f. mature female flower, lateral
view; g. ditto, opened, showing glabrous ovary with minute stigma; h. twig with infructescence
with ripe fruits [a: BS 39770 (Ramos), b: PNH 6236 (Sulit); c—e: Elmer 12337; f, g: Elmer 17220,
h: PNH 17461 (Conklin)]. — Scale bar for a, b, h = 2 cm; for c, d, f, g = 1.7 mm; for e = 0.85 mm.

94 Flora Malesiana, Ser. I, Vol. 14 (2000)

or thinly haired, less than 0.1 mm; in male about 3 times branched, rather many-flow-
ered, 7-14 by 4-10 cm, peduncle 1-2 cm long; bracts elliptic-oblong, 2-4 mm, finely
pubescent, caducous; flowers in male in loose clusters of 4-8 each, glabrous, perianth
3-lobed, pedicel not articulated. Male flowers (+ immature): pedicel slender, 1—-1.5 mm
long; buds globose, 1.5 mm diameter, cleft c. 1/2, lobes 0.2(—0.3) mm thick; androecium
subglobose to short-ellipsoid, 1.2 by 1 mm, apex broadly rounded, in cross section sub-
circular (Plate 2: 4/); thecae 22, almost completely sessile, free apices 0.1(—0.2) mm,
curved over and more or less into the rather narrow apical cavity 0.3 mm deep; column
broad, androphore narrow, 0.2(—0.3) mm long. Female flowers and fruits not seen.

Field-notes — Leaves leathery, glabrous. Flower buds green.

Distribution — Malesia: Known from only one collection in N Aceh, Sumatra.

Habitat & Ecology — Montane forest, possibly on limestone; 1200-1800 m altitude;
male fl. (immature) in Jan.

Note — Horsfieldia atjehensis is in many respects closely related to and + intermedi-
ate between H. amygdalina (Wall.) Warb. (from continental SE Asia), H. glabra, H.
macrothyrsa, and H. sparsa, but is still markedly distinct from these species (De Wilde,
l.c.: 188).

9. Horsfieldia basifissa W.J. de Wilde

Horsfieldia basifissa W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 109. — Type: NGF 10242 (White),
New Guinea.
Horsfieldia polyantha auct. non Warb.: J. Sinclair, Gard. Bull. Sing. 28 (1975) 95, p.p.

Tree 10-25 m. Twigs faintly ridged or not, 2—4(—-8) mm diameter, with grey-brown
hairs 0.1 mm, early glabrescent; bark finely striate, not flaking; lenticels inconspicuous.
Leaves membranous or thinly chartaceous, elliptic-oblong to oblong-lanceolate, 10-22
by 3-8 cm, base attenuate, apex acute-acuminate; upper surface drying olivaceous to
brown, often with paler markings, sometimes faintly pale pustulate, lower surface
glabrescent, hairs 0.1 mm; dots absent; midrib above flattish; nerves 10-15 pairs not
particularly contrasting, above thin, flattish or sunken, beneath with lines of interarching
neither regular nor prominent; venation lax, rather faint; petiole 5-10 by 1.5—2.5 mm;
leaf bud 10 by 1.5 mm, with hairs 0.1 mm. Inflorescences in male 3 (or 4) times branched,
many-flowered, 4-10 by 2.5—6 cm, peduncle 0.2—2 cm long; in female: 5 by 3.5 cm;
with dense to sparse stellate hairs 0.1-0.2 mm; bracts elliptic-oblong, acute, 1—2(-4)
mm long, caducous; flowers generally 1—3 together; perianth 2-lobed, glabrescent ex-
cept at the very base, pedicel with hairs 0.1 mm long, not articulated. Male flowers:
pedicel 1.5—-3 mm long; buds as seen laterally + circular, slightly broader than long,
slightly laterally compressed, not or but slightly collapsed on drying, 2.2—2.7 by 2.6-3
mm, cleft to the base, lobes 0.1-0.2 mm thick; androecium laterally much flattened,
above broadly rounded, 1.5—1.7 by 2 mm (Plate 1: 26); thecae 24—28(-32), erect, at apex
free for 0.1 mm long, column at apex narrowly hollowed for 1/3—2/3; androphore to 0.1
mm, broadly attached. Female flowers (immature): pedicel 1.5—2 mm long, buds broadly
ovoid, 1.5 by 1.4 mm, cleft nearly to the base; ovary ovoid, 1.1 by 0.6 mm, glabrous,
style and stigma minutely 2-lobed. Fruits 1-20 per infructescence, globose or subglobose,
1.1-1.4 cm diameter, glabrous, drying light to dark brown, with or without coarse, paler
coloured lenticels or warts; pericarp 1.5—3 mm thick, woody-granular; fruiting pedicel
3—4 mm; perianth not persistent.

De Wilde — Myristicaceae (Horsfieldia) 95

Field-notes — Slender tree, branches horizontal. Flowers yellow. Fruits green, turn-
ing orange.

Distribution — Malesia: New Guinea (NE Papua Barat, including Memberamo River
area; N Papua New Guinea: Sepik, Madang Prov.).

Habitat & Ecology — Primary and degraded forest, marshy forest, locally common,
recorded from Pometia—Intsia forest; on clays and marls; 0-200 m altitude; fl. Sept.;
fr. Mar., June, Oct.

Note — Apart from H. angularis (see the note under that species) H. basifissa 1s
possibly closely related to H. parviflora, both have glabrous fruits. The globose fruits
are often very similar to those of H. pilifera or H. sinclairii; in these two species, how-
ever, the fruits are always hairy, at least towards the base. Horsfieldia basifissa has
much in common with H. laevigata var. novobritannica, which also has the androecium
deeply hollowed inside; the latter has a more hairy perianth. The female flowers of var.
novobritannica are not known, but its globose fruits are larger than those of H. basifissa
and somewhat hairy at the base. Horsfieldia basifissa is characterized by the subglabrous
male flowers with a very deeply cleft perianth, glabrous ovary, and glabrous, globose
fruits.

10. Horsfieldia borneensis W. J. de Wilde

Horsfieldia borneensis W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 27; Blumea 32 (1987) 468;
Tree Fl. Sabah & Sarawak 3 (2000) 363. — Type: S 1461/0 (Bojang), Sarawak.

Tree 10—30 m. Twigs subterete, (1.5—)2—4(—10) mm diameter, sometimes blackish,
early to rather late glabrescent, hairs rusty, 0.2—0.4 mm; bark finely striate, not distinctly
lenticellate, sometimes finely cracking or slightly flaking. Leaves chartaceous to thinly
coriaceous, elliptic-oblong to oblong-lanceolate, 7-18 by 2—6 cm, base (short-)attenu-
ate, apex acute to (short-)acuminate; upper surface drying dull olivaceous to (partially)
blackish brown, glabrous, lower surface drying pale brown to chocolate, glabrescent,
hairs densely branched dendroid 0.3-0.4 mm (especially on midrib); dots present and
obvious; midrib slightly raised above, (late) glabrescent; nerves 10—16 pairs, slender
above, flat or sunken (slightly raised only close to the midrib) or in thinner-leaved speci-
mens slightly raised, glabrous, lines of interarching + regularly shaped, not distinct;
venation hardly or not visible on both surfaces; petiole 12-25 by 1.5—2.5 mm, some-
times late glabrescent; leaf bud 10-17 by 24 mm, with dense hairs 0.3 mm long. Inflo-
rescences behind the leaves, with dense short-woolly rusty hairs up to 0.7 mm long; in
male: fairly large, many-flowered, about 4 times branched, (8—)13—20 by (5—)10—-18 cm,
peduncle 1.5—3.5 cm long; in female: 8—10(—13) by 4—5 cm, less branched; bracts ellip-
tic to elliptic-oblong, pubescent as the inflorescences, 1.5—5 mm long, caducous; flow-
ers (male) in loose clusters of 2-6, glabrous; perianth 3-lobed, pedicel distinctly articu-
lated. Male flowers: pedicel 1—1.5 mm; buds subglobose to broadly ellipsoid or broadly
obovoid, 1.3—1.8 by 1.2-1.7 mm, cleft c. 1/3 (to nearly 1/2), not collapsing on drying,
lobes 0.2—0.3 mm thick; androecium broadly ellipsoid to subglobose, 0.7—1.2 by 0.6—
1.3 mm, the apex broadly rounded, slightly impressed in the centre with cavity to c. 1/4,
base rounded, in cross section circular (Plate 3: 76); thecae 16-20, almost completely
sessile and mutually closely appressed, at apex incurved over the cavity, free apices
about none; column broad; androphore narrow, at most 0.1 mm long. Female flowers:

96 Flora Malesiana, Ser. I, Vol. 14 (2000)

pedicel thickish, glabrous, 1-1.5 mm long, distinctly articulated; buds subglobose or
broadly ellipsoid, 3-3.5 by 3 mm, glabrous, cleft c. 1/3; ovary subglobose to broadly
ovoid-ellipsoid, 2(—2.5) by 2 mm, glabrous, stigma 2-lobed, 0.2 mm high, descending
1.5 mm down the ovary. Fruits 1-7 per infructescence, ovoid, 4—6 by 3—4.5 cm, gla-
brous, somewhat laterally flattened and slightly flanged, apex and base rounded, drying
brown and often with a glaucous tinge, smooth; pericarp 10-15 mm thick; fruiting pedi-
cel stout, 4-6 mm long; perianth not persistent.

Field-notes — Bark usually dark brown, reddish, or blackish, rough, deeply fissured,
flaking in squares, strips or flakes up to 5 cm wide, up to | cm thick (strips with rounded
edges, appearing smooth); living bark 5-10 mm thick, red-brown, the sap red; sapwood
10 cm, reddish white to pale red; heartwood red-brown. Fruits bluish green, turning green-
yellow to yellow or reddish, pericarp pink inside.

Distribution — Malesia: Borneo (Sarawak, Sabah, E & NE Kalimantan).

Habitat & Ecology — Primary lowland dipterocarp forest, swamp forest; on sandy
soils, flat clayey soil, sandstone, sandy ridges; 0-200 m altitude; fl. Apr., Aug., Sept.;
fr. throughout the year.

Notes — 1. Horsfieldia borneensis and H. wallichii both have characteristic blackish
dots on the lower leaf surface, a dull upper leaf surface with largely sunken nerves, and
similar fruits (although the perianth in H. wallichii is persistent). However, H. wallichii,
which also occurs in Borneo, is generally stouter and has much larger leaves, often with
a persistent indumentum. Above all, it differs in general appearance, shape, and struc-
ture of the androecium, and the pedicel which is not articulated.

2. Horsfieldia borneensis, with its dotted lower leaf surface, belongs to a group of
species including H. wallichii and H. pulcherrima, while the structure of its androecium
links it to species such as H. flocculosa, H. grandis, and H. pulcherrima; compare also
H. punctatifolia.

11. Horsfieldia brachiata (King) Warb.

Horsfieldia brachiata (King) Warb., Mon. Myrist. (1897) 325; Gamble, Mat. Fl. Malay Penins. 5,
23 (1912) 218: Ridl., Fl. Malay Penins. 3 (1924) 59; W.J. de Wilde, Gard. Bull. Sing. 39, 1
(1986) 3: Tree Fl. Sabah & Sarawak 3 (2000) 363. — Myristica brachiata King, Ann. Roy. Bot.
Gard. Calc. 3 (1891) 311, pl. 144. — Horsfieldia subglobosa (Miq.) Warb. var. brachiata (King)
J. Sinclair, Gard. Bull. Sing. 16 (1958) 431, f. SIE. — Horsfieldia brachiata (King) Warb. var.
brachiata: J. Sinclair, Gard. Bull. Sing. 28 (1975) 9. — Lectotype: Griffith 4351, Peninsular Ma-
laysia.

Tree 10-35 m. Twigs + angular or subterete, or more or less flattened, distinctly lined
or ridged (sometimes lines evident only in part of the material), 2—7(—18) mm diameter,
generally early glabrescent, hairs rusty, (0.1—)0.2-0.4 mm long; bark finely to coarsely
striate, not flaking; lenticels present but not much contrasting. Leaves membranous, el-
liptic-oblong to oblong(-lanceolate), 12—26(—30) by 4-9(-11) cm, base cuneate, apex
acute-acuminate; upper surface glabrous, drying olivaceous to brown or sometimes black-
ish, lower surface drying light brown, early glabrescent, midrib sometimes later gla-
brescent: dots absent; midrib raised above, glabrous; nerves 12-20 pairs, raised above,
lines of interarching usually not distinct; venation lax, usually not distinct above; petiole
8—13(—20) by 2-3 mm, glabrescent; leaf bud 8-15 by 3-4 mm, hairs 0.2—0.4 mm long.

De Wilde — Myristicaceae (Horsfieldia) 97

Inflorescences with sparse to dense dendroid hairs 0.2—0.5 mm, sometimes glabrescent;
in male: 3 or 4 times branched, many-flowered, 7—18(—22) by 5—16(—18) cm, peduncle
0.6—1.8 cm long, the flowers in loose clusters of 3—6; in female: many-flowered, 3—8 by
2-6 cm; bracts oblong-lanceolate, acutish, 3-5 mm long, pubescent, caducous; flowers
with perianth 3- (or 4-)lobed, glabrous, pedicel pubescent in various degrees, hairs 0.1—
0.2 mm long (in female glabrescent), articulated. Male flowers: pedicel (1—)1.5—2.5 mm
long; buds (depressed-)globose to broadly obovoid, in cross section rounded or slightly
angular, 1—1.5 by 1.2—1.8 mm, base rounded to short-cuneate, not or but little collapsing
on drying, cleft 1/2—2/3, lobes 0.2—0.4(—0.5) mm thick; androecium depressed-globose
to obovoid in outline, (0.5—)0.7—1 by 0.8—1.2 mm, + rounded or usually + triangular in
cross section (Plate 3: 64); thecae 12—20, anthers 0.5—0.7 mm long, mutually free for
about halfway, usually curved towards the centre, column largely hollowed out, at base
continuing into the 0.2—0.3 mm long androphore, slightly tapering or not. Female flow-
ers: pedicel 1-1.5 mm long; buds broadly ellipsoid, 2.2—2.5 by 1.8—2 mm, cleft c. 1/3;
ovary ovoid, 1—1.4 by 0.8—1.2 mm, glabrous, the stigma 2-lobed, 0.2 by 0.4 mm. Fruits
4—12(-20) per infructescence, broadly ellipsoid, apex narrowly rounded, base (broadly)
rounded, 2—2.8(—3) by 1.8—2.2(—2.6) cm (see note 3), glabrous, drying brown to dark
brown, neither warted nor lenticellate, pericarp 1.5—4 mm thick; fruiting pedicel 1.5—3
mm long; perianth not persistent.

Field-notes — Usually a slender tree with straight bole, once recorded as with but-
tresses to 50 cm high; bark + smooth, pale to dark brown, generally with shallow vertical
fissures | cm apart, sometimes + laminated, scaly, or cracked; living bark 8-10 mm
thick, pinkish to reddish brown, exuding reddish sap; wood whitish to pale brown; no
heartwood; twigs with raised lines. Flowers greenish yellow to dark yellow, scented.
Fruits yellow(-green) or yellow-orange.

Distribution — Peninsular Thailand; Malesia: Sumatra, Peninsular Malaysia (Kedah,
Kelantan, Perak, Trengganu, Pahang, Malacca, Johore), Borneo (Sarawak, including one
deviating collection, see note 3; Sabah; C, E & NE Kalimantan; Brunei); not found in
Singapore and large parts of Kalimantan.

Habitat & Ecology — Primary and degraded lowland rain forest; often near streams
in flatland; marshy, riverside, and peaty forests, forest on alluvial plains, poor forest on
soil with stagnant water, but also on hillsides; on alluvial soils, brown and sandy soil (in
Tristania forest, Sabah), sandstone, peaty soils, loam soil with lime; 0—400 m altitude;
fl. & fr. throughout the year.

Notes — |. Horsfieldia brachiata is close to H. polyspherula, and in most cases
easily recognized by its weak to strong raised lines on the twigs. Its fruits are rather uni-
form in shape and size, 20—28 mm long, and thus + intermediate between those of H.
polyspherula var. polyspherula and var. sumatrana (see there). The leaves of H. brachiata
are rather like those of H. polyspherula var. sumatrana, namely generally membranous
and drying pale, dull olivaceous above and pale cinnamon below. Its flowers are rather
uniform, with mature male buds 1.2—1.8 mm in diameter, and the androecium usually
+ triquetrous with 6-10 stamens (but see note 2), and they do not differ from those of
H. polyspherula s.\. Sterile and flowering collections in which the apical and lower twig
parts are not sufficiently represented may be difficult to place. Horsfieldia brachiata
generally has stouter inflorescences than H. polyspherula and is quite common in ever-
green forests in Peninsular Thailand; H. polyspherula has not been found there.

98

Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Horsfieldia) 99

2. The species usually has 12—16 thecae in the androecium, but material from Penin-
sular Thailand may have 18 or 20 thecae.

3. § 34908 from Sarawak (Kapit, 5th Div.) is a stout specimen, in bad condition, with
female flowers; at L there is a single fruit measuring 40 by 30 mm, with the pericarp
+ woody, 5—7 mm thick. Horsfieldia brachiata 1s not common in Sarawak and this large-
fruited specimen probably represents a separate taxon. It was collected in a kerangas-
mossy forest at c. 800 m altitude, higher than any other specimen of the species.

12. Horsfieldia carnosa Warb.

Horsfieldia carnosa Watb., Mon. Myrist. (1897) 348, 619; Merr., Enum. Born. (1921) 268; J. Sinclair,
Gard. Bull. Sing. 28 (1975) 21; W.J. de Wilde, Gard. Bull. Sing. 38, 2 (°1985’, 1986) 222, f. 26;
Tree Fl. Sabah & Sarawak 3 (2000) 364. — Myristica carnosa (Warb.) Boerl., Handl. 3 (1900)
87. — Lectotype: Beccari 1242 (FI acc. 7625), fr.; Sarawak.

Tree 4-10 m. Twigs 3—10(—16) mm diameter, early glabrescent, hairs grey-brown, 0.1
mm; bark coarsely or finely striate, tending to flake, drying somewhat pale, yellow-
brown or light grey-brown, generally contrasting with the blackish brown of the peti-
oles; lenticels usually not conspicuous. Leaves chartaceous-coriaceous, rarely + mem-
branous, (elliptic-)oblong, 13—35 by 5—11 cm, base long- or sometimes short-attenuate,
apex acute-acuminate; upper surface drying bright dark brown, finely wrinkled-granu-
late, glabrous, lower surface early glabrescent (glabrous); dots absent; midrib above
flat; nerves 13-18 pairs, flat above, lines of interarching indistinct; venation lax, faint or
invisible on both surfaces; petiole 10-16 by 2-4 mm, glabrous or early glabrescent; leaf
bud 9-13 by 2-3 mm, hairs grey-brown, 0.1 mm long. /nflorescences with dense to
sparse hairs 0.1 mm or less; in male: below the leaves, many-flowered, 3 or 4 times
branched, 6-17 by 5-14 cm, peduncle 1—3 cm long; in female: ramiflorous, rather many-
flowered, 1-2 cm long; bracts elliptic to oblong, 4-10 mm long, pubescent, caducous;
flowers glabrous, in male in loose clusters of 3—9, perianth 3-lobed, pedicel not articu-
lated. Male flowers: pedicel 1—1.5 mm long; buds (sub)globose, 1.9—2.1 by 1.8—2 mm,
cleft 1/3—1/2, not collapsing on drying, lobes 0.2 mm thick; androecium (sub)globose,
1-1.2 by 1-1.3 mm, circular in cross section (Plate 2: 6/); thecae 18—22, completely
sessile, without free apices, incurved, concealing a small apical cavity 0.2—0.4 mm deep;
column broad, + spongy, androphore rather narrow, 0.2—0.5 mm long, completely hid-
den by the anthers. Female flowers: pedicel 1—1.5 mm long; buds ellipsoid, 3.5 by 2.5
mm, cleft c. 1/3, lobes 0.3—-0.4 mm thick; ovary ellipsoid, 2 by 1.5 mm, glabrous, stig-
ma consisting of two broad lips 0.2 mm high. Fruits 2-11 per infructescence, ellipsoid,
1.6—2 by 1.2-1.5 cm, glabrous, drying brown, the surface finely granulated; pericarp
1.5 mm thick; fruiting pedicel 1-2 mm long; perianth not persistent. — Fig. 11.

Fig. 11. Horsfieldia carnosa Warb. a. Twig with leaf and male inflorescences; b. apical part of leafy
twig; c. mature male flower, lateral view; d. ditto, opened, showing androecium; e. androecium,
longitudinal section, schematic; f. twig with female inflorescence axillary to leaf scar; g. female
flower at anthesis, lateral view; h. ditto, longitudinally opened, showing glabrous ovary with broad
2-lobed stigmas; 1. older twig with infructescences, fruits mature, aril complete but torn on drying
[a: van Niel 5419; b—e: S 18011; fh: SAN 63191; i: SAN 17438]. — Scale bar for a, b, f, 1 = 2 cm:
for c—e, g,h = 0.85 mm.

100 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Small tree, trunk slender; the bark often flaking or shallowly fissured;
inner bark yellow, thin, sap watery, clear, not reddish; sapwood whitish, twigs light brown.
Flowers green-yellow, anthers whitish. Fruits (immature) greenish yellow, aril orange.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah; W Kalimantan: Mt Klam).

Habitat & Ecology — Heath forest, wet kerangas, peat swamp forest, Agathis—
Casuarina forest; on white sandy soils; 0-100 m altitude; fl. mainly July—Nov.; fr. through-
out the year. An extensive note on the ecology is given by Sinclair, l.c.

Note — Horsfsieldia carnosa is a well-characterized species, a small tree of kerangas
or peat swamp forest, on white sandy soils. It is distantly related to H. glabra, which is
distinguished by a less stout habit, dark twigs, bark not tending to flake, smaller and
usually membranous leaves, globose male flowers, pedicels + articulated, globose or
ellipsoid androecium with short androphore, and somewhat longer, not densely clus-
tered fruits, 1.8—2.4 cm long.

13. Horsfieldia clavata W.J. de Wilde

Horsfieldia clavata W. J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 92, f. 13d-f. — Type: Hoogland
3663, New Guinea.

Shrub or tree, 3-6 m. Twigs 1.5—-3 mm diameter, glabrescent, hairs grey-rusty, eal
mm long; bark finely striate, not flaking; lenticels absent or inconspicuous. Leaves mem-
branous, elliptic or oblong, 7-18 by 3-6 cm, base short- to long-attenuate, apex acute-
acuminate (in Hoogland 3523 2 cm caudate); upper surface drying olivaceous, lower
surface with persistent, scattered, stellate-dendroid scale-like hairs 0.1—0.2 mm long,
especially on midrib, the nerves not much contrasting; dots absent; midrib slender above,
raised; nerves 10-20 pairs (including some intersecondary nerves), above thin and flat
or slightly raised, beneath much raised (not much contrasting in colour), lines of inter-
arching regularly looping, distinct; venation lax, rather indistinct; petiole 7-14 by 1—1.5
mm: leaf bud 7-10 by 1-1.5 mm, hairs 0.1 mm. /nflorescences with scale-like hairs 0.1
mm or less, among the leaves, delicate, 1 or 2 (or 3) times branched, lowest branch from
near the base; in male 2-3 by 1.5—2 cm, rather many-flowered; in female: 1—2 cm long,
2- or 3-flowered; bracts densely pubescent, 1—-1.5 mm long, caducous; flowers solitary
or 2 or 3 together; perianth 2-lobed, with stellate-dendroid hairs 0.1 mm; pedicel not
articulated. Male flowers: pedicel 2.5-3.5 by 1-1.5 mm, pubescent; buds clavate, with
tapering pedicel, together 4-5.5 by 1.5-2.2 mm; perianth rounded above, 1.5—2 by 1.5—
2.2 mm, cleft c. 1/10, lobes 0.2 mm thick, lower down perianth wall 0.5—0.7 mm thick;
androecium clavate, 1.5 by 0.7 mm, anthers 3 (or c. 6 thecae), 0.3 mm long, + stellate,
sessile. column not hollowed out; androphore thickish subcylindrical, slightly bullate-
striate, glabrous (Plate 1: 18). Female flowers: pedicel + slender, 2 mm long; buds ellip-
soid, 1.8(—2) by 1.2 mm, cleft c. 1/4; ovary ovoid, | by 0.6 mm, with dense stellate scale-
like hairs 0.1 mm or less, style 0.4 mm long, stigma 2-lobed, 0.2 mm long. Fruits | (or 2)
per infructescence, broadly ellipsoid-ovoid, 1.3 by 1 cm (excluding pseudostalk), base
broadly rounded, apex + acuminate, beak 2 mm, with hairs 0.1 mm long, drying brown-
ish, without lenticels; pericarp | mm thick; fruiting pedicel 6-10 mm, the pseudostalk
1.5—2 mm long; perianth not persistent. — Fig. 14d-f.

Field-notes — Shrub or treelet. Flowers yellow. Fruits orange or red.

Distribution — Malesia: Papua New Guinea (Northern Prov.).

De Wilde — Myristicaceae (Horsfieldia) 101

Habitat & Ecology — Locally common in regrowth in tall lowland forest on well-
drained soil; 0-50 m altitude; fl. & fr. Aug.

Note — Horsfieldia clavata is related to H. squamulosa and H. crux-melitensis which
have a similar clavate androecium. Horsfieldia squamulosa differs in its slender, male
pedicels. The pedicel, and hence the whole male flower of H. crux-melitensis is simi-
larly club-shaped as in the present species, but about twice as large; its leaves are also
larger and darker, and both male and female flowers have much thickened pedicels.

14. Horsfieldia coriacea W.J. de Wilde

Horsfieldia coriacea W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 50. — Type: bb Cel. III-27,
Sulawesi.

Tree 8-25 m. Twigs 2.5-4(—10) mm diameter, early glabrescent, hairs greyish brown,
0.1 mm; bark finely striate, not flaking; lenticels conspicuous or not. Leaves membra-
nous to thinly chartaceous, (elliptic-)oblong, 14—27 by 5—10 cm, base attenuate, apex
acute-acuminate; upper surface glabrous, drying olivaceous brown to blackish brown,
the midrib glabrous but towards the base in younger leaves finely pubescent, lower sur-
face glabrous; dots absent; midrib moderately raised above; nerves 13—18 pairs, raised
above, lines of interarching not distinct; venation lax, barely visible on either surface;
petiole 12-16 by 2.5-—3.5 mm; leaf bud 12—17 by 2—3 mm, hairs dense, grey-brown to
rusty, 0.1 mm long. /nflorescences behind the leaves, with sparse hairs 0.1 mm; in male:
2 or 3 times branched, few-flowered, 4-10 by 3—5 cm, peduncle 1—2 cm long, the flow-
ers in loose clusters of 3—5; in female (from infructescences): 2—5 cm long; bracts not
seen, caducous; flowers glabrous, perianth 3- or 4-lobed, pedicel not articulated. Male
flowers: pedicel 1.5—2 mm long; buds subglobose to broadly ovoid, 2—2.5 by 2—2.3 mm,
apex shortly rounded to subacute, not or only slightly collapsing on drying, cleft 1/2—
2/3, lobes 0.4—0.5 mm thick, coriaceous; androecium subellipsoid, 1.5—1.6 by 0.8—0.9
mm, in cross section + blunt-triangular (Plate 3: 88); thecae 10 or 12, at the base curved,
and towards the apex erect or somewhat curved, 1.6 mm long, free apices 0.1—0.2(—0.3)
mm, apical cavity narrow, 0).2—0.3(—0.5) mm deep, androphore narrow, 0.1—0.2 mm long,
hidden by the anthers. Female flowers not seen. Fruits 1—5 per infructescence, ellipsoid,
44.2 by 2.5-3.2 cm, glabrous, drying rust-brown, finely granulate and with at most a
few tubercles or lenticels; pericarp rather coriaceous, 3.5-8 mm thick; fruiting pedicel
2—4 mm long; perianth not persistent.

Field-notes — Bark and leaves with aromatic scent; branches horizontal; cauliflorous.
Flowers yellow, strongly scented; perianth fleshy. Ripe fruits orange.

Distribution — Malesia: Endemic in C Sulawesi.

Habitat & Ecology — Primary and disturbed forest (with Jmperata, Gleichenia, and
Melastoma) on ultrabasic soil; 100-700 m altitude; fl. Mar., Nov.; fr. Apr., July.

Notes — 1. In most flowers there are a few minute wart-like appendages 0.1 mm high
around the insertion of the androecium.

2. Horsfieldia coriacea, vegetatively and in fruit, resembles H. costulata, which has a
much larger distribution in Sulawesi and the Philippines. However, the latter differs in
thinner membranous leaves, drying generally more olivaceous, with the midrib on the
upper surface entirely glabrous, the lateral nerves usually forming a greater angle with
the midrib, and less conspicuous lenticels on the twigs. The fruits are generally larger

102 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Horsfieldia) 103

with a pericarp 8—10(—15) mm thick. Furthermore, the male flowers are quite different,
those of H. costulata being arranged in rather dense clusters of 5—10.

3. Horsfieldia coriacea seems closely related to H. majuscula (Sumatra, Peninsular
Malaysia) and H. xanthina (Borneo), both also having an elongate androecium, but with
a broader and tapered androphore, not hidden by the anthers; in H. majuscula the pedicel
is articulated.

15. Horsfieldia corrugata Foreman

Horsfieldia corrugata Foreman, Contr. Herb. Austral. n. 10 (1974) 45, f. 1; W.J. de Wilde, Gard.
Bull. Sing. 38, 1 (1985) 130, f. 20a—c. — Type: LAE 52461, Papua New Guinea.

Tree 5-12 m. Twigs (3—)4—5(—12) mm diameter, early glabrescent, hairs greyish to
rusty, 0.1 mm; bark striate, not flaking, lenticels large, usually not much contrasting in
colour. Leaves thinly coriaceous, elliptic-oblong, 12—29(—32) by 4.5—8.5(—10) cm, base
attenuate, apex acute-acuminate; upper surface drying dark brown, minutely pustulate
or not, lower surface early glabrescent; dots absent; midrib slender to rather broad, flat-
tish above; nerves 12-18 pairs, thin and flat above, beneath lines of interarching with
irregular loops, distinct or not, venation lax, indistinct; petiole 6-18 by 2—3.5 mm; leaf
bud 10-20 by 1.5-3 mm, hairs 0.1 mm. /nflorescences below the leaves, with rusty
stellate hairs 0.1 mm long or less; in male: 2 or 3 times branched, rather slender, (4—)6—
14 by 2-9 cm, peduncle 1—2.5 cm long; in female: up to 5 cm long, peduncle | cm long;
bracts pubescent, 1.5—4 mm long, caducous; flowers (in male) solitary or in loose clus-
ters of 2-5, glabrous or glabrescent, hairs scattered, less than 0.1 mm, perianth 2-lobed,
pedicel + tapering, not articulated. Male flowers: buds in lateral view subcircular, 3—3.5
by 3(-4) mm, apex broadly rounded, the lower half + tapering into the thickish tapering
pedicel, (2—)3—4 mm long; perianth cleft 1/2 to nearly 2/3, lobes 0.2—0.3 mm thick, often
with a few coarse blackish brown wart-like dots; androecium thickish, not much later-
ally compressed, above broadly rounded, (1.5—)2—2.2 by 2—2.2(—3) mm (Plate 2: 34);
thecae 16-24, erect, 2 mm long, free apical parts 0.1—0.2 mm, column narrowly hol-
lowed for 1/5—1/4, androphore 0.2—0.3 mm long. Female flowers: pedicel 4-5 mm long,
minutely pubescent; buds narrowly ovoid, almost glabrous, with a few coarse, dark brown
wart-like dots, 4.5 by 3 mm, cleft 1/4—1/3, lobes 0.3—0.4 mm thick, coriaceous; ovary
ovoid, somewhat dented or corrugated, 2.5—3 by 2.5 mm, with dense hairs less than 0.1
mm long, style and 2-lobed stigma glabrous, 0.8—1 mm long. Fruits 1(—4) per
infructescence, ramiflorous, broadly ellipsoid to subglobose, somewhat flattened, 6—7.5
by 4.5-6.5 cm, coarsely flanged and corrugated, drying blackish brown, with scattered,
coarse, paler coloured tubercles, glabrescent, at base sometimes a short pseudostalk,
apex acutish, pericarp + woody-corky, 10—20 mm thick; fruiting pedicel 5—10 mm long;
perianth not persistent. — Fig. 12a—c.

Fig. 12. Horsfieldia corrugata Foreman. a. Longitudinally opened male flower showing androecium;
b. ditto, female flower, showing pubescent ovary and narrow 2-lobed style; c. fruit. — H. pachycarpa
A.C.Sm. d. Leafy twig with infructescence; e. longitudinally opened male flower showing androe-
cium; f. ditto, female flower with pubescent ovary with short 2-lobed stigma; g. almost mature fruit
[a: Carr 14123: b: LAE 60020; c: Carr 14334; d: LAE 62196; e: LAE 51940; f: Clemens 5378:
g: NGF 38895]. — Scale bar for c, d, g = 2 cm; for a, b, e, f = 1.7 mm.

104 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Wood very light brown. Flowers yellow or orange. Fruits green, strongly
wrinkled or corrugated, and strongly ridged.

Distribution — Malesia: Papua New Guinea (Central, Northern, Milne Bay Prov.).

Habitat & Ecology — Primary and degraded rain forest of mountainous terrain on
slopes and ridges, fagaceous forest; 1200-1900 m altitude; fl. & fr. July to Dec.

Note — When in flower, H. corrugata may be difficult to distinguish from, e.g., H.
pachycarpa, H. tuberculata, or certain forms of H. laevigata. However, the few coarse
and conspicuous blackish brown wart-like dots on the perianth, found in male and fe-
male flowers, help to characterize H. corrugata. The large, corrugated and ridged thick-
lobed fruits are also distinctive, those of the other species may be similar but not ridged.

16. Horsfieldia coryandra W.J. de Wilde

Horsfieldia coryandra W.J. de Wilde, Blumea 32 ( 1987) 464. — Horsfieldia squamulosa auct. non
W J. de Wilde: W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 93, p.p. — Type: NGF 46892
(male fl.; fr.), Papua New Guinea.

Shrub or treelet, 1.5—6 m Twigs 1.5—2 mm diameter, with rusty hairs to 0.1 mm long,
glabrescent; bark finely striate, neither cracking nor flaking; lenticels few and incon-
spicuous or absent. Leaves membranous, elliptic-oblong to lanceolate, 5-20 by 0.7—5
cm, base (long-)attenuate or acute, apex long-acuminate, acumen often slender, to 3 cm
long, or gradually narrowed from slightly above the middle, blade above glabrous, dry-
ing dark brown (sometimes slightly olivaceous), beneath glabrescent with scattered hairs
0.1 mm or less remaining on and near the midrib, drying brown or olivaceous brown;
dots absent: midrib above slender, flat or raised; nerves 1 1—19 pairs (usually with some
intersecondary nerves not reaching the marginal nerve), above indistinct, flat or sunken,
beneath distinct, with lines of interarching usually distinct; venation coarse and distinct;
petiole 6-14 by 0.5—1.5 mm, glabrescent; leaf bud 7-12 by 1(-1.5) mm, with dense
rusty hairs 0.1 mm or less. Inflorescences among the leaves, 2 or 3 times branched,
peduncle 0.2—1 cm long, with grey-brown hairs 0.1 mm or less, rather few-flowered; in
male: 2-3 by 1.5—3 cm, flowers solitary or 2 or 3 together; in female: 1—1.5 cm long, not
or little branched, few-flowered; bracts oblong, 1-2 mm long, caducous; flowers 2-lobed,
with sparse hairs 0.1(—0.2) mm or less, glabrescent in the apical part, pedicel not articu-
lated. Male flowers: pedicel slender, (2.5—)4-6.5 mm long; buds slightly compressed
elliptic(-obovate) to elliptic-oblong, 2-3 by 1.5—2 mm, cleft 1/6 to nearly 1/4; perianth
0.50.7 mm thick, towards the apex of the lobes 0.2—0.3 mm; androecium club-shaped,
the apex subacute (to bluntish), 1.5—2.5 by (0.6—-0.8 mm; anthers 0.5—0.7(—1) mm long,
mutually touching, at apex free for c. 0.3 mm (Plate |: 15); thecae 8, column not hollow-
ed at apex; androphore glabrous or basally with scattered pale brown hairs less than
0.1 mm, the upper part with somewhat warted or wrinkled surface. Female flowers:
pedicel 2.5—3.5 mm; buds ellipsoid or + fusiform, 2—2.5 by 1.5-2 mm, cleft c. 1/3;
perianth (and lobes) 0.2—0.3 mm thick; ovary ovoid, 1.5 by 1.4 mm, densely minutely
pubescent, style erect with 2 + acute lobes together 0.4 mm long. Fruits 1 or 2 per in-
fructescence, broadly ellipsoid-ovoid, including the 0.5-2 mm long apiculum 1.2—-1.5
by 0.8-1 cm, pseudostalk 1.5—2.5 mm, all with hairs 0.1 mm or less, pericarp 0.5(—1)
mm thick, drying blackish, without lenticels; fruiting pedicel (5—)10—12 mm long, not or
hardly broadened towards the apex; perianth not persistent.

De Wilde — Myristicaceae (Horsfieldia) 105

Field-notes — Bark smooth, greenish brown or dark green, underbark red; exudate
red; inner bark brown; wood cream turning brown on exposure. Flowers yellow or or-
ange. Fruits (yellow-)green to orange; aril complete (orifice very small and folded away),
thin, red.

Distribution — Malesia: Papua New Guinea (Milne Bay Prov. incl. Normanby L.,
Northern Prov., Morobe Prov.).

Habitat & Ecology — Understorey shrub or low tree, sometimes gregarious. Lower
hill forest; Castanopsis forest on steep slopes, Eucalyptus-dominated forest, ridge for-
est, on riverbanks; forest on limestone; 200-500 m altitude; fl. & fr. throughout the year.

Note — Close to H. squamulosa, with similar, rather ellipsoid perianth, but differing
in some small features in flowers and fruits.

17. Horsfieldia costulata (Miq.) Warb.

Horsfieldia costulata (Miq.) Warb., Mon. Myrist. (1897) 350; W.J. de Wilde, Gard. Bull. Sing. 39,
1 (1986) 38. — Myristica costulata Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 48. — Type: de
Vriese & Teijsmann s.n., Sulawesi.

Horsfieldia pachythyrsa Warb., Mon. Myrist. (1897) 618; Koord., Meded. Lands PI. Tuin 19 (1898)
70, ‘crassithyrsa’.— Myristica pachythyrsa (Warb.) Boerl., Handl. 3 (1900) 86; 87, ‘crassithyrsa’.
— Horsfieldia minahassae auct. non (Warb.) Koord.: Koord., Meded. Lands Pl. Tuin 19 (1898)
70, p.p., quoad Koorders 18158. — Syntypes: Koorders 15156 (male, L lecto), 18/58 (L), 15170
(female, L), Sulawesi.

Horsfieldia confertiflora Merr., Philipp. J. Sci., Bot. 13 (1918) 285. — Type: FB 3183 (Ahern’s
Coll.), Philippines.

Horsfieldia megacarpa Merr., Philipp. J. Sci., Bot. 13 (1918) 286. — Type: BS 16527 (Ramos),
Philippines.

Horsfieldia villamilii Elmer ex Merr., Enum. Philipp. Flow. Pl. 2 (1923) 182, nom. nud.

Horsfieldia vulcanica Elmer ex Merr., Enum. Philipp. Flow. Pl. 2 (1923) 182, nom. nud.

Tree 9-30 m. Twigs 2.5—5(—10) mm diameter, early glabrescent, with grey-brown to
light rusty hairs, 0.1(—0.2) mm; bark finely to coarsely striate, not flaking; lenticels small,
generally inconspicuous. Leaves membranous or subchartaceous, elliptic-oblong to ob-
long-lanceolate, 15—30 by 5—13 cm, base narrowly rounded to attenuate, apex acute-
acuminate; upper surface drying olivaceous to dark brown, sometimes with whitish marks
as in H. irya; lower surface early glabrescent; dots absent; midrib above flat or slightly
raised, early glabrescent; nerves 14—21 pairs, above thin, flat or raised, lines of interarching
generally indistinct; venation lax, faint on both surfaces; petiole 7-14 by 2—4 mm; leaf
bud 8-14 by 2—2.5 mm, with dense (grey-)rusty hairs 0.1(—0.2) mm. Inflorescences mostly
behind the leaves, with dense or sparse hairs 0.1—0.2 mm; in male: 3 or 4 times branched,
many-flowered, 6-14 by 5—13 cm, peduncle 1—3 cm long; in female: 2—6 cm long, shortly
branched; bracts broadly triangular to elliptic-oblong, 2—4(—5) mm long, short-pubes-
cent, caducous; flowers in male in clusters of 5—10 each, in female fewer, glabrous, pe-
rianth 3- (or 4-)lobed, pedicel not articulated. Male flowers: pedicel slender, 0.4—0.6
(—0.7) mm long; buds (+ depressed-)globose, 1.5—1.8 by 1.5—2 mm; cleft c. 1/2, lobes
0.2 mm thick; androecium (depressed-)globose or broadly ovoid 0.5—0.8 by 0.7-1.1
mm, circular in cross section (Plate 3: 83); thecae 14-20, completely sessile (free apices
to 0.1 mm), incurved, apical cavity narrow, (0.1—)0.2 mm deep; androphore rather stout,
0.2—0.4 mm long, completely or partly hidden by the anthers. Female flowers: pedicel

Flora Malesiana, Ser. I, Vol. 14 (2000)

106

De Wilde — Myristicaceae (Horsfieldia) 107

0.5—1 mm long; buds subglobose, 2.3—2.5 mm diameter, cleft 1/3—1/2; ovary ovoid,
glabrous, 1.2-1.5 mm diameter, stigma minutely 2-lobed, 0.1—0.2 mm. Fruits 1—3 per
infructescence, subglobose to broadly ellipsoid or obovoid, 3.5—6 by 3—4 cm, glabrous,
finely granulate, drying bright brown to blackish brown; pericarp (4—)8—10 mm thick;
fruiting pedicel 2-4 mm long; perianth not persistent.

Field-notes — Tree with or without low buttresses, 30 by 10 cm; bark fissured or with
longitudinal grooves, often peeling off, sap first clear, turning red to brown-red; heart-
wood reddish. Flowers yellow. Fruits yellow to red, on the larger branches.

Distribution — Malesia: Philippines (incl. Palawan), Sulawesi.

Habitat & Ecology — Mixed rain forest, primary dipterocarp forest; recorded from
alluvial soil and volcanic soil, with Eucalyptus deglupta dominance; 250—1200 m alti-
tude; fl. & fr. throughout the year, but fl. mainly July—Sept.

18. Horsfieldia crassifolia (Hook. f. & Thomson) Warb.

Horsfieldia crassifolia (Hook.f. & Thomson) Warb., Mon. Myrist. (1897) 323, p.p.; J. Sinclair,
Gard. Bull. Sing. 16 (1958) 386, f. 34, pl. X-A; 28 (1975) 23; J.A.R. Anderson, Gard. Bull. Sing.
20 (1963) 195; W. J. de Wilde, Gard. Bull. Sing. 38, 2 (°1985’, 1986) 219, f. 25; Tree Fl. Sabah
& Sarawak 3 (2000) 366.— Myristica crassifolia Hook. f. & Thomson, Fl. Ind. (1855) 160;
A.DC., Prodr. 14, 1 (1856) 204; Miq., Fl. Ind. Bat. 1, 2 (1858) 68; Hook. f., Fl. Brit. India 5
(1886) 108; King, Ann. Roy. Bot. Gard. Calc. (1891) 308, pl. 140. — Myristica irya Gaertn. var.
crassifolia Mig. ex Hook. f., Fl. Brit. India 5 (1886) 108, pro syn. — Type: Griffith 4350 (see
notes by Sinclair, 1975: 25), Peninsular Malaysia.

Myristica horsfieldia auct. non Blume: Wall., Cat. (1832) n. 6806, p.p. (other parts are Horsfieldia
polyspherula and H. wallichii).

Myristica subglobosa Mig., F1. Ind. Bat., Suppl. 1 (1861) 383, p.p. (other part is Horsfieldia irya).

Myristica paludicola King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 328, pl. 169. — Horsfieldia fulva
(King) Warb. var. paludicola (King) Warb., Mon. Myrist. (1897) 299. — Syntypes: King’s coll.
4267, 4706, 6688, Wray 3071, Peninsular Malaysia.

Tree 10-25 m. Twigs 2-6(—8) mm diameter, rather early glabrescent, hairs yellow-
brown or rusty, woolly, 0.2—0.5 mm; bark coarsely striate, not flaking; lenticels sparse to
dense, distinct or not. Leaves coriaceous, elliptic to oblong, 10—20(—28) by 3.5—7(—10)
cm, base rounded to attenuate, apex rounded to subacute or rarely emarginate; upper
surface drying dull greenish brown to dark brown, lower surface + covered with sub-
persistent interwoven or spaced stellate scale-like hairs 0.1 mm high (when very young
with dendroid emergents), or glabrescent and then showing distinct hair scars, and scat-
tered dots and dashes; midrib above flattish; nerves 11—16 pairs, above thin and flat or
sunken; venation faint on both surfaces; petiole 9—20(—30) by 1.5—4.5 mm, leaf bud

Fig. 13. Horsfieldia crassifolia (Hook. f. & Thomson) Warb. a. Branch with leafy twig and infruc-
tescence with mature fruits; note persistent perianth and completely closed aril; b. twig with male
inflorescence; c. mature male flower, lateral view; d. ditto, longitudinally opened, showing androe-
cium; e. older twig with female inflorescence axillary to leaf scar; f. opened mature female flower
showing glabrous ovary and 2-lobed stigma; note that the flower is considerably larger than the
male flower; g. part of lower leaf surface with persistent indumentum and irregularly shaped dark-
coloured dots [a: Mondi 51; b—d: S 9226; e-g: SAN 27183]. — Scale bar for a, b, e = 2 cm; for c,
d, f = 0.85 mm; for g = 0.4 mm.

108 Flora Malesiana, Ser. I, Vol. 14 (2000)

7-12 by 2-3 mm with hairs 0.2-0.5 mm. Inflorescences among or behind the leaves, late
glabrescent or with persistent, dense, woolly dendroid hairs 0.2—0.5 mm; in male: 3-5
times branched, broadly paniculate, many-flowered, 6-20 by 4-15 cm, peduncle 0.5-2
cm long; in female: 3-14 cm long; bracts elliptic-lanceolate, 2—5(—7) mm long, pubes-
cent, caducous: flowers (male) in loose clusters of 2-7, glabrous; perianth 2-lobed, pedi-
cel slender, not articulated. Male flowers: pedicel (0.3—)1 mm long; buds globose or
slightly transversely ellipsoid, (0.8—)1-1.3 by I1-1.5 mm, cleft 1/3—1/2(—2/3), lobes 0.2—
0.3 mm thick; androecium globose or + transversely ellipsoid, barely laterally compres-
sed, 0.4—0.5 by 0.5—0.8 mm (Plate 2: 60); thecae (6—)8-12, widely spaced, connectives
broad (and androecium angular), the anthers free for almost the upper half or more;
androphore 0.2(—0.3) mm long, slender. Female flowers: pedicel 1.5—2.5 mm long; buds
broadly obovoid, 2-3 by 2—2.5 mm, cleft 1/5—1/3, lobes 0.6—1 mm thick; ovary obovoid,
glabrous, 1.5 by 1.2-1.5 mm, stigma of 2 sessile small lobes 0.1—0.2 mm high, running
out into a faint ridge at one side of the ovary. Fruits (1—)2—10 per infructescence, ovoid
to obovoid, 1.5—2.2 by 1.2—1.8 cm, glabrous, drying dark brown, with at most few lenti-
cel-like tubercles; pericarp 1.5—2 mm thick; fruiting pedicel 2-5(—7) mm long; perianth
persistent. — Fig. 13.

Field-notes — A few stilt-roots or low buttresses occasionally recorded; bark greyish,
fissured, flaking in small rectangular scales. Flowers yellow, strongly scented.

Distribution — S Thailand; Malesia: Sumatra (including Indragiri, Riau, Bangka,
Belitung), Peninsular Malaysia (Perak, Trengganu, Selangor, Negri Sembilan, Malacca,
Johore), Singapore, whole of Borneo.

Habitat & Ecology — Mostly in marshy forest, freshwater and peat-swamp forest; on
sandy soils, 0-200 m altitude; fl. & fr. throughout the year.

Notes — 1. The lower leaf surfaces of Borneo material of H. crassifolia are earlier
glabrescent as compared to those in Sumatra and Peninsular Malaysia.

2. Horsfieldia crassifolia may be confused with H. fulva, a species also with more or
less coriaceous leaves and a perianth persistent on the fruit, but with a 3-lobed perianth.
Sterile specimens of H. crassifolia may be recognized by the coriaceous leaves, which
have usually persistent scale-like hairs on the lower surface, and sparse to rather dense
irregularly shaped dark dots and dashes. Sinclair (1975: 26) remarked that the species
can easily be recognized from a distance by the rusty or cinnamon-brown colour of the
lower leaf surface. The species is very constant in habit, characters, and habitat.

19. Horsfieldia crux-melitensis Markgr.

Horsfieldia crux-melitensis Markgr., Bot. Jahrb. Syst. 67 (1935) 148, ‘crux melitensis’; J. Sinclair,
Gard. Bull. Sing. 28 (1975) 26, ‘cruxmilitensis’; W. J. de Wilde, Gard. Bull. Sing. 38, 1 (1985)
90, f. 13a—c; Blumea 32 (1987) 459. — Type: Schlechter 19246, Papua New Guinea.

Shrub or treelet 2-7 m. Twigs 1.5—3(—4) mm diameter, glabrescent, hairs rusty, 0.1
mm; bark finely striate, not flaking, lenticels absent or inconspicuous. Leaves membra-
nous, elliptic to obovate(-oblong), 12-27 by 5.5—-11.5 cm, base attenuate, apex acute or
broadly rounded, the tip acute-acuminate; upper surface drying dark brown, without or
with very minute paler dots, lower surface with hairs rather sparse, 0.1 mm, persistent,
especially on the midrib, or late glabrescent, nerves not contrasting in colour; dots ab-
sent; nerves 10-15 pairs, sometimes with intersecondary nerves, thin and flat above,

De Wilde — Myristicaceae (Horsfieldia) 109

Fig. 14. Horsfieldia crux-melitensis Markgr. a. Leaf; b. opened male flower showing club-shaped
androecium; c. opened female flower showing pubescent ovary with minute narrow 2-lobed style.
— H. clavata W.J. de Wilde. d. Twig with infructescence with mature fruit; e. mature male flower,
lateral view; f. ditto, opened, showing club-shaped androecium. — H. squamulosa W.J. de Wilde.
g. Habit of leafy twig with male inflorescences [a, c: LAE 73830; b: Schlechter 19246; d: Hoogland
3623; e, f: Hoogland 3663; g: Brass 7221]. — Scale bar for a, d, g = 2 cm; for b, c, e, f= 1.7 mm.

110 Flora Malesiana, Ser. I, Vol. 14 (2000)

much raised beneath, lines of interarching distinct, irregular; venation lax, rather indis-
tinct; petiole 10-16 by 1.5—2.5 mm; leaf bud 7-12 by 1-2 mm, hairs 0.1—0.2 mm. Inflo-
rescences (male) among the leaves, 1.5—5 by 1.5—4 cm, peduncle 0.6—1.5 cm long, rather
few-flowered, with woolly stellate-dendroid hairs 0.1—0.2 mm, 2 (or 3) times branched;
female inflorescences smaller; bracts subulate, to 5 mm, pubescent, caducous; flowers
solitary or up to 3 together, + clavate; perianth 2-lobed, with scattered stellate hairs 0.1
(—0.2) mm, densest towards base, pedicel tapering, pubescent, not articulated. Male flow-
ers: pedicel 7-8 by 2-3 mm, buds subglobose, 2—3 by 2—3.2 mm, gradually passing into
the tapered pedicel, forming a long club-shaped flower 9-11 by 23.2 mm, apex broadly
rounded, somewhat compressed, cleft 1/8—-1/5, lobes 0.3—0.6 mm long, 0.2—0.3 mm
thick (lower down 0.6—0.8 mm thick); androecium clavate, 1.5—2.5 by 0.7-1.2 mm,
apex + rounded, central column solid (Plate 1: /7): thecae 6—10, 0.2—0.3 mm long,
radiating, connate, androphore thick, subcylindrical, the surface + wrinkled-bullate, more
striate basally, glabrous. Female flowers: pedicel 6 by 2.6 mm, buds 2.2 by 3 mm, cleft
c. 1/10; ovary obovoid, 2 by 1.5 mm, densely pubescent, hairs 0.1 mm long, style with
2-lobed stigma 0.2—0.3 mm long. Fruits 1—4 per infructescence, ellipsoid-ovoid, 1.4 by
0.8—0.9 cm, apex apiculate for 2 mm, base broadly rounded; pericarp 0.5(—1) mm thick,
blackish, finely pubescent, partly glabrescent leaving a fine brown punctation, without
lenticels; seed ellipsoid; fruiting pedicel elongate, thickened towards the apex; perianth
not persistent. — Fig. 14a—c.

Field-notes — Stem 6—8 cm diameter; bark grey or dark green, wood cream or white.
Flowers cream or orange. Fruits yellowish or red.

Distribution — Malesia: Papua New Guinea (Morobe Prov.).

Habitat & Ecology — Mixed rain forest, lowland forest, common in wet areas; alti-
tude 0—50 m; fl. Jan.—May; fr. Mar.

20. Horsfieldia decalvata W.J. de Wilde

Horsfieldia decalvata W. J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 125. — Type: Idjan & Mochtar
18], Halmahera.

Tree 10-15 m. Twigs + flattened, sometimes faintly ridged, 1.5—4(-9) mm diameter,
early glabrescent, hairs brown, 0.1 mm; bark finely striate, not flaking; lenticels small,
not conspicuous. Leaves membranous, elliptic-oblong to oblong-lanceolate, 11-25 by
3.5-7 cm, base (long-)attenuate, apex acute-acuminate; upper surface drying dark brown,
lower surface early glabrescent, hairs 0.1 mm; dots absent; midrib above flat; nerves
12-16 pairs, above thin and flattish, on lower surface lines of interarching not very dis-
tinct; venation on upper surface fine, + distinct or not; petiole 5—10 by 1.5—2.5 mm; leaf
bud 6-10 by 1-2 mm, with dense dull brown hairs 0.1 mm long. Inflorescences + behind
the leaves, 2 (or 3) times branched; in male: 4—6 by (1—)2—4 cm: branches subglabrescent,
hairs 0.1 mm; peduncle 1—2 cm; in female: up to 2.5 cm long, caducous; bracts not seen;
flowers (in male) solitary or 2 or 3 together, minutely pubescent (hairs 0.1 mm); perianth
2-lobed. Male flowers: pedicel 1.5—2 mm long, not articulated: buds subglobose to short
pyriform, moderately compressed, 2.3 by 2.3 mm, apex broadly rounded, tapering ba-
sally into the pedicel, cleft c. 2/3, lobes 0.2-0.3 mm thick: androecium moderately flat-
tened, 1.6 by 1.4 mm, upper part rounded (Plate 2: 32): thecae c. 12 (anthers c. 6 with
+ broad connectives), 1.6 mm long, erect, free apical parts to 0.1 mm, androphore up to

De Wilde — Myristicaceae (Horsfieldia) itl

0.1 mm long, column cleft 1/5—1/4. Female flowers not seen; immature fruits (ovaries)
densely finely pubescent. Fruits 2-5 per infructescence, short-ellipsoid to subglobose,
1.1-1.2 by 1-1.1 cm, with hairs 0.1 mm or less, drying brown, with scattered small tuber-
cles; pericarp | mm thick; fruiting pedicel slender, 4 mm long; perianth not persistent.

Field-notes — Flowers brown. Fruits yellow.

Distribution — Malesia: Moluccas.

Habitat & Ecology — Forest at low altitudes, 0-100 m; fl. Sept.; fr. May—Nov.

Note — Horsfieldia decalvata superficially resembles several other species, includ-
ing the widespread H. laevigata, H. moluccana, and H. tuberculata, but is distinguish-
able by its finely pubescent, pear-shaped male flowers, erect anthers, pubescent ovaries,
and small, subglobose, finely pubescent fruits. In Horsfieldia laevigata the male bud is
more spherical in outline, and the fruit is much larger and pubescent. Horsfieldia moluc-
cana has incurved anthers and glabrous fruits. Horsfieldia tuberculata also has pear-
shaped flowers which are generally glabrous, and it has larger, glabrous fruits.

21. Horsfieldia discolor W.J. de Wilde

Horsfieldia discolor W.J. de Wilde, Blumea 32 (1987) 469; Tree FI. Sabah & Sarawak 3 (2000)
367. — Type: Kostermans 7414, fr., E Kalimantan.

Tree 10—18 m. Twigs grey-brown or straw, (2—)3—4 mm diameter, thinly with dull to
dark rusty scaly-stellate hairs 0.1 mm high, soon glabrescent; bark (coarsely) striate,
pale to grey-brown, neither cracking nor flaking; lenticels absent to many (see the note),
inconspicuous. Leaves thinly chartaceous, extremely brittle when dry, oblong(-lanceo-
late), (9-)13-18 by (3.5—)4-7.5 cm, base attenuate, apex + short-attenuate or acute-
acuminate; upper surface drying olivaceous to dark brown, lower surface early glabrescent
(glabrous), drying bright brown; dots absent; midrib above flat, very slender; nerves 10—
13 pairs, above flat, inconspicuous, lines of interarching indistinct; venation indistinct
on both surfaces; petiole 10-15 by 2-3 mm, drying dark brown, contrasting with the
pale twigs; leaf bud 10 by 2 mm, with dense, dull, scale-like hairs to 0.1 mm. Male and
female inflorescences and flowers not known (but see the note). Infructescences rather
stout, short, little or not branched, 1.5—3.5 cm long, early glabrescent, with 1 or 2 (or 3)
fruits. Fruits subglobose or ellipsoid, glabrous, drying (blackish) brown, somewhat wrin-
kled and at most with few wart-like lenticels, 5.5—8(—9) by 4—6.5 cm; pericarp 10—20
mm thick; fruiting pedicel 2-5 mm long; aril laciniate at apex for 1/4—1/5; perianth not
persistent.

Field-notes — Low to medium tree; bark red-brown, light brown, or black, rather
smooth with longitudinal surface marks or fissures, strips 1-2 cm wide, 0.5—1 mm thick,
once recorded as lenticellate; living bark red-brown or orange-brown, 5—7 mm thick,
soft, exuding clear pink (or red) sap; (sap)wood soft, white or yellow, with reddish streaks.
Fruits orange-yellow to orange-red, with the inner layer pink; seed (Sinclair 9278) 5.5
by 3 cm, brown.

Distribution — Malesia: Borneo (E Kalimantan, type; probably Sarawak and Sabah,
see specimens discussed by De Wilde, I.c.).

Habitat & Ecology — Forest on loam-soil over sandstone, 700—900 m altitude; fr.
July (E Kalimantan), Mar., June, Nov. Doubtful specimens from alluvial forest (riverbank
forest) and kerangas on sandy soils.

112. Flora Malesiana, Ser. I, Vol. 14 (2000)

Note — Horsfieldia discolor is known only in fruit, and characterized by the pale
colour of the young twigs that contrasts with the dark brown or blackish dry petioles.
It is closely related to H. obscura, known only from two male flowering specimens
which differ in the twigs turning brown on drying, and densely set with rather conspic-
uous lenticels. In H. discolor lenticels are almost absent, or in some tentatively included
specimens few to abundant, but they are never conspicuous. Furthermore, H. obscura
occurs on limestone, whereas H. discolor seems associated with sandstone and alluvial
soils.

22. Horsfieldia disticha W. J. de Wilde

Horsfieldia disticha W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 10; Blumea 32 (1987) 467; Tree
Fl. Sabah & Sarawak 3 (2000) 368. — Type: Sinclair (& Kadim) 10453, Brunei.

Tree 20 m. Twigs sometimes faintly angular, 2—3(—4) mm diameter (in fruit-bearing
part 11-13 mm diam.), dark grey-brown or reddish brown, very early glabrescent, hairs
dull rusty, 0.2-0.3 mm; bark longitudinally cracking; lenticels rather sparse but distinct.
Leaves chartaceous, oblong-lanceolate, 8—13.5 by 2.5-3.5 cm, base + rounded to short-
attenuate, apex acuminate (to 15 mm); upper surface drying olivaceous-brown, glabrous,
lower surface pale chocolate, glabrous; dots absent; midrib raised above, glabrous; nerves
9-13 pairs, raised above, lines of interarching indistinct; venation faint on both surfaces;
petiole 10-15 by 1-1.5 mm; leaf bud 10 by 2 mm, with dense dull rusty hairs 0.2—0.3
mm long. Male and female flowers not seen, but perianth apparently 3-lobed as judged
from the perianth scars on the fruits. Infructescences 3-6 by 2-4 cm, borne on the older
wood behind the leaves, glabrous (glabrescent), 3—6-fruited. Fruits ovoid-ellipsoid, 2.8—
3.2 by 2.1-2.5 cm, glabrous, drying dark brown, not tubercled nor lenticellate; pericarp
hard-woody, 8-10 mm thick; fruiting pedicel 10-15 mm long; perianth not persistent.

Field-notes — Bark with longitudinal, shallow furrows. Leaves dark green above,
paler beneath, dull on both surfaces. Fruits unripe, pear-shaped.

Distribution — Malesia: Borneo (Brunei).

Habitat & Ecology — Lowland forest, at side of new road, Andulau Forest Reserve
(West); fr. Aug.

Note — From both the general habit of the specimens, and especially the leaves with
the nerves raised above, H. disticha belongs to the H. polyspherula group. Horsfieldia
disticha differs from H. ridleyana of this group in its general habit, the raised nerves, the
longitudinally cracking of the bark on the twigs, and the much larger fruits with a con-
spicuously thick pericarp. The twigs have remarkedly conspicuous, pale, scattered lenti-
cels, contrasting with the dark bark. The available male inflorescences are too immature
to describe, but they possibly will not grow longer than 3 cm. They have ovate to elliptic
bracts 4(—5) mm long which have, like the inflorescences, dense shaggy rusty-red hairs
up to 0.5 mm long. The immature male buds including pedicels hardly reach 1 mm; they
are 3-lobed, glabrous, and possibly will resemble those of, e.g., H. polyspherula.

23. Horsfieldia elongata W.J. de Wilde

Horsfieldia elongata W.J. de Wilde, Blumea 32 (1987) 465, f. 1. — Type: Stone 10779, Peninsular
Malaysia.

De Wilde — Myristicaceae (Horsfieldia) 113

Fig. 15. Horsfieldia elongata W.J. de Wilde. a. Habit of branchlet with male inflorescences; b. ma-
ture male flower bud; c. ditto, longitudinally opened, showing androecium; d. androecium, longi-
tudinal section, half-schematic [all Stone 10779]. — Scale bar for a = 2 cm; for b—d = 0.85 mm.

Tree c. 8 m. Twigs 1.5—2 mm diameter, hairs pale brown, 0.1 mm, early glabrescent,
bark (coarsely) striate, drying greyish, not flaking; lenticels present. Leaves + membra-
nous, oblong-lanceolate, 13—21 by 3.5—6 cm, base long-attenuate, apex rounded or blunt-
ish to subacute; upper surface glabrous, drying dull, brown-olivaceous, lower surface
slightly paler, glabrous (very early glabrescent); dots absent; midrib above slender, flat;
nerves 15-18 pairs, sunken or flattish, inconspicuous; venation lax, indistinct; petiole
10—20 by 1.5—2.5 mm, drying blackish; leaf bud 10—12 by 1.5—2 mm, with dense pale
brown hairs 0.1 mm long. /nflorescences among the leaves, with pale brown stellate
hairs 0.1 mm; in male: small, + slender, rather few-flowered, once or twice branched,

114 Flora Malesiana, Ser. I, Vol. 14 (2000)

1.5-2.5 by 1-2 cm, peduncle 0.1—0.5 cm long; bracts + elliptic, densely pale brown vil-
lous by hairs 0.1 mm, caducous; flowers solitary or 2 or 3 together, glabrous or glabrescent,
hairs scattered, less than 0.1 mm, perianth 3- or 4-lobed, pedicel not articulated. Male
flowers: pedicel (0.5—)1—2 mm long; buds (+ ovoid-)ellipsoid, 2—2.4 by 1.5—1.8 mm,
cleft 1/3—1/2(—2/3), lobes 0.1—0.2 mm thick; androecium nearly sessile, ellipsoid, 1.8-2
by (1.3—)1.5 mm, in cross section subcircular (Plate 3: 5/); thecae appressed, 16 or 18,
almost completely sessile, free apex of anthers 0.1(—0.2) mm; column with narrow exca-
vation 0.2(—0.3) mm deep; androphore narrow, 0.2—0.3 mm long. Female inflorescences,
flowers, and fruits not seen. — Fig. 15.

Field-notes — Small tree, flowers yellowish.

Distribution — Malesia: Peninsular Malaysia (Pahang, Fraser’s Hill, start of Pine
Tree Hill trail).

Habitat & Ecology — Known only from the type. About 1300 m altitude; fl. June
1972.

Note — Horsfieldia elongata belongs to the group of species with pallid bark of
twigs and often dispersed leaves, like H. atjehensis, H. pallidicaula, H. sparsa, and H.
sucosa; the present species has distichous leaves. It keys out besides H. pallidicaula
from Borneo, to which it seems closely related.

24. Horsfieldia endertii W. J. de Wilde

Horsfieldia endertii W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 24; Tree Fl. Sabah & Sarawak
3 (2000) 368. — Type: Endert 3996, E Kalimantan, W Kutei.

Tree 4-25 m. Twigs 2.5-4(—8) mm diameter, blackish brown, early or late glabrescent,
hairs harsh, deep rust coloured, 0.3—0.6 mm; bark coarsely striate, sometimes tending to
flake: lenticels coarse and conspicuously pale. Leaves (strongly) coriaceous, elliptic-
oblong to oblong-lanceolate, 8-17(—26) by 3-6(-10) cm, base rounded to (short-)atten-
uate, apex rounded to subacute; upper surface glabrous (glabrescent), but base of midrib
late glabrescent, drying olivaceous, yellowish, or dark-brown, lower surface pale brown
to chocolate, not much contrasting with the upper surface; dots absent, but usually with
conspicuous pale yellowish hair scars (lens!); midrib above relatively broad, raised; nerves
8-15 pairs, above sunken, flattish, or slightly raised, lines of interarching fairly regular,
sometimes distinct: venation lax, indistinct on both surfaces; petiole 6-16 by 2-3.5(—4)
mm, leaf bud 10-20 by 3—4 mm, with hairs 0.3—0.6 mm. Inflorescences below the leaves,
with dense, shaggy hairs 0.5—1 mm long; in male: 2 or 3 times branched, not many-
flowered, (1.5—)3-10 by (1—)2—5 cm, peduncle 0.5—2 cm long, the flowers in clusters of
(1—)2-6; in female: + few-flowered, 2—4 cm long; bracts broadly ovate-ellipsoid, 3—7
mm long, densely pubescent, caducous; flowers with perianth 3- (or 4-)lobed, glabrous
(glabrescent), pedicel puberulous in the lower half, not articulated. Male flowers: pedi-
cel 2-3 mm long; buds + obovoid (immature) to ellipsoid, 2.5-3.5 by 2-2.5 mm, gla-
brous, cleft 1/3—1/2, slightly collapsing on drying, lobes 0.3 mm thick; androecium
+ sessile, obovoid to truncate-ellipsoid, 2-2.8 by 1.4-1.6 mm, in cross section sub-
triangular (Plate 3: 74); thecae 24-28, 2—2.8 mm long, almost completely sessile (an-
thers free at apex for only 0.1 mm or less), column with a narrow apical hollow or slit to

De Wilde — Myristicaceae (Horsfieldia) IS

1/5—1/3; androphore narrow, (0—)0.1—0.3 mm long, hidden by the anthers. Female flow-
ers: pedicel 1.5—2 mm long, basally with hairs 0.2 mm; buds broadly ovoid-ellipsoid,
2.5—3 by 2-2.5 mm, cleft 1/3—1/2; ovary ovoid-ellipsoid, 2 by 1.5 mm, glabrous, stigma
0.2 mm high, minutely 2-lobed. Fruits 1—4(—8) per infructescence, ellipsoid, apex rounded
to acutish, 3—4.2 by 1.6—2.4 cm, glabrous, drying grey-brown to dark brown, without or
with a few tuberculate lenticels; pericarp 2-4 mm thick; fruiting pedicel 3—5 mm long;
perianth not persistent.

Field-notes — Tree sometimes dwarfed; bark cracked or finely fissured, brown(-black);
inner bark reddish; cambium and sapwood whitish; exudate turning reddish. Flowers
yellow. Fruits orange-yellow, pink(-red), or orange-red.

Distribution — Malesia: Borneo (Sarawak, Sabah, E Kalimantan).

Habitat & Ecology — Montane; in ridge- or mossy forest, montane dwarfed forest on
wind-swept crests; on sandy soil, black or brownish soil; 1200-2100 m altitude; fl. & fr.
throughout the year.

Notes — |. A similar species, H. xanthina, differs in the androecium and in a much
shorter indumentum; H. montana differs in its globose male flowers of only 2 mm long.
Horsfieldia endertii is characterized by ellipsoid male flowers 1.5—3.5 mm long, with an
androecium which is distinctly longer than broad, not articulated pedicels, very coriaceous
leaves, harsh-haired leaf buds, and inflorescences; on the lower leaf surface there are
practically always pale-yellowish hair scars, visible with a lens. The species usually has
+ rounded leaf tips, as in H. montana, but the leaves in the latter usually dry blackish and
are only 4-14 cm long and thinner.

2. The type, Endert 3996, with male flowers, is the only specimen from Kalimantan.
The remaining specimens from Sarawak and Sabah somewhat differ in general appear-
ance, their twigs are often less roughly hairy towards the apex, and their leaves are
somewhat more coriaceous with the lateral nerves ascending at a slightly sharper angle
to the midrib.

25. Horsfieldia flocculosa (King) Warb.

Horsfieldia flocculosa (King) Warb., Mon. Myrist. (1897) 297; J. Sinclair, Gard. Bull. Sing. 16
(1958) 398, f. 38; W.J. de Wilde, Gard. Bull. Sing. 38, 2 (1985’, 1986) 207. — Myristica floccu-
losa King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 302, pl. 131. — Type: King’s coll. 8618, Penin-
sular Malaysia, Selangor.

Tree 10—28 m. Twigs 6—10(—12) mm diameter, with dense felty-woolly yellow-brown
or pale brown hairs (1—)1.5—3 mm, late glabrescent; bark coarsely striate, soon longitu-
dinally cracking, eventually transversely cracking and + flaking; lenticels absent or in-
distinct. Leaves chartaceous, oblong(-lanceolate), 18—40(—45) by 6—13(—18) cm, base
broadly or narrowly rounded or subcordate, apex acute-acuminate; upper surface drying
olivaceous to (light) brown, finely pustulate and wrinkled, glabrous, lower surface with
dense woolly hairs 1.5—2 mm long; dots absent; midrib flat above; nerves 15—20 pairs,
above sunken, lines of interarching distinct and regular; venation lax, faint above, ob-
scured by the indumentum on the lower surface; petiole 7—14(—20) by 5—7 mm, densely
pubescent; leaf bud stout, 10-15 mm long. /nflorescences behind the leaves, densely

116 Flora Malesiana, Ser. I, Vol. 14 (2000)

woolly-pubescent; in male: stout, many-flowered, 3 or 4 times branched, (8—)12—20 by
5-14 cm, peduncle 3-5 mm diameter, 0.1—1 cm long; in female: little branched, 1.5—3
cm long, few-flowered; bracts ovate to lanceolate, densely pubescent, 5—20 mm long,
caducous: flowers (male) solitary or 2—3(—4) together, not densely clustered, glabrous,
perianth (3- or) 4-lobed in male, (2- or) 3-lobed in female, pedicel not articulated. Male
flowers: pedicel slender, (1.5—)3—4 mm long; buds broadly ellipsoid to obovoid, some-
times subglobose, not or slightly compressed, (2—)2.2—3 by 2—2.7 mm, cleft 1/5—1/4,
lobes 0.2 mm thick; androecium + ellipsoid or broadly obovoid, subtruncate, 1.2-1.5
(-2) by 1-1.3(-1.5) mm, not or only little compressed, base broadly rounded, often
faintly 4-angular in cross section (Plate 2: 52); thecae 20—26, entirely sessile, apically
slightly incurved, column broad, with shallow apical hollow (0).2—0.3 mm deep; androphore
short, 0.1 by 0.4 mm, usually hidden by the slightly sagged anthers. Female flowers:
pedicel stoutish, 2 mm long; buds broadly ellipsoid, 3.5 by 3 mm, cleft 1/6—1/5, lobes
0.4—0.5 mm thick; ovary broadly ovoid, 1.8 by 2 mm, glabrous except for a few minute
hairs (always?) on the suture below the stigma, stigma minute, faintly 2-lobed, 0.3 mm
long. Fruits (almost mature, according to Sinclair) subglobose to slightly ellipsoid, gla-
brous, 3 by 2.5 cm; pericarp 5 mm thick; fruiting pedicel 5 mm long; perianth (at first)
persistent.

Field-notes — Buttresses absent; bark superficially fissured, blackish brown; inner
bark pinkish brown, laminated; sapwood whitish; exudate watery, red. Young leaves
flocculose: lamina somewhat bullate, thickish, glossy medium green above, golden be-
low. Flowers yellow, or waxy light yellow; perianths are described by Sinclair (1958)
as “covered with circles which are hyaline in the centre and brown round the circum-
ference.”

Distribution — Malesia: E Sumatra (Jambi), Peninsular Malaysia (Selangor, Negri
Sembilan, Pahang, and Johore).

Habitat & Ecology — Lowland rain forest, also swampy forest, regenerating forest;
0-300 m altitude; fl. mainly Feb.—June.

Note — Horsfieldia flocculosa is related to H. superba, H. fulva (there too female
and male flowers are + similar in shape), and also to the H. grandis group, but it is quite
distinguishable by its flocculose indumentum; the margins of the dry leaves are revo-
lute.

26. Horsfieldia fragillima Airy Shaw

Horsfieldia fragillima Airy Shaw, Kew Bull. 1939, n. 10 (1940) 542: W.J. de Wilde, Gard. Bull.
Sing. 39, 1 (1986) 29, f. 29; Tree Fl. Sabah & Sarawak 3 (2000) 369. — Type: Richards 2602,
Sarawak, 4th Div., Mt Dulit.

Tree 10—30 m. Twigs sometimes faintly angular, 2.5—7(—15) mm diameter, early gla-
brescent, hairs 0.3 mm; bark coarsely striate, + flaking; lenticels distinct, on older wood

Fig. 16. Horsfieldia fragillima Airy Shaw. a. Leafy twig apex; b. older twig with male inflorescence
axillary to leaf scar; c. mature male flower, lateral view; d. ditto, longitudinally opened, showing
androecium: e. androecium, longitudinal section; f. twig with infructescences, fruit ma-ture; note
persistent perianth [a—e: § 34358; f: S 16985]. — Scale bar for a, b, f = 2 cm; for c—e = 0.85 mm.

117

De Wilde — Myristicaceae (Horsfieldia)

118 Flora Malesiana, Ser. I, Vol. 14 (2000)

inconspicuous. Leaves sometimes (partly) in 3 rows, membranous to thinly (rarely thickly)
coriaceous, oblong(-lanceolate), usually + tapering below the middle, 20—45 by 6.5-
12(—15.5) cm, base narrowly (rarely broadly) rounded to attenuate, apex acute(-acum1-
nate); upper surface drying olivaceous to brown, lower surface early glabrescent, with
some minute indumentum remaining at the base of the midrib; dots absent; midrib raised
above. at base broad and flat; nerves 20—30 pairs, raised above, lines of interarching
+ indistinct: venation lax, faint above; petiole stout, faintly pulvinate, 4—13(—20) by 3-8
mm: leaf bud 12-20 by 3-5 mm, with dense hairs 0.3 mm. Inflorescences behind the
leaves, with sparse hairs (0.1—)0.2—0.4 mm long; in male: large, very many-flowered, in
clusters of 2-6, 4 or 5 times branched, 15-30 by 10—20 cm, peduncle 2.5—6 cm long; in
female (from infructescences): 6-13 cm long, stout, rather few-flowered; bracts up to 12
by 5 mm, tomentulose, caducous; flowers glabrous, perianth 3- (or 4-)lobed; pedicel
glabrous, not articulated. Male flowers: pedicel 1-1.5(—2) mm long; buds (somewhat
depressed) globose, 1.42 by 2—2.5 mm, cleft c. 1/2, lobes (0.2(-0.3) mm thick; androecium
depressed-globose, synandrium almost saucer-shaped, deeply and broadly depressed in
the centre, 0.5-1 by 1-1.5 mm (Plate 3: 77); thecae 14-18, almost completely sessile,
free apices up to 0.1 mm, incurved; column broadly hollowed to about halfway, andro-
phore narrow, up to 0.1 mm long. Female flowers (from fruit): buds 4-5 by 3—4 mm.
Fruits up to 8 per infructescence, on the older wood, broadly ellipsoid, possibly slightly
flattened, 6-8 by 4—6 cm, glabrous, drying dark brown, wrinkled and usually with small
or large warts; pericarp 10-20 mm thick; fruiting pedicel 2-4 mm long; perianth persist-
ent. — Fig. 16.

Field-notes — Trunk without buttresses, and then often with broad gullies, rounded-
fluted, or with short and rounded buttresses only; bark chocolate, reddish, or blackish
brown, furrowed or not, + fissured, mostly flaky; inner bark 3 mm thick, pale pink or
reddish to yellowish; sapwood soft, whitish or (yellow- pink. Branches somewhat droop-
ing. Flowers yellow. Fruits ramiflorous, to 10 by 7 cm (seed 3 by 2 cm), greenish yellow
turning rose-pink to red; pericarp to over 2 cm thick.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, W & C Kalimantan; pos-
sibly E Kalimantan).

Habitat & Ecology — Primary lowland dipterocarp forest; hill slopes, and flat land,
once found in seasonal swamp; common on leached clayey loam over sandstone, and
sandy clay, often along or near streams and in riverine forest; 0—400 m altitude; fl. Feb.—
May; fr. throughout the year.

Uses — Fruits edible, very acid and resinous.

Notes — 1. Horsfieldia fragillima may be confused in the sterile state superficially
with H. laticostata, as the former may have a similar broad basal part of the midrib.
Horsfieldia laticostata differs in the male flowers, in much smaller fruits, and in differ-
ent texture and colour of the leaves. Horsfieldia fragillima seems related to H. splendida
and H. pulcherrima, which have similar androecia (though with a narrower hollow) and
not articulated pedicels, and they differ from the present species in general habit, includ-
ing their markedly hairy lower leaf surface.

2. Argent & Sidiyasa et al. 93176, from C Kalimantan, is an exceedingly stout speci-
men with large coriaceous leaves, up to 15.5 cm wide. The mature aril is bright yellow
(arils are red in almost all Myristicaceae).

De Wilde — Myristicaceae (Horsfieldia) 119

27. Horsfieldia fulva (King) Warb.

Horsfieldia fulva (King) Warb., Mon. Myrist. (1897) 297; J. Sinclair, Gard. Bull. Sing. 16 (1958)
396, f. 37; 28 (1975) 33; W.J. de Wilde, Gard. Bull. Sing. 38, 2 (°1985’, 1986) 199. — Myristica
fulva King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 297, pl. 124. — Syntypes: Maingay 1304 (2426),
Scortechini 154a (lecto), Peninsular Malaysia.

Tree 10-20 m. Twigs 3—5(—10) mm diameter, late glabrescent, hairs dense, 0.2—0.3
mm long; bark grey(-brown), finely striate, not flaking; lenticels many, not conspicuous.
Leaves chartaceous to coriaceous, elliptic-oblong to (obovate-)oblong, 13—21 by 4—9.5
cm, base attenuate, apex acute-acuminate or bluntish; upper surface drying dull olivaceous
to brown, lower surface pale brown, glabrous or with persistent hairs 0.2—0.3 mm long
on and near the midrib and lateral nerves; dots absent; midrib + raised above, early
glabrescent; nerves | 1—14(—18) pairs, above flat or + sunken, lines of interarching regu-
lar, not distinct; venation hardly visible on both surfaces; petiole 8—13 by 2.5-3.5 mm;
leaf bud 10-15 by 2.5—3 mm, with dense hairs 0.2—0.3 mm long. /nflorescences behind
the leaves, with hairs 0.2—0.3 mm; in male: about 3 times branched, many-flowered,
3-10 by 2-6 cm, peduncle 0.2—0.7 cm long; in female: few-flowered, 1—2 cm long;
bracts not seen, caducous; flowers solitary or in loose clusters of up to 5, glabrous,
perianth 3-lobed, pedicel + articulated. Male flowers: pedicel 1-3 mm long, slender;
buds ellipsoid(-obovoid), 3—4(—5) by 2—2.5 mm, apex rounded, base + attenuate to
rounded, cleft 1/5—1/4, lobes 0.2—0.3 mm thick; androecium long-ellipsoid, apex trun-
cate to rounded, subcylindrical to + triangular, 2-3 by 1.1—3 mm (Plate 2: 47); thecae
20-24, almost entirely sessile, mutually appressed, free apices to 0.1 mm, column at
apex narrowly hollowed or cleft to 0.3 mm, androphore narrow, to 0.1 mm long. Female
flowers (Sinclair 1958): pedicel stout, 3 mm long; buds long-ellipsoid, S—6 mm long;
ovary 2.5—3 mm long, glabrous, stigma sessile, bilobed. Fruits up to 3 per infructescence,
ovoid-ellipsoid, apex obtuse to acutish, base broadly rounded, 2.2—2.4(—3) by 1.6—2
(—2.5) cm, glabrous, drying bright brown, without lenticels or warts; pericarp 3 mm
thick; fruiting pedicel 3—5 mm long; perianth persistent (always?).

Field-notes — Bark yellowish brown, thin, shallowly longitudinally fissured but not
flaking; inner bark orange; wood white; sap watery, pale pink, not copious. Flowers
orange. Fruits yellow.

Distribution — Malesia: Sumatra (Jambi Prov.), Peninsular Malaysia (Perak, includ-
ing Pulau Rumbia; Selangor, Negri Sembilan, Malacca).

Habitat & Ecology — Lowland rain forest; undulating country, on ridges; 0-200 m
altitude; fl. & fr. throughout the year.

Note — Horsfieldia fulva is easily recognized by the dull parchment-like leaves when
dry; the nerves are flat above or somewhat sunken, and the venation is not or hardly
visible; it is one of the few species with a 3-lobed perianth which is elongate and rather
large, 3 mm long or more. The species is closely related to H. superba, which is larger in
size in almost all aspects, and has a persistent indumentum on the lower leaf surface.

28. Horsfieldia glabra (Blume) Warb.

Horsfieldia glabra (Blume) Warb., Mon. Myrist. (1897) 313, t. 21, p.p. — Myristica glabra Blume,
Bijdr. (1826) 576. — Syntype: Blume s.n., Java.

120 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 6—25 m. Twigs 2—4 mm diameter, (blackish) brown, early glabrescent, hairs
(grey-)brown, up to 0.2 mm long; bark finely striate, not flaking; lenticels conspicuous.
Leaves in 2 or 3 rows, membranous to thinly coriaceous (very brittle when dry), elliptic
or obovate to oblong-lanceolate, 8-27 by 3—-10.5 cm, base usually long-attenuate, apex
acute(-acuminate); upper surface drying olivaceous to dark brown; lower surface early
glabrescent; dots (not dashes) present (lens! ); midrib flat above; nerves 8-16 pairs, thin,
flat. or sunken above (in var. glabra sometimes + raised), lines of interarching indistinct;
venation lax. indistinct on both surfaces; petiole 10-15 by 1.5—2.5 mm; leaf bud 7-12 by
1.5-2.5 mm, slender, in tristichous twigs slightly thicker, with (grey-)brown hairs 0. 1—
0.2 mm. Inflorescences among or behind the leaves, with dense to sparse hairs to 0.1 mm
long, sometimes glabrescent; in male: (2 or) 3 (or 4) times branched, many-flowered,
5-10 by 4-7 cm, peduncle 0.4—1.5 cm long; in female: | or 2 times branched, 2—4 by
1-3 cm; bracts elliptic-lanceolate, 2-5 mm long, short-pubescent, caducous; flowers in
male (2—)3—5 in loose clusters, in female solitary or 2 or 3 together, glabrous; perianth
(2-) 3- or 4-lobed, pedicel articulated or not, usually mixed in one inflorescence. Male
flowers: pedicel slender or thickish; buds globose or broadly ellipsoid or obovoid, 1.5—
2.5 mm long, cleft 1/2—2/3, lobes 0.2-0.3 mm thick; androecium (depressed-)globose,
or ellipsoid or short-obovoid (Plate 3: 92a, b); thecae 18—30, almost completely sessile;
androphore narrow, (0—)1—2 mm long. Female flowers: pedicel 0.5—1.5 mm long; buds
ellipsoid, 2.5-3 by 2.2-2.5 mm, cleft 1/3 to c. 1/2; ovary ovoid, 1.5—2 by 1.2-1.5 mm,
glabrous, the stigma narrowly 2-lobed, 0.1 mm high (var. glabra), or broad-lipped, 0.2
(0.3) mm high (var. oviflora). Fruits 2-6 per infructescence, ovoid-ellipsoid, base
(broadly) rounded, 1.8-2.4 by 1.4-1.9 cm, glabrous, drying blackish brown, without
lenticel-like tubercles; pericarp 1—2.5 mm thick; fruiting pedicel 1-2.5 mm long; peri-
anth not persistent.

Distribution — Malesia: W, C & S Sumatra, Mentawai I. north to Simeulué (Sima-
loer I.), Java (3 varieties), Bali.

Note — In H. glabra dots are always present on the lower leaf surface, a character
regarded by Sinclair (1958: 413) as exclusive for the related H. punctatifolia which it
resembles vegetatively.

KEY TO THE VARIETIES

la. Male buds globose or subglobose, [1.7—2.5 mm long], cleft 1/2—2/3; androecium
globose or depressed-globose, circular or faintly triangular in cross section. [Pedicel

+ slender. Leaves membranous to chartaceous, nerves flat or slightly raised above.
Fruits 18-24 mm long. — Distribution as the species; 0—600(—1500) m altitude. |

oh MATE RE ies Sti Lay ee SE Peck rosette el eine eS eaece a. var. glabra

b. Male buds broadly ellipsoid to broadly obovoid, cleft about halfway; androecium

ellipsoid to obovoid, blunt-triangular in cross section .....+.-++++++++5+05: 2
2a. Male buds ellipsoid, 1.5 mm long; androecium ellipsoid. Pedicel slender. Leaves
membranous, nerves flat. Fruits not seen. —E Java .......... b. var. javanica

b. Male buds broadly ellipsoid-obovoid, 2—2.5 mm long; androecium broadly ellip-
soid-obovoid. Pedicel rather short and thickish. Leaves chartaceous to subcoriaceous,
nerves flat or sunken above. Fruits 18-20 mm long. — W & C Java; 600-1500 m
altitude! o- aliscink all eerie 5 coved sk ee eps a c. var. oviflora

De Wilde — Myristicaceae (Horsfieldia) 121

a. var. glabra

Horsfieldia glabra (Blume) Warb., Mon. Myrist. (1897) 313, t. 21, p.p.; J. Sinclair, Gard. Bull.
Sing. 16 (1958) 411, p.p., for the basionym only; 28 (1975) 35, p.p.; Backer & Bakh. f., Fl. Java
1 (1964) 138; W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 57. — Myristica glabra Blume,
Bijdr. (1826) 576; Rumphia | (1837) 191, t. 64 f. 1; Miq., Pl. Jungh. (1852) 172; Fl. Ind. Bat. 1,
2 (1858) 65 (excl. Myristica integra Wall.); Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 49 (excl. var.
sumatrana). — Pyrrhosa glabra (Blume) Hassk., Cat. Pl. Hort. Bog. (1844) 174.

Myristica glabra Blume var. grandifolia Migq., Fl. Ind. Bat. 1, 2 (1858) 65; Suppl. 1 (1860) 156. —
Type: Teijsmann s.n., W Coast Sumatra.

Leaves membranous to chartaceous, up to 27 cm long; nerves flat or slightly raised
above. Male flowers: pedicel slender, clearly marked-off from the perianth; buds glo-
bose, 1.7—2.5 mm diameter, cleft 1/2—2/3, perianth 3- or 4-lobed, lobes 0.2—0.3 mm
thick; androecium globose or depressed-globose, 0.8—1.3(—1.5) by 1.3—1.6 mm, circular
or faintly triangular in cross section (Plate 3: 92a); thecae 20-30, almost completely
sessile, free apices to 0.1 mm, curved over towards the apex and more or less into the
rather broad, 0.3—0.5 mm deep, apical cavity; column broad; androphore narrow, up to
0.2 mm long. Female flowers: stigma minutely 2-lobed, 0.1 mm long. Fruits 1.8—2.4 cm
long.

Field-notes — Bark smooth to rough, shallowly longitudinally fissured. Flowers yel-
low, smelling of Peru-balsam. Fruits glossy greenish orange, pericarp to 4 mm thick.

Distribution — As the species.

Habitat & Ecology — Primary and degraded forest, also in coastal forest on lime-
stone; 0O—800(—1500) m altitude; fl. & fr. throughout the year.

Notes — 1. Phyllotaxis in Java di- or tristichous, sometimes mixed in one collection;
in Sumatra distichous.

2. The male buds of the Sumatra specimens are faintly ellipsoid, strictly globose in
Java.

b. var. javanica W.J. de Wilde

Horsfieldia glabra (Blume) Warb. var. javanica W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 59.
— Type: Koorders 5210, Java.

Leaves membranous, 9—12 cm long, nerves above flat. Male flowers: pedicel slender;
buds ellipsoid, 1.5 by 1.2—1.3 mm, cleft c. 1/2, perianth 3-lobed, lobes 0.1—0.2 mm thick;
androecium ellipsoid, apex subtruncate, 1|—1.2 by 0.7—0.8 mm, blunt-triangular in cross
section (Plate 3: 92b); thecae 18-30, completely sessile, free apices up to 0.1 mm, at
apex not or only slightly incurved; apical cavity small and narrow, 0.1—0.2 mm deep;
column solid; androphore narrow, up to 0.1 mm long. Female flowers and fruits not
seen.

Distribution — Malesia: Java (possibly only in E Java), Bali (collection from
submontane area).

Note — According to the flowers this variety seems related to H. penangiana, a spe-
cies also with dotted leaves and a perianth of similar size and shape (partly), but with the
androecium round in cross section and with fewer thecae, 10—18(—20); in H. penangi-
ana the twigs are more slender and the leaves generally smaller.

122 Flora Malesiana, Ser. I, Vol. 14 (2000)

c. var. oviflora W.J. de Wilde

Horsfieldia glabra (Blume) Warb. var. oviflora W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 59. —
Type: bb Ja. 3827 (L), Central Java.

Leaves chartaceous to subcoriaceous, 8-15 cm long; nerves flat or sunken above.
Male flowers: pedicel 1 mm long, thickish; buds broadly obovate-ellipsoid, 2—2.5 by
1.7-2.3 mm, cleft c. 1/2, perianth 3-lobed, lobes 0.2—0.3 mm thick; androecium ellip-
soid-obovoid, apex subtruncate, 1.2—1.8 by 0.8—1 mm, blunt-triangular in cross section;
thecae 20—30, + completely sessile, free apices 0.1-0.2 mm, at apex little incurved;
apical cavity narrow to rather broad and deep, reaching to nearly halfway the central
column, 0.4—1 mm deep; androphore narrow, 0.1 mm long. Female flowers: stigma
broadly 2-lobed, 0.2-0.3 mm high. Fruits 1.8—2 by 1.4-1.6 cm.

Field-notes — Bark smooth. Flowers yellow, smelling of Peru-balsam.

Distribution — Malesia: W and C Java.

Habitat & Ecology — Forest at 600-1500 m altitude; fl. throughout the year; fr. June.

Note — Phyllotaxis of all specimens seen is tristichous.

29. Horsfieldia gracilis W. J. de Wilde

Horsfieldia gracilis W.J. de Wilde, Gard. Bull. Sing. 38, 2 ((1985’, 1986) 211; Blumea 41 (1996)
377, f. 1d—g; Tree Fl. Sabah & Sarawak 3 (2000) 371. — Type: S 16604 (Ilias Paie), Sarawak.

Tree c. 5 m. Twigs 1.5—2.5 mm diameter, late glabrescent, hairs light brown, 0.3—0.5
mm: bark striate, neither cracking nor flaking; lenticels minute, inconspicuous or ab-
sent. Leaves membranous, oblong-lanceolate, 12-21 by 4—6(—6.5) cm, base (rounded-)
attenuate, apex acute-acuminate; upper surface drying dull olivaceous, lower surface
pale, olivaceous brownish, with subpersistent scattered pale hairs 0.3(—0.5) mm, or
subglabrous; dots absent; midrib raised above; nerves 12-17 pairs, slightly raised to flat
above, lines of interarching regular, slightly impressed, obvious; venation lax, indis-
tinct; petiole 8-14 by 1.5(-2) mm, late glabrescent, leaf bud 6—7 by 1.5 mm, with dense
pale brown hairs 0.3—0.5 mm long. Inflorescences among the leaves, pubescent, hairs
0.3—0.5 mm long; in male: paniculate, 3-5 cm long, many-flowered; in female: not or
only once branched, 2-5-flowered, 1—1.5 cm long; bracts not seen; flowers (male) in
clusters of 2-6, glabrous, perianth 3-lobed (female) or 4-lobed (male). Male flowers:
pedicel 0.5 mm long, articulated; buds depressed-globose, 0.6 by | mm, cleft slightly
over 1/2: androecium depressed-globose, 0.5 by 0.9—1 mm (Plate 2: 55); thecae 32-36,
central column slightly hollowed-out at apex. Female flowers: pedicel 1 mm long, not
articulated; perianth (from fruit) 2 by 1.5 mm, cleft c. 1/2; pistil not seen. Fruits 1-3 per
infructescence, ellipsoid, apex and base subobtuse, 1.4—1 5 by I-1.1 cm, glabrous, dry-
ing dark brown, finely granulate, lenticel-like tubercles absent; pericarp 0.5—1 mm thick;
fruiting pedicel 1-1.5 mm; perianth persistent. — Fig. 24d-g.

Distribution — Malesia: Borneo (Sarawak: Miri Dist.; Brunei).

Habitat & Ecology — Primary lowland forest.

Note — Related to H. paucinervis, but H. gracilis is distinguishable by its more slen-
der habit, thin membranous leaves and (sub)persistent rough indumentum on leaf bud,
twig apex and lower leaf surface, especially on the midrib and nerves, and by the small
fruits with persistent perianth. Superficially the species may be taken for H. macilenta or
H. tenuifolia. For a more extensive discussion, see De Wilde 1996.

De Wilde — Myristicaceae (Horsfieldia) 123

30. Horsfieldia grandis (Hook. f.) Warb.

Horsfieldia grandis (Hook. f.) Warb., Mon. Myrist. (1897) 301; J. Sinclair, Gard. Bull. Sing. 16
(1958) 400, f. 39; 28 (1975) 48; W.J. de Wilde, Gard. Bull. Sing. 38, 2 ((1985’, 1986) 203; Tree
Fl. Sabah & Sarawak 3 (2000) 372. — Myristica grandis Hook. f., Trans. Linn. Soc. 23 (1860)
157. — Type: Low s.n., Sabah.

Myristica rubiginosa King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 302, pl. 130. — Type: King’s coll.
1233, Singapore.

Tree 6—25 m. Twigs (3—)4-10 mm diameter, late glabrescent, the hairs dense, harsh,
1—1.5 mm long; bark coarsely striate, sometimes slightly cracking, or flaking; lenticels
often present, somewhat elongate, inconspicuous. Leaves membranous or chartaceous,
+ bullate, elliptic-oblong to oblong-oblanceolate, 12—40 by 5—20 cm, base + attenuate to
subcordate, apex acute or acute-acuminate; upper surface drying olivaceous, minutely
paler pustulate, with harsh, persistent sparse hairs, sometimes glabrescent but always
scabrous by harsh hair bases, lower surface with persistent dense to sparse harsh-woolly
hairs 1-2 mm long; dots absent; midrib somewhat raised above, harsh-pubescent; nerves
(8—)10—16(-19) pairs, flattish or usually sunken above, lines of interarching distinct;
venation lax, distinct on both surfaces (especially below); petiole 6-15 by 2.5-6 mm,
pubescent; leaf bud short-conical, 7-15 mm long, densely pubescent. /nflorescences
with dense yellow-rusty hairs, 1.5—3 mm long; in male: many-flowered, 3 or 4 times
branched, rather lax, 6—25 by 2.5—10(—15) cm, peduncle up to 1.2 cm long; in female:
1.5—5 cm long, the buds often of different age; bracts oblong to lanceolate-linear, (1—)3—
12 mm long, caducous; flowers solitary or usually (in male) in loose clusters, often ag-
gregated into compound clusters corresponding to the main ramifications of the inflo-
rescences, glabrous; perianth 3- or 4-lobed, pedicel slender, not articulated. Male flowers:
pedicel (0.5—)1—2 mm long; buds globose or depressed-globose, 1.2—1.8(—2) mm diam-
eter, apex (broadly) rounded, base rounded, cleft c. 1/3 to nearly 1/2, lobes 0.2 mm
thick; androecium depressed globose, not laterally compressed, 0.5—1 by 0.8—1.5 mm
(Plate 2: 49); thecae 16—20, almost completely sessile, incurved towards the apex, col-
umn broad, hollow 0.2 mm deep; androphore narrow, 0.2—0.4 mm long. Female flowers:
pedicel 0.3—0.5 mm long; buds subglobose to broadly ellipsoid, 2—3.2 by 2—2.8 mm,
cleft 1/4—1/3; ovary globose to broadly ellipsoid, 1.5—2 by 1.5—1.8 mm, glabrous, stigma
sessile, faintly 2-lobed, 0.2 mm high. Fruits 2-10 per infructescence, + clustered, obovoid-
ellipsoid, apex rounded, 1—1.4 by 0.8—1.1 cm, glabrous; pericarp 1.5 mm thick, drying
dark brown or reddish brown, without lenticels or warts; seed almost globose, 8—10 mm
diameter; fruiting pedicel | mm long; perianth persistent.

Field-notes — Slender tree, without buttresses; branchlets few; bark usually longitudi-
nally fissured, scaly, flaky, or cracked, strips | mm thick, | cm wide, hard and thin, some-
times smooth, non-flaking; inner bark 2—3 mm thick, slash rich red-brown, with reddish
watery exudate; sapwood soft, whitish to yellowish pink, wood pale brown. Flowers
yellow, with faint odour. Fruits yellowish or + orange; pericarp inside pink; seed grey.

Distribution — Malesia: Sumatra (Palembang; Riau, not seen), Peninsular Malaysia
(Pahang, Johore), Singapore, Borneo (Sarawak, Brunei, Sabah, E Kalimantan).

Habitat & Ecology — Understorey or middlestorey tree of primary or degraded forest
or ridge forest; on clayey soil, sandstone; 0-600 m altitude; fl. & fr. throughout the year.

Note — The perianths of H. grandis are usually 4-, less frequently 3-lobed. There are
16-20 thecae which are tightly appressed and difficult to count.

124 Flora Malesiana, Ser. I, Vol. 14 (2000)

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De Wilde — Myristicaceae (Horsfieldia) 125

31. Horsfieldia hellwigii (Warb.) Warb.

Horsfieldia hellwigii (Warb.) Warb., Mon. Myrist. (1897) 343. — Myristica hellwigii Warb., Bot.
Jahrb. Syst. 18 (1893) 192. — Type: Hellwig 416 (B lost), Finschhafen, Papua New Guinea.

Tree 5—30 m. Twigs subterete or + ridged, often hollow, (4—)5—15 mm diameter, with
rusty woolly or felty hairs 0.5—1(—1.5) mm long, early to late glabrescent; bark striate,
not flaking; lenticels usually distinct. Leaves membranous or chartaceous, elliptic-ob-
long to oblong-lanceolate, sometimes + parallel-sided, 17—40(—50) by 5—14 cm, base
+ rounded or (short-)attenuate, apex acute-acuminate (in New Britain rarely short-cau-
date); upper surface drying dull olivaceous, minutely pustulate, lower surface late gla-
brescent or with hairs of mixed or subequal size 0.3—1 mm long, when shed not leaving
thickened hair bases; dots absent; midrib flat or slightly sunken above, usually with per-
sistent indumentum towards base; nerves 12-33 pairs, + flat above (generally at c. 45°
to the midrib), lines of interarching thin, + irregular beneath; venation lax, faint above;
petiole (2—)5—8 by 2.5—-5 mm, not or hardly winged; leaf bud stout, 3—7 cm long, with
dense hairs 0.5—-1.5 mm long. Inflorescences among the lower leaves, woolly-pubes-
cent, hairs 0.5—1 mm; in male: 2 or 3 times branched, 3.5—15 by 2-10 cm; peduncle up to
3 cm; flowers (male) in clusters of 3—6; in female: up to 8 by 5 cm, flowers in clusters of
up to 4; bracts elliptic to broadly ovate, acute, 3-7 mm long, caducous; flowers with
perianth 2-lobed (not known in var. /ignosa), glabrous or glabrescent except at base,
pedicel with hairs 0.2-0.5 mm, not articulated. Male flowers: pedicel slender, 1—3(-4)
mm; buds somewhat laterally compressed, subcircular, 1.8—3.2 by 2.3—3.5 mm, apex
and base broadly rounded, cleft (1/3—)1/2—2/3, lobes 0.2 mm thick; androecium:
synandrium laterally compressed, (1.2—)1.5—2 by (1.3—)1.7—2 mm, 0.8—1 mm thick, apex
+ broadly rounded-truncate, column narrowly hollowed for the upper 1/4(—1/5), anthers
(10—)12-18 (i.e., with 12-18 thecae at each side), erect, 1.2—-2 mm long, completely
sessile, mutually touching, free apices up to 0.1 mm, not incurved (Plate 2: 38); andro-
phore + broad, up to 0.1 mm long. Female flowers: pedicel 1-2 mm long; buds subglobose
or broadly ovoid, 2.8-3.5 by 3 mm, cleft 1/3—1/2; ovary subglobose, 2 mm diameter,
with dense hairs 0.2—0.4 mm long, stigma subsessile, faintly 2-lobed, 0.1 mm high.
Fruits 3—15 per infructescence, subglobose or broadly ellipsoid to fusiform, apex rounded
or acutish, 1.2—2.8 by 1—1.8 cm, densely pubescent or partly glabrescent, hairs 0.5 mm,
usually finely lenticellate-tuberculate; pericarp 1-3 mm (in var. lignosa 4-8 mm) thick;
fruiting pedicel 1-4 mm long; perianth not persistent. — Fig. 17.

Field-notes — Bole straight, not buttressed; bark finely longitudinally fissured; wood
rather soft and light, whitish or straw, heartwood pinkish. Crown narrow, dense; branches
often tending to be whorled, horizontal, later on drooping; leaves drooping. Flowers yel-
low. Fruits green turning yellow or orange.

Fig. 17. Horsfieldia hellwigii (Warb.) Warb. var. hellwigii. a. Apical part of leafy twig; b. twig with
male inflorescence; c. mature male flower, lateral view; d. ditto, longitudinally opened, showing
androecium; e. twig with female inflorescences axillary to leaf scars; f. female flower at anthesis;
g. ditto, longitudinally opened, showing pubescent ovary and small 2-lobed stigma; h. infructes-
cence with mature fruits [a: NGF 26412; b—d: NGF 4019; e—g: Hoogland 3431; h: LAE 67101). —
Scale bar for a, b, e, h = 2 cm; forc, d, f, g = 1.7 mm.

126 Flora Malesiana, Ser. I, Vol. 14 (2000)

Distribution — Malesia: Papua New Guinea (including New Britain and New Ire-
land); not yet found in Papua Barat.

Habitat & Ecology — Primary, degraded, and secondary forest, including ridge, gal-
lery, and monsoon forest; in primary forest an understorey or second storey tree, in
secondary forest occasionally common; on alluvial soils, also limestone; 0—1200 m alti-
tude; fl. throughout the year; fr. predominantly July—Dec.

Uses — Fruits sometimes recorded as edible.

KEY TO THE VARIETIES

la. Fruits slightly asymmetrically subglobose; pericarp woody, 4—8 mm thick, the sur-

face not wrinklediondrying 242-2... eee ee eee c. var. lignosa

b. Fruits subglobose, ellipsoid, or fusiform; pericarp 2 mm thick, the surface usually
somewhat wrinkled on drying +........5.....--2 sees ee eet tee eee 2

2a. Fruits broadly ellipsoid to nearly globose, 12-15 mm long .. b. var. brachycarpa
b. Fruits broadly ellipsoid to fusiform, 16-28 mm long ......----- a. var. hellwigii

a. var. hellwigii

Horsfieldia hellwigii (Warb.) Warb., Mon. Myrist. (1897) 343; Markgr., Bot. Jahrb. Syst. 67 (1935)
150: A.C. Sm., J. Arnold Arbor. 22 (1941) 61; J. Sinclair, Gard. Bull. Sing. 28 (1975) 49 (for the
type variety only); W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 138, 141, f. 21. — Myristica
hellwigii Warb., Bot. Jahrb. Syst. 18 (1893) 192.

Horsfieldia glabrescens Warb. in K. Schum. & Lauterb., Fl. Schutzgeb. Siidsee (1900) 325.— Type:
Tappenbeck 74, Papua New Guinea, Madang.

Lower leaf surface usually rather densely pubescent with hairs 0.3—1 mm. Male flow-
ers: buds 2.2-3.2 by 2.4—3.5 mm, cleft 1/2—2/3. thecae (24—)28-—36. Fruits broadly
ellipsoid to fusiform, 1.6—2.8 by 1-1.8 cm; pericarp 2—3 mm thick, usually somewhat
wrinkled on drying.

Distribution — As for the species.

Note — NGF 40599 from W New Britain, and LAE 66008 from nearby Umboi I.
slightly differ one from the other, the first is much more hairy. Both deviate from normal
H. hellwigii var. hellwigii by more pear-shaped, 1.e., at base more tapering male buds.
Possibly the specimens are hybrids, e.g., with H. tuberculata, or they may represent a
separate taxon.

b. var. brachycarpa W.J. de Wilde

Horsfieldia hellwigii (Warb.) Warb. var. brachycarpa W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985)
142. — Type: Lauterbach 1191, Papua New Guinea.

Lower leaf surface rather sparsely hairy, the hairs 0.3 mm long. Male flowers: buds 2
mm diameter (1.9 by 2.3 mm), cleft c. 1/3; thecae 20-24. Fruits subglobose to broadly
ellipsoid, 1.2-1.5(-1.7) by 1-1.3 cm; pericarp 1-2 mm thick, usually somewhat wrin-
kled on drying.

Distribution — Malesia: Papua New Guinea (Sepik, Madang, Morobe Prov.).

Habitat & Ecology — Twice collected in levee-forest; 0-100 m altitude.

De Wilde — Myristicaceae (Horsfieldia) 127

c. var. lignosa W.J. de Wilde

Horsfieldia hellwigii (Warb.) Warb. var. lignosa W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 142.
— Type: LAE 56060 (Leach), Papua New Guinea, Milne Bay Prov.

Lower leaf surface rather sparsely hairy, the hairs 0.2-0.3 mm long (on leaf bud 0.5
mm long); flowers unknown. Fruits somewhat asymmetrically subglobose, slightly flat-
tened or not, 1.6—1.9 cm diameter (immature); pericarp very woody, 4-8 mm thick, the
surface not wrinkled on drying, with dense rusty hairs 0.3(—0.5) mm long.

Distribution — Malesia: SE Papua New Guinea (Central and Milne Bay Prov.).

Habitat & Ecology — Lowland and montane forest; 300-1150 m altitude; fr. June,
Sept.

32. Horsfieldia hirtiflora W.J. de Wilde

Horsfieldia hirtiflora W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 1, f. 27. — Type: Rahmat si
Boeea 9257, northern Sumatra.

Tree 10 m. 7wigs subterete, in the apical part + angular and ridged, lower down lined
or ridged, 4—6(—15) mm diameter, glabrescent, hairs 0.3—0.4 mm; bark finely to coarsely
striate, not flaking; lenticels not conspicuous. Leaves in 2 or 3 rows, membranous,
(obovoid-)oblong, 17-30 by 6-11 cm, base + long-cuneate, apex shortly acute-acumi-
nate; upper surface drying brown-olivaceous, lower surface brown, glabrous but midrib
+ late glabrescent; dots absent; midrib raised or flattish above, glabrescent; nerves 15—
20 pairs, above slender, raised, glabrous, lines of interarching + sunken, distinct; vena-
tion lax, indistinct; petiole 15-25 by 2.5-3.5 mm, late glabrescent; leaf bud 8-10 by
4-5 mm, with hairs 0.3—0.4 mm. Inflorescences behind the leaves, with dense rusty hairs
0.3-0.6 mm: in male: 3 or 4 times branched, rather many-flowered, 10-12 by 5—6 cm,
peduncle 3.5-4 cm, flowers in clusters of 3-5; in female: few-flowered, 4 cm long;
bracts elliptic-oblong, acute, 5-10 mm long, densely pubescent, caducous; flowers with
hairs 0.1—0.3 mm long, perianth 3- or 4-lobed, pedicel articulated. Male flowers: pedicel
1.5-2.5 mm long; buds subglobose-obovoid, + collapsing on drying, 2.2—2.5 by 2-2.5
mm, apex broadly rounded, base + rounded, cleft c. 1/2 to nearly 2/3, lobes 0.3-0.4 mm
thick; androecium globose to obovoid, (1.2—)1.5 mm diameter, subcircular or blunt-
angular in cross section (Plate 3: 63); thecae 12 or 14, acutish, 0.7—0.8 mm long, free for
about halfway, somewhat curved towards the centre, column largely hollowed out, the
basal part of the androecium consisting of a tapering androphore 0.5—0.7 mm long. Fe-
male flowers not seen. Fruits 1-3 per infructescence, broadly ellipsoid-obovoid, apex
broadly rounded, base narrowly rounded and short-attenuate, 5—5.7 by 3.8-4.5 cm, gla-
brous, drying dark brown, sparsely coarsely tubercled: pericarp 10(—15) mm thick; fruit-
ing pedicel 2-3 mm long, pubescent; perianth not persistent. — Fig. 18.

Field-notes — Tree 10 m tall, diameter 12 cm; bark rough, hard, black. Fruits yellow,
subglobose, 8 cm diameter.

Distribution — Malesia: N Sumatra (Tapanuli, E Coast).

Habitat & Ecology — Rain forest; on sandstone; 100-500 m altitude; fl. June—July;
fe. Dec.

Notes — 1. Horsfieldia hirtiflora is related to H. brachiata with which it has the dis-
tinctly ridged twigs in common, but H. hirtiflora differs from H. brachiata in the coarser
habit, larger and hairy flowers, and large fruits.

128 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Horsfieldia) 129

2. In some specimens the leaves towards the apex of the twigs are not distichous, but
+ arranged in three rows.

33. Horsfieldia inflexa W.J. de Wilde

Horsfieldia inflexa W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 63, f. 8. — Type: LAE 52866
(Streimann & Martin), Papua New Guinea.

Tree 10-21 m. Twigs ridged and angular, especially in the apical part, 2—5(—7) mm
diameter, early glabrescent, hairs grey-brown to 0.1 mm; bark faintly striate, not flak-
ing; lenticels abundant, inconspicuous. Leaves thinly chartaceous, (elliptic-)oblong, 8—
20 by 2.5—7.5 cm, base attenuate, apex bluntish to acute-acuminate; upper surface dry-
ing olivaceous to blackish brown, sometimes indistinctly pale-pustulate, lower surface
early glabrescent; dots present (lens!); midrib slender, slightly raised above; nerves 8—
13 pairs, thin, flat, indistinct, lines of interarching beneath fairly regular but indistinct;
venation lax, indistinct on both surfaces; petiole 14-30 by 1.5—2.5 mm; leaf bud 10-14
by 1.5—2 mm, with hairs 0.1 mm long or less. Inflorescences glabrescent or with scat-
tered scale-like hairs less than 0.1 mm; in male: 2 (or 3) times branched, 3—10 by 1.5—4.5
cm, the primary branches + spike-like; in female: 3-5 cm long; peduncle 0.1—1.2 cm
long; bracts elliptic, 1-3 mm long, with fimbriate margin, caducous; flowers solitary or
up to 4 together, glabrous; perianth 2-lobed, pedicel not articulated. Male flowers: pedi-
cel 1-2 mm long; buds subglobose, slightly broader than long, not or but little laterally
compressed, together with the pedicel sometimes slightly pear-shaped, 2.2—2.5 by 2.5—3
mm, apex broadly rounded, basal part rounded or shortly tapering, cleft 9/10, lobes 0.2—
0.5 mm thick, not collapsing on drying; androecium blunt-quadrangular in outline, some-
times broader than long, laterally compressed, 1.5 by 1.5—2 mm, apex broadly rounded
(Plate 1: 4); thecae 20—24(—28?), 1.5—2 mm long, free apices long, 0.7—1 mm, at one side
of the androecium strongly incurved into the central hollow, almost reaching the base;
androphore up to 0.1 mm long. Female flowers: pedicel 1—1.5 mm long; buds ovoid-
ellipsoid, 2—2.5 by 2 mm, cleft c. 2/3; ovary ovoid, 1.5 by 1.2 mm, glabrous, stigma
sessile, minutely 2-lobed, 0.2 mm. Fruits solitary or up to 6 per infructescence, ellip-
soid, apex rounded to subacute, base + rounded, 2 by 1.4—1.5 cm, glabrous, drying dark
brown, without or with only small tubercles or lenticels; pericarp 1.5 mm thick; fruiting
pedicel 2-6 mm long; perianth not persistent. — Fig. 19.

Field-notes — Slender tree, buttresses absent; bark shallowly longitudinally fissured,
not or slightly peeling off; slash reddish brown, sapwood whitish red or cream, heart-
wood not differentiated. Flowers green, turning yellow, fragrant. Fruits yellowish or-
ange.

Distribution — Malesia: northern part of New Guinea: Papua Barat (Bird’s Head,
Geelvink Bay, Jayapura); northern Papua New Guinea (West Sepik Prov.).

Fig. 18. Horsfieldia hirtiflora W.J. de Wilde. a. Leafy twig with immature male inflorescence; note
lined twig, dispersed leaves, and bracts present in inflorescence; b. submature male flower, lateral
view; c. ditto, opened, showing androecium; d. immature androecium, longitudinal section, sche-
matic; e. twig with infructescence, fruit mature [a—d: Rahmat si Boeea 9257; e: Kostermans 22048}.
— Scale bar for a, e = 2 cm; for b—d = 0.85 mm.

130 Flora Malesiana, Ser. I, Vol. 14 (2000)

) Jugs

ib
IL

{

»

Fig. 19. Horsfieldia inflexa W.J. de Wilde. a. Twig with male inflorescences; note lined twig:
b. opened mature male flower showing androecium; c. androecium, longitudinal section, schematic;
d. twig with female inflorescences axillary to fallen leaves; e. mature female flower, lateral view;
f. ditto, opened, showing glabrous ovary with broad-lipped stigma; g. twig with infructescences,
h. part of lower leaf surface with scattered dark-coloured non-traumatic cork warts as blackish dots
[a—c: LAE 52866; d—f: van Royen 3166; g, h: LAE 52862]. — Scale bar for a, d, g = 2 cm; for b, c,
e, f = 0.85 mm; for h = 1.8 mm.

De Wilde — Myristicaceae (Horsfieldia) 131

Habitat & Ecology — Primary and old degraded forest on alluvial soils, e.g., sandy
clay; also in hilly forest and swamp forest; 0-400 m altitude; fl. throughout the year;
fr. Sept.—Nov.

Notes — 1. This is the only Horsfieldia in New Guinea with dark dots on the lower
leaf surface.

2. Horsfieldia inflexa resembles H. moluccana in general shape and texture of the
leaves, and the relatively long petioles. It can also be confused with the related species
H. angularis and H. basifissa. Horsfieldia moluccana generally has more pear-shaped
male buds and differs from H. inflexa in terete or faintly lined twigs and not dotted
leaves. Horsfieldia angularis is distinguishable by not dotted leaves, hairy flowers, an
androecium with a narrow central crevice and straight anthers. Horsfieldia basifissa has,
in contrast, terete or only faintly ridged twigs, and differs further in various characters of
the male flower including the androecium.

3. Some specimens of H. inflexa from Japen I. resemble H. parviflora by rather
unbranched spike-like inflorescences; H. parviflora has more pronounced pear-shaped
flowers and paler, not angular twigs. In most of the specimens of H. inflexa, the male
inflorescences are distinctly branched with the lateral branches almost unbranched and
spike-like.

34. Horsfieldia iriana W.J. de Wilde

Horsfieldia iriana W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 99. —Type: Zippelius (139d), West
New Guinea.

Horsfieldia novo-guineensis Warb., Mon. Myrist. (1897) 271, p.p., only the type of Horsfieldia
iriana, not the lectotype, which is H. aruana.

Myristica nesophila auct. non Migq.: Miq., Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 206; 2 (1865) 49,
p.p., as based on Zippelius (139d), not the lectotype of Myristica aruana Blume = Hors-
fieldia aruana.

Tree c. 10 m. Twigs 2-angular, lower down ridged, 3—5 mm diameter, early glabrescent,
hairs grey-rusty, 0.2 mm long; bark rather smooth, not flaking; lenticels small, distinct.
Leaves thinly coriaceous, oblong-lanceolate, 17—28 by 4.5—8 cm, base long-attenuate,
apex acute-acuminate; upper surface drying olivaceous, lower surface early glabrescent,
at first with stellate-scaly hairs 0.1-0.2 mm long; dots absent; midrib broad and flat
above; nerves 13-16 pairs, thin, flat above, lines of interarching indistinct; venation lax,
slightly raised, indistinct above; petiole 6-11 by 2—3.5 mm; leaf bud 10-15 by 1.5-2
mm, with hairs 0.1—0.2 mm long. Inflorescences in male: among the leaves, with sparse
stellate hairs 0.2(—0.3) mm long, (2 or) 3 (or 4) times branched, rather many-flowered,
6-8 by 4-6 cm, peduncle 0.5—1 cm long; bracts not seen, caducous; flowers in loose
clusters of 2-5, perianth 2-lobed, glabrous except some minute hairs towards the base;
pedicel sparsely pubescent, not articulated. Male flowers: pedicel not tapered, slender,
2.5—3 mm long; buds in lateral view circular to slightly longitudinally ellipsoid, not or
slightly laterally compressed, blackish brown, not collapsing on drying, 2.3—2.8 by 2.2—
2.8 mm, apex subacute or narrowly rounded, base rounded, cleft 3/4—5/6, lobes 0.2 mm
thick; androecium laterally compressed, 1.5 by 1.5—2 mm, apex (broadly) rounded (Plate
1: 22); thecae 20-28, erect, free apices 0.1(—0.2) mm long, column narrowly hollowed
for c. 1/4; androphore absent. Female flowers and fruits not seen.

132 Flora Malesiana, Ser. I, Vol. 14 (2000)

Distribution — Malesia: SW Papua Barat, known only from the type.

Habitat & Ecology — Possibly from coastal lowland forest.

Note — The only specimen known, Zippelius (139d), is part of the heterogeneous
Zippelius material which served for the description of Myristica aruana Blume and
Horsfieldia novo-guineensis Warb. For details see de Wilde (1985).

35. Horsfieldia irya (Gaertn.) Warb.

Horsfieldia irya (Gaertn.) Warb., Mon. Myrist. (1897) 317 (incl. var. or forma ceylanica, javanica,
malayana, moluccana, siamensis, wallichii), t. 22; J. Sinclair, Gard. Bull. Sing. 16 (1958) 382,
f. 33, pl. IX-A; 28 (1975) 61; Backer & Bakh. f., Fl. Java 1 (1964) 138; W.J. de Wilde, Gard.
Bull. Sing. 38, 1 (1985) 55, f. 6; Tree Fl. Sabah & Sarawak 3 (2000) 373. — Myristica irya
Gaertn., Fruct. 1 (1788) 195, t. 41; Hook. f. & Thomson, FI. Ind. (1855) 159; A. DC., Prodr. 14,
1 (1856) 202 (excl. M. exaltata, p.p., see under Endocomia, Blumea 30 (1984) 173); King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 308, pl. 141, 141-bis. — Type: Gaertner’s drawing, Sri Lanka.

Myristica javanica Blume, Bijdr. (1826) 576; Rumphia | (1835) 190, t. 62. — Type: Blume’s au-
thentic specimens not found in L [Rumphia, t. 62, male fl., fr.].

Myristica spherocarpa Wall., Pl. As. Rar. (1830) 79, t. 89. — Myristica irya Gaertn. var. wallichii
King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 309, pl. 141-bis 3-5. — Type: Wallich Cat. n. 6796.

Myristica micrantha Wall., Cat. (1832) n. 6807, nom. nud. — Type: Wallich Cat. n. 6807.

Myristica lemanniana A.DC., Ann. Sc. Nat. Bot. 4, 4 (1855) 31, t. 4; Prodr. 14, 1 (1856) 203. —
Horsfieldia lemanniana (A. DC.) Warb., Mon. Myrist. (1897) 326. — Type: Lemanns.n., Penin-
sular Malaysia, Malacca.

Myristica subglobosa Miq., Fl. Ind. Bat., Suppl. 1 (1861) 383. — Myristica globularia Blume vat.
subglobosa (Miq.) Migq., Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 206. — Horsfieldia subglobosa
(Mig.) Warb., Mon. Myrist. (1897) 328 (for the original syntype only). — Syntypes: Diepenhorst
HB 2148, Teijsmann HB 3189, both Sumatra.

Myristica vrieseana Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 49. — Myristica irya Gaertn. var.
longifolia King, Ann. Roy. Bot. Gard. Cale. 3 (1891) 309, pl. 141-bis 1 & 2. — Type: de Vriese
sii).

Horsfieldia labillardieri Warb., Mon. Myrist. (1897) 283, t. 21. — Myristica labillardieri (Warb.)
Boerl., Handl. 3 (1900) 85. — Type: Labillardiére s.n., male fl. (B lost; iso FI, see note by
Sinclair 1975: 89), Java.

Horsfieldia acuminata Merr., Philipp. J. Sci. 17 11920", 1921) 253; Enum. Philipp. Flow. Pl. 2
(1923) 181. — Type: FB 27507 (de Mesa), Philippines.

Horsfieldia nunu Kaneh., Trop. Woods 29 (1932) 5, nom. nud.; Bot. Mag. Tokyo 46 (1932) 451; Fl.
Micr. (1933) f. 32; Enum. Micron. Plants, in J. Dept. Kyushu Imp. Univ. 4, 6 (1935) 319. —
Syntypes: Kanehira 1303, 1304, Pacific area.

Horsfieldia amklaal Kaneh., Bot. Mag. Tokyo 47 (1933) 670; FI. Micr. ( 1933) 109, £. 31, pl. 16:—
Syntypes: Kanehira 1944, 1978, 2058, 2059, Pacific area.

Horsfieldia congestiflora A.C.Sm., J. Arnold Arbor. 22 ( 1941) 64.— Type: Brass 8010, New Guinea.

Fig. 20. Horsfieldia irya (Gaertn.) Warb. a. Leafy twig apex; note whitish blotched leaves; b. twig,
ridged, with male inflorescence; c. mature male flower bud, lateral view; d. male flower, longitudi-
nal section, showing androecium; e. androecium, longitudinal section, schematic; f. twig with fe-
male inflorescence; g. mature female flower; h. ditto, opened, showing glabrous ovary with minute
2-lobed stigma; i. twig with infructescence; note spherical fruits [a, b: NGF 22319; c—e: Kostermans
24385: fi: FRI 3044]. — Scale bar for a, b, f, i= 2 cm; for g = 1.7 mm; for c, h= 0.85 mm; d, e =
0.4 mm

De Wilde — Myristicaceae (Horsfieldia) 133

134 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 10—25(—40) m. Twigs towards apex often drying flattened, usually thinly ridged,
(2-)3—10 mm diameter, glabrescent, indumentum minute to conspicuous (New Guinea
and Pacific Islands), grey to rusty, dendroid hairs of mixed size, 0.1—0.5(-1) mm long;
bark (coarsely) striate, + blackish, not flaking; lenticels conspicuous. Leaves often
+ curved towards the apex, membranous, elliptic-oblong to lanceolate, 10—30(—35) by
3-7(-9) cm, base rounded to attenuate, apex acute-acuminate; upper surface drying dull,
greenish to blackish, almost always with irregular whitish marks of unknown origin,
lower surface early glabrescent; dots absent; midrib flattish above; nerves 10-20 pairs,
thin and flattish above, lines of interarching usually indistinct; venation lax, faint above,
thin though distinct beneath; petiole 7-16 by 1.5—3(—4) mm; leaf bud 10(—15) by 2-3
mm, with hairs 0.1—0.5(—1) mm. Inflorescences with dense hairs 0.1—0.5(-1) mm long,
persistent or glabrescent; in male: 3 or 4 times branched, many-flowered, 4-18 by 3-7
(10) cm; in female: 2-6(—8) cm long, 2 (or 3) times branched; peduncle 0.54.5 cm
long; bracts + acute, 1.5—4 mm long (sometimes larger, leaf-like), caducous; flowers in
male in clusters of 3-10, in female fewer; perianth 2-lobed, glabrous, at base glabrescent;
pedicel pubescent or glabrescent, not articulated. Male flowers: pedicel slender, 0.1-1.5
mm; buds subglobose or + transversely ellipsoid, somewhat laterally compressed or not,
1-1.5 by 1-1.5 mm (largest in Indochina), apical part broadly rounded, base rounded
or short-tapering, cleft 1/2-2/3, lobes 0.2 mm thick; androecium + broadly obovoid,
0.8-1.2 by 1-1.5 mm (Plate 1: 2); thecae 12-18(—20, Indochina), not closely touching,
0.5—0.8(-1) mm long, free apical part 0.2-0.3 mm, incurved, column and androphore
broadly concave, more or less saucer- or cup-shaped, 0.4—0.5 by (0.5—)0.6—1 mm,
tapered towards the base. Female flowers: pedicel 1-4 mm long; buds obovoid or ellip-
soid, 1.5—2.3 by 1.3-2 mm, cleft 1/4-1/3; ovary broadly obovoid, glabrous, 1.2—1.5 by
1-1.3 mm, stigma minute, 0.1 mm. Fruits 2-8 per infructescence, globose, 1.5—2.2 cm
diameter, glabrous, finely granular, without tubercles or lenticels, drying dark brown to
blackish; pericarp 1-2 mm thick; seeds globose; fruiting pedicel 5—10 mm long; peri-
anth not persistent. — Fig. 20.

Field-notes — Tall tree with straight bole; crown with several big limbs, or narrow
and with slender drooping branches near the apex; bole often fluted or with prop-roots,
or usually with buttresses up to 3 m high, 2 m out, and up to 10 cm thick, sometimes
without buttresses: the bark often fissured, cracked, or mostly flaking or peeling off in
small pieces, sometimes smooth; inner bark whitish, to 7 mm thick; sapwood whitish or
pinkish; heartwood absent or only slightly darker; the wood rather soft. Flowers (dark)
(orange-)yellow, once reddish; strongly sweet-scented, or without scent. Fruits yellow-
ish or red; up to nearly 3 cm diameter.

Distribution —Sri Lanka, Burma, Andaman and Nicobar Islands, S Indochina (Cochin-
china), Cambodia, Thailand, Caroline Is., Solomon Is.; in Malesia: Sumatra, Peninsular
Malaysia, Singapore, Borneo, Java, Sulawesi, Moluccas, C(?) & S Philippines, New
Guinea: no collections seen from the Lesser Sunda Islands and N Philippines.

Habitat & Ecology — Primary and (old) secondary forest; most frequent in (periodi-
cally) swampy coastal or riverine areas, on alluvial (sandy, loamy, or clayey) soils, but
also found more inland: 0—450 m altitude; fl. & fr. apparently throughout the year, but
probably this is correlated with regional and local climatic conditions. The seeds contain
an air chamber, facilitating dispersal by floating.

Uses — Rarely recorded that fruits are edible; fruits eaten by monkeys in Sri Lanka.

De Wilde — Myristicaceae (Horsfieldia) 135

Note — Horsfieldia irya is a homogeneous species, well characterized by its small
subglobose male flowers of 1 mm diameter (in Indochina up to 1.6 mm diam.), with
tyical broad and deeply concave androecium with tapered, relatively large androphore.
The fruits (and seeds) are globose, glabrous (ovary glabrous). The twigs are usually thin-
ly ridged. Characteristic are the irregular whitish marks of unknown origin, almost al-
ways present on the older leaves in dried specimens. The indumentum 1s variable: short-
haired, sometimes seemingly glabrous specimens are predominant in Sri Lanka, SE Asia,
W Malesia, and Moluccas; in New Guinea and the Solomons most specimens have con-
spicuous, often woolly hairs to 1 mm long on twig apex, leaf bud, and inflorescences.

36. Horsfieldia iryaghedhi (Gaertn.) Warb.

Horsfieldia iryaghedhi (Gaertn.) Warb., Mon. Myrist. (1897) 332, t. 21 f. 1-4; J. Sinclair, Gard.
Bull. Sing. 28 (1975) 68; W.J. de Wilde, Gard. Bull. Sing. 37, 2 1984’, 1985) 167, f. LA-1, 2-1,
3. — Myristica iryaghedhi Gaertn., Fruct. 1 (1788) 196, t. 41, f. 4. — Type: Gaertner’s drawing.

Myristica glomerata Thunb., Act. Holm. sive Kongl. Vet. Acad. Nya Handl. (1799) 88, t. 2 f. 1 [non
Mig. (1852)]. — Type: Herb. Thunberg, fragment of male inflorescence.

Horsfieldia odorata Willd., Sp. Pl. 4 (1805) 872. — Type: not known (see Sinclair 1975).

Myristica horsfieldii Blume, Bijdr. (1826) 577, ‘horsfieldia’; Rumphia | (1837) 192, t. 63; King,
Ann. Roy. Bot. Gard. Calc. 3 (1891) 296, pl. 122, 123. — Pyrrhosa horsfieldii (Blume) Hassk.,
Cat. Pl. Hort. Bog. (1844) 174. — Type: Blume s.n., Java.

Myristica notha auct. non Wall.: Koenig ex Blume, Rumphia | (1837) 192, nom. nud. pro syn.

Myristica (‘Cnema’) glomerata Miq., Pl. Jungh. (1852) 170 [non Thunb. (1799)]. — Type: Junghuhn
s.n., Java.

Myristica odorata Reinw. ex de Vriese, Pl. Ind. Bat. Or. 2 (1857) 95 [non Willd. (1805)], nom. nud.
pro syn.

For more references, see J. Sinclair, l.c..

Tree 5-25 m. Twigs 2-5 mm diameter, early or late glabrescent, at first with woolly
rust-yellow hairs 0.5 mm; bark coarsely striate or longitudinally cracking, rarely some-
what flaking; lenticels present, sometimes inconspicuous. Leaves chartaceous, ovate-
elliptic to oblong-lanceolate, 10-28 by 4-12 cm, base rounded to attenuate, apex acute-
acuminate, upper surface glabrous, drying dark brown or olivaceous, lower surface early
or late glabrescent, papillose; dots absent; midrib late glabrescent, slightly raised above;
nerves 9-16 pairs, flat or sunken above, lines of interarching fairly distinct; venation
+ trabeculate, distinct to hardly visible above; petiole late glabrescent, 15—20(—25) by 2—
3 mm; leaf bud 10-15 by 3-4 mm, with dense hairs 0.5 mm long. Inflorescences with
dense woolly hairs 0.3—0.5 mm; in male: 6-15 by 4-10 cm, usually twice branched, the
branches few, thickish, peduncle 0.3—2 cm; flowers in 10-25 subglobose dense clusters
5-10 mm diameter, spaced along the branches, each cluster with 80—100 (sub)sessile
flowers; in female: smaller, little-branched, 1.5—4 by 1.5—2 cm, the flowers solitary or
few together; bracts subtriangular to elliptic, the larger ones with distinct midnerve, 1-3
mm long, pubescent, caducous; flowers subsessile, perianth in male 3- (or 4-)lobed, in
female 3- (or 2-)lobed, glabrous except at the base (hairs 0.2-0.5 mm), pedicel pubes-
cent, not articulated. Male flowers: pedicel 0.2-0.3 mm long; buds narrowly + obovoid-
oblong or obconical, 3—5S- (or 6-)angular, 22.5 by 1-1.6 mm, apex broadly rounded,
gradually tapering to the base, cleft (1/4-)3/4, lobes 0.2(—0.3) mm thick; androecium
elongate, narrowly obconical or ellipsoid-oblong, apex + truncate, base tapering, 1—-1.2

Flora Malesiana, Ser. I, Vol. 14 (2000)

3

De Wilde — Myristicaceae (Horsfieldia) 137

by 0.4-0.5 mm (Plate 1: /); thecae 6-10, largely sessile, erect, free apices 0.2 mm, col-
umn narrow, hollow to halfway, androphore narrow, 0.2—0.5 mm long. Female flowers:
pedicel up to 0.2 mm long; buds broadly ellipsoid, 3 by 2 mm, cleft c. 1/2; ovary broadly
ellipsoid, 2 by 1.5 mm, densely short-pubescent, stigma minutely 2—4-(or more-)lobu-
late. Fruits 3-8 in a cluster, ellipsoid or ellipsoid-oblong, 2.5—4.2 by 1.7—2.4 cm, with
dense rust-yellowish stellate-dendroid hairs 0.5 mm, sometimes partly glabrescent;
pericarp 1.5—2(—3) mm thick, not tuberculate; fruiting pedicel 0.5-2 mm long; perianth
not persistent. — Fig. 21.

Field-notes — Tree at base to 50 cm diameter, branches drooping, sometimes branched
from the base; stilt-roots sometimes present when growing in wet soil; bark hard, outer
bark light- or black-brown, or greyish red, fissured, gritty, or peeling off in large pieces,
or smooth; living bark 5-10 mm, beefy red or red brown, with white lines, exuding a
clear light reddish brown sap; wood white to light yellow with red streaks. Leaves grey-
green beneath. Perianth (dark) yellow, or orange-brown; pistil with brown hairs. Male
flowers 3 mm long, strongly scented, reminiscent of Michelia champaca. Fruits yel-
low(-brown) with rusty indumentum.

Distribution — Sri Lanka; in Malesia introduced in gardens in Peninsular Malaysia
(Penang I.), Singapore, and Java.

Habitat & Ecology — In Sri Lanka in lowland rain forest, wet evergreen or interme-
diate forest; also in disturbed forest; 0-500 m altitude; fl. & fr. throughout the year.

Uses — Formerly in Java a wax was obtained after cooking the fruits. The wood is
moderately heavy and even-grained (see Sinclair 1975: 72).

Note — Horsfieldia iryaghedhi deviates from all other Horsfieldias, and can be placed
in a separate section. It is distinguishable by the leaves being papillate beneath, the male
flowers arranged in compact heads, and a many-lobulate stigma; according to Warburg
(1.c.: 334) the seeds contain some starch, possibly unique for this species.

37. Horsfieldia laevigata (Blume) Warb.

Horsfieldia laevigata (Blume) Warb., Mon. Myrist. (1897) 351, t. 21 (excl. specimens from Java);
W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 115, f. 17 a-i. — Myristica laevigata Blume,
Rumphia 1| (1837) 191, t. 64 f. 3, anal. 1-4; A. DC., Prodr. 14, 1 (1856) 202; Migq., Fl. Ind. Bat.
1, 2 (1858) 65, p.p. — Type: Commerson (s.n.) 238, female, cult., Mauritius.

For more references and synonyms see the varieties.

Tree 4—25 m. Twigs faintly ridged or not, 1.5—5 mm diameter, early glabrescent, hairs
grey to brown, 0.1—0.2 mm; bark striate, not flaking; lenticels conspicuous or not. Leaves
membranous or thin-chartaceous, elliptic to oblong-lanceolate, 10—30 by (3—)4—12 cm,
base attenuate, apex acute-acuminate; upper surface drying dull olivaceous to brown,

Fig. 21. Horsfieldia iryaghedhi (Gaertn.) Warb. a. Habit of twig with leafy shoot and female inflo-
rescences; b. twig with male inflorescence; c. subspherical male flowerhead, flowers partly re-
moved, one flower opened; d. mature male flower bud, lateral view; e. ditto, longitudinal section
showing androecium; f. mature female flower bud; g. ditto, opened, showing pubescent ovary
and sessile, minutely lobulate stigma; h. infructescence; i & j. seeds with and without seed coat
[a, f, g: Anonymous, Hort. Bot. sub XIII-E-9; b—e: Jayasurya & Bandaranaike 1869; h—}: Koster-
mans 26670). — Scale bar for a, b, h—j = 2 cm; for c = 3.3 mm; for d—g = 0.85 mm.

Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Horsfieldia) 139

usually minutely pale pustulate or irregularly blotched (as in H. irya), lower surface
(largely) glabrescent, hairs 0.1—0.2 mm; dots absent; midrib + flat above; nerves (10—)
12-30 pairs, slightly raised above, lines of interarching irregular, faint beneath; venation
lax, faint or distinct on both surfaces; petiole 5—15 by 1.5—3 mm; leaf bud 10-15 by 1.5—
2 mm, with hairs 0.1—0.2 mm. Inflorescences subglabrescent or with dense or sparse
scale-like stellate hairs 0.1—0.2(—0.5) mm; in male: 2—4 times branched, many-flowered,
5-20 by 3-10 cm, peduncle 1—4 cm; in female: 2-10 cm long; bracts 2-3 mm long,
caducous; flowers (male) in loose clusters of 2—5; perianth 2-lobed, with sparse or dense
hairs 0.1—0.2 mm long, pedicel not tapering, generally pubescent, not articulated. Male
flowers: pedicel slender, 1.5—-3(—4) mm long; buds flattened, subcircular (sometimes
slightly longer than broad, or broader than long, e. g., in some specimens from the Papuan
Islands; see note 2), 1.7—2.8(—3) by 1.7—3(-3.3) mm, cleft 1/2—3/4(-4/5), lobes (0.1—)
0.2-0.3 mm thick; androecium flattened, quadrangular to + reniform in outline, apex
rounded or subtruncate, 1.1—1.5 by 1.1—1.8(—2.2) mm (Plate 1: 29); thecae 18-32, erect,
free apex up to 0.2 or 0.40.6 (var. novobritannica) mm long, column narrowly hollow-
ed for 1/4(—1/2) or in var. novobritannica for c. 9/10; androphore to 0.1(—0.2) mm, broadly
attached. Female flowers: pedicel 2—2.5 mm long; buds broadly ellipsoid to globose,
2.5-3.1 by 2.8-3.1 mm, cleft 1/3—2/3; ovary ovoid or subglobose, 2—2.3 by 1.7—2.2 mm,
hairs 0.1 mm, style to 0.3 mm long, stigma hardly 2-lobed, 0.1—0.2 mm. Fruits (1—-) 2-15
per infructescence, ellipsoid or nearly globose, apex rounded to acutish, base rounded,
(1.6-)1.8—2.8(—3) by 1.4-2(—2.2) cm, glabrescent but always with minute hairs (0.1 mm)
remaining at least at base (lens!), usually with coarse pale tubercles or lenticels; pericarp
2-3 mm thick (or 4—6 mm in some forms from SW New Guinea and New Britain); seeds
ellipsoid; fruiting pedicel 3-6 mm long; perianth not persistent. — Fig. 22.

Distribution — Malesia: Moluccas, New Guinea (including Papuan Islands and Bis-
marck Archipelago); see further under the varieties.

Note — A variable, complex species; one prominent form is segregated as a variety.

KEY TO THE VARIETIES

la. Hairs on inflorescences 0.1—0.2(—0.3) mm long, sometimes almost absent. Anthers
at apex free for only 0.1-0.2 mm; the column hollowed for 1/4(—1/2, or slightly
deeper). Fruits generally ellipsoid, pericarp 2-3 mm, rarely (SW New Guinea) 4-6
mm thick. — Moluccas, New Guinea, including New Britain ... a. var. laevigata

b. Hairs of inflorescences more woolly, 0.3—0.5 mm long. Anthers at apex free for 0.4—
0.6 mm; the column hollowed for c. 9/10. Fruits generally subglobose or short-ellip-
soid, pericarp 2-5 mm thick. — New Britain ........... b. var. novobritannica

Fig. 22. Horsfieldia laevigata (Blume) Warb. var. laevigata. a. Twig apex with leaves; b. opened
mature male flower, showing androecium; c. androecium, longitudinal section, schematic; d. twig
with female inflorescence; e. opened female flower, with finely pubescent ovary and minute 2-lobed
stigma. — H. laevigata var. novobritannica (J. Sinclair) W.J. de Wilde. f. Twig with male inflores-
cence; g. opened mature male flower, showing androecium; h. androecium, longitudinal section,
schematic; i. infructescence with mature fruits. — H. pilifera Markgr. j}. Opened male flower, show-
ing androecium [a—c: LAE 52086; d,e: Commerson (s.n.) 238, f-h: Floyd 6430; i: NGF 10811 (White);
j: Ledermann 6675]. — Scale bar for a, d, f, i = 2 cm; for b, e, j = 1.7 mm; for c, g, h = 0.85 mm.

140 Flora Malesiana, Ser. I, Vol. 14 (2000)

a. var. laevigata

Myristica nesophila Miq., Ann. Mus. Bot. Lugd.-Bat. | (1864) 206, p.p. (excluding specimens from
Batjan). — Horsfieldia nesophila (Miq.) Warb., Mon. Myrist. (1897) 281, t. 21. — Lectotype:
de Vriese s.n. (male) (L), Seram.

Horsfieldia polyantha Warb., Mon. Myrist. (1897) 281, t. 23: J. Sinclair, Gard. Bull. Sing. 28 (1975)
95 (for the greater part, incl. type). — Myristica polyantha (Warb.) Boerl., Handl. 3 (1900) 85.
— Type: Beccari s.n. (Acc. Nos. 7619, 7619-A), Aru Is., Wokam.

Twigs 2-5(-9) mm diameter. Leaves 10-30 by 4-12 cm. Inflorescences with dense to
sparse hairs 0.1—0.2(—0.3) mm, sometimes almost glabrous. Male flowers: buds 1.7—3.3
mm diameter, cleft 1/2—3/4(—5/6); thecae (18—)20-32, free apex (0—)0.1—0.2 mm; col-
umn hollowed for 1/4(—1/2, or slightly deeper). Infructescences up to 10 cm long. Fruits
ellipsoid, 18-28 mm long, drying blackish or brown, usually with coarse wart-like
lenticels; pericarp 2-3(—6) mm thick. — Fig. 22a-e.

Field-notes — Bole straight, without buttresses; bark often shallowly vertically fissured,
not peeling off; branches horizontal, or drooping; wood whitish. Flowers greenish or
brownish yellow. Fruits yellow to orange.

Distribution — As the species.

Habitat & Ecology — Primary and degraded rain forest on ridges and plains, riverine
forest, swampy scrub and forest, edges of sago-swamps; on a great variety of soils,
including black volcanic soil (Moluccas), 0—1000(—1300) m altitude; fl. & fr. through-
out the year.

Uses — Fruits edible; wood of medium weight and hardness; used for house con-
struction.

Notes — 1. After the separation of var. novobritannica the remaining type variety is
still very variable, especially in the leaf texture and the length of hairs on the leaf buds,
and in the shape of the male buds.

2. The width of the male bud varies between 1.7 and 3.3 mm. Specimens with smaller
flowers have been placed in H. pilifera. The shape of the bud in lateral view is generally
subcircular. Certain specimens, e.g. from the Papuan Islands and Gulf Prov., may have a
bud which is rather markedly broader than long, and these flowers may resemble those
of, e.g., H. spicata. In specimens from the Moluccas the bud is often slightly longer than
broad. The degree to which the male bud opens is in most specimens halfway to 2/3,
specimens from the Papuan Islands and the Bismarck Archipelago may have male buds
cleft as deep as 4/5—5/6, a feature reminiscent of H. spicata. In these broad, deeply cleft
male flowers, the androecium is broad, rather reniform as seen laterally.

3. The fruits of some specimens from SW New Guinea have thick corky pericarps,
4—6 mm thick, but the leaves agree with those of var. laevigata. Forms with larger and
thick-walled fruits appear difficult to separate from H. pachycarpa, see the notes under
that species.

b. var. novobritannica (J. Sinclair) W. J. de Wilde

Horsfieldia laevigata (Blume) Warb. vat. novobritannica (J. Sinclair) W.J. de Wilde, Gard. Bull.
Sing. 38, 1 (1985) 120, f. 17 fa. — Horsfieldia hellwigii (Warb.) Warb. var. novobritannica J.
Sinclair, Gard. Bull. Sing. 28 (1975) 54. — Type: NGF 6430 (Floyd), Papua New Guinea.

De Wilde — Myristicaceae (Horsfieldia) 14]

Horsfieldia novae-lauenburgiae Warb., Mon. Myrist. (1897) 278; K.Schum., Notizbl. Bot. Gart.
Berlin 2 (1898) 117; K.Schum. & Lauterb., Fl. Schutzgeb. Siidsee (1900) 324; Markgr., Bot.
Jahrb. Syst. 67 (1935) 151. — Type: Warburg 20713 (B lost; G iso, n.v.; identity not sure),
Bismarck Archipelago, Neu Lauenburg Group, Ulu Is.

Horsfieldia ralunensis auct. non Warb.: Kaneh. & Hatus., Bot. Mag. Tokyo 52 (1938) 355 (speci-
men Kanehira 3969, n.v.).

Twigs 2-6(—8) mm diameter. Leaves (12—)17-30 by (3—)5—8 cm. Inflorescences with
rather dense woolly indumentum of hairs 0.3—0.5 mm long. Male flowers: buds 1.8—2
by 2.2 mm, cleft c. 4/5; thecae c. 28, almost completely filling the perianth, free apices
0.4—0.6 mm; column hollow for c. 9/10. Infructescences up to 8 cm long. Fruits broadly
ellipsoid to almost globose, 18-22 by 16-20 mm, drying grey-brown, with coarse wart-
like lenticels; pericarp rather hard, 2—3(—5) mm thick. — Fig. 22 f-i.

Field-notes — Fruits globose, golden brown, yellow, or orange when mature.

Distribution — Malesia: Papua New Guinea (Bismarck Archipelago: New Britain).

Habitat & Ecology — Primary and disturbed rain forest; 0—1000 m altitude; fl. Aug.,
fr. throughout the year.

Note — The staminal column is reminiscent of that in H. irya, and probably the vari-
ety originated by hybridization with the latter. Inflorescences of H. irya from this region
may be similarly woolly hairy. The leaves of var. novobritannica may show whitish
markings, similar to those found in H. irya.

38. Horsfieldia lancifolia W. J. de Wilde

Horsfieldia lancifolia W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 123, f. 18. — Type: bb Cel./II-
464, Sulawesi.

Tree 10—30 m. Twigs 1—2.5(-5) mm diameter, sometimes faintly lined, with hairs 0.1
mm or less, glabrescent; bark finely striate, not flaking; lenticels small, inconspicuous.
Leaves (thinly) chartaceous, oblong or oblong-lanceolate, 5-16 by 1.5—3.5(—4.5) cm,
base (long-)cuneate, apex acute-acuminate; upper surface drying olivaceous-brown, some-
times with small whitish marks, lower surface early or late glabrescent, hairs 0.1 mm or
less; with or without dots of mixed size; midrib slightly raised above; nerves 9-17 pairs,
flat, inconspicuous above, lines of interarching + regular, indistinct; venation lax, incon-
spicuous; petiole 10-20 by 1-1.5 mm; leaf bud 10-20 by 1.5—2 mm, with hairs 0.1 mm.
Inflorescences with sparse hairs 0.1—0.2 mm long; in male: 2 or 3 times branched, not
many-flowered, 5 by 3.5 cm, peduncle | cm long, flowers solitary or 2 or 3 together; in
female: 1-3 cm long, little branched, 4-10-flowered; flowers with sparse hairs 0.1—0.2
mm or less, perianth 2-lobed; pedicel not articulated. Male flowers: pedicel 1.5—-2 mm
long, tapered; buds obovoid to pear-shaped, somewhat laterally compressed, drying black-
ish, not or little collapsing on drying, 2.5—3 by 2—2.3 mm, apex broadly rounded, base
tapering into the pedicel, cleft c. 1/6, lobes (at the base) 0.4—0.5 mm thick; androecium
(synandrium) long-obovoid, laterally flattened, 1.5—1.8 by 1—1.2 mm, subtruncate (Plate
1: 31); thecae 12-16, erect, free apices 0.3-0.4 mm; column solid, except for the apical
part between the free apices of the anthers; androphore slender, 0.6—0.8 mm long. Fe-
male flowers: pedicel 1-2 or 3-4 mm long; buds subglobose to obovoid, variable in size,
2-3 by (1.8—)2-3.5 mm, cleft 1/4-1/6; ovary ovoid, 1.4—2.2 by 1-2 mm, densely minutely
pubescent; style and stigma(s) minute, 0.1 mm. Fruits 1-4 per infructescence, ellipsoid

Fig. 23. Horsfieldia lancifolia W.J. de Wilde. a. Habit of leafy twig with male inflorescences;
b. mature male flower, lateral view; c. ditto, opened, showing androecium; d. androecium, longitu-
dinal section, schematic; e. twig with female axillary inflorescences, leaves fallen; f. mature female
flower, lateral view; g. ditto, opened, showing finely pubescent ovary; h. twig with infructescences
with mature fruits [a—d: bb Cel./II-464; e—g: de Vogel 5267; h: van Balgooy 3973]. — Scale bar for
a, e, h = 2 cm; for b—d, f, g = 1.7 mm.

De Wilde — Myristicaceae (Horsfieldia) 143

(and + contracted at base) or pear-shaped, 2.5—3.5 by 1.8—2.4 cm, glabrescent, granulate
or with small lenticels or tubercles, drying brown; pericarp thick-woody, 4—8 mm thick;
seeds broadly ellipsoid, 17 by 14 mm; fruiting pedicel 3-10 mm long; perianth not per-
sistent. — Fig. 23.

Field-notes — Flower buds brown; fruits glossy green, turning yellow-green to or-
ange.

Distribution — Malesia: C & S Sulawesi.

Habitat & Ecology — Forest on ultrabasic rock (iron, nickel), laterite, also limestone
ridges; 200-1200 m altitude; fl. throughout the year, fr. May—July.

Note — Horsfieldia lancifolia can be easily recognized among the Horsfieldias from
Sulawesi and the Moluccas by the pear-shaped male buds, rather large fruits with thick
pericarp, and small, narrow leaves. The androphore is proportionally long as compared
to other species.

39. Horsfieldia laticostata (J. Sinclair) W.J. de Wilde

Horsfieldia laticostata (J. Sinclair) W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 15; Tree Fl. Sabah
& Sarawak 3 (2000) 374. — Horsfieldia brachiata (King) Warb. var. laticostata J. Sinclair,
Gard. Bull. Sing. 28 (1975) 12. — Type: Sinclair 10265, Sarawak.

Tree 12-35 m. Twigs 3-8 mm diameter, early glabrescent, hairs grey-brown or rusty,
0.1-0.4 mm; bark coarsely striate, when older almost flaking; lenticels usually distinct.
Leaves thinly to thickly coriaceous, (obovate-)oblong, (15—)20-33 by 6-12 cm, base
attenuate, apex (short-)acute-acuminate; upper surface drying dull olivaceous-brown,
glabrous, lower surface chocolate to rusty, contrasting with the upper surface, early gla-
brescent; dots absent; midrib raised above, towards the base + flat and conspicuously
broad, 3-4(—5) mm wide at transition to petiole, sometimes with remnants of indumentum
towards the base; nerves 1 1—20(—24) pairs, raised but towards the blade margin sunken,
lines of interarching + distinct, sunken; venation lax, flat or sunken, distinct or not;
petiole stout, 6-15 by 5-8 mm; leaf bud 15-23 by 3-5 mm, with dense grey-brown or
rusty hairs 0.1—0.4 mm long. Inflorescences below the leaves, with sparse hairs 0.2—0.4
mm long; in male: stout, 4 or 5 times branched, many-flowered, 10-25 by 8-22 cm,
main axis towards base 3-4 mm diameter, peduncle 0.5—3.5 cm long; in female: 3—7 cm
long, 2 or 3 times branched, rather few-flowered; bracts elliptic, acute, 2-8 by 1-4 mm,
with hairs 0.3 mm, + glabrescent, caducous; flowers (male) in clusters of up to 10, peri-
anth 3-lobed, glabrous, pedicel glabrous except few hairs 0.1—0.2 mm long towards the
base, distinctly articulated. Male flowers: pedicel 1-2 mm long, slender; buds broadly
obovoid to subglobose, 1.3—1.5 mm diameter, apex broadly rounded, base rounded or
sometimes + attenuate, glabrous, cleft c. 1/2, not collapsing on drying, lobes 0.2—0.3
mm, towards base of perianth 0.5 mm thick; androecium including androphore broadly
obovoid in outline, 0.7 mm diameter, triangular in cross section (Plate 3: 70); thecae 12,
suberect, + acute, mutually largely free, 0.3(—0.4) mm long; androphore tapering, 0.3
(-0.4) mm long. Female flowers: pedicel 0.5—1 mm long; buds broadly ellipsoid-glo-
bose, 2.5 mm diameter, glabrous or with a few minute hairs 0.1 mm long, cleft c. 1/3;
ovary broadly ovoid, 1.5 mm long, glabrous, stigma 0.1 mm long, shallowly 2(-4)-
lobed. Fruits 1-4 per infructescence, ellipsoid, 2.8-4 by 2.2-2.5 cm, glabrous, with or
without warts or lenticels; pericarp 4-5 mm thick; fruiting pedicel 2-4 mm long; peri-
anth not persistent.

144 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Tree without buttresses. Branches predominantly horizontal. Bark hard,
fissured or flaky, (reddish) brown; inner bark reddish, laminated, sapwood whitish. Flow-
ers yellow, smelling of Peru balsam. Fruits yellow, orange, or red; fruiting abundantly
throughout the crown along the smaller branches. Seeds glossy white, spotted.

Distribution — Malesia: Borneo (Sarawak, Sabah, C & NE Kalimantan).

Habitat & Ecology — Primary peat swamp forest, forest on sandy acid soils, water-
logged sand soils, kerangas; 0-400 m altitude; fl. & fr. throughout the year.

Note — Horsfieldia laticostata is closely related to the variable H. polyspherula,
especially to the varieties sumatrana and maxima. The species seems confined to forests
on poor soils, peaty and sandy grounds. It is characterized by stout twigs, large coriaceous
leaves with the midrib very broad and flat at the transition to the petiole, the usually
stout, broad petioles, fruits of moderate size (30—40 mm long), and large, stout, male in-
florescences. The male flowers (perianth and androecium) are very similar to those of
H. polyspherula.

40. Horsfieldia leptantha W.J. de Wilde

Horsfieldia leptantha W.J. de Wilde, Gard. Bull. Sing. 38, | (1985) 137. — Type: BW 12194 (Vink),
Papua Barat, Fak-Fak.

Tree 8-30 m. Twigs 4-8 mm diameter, early to late glabrescent, hairs rusty, 0.5—1 mm
long: bark finely or coarsely striate, not flaking; lenticels usually present. Leaves mem-
branous to chartaceous, elliptic to oblong-lanceolate, 13-35 by 5—13.5 cm, base (short-)
attenuate, apex acute-acuminate; upper surface drying dull olivaceous, smooth or finely
pustulate, lower surface early or late glabrescent, or with persistent stellate-dendroid
woolly or mealy hairs of mixed size, 0.5—1 mm long, when shed not leaving rough hair
bases: dots absent; midrib above flattish, glabrescent except towards base; nerves 10-
20(—30?) pairs, at 45—50° to the midrib, 5-20 mm apart, thin and sunken above, lines of
interarching irregular, prominent or not below; venation lax, distinct or not; petiole 5—12
by 2.54 mm; leaf bud 2.5—5 cm long, with dense hairs 0.5—1 mm long. Inflorescences
with woolly hairs 0.5—1 mm long; in male: rather many-flowered, 2 or 3 times branched,
11 by 7 cm, peduncle 0.5—2 cm long; bracts broadly elliptic, obtuse, 2-3 mm long;
flowers in loose clusters of 2—6, with hairs 0.2—0.3(—0.5) mm long; perianth 2-lobed,
pedicel slender, not articulated. Male flowers: pedicel 1-1.5 mm long; buds somewhat
compressed, in lateral view subcircular, 2.5 by 2.7-3 mm, membranous, not collapsing
on drying, cleft 3/4-5/6, lobes 0.2 mm thick; androecium compressed, 1—1.3 by 1.3—-1.6
mm, apex broadly rounded-truncate (Plate 2: 37); thecae 20-28, erect, touching each
other, 1—1.1 mm long, free apices 0.2-0.3 mm, not or faintly incurved, column narrowly
hollowed for 0.3 mm; androphore + absent. Female flowers not seen. Infructescences
6-12 cm long, branched. Fruits 2-8 per infructescence, broadly ellipsoid to subglobose
(seeds ellipsoid), 2-2.4 by 1.6—-2 cm, largely glabrescent but with minute vestigial hairs
towards the base, tubercles coarse, pale, lenticel-like, pericarp woody-coriaceous, 4-7
mm thick; fruiting pedicel 5—10 mm long; perianth not persistent.

Field-notes — Bark greenish black; blaze with pinkish red serous sap.

Distribution — Malesia: Papua Barat (Bird’s Head); Papua New Guinea (West Sepik
Prov.).

De Wilde — Myristicaceae (Horsfieldia) 145

Habitat & Ecology — Degraded and primary forest, ridge forest, on clay soil over
limestone; 0—600 m altitude; flowering and fruiting apparently not seasonal.

Note — The male buds are deeply cleft, to 3/4—5/6, much deeper than in the other
members of the group of species with H. hellwigii, to which H. leptantha belongs. The
fruits with coarse lenticels and tubercles are reminiscent of those in H. laevigata, and
possibly there is a close relationship with this species.

41. Horsfieldia macilenta W.J. de Wilde

Horsfieldia macilenta W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 13; Tree Fl. Sabah & Sarawak
3 (2000) 375. — Type: SAN 37103 (Aban), Sabah.

Tree 4-17 m. Twigs 1-3 mm diameter, rather late glabrescent, hairs pale rusty, stellate-
dendroid, (0.2—)0.5 mm long; rather bark finely striate, not cracking; lenticels small,
inconspicuous. Leaves membranous, oblong(-lanceolate), 10—18(—27) by 3—6.5 cm, base
(rounded to short-)attenuate, apex acute-acuminate; upper surface drying dull olivaceous,
glabrous, lower surface light brown, glabrous except midrib and the very base; dots
absent; midrib raised above, on lower surface late glabrescent; nerves 10—15(—18) pairs,
raised, lines of interarching fairly regular, distinct or not; venation lax, + indistinct; peti-
ole 10-15 by 1.5—2 mm, late glabrescent; leaf bud 6-8 by 1.5 mm, with dense hairs
(0.2-)0.5 mm. Inflorescences with sparse woolly hairs (0.2—)0.5 mm; in male: many-
flowered, 2 or 3(—4) times branched, 4-10 by 3—5(—7) cm, peduncle 0.3—1.2 cm long; in
female: similar to male, or shorter; bracts oblong to lanceolate, 2-5 mm, pubescent,
caducous; flowers in male in clusters of 5—8, in female fewer, glabrous or sometimes a
few hairs less than 0.1 mm towards the base of the pedicel; perianth 3-lobed, pedicel
articulated. Male flowers: pedicel 0.8—1.3 mm long; buds globose to depressed globose,
0.7-1.2 by 1-1.2 mm, cleft c. 1/2, lobes 0.2—0.3 mm thick; androecium subglobose (in-
cluding androphore) 0.4—0.5 by 0.5—0.7 mm, + triangular in cross section (Plate 3: 69);
thecae 10—14, suberect, 0.3—0.4 mm long, free for at least the upper half, column largely
hollow, passing into the broad tapering androphore 0.2—0.3 mm long. Female flowers:
pedicel 1.5 mm; buds subglobose, 1.5 mm diameter, cleft c. 1/2; ovary globose, gla-
brous, 1—1.2 mm diameter, stigma minutely 2-lipped, 0.1—0.2 mm long. Fruits (in
Sumatra) 2 or 3 together in a short infructescence 2—3 cm long, glabrous, ellipsoid, 2.2—
2.4 by 1.5—-1.6 cm, apex rounded, base contracted into a 2-4 mm long narrowed part;
pericarp 2 mm thick, not or sparingly tubercled; fruiting pedicel 5 mm long; perianth
not persistent.

Field-notes — Outer bark whitish or green-yellow, inner bark red with red sap; cam-
bium yellowish. Flowers (greenish) yellow.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Borneo (Sarawak, Sabah).

Habitat & Ecology — Primary lowland mixed dipterocarp forest, swamp forest; up to
200 m altitude; fl. June—Nov.

Note — According the structure of the male flowers, the present species is related to
H. polyspherula, especially the slender specimens of var. polyspherula. Horsfieldia maci-
lenta differs from H. polyspherula in the more slender twigs, thinly membranous leaves,
and slender and tiny inflorescences.

146 Flora Malesiana, Ser. I, Vol. 14 (2000)

42. Horsfieldia macrothyrsa (Miq.) Warb.

Horsfieldia macrothyrsa (Miq.) Warb., Mon. Myrist. (1897) 307; K. Heyne, Nutt. Pl. Ned. Indié
(1927) 637; W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 55. — Myristica macrothyrsa Miq.,
Pl. Jungh. (1852) 172; A. DC., Prodr. 14, 1 (1856) 203; Mig., Fl. Ind. Bat. 1, 2 (1858) 66; Suppl.
1 (1860) 156. — Type: Junghuhn (559), Sumatra, Tapanuli.

Tree 4-15 m. Twigs (2-)2.5—6 mm diameter, (blackish) brown, early glabrescent,
hairs (grey-)brown, 0.1 mm long; bark finely striate, not flaking; lenticels usually dis-
tinct. Leaves membranous, (elliptic-)oblong, 12—28 by 4—12 cm, base attenuate, apex
acute-acuminate; upper surface drying olivaceous to brown; lower surface early
glabrescent; dots (no dashes) present (lens!); midrib + flat above; nerves 9-17 pairs,
thin, flat or + raised above (see note 2), lines of interarching indistinct; venation lax,
indistinct on both surfaces; petiole 12-20 by 2-4 mm; leaf bud 10-15 by 1.5-3 mm,
with dense (grey-)brown hairs 0.1 mm. /nflorescences with sparse hairs 0.1 mm or
less: in male: about 3 times branched, + few-flowered, 7-20 by 5-12 cm, peduncle
(0.5—4.5 cm long; in female (according to infructescences): about twice branched, 3—6
cm long; bracts elliptic-oblong, 2-4 mm long, short-pubescent, caducous; flowers in
male in loose clusters of 2—4, in female 1—3 together, glabrous; the perianth 3- (or 4-)

sistent.

Field-notes —- Small tree; bark fissured or somewhat peeling; sap dark red-brown;
wood white to yellowish with red veins. Flowers greenish to yellow, aromatic. Fruits
2.5-3.5 cm long, ellipsoid, greenish to light yellow, pericarp light yellow inside; aril
green (almost mature, Lérzing 11703); seeds pale yellow.

Distribution — Malesia: northern and Central Sumatra.

Habitat & Ecology — Lower and mid-montane forest, riverine forest; 400-1600 m
altitude; fl. & fr. throughout the year.

Notes — 1. Related to H. glabra, with similar fruits and flowers. Horsfieldia
macrothyrsa has markedly larger male flowers, and about twice as many anthers. It has
a different distributional area; H. glabra is not found in C and N Sumatra.

2. The specimens of H. macrothyrsa rather differ from each other in various aspects.
Plants from Mt Sago (= Mt Malintang), C Sumatra, are less robust in general habit, the
leaves and inflorescences are smaller, the male buds 3—3.5 mm diameter. Specimens
from Sibolangit Botanic Garden jungle are stout, with large leaves (to 28 cm long), large
male buds (4.3 by 4 mm), and deviate furthermore by the slightly raised nerves on the
upper leaf surface.

De Wilde — Myristicaceae (Horsfieldia) 147

43. Horsfieldia majuscula (King) Warb.

Horsfieldia majuscula (King) Warb., Mon. Myrist. (1897) 315; Gamble, Mat. Fl. Malay Penins. 5,
23 (1912) 215; Ridl., Fl. Malay Penins. 3 (1924) 57; W.J. de Wilde, Gard. Bull. Sing. 39, 1
(1986) 48. — Myristica majuscula King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 310, pl. 143. —
Lectotype: King’s coll. 5039, Peninsular Malaysia.

Horsfieldia bartlettii Merr., Papers Mich. Acad. Sc. Arts & Letters 24, 1 for 1938 (1939) 71. —
Type: Rahmat si Boeea 8772, Sumatra.

Tree 6—25 m. Twigs 2.5—5 mm diameter, grey-brown, early glabrescent, hairs grey-
ish brown, 0.1(—0.2) mm; bark coarsely longitudinally striate, not flaking; lenticels small,
rather inconspicuous. Leaves membranous or thinly chartaceous, (elliptic-)oblong, 15—
27 by 5.5—9.5 cm, base attenuate, apex acute-acuminate; upper surface glabrous, drying
olivaceous-brown, lower surface brown, not conspicuously contrasting in colour with
upper surface, early glabrescent; dots absent; midrib raised above; nerves 11-19 pairs,
raised above; venation lax, indistinct on both surfaces; petiole 10-20 by 2—2.5 mm, leaf
bud 10-15 by 2-3 mm, with grey brown hairs 0.1(—0.2) mm. Inflorescences with rather
dense stellate hairs 0.1 mm long; in male: 2 or 3 times branched, not very many-flow-
ered, 8 by 4—5 cm, peduncle 1—1.5 cm; flowers in clusters of 2—5; in female: 2-5 cm
long, twice branched, fewer-flowered; bracts ovate-oblong, 4 by 2 mm, with dense hairs
0.2 mm long, caducous; flowers with perianth 3- (or 4-)lobed, glabrous, pedicel gla-
brous or sparsely pubescent (hairs 0.1 mm), articulated. Male flowers: pedicel 2-3 mm
long; buds broadly ellipsoid or obovoid, 2—3 by 1.8—2.5 mm, apex rounded, base short-
attenuate, slightly tapering, cleft 1/3 to 1/2, not or slightly collapsing on drying, lobes
thick, 0.3 mm, towards the base to 0.8(—1) mm thick; androecium (including the broad,
shortly tapering, 0.2-0.5 mm long androphore) ellipsoid to ellipsoid-oblong, (1.4—)
1.7-2 by 0.8-1 mm, in cross section subtriangular to subelliptic (Plate 3: 87); thecae
14-18, slightly curved, 1.2—-1.8 mm long, free apices (0.2—)0.3—0.5 mm. Female flow-
ers: pedicel 2-3 mm long; buds broadly obovoid, 2.5 by 2.5 mm, cleft c. 1/3; ovary
ellipsoid, 1.8—2 by 1.3-1.5 mm, glabrous, stigma 2-lipped, 0.3 mm high. Fruits 1—5 per
infructescence, ellipsoid, 4.3-6.5 by 3—4.5 cm, glabrous, drying rusty-brown, finely
granulate, not or inconspicuously warted or lenticellate; pericarp 4—11 mm thick; fruit-
ing pedicel 2-7 mm long; perianth not persistent.

Field-notes — Tree without buttresses; bark shallowly fissured or forming shallow,
rectangular flakes; slash wood whitish, red-flecked. Flowers yellow. Fruits up to 5 per
infructescence, up to 7 by 6 cm, yellow to bright red; pericarp 1.5 cm thick.

Distribution — Malesia: Sumatra (W & E Coast, Palembang, Jambi, Aceh), Peninsu-
lar Malaysia (Perak, Pahang, Kelantan).

Habitat & Ecology — Dry-land forest and freshwater swamp-forest of lowland and
montane areas; rocky stream banks, river valleys; up to 1200 m altitude; fl. & fr. throughout
the year.

Note — Horsfieldia majuscula can be confused with H. polyspherula var. sumatrana
and var. maxima. Variety sumatrana differs in usually coarser hairs on the leaf buds,
more contrasting colour of the two surfaces of dry leaves, smaller globose male peri-
anths, essentially different androecium, and smaller fruits, up to 35 mm long. Variety
maxima, known only from Borneo, differs in the same characters, but has fruits of a
similar size as those of H. majuscula, with the pericarp up to 15 mm thick.

148 Flora Malesiana, Ser. I, Vol. 14 (2000)

44. Horsfieldia moluccana W.J. de Wilde

Horsfieldia moluccana W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 65. — Type: Kostermans
673a, Morotai.

Tree 8-20(—30) m. Twigs sometimes slightly angular, 2-5 mm diameter, early
glabrescent; hairs 0.1—0.3 mm; bark striate, not flaking, lenticels conspicuous or incon-
spicuous. Leaves thinly chartaceous, (elliptic-)oblong, (6—)8—25 by 2.5—8.5 cm, base
attenuate, apex acute-acuminate; upper surface drying olivaceous to brown, usually mi-
nutely pale-pustulate, lower surface early glabrescent; dots absent; midrib slender, flat
above: nerves 6—15 pairs, thin, flat, indistinct above, lines of interarching indistinct;
venation + fine, indistinct on both surfaces; petiole 10-26 by 1-2 mm; leaf bud 6-12 by
1-2 mm, with hairs 0.1—0.3 mm. Inflorescences with sparse stellate hairs 0.1 mm or less;
in male: (1—)2—3 times branched (sometimes hardly branched, spike-like, see note Ny);
5-11 by 2-5 cm; in female: up to 5 cm long; peduncle 0.5—2 cm; bracts + oblong, 1.5—4
mm long, thinly pubescent, caducous; flowers solitary or to 4 together, glabrous; peri-
anth 2-lobed, pedicel not articulated. Male flowers: buds (including pedicel) pear-shaped,
little to much compressed, about as broad as or slightly broader than long, 1.5—2.5(-3)
by 2.2-3.8 mm, apex broadly rounded, the lower 1/3 tapering into the pedicel 2—3(—3.5)
mm. cleft 2/3—4/5, lobes 0.2-0.3 mm thick; androecium compressed, broadly transversely
ellipsoid or kidney-shaped, broadly rounded above, 1.1-1.5 by 1.4-2.8 mm (Plate 1: 5);
thecae (14—)20—36, contiguous, 1.5-2 mm long, free apices 0.1—0.5 mm, only at one
side of the androecium strongly incurved; column hollow for at least 2/3; androphore to
0.1 mm long. Female flowers: pedicel 2—2.5 mm long; buds broadly ovoid-ellipsoid,
1.8-2.2 by 2-2.2 mm, cleft 1/2-4/5; ovary ovoid, glabrous, 1-1.5 by 1 mm, stigma
minutely 2-lobed, 0.1 mm high. Fruits solitary or 2-6 per infructescence, ellipsoid, apex
rounded to subacute, 1.3—2.8 by 1.1—1.7 cm, glabrous, without or with sparse tubercles;
pericarp 1-2 mm thick; fruiting pedicel 2-5 mm long; perianth not persistent.

Distribution — Malesia: northern Moluccas, West New Guinea.

Note — A species with 4 varieties, closely related to H. spicata and H. tuberculata.
Horsfieldia spicata has the generally deeply asymmetrically incurved anthers in com-
mon with H. moluccana, but the former differs in pale twigs and (almost) spike-like
male inflorescences. Horsfieldia tuberculata has the staminal column hollowed at the
apex for 1/5—1/3.

KEY TO THE VARIETIES

la. Hairs of leaf buds woolly-rusty, 0.2-0.3 mm long. — Bird’s Head: Fak-Fak jo.3. >
re ce ac Perle om eee Let arreE aromas oes c. var. pubescens

b. Hairs of leaf buds up to 0.1 mm... ©... 0-2. eee se ee 2
2a. Petiole 10-15 mm long. [Male buds 3 mm wide.] — Morotai, Obi [Soiree
ea Ea, rue ner: 5... AE Ome need cao Lore BFS 2 a. var. moluccana

b. Petiole (10—)15—25 mm long, generally longer in proportion to the smaller blade. —
West New GUilled. osc meee ae de soa cise seas aus = + ne pear 5

3a. Leaf blades 7-15 cm long. Fruits 1.3—1.8 cm long. Male buds 2—2.5 mm WIG wenn:
satel ends Syemeaetemeted in IR sc" a sor yn ane ea wire me a ec Ser aaehih as b. var. petiolaris

b. Leaf blades 13-23 cm long. Fruits 2.2—3 cm long. Male buds (3—)3.5—3.8 mm wide
ESN! DEAL SOLED, ECUTED CER AEP R etre ne facets orcs = Sincere ae emerereno cana ace ohatene d. var. robusta

De Wilde — Myristicaceae (Horsfieldia) 149

a. var. moluccana

?Horsfieldia olivaeformis Warb., Mon. Myrist. (1897) 352, t. 23; Markgr., Bot. Jahrb. Syst. 67
(1935) 152, p.p.; W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 66. — Myristica olivaeformis
(Warb.) Boerl., Handl. 3 (1900) 87. — Type: Beccari 17] (FI, n.v., see note 2), Papua Barat,
Sorong, Bird’s Head.

Leaf blades 9—22 by 4-8 cm; petiole (8—)10-20 mm. Indumentum of leaf bud com-
posed of hairs 0.1 mm long or less. Male flowers: buds 2—3 by 2.7-3.3 mm, pedicel 2-3
mm long. Fruits 1.5 cm long.

Field-notes — Straight tall tree, to 30 m, once with prop-roots up to 1.5 m; bark peel-
ing off or not, exudate watery, turning pink, later brownish. Flowers yellow, once red.

Distribution — Malesia: northern Moluccas (Morotai, Obi).

Habitat & Ecology — Well-drained forests on clayey soil, volcanic soil, alluvial soil
rich in humus, also flat land behind the mangrove; also found on limestone, at the base
of serpentine-rock, or on porous nickel-containing soil; 0-600 m altitude; fl. & fr. through-
out the year.

Notes — 1. Can be confused with H. parviflora which has more roundish (not pear-
shaped) male buds and shorter petioles. Horsfieldia spicata is closely related but differs
in the paler colour of the dried twigs, the generally spike-like male inflorescences, and
the more membranous leaves.

2. [have not seen the type of H. olivaeformis, a name which may have priority but is
not accepted here (De Wilde, I.c.).

b. var. petiolaris W.J. de Wilde

Horsfieldia moluccana W.J. de Wilde var. petiolaris W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985)
67. — Type: van Royen 5388, West New Guinea.

Leaf blades 6—15(—19) by 2.5—6.5(—7) cm; petiole proportionally long, 10-20 mm.
Indumentum of leaf bud composed of hairs up to 0.1 mm long. Male flowers: buds 1.5—
2.2 by 2.2-2.5 mm, pedicel 2—3.5 mm long. Fruits 1.5—1.8 cm long.

Field-notes — Bark flaking. Flowers greenish. Fruits yellow or orange-yellow.

Distribution — Malesia: Papua Barat (Bird’s Head; islands in the Geelvink Bay:
Noemfoer, Meos Waar, Japen I.; Waigeo I.).

Habitat & Ecology — Locally common in forests on sandy or stony-clayey soils;
Calophyllum—Ficus forest; 0-100 m altitude; fl. & fr. throughout the year.

Uses — Fruits once reported as edible and sour.

c. var. pubescens W.J. de Wilde

Horsfieldia moluccana W.J. de Wilde var. pubescens W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985)
68. — Type: BW 15370 (Vink), West New Guinea, Bird’s Head.

Leaf blades (6—)8—14 by (2.5—)3—5 cm; petiole (9—)11—18 mm long. Indumentum of
leaf bud + woolly, composed of hairs 0.2—0.3 mm long. Flowers not seen. Fruits 1.3 cm
long.

Field-notes — Tree to 16 m; buttresses up to | m high, 0.5 m wide; the bark little or
strongly peeling; inner bark with much red and clear exudate; wood white. Fruits light
green.

150 Flora Malesiana, Ser. I, Vol. 14 (2000)

Distribution — Malesia: Papua Barat (Bird’s Head).

Habitat & Ecology — Common in primary and degraded forest on clayey soil or
sandy clay over limestone; most collections from limestone; 50—300 m altitude; fr. Mar.,
May.

Note — This variety appears to be almost identical with var. petiolaris, except for the
more woolly indumentum. In Horsfieldia, the nature of the indumentum is usually of
taxonomic significance.

d. var. robusta W.J. de Wilde

Horsfieldia moluccana W.J. de Wilde var. robusta W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 68.
— Type: van Royen & Sleumer 6682, West New Guinea.

Leaf blades 12—22(-25) by 3.5—-8(-9) cm; petiole 12-26 mm. Indumentum of leaf
bud consisting of hairs 0.1 mm long. Male flowers: buds 2.5—3 by (3-)3.5-3.8 mm,
pedicel 3 mm. Fruits 2.2-3 cm long.

Field-notes — Bark flaking. Flowers yellow, fragrant. Fruits yellow.

Distribution — Malesia: Papua Barat (Bird’s Head, Batanta 1):

Habitat & Ecology — Degraded and coastal forest, on limestone; 0-15 m altitude;
fl. & fr. throughout the year.

Note — Var. robusta is a form similar to var. petiolaris but coarser in all aspects:
leaves, flowers, and fruits are larger. It superficially resembles H. tuberculata, which
differs in the generally shorter petioles, and the androecium with the anthers not strongly
inflexed into the cavity

45. Horsfieldia montana Airy Shaw

Horsfieldia montana Airy Shaw, Kew Bull. 1939, n. 10 (1940) 542; W.J. de Wilde, Gard. Bull.
Sing. 39, 1 (1986) 35; Tree Fl. Sabah & Sarawak 3 (2000) 375. — Type: Richards (Native Coll.)
2509, Sarawak, Dulit Range.

Tree (3—)7—24 m. Twigs terete or faintly angular, 1.5—4(—7) mm diameter, early to
late glabrescent, hairs short or long-shaggy, 0.2—1 mm long; bark coarsely striate, flak-
ing or not, lenticels inconspicuous or absent. Leaves chartaceous or coriaceous, elliptic
(-oblong), 4-14 by 2-6 cm, base (short-)attenuate, apex rounded or (sub)acute; upper
surface glabrous (base of midrib late glabrescent), drying olivaceous to blackish, lower
surface glabrous but very base and midrib late glabrescent, drying chocolate, not much
contrasting with upper surface; dots absent; midrib raised above, nerves 6—11 pairs,
flattish to raised above, lines of interarching indistinct; venation lax, faint on both sur-
faces; petiole 5—16 by 1.5—2.5 mm, late glabrescent; leaf bud 5—10 by 1.5—4 mm with
hairs 0.2-1 mm long. Inflorescences behind the leaves, sparsely or densely pubescent,
hairs 0.2—1 mm long (see note); in male: 3 or 4 times branched, sometimes many-flow-
ered, 4-12(-16) by 3-10 cm, peduncle 0.5—2 cm long, the flowers in loose clusters of
3-10 each: in female: few-flowered, 2—6 cm long; bracts ovate to ellipsoid, 2.5—6 mm
long, pubescent, caducous; flowers glabrous, perianth 3- or 4-lobed, pedicel in male not
articulated, in female + articulated (always?). Male flowers: pedicel 1-1.5 mm; buds
(sub) globose, (1.2—)1.4-2 mm diameter, cleft c. 1/3 to nearly 1/2, not or slightly collaps-
ing on drying, lobes (0.2—)0.3 mm thick; androecium (depressed) globose, apex rounded,

De Wilde — Myristicaceae (Horsfieldia) 15]

base rounded, + truncate, or sagged, (0.5—)0.6—1.1 by 0.8—1.1 mm, subcircular in cross
section (Plate 3: 80); thecae (16—)18—26, almost completely sessile, 0.8—1.2 mm long,
+ curved, free apices to 0.1 mm; column broad, with shallow apical cavity or slit 0.1—0.2
mm deep; androphore narrow, 0.3—0.5 mm long, hidden by the anthers, thus making the
androecium look sessile. Female flowers: pedicel 1.5 mm long; buds ellipsoid-obovoid,
2 by 1.8 mm, cleft c. 1/3; ovary ellipsoid, 1.5 by 1.3 mm, glabrous, stigma minutely
2-lobed, 0.1—0.2 mm high. Fruits 2-9 per infructescence, ellipsoid, base rounded, slight-
ly contracted towards the fruiting pedicel, apex rounded or + acute, 2—2.7 by 1.3-1.7
cm, glabrous, with or without few small lenticel-like tubercles, pericarp 1.5—2 mm thick;
fruiting pedicel 2—4 mm long, at base + articulated; perianth not persistent.

Field-notes — Low or medium-sized tree, without buttresses; bark (slightly) longitu-
dinally, shallowly fissured, or sometimes flaky, chocolate, red-brown, or dark grey; in-
ner bark pale yellowish, reddish, or brownish, with red watery exudate or not; sapwood
pale orange or whitish. Flowers yellow, sweet-scented; androecium orange. Fruits yel-
low or red.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah).

Habitat & Ecology — Crest forest, kerangas on ridges, montane forest, moss forest,
Agathis forest; on black soil or sandy soil; (800—)1300—2000 m altitude; fl. & fr. through-
out the year.

Note — Specimens from Kinabalu and vicinity differ markedly from those from
Sarawak and Brunei in the nature of the indumentum. Specimens from the latter two
areas (including the type) have compact hairs 0.2(—0.5) mm long; the Kinabalu speci-
mens have shaggy hairs 0.5—1 mm long, sometimes with longer emergents.

46. Horsfieldia motleyi Warb.

Horsfieldia motleyi Warb., Mon. Myrist. (1897) 304; Merr., Enum. Born. (1921) 268; J. Sinclair,
Gard. Bull. Sing. 28 (1975) 81; W.J. de Wilde, Gard. Bull. Sing. 38, 2 1985’, 1986) 209; Tree
Fl. Sabah & Sarawak 3 (2000) 376. — Myristica motleyi (Warb.) Boerl., Handl. 3 (1900) 85. —
Type: Motley 355, SE Kalimantan, Banjermasin.

Horsfieldia macrobotrys Merr., Univ. Calif. Publ. Bot. 15 (1929) 76. — Type: Elmer 21882, Sabah,
Tawau.

Tree 12-35 m. Twigs (2.5—)3—5 mm diameter, at first with dense rusty hairs 0.5(—1)
mm, late glabrescent; bark finely striate, not flaking; lenticels inconspicuous. Leaves
membranous, elliptic to oblong, 9-27 by 4—12 cm, base rounded to attenuate, apex + ob-
tuse to acute-acuminate; upper surface drying olivaceous(-brown), glabrous, lower sur-
face with persistent rather dense stellate-dendroid hairs of mixed size, 0.3-1 mm long;
dots absent; midrib flattish above, sometimes late glabrescent; nerves 9-21 pairs, flat or
sunken, often late glabrescent above; lines of interarching + indistinct; venation lax,
faint above; petiole 13-22 by 1.5—4 mm, late glabrescent; leaf bud 15-20 by 3-4 mm,
with dense hairs 0.5 mm long. Inflorescences with dense stellate-dendroid yellow-
brown to rusty hairs 0.5—1 mm long; in male: 4 or 5 times branched, many-flowered,
12-20 by 10-14 cm, peduncle 2—4 cm, flowers + densely clustered (immature buds
+ angular); in female: much branched, rather many-flowered, 3—6(—10) cm long; bracts
broadly ellipsoid, acutish, 2-5 mm long, densely pubescent, caducous; flowers (male) in
clusters of 5—20, with hairs 0.1—0.3 mm long, perianth 3-lobed, pedicel not articulated.

152 Flora Malesiana, Ser. I, Vol. 14 (2000)

Male flowers: pedicel slender, 1—1.5 mm long; buds subglobose or broadly obovoid,
0.8—1 by 0.7-1(-1.1) mm, pubescent or late glabrescent, cleft c. 1/2, lobes 0.1 mm thick;
androecium broadly obovoid, 0.5—0.7 by (0.3—)0.5—0.6 mm, synandrium at apex some-
what flattish, depressed in the centre, subcircular in cross section (Plate 2: 53); thecae
10, completely sessile, towards the apex not or hardly incurved, 0.25—0.4 mm long,
column broad, with broad, apical hollow to c. 1/10, androphore conspicuous, continuous
with the anthers, + tapering, 0.2-0.3 mm long. Female flowers: pedicel 1.5 mm long;
buds ellipsoid, 2.4 by 1.8-2 mm, pubescent, cleft 1/4—1/3; ovary broadly ellipsoid, 1.5
by 1.2 mm, grooved at one side, glabrous, stigma broad, shallowly 2-lobed, 0.2 mm
high. Fruits 5-15 per infructescence, broadly ellipsoid, 1.9-2.3 by 1.6-1.8 cm, glabrous,
smooth; pericarp 2 mm thick; fruiting pedicel 2-4 mm long; perianth not persistent.
Field-notes — Bole sometimes with steep buttresses to 5 m high, 30 cm out, 9 cm
thick, merging into the stem; bark fissured, often scaly, strips 3-4 cm wide, to 10 mm
thick, black-brown, chocolate, or red-brown; living bark 10-12 mm thick, undulate in
cross section, dark brown, brown-red, or red-laminated; cambium pinkish; sapwood (red-
dish) white, heartwood pinkish or brown; exudate of bark a red watery sap, sometimes
sticky, appearing fast. Flowers (dark) yellow. Fruits orange-red, with sticky exudate.
Distribution — Malesia: Borneo (Sarawak, Sabah, W, S, SE and E Kalimantan).
Habitat & Ecology — Primary forest, disturbed or poor forest, Dryobalanops forest,
hill dipterocarp forest, usually on dry sandy soils, sandy clay, also loam with lime; often
on ridge tops; 0-750 m altitude; fl. & fr. throughout the year.
Note — Horsfieldia motleyi, especially in a young stage, may be confused with
Endocomia rufirachis, a species also with pubescent flowers.

47. Horsfieldia nervosa W.J. de Wilde

Horsfieldia nervosa W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 16; Tree Fl. Sabah & Sarawak 3
(2000) 377. — Type: S 16652 (Ilias Paie & Ashton), Sarawak.

Tree 13-16 m. Twigs 3.5—6 mm diameter, grey-brown, late glabrescent, hairs rusty,
(0.30.6 mm long; bark coarsely striate, slightly flaking or not; lenticels + inconspicu-
ous. Leaves thinly coriaceous, oblong, 16-28 by 5.5—9 cm, base short-attenuate to rounded,
apex acute-acuminate; upper surface drying olivaceous, largely glabrous but midrib and
nerves late glabrescent; lower surface chocolate, contrasting in colour with upper sur-
face, glabrescent but hairs 0.5 mm long, often (partly) persistent on midrib and nerves,
dots absent: midrib slender (at transition of petiole 1.5 mm wide), raised above; nerves
16-19 pairs, raised above, glabrescent, lines of interarching flattish, indistinct; venation
lax, indistinct, petiole 15—22 by 3.5—4.5 mm, late glabrescent; leaf bud 16-20 by 3.5—-4.5
mm, with dense rusty hairs 0.5 mm long. Inflorescences behind the leaves, with dense
woolly-shaggy hairs up to 0.6 mm long; in male: about 4 times branched, many-flow-
ered, 10 by 10 cm, peduncle 1-2 cm long, the flowers in clusters of 2—6; in female:
fewer-flowered, 4 by 3.5 cm; bracts pubescent, caducous; flowers with perianth 3-lobed,
glabrous, pedicel towards base with hairs 0. 1-0.2 mm, articulated. Male flowers: pedi-
cel 1.5 mm long; buds globose, 1.2 mm diameter, cleft nearly 1/2, not collapsing on
drying, lobes 0.2 mm thick, at base of perianth + fleshy-coriaceous, 0.5—0.6 mm thick;
androecium including androphore + broadly obovoid, 0.6—0.7 by 0.6 mm, triquetrous in
cross section (Plate 3: 7/); thecae 10 or 12, at least the upper half free, + erect, 0.3-0.4

De Wilde — Myristicaceae (Horsfieldia) 153

mm long; column deeply hollowed; androphore + tapering to the base, 0.3 mm long.
Female flowers: pedicel 1.5(—2) mm long, + pubescent with hairs 0.1—0.2 mm; buds
subglobose, 2.5 mm diameter, subglabrous, cleft slightly over 1/3; ovary broadly ovoid,
1.5 mm diameter, glabrous, stigma 0.3 by 0.6 mm, 2-lobed, the lobes shallowly 5- or 6-
lobulate. Fruits not seen.

Field-notes — Buttresses thin, small, to 35 cm tall; bark pale ochre and brown-mot-
tled, smooth. Flowers pale yellow.

Distribution — Malesia: Borneo (Sarawak: Ist Div.; two collections only).

Habitat & Ecology — Primary forest on yellow podzolic soil; c.70 m altitude; fl. Nov.

Note — This species is close to H. polyspherula, which has smaller leaves with fewer
lateral nerves and attenuate base. In H. nervosa the nerves on the upper leaf surface are
strong and markedly raised, and midrib and nerves on both surfaces remain covered
with indumentum for a long time.

48. Horsfieldia obscura W. J. de Wilde

Horsfieldia obscura W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 44; Blumea 32 (1987) 468; Tree
Fl. Sabah & Sarawak 3 (2000) 378. — Type: Kostermans 13773, E Kalimantan.

Tree c. 20 m. Twigs 2—3(—4) mm diameter, early glabrescent, hairs greyish brown to
rusty, 0.1—0.2 mm long; bark finely striate, not flaking; lenticels not much contrasting.
Leaves membranous, (elliptic-)oblong, 10—15 by 4~7 cm, base short-attenuate, apex short
acute-acuminate; upper surface dark brown, lower surface pale or bright brown, gla-
brous; dots absent; midrib + flat above; nerves 10-13 pairs, thin and flat above, lines of
interarching invisible; venation lax, very faint on both surfaces; petiole 11-15 by 1.5—
2.5 mm, glabrous; leaf bud slender, 7-10 by 1.5—2 mm, with dense (grey-)rusty hairs
0.1—0.2 mm. Inflorescences behind the leaves, with + sparse hairs 0.1 mm; in male: 3
(or 4) times branched, + many-flowered, 7-10 by 5—8 cm, peduncle (0.2—)0.6—1.2 cm
long; in female: 3.5 cm long (stout, as seen in infructescences); bracts caducous, not
seen; flowers in male in clusters of 3—5, perianth 3- (or 4-)lobed, glabrous, pedicel gla-
brous or with some minute hairs towards the base, not or faintly articulated. Male flow-
ers: pedicel 1-2 mm long, slender; buds subglobose, 2—2.2 by 2—2.3 mm, cleft 2/3—4/5,
lobes 0.2—0.3 mm thick; androecium (depressed) globose-ovate, apex narrowly, base
broadly rounded, | by 1—1.2 mm, in cross section (sub)circular (Plate 3: 85); thecae 16—
20, almost completely sessile, free apices sometimes + sterile, up to 0.1(—0.2) mm long,
concealing the narrow, 0.1—0.2 mm deep apical cavity of the column; androphore nar-
row, to 0.2 mm long, hidden by the anthers. Female flowers not seen. Fruits (of S 36305,
see note 2) 2 or 3 ina short, stout infructescence, broadly ellipsoid, 5—5.5 by 3.5—4 cm,
glabrous, with a few, small tubercles, pericarp 7-10 mm thick, perianth not persistent.

Field-notes — Trunk irregular; bark red-brown, rough, 5 mm thick, peeling off irreg-
ularly in strips; living bark 10 mm, red to brown-red; sap red; wood reddish to brown-
red. Flowers (dark) yellow, smelling of Peru-balsam. Fruits bright orange.

Distribution — Malesia: Borneo (E Kalimantan, possibly Sarawak, see note 2), Phil-
ippines (Palawan).

Habitat & Ecology — Ridge forest on limestone, coral limestone, yellow sandy soil
in lowland dipterocarp forest; 150-730 m altitude; fl. (E Kalimantan) Aug., Nov.; fr.
(Sarawak) May.

154 Flora Malesiana, Ser. I, Vol. 14 (2000)

Notes — 1. Horsfieldia obscura keys out beside H. subalpina (accepted for Borneo
as subsp. kinabaluensis and restricted to the montane forest in the Kinabalu area). The
latter differs from H. obscura in the more elongate male buds, cleft to nearly halfway,
short-ellipsoid androecium, more rigid leaves, and smaller fruits with less thick pericarp.
Horsfieldia obscura appears to be closely related to typical H. subalpina, from Peninsu-
lar Malaysia, which differs in habit (larger leaves, more distinct lenticels on twigs, larger
inflorescences), in male flowers cleft to about halfway, and smaller fruits.

2. Some doubtful specimens in fruit, all from Borneo, key out beside H. subalpina,
H. obscura, or H. discolor, These specimens are discussed by De Wilde (I.c.), and may
represent new species.

49. Horsfieldia obscurinervia Merr.

Horsfieldia obscurinervia Merr., Philipp. J. Sci., Bot. 12 (1917) 265; Enum. Philipp. Flow. Pl. 2
(1923) 182; W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 71. — Type: FB 26503 (de Mesa &
Magistrado), Philippines.

Horsfieldia ramosii Merr., Philipp. J. Sci. 17 1920", 1921) 254: Enum. Philipp. Flow. Pl. 2 (1923)
182. — Type: BS 35047 (Ramos), Philippines.

Tree c. 11 m. Twigs 1.5—4 mm diameter, at first with greyish hairs less than 0.1 mm
long, glabrescent; bark finely striate, not flaking; lenticels rather distinct. Leaves
chartaceous, (oblong-)lanceolate, 5-14 by 2—4 cm, base attenuate, apex acute-acumi-
nate; upper surface drying olivaceous to brown, + glossy, lower surface early glabrescent,
dots absent; midrib flat or slightly raised above; nerves 7—15 pairs, slender, inconspicu-
ous on both surfaces, lines of interarching inconspicuous; venation hardly visible; peti-
ole 6-15 by 1-1.5 mm; leaf bud 6-10 by 1.5-2 mm, with hairs 0.1 mm long or less.
Inflorescences subglabrous or with sparse hairs 0. 1—0.3 mm long; in male: 2 or 3 times
branched, rather few-flowered, 3—4 by 2—3 cm, peduncle 0.5—1 cm; bracts and bracteoles
not seen, caducous; female not seen; flowers solitary or 2 or 3 together, glabrous, peri-
anth 2-lobed, pedicel not articulated. Male flowers: pedicel slender, 1 mm long; buds
slightly compressed, subobtriangular, 2 by 2.2 mm, apex rounded, base + cuneate, rather
firm, not collapsing on drying, bright brown, cleft c. 1/2, lobes 0.2—0.3 mm thick; an-
droecium + obtriangular or obovoid, narrowed to the base, sli ghtly laterally compressed,
thickish, 1.5 by 1.2 mm, column largely hollow; anthers 11 or 12 (i.e., 11 or 12 thecae on
each side), almost completely connate (the free apices 0.1 mm), forming a deep thick-
walled cup, the anthers at one side deeply inflexed and almost completely filling the cup
(Plate 1: 7); androphore narrow, 0.1 mm long. Female flowers: not seen. Infructescences
2-3 cm long, once or twice branched, with 2-5 fruits. Fruits short-ellipsoid, 1.1—1.3 by
0.9-1.1 cm, glabrescent except for minute dendroid hairs at base (hence ovary pubes-
cent), finely granulate, not tuberculate, drying (reddish) brown; pericarp |—1.5 mm thick;
fruiting pedicel 2-5 mm long; perianth not persistent.

Field-notes — Small tree. Flowers yellow.

Distribution — Malesia: Philippines (Luzon).

Habitat & Ecology — On low hills at c. 20 m altitude; fl. July; fr. Nov., Dec.

Note — Related to H. parviflora from the Moluccas, with an almost similar flower
structure but differing in the larger membranous leaves, larger male perianths (2.5—4
mm width), 36—50 thecae, thinner-walled and deeper androecium cup, with the anthers
usually inflexed at both sides of the androecium, and glabrous ovary and fruit.

De Wilde — Myristicaceae (Horsfieldia) 155

50. Horsfieldia obtusa W. J. de Wilde

Horsfieldia obtusa W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 9; Tree Fl. Sabah & Sarawak 3
(2000) 379. — Type: BS 821 (Native Coll.), Sarawak.

Tree, height not recorded. Twigs terete or faintly angular, 2.5—-4(—6) mm diameter,
grey-brown, rather late glabrescent, hairs dark rusty, 0.2—0.4 mm long; bark coarsely
striate, finely longitudinally cracking; lenticels small and inconspicuous. Leaves thinly
coriaceous, oblong, 8—10 by 3—3.5 cm, base attenuate, apex rounded; upper surface gla-
brous (except midrib), dark olivaceous on drying, lower surface early glabrescent, pur-
plish brown on drying, with pale yellowish enlarged hair scars, but dots absent; midrib
moderately raised, late glabrescent above; nerves c. 10 pairs, + flat above, lines of in-
terarching regular, clearly visible; venation lax, hardly visible; petiole 10 by 2 mm, late
glabrescent; leaf bud 12 by 3 mm, with dense dark rusty hairs 0.2-0.4 mm. /nflores-
cences behind the leaves, densely pubescent, hairs rusty, 0.2—0.4(—0.5) mm long; in male:
about 3 times branched, many-flowered, 5—9 by 3—5 cm, peduncle 0.3—1 cm long, with
flowers in clusters of 3-6; female not seen; bracts caducous; flowers glabrous, perianth
3- (or 4-)lobed, pedicel articulated. Male flowers: pedicel 1-1.5 mm long, slender; buds
subglobose, 1.3-1.5 by 1.5-1.6 mm, cleft nearly 1/2, firm, not collapsing on drying,
lobes at the apex 0.2 mm, at base 0.4 mm thick; androecium (including androphore)
broadly obovoid, 0.8 by 1 mm, in cross section 3- (or 4-)quetrous (Plate 3: 67); thecae
18 or 20, suberect, 0.5 mm long, free in the upper half, acutish; androphore + tapering to
base, 0.3 mm long; column hollowed to about the base of the anthers. Female flowers
and fruits not known.

Distribution — Malesia: Borneo (Sarawak; known only from the type, precise local-
ity not indicated).

Habitat & Ecology — Not known.

Note — Horsfieldia obtusa obviously belongs in the group of species with H. poly-
spherula on account of the articulated pedicel and angular androecium with the distally
free anthers. The species is distinguishable by generally flat nerves and rather large male
flowers with 18 or 20 thecae. By the rather small coriaceous leaves with rounded apex it
is reminiscent of H. montana which has quite different male flowers. Horsfieldia xanthina
has similar large, pale yellowish hair scars on the leaves, but differs in other ways.

51. Horsfieldia olens W. J. de Wilde

Horsfieldia olens W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 80. — Type: NGF 31966, Papua
New Guinea.

Tree 10—35 m. Twigs 2—4 mm diameter, quadrangular by ridges from both sides of
the petiole bases (lower down ridged or lined), early glabrescent, hairs greyish or brown
0.1 mm; bark striate, not flaking; lenticels usually distinct. Leaves chartaceous, oblong
(-lanceolate), 7-14 by 2.5—6 cm, base attenuate, apex rounded, obtuse, or bluntly short-
acute-acuminate, upper surface brown to blackish on drying, sometimes with few minute
whitish dots or pustules; lower surface early glabrescent, hairs minute, 0.1 mm or less;
dots absent but irregularly shaped pustules of a different nature present; midrib raised
above; nerves 7—10 pairs, above and below thin, flattish, inconspicuous, lines of inter-
arching regular, thin; venation lax, faint on both surfaces; petiole 12—20 by 1.5—2 mm;

156 Flora Malesiana, Ser. I, Vol. 14 (2000)

leaf bud 6-10 by 1.5 mm, with hairs 0.1 mm long. /nflorescences glabrescent or with
sparse hairs 0.1—0.2 mm, short and stout, 2 or 3 times branched, peduncle 0.1—1 cm
long, rather many-flowered; in male and female (according to the infructescence): 2—6
by 1.5—4 cm, bracts not seen, caducous; flowers (male) in loose clusters of 3—5(—7),
glabrous; perianth 3- (or 4-)lobed, pedicel slender, + articulated or not. Male flowers:
pedicel 2.5—4 mm long; buds broadly ellipsoid to globose, not angular, 1.8—2.3 mm
diameter, cleft c. 5/6 or nearly to the base, lobes 0.2 mm thick; androecium + obovoid,
blunt-triangular in cross section, 1.5 by 1 mm (hence not filling the perianth) (Plate 1:
12): thecae 20-24, free apices 0.3—0.8 mm, incurved, those of one side clasping the
others: column broad, hollowed for 1/4 to 1/3; androphore broad, up to 0.1 mm long.
Female flowers not known. Fruits 2—6 per infructescence, ellipsoid, 1-1.6 by 0.8—1.2
cm, apex minutely pointed, base subattenuate, glabrous, without or with sparse small
tubercles or lenticels; pericarp 1-1.5 mm thick; fruiting pedicel 1—4 mm long; perianth
not persistent.

Field-notes —- Sometimes with small stilt-roots; bark longitudinally fissured, (red-)
brown or blackish brown, inside reddish, with some reddish exudate, with a very offen-
sive smell or a strong disinfectant smell; wood whitish to yellow. Leaves + leathery,
bluish green above (once recorded). Flowers yellow. Fruits orange(-yellow).

Distribution — Malesia: New Guinea (Papua Barat, Digul; Papua New Guinea, West-
ern Prov.).

Habitat & Ecology — Swamp edges, in fringes (with Acacia) of savanna and rain
forest; ridge forest; primary forest on level land inundated in the wet season, swampy
forest on peaty soil; Myrtaceae—Vatica~Campnosperma forest on well-drained podzolic
ground; 0—200 m altitude; fl. June; fr. Feb.—Aug.

Note — Apparently related to H. sepikensis, one of the few other New Guinean spe-
cies with 3-lobed perianths. That species differs in non-angular twig apices, more elon-
gate perianth, more slender inflorescences, membranous leaves, and apparently a differ-
ent ecology. Horsfieldia olens is mostly found in dry or wet habitats on poor peaty or
podzolic soils.

52. Horsfieldia oligocarpa Warb.

Horsfieldia oligocarpa Warb., Mon. Myrist. (1897) 354, t. 22; W.J. de Wilde, Gard. Bull. Sing. 39;
1 (1986) 22: Tree Fl. Sabah & Sarawak 3 (2000) 379. — Myristica oligocarpa (Warb.) Boerl.,
Handl. 3 (1900) 87. — Horsfieldia polyspherula (Hook. f.) J. Sinclair var. oligocarpa (Warb.)
J. Sinclair, Gard. Bull. Sing. 28 (1975) 104. — Lectotype: Beccari 2066 (FI acc. 7620), fr., Sara-
wak.

Tree 4-20 m. Twigs 2-3(—4) mm diameter, grey-brown, later on pale yellowish or
whitish brown, contrasting with the dark colour of the dry petioles, early glabrescent,
hairs 0.1 mm long; bark striate, not flaking; lenticels small, at first distinct. Leaves
chartaceous, (elliptic-)oblong, 7-16 by 2.5—6 cm, base attenuate, apex acute-acuminate,
upper surface glabrous, drying dull, olivaceous greenish, lower surface bright rusty or
chocolate, much contrasting with the upper surface, glabrous; dots absent; midrib raised
above, glabrous; nerves 8—11(—15) pairs, slender, raised above, lines of interarching and
venation indistinct at both surfaces; petiole 6-12 by 1.5—2.5 mm, drying blackish brown,
leaf bud 7-10 by 2-3 mm, with dense grey-rusty hairs 0.2 mm long, the indumentum

De Wilde — Myristicaceae (Horsfieldia) 157

early shed in the form of small crust-like pieces. Inflorescences with sparse stellate hairs
0.2—0.3 mm long, + glabrescent; in male: 2 or 3 times branched, 3—6 by 1.5—4 cm, pe-
duncle 0.5—1 cm long, few-flowered, the flowers in clusters of 3—8; in female (accord-
ing to infructescences): 1.5—4 cm long; bracts not seen, caducous; flowers glabrous,
perianth 3-lobed, pedicel articulated. Male flowers: pedicel 1—1.5 mm long, slightly
tapering; buds globose to broadly obovoid, 1—1.7 mm diameter, base + rounded, coria-
ceous, not collapsing on drying, cleft nearly 1/2, lobes towards base to 0.5 mm thick;
androecium (including androphore) + obovoid, 1 by | mm, triangular in cross section
(Plate 3: 73); thecae 12 or 14, largely free, 0.5 mm long, apex acutish, at base attached to
a short column continuous with tapering androphore 0.5 mm long. Female flowers: not
seen. Fruits 1—4 per infructescence, ellipsoid, 1.8—2.7 by 1.4—1.9 cm, glabrous, drying
(dark) brown, hardly lenticellate, pericarp 2-3 mm thick; fruiting pedicel 2-5 mm long;
perianth not persistent.

Field-notes — Shrub or tree, buttresses absent; bark grey and dark brown, or reddish
brown, fissures 0.8—1.3 cm wide, 22 cm long; inner bark pink brown, soft, 1 cm thick, sap
red; sapwood pink-yellow, soft. Timber firm. Leaves pale green, dull, not glaucous be-
neath. Flowers light brown. Fruits pale yellow to orange, pear-shaped, + pointed at apex.

Distribution — Malesia: Borneo (Sarawak, Brunei).

Habitat & Ecology — Heath forest and forest on poor soils: white (podzolic) sand,
yellow sand, sandstone, terraces, sand and peat, once recorded from a ridge; altitude
0—50(—100) m; fl. June, Aug.; fr. Aug.—Oct.

Note — Close to H. polyspherula, especially var. polyspherula, with fruits of about
the same size. Horsfieldia oligocarpa stands out by the overall pale colour, the pale
twigs, the leaves above a dull pale green on drying, contrasting strongly with the bright
brown or copper lower leaf surface (more contrasting than usually in H. polyspherula),
the small, not many-flowered, + glabrescent inflorescences, and the markedly coriaceous
(? always) flowers.

53. Horsfieldia pachycarpa A.C. Sm.

Horsfieldia pachycarpa A.C. Sm., J. Arnold Arbor. 22 (1941) 64; W.J. de Wilde, Gard. Bull. Sing.
38, 1 (1985) 133, f. 20d—g. — Type: Brass 610, West New Guinea.
Horsfieldia praetermissa J. Sinclair, in sched. (Carr 13262, etc.).

Tree 5—25 m. Twigs faintly ridged or not, 3—5 mm diameter, early glabrescent, hairs
0.1 mm; bark coarsely striate, not flaking; lenticels usually coarse and distinct. Leaves
membranous or thinly coriaceous, elliptic-oblong to lanceolate, 17—30 by 4-11 cm, base
attenuate, apex acute-acuminate; upper surface usually minutely pustulate, lower sur-
face early glabrescent; dots absent; midrib flattish above; nerves 14—18 pairs, flat above,
inconspicuous, lines of interarching below not very regular, distinct or not; venation lax,
indistinct; petiole 6-12 by 2-5 mm, leaf bud 10-15 by 2—3 mm, with hairs 0.1 mm.
Inflorescences subglabrous or with sparse stellate hairs 0.1—0.2(—0.3) mm, 2—4 times
branched; in male: 7-15 by 6—10 cm; in female up to 10 by 4 cm, peduncle 0.5—2.5 cm;
bracts caducous, not seen; flowers in loose clusters up to 5, with sparse hairs 0.1 mm,
sometimes glabrescent towards the apex, perianth 2-lobed, pedicel not articulated. Male
flowers: buds short-pear-shaped, somewhat compressed, about as broad as long or slightly
longer than broad, 2.5—3.2 by 2.8—3.2 mm, apex bluntish to broadly rounded, the lower

158 Flora Malesiana, Ser. I, Vol. 14 (2000)

2/3 + gradually passing into the thickish tapering pedicel 2-5 mm long, cleft c. 1/2,
lobes towards apex 0.2 mm thick, the perianth towards base thicker, firm-fleshy or coria-
ceous. 0.3—0.8 mm thick; androecium + flattened, 1.8—2.1 by 1.1-1.8 mm, apex broadly
rounded (Plate 2: 35); thecae 10 or 12, (or 18 or 207), erect, 1.5—2 mm long, free apical
parts 0.1—0.2 mm; androphore rather slender, (.2—0.5 mm long; column narrowly hol-
lowed for 1/6—1/3. Female flowers: pedicel 2-4 mm long; buds ovoid-ellipsoid, 2.5—4
by 2.5—3.2 mm, cleft c. 1/3; ovary ovoid, densely minutely pubescent, 2.5—3 by 1.8—2.5
mm, stigma sessile, minutely 2-lobed, 0.1 mm. Fruits 1-6 per infructescence, (broadly)
ellipsoid, often + ridged towards the base, rounded or tapering into a short pseudostalk,
apex rounded, 3.5—4.5 by 2-3 cm, minutely pubescent towards the base, or glabrescent,
drying blackish (brown), usually with conspicuous coarse, paler coloured lenticel-like
tubercles; pericarp + woody, (4—)5—10 mm thick; fruiting pedicel 5-10 mm; perianth
not persistent. — Fig. 12d-g.

Field-notes — Branches often horizontal; bark shallowly vertically fissured; exudate
watery, clear or + reddish; wood pink, cream, or whitish. Flowers yellow, fragrant. Fruits
ramiflorous, glossy green turning yellow to orange, eaten by cuscus.

Distribution — Malesia: New Guinea (W Papua Barat: Snow Mountains, Idenburg
River.; Papua New Guinea: West Sepik, Western, Eastern, and Southern Highlands, Ma-
dang, Morobe, Central Provinces).

Habitat & Ecology — Primary and degraded montane forest, fagaceous (Castanopsis-
Lithocarpus) forest; forest clearings; often on ridges; (450—)1000—2000 m altitude; fl.
& fr. throughout the year.

Notes — 1. Horsfieldia pachycarpa differs from the resembling H. tuberculata in the
thick woody-fleshy, largely pubescent perianth, and the pubescent ovary and fruit. Fruit-
ing specimens may also resemble H. laevigata, a species obviously closely related. The
male flowers of H. pachycarpa are + pear-shaped, and of a more fleshy-woody consist-
ency; those of H. laevigata are much more globose, of a more membranous-herbaceous
consistency, with the pedicel more slender.

On account of the somewhat resembling male flowers, H. pachycarpa seems related
to H. corrugata, a species from similar montane habitats, differing in the much larger
corrugated fruits and the flowers which probably always have large, thickened, blackish
dots, absent in H. pachycarpa.

Horsfieldia tuberculata var. crassivalva (from the Louisiade Archipelago), known
only from fruit with also a thick pericarp, is similar as well.

2. Some deviating specimens (almost all collected above 1000 m altitude) have been
discussed by De Wilde (1.c.: 134); the female perianths measure 2-2.8 by 1.8—2.3 mm,
the fruits 2.5-3.5 by 1.7—2 cm, and they have a thick woody pericarp. In appearance and
size of the fruits these specimens seem intermediate with the widespread and common
H. laevigata.

54. Horsfieldia pachyrachis W. J. de Wilde

Horsfieldia pachyrachis W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 5. — Type: bb 28128, West
Kalimantan.

Tree. Twigs 5-7 mm diameter, blackish brown, glabrescent, hairs rusty to grey-brown,
0.2 mm: bark of older twigs not seen; lenticels conspicuous. Leaves thinly chartaceous,

De Wilde — Myristicaceae (Horsfieldia) 159

(obovate-)oblong, 16-26 by 6.5—9 cm, base (long-)attenuate, apex broadly acutish; up-
per surface drying olivaceous to blackish brown, glabrous, lower surface drying dark
brown, early glabrescent but on the midrib towards the base vestigial hairs 0.2-0.3 mm;
dots absent; midrib raised above, near the transition of the petiole 2.5—3 mm wide; nerves
14-17 pairs, raised above, lines of interarching not very distinct; venation lax, + flat,
indistinct; petiole 7-11 by 3.5—4.5 mm; leaf bud 15 by 4 mm, with hairs 0.2 mm. Inflo-
rescences behind the leaves, with dense or sparse dendroid hairs 0.2—0.3 mm, glabrescent;
in male: stout and compact, 4 or 5 times branched, many-flowered, 14 by 10 cm (not
fully expanded), the main axis stout, towards the base 5-8 mm diameter, length of
peduncle not known (broken off); bracts caducous, those of uppermost ramifica-
tions + elliptic, subacute, densely pubescent, 3-5 by 2—4 mm; flowers in clusters of
4-10, perianth 3-lobed, glabrous, pedicel with sparse hairs 0.1—0.2 mm long towards
base, articulated. Male flowers (submature): pedicel 1.5—2 mm long; buds broadly glo-
bose to broadly obovoid, 1.5 by 2 mm, apex broadly rounded, slightly depressed, base
+ narrowly rounded, glabrous, cleft nearly 1/2, hard-fleshy, not collapsing on drying,
lobes 0.2—0.3 mm thick, perianth towards the base 0.5—0.7 mm thick; androecium sub-
globose-obovoid, 0.6 by 0.6 mm, + sharp-triangular in cross section (Plate 3: 65); thecae
10-14, slightly incurved, connate, concealing the apical cavity reaching to 1/4—1/2 of
the column; androphore slightly tapering, 0.1—0.2 mm long. Female flowers and fruits
not seen.

Distribution — Malesia: Borneo (W Kalimantan: Melawi, Bukit Kelawai, known
only from the type).

Habitat & Ecology — Altitude 80 m; fl. May; no further ecological data known.

Note — The male flowers resemble those of the group of species with H. polyspherula
(especially var. sumatrana) or H. laticostata, particularly the distinctly triquetrous androe-
cium. However, in the H. polyspherula-group the (sub)erect anthers are mutually free
for at least about halfway, whereas in H. pachyrachis the anthers are (almost) com-
pletely connate. In general habit and the colour of the leafy twig, H. pachyrachis some-
what resembles the H. polyspherula-group. The connate anthers (and the appearance of
the leaves) point to the group of species keyed out around H. fragillima.

55. Horsfieldia pallidicaula W.J. de Wilde

Horsfieldia pallidicaula W.J. de Wilde, Gard. Bull. Sing. 38, 2 ((1985’, 1986) 191; Tree Fl. Sabah
& Sarawak 3 (2000) 380. — Type: Jacobs 5413, Sarawak.

Tree 7-20 m. Twigs 2-10 mm diameter, pale, whitish or greyish brown, contrasting
with the blackish drying colour of the petioles (which often are greyish and sometimes
flaky in the lower half), early glabrescent, hairs grey-brown, 0.1—0.3 mm long; bark
finely striate, slightly flaking or not; lenticels inconspicuous or absent. Leaves either in
2, 3, or 5 rows, membranous, oblong(-lanceolate), 10—30 by 4—9.5 cm, base long-atten-
uate, apex acute-acuminate; upper surface drying (blackish) brown, lower surface dry-
ing (grey-)brown, early glabrescent; dots absent; midrib slightly raised above, glabrous;
nerves 10—18 pairs, slender, flat or slightly raised above, lines of interarching indistinct;
venation lax, indistinct; petiole 10-25 by 1.5-3 mm; leaf bud stout or slender, 7-10 by
2—4 mm, with brown-grey or rusty hairs (0.1—)0.2(—0.3) mm long. /nflorescences gen-
erally behind the leaves, glabrescent, hairs sparse, 0.1—0.2 mm; in male: (2 or) 3 times

160 Flora Malesiana, Ser. I, Vol. 14 (2000)

branched, many-flowered, 3—9 by 2.5—7 cm, peduncle up to 0.7 cm long (inflorescences
often branched from near the base), the flowers in loose clusters of 2—4; in female:
rather few-flowered, 1-3 cm long; bracts lanceolate, acute, finely pubescent, 1-3 mm
long, caducous; flowers glabrous, perianth 3- or 4-lobed, in female sometimes 2-lobed,
pedicel not articulated. Male flowers: pedicel 1-2 mm, slender; buds globose or broadly
obovoid, 1.5—2(—2.2) mm diameter, base rounded to short-attenuate, cleft c. 1/3 to near-
ly 1/2, not or but slightly collapsing on drying, lobes 0.2—0.3 mm thick; androecium
short-ellipsoid or depressed-globose, 0.8—1.2 by 0.8—1.2 mm, circular in cross section
(Plate 2: 43); thecae 16-20, completely sessile (free apices 0.1 mm long), incurved to-
wards the apex of the androecium; column broad with narrow apical cavity, 0. 1—0.2 mm
deep; androphore narrow, 0—0.2 mm long. Female flowers: pedicel 1—1.5 mm long;
buds broadly ellipsoid or subglobose, 2.5—3 by 2.5 mm, cleft 1/4-1/3; ovary broadly
ovoid, 1.7-2 by 1.5-2 mm, glabrous, stigma lobes 0.1—0.2 mm long. Fruits 1-5 per
infructescence, broadly ellipsoid, 1.5—6 by 1-3 cm, glabrous, drying blackish, finely
granulate and sometimes tubercled; pericarp variable; fruiting pedicel 1—3 mm long;
perianth persistent under the fruits (always?) (see under the varieties).

Distribution — Malesia: Borneo (Sabah and Sarawak).

Note — The three varieties are sympatric and similar in the vegetative characters.
They are mainly distinguished by the remarkable differences in fruit size, and hence
flowering material cannot be named with certainty.

KEY TO THE VARIETIES

la. Fruits 1.5-1.7 by 1-1.2 cm; pericarp 1-1.5 mm thick. Leaves up to 23 cm long....
np aS see Mat ARN ae A A NAA int og Rh rhea c. var. microcarya

b. Fruits larser) 2-6 by 1/8=3' cms: Pere... eo 2
2a. Fruits 2—2.2 by 1.8 cm; pericarp 2-3 mm thick. Leaves up to 25cm long........-.
dra amie ar aL [i Ay Ral ie soil 9 aan nr te at a. var. pallidicaula

b. Fruits 3.7-6 by 2.7-3 cm; pericarp 6—-8(—10) mm thick. Leaves 25-30 cm lone...
| AAT ere CSN Re Ske SNMP tate te Oe ese Sota cca aneED eee b. var. macrocarya

a. var. pallidicaula

Leaves 10-25 cm long, up to 7.5 cm wide. Fruits 2-2.2 by 1.8 cm; pericarp 2-3 mm
thick; fruiting pedicel and perianth not seen.

Field-notes — Bark smooth, dark grey; sapwood white-red. Perianth yellow or yel-
low-green, androecium pale pink-yellow, pollen white.

Distribution — Malesia: Borneo (Sarawak, West Sabah; West Kalimantan, doubtful,
see note 2).

Habitat & Ecology — Primary lowland and lower montane forest; sandstone; 0-700
m altitude; fl. throughout the year; fr. June.

Notes 1. Horsfieldia pallidicaula vegetatively resembles the related species H.
sucosa and H. sterilis, both essentially differing in the male flowers.

2. Winkler 1435, from West Kalimantan, deviates in the very thin membranous leaves
which dry greenish; the flowers are smaller and differ in details (see De Wilde, l.c.: 193).
The specimen was collected in a marshy forest at c. 50 m altitude.

De Wilde — Myristicaceae (Horsfieldia) 161

b. var. macrocarya W.J. de Wilde

Horsfieldia pallidicaula W.J. de Wilde var. macrocarya W. J. de Wilde, Gard. Bull. Sing. 38, 1 (1985)
193; Tree Fl. Sabah & Sarawak 3 (2000) 381. — Type: Ding Hou 474, Sarawak.

Leaves 23-30 by 6—9.5 cm. Fruits 3.5—6 by 2.5—3 cm; pericarp 6—8(—10) mm thick;
fruiting pedicel 1-2 mm long, persistent perianth 2- or 3-lobed.

Field-notes — Bark smooth to + flaky, or longitudinally fissured. Fruits and aril pink.

Distribution — Malesia: Borneo (Sarawak: 4th and 5th Div.).

Habitat & Ecology — Lowland mixed dipterocarp forest; yellow sandy clay soil; alti-
tude 30-200 m; fr. July, Oct.

c. var. microcarya (J. Sinclair) W.J. de Wilde

Horsfieldia pallidicaula W.J. de Wilde var. microcarya (J. Sinclair) W.J. de Wilde, Gard. Bull. Sing.
38, 1 (1985) 193: Tree Fl. Sabah & Sarawak 3 (2000) 381. — Horsfieldia bracteosa Henderson
var. microcarya J. Sinclair, Gard. Bull. Sing. 28 (1975) 20. — Type: SAN 16971 , Sabah.

Leaves to 20 cm long, up to 7.5 cm wide. Fruits 1.5—1.7 by 1—1.2 cm; pericarp 1—1.5
mm thick; fruiting pedicel 2 mm long, persistent perianth 3-lobed.

Distribution — Malesia: Borneo (West Sabah, E Kalimantan).

Habitat & Ecology — Lowland forest at c. 100 m altitude; fr. July.

56. Horsfieldia parviflora (Roxb.) J. Sinclair

Horsfieldia parviflora (Roxb.) J. Sinclair, Gard. Bull. Sing. 28 (1975) 82; W.J. de Wilde, Gard.
Bull. Sing. 38, 1 (1985) 69. — Myristica parviflora Roxb., Fl. Ind. 3 (1832) 847; (ed. 1874) 744;
Icones 2574. — Type: Roxburgh’s description and figure; Calcutta, culta.

Myristica tingens Blume, Rumphia | (1837) 190. — Horsfieldia sp. Merr., Int. Rumph. (1917) 231.
— Based on Palala minima, P. tertia, P. tingens Rumph., Herb. Amb. 2, 10 (1741) 27, t. 7 f. A,
B; see Sinclair, 1975: 161.

Myristica globularia Blume, Rumphia | (1837) 191, t. 64 f. 2 (non Lamarck). — Pyrrhosa globularia
(Blume) Hassk., Cat. Pl. Hort. Bog. (1844) 174. — Horsfieldia globularia (Blume) Warb., Mon.
Myrist. (1897) 288, t. 21 1-4. — Palala globularia (= P. quinta) Rumph., Herb. Amb. 2, 10
(1741) 28, t. 9 f. a-b; see Sinclair, 1975: 165. — Type: Blume’s figure, and Zippel s.n. (Ambon,
‘mas’ ), a sterile specimen.

Myristica bivalvis Hook. f., Fl. Brit. India 5 (1886) 107; King, Ann. Roy. Bot. Gard. Calc. 3 (1891)
307, pl. 139. — Horsfieldia bivalvis (Hook. f.) Merr., Philipp. J. Sci., Bot. 2 ((1916’, 1917) 271;
J. Sinclair, Gard. Bull. Sing. 16 (1958) 379, f. 32, pl. VHI-B. — Type: Murton 149, Java, culta.

Horsfieldia globularia (Blume) Warb. var. minahassae Warb., Mon. Myrist. (1897) 617. —
Horsfieldia minahassae (Warb.) Koord., Meded. Lands Pl. Tuin 19 (1898) 70. — Syntypes:
Koorders 18123, 18124, 15146, 18164 (lecto), Sulawesi.

Tree 10-20 m. Twigs somewhat flattened but not angular, 2-5 mm diameter,
glabrescent, hairs 0.1 mm; bark finely striate, not flaking; lenticels abundant, small, not
conspicuous. Leaves membranous, (oblong-)lanceolate, 8—23 by 2.5—7.5 cm, base at-
tenuate, apex acute-acuminate; upper surface drying olivaceous or brown, dull (some-
times with fine pale pustules or larger irregularly shaped marks), lower surface early
glabrescent; dots absent; midrib flat above; nerves 10—15 pairs, flat, inconspicuous above,
lines of interarching indistinct; venation fine, indistinct; petiole 6-16 by 1.5—2.5 mm;
leaf bud 6-13 by 1—2 mm, with hairs 0.1 mm long. /nflorescences with sparse or dense
hairs 0.1—0.3 mm; in both male and female: 3 or 4 times branched, many-flowered, 4—10

162 Flora Malesiana, Ser. I, Vol. 14 (2000)

by 4-8 cm, peduncle 1—2 cm long; bracts pubescent, elliptic to oblong, 2-5 mm long,
caducous: flowers in loose clusters of 2—4, perianth 2-lobed, glabrous or in female some-
times minutely pubescent at base, pedicel with sparse hairs 0.1 mm long or less, not
articulated. Male flowers: pedicel slender, 1-2 mm long; buds somewhat laterally com-
pressed, obtriangular to transversely ellipsoid, 2.2-3 by 2.5—4 mm, apex broadly rounded,
at base short-attenuate, firm, not collapsing on drying, bright brown or with a grey-blue
tinge, cleft c. 1/2, lobes (0.1—)0.2—0.3 mm thick; androecium transversely ellipsoid or
+ obtriangular, not or slightly laterally compressed, largely hollow, (1-)1.6—2.2 by 1.6—
3 mm (Plate 1: 6); thecae 36—50, completely connate, forming a thin-walled cup, the
anthers (sometimes only on one side of the androecium) completely inflexed from the
middle and reaching nearly the bottom of the cup, free apices 0.1 mm; androphore nar-
row, 0.1—0.3 mm long. Female flowers: pedicel 1-2 mm, with sparse hairs 0.1 mm; buds
ellipsoid, (2.5—)3—3.5 by 2.5 mm, cleft c. 1/3; ovary ovoid-ellipsoid, 2—2.3 by 1.5 mm,
glabrous, style and stigma 2- (or 3-)lobed, 0.1—0.2 mm long. Fruits 2-10 per infruc-
tescence, ellipsoid to nearly globose, 1.1—1.6(—2) by 1-1.3 cm, glabrous, finely granu-
late, not or hardly tuberculate; pericarp 1—-1.5 mm thick; fruiting pedicel 24 mm long;
perianth not persistent.

Field-notes — Tree without buttresses; bark smooth or fissured; wood whitish. Flow-
ers yellow, fragrant; anthers yellowish white. Fruits behind the leaves, yellow or light
brown; aril bright red, once recorded as yellow (unripe’”).

Distribution — Malesia: Sulawesi (including Kabaena I.), Moluccas (Seram, Am-
bon), Singapore (running wild in the Garden’s Jungle of the Botanic Garden).

Habitat & Ecology — Forests; once on sandy loam; 0—600 m altitude; fl. & fr. through-
out the year.

Notes — 1. Elbert 3457 (Kabaena I., limestone; S Sulawesi) has rather large fruits,
1.6 cm long, with a distinct pseudostalk 1.5—2 mm long; fruit of Darnaedi 2287 (S Cen-
tral Sulawesi) is 2 cm long.

2. Horsfieldia parviflora is easily distinguishable by the smooth, rather inflated male
buds, + obtriangular to transversely ellipsoid in lateral view, not collapsing on drying,
usually of a bluish or reddish brown tinge.

57. Horsfieldia paucinervis Warb.

Horsfieldia paucinervis Warb, Mon. Myrist. (1897) 345, t. 22: Merr., Enum. Born. (1921) 268;
J. Sinclair, Gard. Bull. Sing. 28 (1975) 93; W.J. de Wilde, Gard. Bull. Sing. 38, 2 ((1985’, 1986)
212: Blumea 41 (1996) 379, f. la—c; Tree Fl. Sabah & Sarawak 3 (2000) 381. — Myristica
paucinervis (Warb.) Boerl., Handl. 3 (1900) 87. — Type: Beccari 3279 (female fl.), Sarawak.

Fig. 24. Some taxonomically related Horsfieldia species. — H. paucinervis Warb. a. Habit of male
flowering twig; b. longitudinally opened male flower and schematic section of androecium; c. twig
with fruit. — H. aff. gracilis W.J. de Wilde. d. Twig with male inflorescence; e. male flower and
section of androecium. — H. gracilis W.J. de Wilde. f. Twig with mature fruit; note persistent peri-
anth. — H. cf. gracilis W.J. de Wilde. g. Male flower and section of androecium. — H. reticulata
Warb. h. Male flower and section of androecium; i. fruit; note persistent perianth [a, b: Purse-
glove P 4403: c: KeBler 319; d, e: S 50154 (Paie); f: S 16604 (Paie); g. Wong KMW 1494; h: BRUN
5654: i: S 36018 (Chai). — Scale bar for a, d, f, i= 2 cm; for b, e, g,h = 1 mm; for ¢ = -2/em:

163

De Wilde — Myristicaceae (Horsfieldia)

164 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 3-8 m. Twigs 2-3 mm diameter, late glabrescent, hairs reddish to yellow-rusty,
shaggy, 1.5—-2 mm long; bark dark grey, striate, not cracking; lenticels inconspicuous.
Leaves thinly chartaceous, elliptic(-oblong), 7-15 by 3-6.5 cm, base rounded to attenu-
ate, apex acute(-acuminate); upper surface drying olivaceous or brown, glabrous, lower
surface dull brown with persistent + sparse dendroid hairs of mixed size, 0.5—1 mm
long; dots (or dashes) present (always?); midrib flat above, with persistent indumentum
or late glabrescent; nerves 5—9(—11) pairs, flat or sunken, lines of interarching indistinct;
venation hardly visible; petiole 6-12 by 1.5—2.5 mm, pubescent: leaf buds 8-10 by 3-4
mm, hairs 1(—2) mm long. Inflorescences with dense woolly hairs (0.7—)1—2 mm long;
in male: many-flowered, 3 or 4 times branched, 5—9 by 3-5 cm, peduncle to 1.5 cm long;
in female: 4—6 cm long; bracts caducous; flowers (in male several in loose clusters)
glabrous, perianth 3-lobed, pedicel not articulated. Male flowers: pedicel slender, 0.5—1
mm long; buds subglobose, 0.8—1 by 1—1.4 mm, cleft c. 1/3 to nearly 1/2, lobes 0.1 mm
thick: androecium small, + depressed-globose, 0.3—0.5 by 0.4—0.8 mm, + circular in
cross section (Plate 2: 56); thecae 8 or 10, almost completely sessile, the tips incurved;
column broad, with a minute apical hollow c. 1/5 deep, androphore narrow, somewhat
tapering, 0.1—0.2 mm long. Female flowers (Warburg, l.c. & Sinclair, l.c.): pedicel 2mm
long, glabrous; buds ovoid-globose, 2 mm diameter, cleft slightly over 1/2; ovary sub-
globose, glabrous, stigma minute. Fruits (immature) oblong, obtuse at both ends, gla-
brous, 1 by 0.7 cm; fruiting pedicel 2 mm long; perianth persistent. — Fig. 24a—c.

Field-notes — Shrubs or slender trees; sap pale pink, watery. Flowers yellow.

Distribution — Malesia: Borneo (Sarawak: Ist, 2nd, 4th Div.; W Kalimantan).

Habitat & Ecology — Coastal kerangas and degraded forest on eroded white sand;
0-100 m altitude; fl. throughout the year.

Note — Horsfieldia paucinervis belongs to the group of species with pubescent leaves
to which also belong H. gracilis, H. reticulata, H. rufo-lanata, H. splendida, and H. to-
mentosa; it seems especially related to H. gracilis.

58. Horsfieldia penangiana J. Sinclair

Horsfieldia penangiana J. Sinclair, Gard. Bull. Sing. 16 (1958) 408, f. 42; 28 (1975) 94; W. J. de
Wilde, Gard. Bull. Sing. 39, 1 (1986) 52; Blumea 41 (1996) 379; Tree Fl. Sabah & Sarawak 3
(2000) 382. — Type: Curtis 2406, Peninsular Malaysia, Penang.

Tree 4-25 m. Twigs 1.5—2(—4) mm diameter, early glabrescent, hairs grey-brown, to
0.1 mm long; bark finely striate, not flaking; lenticels small, conspicuous or not. Leaves
membranous to (thinly) coriaceous, (elliptic-)oblong, 5—12(-17) by 2-4 cm, base at-
tenuate, apex acute-acuminate or rounded, upper surface drying olivaceous to blackish
brown, lower surface early glabrescent; dots present (lens!): midrib flattish or raised
above: nerves 8-11 pairs, flat, inconspicuous above, lines of interarching indistinct; ve-
nation lax, hardly visible; petiole 8-13 by 1-2 mm; leaf bud 6-9 by 1-1.5 mm, with
dense greyish brown hairs 0.1 mm. Inflorescences with sparse hairs 0.1 mm; in male:
3 or 4 times branched, moderately to many-flowered, 2—7 by 1.5—4.5 cm, peduncle 0.2—
2 cm long; in female (from infructescences): 2-5 cm long; bracts ovate-oblong, short-
pubescent, 1.5—2.5 mm long, caducous; flowers in male in loose clusters of 2-5, gla-
brous, perianth 2—4-lobed, pedicel + articulated or not. Male flowers: pedicel 0.8—2 mm,
slender, buds variable in shape, subglobose or ellipsoid, + circular to faintly triangular in

De Wilde — Myristicaceae (Horsfieldia) 165

cross section, 1.2—1.8 by 1-1.5 mm, cleft c. 1/3—1/2, lobes 0.2 mm thick; androecium
either broadly ellipsoid, globose, or depressed-globose, 0.6—0.7 by 1 mm, or mostly
ellipsoid, 0.7—1.5 by 0.6—1.2 mm (Plate 3: 89); thecae 10—18(—20) or 24(—30), almost
completely sessile, free apices to 0.3 mm, erect or slightly incurved, column with nar-
row shallow hollow, androphore up to 0.1 mm long. Female flowers not seen. Fruits
2-6 per infructescence, ovoid-ellipsoid, 1.1—2 by 0.9—1.6 cm, apex (narrowly) rounded,
base rounded, glabrous, finely wrinkled or granulate, without tubercles; pericarp 1.5
mm thick; fruiting pedicel 3—4 mm long, articulated; perianth not persistent.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Borneo (Sarawak, E Kali-
mantan); 2 subspecies.

Note — Horsfieldia penangiana can be recognized by the slender twigs with + small,
dotted leaves, very short hairs on leaf buds and inflorescences, globose or ellipsoid male
buds, and by an ellipsoid androecium which is (sub)circular in cross section. In habit
it resembles Gymnacranthera eugeniifolia. Sterile specimens may also recall H. ridleyana.
Taxonomically H. penangiana seems close to H. glabra, especially its var. javanica.

KEY TO THE SUBSPECIES

la. Leaf apex acute or acute-acuminate. — Sumatra, Peninsular Malaysia...........
emmrnterss -13)¢25274 hod) ie PL te ly tdi cistenes ly bei a. subsp. penangiana
beleatapex blunt or rounded. Bormeo..,:... 3.4..-.--s2-.4e: b. subsp. obtusifolia

a. subsp. penangiana

Myristica griffithii auct. non Hook. f.: King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 31, p.p., quoad
Curtis 2406, 2458. — Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. griffithii
auct. non (Hook. f.) Warb.: Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 226, p.p.

Leaf blades elliptic-oblong, 6—12 cm long, apex acute or acute-acuminate. Male buds
(2- or) 3- or 4-lobed, broadly ellipsoid or (sub)globose, 1.2—1.8 mm long, thecae 10-
18(—20). Fruits 2 by 1.6 cm.

Distribution — Malesia: Sumatra, Peninsular Malaysia.

Habitat & Ecology — Primary dryland forest, ridge-top or montane forest; 0—1300 m
altitude; fl. June; fr. Aug.

Note — Perianths are generally 3- (or 4-)lobed, but occasionally 2-lobed; the shape
of the mature male bud is also variable, and the pedicel is articulated or not.

b. subsp. obtusifolia W. J. de Wilde

Horsfieldia penangiana J. Sinclair subsp. obtusifolia W.J. de Wilde, Blumea 41 (1996) 379; Tree
Fl. Sabah & Sarawak 3 (2000) 382. — Type: S 47061 (Awa & Othman), Sarawak.

Leaf blade (elliptic-)oblong, 6—14(—17) cm long, apex blunt or rounded. Male buds 3-
lobed, globose, (1.5—)2 mm diameter, thecae 24(—30). Fruits 1.5 by 1 cm.

Field-note — Almost mature male flower buds green; mature fruits dark green.

Distribution — Malesia: Borneo (Sarawak, E Kalimantan).

Habitat — Forest of ridge tops, hills; (SO—)300—800 m altitude.

Note — Fruiting pedicels are articulated at the base, the male flower pedicels are
partly articulated.

166 Flora Malesiana, Ser. I, Vol. 14 (2000)

59. Horsfieldia perangusta W. J. de Wilde

Horsfieldia perangusta W.J. de Wilde, Blumea 41 (1996) 379. — Type: KEP FRI 7934 (Cockburn),
Peninsular Malaysia.

Slender tree, 10—12 m. Twigs 3(—4) mm diameter, at first with sparse, shaggy, dark
rusty-red hairs | mm long, + early glabrescent; bark dark brown, coarsely striate, not
cracking, not or inconspicuously lenticellate. Leaves chartaceous or thinly coriaceous,
elliptic-oblong, 20-25 by 7-10 cm, base cuneate, apex acute-acuminate; upper surface
drying olivaceous, nearly glossy, below + pale cinnamon, at first with + sparse, + ap-
pressed, shaggy pale hairs 0.5—1 mm, glabrescent except on and near the midrib and
blade base, margin conspicuously rolled-in; dots absent; midrib sunken above, slender
(appearing as a thin line); nerves 10-12 pairs, inconspicuous above, lines of interarching
visible only below; venation + invisible on both surfaces; petiole 17-20 by 3-4 mm,
glabrescent; leaf bud 10 by 3 mm, with dense red-rusty hairs | mm long. /nflorescences
(from infructescences) on the older twigs below the leaves (twig diameter 5—8 mm),
sparsely branched, glabrous (glabrescent), 1-2 cm long. Flowers not known. Fruits 1-3
per infructescence, ellipsoid, 25-30 by 18-20 mm, apex blunt, base rounded; pericarp
2-3(—4) mm thick, + glabrous, (possibly) with remnants of minute, rusty, scurfy hairs at
very base (hence, ovary minutely pubescent); fruiting pedicel at first with sparse minute
indumentum, 2 by 3 mm; seed ellipsoid, 20 by 14-15 mm, light brown; perianth not per-
sistent.

Field-notes — Slender tree to 12 m, girth c. 35 cm; bark dark chocolate brown, smooth,
rugose; slash inner bark red-orange, fibrous, laminated; slash wood white. Fruits orange,
seed with a deep orange-red leathery aril.

Distribution —Malesia: Peninsular Malaysia (Pahang, Udu Endau Forest Reserve,
compt. 285), known only from the type.

Habitat & Ecology — Ridge-top forest dominated by palms; 400 m altitude; fr. Mar.

Note — The present species possibly is close to H. fulva, which differs in its leaves
becoming dull and finely wrinkled above on drying. Horsfieldia perangusta has con-
spicuously incurved blade margins, and a narrow line-shaped midnerve above. Flowers
are not known. Male buds of H. fulva are conspicuously elongate, a feature uncommon
in Horsfieldia.

60. Horsfieldia pilifera Markgr.

Horsfieldia pilifera Markgr., Bot. Jahrb. Syst. 67 (1935) 154; W.J. de Wilde, Gard. Bull. Sing. 38,
1 (1985) 121, f. 17). — Type: Ledermann 10450, Papua New Guinea.

Horsfieldia novo-guineensis Warb., Mon. Myrist. ( 1897) 271, p.p. (Hollrung 657, syntype; lectotype
= Horsfieldia aruana).

Tree (5—)10—20 m. Twigs lined or faintly ridged or not, 1.5—4(-10) mm diameter,
early glabrescent, hairs grey to brown, 0.1 mm; bark striate, not flaking; lenticels mostly
inconspicuous. Leaves membranous, elliptic to oblong-lanceolate, 7-27 by 2.5—8.5 cm,
base attenuate, apex acute-acuminate; upper surface usually minutely whitish pustulate,
lower surface glabrescent, hairs stellate, 0.1 mm long; dots absent; midrib hardly raised
above; nerves 7-16 pairs, above thin, flat, lines of interarching faint; venation lax, faint
on both surfaces; petiole 6-12 by 1.5—2.5 mm; leaf buds 10 by 1-2 mm, with hairs 0.1

De Wilde — Myristicaceae (Horsfieldia) 167

mm long. Inflorescences with sparse or dense rather woolly hairs 0.1—0.3 mm; in male:
2-4 times branched, many-flowered, 5—12 by 4—8 cm, peduncle 1—2.5 cm; female 4—12
cm long; bracts 1—-2(—3) mm long, caducous; flowers (male) in clusters of 2—5; perianth
2-lobed, with sparse or dense stellate hairs 0.1 mm long; pedicel slender, finely pubes-
cent, not articulated. Male flowers: pedicel 1-2 mm long; buds little to much flattened,
subcircular or slightly transversely or longitudinally elliptic, (1—)1.2—1.8 by 1.2-1.9 mm,
apex broadly rounded, base rounded or short-attenuate, cleft 1/3—1/2, lobes 0.1 mm thick;
androecium flattened, subquadrangular in outline, 0.7—1.2 by 0.6—-1.1 mm, apex broad-
ly rounded (Plate 1: 30); thecae 16-20, (sub)erect, sessile, free apices to 0.1 mm long,
column narrowly hollowed for 1/5—1/4; androphore up to 0.1 mm, broadly attached.
Female flowers: pedicel 2 mm long; buds broadly ellipsoid, 2.8 by 2.5 mm, cleft 1/2—
2/3; ovary globose to ovoid, 1.5 by 1.3 mm, with hairs 0.1 mm or less, style and stigmas
0.1 mm long. Fruits (2—)5—20 per infructescence, globose or short-ellipsoid, 1.1—1.6 by
1.1-1.6 cm, glabrescent but with minute persistent hairs towards the base (lens!), with-
out or with few tubercles; pericarp often + woody, thickest at one side, 1-3 mm thick;
seed ellipsoid; fruiting pedicel 1-5 mm long; perianth not persistent. — Fig. 22).

Field-notes — Bark longitudinally fissured; sap watery, turning pink or red; wood
straw to brown, of moderate weight and hardness. Flowers yellow. Fruits hard, glossy
green, turning dark yellow, orange or red.

Distribution — Malesia: northern half of New Guinea (Papua Barat: Bird’s Head,
Japen I., Jayapura; Papua New Guinea: Sepik, Madang, Morobe Prov.).

Habitat & Ecology — Primary and degraded forest, on sandy loam soil, mixed forest
with Anisoptera at c. 100 m; 0-1000 m altitude; fl. & fr. throughout the year. Fruiting
once reported as very prolific.

Note — Close to H. laevigata, but noticeable for the smaller male buds and the smaller
(sub)globose fruits without or with few (coarser and paler coloured) lenticel-like tuber-
cles. The male buds are accepted as variable in outline, varying from lengthwise to
+ transversely ellipsoid. The only known female flowers (Schlechter 16933) are some-
what smaller than those of H. laevigata, but they probably do not differ significantly in
the two species. In fruit (sometimes globose) H. pilifera may be confused with H. basifissa,
a species with quite different male flowers.

61. Horsfieldia polyspherula (Hook.f. emend. King) J. Sinclair

Horsfieldia polyspherula (Hook. f. emend. King) J. Sinclair, Gard. Bull. Sing. 16 (1958) 422, f. 47,
pl. XII-B; W.J. de Wilde, Tree Fl. Sabah & Sarawak 3 (2000) 383. — Myristica polyspherula
Hook. f., Fl. Brit. India 5 (1886) 108, p.p. (see notes by Sinclair, 1958: 425; 1975: 103). —
Lectotype (Sinclair, 1975: 103): Griffith 4354, Peninsular Malaysia.

For more references and synonyms see the varieties.

Tree 4—35(—40) m. Twigs terete or subterete, 2-5 mm diameter, never lined or ridged,
early or late glabrescent, hairs rusty, mealy, 0.1—0.6 mm long; bark finely or coarsely
striate, neither cracking nor flaking, lenticels distinct or not. Leaves thickly mem-
branous to chartaceous, very brittle, (elliptic-)ovate-oblong to oblong-lanceolate, 7—28
by 2.5—9 cm, base + rounded or usually attenuate, apex acute-acuminate; upper surface
drying olivaceous or (greenish) brown, lower surface light brown or chocolate-brown,
usually much contrasting with the upper surface, early glabrescent but midrib often late

168 Flora Malesiana, Ser. I, Vol. 14 (2000)

glabrescent; dots absent; midrib slender; nerves 6—15(—20) pairs, distinctly raised above,
lines of interarching usually not distinct; venation lax, faint; petiole 6-15 by 1.5—3 mm,
sometimes late glabrescent; leaf bud 6-17 by 1.5-3 mm, with rusty dendroid hairs 0.1—
0.6 mm long. Inflorescences with sparse to dense woolly hairs 0.6 mm long, sometimes
glabrescent; in male: 3—5 times branched, many-flowered, 4—15(—20) by 3-12 cm, pe-
duncle 0.3—1.5 cm long, the flowers in clusters of up to 8 each; in female: few- to many-
flowered, up to 8 cm long; bracts oblong to lanceolate, 1.5—7 mm long, densely pubes-
cent, caducous; flowers with perianth 3-lobed, glabrous or glabrescent, pedicel glabrous
or minutely pubescent towards the base, articulated. Male flowers: pedicel slender, some-
times tapering, 0.8—1.5(—2) mm long; buds globose or broadly obovoid, 1—1.8 mm diam-
eter, apex (broadly) rounded, base rounded or + tapering into the pedicel, cleft 1/2—2/3,
not collapsing on drying, lobes 0.2—0.4 mm thick, at base up to 0.6 mm thick; androecium
(including androphore) + broadly obovoid in outline, 0.5—0.8 by 0.6—1 mm, triangular
in cross section (Plate 3: 72); thecae (6 or) 8-14, free for at least halfway, + curved or
suberect, 0.3—0.5 mm long, apex acutish, at base column short bowl-shaped, narrowed
into + tapering androphore, (0.2—)0.3—0.4 mm long. Female flowers: pedicel 1—-1.8 mm
long, minutely pubescent; buds broadly ellipsoid-obovoid, 2—3 by 1.8—2.8 mm, glabrous
or with sparse hairs 0.1 mm, cleft 1/3—2/3; ovary ovoid-ellipsoid, 1.2—1.5 by 1—-1.5 mm,
glabrous, the stigma shallowly 2-lobed, 0.2-0.3 by 0.5 mm. Fruits 1—6 per infructes-
cence, subglobose or ellipsoid, apex rounded, base rounded or slightly attenuate, 1.9—6
by 1.4—5 cm, glabrous, not lenticel-like tuberculate; pericarp 2-15 mm thick; fruiting
pedicel 1-4 mm long; perianth not persistent.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Borneo, Philippines (Luzon,
Mindanao, Palawan; only var. polyspherula); 0—1100 m altitude.

Note — Horsfieldia polyspherula is a variable species of a complex to which belong
closely related species such as H. laticostata, H. oligocarpa, and H. tenuifolia (all from
Borneo), H. brachiata (Sumatra to Borneo), and H. majuscula (Peninsular Malaysia).
Variability is found in the thickness of twigs, size and texture of leaves, density of
indumentum on leaf buds and twig apex, and especially in fruit size and thickness of the
pericarp. On the basis of mainly fruit characters, arbitrarily three rather heterogeneous
varieties can be recognized. There are slight differences in the size of the male perianth,
mainly on account of the varying thickness of its lobes; the number of thecae is possibly
the same in all three varieties.

The anthers have two rather widely separated, almost mutually free thecae, giving the
impression as if there are twice as many anthers than actually are present.

KEY TO THE VARIETIES

la. Fruits (when dry) (3.5—)4—6 cm long, pericarp 5-15 mm thick. Male flowers not

known. Leaves 9-20 cm long, nerves 9-15 pairs........--+--- b. var. maxima

b. Fruits up to 3.5 cm long, pericarp 2-5 mm thick ......--.--.++ss+eeereeees 2
2a. Fruits (2.5—)2.8-3.5 cm long. Male buds 1.2—1.8 mm diameter, thecae 12 or 14.
Leaves 13-28 cm long, nerves 11—15(—20) pairs.........---- c. var. sumatrana

b. Fruits (1.7—)1.9-2.5(—2.8) cm long. Male buds 1—1.5 mm diameter; thecae (6 or) 8—
14. Leaves 7-19 cm long, nerves 6-15 pairs.......------- a. var. polyspherula

De Wilde — Myristicaceae (Horsfieldia) 169

a. var. polyspherula

Horsfieldia polyspherula (Hook. f. emend. King) J. Sinclair, Gard. Bull. Sing. 16 (1958) 422, f. 47,
pl. XII-B; 28 (1975) 101, p.p., for the type variety only; W.J. de Wilde, Gard. Bull. Sing. 39, 1
(1986) 17; Blumea 41 (1996) 381; Tree Fl. Sabah & Sarawak 3 (2000) 384. — Myristica
polyspherula Hook. f., Fl. Brit. India 5 (1886) 108, p.p. (see notes by Sinclair, 1958: 425; 1975:
103); King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 312, pl. 146, emend.

Myristica globularia auct. non Blume: Hook. f. & Thomson, Fl. Ind. (1855) 160; A. DC., Prodr. 14,
1 (1856) 202, p.p., for the specimens from Malacca.

Horsfieldia lemanniana auct. non (A.DC.) Warb.: Warb., Mon. Myrist. (1897) 326, p.p. (type of
basionym Myristica lemanniana excluded).

Twigs 2—3 mm diameter; leaf buds covered with hairs (0.2—)0.3—0.6 mm long. Leaves
7-19 by 2.5-6 cm, lateral nerves 6—15 pairs. Male buds 1—1.5 mm diameter; thecae
(6 or) 8-14. Female perianth 2—2.5 mm long. Fruits (1.7—)1.9—2.5(—2.8) by 1.4—2 cm,
pericarp 2—4 mm thick.

Field-notes — Slender tree with narrow crown, the branches often almost in whorls;
bark fissured, rarely flaky; inner bark yellow or reddish, fibrous; wood whitish to ochre-
brown. Flowers at first jade-green, at anthesis (orange-)yellow with faint sweet odour
when crushed. Fruits greenish yellow to orange.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo (Sarawak,
rare; Sabah, E Kalimantan), Philippines (Luzon, Mindanao, rare; Palawan).

Habitat & Ecology — Lowland forest, usually on sandy soils; also Casuarina forest
(E Borneo), fresh-water swamp forest, ridge-top forest, kerangas (Sarawak, rare); alti-
tude 0-900 m; fl. & fr. throughout the year, but most fl. collections June—Sept..

Notes — 1. The leaves, as in var. sumatrana, are usually brittle when dry and often
fragmented in herbaria; they are usually greenish above and contrast well with the light
to chocolate-brown colour of the lower surface. The hairs of the leaf buds and young
twig apex are rather long, 0.2—0.6 mm, slightly longer than in the other varieties.

2. Because var. polyspherula is mainly characterized by its smaller fruits, it is often
difficult to tell whether specimens with fruits just over 25 mm long belong to var. suma-
trana or to var. polyspherula.

3. Specimens from kerangas in Sarawak may deviate by a stouter habit and less con-
trasting colour of the dry leaves.

b. var. maxima W.J. de Wilde

Horsfieldia polyspherula (Hook. f. emend. King) J. Sinclair var. maxima W.J. de Wilde, Gard. Bull.
Sing. 39, 1 (1986) 22; Tree Fl. Sabah & Sarawak 3 (2000) 383. — Type: S 36228 (Chai), Sarawak,
7th Div.

Twigs 2—4 mm diameter; leaf buds covered with hairs 0.2—0.5 mm long. Leaves 9-20
by 3.5—7 cm, lateral nerves (6—)9—16 pairs. Flowers not seen. Fruits (3.5—)4—6 by 3-5
cm, pericarp (5—)8—15 mm thick.

Field-notes — Buttresses present or absent; bark with shallow boat-shaped fissures;
inner bark pinkish; wood medium soft, whitish yellow. Fruits yellow to red.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, E Kalimantan).

Habitat & Ecology — Mixed forest, Agathis forest, kerangas; on sandy water-logged
soil, sandy loam or yellow clay-loam; 50—500(—1000) m altitude; fr. throughout the year.

170 Flora Malesiana, Ser. I, Vol. 14 (2000)

Note — This variety differs mainly in its conspicuously large and almost globose
fruits with a very thick pericarp. It may be confused with H. majuscula from Peninsular
Malaysia, which differs from H. polyspherula in the size of the male flowers (flowers of
var. maxima are unknown), a different androecium, leaf colour, and somewhat in the
fruits with a thinner pericarp.

Fruiting specimens of H. polyspherula var. maxima may also be confused with H.
punctatifolia, which differs in its dotted leaves.

c. var. sumatrana (Miq.) W.J. de Wilde

Horsfieldia polyspherula (Hook.f. emend. King) J. Sinclair var. sumatrana (Miq.) W.J. de Wilde,
Gard. Bull. Sing. 39, 1 (1986) 20; Tree Fl. Sabah & Sarawak 3 (2000) 384. — Myristica glabra
Blume var. sumatrana Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 49. — Horsfieldia brachiata
(King) Warb. var. sumatrana (Miq.) [J. Sinclair ex Whitmore, Tree Fl. Malaya 1 (1972) 325,
nom. inval., basionym wrongly cited and without literature ref.] J. Sinclair, Gard. Bull. Sing. 28
(1975) 13, p.p. (excl. syn. Horsfieldia majuscula and bartlettii). — Type: Korthals s.n., W Sumatra.

Myristica integra Wall., Cat. (1832) n. 6799, nom. nud.

Myristica collettiana King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 312, pl. 147. — Syntypes: King
3620, 3899 (lecto), 6672, 6737, Peninsular Malaysia.

Horsfieldia subglobosa auct. non (Migq.) Warb.: Warb., Mon. Myrist. (1897) 328; Gamble, Mat. Fl.
Malay Penins. 5, 23 (1912) 220; Ridl., Fl. Malay Penins. 3 (1924) 60; J. Sinclair, Gard. Bull.
Sing. 16 (1958) 425, 426, f. 48-50, 51A—D, p.p. (excl. syn. Horsfieldia majuscula).

Twigs 2-4 mm diameter; leaf buds covered with hairs 0.1—0.4 mm long. Leaves 13—
28 by 4.5—-9 cm, lateral nerves 11—-15(—20) pairs. Male buds 1.2—1.8 mm diameter,
thecae 12 or 14. Female buds 3 mm long. Fruits (2.5—)2.8—-3.5 by 2.2-2.7 cm, pericarp
3—5 mm thick.

Field-notes — Tree usually slender, bole straight, without buttresses, bark dark brown,
smooth, shallowly to fairly fissured, sometimes flaky (strips 10-20 mm wide); inner
bark reddish, fibrous, laminated, kino profuse, colourless then deep red; slash wood
(sapwood) and cambium pale; heartwood pinkish. Flowers (waxy) yellow. Fruits glossy
green turning (greenish) yellow, or orange.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo.

Habitat & Ecology — Lowland mixed dipterocarp forest, ridge and montane forest,
(peat) swamp forest, and kerangas; on sandy(-loamy) soils, red or yellow clayey soil;
0—1100 m altitude; fl. & fr. throughout the year.

Notes — 1. The hairs of the leaf buds in specimens from Sumatra (including the type
of var. sumatrana) are quite short, 0.1 mm long; in specimens from Peninsular Malaysia
and Borneo they are usually longer and rough, 0.4 mm long.

2. Specimens with fruits less than 30 mm long, smallish leaves, and rather slender
twigs may be difficult to distinguish from var. polyspherula.

3. Specimens from East Kalimantan may deviate in habit by thinner, membranous
leaves, which dry to a paler brownish colour; the flowers and fruits are not different,
although Kostermans (/26/3) noted that the fruit is wine-red, a colour as yet unrecorded
for other fruiting collections.

4. King’s collector (Goping) 6004 (syntype of H. majuscula, not the lectotype) and
Nur 34117 (both from Peninsular Malaysia, with male flowers) belong to the H. poly-
spherula-complex on account of the subglobose male buds and details of the androecium.

De Wilde — Myristicaceae (Horsfieldia) 17]

They agree with var. sumatrana, but differ by the larger buds, 1.8—-2 mm diameter, with
larger androecium, | by 0.8 mm; there are 10 or 12 thecae.

62. Horsfieldia psilantha W. J. de Wilde

Horsfieldia psilantha W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 112. — Type: NGF 43642
(Womersley), male fl., Papua New Guinea, Long Island.

Tree 5—25 m. Twigs 3—6(—15, in fruiting twigs) mm diameter, glabrescent, hairs red-
dish or grey-brown, 0.1—0.3 mm; bark finely striate, not flaking, lenticellate. Leaves
membranous, (oblong-)lanceolate, 20—40 by 4.5—-12.5 cm, base attenuate, apex acute-
acuminate; upper surface drying olivaceous, usually minutely pale-punctate; lower sur-
face (late) glabrescent, hairs scattered, 0.1—0.3 mm, on and near the midrib; dots absent;
midrib + flat above; nerves 14—24 pairs, thin, + flat above, lines of interarching + irreg-
ular but distinct; venation lax, indistinct; petiole 5—20 by 2—3.5 mm; leaf bud 20—25
by 2.5—3.5 mm, with hairs 0.1-0.3 mm. /nflorescences with sparse woolly stellate-den-
droid hairs 0.2—0.4 mm; in male and female (from infructescences): 3 or 4 times branched,
many-flowered, 10-16 by 8-12 cm, peduncle 1—4 cm; bracts not seen, caducous; flow-
ers in loose clusters of 2—5, glabrous, perianth 2-lobed, pedicel slender, not articulated.
Male flowers: pedicel 2—4.5 mm long; buds somewhat laterally compressed, subcircular,
(2-)2.5-3 by (2.5—)3—3.5(—4) mm, apex broadly rounded, base rounded to short-attenu-
ate, cleft 1/2—2/3: lobes 0.2—0.3 mm thick; androecium laterally much compressed, apex
rounded-truncate, 1.4—1.8 by 1.5—2.2 mm (Plate1: 28); thecae 24-28, mutually appressed,
erect, free apices 0.1 mm; androphore up to 0.2 mm; column narrowly hollowed for
1/4-1/2. Female flowers not seen. Fruits up to 10 per infructescence, ellipsoid, 1.7—2.2
by 1.2-1.7 cm, with rusty hairs 0.2 mm long, sometimes only remaining at the very base;
pericarp 1-2 mm thick, without or with scattered small lenticels or wartlets; fruiting
pedicel 2-8 mm long; perianth not persistent.

Field-notes — Slender tree, branches often drooping, without or with a few buttress-
roots; bark blackish or dark grey-brown, longitudinally fissured; inner bark cream or
pink, exudate pink or colourless; sapwood straw- or cream-coloured. Flowers orange-
yellow. Fruits yellow to orange.

Distribution — Malesia: NE Papua New Guinea (Madang Prov.: Bagabag I., Long I.;
New Britain; New Ireland).

Habitat & Ecology — Forest, including beach forest, shaded degraded forest; altitude
0-200 m; fl. May & Oct.; fr. throughout the year.

Note — Horsfieldia psilantha is related to and resembles H. laevigata, H. tuberculata,
and H. whitmorei. Horsfieldia laevigata, a variable and widespread species as well, al-
ways has hairy perianths, though sometimes only scattered hairs are present; it usually
has smaller leaves, and the fruits usually have many more and coarser lenticel-like
tubercles. Horsfieldia tuberculata, variable and widespread, has similarly glabrous flow-
ers, but the shape of the perianth is more tapered at the base, while in the present species
it is more circular in lateral view, with the base not or but slightly tapered; H. tuberculata
furthermore has glabrous fruits and ovaries. The leaves of H. whitmorei, a species from
the Solomon Islands, sometimes have similar, rather regularly looping, submarginal veins,
and similar fruits, but the male perianth is smaller, only 2 mm diameter or less, cleft to
c. 9/10, with the base and pedicel hairy.

172, Flora Malesiana, Ser. I, Vol. 14 (2000)

63. Horsfieldia pulcherrima W. J. de Wilde

Horsfieldia pulcherrima W.J. de Wilde, Gard. Bull. Sing. 38, 2 (‘1985’, 1986) 206. — Type: FRI
8008 (Cockburn), Peninsular Malaysia.

Tree 7-27 m. Twigs 5-8 mm diameter, with dense felty to woolly rusty or reddish
brown hairs 1—1.5 mm, late glabrescent; bark coarsely striate, + cracking and flaking;
lenticels only in the older wood, large, not contrasting in colour. Leaves chartaceous,
(elliptic-)oblong, (18—)24—36 by 8-14 cm, base rounded to attenuate, apex acute-acu-
minate; upper surface drying dull olivaceous to blackish brown, early glabrescent, lower
surface with dense rusty or red-brown hairs 1-1.5 mm long; dots present; midrib slen-
der, flat above, late glabrescent; nerves 18—23 pairs, slender, flat or sunken, lines of in-
terarching not very distinct; venation lax, faint; petiole 15-25 by 5—7 mm, pubescent;
leaf bud 25-30 by 7-9 mm, with hairs 1-1.5 mm long. /nflorescences behind the leaves,
with dense + shaggy hairs 0.5—1.5 mm long; in male: 5—8 by 5 cm, many-flowered, 3 or
4 times branched, peduncle 0.2—1 cm long; in female: a short, irregularly shaped, woody
knob (as in some Knemas), 1 cm long; flowers (male) solitary or in loose clusters of
up to 15, glabrous, perianth 3- (or 4-)lobed, pedicel not articulated; bracts ovate-oblong,
acute, densely pubescent outside, 3-7 mm long, caducous. Male flowers: pedicel 1—1.5
mm long, slender; buds somewhat depressed-globose, 1 by ].2-1.3(-1.4) mm, cleft c.
1/3 to nearly 1/2, lobes 0.2 mm thick; androecium depressed-globose, at apex with a
3- (or 4-)radiate crack, 0.5—0.6 by 0.8—1 mm (Plate 2: 51); thecae 24 or 26, closely pres-
sed, completely sessile, the apical parts overarching the apical hollow; column broadly
saucer-shaped, with a broad hollow to + halfway; androphore narrow, 0.1-0.2(-0.3) mm
long, largely hidden by the anthers. Female flowers (from remnants under fruits): 3 mm
long, with sparse hairs 0.3 mm long, cleft nearly 1/2, lobes 0.3 mm thick; ovary pubes-
cent. Fruits 1-3(—15) per infructescence, broadly ellipsoid to globose, 1.6-1.8 by 1.5-
1.7 cm, with dense shaggy rusty hairs 2 mm long; pericarp + woody, 1.5 mm thick; seeds
broadly ellipsoid, 10 mm long; fruiting pedicel to | mm long; perianth not persistent.

Field-notes — Slender tree, bole straight; bark grey-brown to blackish, shallowly
fissured, occasionally flaking. Inner bark red, laminated, with some red exudate. Slash
wood pale (yellow), wood brown. Fruits yellowish brown hairy.

Distribution — Malesia: Sumatra (Jambi), Peninsular Malaysia (Pahang, Johore).

Habitat & Ecology — Lowland primary forest, swamp forest; 50—600 m altitude;
apparently a rare species; fl. Mar.; fr. June, Sept.

Notes — 1. When sterile, H. pulcherrima may be confused with H. flocculosa, H. su-
perba, or H. wallichii, but the male flowers are quite different, similar to those of
H. grandis. The species is peculiar because of its rather small, subglobose, densely to-
mentose fruits; in related species the fruits (and ovary) are glabrous.

2. Horsfieldia pulcherrima may superficially resemble and may be confused with the
stout-leaved Gymnacranthera bancana, but the latter has a different indumentum, and
lacks the blackish dots on the lower leaf surface.

64. Horsfieldia pulverulenta Warb.

Horsfieldia pulverulenta Warb., Mon. Myrist. (1897) 342, t. 23; Markgr., Bot. Jahrb. Syst. 67 (1935)
150 (sub H. ralunensis); W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 135. — Myristica
pulverulenta (Warb.) Boerl., Handl. 3 (1900) 87. — Horsfieldia hellwigii (Warb.) Warb. var.

De Wilde — Myristicaceae (Horsfieldia) 173

pulverulenta (Warb.) J. Sinclair, Gard. Bull. Sing. 28 (1975) 56, p.p., excl. BW 12194 (Vink) =
Horsfieldia leptantha. — Syntypes: Beccari 759, 925, Papua Barat, Bird’s Head, Andai, Mt
Arfak.

Horsfieldia hellwigii (Warb.) Warb. var. hellwigii x var. pulverulenta (Warb.) J. Sinclair, Gard. Bull.
Sing. 28 (1975) 58. — Syntypes: Saunders 202 (L lecto), 358, 398, 483, Papua New Guinea.

Tree 15-25 m. Twigs 4-10 mm diameter, early to rather late glabrescent, hairs dark
rusty, 0.5—1.2 mm; bark finely striate or not, not flaking, lenticels usually present. Leaves
coriaceous, elliptic to oblong-lanceolate, 14-35(—40) by 4—10.5(—13) cm, base rounded
or short-attenuate, apex acute-acuminate, often to 2(—3, in sapling shoots) cm caudate;
upper surface drying dull olivaceous-brown, minutely rugose-pustulate, lower surface
late glabrescent or with persistent + evenly sized, spaced, rather harsh, dark brown hairs
1-1.5 mm long, when shed rough with thickened hair bases; dots absent; midrib + flat
above, late glabrescent, but indumentum usually persistent towards base; nerves 11—30
pairs, + straight, 50—70° to the midrib, 5-15 mm apart, thin and sunken above, lines of
interarching regular and prominent; venation lax, sunken and clearly visible above, giv-
ing the blade a + bullate appearance; petiole 2-12 by 2.54.5 mm, not or hardly winged;
leaf bud stout, 3—6 cm long, with harsh hairs 1—1.5 mm long. Inflorescences with woolly
hairs 1-1.5 mm long, 2 or 3 times branched, rather many-flowered; in male and female:
4-10 by 2-9 cm, peduncle up to 1.5 cm long; bracts (broadly) ovate, acutish, 3-5 mm
long, caducous; flowers + solitary (in female) or in loose clusters of 2—6, perianth
2-lobed, with stellate(-dendroid) hairs 0.1—0.3 mm, pedicel slender, with coarse hairs
0.4-0.7 mm, not articulated. Male flowers: pedicel 2—3.5 mm long; buds somewhat com-
pressed, in lateral view subcircular, 1.5—3 by 3-4 mm, the basal part thick and coriaceous,
the remainder collapsed when dry and perianth then often saucer-shaped or wrinkled,
above the anthers opening by a minute pore-like slit less than 1 mm wide; lobes and
apical part of perianth 0.2(—0.4) mm thick; androecium consisting of a coriaceous + el-
lipsoid column 0.8—1.1 mm long, with 2 small anthers, each 0.2(—0.3) mm at the apex
(Plate 2: 36). Female flowers: pedicel (1.5—)3—5 mm long; buds broadly ellipsoid-ovoid,
3.8—4 by 3.5—4 mm, cleft 1/5—1/10, with a minute pore-like slit above the stigmas; ovary
ovoid-subglobose, 2.5—3 by 2.5 mm, with dense hairs 0.1—0.3 mm; style erect, glabrous,
0.2-0.8 mm long; stigma 2-lobed, 0.2-0.3 mm long. Fruits 3-10 per infructescence,
ellipsoid, apex acute, sometimes acuminate, 3—5 by 2—3 cm, minutely pubescent at least
at the base, pericarp woody-coriaceous, 4—7 mm thick, usually with paler, small or coarse
lenticels or tubercles; fruiting pedicel 2-7 mm long; perianth not persistent.

Field-notes — Bole with slight buttresses or not buttressed. Bark often strongly peel-
ing in small, oblong, thin scales, black-brown. Wood whitish or straw, moderately hard
and heavy. Flowers (greenish) yellow or orange-yellow. Fruits (greenish) yellow or yel-
low-brown.

Distribution — Malesia: New Guinea (Papua Barat: Bird’s Head, Jayapura, Geelvink
Bay, Mimika; Papua New Guinea: West & East Sepik, Madang, Western Prov.; also a
deviating specimen from Gulf Prov.).

Habitat & Ecology — Lowland primary and (old) degraded rain forest, ridge-side or
swamp forest; on clay, stony-sandy soil; 0-500 m altitude; fl. & fr. throughout the year.

Notes — 1. Horsfieldia pulverulenta belongs to the group with H. hellwigii, which
has stout twigs, and conspicuous coarse hairs on the leaf buds and apex of twigs, (0.5—)
1-1.5 mm long. It is distinguished by its woody perianths, in male usually collapsed

174 Flora Malesiana, Ser. I, Vol. 14 (2000)

around the small androecium. The androecium is deviating in shape within Horsfieldia
and consists of an ellipsoid woody body, at the apex with only 2 apparently much re-
duced anthers (or thecae?) just below the apical pore-like slit of the perianth.

The female flowers of H. pulverulenta are larger than those of H. hellwigit.

2. Schodde & Craven 4662 (Gulf Province, male flowers) deviates in non-coriaceous,
woolly (not harsh) hairy leaves, and densely hairy perianths. The perianth is light brown
on drying, not blackish brown as is usual in #7. pulverulenta, and collapses only slightly;
it has an androecium comparable to that of H. pulverulenta. The marginal nerve of the
leaves loops very regulary, as in H. pulverulenta. The specimen may represent a separate
taxon.

65. Horsfieldia punctata W. J. de Wilde

Horsfieldia punctata W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 37. — Type: FRI 9014 (Bur-
gess), Peninsular Malaysia.

Tree 10-25 m. Twigs 2.5—5 mm diameter, early or late glabrescent, hairs rusty, 0.1—
0.4 mm long; bark coarsely striate, with a tendency to crack longitudinally and to flake;
lenticels + absent. Leaves coriaceous, (elliptic-)oblong, 4.5—12 by 2-5 cm, base (short-)
attenuate, apex rounded to subacute; upper surface drying olivaceous-brown, lower sur-
face rufous, glabrous, but midrib remaining pubescent for some time; dense dots present;
midrib somewhat raised above; nerves 5—12 pairs, thin and almost flat above, late
glabrescent, lines of interarching indistinct; venation invisible on both surfaces; petiole
6-12 by 1.5—2.5 mm, late glabrescent; leaf bud 7-13 by 2-3.5 mm, with dense grey-
brown to rusty hairs 0.1—0.4 mm. Inflorescences with dense rusty hairs 0.2—0.3 mm; in
male: usually twice branched, not many-flowered, 2-3 by 0.5—1 cm, peduncle | cm long;
in female: once branched, 2-3 cm long, glabrescent; bracts elliptic-oblong, 1-3 mm,
pubescent, caducous; flowers in male in clusters of 3—5, perianth 3-lobed, glabrous,
pedicel towards the base with hairs 0.1 mm or less, not articulated. Male flowers
(submature): pedicel | mm long, slender; buds globose, 0.8 by 0.8—1 mm, cleft (2/3)
3/4. lobes 0.2 mm thick; androecium (depressed-)globose, 0.4—0.5 by 0.5 mm, + circu-
lar in cross section (Plate 3: 82); thecae 10 or 12 (see note 1), sessile, towards the apex
incurved and concealing a shallow, narrow, apical cavity in the column, 0.2(—0.3) mm
deep; androphore narrow, 0.1 mm long. Female flowers not seen. Fruits 2-4 per in-
fructescence, ovoid, 2—2.3 by 1.7—1.9 cm, glabrous, finely granulate; pericarp 1.5 mm
thick; fruiting pedicel 2 mm long; perianth not persistent.

Field-notes — Bole straight, buttresses absent; bark deep- to mid-brown, grid-cracked
with rather chunky scales or finely fissured with firm ridges, or bark thick and corky,
finely longitudinally fissured; outer cut of slash bark brown, inner bark bright red, lay-
ered, separated by blade line; slash wood white to fawn, speckled red; exudate red, blood-
like. Fruits greenish yellow, slightly glaucous.

Distribution — Malesia: Peninsular Malaysia (Cameron Highlands; Fraser’s Hill,
Genting, Gunung Bunga Bua).

Habitat & Ecology — Lower montane forest on granite, ridge forest; at c. 1000 m
altitude; fl. Mar.; fr. Nov.

Notes — 1. The available flowers are clearly immature and hence the anthers are
difficult to count; possibly there are c. 11 thecae and in reality only 5 or 6 anthers.

De Wilde — Myristicaceae (Horsfieldia) 175

2. Horsfieldia punctata is obviously closely related to H. glabra from S Sumatra and
Java, and to H. montana, H. punctatifolia and H. subalpina. Horsfieldia montana (from
Borneo) is very similar in general habit as well as in male flowers and fruit shape, but
lacks the characteristic dots on the leaves; H. punctatifolia, a species with a wider distri-
bution in Sumatra, Peninsular Malaysia, and Borneo, differs in membranous leaves and
larger fruits with a very thick-leathery pericarp. Horsfieldia subalpina also seems re-
lated, but lacks the leaf dots, has larger and less coriaceous leaves, and larger fruits.
Horsfieldia punctatifolia has glabrous pedicels and in H. subalpina the pedicels are finely
puberulous in the lower part, similar as in the present species. Generally H. glabra has a
much shorter indumentum.

66. Horsfieldia punctatifolia J. Sinclair

Horsfieldia punctatifolia J. Sinclair, Gard. Bull. Sing. 16 (1958) 413, f. 44, pl. XI-B; 28 (1975) 105;
W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 54; Tree Fl. Sabah & Sarawak 3 (2000) 384. —
Type: SF 4221/1 (Sinclair) (= Sinclair 7987), fr., Singapore.

Tree 7-30 m. Twigs 2.5—4 mm diameter, early glabrescent, hairs grey-brown, up to
0.1 mm long; bark finely to coarsely striate, not flaking; lenticels conspicuous or incon-
spicuous. Leaves membranous, (elliptic-)oblong, 9-21 by 3—9 cm, base attenuate, apex
acute-acuminate; upper surface drying dull olivaceous or dark brown, lower surface
glabrous; scattered dots (0.1 mm diameter) present; midrib + flat above; nerves 11-16
pairs, thin, flat above, lines of interarching not distinct; venation lax, hardly visible on
both surfaces; petiole 10-17 by 1.5-3 mm, glabrous; leaf bud 8-12 by 1.5—2 mm, with
dense grey-brown hairs up to 0.1 mm long. Inflorescences with sparse hairs up to 0.1
mm, or glabrescent; in male: about 3 times branched, rather many-flowered, 4-10 by
2-8 cm, peduncle 0.5—2 cm long; in female: about twice branched, 3-6 by 1.5—4 cm,
fewer-flowered than in male; bracts + oblong, 2-4 mm, short-pubescent, caducous; flowers
in male in loose clusters of 2—5, glabrous, perianth 3- or 4-lobed, pedicel not articulated.
Male flowers: pedicel 1-4 mm, slender; buds depressed-globose, 1.4—2 by 1.6—2.2 mm,
cleft 3/4—4/5, lobes 0.2—0.3 mm thick; androecium depressed-globose to depressed broad-
ly ovoid, 0.6—0.7 by 1-1.5 mm, nearly circular in cross section (Plate 3: 90); thecae
14-18, completely sessile (free apices up to 0.1 mm), the apices of anthers concealing
the apical hollow, column broad with narrow hollow up to 0.2 mm deep, in Sumatra up
to 0.4 mm deep; androphore narrow, 0.1—0.2 mm long. Female flowers: pedicel 1-2 mm
long; buds ellipsoid, 2.8-3.5 by 2.5-3 mm, cleft c. 1/2; ovary broadly ovoid, 1.5—2 by
1.3-2 mm, glabrous, stigma irregularly 2-lipped, 0.2-0.3 mm high. Fruits 1—3 per
infructescence, (broadly) ellipsoid, glabrous, 4.5—8 by 3—4.5 cm, drying blackish brown,
without or with few small warts; pericarp 10-20 mm thick; fruiting pedicel 4-6 mm
long; perianth not persistent.

Field-notes — With or without low-rounded or steep thick buttresses; bark + smooth,
shallowly fissured or cracked, or brittle-scaly; inner bark pinkish to red, with reddish
watery exudate; cambium white; slash wood white, yellowish, or pinkish; heartwood
dark brown. Perianth (bright) yellow, with a turpentine odour. Fruits yellow to red, apri-
cot, orange(-brown) flushed pink; pericarp pink inside.

Distribution — Malesia: Sumatra (North, and E Coast Prov.), Peninsular Malaysia,
Singapore, Borneo (Sarawak, Sabah; W, C & E Kalimantan).

176 Flora Malesiana, Ser. I, Vol. 14 (2000)

Habitat & Ecology — Primary forest, including hillside, ridge-top, and pole marshy
forest, and kerangas; ona variety of soil types including grey and brown soil, sandy clay,
tertiary sandstone, dacite hill; 0-1100 m altitude; fl. & fr. throughout the year.

Note — Horsfieldia punctatifolia is characterized by the dots on the leaves, deeply
cleft male buds, few anthers, and large fruits with thick pericarp.

67. Horsfieldia ralunensis Warb.

Horsfieldia ralunensis Warb, Mon. Myrist. (1897) 336; K.Schum., Notizbl. Bot. Gart. Berlin 2
(1898) 117; K. Schum. & Lauterb., Fl. Schutzgeb. Siidsee (1900) 324; Markgr., Bot. Jahrb. Syst.
67 (1935) 150; W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 143. — Type: Warburg 20709 (B
lost, n.v.), New Britain (Neu Pommern), Gazelle Peninsula, Ralun.

Tree 5—18 m. Twigs 5—10 mm diameter, early or late glabrescent, hairs light rusty or
yellow-brown, 0.5—1 mm long; bark finely striate, not flaking; lenticels present but not
distinct. Leaves membranous, (oblong-)lanceolate, 30-60 by 7-11 cm, base nearly
rounded or (short-)attenuate, apex long acute-acuminate, usually 1-2 cm caudate; upper
surface drying dull olivaceous, minutely palely punctate-pustulate, lower surface with
persistent pale brown rather soft dendroid hairs of variable size 0.5—1 mm, when shed
not leaving thickened rough hair bases; dots absent; midrib flat above, glabrescent ex-
cept at the very base; nerves 30—40 pairs, generally + straight, 8—15(—20) mm apart, thin
and flat above, lines of interarching rather distinct, + irregularly looping; venation lax,
indistinct; petiole 5—16 by 3—4 mm, not or hardly winged; leaf bud 4—6 cm long, with
dense velvety hairs (0.5—)1 mm. Inflorescences with woolly hairs 1—1.5 mm long; in
male and female: 2 or 3 times branched, rather many-flowered, 4—20 by 3-10(-12) cm,
peduncle 0.5—5 cm long; bracts broadly ellipsoid, subacute, 5-10 mm long; flowers in
loose clusters of 3—6, perianth 2-lobed, glabrous except at the very base, pedicel with
hairs 0.3—0.8 mm long, not articulated. Male flowers: pedicel 1(—2) mm long; buds
obovoid-ellipsoid, at apex + acute, 2—2.3 by 1.5-1.7 mm, cleft c. 1/4, lobes 0.2 mm
thick. Androecium 1-1.1 by 1.2-1.3 mm, somewhat flattened, apex broadly rounded,
column narrowly hollowed for c. 1/4 (Plate 2: 39); thecae 20-24, almost completely
sessile, erect, mutually touching, 1-1.2 mm long, free apices c. 0.2 mm, androphore
slender, 0.3—0.4 mm long. Female flowers: pedicel 1—-1.5 mm; buds obovoid, 4 by
2.5—3 mm, cleft c. 1/4; ovary ovoid, 2 by 1.5 mm, with dense hairs 0.5 mm, style and
stigmas minutely 2-lobed, + elongate, 0.2 mm long. Fruits 2-10 per infructescence, sub-
ellipsoid, apex + obtuse, base broadly rounded, 2.5—3 by 1.5-1.9 cm, with hairs 0.5 mm
long, and with lenticel-like tubercles; pericarp 3-5 mm thick; fruiting pedicel 1-3 mm
long; perianth not persistent.

Field-notes — Small or medium-sized tapered understorey or subcanopy tree, bole
straight, sometimes slightly buttressed; branches horizontal but drooping apically; bark
dark, mottled, or with short vertical fissures; wood straw, moderately soft. Inflorescences
erect, flowers yellow. Mature fruits green or brown-green.

Distribution — Malesia: Papua New Guinea (W & E New Britain, Gazelle Penin-
sula; New Ireland).

Habitat & Ecology — Lowland rain forest; well-drained pumice terrain, sandy soil,
ridge forest, at edge of swamp; 0-100 m altitude; fl. & fr. apparently not seasonal.

De Wilde — Myristicaceae (Horsfieldia) Wa,

Note — Related to H. hellwigii, H. leptantha, and H. pulverulenta, the first also occur-
ring in New Britain; H. ralunensis can be distinguished from H. hellwigii by the larger
fruits, smaller and narrower male flowers (with a somewhat different androecium), and
generally more elongate leaves.

68. Horsfieldia reticulata Warb.

Horsfieldia reticulata Warb., Mon. Myrist. (1897) 304, t. 22; Merr., Enum. Born. (1921) 268;
J. Sinclair, Gard. Bull. Sing. 28 (1975) 107; W.J. de Wilde, Gard. Bull. Sing. 38, 2 (1985’,
1986) 218; Blumea 41 (1996) 381, f. 1h, i; Tree Fl. Sabah & Sarawak 3 (2000) 385. — Myristica
reticulata (Warb.) Boerl., Hand]. 3 (1900) 85. — Type: Beccari 3475, Sarawak.

Horsfieldia affinis W.J. de Wilde, Gard. Bull. Sing. 38, 2 ((1985’, 1986) 217. — Type: S 247185
(Wright), Sarawak.

Tree 7-20 m. Twigs 2.5—6 mm diameter, late glabrescent, hairs rusty, 0.5—1 mm long;
bark striate, when old cracked but not flaking; lenticels absent or small and inconspic-
uous. Leaves membranous or chartaceous, rarely bullate, elliptic-oblong to lanceolate,
8-35 by 4-11 cm, base rounded to attenuate, apex acute-acuminate; upper surface dry-
ing (blackish) brown, glabrous, lower surface with (sub)persistent rather sparse (stellate
or) dendroid hairs 0.5—1 mm long; dots absent; midrib + flat above, often late glabrescent;
nerves 9-20 pairs, slender, (sunken or) raised, lines of interarching regular, distinct;
venation lax, faint or distinct; petiole (7—)12—20 by 2—4 mm, pubescent; leaf bud 10—15
by 3—4 mm, with dense hairs 0.5—1 mm. Inflorescences with dense shaggy hairs 0.5—2
mm long; in male: many-flowered, 4 or 5 times branched, 12—25 by 8—14 cm, peduncle
1.5—6 cm long; in female: fewer flowered than in males, 5—6 cm long; bracts triangular
to oblong, acute, 3-9 mm long, densely pubescent, caducous; flowers in male in loose
clusters of 2—6, glabrous, perianth 3- or 4- (or 5-)lobed, pedicel articulated. Male flow-
ers: pedicel slender, 0.5—2 mm long; buds depressed-globose, |.2—2 by 1.5—2(—2.5) mm,
cleft 1/4—1/3, lobes 0.1—0.2 mm thick; androecium (depressed) globose, slightly impres-
sed at the centre, 0.6—0.8 by 1—1.5 mm (Plate 2: 59); thecae 20—24(—30), completely
sessile, incurved towards the apex; column broad, with hollow up to + halfway; androphore
slender or broad, 0.2—0.4 mm long, sometimes hidden by the anthers. Female flowers
(from remnants under the fruits): buds 2.5—3 by 2—2.5 mm, cleft over halfway, lobes 3,
probably with very weak indumentum. Fruits 4—7 per infructescence, ellipsoid, 2.3—2.7
by 1.7—2.1 cm, glabrous, with some paler wart-like tubercles; pericarp 1.5—2 mm thick;
fruiting pedicel 3—4 mm long; perianth persistent. — Fig. 24h, i.

Field-notes — Small tree, crown pyramidal, buttresses absent; bark rough, dark brown,
lenticellate; outer bark 1-2 mm thick, brown-red, inside red. Flowers dark or golden
yellow, fragrant or with strong sweet smell of balsam of Peru. Fruits ramiflorous, in
bunches behind the leaves, pinkish orange, + sour, eaten by the Dayaks.

Distribution — Malesia: Borneo (Sarawak; Brunei; W, C, E & SE Kalimantan).

Habitat & Ecology — Forest on alluvial soils, riverbanks; sandy clay soil or ridges,
sandstone; 0—600 m altitude; fl. May—Nov.; fr. July.

Notes — |. Horsfieldia reticulata may be confused with H. motleyi, which differs in
its inarticulated pedicels, much smaller pubescent flowers with a quite different
androecium, the leaves dull olivaceous brown on drying, and sunken lateral nerves.

2. In H. reticulata the male flower, including the androecium, is very similar to that of
H. flocculosa, H. grandis, and H. tomentosa, obviously all related species.

178 Flora Malesiana, Ser. I, Vol. 14 (2000)

69. Horsfieldia ridleyana (King) Warb.

Horsfieldia ridleyana (King) Warb., Mon. Myrist. (1897) 331; Gamble, Mat. Fl. Malay Penins. 5,
23 (1912) 221: Ridl., Fl. Malay Penins. 3 (1924) 60; Burkill, Dict. Econ. Prod. Malay Penins.
(1935) 1199: J. Sinclair, Gard. Bull. Sing. 16 (1958) 432, f. 52; 28 (1975) 108; W.J. de Wilde,
Gard. Bull. Sing. 39, 1 (1986) 7; Tree Fl. Sabah & Sarawak 3 (2000) 386. — Myristica ridleyana
King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 311, pl. 145. — Syntypes: Cantley 1798, King’s coll.
10917: Scortechini 862 (male fl., lecto), Peninsular Malaysia.

Horsfieldia kerangasicola J. Sinclair, in sched. (Borneo material).

Tree 5-25 m. Twigs sometimes + angular, 1.5—3.5(—5) mm diameter, early glabrescent,
with rusty hairs 0.2—0.4 mm long; bark finely to coarsely striate, not flaking; lenticels
either small and distinct, or absent. Leaves thinly chartaceous to coriaceous, elliptic-
oblong to lanceolate, 5—15(-16) by 2—4.5 cm, base attenuate, apex (acute-)acuminate;
upper surface glabrous, drying olivaceous-brown, lower surface glabrous, light brown
or reddish brown, sometimes contrasting with upper surface; dots absent; midrib flat,
sunken or slightly raised above, inconspicuous; nerves 7-15 pairs, sunken or + flat above,
inconspicuous; venation lax, faint; petiole 7-15 by 1.5-2.5 mm, early glabrescent; leaf
bud 6-14 by 1-2 mm, with dense hairs 0.2—0.5 mm long. Inflorescences with sparse
hairs 0.1—0.3 mm, sometimes glabrescent; in male: 3 (or 4) times branched, rather many-
flowered, 2-6 by 2-4 cm, peduncle (0.2—)0.5—1.5 cm long, the flowers in loose clusters
of 3—6: in female: fewer-flowered, 1.5—4 cm long; bracts oblong-lanceolate, 2-4 mm
long, caducous; flowers glabrous, perianth 3- or 4- (rarely 2-)lobed, pedicel slender, ar-
ticulated. Male flowers: pedicel 1-2 mm long; buds subglobose or short-ellipsoid, 1—1.2
by 0.8—1.3 mm, base rounded or short-attenuate, cleft c. 1/3 to nearly 1/2, not or slightly
collapsing on drying, lobes 0.2—0.4 mm thick; androecium (including androphore) broad-
ly obovoid, 0.6—-0.8 by 0.4-0.7 mm, in cross section 3- or 4-angular (or + ellipsoid in
2-lobed flowers) (Plate 3: 66); thecae 8-12, mutually almost entirely free (Peninsular
Malaysia, Borneo), or free only for nearly the upper half (part of the material from
Peninsular Malaysia), suberect, + acute, 0.3—0.4 mm long; androphore + tapering, nearly
as long as the anthers, 0.2—0.4 mm long. Female flowers: pedicel 1-2 mm long; buds
ellipsoid, 1.5—-1.8 by 1.5 mm, cleft 1/3 to nearly 1/2; ovary ellipsoid, 1.2—-1.3 by 0.6—0.7
mm, + grooved at one side, glabrous, stigma shallowly 2-lipped, 0.1-0.2 by 0.6 mm.
Fruits 1-6 per infructescence, ellipsoid, 1.5—2 by 1-1.4 cm, glabrous, without len-
ticel-like tubercles; pericarp 1.5 mm thick; fruiting pedicel 2—5 mm long; perianth not
persistent.

Field-notes — Slender or crooked trees, once with many buttresses; bark dark brown
to red brown, shallowly (rectangular) fissured, cracked, flaky, or dippled; slash bark
reddish, fibrous, laminated; sapwood pale, creamy pink. Flowers yellow; stamens pink;
ovary pale green. Fruits glossy, green turning yellow-green.

Distribution — Malesia: Peninsular Malaysia (Perak, Kelantan, Trengganu, Pahang,
Selangor, Malacca), Borneo (Sarawak, Brunei, Sabah); apparently not in Sumatra (see
note 3).

Habitat & Ecology — Forest on poor soils: heath forest (with Dacrydium beccarit),
Gymnostoma-forest, kerangas, ridge-forest, quartzite conglomerate-ridges; sandstone with
very shallow soil, with Gymnostoma, Tristania, Cotylelobium; sandstone ridges with
Dipteris, sandstone summits with Dacrydium; 0—1100 m altitude; fl. throughout the year,
in Borneo most collections June—Oct.; fr. throughout the year.

De Wilde — Myristicaceae (Horsfieldia) 9

Notes — 1. Some collections from Pahang have proportionally many 2-lobed flowers
in the inflorescences. Such flowers slightly differ in a shorter androecium which is not
conspicuously triquetrous but rather subellipsoid in cross section, with shorter androphore.

2. Specimens from Peninsular Malaysia generally have less coriaceous leaves and a
thicker indumentum on the leaf buds (0.4 mm) as compared with most of the specimens
from Sarawak, Brunei, and Sabah. Specimens from Borneo generally occur in heath
forest or kerangas, those from Peninsular Malaysia grow on ridges and hill slopes.

3. The only (sterile) specimen (bb 6479) from Sumatra may represent H. ridleyana,
but it also may be H. triandra.

4. Vegetatively H. ridleyana may be confused with H. penangiana. In Borneo H.
ridleyana may resemble H. oligocarpa, but the latter has distinctly raised nerves on the
upper leaf surface.

70. Horsfieldia rufo-lanata Airy Shaw

Horsfieldia rufo-lanata Airy Shaw, Kew Bull. 1939, n. 10 (1940) 440; J. Sinclair, Gard. Bull. Sing.
28 (1975) 111; W.J. de Wilde, Gard. Bull. Sing. 38, 2 ((1985’, 1986) 216; Tree Fl. Sabah &
Sarawak 3 (2000) 387. — Type: Richards 1667, Sarawak.

Tree 8-17 m. Twigs 3.5—5 mm diameter, late glabrescent, hairs dark rusty to reddish
brown, 1—1.5 mm long; bark coarsely striate, later on + longitudinally cracking, not
flaking; lenticels indistinct. Leaves chartaceous, sometimes slightly bullate, elliptic to
oblong, 10—23 by 5—10(—12) cm, base nearly rounded to subattenuate, apex (acute-)acu-
minate; upper surface drying olivaceous-brown, glabrous except for the nerves, lower
surface with dense to sparse, rather harsh hairs of variable size (ranging from sessile-
stellate, 0.3 mm, to short-armed dendroid emerging hairs up to 1.5 mm long); dots ab-
sent; midrib + raised above, late glabrescent; nerves 11—16 pairs, raised, pubescent or
late glabrescent, lines of interarching distinct, regular; venation lax, sunken, distinct or
not; petiole 10-16 by 3—4.5 mm, pubescent; leaf bud 15—20 by 3-5 mm with hairs 1—1.5
mm. Inflorescences with dense woolly-shaggy hairs 2-3 mm long; in male: many-flow-
ered, 3 or 4 times branched, 8—12 by 5—7 cm, peduncle 1.5—3 cm; in female: few-flow-
ered, 3 cm long; bracts densely shaggy pubescent, oblong-triangular to lanceolate, 3-8
mm long; flowers in male rather few-flowered in loose clusters, glabrous, perianth 3- or
4-lobed, pedicel indistinctly articulated. Male flowers: pedicel rather stout, 0.5—1 by 0.5
mm; buds subglobose, 2 by 2.3 mm, rather firm, cleft nearly 1/2, lobes 0.2(-0.3) mm
thick; androecium globose, 1.2 mm diameter (Plate 2: 58); thecae c. 30 (closely appressed),
completely sessile, incurved towards apex; column broad, with small apical hollow to
c. 1/5; androphore slender, 0.2 mm long. Female flowers not known. Fruits 2—4 per
infructescence, broadly ovoid-ellipsoid, 2—2.2 by 1.5—1.6 cm, glabrescent, with rem-
nants of minute hairs 0.1 mm at the base, without distinct lenticels or tubercles, pericarp
2 mm thick; fruiting pedicel 1-3 mm long; perianth not persistent.

Distribution — Malesia: Borneo (Sarawak: 4th & Sth Div.; Sabah).

Habitat & Ecology — Montane forest, upper dipterocarp forest on sandstone; altitude
900-1400 m; fl. June, Sept.; fr. Sept.

Note — A montane species closely related to H. splendida and especially to H. retic-
ulata. Horsfieldia rufo-lanata differs in the longer woolly hairs of the inflorescences,

180 Flora Malesiana, Ser. I, Vol. 14 (2000)

the almost globose (not depressed-globose) male perianth, the near-globose androecium,
more thecae, 26—30(—40) as against 20-24 and 16-20 in H. reticulata and H. splendi-
da, respectively. The nerves on the upper leaf surface of H. rufo-lanata apparently re-
main pubescent for a much longer time, and are generally raised; in contrast with H.
splendida, the blades are usually smaller and more elliptic and have stiffer hairs on the
lower surface: H. reticulata has a much less conspicuous indumentum on the lower leaf
surface.

71. Horsfieldia sabulosa J. Sinclair

Horsfieldia sabulosa J. Sinclair, Gard. Bull. Sing. 27 (1974) 133, W.J. de Wilde, Gard. Bull. Sing.
38 (‘1985’, 1986) 185, f. 22; Tree Fl. Sabah & Sarawak 3 (2000) 387. — Type: Sinclair & Kadim
10491, Brunei.

Tree 10—37 m. Twigs stout, with wart-like thickenings marking the leaf scars, (3—)5-—
10(—20) mm diameter, rather late glabrescent, hairs grey-brown to rusty 0.5—1(—1.5) mm
long; bark grey-brown, lower down grey-blackish, rough, longitudinally cracking and
often + flaking; lenticels inconspicuous. Leaves in 3—5 rows, generally clustered to-
wards the apex of the twigs, coriaceous, elliptic-oblong to (sub)lanceolate, 9-21 by 2-6
cm, base attenuate, apex acute(-acuminate); upper surface of mature leaves glabrous,
+ glossy or not, olivaceous-brown, lower surface pale brown, early glabrescent; + round
to line-shaped dark brown or blackish dots present; midrib flat, glabrous above; nerves
12-20 pairs, flat above and below, lines of interarching invisible; petiole long in propor-
tion to the blades, 25—50 by 2—4 mm, late glabrescent, hairs 0.5—1.5 mm; leaf bud stout,
10-15 by (3—-)4-5 mm, with dense grey-brown to rusty hairs 0.5—1.5 mm long. /nflores-
cences behind the leaves, with dense hairs 1—-1.5 mm long; in male: 2 or 3 times branched,
many-flowered, 10-13 by 5—6 cm, the peduncle 1.5—2.5 cm long; in female: + fewer-
flowered, 3—5 cm long; bracts elliptic to lanceolate, + boat-shaped, 4-15 mm long, with
dense hairs 0.5—-1.5 mm, glabrous inside, late caducous; flowers in loose clusters of
5-10, glabrous; perianth 3- (or 4-)lobed, pedicel articulated. Male flowers: pedicel 1.5
(2) mm long; buds (depressed-)globose, 2 by 2-2.3 mm, cleft (1/2-)1/3, not collapsing
on drying, the lobes 0.2 mm thick; androecium + globose, 1-1.2 by 1.7-1.8 mm
(Plate 2: 40); thecae 24(—30), + completely sessile, column broad with shallow apical
cavity; androphore narrow, 0.2 mm long. Female flowers ( Sinclair, l.c.: 134): immature
ovary glabrous. Fruits 1-6 per infructescence, ovoid, 3-5 by 2.5—4 cm, glabrous, drying
dark brown or somewhat bluish black, not warted; pericarp hard, 5—8(—10) mm thick;
fruiting pedicel 2-8 mm long; perianth not persistent. — Fig. 25.

Field-notes — Tall tree, buttresses absent; bark dark grey, longitudinally fissured; sap
red, copious; inner bark reddish brown, laminated, fibrous; sapwood soft, pinkish, with
4 hollow centre 2.5 cm wide. Leaves glossy above, very glaucous beneath, midrib green-
ish yellow. Fruits yellow, inside apricot; ramiflorous with many fruits on each branch.

Distribution — Malesia: Borneo (Sarawak: Serian and Bintulu District, Mulu Na-
tional Park; Brunei; Sabah: Sipitang).

Habitat & Ecology — Mixed forest on sandy or peaty soil, deep yellow sands, sandy
loam, or heavy yellow clay soil; in Agathis forest, ridge forest; 0-100 m altitude; fl.
May-July; fr. May—Dec.

De Wilde — Myristicaceae (Horsfieldia) 181

Notes — 1. A very characteristic species on account of its thick twigs with bunched
leaves, the latter with long petioles in (3—)5 rows, ramiflorous. Horsfieldia wallichii has
similar fruits, and almost similar dark brown dots on the lower leaf surface.

2. According to Koster & Baas (Blumea 27, 1981, 152) the leaves are unique in hay-
ing an iso-bilateral mesophyll.

Fig. 25. Horsfieldia sabulosa J. Sinclair. a. Shoot apex with densely bunched leaves with dispersed
phyllotaxis; indumentum of young leaves partly fallen; b. older wood with leaf scars in dispers-
ed phyllotaxis and immature male inflorescences; note bracts; c. smaller bract; d. immature male
flower, lateral view; e. ditto, longitudinally opened, showing androecium; f. longitudinal section of
androecium, schematic; g. old wood with infructescence, fruits mature, seeds completely covered
by aril [a—f: SAN 15146; g: BRUN 0828]. — Scale bar for a, b, g = 2 cm; for c—f = 0.85 mm.

182 Flora Malesiana, Ser. I, Vol. 14 (2000)

72. Horsfieldia samarensis W.J. de Wilde

Horsfieldia samarensis W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 76. — Type: PNH 147374
(Gutierrez), Philippines, Samar I.

Tree 5 m. Twigs 1.5—3.5 mm diameter, early glabrescent, hairs greyish, less than 0.1
mm: bark finely striate, not flaking; lenticels rather inconspicuous. Leaves membranous,
(oblong-)lanceolate, 7-11 by 2—3 cm, base attenuate, apex acute-acuminate, upper sur-
face drying dull olivaceous, lower surface bright brown, early glabrescent, hairs sparse,
less than 0.1 mm: dots absent; midrib slightly raised above; nerves 10-13 pairs, above
slender, flat: lines of interarching faint though regularly looping; venation inconspicu-
ous on both surfaces; petiole 8-10 by 1-1.5 mm, leaf bud 10 by | mm, with hairs less
than 0.1 mm. Inflorescences glabrescent or thinly set with stellate hairs 0.1 mm or less,
slender: in male: 2 or 3 times branched, 3-4 by 1.5—2.5 cm, rather few-flowered, pedun-
cle 1-1.5 cm; flowers solitary or in loose clusters of 2 or 3, glabrous, perianth 2-lobed,
pedicel not articulated, bracts not seen, caducous. Male flowers: pedicel slender, 1.5—3
mm long; buds slightly laterally compressed, transversely ellipsoid, 2-2.2 by 2.5—2.7
mm, above and below broadly rounded, drying brown, not collapsing on drying, cleft
2/3—4/5, lobes firm, 0.2(—0.3) mm thick; androecium + flattened, broadly obovoid to
transversely short-ellipsoid, 1.2-1.3 by 1.4—1.5 mm, (.8—0.9 mm thick (Plate 1: /0);
thecae 28 or 30, closely set, basally connate and forming a saucer-shaped cup into which
the anthers at one side inflect deeply nearly to the base, clasped and covered by the other
anthers, the inflexed parts of the anthers mutually free; androphore narrow, (0—)0.1 mm
long. Female flowers and fruits not seen.

Field-notes — Tree c. 5 m tall, dbh c. 6 cm. Flowers green.

Distribution — Malesia: Philippines (Samar I.; known only from the type).

Habitat & Ecology — North slope, 200-270 m altitude; fl. May.

Note — According to the male flower structure related to H. talaudensis, but differ-
ing in several points. Horsfieldia talaudensis is more robust, has stouter and more densely
hairy male inflorescences, shorter (than the perianth) and densely hairy pedicels, some-
what larger buds and the androecium with anthers inflexed into the cavity at both sides.

73. Horsfieldia schlechteri Warb.

Horsfieldia schlechteri Warb. in K. Schum. & Lauterb., Nachtr. Fl. Schutzgeb. Siidsee (1905) 267;
Markgr., Bot. Jahrb. Syst. 67 (1935) 153; W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 106,
f. 15a-f. — Horsfieldia subtilis (Mig.) Warb. var. schlechteri (Warb.) J. Sinclair, Gard. Bull.
Sing. 28 (1975) 137. — Type: Schlechter 14500, Papua New Guinea, Torricelli Mts.

Tree or shrub, 3-15 m. Twigs 1-4 mm diameter, hairs rusty-grey, 0.1 mm long, early
glabrescent; bark finely striate, not flaking; without or with few lenticels. Leaves mem-
branous or thinly chartaceous, elliptic-oblong to oblong-lanceolate, 6-19 by 1—6.5 cm,
base attenuate, apex acute-acuminate; upper surface drying (dark) olivaceous-brown,
finely pale pustulate or not, lower surface early glabrescent, hairs 0.1 mm or less; dots
absent: midrib slender, raised above; nerves 6—14 pairs, raised or + flat above, indistinct;
venation lax, hardly distinct on both surfaces; petiole 5—12 by 0.7—1.5 mm; leaf bud 8 by
| mm, with hairs 0.1 mm long. Inflorescences glabrescent or with sparse to dense stellate
hairs 0.1 mm long or less; in male: (1 or) 2 (or 3) times branched, 1.5—7 by 1—4 cm,

De Wilde — Myristicaceae (Horsfieldia) 183

peduncle 0.3—1.5 cm, slender; in female: few-flowered, |—4 cm long; bracts 0.5—1 mm
long, densely woolly pubescent, caducous; flowers solitary or 2 or 3 together, perianth
2-lobed, (almost) glabrous; pedicel slender, sometimes with minute hairs, not articu-
lated. Male flowers: pedicel slender, (1—)2—3.5 mm long, slightly broadened towards the
perianth, glabrous or with some scattered minute hairs at base; buds broadly obovoid or

Fig. 26. Horsfieldia schlechteri Warb. a. Leafy twig with male inflorescences; b. mature male
flower; c. ditto, opened, showing stiped androecium; d. opened mature female flower showing gla-
brous ovary; e. twig with infructescence, fruits mature; f. mature fruit. — H. subtilis (Miq.) Warb.
var. subtilis. g. Opened male flower showing androecium, androphore hidden by the anthers;
h. twig with mature infructescence [a—c: Kostermans & Soegeng 359; d & e: BW 4307, f: BW 2900:
g: LAE 70256; h: Hoogland 3503). — Scale bar for a, e, f, h = 2 cm; for b—d, g = 0.85 mm.

184 Flora Malesiana, Ser. I, Vol. 14 (2000)

mostly subglobose, not or only somewhat compressed laterally, 1-1.5 by (I—)1.5—2 mm,
when dry wrinkled but not or slightly collapsed, apex rounded, base shortly tapered,
cleft 1/3(—1/2), lobes 0.10.2 mm thick; androecium + flattened especially in the upper
part, rather thick at base, broadly ellipsoid in outline, apex and base broadly rounded,
0.6-0.8 by 0.9-1 mm (Plate 1: 25); thecae 16, mutually appressed, the free apices up to
0.1 mm; androphore 0.4—0.5 mm; column hollow for c. 1/5. Female flowers: pedicel
1.5—2 mm long; buds ellipsoid-obovoid, 1.5—2 by 1.5 mm, cleft to c. 1/4; ovary ellip-
soid, 1-1.2 by 0.8 mm, glabrous or with a few minute hairs at apex, stigma minutely
2-lobulate, 0.1 mm long. Fruits | (or 2) per infructescence, ellipsoid, apex rounded, not
pointed or for only 1 mm, base rounded or attenuate into a pseudostalk (1.5—)2—6 mm
long, excluding pseudostalk 1.3—2(—2.5) by (0.9-1.2 cm, glabrous, drying blackish, with
or without paler pustules (or lenticels); pericarp 1-2 mm thick; fruiting pedicel 5—10
mm long; perianth not persistent. — Fig. 26a-f.

Field-notes — Bark dark brown or blackish, with longitudinal fissures. Perianths
(orange-) yellow. Fruits green-yellow or (yellow-)orange.

Distribution — Malesia: New Guinea (Papua Barat: Jayapura District; N Papua New
Guinea: W Sepik Prov.).

Habitat & Ecology — Forest on stony slopes, foothills, ridges; mossy forest, lowland
and lower montane forest; on stony clay, sandy soil, and limestone; 20—500(—1500) m
altitude; fl. & fr. throughout the year.

74. Horsfieldia sepikensis Markgr.

Horsfieldia sepikensis Markgr., Bot. Jahrb. Syst. 67 (1935) 147; W.J. de Wilde, Gard. Bull. Sing.
38. 1 (1985) 81, f. 10. — Horsfieldia spicata (Roxb.) J. Sinclair var. sepikensis (Markgr.)
J. Sinclair, Gard. Bull. Sing. 28 (1975) 125, p.p. — Type: Ledermann 8016, Papua New Guinea.

Tree 10-25 m. Twigs faintly ridged or not, 2—4 mm diameter, early glabrescent, hairs
greyish brown, 0.1 mm long or less; bark striate, not flaking; lenticels small but distinct.
Leaves membranous, oblong(-lanceolate), 8-17 by 3.5—6 cm, base attenuate, apex acute-
acuminate with the very tip usually + blunt; upper surface drying dark brown, lower
surface early glabrescent, hairs less than 0.1 mm; dots absent; midrib slightly raised
above; nerves 8-12 pairs, above thin and flat; venation lax, very faint on both surfaces;
petiole relatively long, 15-24 by 1-2 mm, leaf bud 10 by 1 mm, with hairs less than
0.1 mm. Inflorescences glabrescent or with sparse hairs (0.1 mm, 2 or 3 times branched,
many-flowered; in male: 7-12 by 4-6 cm, peduncle 0.5—1.5 cm long; in female: 2-4 by
1.5—2 cm: bracts not seen, caducous; flowers (male) in loose clusters of 2-7, glabrous,
perianth 3- (or 4-)lobed, finely punctate, pedicel not articulated. Male flowers: pedicel
slender, 2-3 mm long; buds broadly ellipsoid-obovoid, + triangular, 1.5—2 by 1.4—1.6

Fig. 27. Horsfieldia sepikensis Markgr. a. Twig with male inflorescence; b. mature male flower,
lateral view; c. ditto, opened, showing androecium; d. androecium, longitudinal section, schematic;
e & f. female flowering twigs, inflorescences axillary to leaves; g. mature female flower bud;
h. ditto, at full anthesis, showing glabrous ovary and large broadly 2-lobed stigma [a—d: Hoogland
& Craven 10255; e—h: Hoogland & Craven 10237). — Scale bar for a, e, f = 2 cm; for b—d, g, h=
0.85 mm.

185

De Wilde — Myristicaceae (Horsfieldia)

186 Flora Malesiana, Ser. I, Vol. 14 (2000)

mm, apex + acute, base + attenuate, cleft c. 5/6, lobes 0.1 mm thick; androecium + ob-
ovoid, + blunt-triangular, 1.5 by 1-1.2 mm (Plate 1: /3); thecae 24-28, tightly appressed
(sometimes touching each other in a fish-bone pattern), free apices 0.2—0.5 mm long,
+ curved into the apical hollow, column broad with hollow to c. 1/3, androphore up to
0.1 mm long, narrow. Female flowers: pedicel 1.5—2 mm long; buds ellipsoid, 2 by 1.6—
1.8 mm, 3- or 4-lobed, cleft almost to the base; ovary ovoid, 1.5-1.8 by 1.2-1.5 mm,
glabrous, style absent, stigmas relatively very large, consisting of two broad fleshy lobes
1 by 0.2 mm, only 0.1 mm high. Fruits not seen. — Fig. 27.

Field-note — Flowers yellow.

Distribution — Malesia: Papua New Guinea (East Sepik Prov.).

Habitat & Ecology — Primary and degraded forest, riverine forest; ridge forest; alti-
tude 0-50 m:; fl. & fr. throughout the year.

Notes — 1. A noteworthy species because of its predominantly 3-lobed flowers. The
only other species with 3- or 4-lobed flowers in New Guinea is the + related H. olens.
Horsfieldia sepikensis is reminiscent of Gymnacranthera paniculata vat. zippeliana,
which differs in the nature of the hairs (on leaf buds and flowers), in hairy perianths cleft
to only about halfway, in a hairy ovary, and a different texture and colour of the leaves
(whitish below).

2. Markgraf (1.c.) described the fruits as globose, 13-15 cm diameter; pericarp 2 mm
thick; seed globose, | cm diameter. The fruiting Ledermann-specimens, formerly in the
Berlin herbarium, have been destroyed.

3. The male flowers have been described by Markgraf (l.c., and probably errone-
ously) as cleft to about halfway.

75. Horsfieldia sessilifolia W. J. de Wilde

Horsfieldia sessilifolia W.J. de Wilde, Gard. Bull. Sing. 38, 2 (*1985°, 1986) 201; Tree FI. Sabah &
Sarawak 3 (2000) 389. — Type: S 27808 (Turkey bin Tran), Sarawak.

Tree c. 30 m. Twigs 8-10 mm diameter, twig apex (or leaf buds) not seen; bark gla-
brous, finely striate and rather densely set with conspicuous lenticels. Leaves chartaceous,
oblong-lanceolate, + parallel-sided, 50 by 14 cm, base broadly rounded to subcordate,
apex acute-acuminate; upper surface drying dark brown, glabrous, lower surface with
densely interwoven bright-brown hairs 0.1 (—0.3) mm; dots absent; midrib + flat above;
nerves 30(—35) pairs, flat or slightly sunken above, lines of interarching distinct; vena-
tion lax, indistinct, on the lower leaf surface hidden by the indumentum; petiole absent
or short, 2-3 by 8 mm; leaf bud not seen. Inflorescences (female) with woolly hairs
0.3—0.5 mm long, about 3 times branched, 4—5 by 3.5 cm, rather many-flowered; pe-
duncle | by 0.8 cm; bracts oblong-lanceolate, 10 mm long, inside subglabrous, outside
pubescent as the inflorescences, caducous; flowers (female) in clusters of 2-5; perianths
3-lobed, minutely pubescent in the lower half. Male flowers: not seen. Female flowers:
pedicel stout, 1 mm long, minutely pubescent; buds obovoid-ellipsoid, stout, 4.5—5 by
4—4.5 mm, coriaceous, with hairs 0.1 mm in the lower half, cleft 1/8—1/10, lobes 0.5 mm
thick; ovary ovoid, 2.5 by 2.5 mm, subglabrous, with few minute whitish hairs less than
0.1 mmon and near the base of the suture; stigma broadly 2-lipped, 0.2-0.3 by 1.5 mm,
the lips not or only faintly lobed. Fruits not seen.

Field-notes — Tree c. 30 m tall, c. 1 m girth, with stilt-roots.

De Wilde — Myristicaceae (Horsfieldia) 187

Distribution — Malesia: Borneo (Sarawak: Sibu District, 3rd Div., Sg. Tutus, Loba
Kabang P.F.); known only from the type.

Habitat & Ecology — Lowland mixed swamp forest, at apparently not more than
100 m altitude; fl. June.

Note — The only specimen known is reminiscent of some other stout-leaved
Horsfieldias, such as H. sylvestris from SE Malesia, and to a lesser extent H. pulcherrima
(Sumatra), H. splendida (Borneo), and H. superba (Peninsular Malaysia, Sumatra), but
H. sessilifolia is distinguishable in almost sessile leaves and the densely short-pubescent
lower leaf surface. Horsfieldia sylvestris is generally less stout; and has similarly
subsessile (but narrower) leaves and 2-lobed perianths. The other stout-leaved species
mentioned here all have distinctly petioled leaves.

76. Horsfieldia sinclairii W. J. de Wilde

Horsfieldia sinclairii W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 110. — Type: NGF 28886
(Streimann & Katik), Papua New Guinea.

Horsfieldia erubescens J. Sinclair, Gard. Bull. Sing. 28 (1975) 6, in sched.

Horsfieldia australiana auct. non S.T. Blake: J. Sinclair, Gard. Bull. Sing. 28 (1975) 6, p.p. — Type
in sched.: NGF 8664 (Womersley & Brass), Papua New Guinea.

Tree 4—25 m. 7Jwigs not or but faintly ridged, 1.5—3 mm diameter, early glabrescent,
with brown hairs 0.1 mm; bark finely striate, not flaking; lenticels mostly inconspicu-
ous. Leaves membranous, elliptic-oblong to oblong-lanceolate, 6-14 by 1.7—4.5 cm,
base attenuate, apex acute-acuminate, upper surface drying brown, sometimes with paler
pustules or blotches; lower surface glabrescent, hairs 0.1 mm; dots absent; midrib slen-
der, flat above, often reddish tinged, contrasting below; nerves 6—14 pairs, thin and flat
above, inconspicuous beneath, lines of interarching faint; venation lax, inconspicuous;
petiole 6-15 by 0.8-1.5 mm; leaf bud 8-15 by 1—2 mm with hairs 0.1 mm long. Jnflores-
cences subglabrous or with sparse hairs 0.1 mm; in male: many-flowered, 2—4 times
branched, 2.5—8 by 1.5-6 cm, peduncle 0.2—1 cm long; in female: up to 5(—10) by 4 cm;
bracts 0.5—2.5 mm long, caducous; flowers (male) in loose clusters of 2—5, glabrous;
perianth 2-lobed (in some specimens mixed with 3-lobed perianths), pedicel slender, not
articulated. Male flowers: pedicel 0.6—1.5 mm long; buds slightly laterally compressed,
subcircular (rarely slightly broader than long), not or somewhat collapsed on drying,
1.1-2 by 1.5-1.8 mm, cleft c. 1/2, lobes 0.1—0.2 mm thick; androecium + flattened, in
lateral view broadly circular to obtriangular, apex broadly rounded, 0.6—1.4 by 1-1.3
mm (Plate 1: 27); thecae 12-20, free apices 0.1—0.3 mm long, the column solid or hol-
lowed to c. 1/3; androphore narrow, up to 0.3 mm long. Female flowers much larger than
male, pedicel | mm long; buds ellipsoid-ovoid, 2—2.4 by 1.8—2.2 mm, cleft c. 1/3; ovary
globose, 1.6 mm diameter, densely minutely pubescent, style + absent, stigma distinctly
2-lobed, 0.2 mm long. Fruits 1-5(—10?) per infructescence, globose to short-ellipsoid,
or obovoid, base rounded or contracted into a pseudostalk up to 2 mm, ridged or not,
drying brown or blackish, 1.5—2.5 by 1.5—2 cm, glabrescent but often with remnants of
indumentum towards base (lens!), without or with few coarse lenticels or tubercles;
pericarp 4—6 mm thick, woody; fruiting pedicel 1-4 mm long; perianth not persistent.

Field-notes — Once reported with buttresses | by 1 ft.; bark rough, fissured or peel-
ing off in irregular flakes leaving concave depressions, exudate reddish; wood cream,

188 Flora Malesiana, Ser. I, Vol. 14 (2000)

straw. or brown. Flowers creamy or yellow(-orange); fragrant. Fruits glossy green, turn-
ing yellow to orange.

Distribution — Malesia: Papua New Guinea (Madang, Morobe, Northern, and Milne
Bay Proy., including Fergusson and Normanby I., Central and Gulf Prov.).

Habitat & Ecology — Understorey tree in primary and disturbed lowland and mon-
tane rain forest, flood-plain forest, along creeks, on slopes, on Castanopsis-dominated
ridges, and in Anisoptera—Hopea-dominated forest; stony places; 0—950(—1100) m alti-
tude; fl. mainly Mar., Apr., June; fr. mainly July, Oct.

Notes — 1. Characterized by slender twigs, small thin leaves with often reddish-
tinged midrib beneath, small subglobose glabrous male perianths, much larger female
flowers with pubescent ovary, and short-ellipsoid, dark brown or blackish, dry fruits,
with 4—6 mm thick woody pericarp.

2. A few deviating specimens which key out near H. sinclairii possibly represent
separate, as yet undescribed taxa and are discussed by De Wilde (1985: 112).

77. Horsfieldia smithii Warb.

Horsfieldia smithii Warb., Mon. Myrist. (1897) 270, t. 21: W.J. de Wilde, Gard. Bull. Sing. 38, 1
(1985) 77. — Myristica smithii (Warb.) Boerl., Handl. 3 (1900) 87. — Type: Smith s.n., May
1797 (in BM sheet 296), Banda.

Tree 10—20 m. Twigs 2.5—5 mm diameter, in apical part flat and 2-angled, somewhat
yellowish, lower down subterete, with two (faint) ridges or lines, glabrescent, hairs 0.1
mm long or less; bark finely striate, not flaking; lenticels rather small and inconspicu-
ous. Leaves membranous, oblong-lanceolate, 10-30 by 4-10 cm, base attenuate, apex
acute-acuminate; upper surface drying olivaceous-brown, with small paler pustules or
not, almost always with larger irregular whitish marks, lower surface early glabrescent,
hairs 0.1 mm long or less; dots absent; midrib above slender, flat; nerves 10-18 pairs,
flat or slightly raised above, inconspicuous, lines of interarching not distinct; venation
lax, indistinct; petiole 10-16 by 1.5-2.5 mm; leaf bud 10 by 2 mm with hairs 0.1 mm or
less. Inflorescences with sparse or dense hairs 0.1 mm or less; in male: (2—)3—4 times
branched, many-flowered, 5—8 by 4-8 cm; in female: 2-3 cm long; peduncle 0.5—1.5
cm: bracts not seen, caducous; flowers (male) 2-4 together, glabrous, perianth 2-lobed,
pedicel not articulated. Male flowers: pedicel slender, 1.5—2 mm; buds laterally com-
pressed, subcircular or transversely ellipsoid, or slightly reniform, dull and usually +
collapsed on drying, 2.5—3 by 3-4 mm, apex broadly rounded, base rounded or
subtruncate, or shortly tapering, cleft 2/3—3/4, lobes (0.1—0.2 mm thick; androecium in-
flated, somewhat compressed, transversely ellipsoid, 1-1.5(—2) by 2.5-3.5 mm; anthers
12-15 (i.e., 12-15 thecae at both sides of the androecium) (Plate 1: //); the thecae slen-
der, almost free, at base attached to the rim of the androphore, 1.5—2 mm long, their
upper halves deeply incurved almost to the base of the androecium, androphore + taper-
ing, 0.50.8 mm long. Female flowers: pedicel 2 mm long, thinly pubescent; buds ovoid-
ellipsoid, 2-3 mm long, cleft c. 1/2; ovary globose-ovoid, 2 by 1.7 mm, glabrous, stigma
0.1 mm long or less. Fruits 1-3 per infructescence, ellipsoid, 1.5—2(-3) by 1.41.6 cm,
glabrous, with scattered small tubercles or lenticels; pericarp 1.5—2 mm thick; fruiting
pedicel 3-5 mm long; perianth not persistent.

Field-notes — Flowers yellow, fruits yellow with whitish dots.

De Wilde — Myristicaceae (Horsfieldia) 189

Distribution — Malesia: Moluccas (Seram, Banda, Damar Is.; possibly Ternate).

Habitat & Ecology — Altitude 0—100 m; fl. May, Oct.; fr. Oct.

Notes — 1. Horsfieldia smithii is closely related to H. palauensis Kaneh. (Pacific
Islands), H. ardisiifolia and H. parviflora, with which it shares the largely free thecae
curved into the hollowed + cup- or saucer-shaped androphore; the more or less incurved
anthers of H. irya are reminiscent, but here the buds are much smaller and the anthers
shorter and free at the apex for only 0.2 mm; in H. moluccana (also with incurved an-
thers) the male buds are + pear-shaped and the anthers largely interconnate.

2. Horsfieldia smithii and related species like H. ardisiifolia, H. irya, H. moluccana,
and H. parviflora often have (somewhat) ridged or angular twigs, and in sterile or fruit-
ing stages may be confused with species with typically ridged or winged twigs, for in-
stance H. angularis, or a few species confined to West Malesia.

78. Horsfieldia sparsa W.J. de Wilde

Horsfieldia sparsa W.J. de Wilde, Gard. Bull. Sing. 38, 2 (°1985’, 1986) 194. — Type: KEP 110406
(Ogata), Peninsular Malaysia.

Horsfieldia sucosa auct. non (King) Warb.: J. Sinclair, Gard. Bull. Sing. 16 (1958) 416, f. 45, pl.
XII-A; 28 (1975) 139.

Tree 15—40 m. Twigs 4-11 mm diameter, pale whitish brown or grey-brown, con-
trasting with the brown-black colour of the dried petioles, early glabrescent, at first with
grey-brown hairs 0.1—0.2 mm; bark rather smooth or coarsely striate, sometimes with a
tendency to flake; lenticels conspicuous only towards the apex. Leaves crowded towards
the apex or not, in 3—5 rows, thickly membranous to chartaceous, oblong(-lanceolate),
(10—)18—24 by 3.5—7 cm, base long-attenuate, apex acute-acuminate; upper surface dry-
ing (dark) brown, lower surface drying bright brown, glabrous (early glabrescent); dots
absent; midrib flat or slightly raised above, glabrous; nerves 12—16 pairs, + flat above,
lines of interarching not distinct; venation lax, hardly visible above; petiole 12-34 by
1.5—-3 mm; leaf bud relatively short and broad, 8—12 by 3-5 mm, with dense hairs 0.1—
0.2 mm long. Inflorescences behind the leaves, subglabrous or glabrescent, with weak
hairs 0.1—0.2 mm; in male: 2 or 3 times branched, many-flowered, 6—12 by 4—6 cm,
peduncle (0.2—)0.5—1.5 cm long, the flowers in clusters of 4—10; in female (from in-
fructescences): few-flowered, 1.5—3 cm long; bracts minute, very early caducous; flow-
ers glabrous, perianth 3- or 4- (or 5-)lobed, pedicel not or incidentally articulated. Male
flowers: pedicel slender, 2-4 mm long; buds (slightly depressed-)globose, 1.5—2 by 2—
2.5 mm, cleft 1/3—1/2, not or but slightly collapsing on drying; lobes 0.1—0.2 mm thick,
androecium (depressed-)globose, 0.8—1 by 1—1.2 mm (Plate 2: 44); thecae 14-18, al-
most completely sessile, incurved at the apex; column broad and solid without or with
minute apical cavity (this rarely in S Thailand); androphore + slender, 0.3—0.4 mm long.
Female flowers (from young fruits): 3 mm long, 3- or 4-lobed; ovary glabrous. Fruits
2-4 per infructescence, ovoid-ellipsoid, 3—5.5 by 2.5—4.5 cm, glabrous, drying black-
ish, finely granulate, sometimes + tuberculate; pericarp 4-20 mm thick; fruiting pedicel
5-7 mm long; perianth not persistent.

Field-notes — Erect tree with clear bole, once recorded as + tapering, once with but-
tresses to | ft; bark (blackish) brown, shallowly fissured or thinly flaking; inner bark
fibrous, pale red; copious watery or sticky pink-red sap; slash wood cream or pale pink-

190 Flora Malesiana, Ser. I, Vol. 14 (2000)

brown, red-flecked. Fruits large, up to 11 cm long; pericarp thick fleshy, yellow or or-
ange red, often flushed pink, smooth, waxy, glossy; seeds rather small, up to 5 cm long.

Distribution — Peninsular Thailand; in Malesia: Sumatra, Peninsular Malaysia, Singa-
pore.

Habitat & Ecology — Primary and degraded dryland forest, periodically inundated
forest; mainly on sandy (loam) soil, sandstone; 0-500 m altitude; fl. Mar., Aug.; fr. main-
ly May-July.

Note — Horsfieldia sparsa is distinguishable from similar species with pale twigs as,
e.g., H. pallidicaula and H. sucosa, by stoutness of the twigs, dispersed and + chartaceous
leaves, sometimes crowded towards the thickish, short, apical leaf bud, and relatively
long petioles; also by the nearly glabrous inflorescences, the 3- or 4-lobed flowers, the
not or only indistinctly articulated male pedicels, and the rather distinctly stalked
androecium (androphore), generally without an apical cavity. Dry fruits reach 5.5 cm,
according to the field labels fresh fruits may reach up to 11 cm, with thick fleshy-juicy
pericarps, which obviously shrink considerably on drying. Horsfieldia pallidicaula dif-
fers in a sessile androecium; H. sucosa has articulated pedicels.

79. Horsfieldia spicata (Roxb.) J. Sinclair

Horsfieldia spicata (Roxb.) J. Sinclair, Gard. Bull. Sing. 28 (1975) 112, 113 (var. spicata), p.p.;
W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 59, f. 7. — Myristica spicata Roxb., Fl. Ind. 3
(1832) 847; (ed. 1874) 744; Warb., Mon. Myrist. (1897) 271, in obs. sub Horsfieldia smithit
Warb. — Type: Roxburgh’s description (see Sinclair, 1975: 122).

Myristica canariformis Blume, Rumphia | (1837) 190. — Horsfieldia canariformis (Blume) Merr.,
Int. Rumph. (1917) 230. — Based on Palala quarta, P. canariformis, P. dentaria Rumph., Herb.
Amb. 2, 10 (1741) 27 t. 8 (see Sinclair, 1975: 162).

Horsfieldia batjanica Warb., Mon. Myrist. (1897) 275, t. 21. — Myristica batjanica (Warb.) Boerl.,
Handl. 3 (1900) 85. — Type: introduced by Teijsmann in Hort. Bog.; original tree still cultivated
in Bogor, and collected sub Kostermans 11186, Rastini (220), Sinclair 10035.

Horsfieldia roxburghii Warb., Mon. Myrist. (1897) 277, t. 21. — Myristica roxburgii (Warb.) Boerl.,
Handl. 3 (1900) 85. — Type: Smith in Hb. Roxburgh (orig. Ternate); culta in Hort. Bog. (male
fl., orig. Ambon) (still in cultivation in Bogor sub no. IV.G.90, collected under Sinclair 10037).

Horsfieldia parviflora auct. non (Roxb.) J. Sinclair: J. Sinclair, Gard. Bull. Sing. 28 (1975) 82, p.p.

Tree 2.5-20 m. Twigs 2-4 mm diameter, early glabrescent, hairs greyish, 0.1 mm or
less; bark pale to whitish brown, contrasting with the blackish colour of dried petioles
and inflorescences, striate, not flaking; lenticels few, coarse. Leaves membranous, (el-
liptic-)oblong, 8-30 by 2.5—10 cm, base attenuate, apex acute-acuminate, often densely
speckled with pale irregular pustules of unknown origin; upper surface drying dull
olivaceous-brown, lower surface early glabrescent, at first with minute scale-like hairs;

Fig. 28. Horsfieldia spicata (Roxb.) J. Sinclair. a. Leafy twig with male inflorescences; b. mature
male flower, lateral view; c. male flower, opened, showing androecium; d. male perianth, inner
side, showing impression of androecium; e. androecium, longitudinal section, schematic; f. twig
with female inflorescence; g. mature female flower, lateral view; h. ditto, opened, showing glabrous
ovary and 2-lobed stigma; i. twig with infructescence [a—d: Beguin 1407, e: Teijsmann s.n.; f—h:
Kostermans s.n. (Hort. Bog. sub IV-H-13); i: de Vogel 3206]. — Scale bar for’a, i, 1 =2em- tor
b—e, g, h = 1.65 mm.

De Wilde — Myristicaceae (Horsfieldia) 19]

192 Flora Malesiana, Ser. I, Vol. 14 (2000)

dots absent (very minute blackish points often present); midrib flattish above; nerves
11-17 pairs, thin and flat above, lines of interarching + indistinct, venation lax, thin,
distinct or not; petiole 10-20 by 1.5-3 mm, blackish on drying; leaf bud 10 by 1.5-2
mm, with dense grey-brown hairs 0.1 mm long or less. /nflorescences early glabrescent,
blackish on drying, slender, spike-like, the lateral branches up to 2(—5) mm, peduncle
1-3 cm, not many-flowered; in male 4-10 by | cm; in female 2-3 cm long; bracts
+ blunt, 0.5—1 mm, caducous; flowers up to 3 together in male, solitary or 2 together in
female; perianth 2-lobed, early glabrescent; pedicel glabrous, not articulated. Male flow-
ers: pedicel tapered, 1.5-3 mm long; buds + obovoid or short pear-shaped, rather com-
pressed, the upper part subcircular to reniform, about as long as broad or slightly broader
than long, 2.3-3 by 3-3.5 mm, base + tapering into the pedicel; buds cleft (nearly) to the
base. lobes 0.2 mm thick; androecium laterally compressed, reniform, apex broadly
rounded, base broadly attached, 1.5—1.8 by 2.5 mm, androphore + absent (Plate 1: 3);
thecae (24—)32—44, closely set, free apices 0.1—0.4 mm, at one side weakly to strongly
incurved into the apical cavity to 1/4-1/2(—3/4, see note 2) of the column. Female flow-
ers: buds subglobose, 2—2.5 by 2.2-2.8 mm, at base passing into the tapered pedicel
1.5-2.5 mm, cleft 1/2—2/3; ovary broadly ovoid, glabrous, 1.5 by 1.5 mm, stigma minute,
faintly 2-lobed, 0.1 mm high. Fruits 1—5 per infructescence, short-ellipsoid, 1.5—2(—2.5)
by 1.2-1.8 cm, glabrous, drying blackish, without pustules; pericarp 1.5 mm thick; fruit-
ing pedicel 4-10 mm long; perianth not persistent. — Fig. 28.

Field-notes — Buttresses up to 50 cm out and high, or absent; bark often not fissured,
peeling off; exudate watery, not or only slightly reddish; sapwood usually cream, gradu-
ally passing into the darker heartwood. Flowers greenish or ochre-yellow. Fruits orange.

Distribution — Malesia: Moluccas (Morotai, Halmahera, Ternate, Bacan, Ambon,
Seram, Buru).

Habitat & Ecology — Forest on alluvial soils, deep clay, soil rich in humus, and
porous volcanic soil over schists; 0—1000 m altitude; fl. & fr. throughout the year.

Uses — According to de Vogel 3206, the outer bark, mixed with ‘Kuleman’ (a differ-
ent species of Horsfieldia), is used for curing hepatitis.

Notes — 1. Horsfieldia spicata is recognizable at first glance by its spike-like inflo-
rescences drying blackish, and by the pale twigs contrasting with the blackish dried
petioles and the inflorescences.

2. Possibly its most closely related species is H. moluccana. Apart from the charac-
ters as used in the key, H. spicata differs from H. moluccana in the more membranous
leaves. Teijsmann s.n. (L), from Ambon, seems intermediate between H. spicata and
H. moluccana, as explained by De Wilde (1. c.: 63).

80. Horsfieldia splendida W. J. de Wilde

Horsfieldia splendida W.J. de Wilde, Gard. Bull. Sing. 38, 2 (1985’, 1986) 213, f. 24; Tree FI.
Sabah & Sarawak 3 (2000) 390. — Type: S 33723 (Othman), Sarawak.

Tree 10—20(—30) m. Twigs 4-7 mm diameter, late glabrescent, at first with dense
woolly, yellow-brown to rusty hairs 1-1.5 mm; bark dark brown or blackish, coarsely
longitudinally striate and fissured, later on cracking and + flaking; lenticels few, indis-
tinct. Leaves membranous to chartaceous, elliptic-oblong to oblong-oblanceolate, 18—
45 by 6.5—17 cm, base almost rounded to attenuate, apex acute-acuminate; upper surface

De Wilde — Myristicaceae (Horsfieldia) 193

drying dull olivaceous-brown, early glabrescent, lower surface with conspicuous rather
dense, flossy, stalked-dendroid hairs, + evenly spaced and of about the same size, hairs
0.5—1.5 mm long; dots absent; midrib late glabrescent above, rather flat; nerves 18—25
pairs, sunken above, lines of interarching distinct, regular; venation lax, distinct; petiole
12-17 by 3.5—5 mm, pubescent; leaf bud 15(—20) by 5 mm, with hairs |—1.5 mm long.
Inflorescences with dense woolly dendroid hairs 1—1.5 mm; in male: many-flowered,
4 (or 5) times branched, 6-16 by 3—12(—16) cm, peduncle 0.5—2 cm; in female: rather
few-flowered, 3—5 cm long; bracts + ovate-triangular, acuminate, 0.4—1 cm long, densely
pubescent, caducous; flowers in male 3-8 in loose clusters, perianth 3- (or 4-)lobed,
glabrous or thinly pubescent, pedicel slender, glabrous or subglabrescent, hairs 0.3—0.5
mm long, not articulated. Male flowers: pedicel 1—-1.5 mm long; buds (depressed) sub-
globose, 1.4-2 by 1.6—2 mm, cleft 1/3 to nearly 1/2, lobes 0.2 mm thick; androecium
depressed-subglobose, + impressed in the centre, circular or faintly triangular in cross
section, 0.5—0.6 by 1—1.3 mm (Plate 2: 57); thecae 16—20, completely sessile, incurved
towards the apex; column broad, solid, with small apical hollow to 1/3(—1/2); androphore
+ slender, 0.3(—0.5) mm long. Female flowers (from remnants under fruits): pedicel
0.5—1 mm long, pubescent; buds 3 by 2.5 mm, 3- or 4-lobed, pubescent; ovary ovoid-
ellipsoid, pubescent; stigma minutely 2-lobed, 0.1 mm high. Fruits 2-7 per infructes-
cence, broadly ovoid-ellipsoid, 2-3 by 1.5—2 cm, pubescent (hairs 0.5 mm), without
lenticels or tubercles; pericarp 3 mm thick; fruiting pedicel | mm long; perianth persist-
ent. — Fig. 29.

Field-notes — Bark of trunk usually blackish, (red-)brown, fissured or flaking: inner
bark soft, light brown or reddish, laminated, exudate reddish; wood yellowish or pale,
soft or medium hard. Flowers yellow, with odour of balsam of Peru. Fruits red or orange.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, W and E Kalimantan).

Habitat & Ecology — Lowland mixed dipterocarp forest, edges of swamp forest,
kerangas (with Dryobalanops fusca dominant), montane forest; brown or yellowish soil,
tuff-plateau; 0—600(—1500) m altitude; fl. & fr. throughout the year.

Notes — 1. Material of this species was formerly mixed up with H. reticulata, a spe-
cies with smaller leaves but with almost similar flowers.

2. Horsfieldia splendida is closely related to H. grandis, H. reticulata, H. rufo-lanata,
and H. tomentosa, and less so to H. flocculosa, all having similar male flowers. The peri-
anth remains persistent under the fruits in these species, except for H. rufo-lanata and
H. tomentosa.

81. Horsfieldia squamulosa W. J. de Wilde

Horsfieldia squamulosa W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 93. f. 13g (excl. specimens
now in H. coryandra); Blumea 32 (1987) 460. — Type: NGF 42995 (Henty & Barlow), Papua
New Guinea.

Shrub or slender treelet, 2-10 m. Twigs 1-2 mm diameter, with rusty hairs 0.1(—0.2)
mm, late glabrescent; bark finely striate, not cracked nor flaking; lenticels absent or
few and inconspicuous. Leaves membranous or thinly chartaceous, oblong to lanceolate,
7-15 by 1.5-4 cm, base acute or attenuate, apex acute-acuminate to long-tapering; blade
drying blackish brown, glabrous above, beneath glabrescent (with few minute hairs re-
maining on midrib); dots absent; midrib very slender, slightly raised above; nerves

194 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Horsfieldia) 195

12-25 pairs, indistinct, flat or slightly raised above, lines of interarching distinct or in-
distinct; venation indistinct; petiole 7-12 by (0.5—)1 mm, glabrescent; leaf bud S—7(—10)
by 0.5—1 mm, with dense minute rusty hairs 0.1 mm. /nflorescences among the leaves,
small, twice branched (lowest branch 2—5 mm from the base), with hairs 0.1 mm or less;
bracts ellipsoid, 0.5-1 mm long, caducous; in male: 2 by 2 cm, few-branched, few-
flowered; in female: 1—1.5 cm long, not or somewhat branched, 1—3-flowered; flowers
solitary or up to 3 together, perianth 2-lobed, thinly with stellate-dendroid hairs 0.1—0.2
mm long, pedicel thinly pubescent, not articulated. Male flowers: pedicel 3—3.5 mm
long: buds slightly flattened, subglobose or broadly ellipsoid or obovoid, 2.2—2.4 mm
diameter; cleft c. 1/8, perianth 0.7—0.8 mm thick, the lobes at apex 0.2 mm thick;
androecium club-shaped, apex subacute to broadly rounded, 1.7—1.8 by 0.6—0.7 mm
(Plate 1: 79); thecae 8, apical, mutually appressed, 0.4 mm long, consolidated with the
androphore, apex slightly free or not; androphore in the lower 2/3 with dense minute
pale brown hairs less than 0.1 mm, the apical part coarsely irregularly warted or wrin-
kled-bullate, glabrous; column not hollow at apex. Female flowers: pedicel 2—2.5 mm
long; buds subglobose, 1.8 mm diameter, lobes 2, cleft c. 1/8; ovary ellipsoid-ovoid, 1.5
by 1(—1.1) mm, densely minutely pubescent; stigma minute, of two suberect lobes 0.1—
0.2 mm. Fruits 1(—3) per infructescence, ovoid-ellipsoid, including the 2-3 mm long
rostrum 1.6—1.7 by 0.9—1 cm, base contracted into pseudostalk 4—S mm; all thinly with
pale brown stellate hairs 0.1 mm or less, drying blackish, without lenticels; pericarp
0.5(-1) mm thick; fruiting pedicel slender, slightly broadened to the apex or not, 8-12
mm long; perianth not persistent. — Fig. 14g.

Field-notes — Shrub or small, slender tree, 2-10 m tall, dbh to 10 cm diameter; nu-
merous excurrent horizontal branches, or branches + whorled, drooping; bark brown,
wood white, exudate red. Flowers yellow. Fruits conical, orange(-yellow); aril not aro-
matic.

Distribution — Malesia: Papua New Guinea (Western Prov., about 5—6° S, 141° E).

Habitat & Ecology — Primary and disturbed forest; on ridge-crests, undulating land,
forest bordering sago-swamp; locally plentiful in forest undergrowth; 100-200 m alti-
tude; fl. & fr. July—Aug.

Notes — 1. Horsfieldia squamulosa is related to H. coryandra, which generally has
similar leaves (though usually with coarser venation), but differs in a more elongate
male bud, (almost) glabrous androphore, and fruits with a pseudostalk of only 2 mm
long. Vegetatively both species much resemble H. schlechteri from the same region, es-
pecially in the foliage.

2. Horsfieldia squamulosa forms a coherent group with H. clavata, H. coryandra,
H. crux-melitensis, and H. clavata (the clavata-group), species of small trees or shrubs
with club-shaped androecium consisting of a relatively large androphore and small ses-

Fig. 29. Horsfieldia splendida W.J. de Wilde. a. Apical part of leafy twig; b. twig with immature
male inflorescence axillary to leaf scar; note bracts; c. mature male inflorescence; d. mature male
flower, lateral view; e. ditto, longitudinally opened, showing androecium; f. androecium, longitudi-
nal section, schematic; g. twig with infructescence, fruits immature; h. mature fruit; note persistent
perianth [a, b: Anderson 12916; c—f: BNB For. Dept. 4782; g: SAN 16927; h; SFN 35606]. — Scale
bar for a—c, g, h = 2 cm; for d—f = 0.85 mm.

196 Flora Malesiana, Ser. I, Vol. 14 (2000)

sile anthers at the apex. The remote species H. pulverulenta, H. sterilis, and H. triandra
have somewhat similar androecia.

82. Horsfieldia sterilis W. J. de Wilde

Horsfieldia sterilis W.J. de Wilde, Gard. Bull. Sing. 38, 2 (‘1985°, 1986) 224; Tree Fl. Sabah &
Sarawak 3 (2000) 390. — Type: SAN 30597 (Aban Gibot), Sabah.

Tree or shrub, 3-12 m. Twigs 2.5—4 mm diameter, sometimes grey-brown and con-
trasting with the blackish colour of the dry petioles, early glabrescent, hairs rusty to
grey-brown, 0.1—0.2 mm; bark finely striate, not tending to crack or flake; lenticels
conspicuous or not. Leaves membranous, elliptic-oblong to oblong(-lanceolate), 13-37
by 4.5-11 cm, base attenuate, apex acute-acuminate; upper surface glabrous, drying dark
brown or blackish, lower surface glabrous; dots absent; midrib flat above, glabrous;
nerves 11-18 pairs, + flat above, lines of interarching not particularly distinct; venation
lax. faint on both surfaces; petiole 7-20 by 23.5 mm; leaf bud 7-12 by 22.5 mm, with
dense pale-rusty hairs 0.1—0.2 mm long. Inflorescences subglabrescent, hairs + sparse,
0.1—0.2 mm long; in male: lax, 10-20 by 5—8 cm with few side-branches, 2 or 3 times
branched, not densely flowered, peduncle 3-13 cm long, with several bract scars, or
branched from the base; in female: elongate, slightly branched, almost spike-like, 5-10
cm long; bracts oblong-lanceolate, 3-8 mm long, acute, with hairs 0.1—0.3 mm espe-
cially at the margins, caducous; flowers (male) in loose clusters of 2—6, glabrous, peri-
anth 2-lobed, pedicel not articulated. Male flowers: pedicel slender, 1-1.5 mm long;
buds broadly obovoid-subglobose, 1.5—1.7 mm diameter, apex broadly rounded or some-
what depressed, base subattenuate, cleft c. 1/4, lobes thick-fleshy, 0.3—0.4 mm thick,
shrinking, not collapsing on drying; androecium broadly obovoid, 0.8 by 0.6—0.7 mm,
apex + rounded, subcircular in cross section; anthers 3 or 4 or 6—8 (or with 6-8 thecae?),
completely sessile at apex of the androecium, (0.4 mm long, connectives broad, + trian-
gular, with narrow thecae (Plate 3: 62); column broad, solid, apical hollow absent or
inconspicuous; androphore broad, tapering to below, (0.4 mm long. Female flowers: buds
obovoid, 22.5 mm long, cleft c. 1/4, ovary subglobose, 1.5 mm diameter, glabrous,
stigma lobes minute. Fruits (immature) 4—10 per spike-like infructescence, ellipsoid,
2.2 by 1.8 cm, glabrous, drying blackish, not tubercled; pericarp 2 mm thick; fruiting
pedicel 3-4 mm long; perianth persistent under immature fruits.

Field-notes — Small trees or shrubs; bark grey-brown, not fissured; inner bark with
orange-red sap; sapwood pale yellow.

Distribution — Malesia: Borneo (C & NE Kalimantan and C & SE Sabah).

Habitat & Ecology — Forest on hillsides and riverbanks; 80—500 m altitude; fl. Apr—
July.

Note — Horsfieldia sterilis is noticeable by its 2-lobed perianth and the subconical
broadly obovoid androecium of which only the apical half bears apparently reduced
anthers: the basal part of the androecium or androphore 1s sterile, broad and tapering.

The structure of the androecium is reminiscent of that of species as H. crux-melitensis
and H. clavata from New Guinea, but in these species the androecium is much more
elongate. In general habit H. sterilis resembles, e.g., H. pallidicaula or H. sucosa, be-
cause of its pale twigs and blackish drying leaves, but H. sterilis differs in more elongate
inflorescences.

De Wilde — Myristicaceae (Horsfieldia) 197

83. Horsfieldia subalpina J. Sinclair

Horsfieldia subalpina J. Sinclair, Gard. Bull. Sing. 16 (1958) 410; 28 (1975) 131; W.J. de Wilde,
Gard. Bull. Sing. 39, 1 (1986) 40; Tree Fl. Sabah & Sarawak 3 (2000) 392. — Type: Wray 467,
Peninsular Malaysia.

Tree 6-30 m. Twigs 2.5—5 mm diameter, early glabrescent, hairs greyish brown, 0.1
mm long or less; bark finely to coarsely striate, not flaking; lenticels usually conspicu-
ous. Leaves membranous or chartaceous, (elliptic-)oblong, 15—27 by 5—10 cm, base at-
tenuate, apex acute(-acuminate), sometimes + blunt; upper surface drying olivaceous to
dark brown, lower surface glabrous; dots absent; midrib + flat above; nerves 9—18(—20)
pairs, thin, flat or slightly raised above, lines of interarching indistinct; venation lax,
faint on both surfaces; petiole S—15 by 2-3 mm; leaf bud 12—20 by 2-3 mm, with dense
grey-brown to rusty hairs 0.1 mm long or less. Inflorescences behind the leaves, with
sparse (subsp. kinabaluensis) or dense hairs 0.1 mm long; in male: rather stout, about
3 (or 4) times branched, many-flowered, 5—14 by 3-10 cm, peduncle (0.6—)1—3 cm long;
female 2 (or 3) times branched, 2—7 by 1.5—4 cm, fewer flowered than male; bracts
ellipsoid to oblong, acutish, densely short-pubescent, 2-5 mm long, caducous; flowers
in male in loose clusters of 2—5, perianth 3- (or 4-)lobed, glabrous, pedicel glabrous
(subsp. kinabaluensis) or partly thinly pubescent (subsp. subalpina), not articulated. Male
flowers: pedicel 1.5—2 mm, slender; buds broad-ellipsoid or subglobose, 1.6—2.3 mm
long, cleft c. 1/2, lobes 0.2(—0.3) mm thick; androecium globose or broadly ellipsoid,
1-1.5 mm long, cross section circular (Plate 3: 84); thecae 16-24, almost completely
sessile, free apices 0.1 mm, curving towards the apex and concealing the narrow apical
hollow, 0.3—0.5 mm deep, of the broad column; androphore narrow, 0.2—0.3 mm long,
largely hidden by the anthers. Female flowers: pedicel 1.5 mm long; buds ellipsoid,
22.5 by 1.8-2.1 mm, glabrous (subsp. kinabaluensis), cleft (1/3—)1/2; ovary subglobose
to broadly ellipsoid, 1.2—-1.5 mm long, glabrous, stigma minutely 2-lobed, 0.1 mm high.
Fruits 2—6 per infructescence, subglobose, broadly ellipsoid, or ellipsoid-oblong, 2.5—5
cm long, glabrous; fruiting pedicel 3—7 mm long; perianth not persistent.

Distribution — Malesia: two subspecies, one in montane Peninsular Malaysia, one in
the Kinabalu area in Sabah.

Note — In the group of species with H. obscura and H. subalpina, a number of fruit-
ing specimens could not be assigned satisfactorily to any of the known taxa. More infor-
mation on their affinities can only be obtained from male flowering specimens which
have vegetative characters closely matching those of the fruiting material. These fruit-
ing collections have been discussed under H. obscura by De Wilde (1.c., 1986: 44 &
Blumea 32, 1987: 469).

KEY TO THE SUBSPECIES

la. Pedicel of male flowers pubescent, at least in the lower half. Male buds subglobose;
androecium subglobose, thecae 18-24. Fruits subglobose to broadly ellipsoid, 2.5—
PREMEMVIOMNO: Sage arses is ut} eetEEet Ss pet ale | Spey ae Sat yet eeenct eM. Gut ol oe a. subsp. subalpina

b. Pedicel of male flowers glabrous. Male buds (broadly) ellipsoid; androecium (broadly )
ellipsoid, thecae,16,or18.Fruits ellipsoid; 3-5);em long. .2)-)....0has,aace bet

198 Flora Malesiana, Ser. I, Vol. 14 (2000)

a. subsp. subalpina

Inflorescences rather densely pubescent. Male pedicel, at least in the lower half thinly
pubescent with hairs 0.1 mm. Male flowers: buds subglobose to broadly ellipsoid, 1.6—
2.3 by 1.6-2.2 mm; androecium subglobose, 1—1.5 by 1—1.2 mm; thecae 18-24. Female
flowers not seen. Fruits subglobose to broadly ellipsoid, 2.5—4 by 22.3 cm, apex and
base rounded, drying brown-blackish, without or with a few tubercles; pericarp 3—4 mm
thick.

Field-notes — Bark smooth or with shallow widely spaced fissures; slash reddish,
with red sap; slash wood whitish. Flowers yellow. Fruits greenish yellow and glaucous,
or yellow; seed white. Fresh fruits may reach 4.5—5 cm length, the dry fruits are only
2.5—4 cm long.

Distribution — Malesia: Peninsular Malaysia (Perak, Pahang, Selangor; Fraser’s Hill,
Genting Highlands).

Habitat & Ecology — Mountain forest, 800-1500 m altitude; fl. Apr., June; fr. Jan.,
Aug.—Dec.

b. subsp. kinabaluensis W. J. de Wilde

Horsfieldia subalpina J. Sinclair subsp. kinabaluensis W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986)
Al: Tree Fl. Sabah & Sarawak 3 (2000) 392. — Type: Clemens 33136, Sabah.

Inflorescences very sparsely pubescent. Male pedicel glabrous. Male flowers: buds
(broadly) ellipsoid, 1.7—2 by 1.5—1.8 mm; androecium broadly ellipsoid, 1.1—1.2 by 0.8—
| mm: thecae 16 or 18. Female flowers as described under the species. Fruits ellipsoid to
broadly ellipsoid, 35.1 by 1.7—2.5 cm, apex + narrowly rounded, base rounded, drying
bright brown to blackish brown, without tubercles; pericarp 4-5 mm thick.

Field-notes — Buttresses absent; bark slightly fissured, reddish brown; inner bark
fibrous, whitish turning brown, or soft, and then yellowish; cambium pale yellow; wood
white to yellowish, medium hard, heartwood not differentiated. Flowers bright yellow.
Fruits yellow-red.

Distribution — Malesia: Borneo (Sabah: Mt Kinabalu and vicinity; one doubtful col-
lection from E Sarawak).

Habitat & Ecology — Montane (oak) forest; clayish soil; 1400-2000 m altitude; fl. &
fr. throughout the year.

84. Horsfieldia subtilis (Mig.) Warb.

Horsfieldia subtilis (Miq.) Warb., Mon. Myrist. (1897) 286, t. 23. — Myristica subtilis Mig., Ann.
Mus. Bot. Lugd.-Bat. 2 (1865) 50. — Type: Zippelius (78), West New Guinea.
For more references and synonyms see the varieties.

Tree 2-10(—20) m. Twigs 1-3(—8) mm diameter, early glabrescent, at first with grey-
brown hairs 0.1 mm long or less; bark finely striate, not flaking; lenticels fine, usually
present. Leaves membranous or in var. calcarea t chartaceous, elliptic to oblong-lanceo-
late, 6-28 by 2—9.5 cm, base attenuate, apex acute-acuminate; upper surface drying dull
olivaceous to brown, with or without fine paler dots, lower surface early glabrescent,
hairs 0.1 mm or less: dots absent; midrib slightly raised above; nerves 6—16 pairs, above
thin, flat or raised above, lines of interarching faint or distinct; venation lax, usually in-

De Wilde — Myristicaceae (Horsfieldia) 199

distinct; petiole 5-13 by 1-2.5 mm; leaf bud 6-12 by 1.5—2 mm, with hairs 0.1 mm.
Inflorescences glabrescent or with sparse stellate scale-like hairs 0.1—0.2 mm long; in
male: 1—2(—3) times branched, few- to many-flowered, 2—8(—9) by 1.5—6 cm, peduncle
up to 1 cm long; female 2—5(—8) cm long; bracts 0.5—2 mm long, glabrescent, with fim-
briate margins, caducous; flowers in male in loose clusters of 1—8(—10), glabrous, peri-
anth 2-lobed, pedicel slender, not articulated. Male flowers: pedicel 1-3 mm long; buds
compressed, in lateral view circular, + broadly transversely subellipsoid, or broadly
obtriangular (rarely slightly longer than broad), collapsing on drying, 1.3-2.4 by 1.8-3
mm, apex subtruncate or (broadly) rounded, base subtruncate to short-cuneate, cleft
1/5—1/3(-1/2), lobes 0.1—0.2 (in var. aucta up to 0.5) mm thick; androecium flattened
towards the apex, base broad and usually almost cylindrical, in lateral view sub-
quadrangular, apex broadly rounded or subtruncate, nearly filling the perianth, (0.7—)1—
1.7 by 1.4-1.6 mm (Plate 1: 24); thecae 18-24, + erect, free apices to 0.1 mm long,
column narrowly hollowed for 1/4-1/3; androphore + slender, 0.2—0.5 mm long, some-
times + hidden by sagged anthers. Female flowers: pedicel 1-5 mm long; buds broadly
ellipsoid, ovoid, or subglobose, 1.8—2.5 by 2-2.5 mm, cleft c. 1/3; ovary ovoid, 1.1—1.5
by 0.8—1.1 mm, glabrous, style with minutely bilobed stigma, 0.2 mm long. Fruits (1—)
5-15 per infructescence, globose or subglobose, 0.9—1.3 by 0.8—-1.3 cm (pseudostalk up
to 1 mm), or in varieties larger, rather ellipsoid, 1.4—2 by 1.1—1.4 cm, with the apex
rounded to acutish, base rounded, without or with pseudostalk up to 3 mm; glabrous,
drying blackish, without or with minute paler tubercles or lenticels; pericarp 1(—2) mm
thick; seeds subglobose to ellipsoid; fruiting pedicel 1-7 mm; perianth not persistent. —
Fig. 26g, h.
Distribution — Malesia: Moluccas (Aru Is.), whole of New Guinea.

KEY TO THE VARIETIES

la. Male perianth buds 2-3 mm wide. Fruits globose or short-ellipsoid, 0.9—1.3 cm long

memome pseudostalkO—l mm long’... 20.0055. a. ieee eae a. var. subtilis

b. Male perianth buds 2.5—3 mm wide (always?). Fruits larger, short-ellipsoid, 1.4—2
cumene meucdme pseudostalk 3 mm lOMg oi)... eee ee ee J;

2a. Leaves chartaceous, elliptic, 6-9 cm long. Pseudostalk of fruits 2-3 mm long. —
Limestone area, Papua Barat, SW Bird’s Head; 200-300m ..... c. var. calcarea

b. Leaves coriaceous or membranous, elliptic-oblong, 10—22(—26) cm long. Pseudostalk
of fruits 0-3 mm long. — Papua New Guinea, possibly also Papua Barat (Bird’s
eae O00 = lL OOO ily. ts sale os. is MSE 9 Geka tas tes J OO ods ea ee 3

3a. Leaves membranous. Perianth (female) glabrous inside ........... b. var. aucta
b. Leaves coriaceous. Perianth (female) hairy inside ............. d. var. rostrata

a. var. subtilis

Horsfieldia subtilis (Mig.) Warb., Mon. Myrist. (1897) 286, t. 23; Markgr., Bot. Jahrb. Syst. 67
(1935) 152; J. Sinclair, Gard. Bull. Sing. 28 (1975) 132; W.J. de Wilde, Gard. Bull. Sing. 38, 1
(1985) 102, f. 15g, h. — Myristica subtilis Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 50.

Horsfieldia aruensis Warb., Mon. Myrist. (1897) 284, t. 23; Markgr., Bot. Jahrb. Syst. 67 (1935)
154. — Myristica aruensis (Warb.) Boerl., Handl. 3 (1900) 85. — Type: Beccari s.n. (FI 7622,
A-C, 7623, n.v.), Aru Is.

200 Flora Malesiana, Ser. I, Vol. 14 (2000)

Horsfieldia lauterbachii Warb., Mon. Myrist. (1897) 285, t. 23: K. Schum. & Lauterb., Fl. Schutzgeb.
Siidsee (1900) 324: Pulle, Nova Guinea 8 (1912) 635; Markgr., J. Arnold Arbor. 10 (1929) 213;
Bot. Jahrb. Syst. 67 (1935) 153. — Type: Lauterbach 805 (B lost, BRSL, n.v.), New Guinea.

Horsfieldia ramuensis Warb. in K. Schum. & Lauterb., Nachtr. Fl. Schutzgeb. Siidsee (1905) 266.
— Syntypes: Rodatz & Klink 20, 24 (both B lost), New Guinea.

Horsfieldia globularia auct. non (Blume) Warb.: K. Schum. & Lauterb., Fl. Schutzgeb. Siidsee (1900)
324.

Horsfieldia nesophila auct. non (Miq.) Warb.: Pulle, Nova Guinea 8 (1912) 635.

Leaves membranous, elliptic to oblong. Male flowers: buds 1.8-3 mm wide. Fruits
(sub)globose, 0.9-1.3 by 0.8-1.1(-1.3) cm, including the pseudostalk up to 1 mm long.
— Fig. 26g, h.

Field-notes — Low trees, usually 3—5 m tall; bole straight; bark greyish black or grey-
brown, finely longitudinally fissured, with broadened lenticels; branches often horizon-
tal or drooping; exudate pinkish, or colourless and turning reddish; wood straw, usually
mottled with + red streaks. Flowers (orange-)yellow. Fruits (greenish) yellow or orange.

Distribution — As for the species.

Habitat & Ecology — Understorey tree of primary and degraded forest, dry or marshy
forest. often tidal (fresh water) or riverine forest; on alluvial, clayey, and sandy clayey
soils, limestone, or coral soils; 0-800 m altitude; fl. & fr. throughout the year. Stems
once reported as inhabited by ants.

Uses — Leaves and twigs burnt as a mosquito repellent.

b. var. aucta W.J. de Wilde

Horsfieldia subtilis (Miq.) Warb. var. aucta W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 105. —
Type: Jacobs 8972, Papua New Guinea.

Leaves membranous, elliptic-oblong, 1 1—20(—26, see note 2) by 3-7.5(—9) cm, apex
for 1-1.5 cm acute-acuminate. Male flowers: buds obtriangular, 2.5—3 mm wide (al-
ways?). Fruits short-ellipsoid, 1.5—2 by 1.1-1.5 cm including pseudostalk up to 2.5 mm
long.

Field-notes — Shrub or low tree, 3-8 m. Male flowers fleshy, dark yellow. Fruits
glossy orange, hard; aril dark orange, or red, at the base black.

Distribution — Malesia: New Guinea (possibly W Papua Barat: Bird’s Head, near
Manokwari, cf. De Wilde, I.c.: 106; Papua New Guinea).

Habitat & Ecology — Montane primary and degraded rain forest, on well-drained
volcanic soil, or peaty soil; 600-1000 m altitude; fl. Sept., Oct.; fr. July—Oct.

Notes — 1. The male flowers are stouter and have a thicker perianth as compared with
those of var. subtilis. The inflorescences are 8—9 cm long, stouter than generally found
in the type variety.

2. Sands et al. 6165, from Bird’s Head, with leaves to 26 by 9 cm and fruits 2 by eS)
cm, is like a giant form of var. calcarea; its twigs measure 4(—5) mm in diameter at the
apex. Possibly it should be recognized as still another, new variety.

c. var. calearea W.J. de Wilde

Horsfieldia subtilis (Mig.) Warb. var. calcarea W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 104.
— Type: BW 15270 (Vink), New Guinea, Bird’s Head.

De Wilde — Myristicaceae (Horsfieldia) 201

Leaves thinly chartaceous, elliptic, 6—9 by 2.5—4 cm, at the apex proportionally long-
acute-acuminate for 1—-1.5 cm. Male flowers not seen. Fruits ellipsoid, 1.7—1.9 by 1.2—
1.4 cm, including pseudostalk 2—3 mm long.

Field-notes — Shrub, 5 m tall; rather common. Ripe fruits orange.

Distribution — Malesia: W Papua Barat (SW Bird’s Head).

Habitat & Ecology — Secondary forest on limestone rock with thin clay cover;
220-300 m altitude; female fl. May; fr. Mar., May.

Note — The dry pericarp in BW 15270 (Vink) suggests that the fruits were rather
fleshy in the fresh state and drying left the pseudostalks distinct, 3 mm long.

d. var. rostrata (Markegr.) J. Sinclair

Horsfieldia subtilis (Miq.) Warb. var. rostrata (Markgr.) J. Sinclair, Gard. Bull. Sing. 28 (1975)
136; W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 106. — Horsfieldia rostrata Markgr., Bot.
Jahrb. Syst. 67 (1935) 152. — Type: Ledermann 8916 (B lost; iso SING, n.v.), Papua New
Guinea.

Leaves chartaceous to coriaceous, cuneate-obovate, 9-13 by 2—4.5 cm, apex shortly
acute-acuminate. Male flowers not known. Female perianth pilose inside (see note).
Fruits ellipsoid, (apex 2 mm rostrate) 2 by 1.2 cm, including the 2-4 mm long pseudostalk.

Distribution — Malesia: NE Papua New Guinea (Sepik Prov., ‘Etappenberg’; known
only from the type).

Habitat & Ecology — Mossy montane forest with much Agathis, c. 850 m altitude;
female fl. & fr. Oct.

Note — Markgraf, l.c., described the perianth of the female flowers (Ledermann 8916,
lost) as pilose inside, which is highly remarkable, and the ovary and fruits as glabrous.

85. Horsfieldia sucosa (King) Warb.

Horsfieldia sucosa (King) Warb., Mon. Myrist. (1897) 322; W.J. de Wilde, Gard. Bull. Sing. 38
(1985, 1986) 188; Tree Fl. Sabah & Sarawak 3 (2000) 393. — Myristica sucosa King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 301, pl. 172. — Lectotype: King’s coll. 4647 (fr., K), Peninsular
Malaysia.

Tree 6—20 m. Twigs 2-10 mm diameter, (whitish) grey-brown or straw-coloured, con-
trasting with the blackish colour of the dried petioles, early glabrescent, at first with
rusty or greyish hairs 0.1—0.3 mm long; bark coarsely striate or not, with a tendency to
flake; lenticels conspicuous only on the very young parts. Leaves in 2 or 3 rows, mem-
branous to thin-chartaceous, elliptic-oblong to oblong-(ob)lanceolate, 14-28 by 4.5-8.5
cm, base (long-)attenuate, apex acute-acuminate; upper surface glabrous, drying green-
ish to blackish brown, lower surface early glabrescent, drying light brown; dots absent,
but scattered smaller blackish points often present; midrib + flat above, glabrous; nerves
13-17 pairs, slender, flat or slightly raised above, lines of interarching indistinct; vena-
tion lax, faint; petiole 10-20 by 2-3 mm; leaf bud 10-15 by 2-3 mm, with rusty to
greyish brown hairs 0.1—0.3 mm long. Inflorescences behind the leaves, thinly pubes-
cent or late glabrescent, hairs 0.1—0.2 mm long; in male: 3 or 4 times branched, many-
flowered, lax or condensed, 7-19 by 5—16 cm, peduncle 1—2 cm long, flowers in clusters
of 3—7; in female rather few-flowered, 1—2 cm long; bracts + ovate-elliptic to lanceolate,

202 Flora Malesiana, Ser. I, Vol. 14 (2000)

acutish, pubescent, 1.5—4 mm long, late caducous; flowers with perianth either mostly
2-lobed (Borneo, see under the subspecies) or mostly 3-lobed (Sumatra, Peninsular
Malaysia), glabrous, pedicel slender, usually glabrous (see under subsp. sucosa), articu-
lated, contrasting with the pubescent branches of the inflorescence. Male flowers: pedi-
cel 1-2 mm long; buds (depressed) globose, apex + flat to broadly rounded, 1.2—1.5 by
1.5-2 mm, cleft 1/3—1/2, when dry not or slightly collapsed at the apex, lobes 0.2—0.3
mm thick; androecium depressed globose, 0.3—0.6 by 0.8-1.2 mm, circular or (in Bor-
neo) broadly ellipsoid in cross section (Plate 2: 42); thecae 14-22, completely sessile
with + incurved apex, column broad, + saucer-shaped, with broad, flattish, apical hollow
to nearly halfway; androphore rather narrow, 0.1—0.3 mm long. Female flowers (subsp.
bifissa): pedicel 1-1.5 mm long, when young with minute hairs 0.1 mm long or less
towards the base; buds broadly ellipsoid, 2.8-3.5 by 2.2-3 mm, 2-lobed, cleft c. 1/4;
ovary 1.5 mm diameter, glabrous, stigma shallowly 2-lobed, 0.1 mm high. Fruits 1—4
per infructescence, broadly ovoid-ellipsoid, apex + narrowly rounded, 2.3-3.5 by 2-2.5
cm, glabrous, drying blackish, finely granulate, sometimes + tuberculate; pericarp 4 mm
thick; fruiting pedicel 1-2 mm long; perianth in Peninsular Malaysia persistent (see
further under the subspecies).

Distribution — Malesia: Sumatra, Peninsular Malaysia, Borneo.

Note — Divided into two geographically separated taxa, mainly based on the number
of lobes of the perianth. It is noteworthy that this character, which can be used for the
division of the genus into sections (see p. 55), occurs here within a group of species with
mainly 3-lobed perianths.

KEY TO THE SUBSPECIES

la. Perianths predominantly 3-lobed. — Sumatra, Peninsular Malaysia...........--
teehee de sc beh tate Py ates Yet ae Nie i meee ae BK MT MRI SMO Xo: eo thiSs a. subsp. sucosa
b. Perianths predominantly 2-lobed. — Borneo ........-----+--- b. subsp. bifissa

a. subsp. sucosa

Horsfieldia sucosa (King) Warb., Mon. Myrist. (1897) 322; J. Sinclair, Gard. Bull. Sing. 16 (1958)
416, p.p. (incl. lectotype, excl. f. 45 = Horsfieldia sparsa), pl. XU-A; 28 (1975) 139, p.p.;
W.J. de Wilde, Gard. Bull. Sing. 38, 2 ((1985’, 1986) 188; Tree Fl. Sabah & Sarawak 3 (2000)
393. — Myristica sucosa King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 301, pl. 172.

Horsfieldia bracteosa Henderson, Gard. Bull. Str. Settl. 7, 2 (1933) 120, pl. 30; J. Sinclair, Gard.
Bull. Sing. 16 (1958) 419, f. 46. — Horsfieldia bracteosa Henderson var. bracteosa: J. Sinclair,
Gard. Bull. Sing. 28 (1975) 18. — Type: SF 24521 (Henderson), Peninsular Malaysia.

Perianths predominantly 3-lobed, rarely a few 2- or 4-lobed; mature male buds 1.2—
1.5 by 1.5—2 mm; androecium 0.4—0.6 by (.8—1.2 mm; thecae 14-18 (Peninsular Ma-
laysia) or 18-22 (Sumatra). Fruits 2.5-3.5 by 22.5 cm with persistent 3-lobed perianth.

Field-notes — Bark smooth, shallowly fissured, or thin-scaly; slash bark laminated,
reddish, with sticky reddish exudate; wood yellowish or pink. Flowers yellow(-green),
scentless. Fruits glossy green, turning yellow or pink.

Distribution — Malesia: Sumatra, Peninsular Malaysia.

Habitat & Ecology — Primary or disturbed dry land and seasonal swamp forest; sandy
soils or sandstone; 0—500 m altitude; fl. & fr. throughout the year.

Oo

De Wilde — Myristicaceae (Horsfieldia) 20

Notes — 1. Dry fruits are at most 3.5 cm long, but fresh fruits 5—-7.5 cm long, with a
thick pericarp.

2. Horsfieldia sucosa is vegetatively very similar to H. pallidicaula and H. sterilis,
the latter two being markedly different in the male flowers; they have a different androe-
cium, and the pedicels are not articulated.

3. It seems that the distributional area excludes that of the related and resembling
species H. pallidicaula.

b. subsp. bifissa W.J. de Wilde

Horsfieldia sucosa (King) Warb. subsp. bifissa W. J. de Wilde, Gard. Bull. Sing. 38, 2 1985’, 1986)
190; Tree Fl. Sabah & Sarawak 3 (2000) 393. — Type: Soegeng 58, Kutei.

Perianths predominantly 2-lobed, the odd 3-lobed perianth present; mature male buds
1.2-1.4 by 1.5-1.7 mm; androecium 0.3—0.4 by 0.8—0.9 mm; thecae 14. Fruits 2—2.5
(4) by 2 cm, the 2-lobed perianth + caducous.

Field-notes — Bark smooth, undulately fissured; inner bark 10 mm thick, laminated,
reddish or yellow brown; wood whitish, pale brown, or reddish. Flowers yellow, the
males very fragrant.

Distribution — Malesia: Borneo (Sarawak, E Sabah, C, E & S Kalimantan).

Habitat & Ecology — Lowland forest; on sandy loam soil, sandy ridge in Shorea
laevifolia forest, also in forest over limestone; 0—800 m altitude; fl. & fr. throughout the
year.

Note — Fruiting material may be difficult to separate from that of H. pallidicaula var.
macrocarya in which the perianth is 3- or 4-lobed.

86. Horsfieldia superba (Hook. f. & Thomson) Warb.

Horsfieldia superba (Hook. f. & Thomson) Warb., Mon. Myrist. (1897) 295; Corner, Wayside Trees
(1940 & 1952) 476; J. Sinclair, Gard. Bull. Sing. 16 (1958) 393, f. 36, pl. X-B; 28 (1975) 141;
W.J. de Wilde, Gard. Bull. Sing. 38 (1985’, 1986) 200. — Myristica superba Hook. f. & Thom-
son, Fl. Ind. (1855) 162; A.DC., Prodr. 14, 1 (1856) 194; Migq., Fl. Ind. Bat. 1, 2 (1858) 62;
Hook. f., Fl. Brit. India 5 (1886) 105; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 298, pl. 124bis,
125bis. — Type: Phillips s.n., Peninsular Malaysia.

Tree 10-30 m. Twigs sometimes faintly angular, 5-8 mm diameter, late glabrescent,
at first with dense, rusty, dendroid hairs 0.5—1 mm long, bark dark grey, striate, some-
times slightly cracking and flaking; lenticels usually many and conspicuous. Leaves
usually in 2, sometimes in 3 rows (see note 2), coriaceous, (elliptic-)oblong, (17—)25-—
40(—70) by (7.5—)10—18(—22) cm, base narrowly subcordate to short-attenuate, apex
+ blunt to acute-acuminate; upper surface drying dull olivaceous-brown with finely
wrinkled structure, glabrescent (except sometimes midrib), lower surface with (bright)
brown rather sparse to dense mealy indumentum of dendroid hairs of mixed size, (0.3—)
0.5—1 mm long, usually with many or few emerged hairs to 1.5 mm long; dots absent;
midrib rather broad, flat above, pubescent or glabrescent; nerves 15—25(—30) pairs, flat
or sunken above, lines of interarching fairly regular, not very distinct; venation faint on
both surfaces; petiole 6-15 by 5—7 mm, late glabrescent; leaf bud 20—30 by 5-10 mm,
with dense hairs 0.5—1.5 mm long. Inflorescences behind the leaves, with + dense, woolly,

204 Flora Malesiana, Ser. I, Vol. 14 (2000)

long-branched yellowish brown dendroid hairs 0.5—1 mm long; in male: rather many-
flowered, 2 or 3 times branched, 7-15 by 2.5-10 cm, peduncle 1—1.5 cm; in female:
+ few-flowered, somewhat branched, 2—5 cm long; bracts broadly ellipsoid, subacute-
acuminate, up to 12 by 10 mm, densely pale brownish pubescent, caducous; flowers in
male up to 5 in acluster, glabrous, perianth 3- or 4-lobed, drying often with a grey bluish
tinge, pedicel not articulated. Male flowers: pedicel slender, 1.5—3.5(—5) mm; buds el-
lipsoid to obovoid-ellipsoid, 6—7(—8) by 4-5 mm, base rounded to short-attenuate, cleft
1/5—1/4. lobes 0.4 mm thick; androecium elongate-ellipsoid, subcylindrical to triangular
in cross section, (4—)4.5—5 by 2.2—2.5 mm, apex subtruncate, with a shallow 3-radiate
crack. base subtruncate (Plate 2: 48); thecae 32-40, completely sessile; androphore rather
narrow, 0.1—0.4 mm long. Female flowers: pedicel stout, 2-3.5 mm long; buds ellip-
soid, 7-8 by 4.5 mm, cleft c. 1/5, lobes rather coriaceous; ovary ovoid, slightly com-
pressed, 4—4.5 by 3 mm, glabrous, stigma shallowly 2-lobed, broad, 0.5 by 1.5 mm.
Fruits 1-3 per infructescence, broadly ovoid-ellipsoid, 3.8-5.5 by 2.8—4.5 cm, glabrous,
+ coarsely warty and wrinkled, pericarp 8—12 mm thick; fruiting pedicel stout, 3-6 mm
long; perianth persistent.

Field-notes — Bole straight; bark longitudinally fissured, dippled or cracked; bark
slash brittle, gritty; slash wood soft, whitish or yellowish. Leaves glossy above, becom-
ing dull on drying. Flowers bright yellow, smell unpleasant or of ripe pears, visited by
bees. Fruits globose, (greenish) yellow, or orange.

Distribution — Malesia: Sumatra (Central, West), Peninsular Malaysia (almost all
states), Singapore.

Habitat & Ecology — Forest on alluvial soils, undulating country, also in swampy
forest: 0-400 m altitude; fl. Aug.—Oct.; most fruits collected May—Aug.

Notes — 1. Related to H. fulva on account of the oblong or obovoid male perianth
and oblong staminal column.

2. By way of exception the leaves, also in fertile twigs, are arranged in three rows.

3. Sterile H. superba may be confused with Gymnacranthera bancana, also with a
stout habit and remaining indumentum on the twig apex and the lower leaf surface, but
its hairs are much more interwoven forming a thin felty mat; in H. superba the lower leaf
surface is covered with harsh, stellate-dendroid hairs.

87. Horsfieldia sylvestris (Houtt.) Warb.

Horsfieldia sylvestris (Houtt.) Warb., Mon. Myrist. (1897) 337, t. 22; J. Sinclair, Gard. Bull. Sing.
28 (1975) 142: W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 84, f. 11. — Myristica sylvestris
Houtt.. Nat. Hist. Pl. 2, 3 (1774) 340. — Type: not indicated; see discussion by J. Sinclair, l.c.:
147, 148.

Myristica salicifolia Willd. in Roem. & Usteri, Mag. Bot. 3, 9 (1790) 26; Sp. Pl. 4 (1806) 871;
Roxb., Fl. Ind. 3 (1832) 846. — Type: not known, see above.

Myristica pinnaeformis Zipp. (msc.) ex Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 49. — Type:
Zippelius s.n. (180), West New Guinea.

Myristica pendulina Hook. f., Fl. Brit. India 5 (1890) 859; King, Ann. Roy. Bot. Gard. Calc. 3
(1891) 329, pl. 170. — Type: Cantley s.n., culta.

Horsfieldia sylvestris (Houtt.) Warb.var. villosa Warb., Mon. Myrist. (1897) 341. — Syntypes: Beccart
696, Warburg 20708, New Guinea.

Myristica edulis F. Muell. in sched. (Hb. von Miiller, d’Albertis 11, MEL, not seen).

De Wilde — Myristicaceae (Horsfieldia) 205

Tree 7—40(—60) m. Twigs hollow, in innovations when dry + angular, flattish, or ridged,
4-14 mm diameter, glabrescent, at first with rusty, + woolly hairs 0.3—1(—1.5) mm; bark
faintly striate, not flaking; lenticels coarse. Leaves (thinly) chartaceous, lanceolate(-lin-
ear), (17—)20—45 by 3—7(—9) cm, base rounded to short-attenuate, apex long acute-acu-
minate; upper surface drying dull, greenish brown to dark brown, minutely pustulate or
not, lower surface late glabrescent or with (partially) persistent hairs of mixed size, 0.1—
1(—1.5) mm; dots absent; midrib + flat above, late glabrescent; nerves 30—42 pairs, thin,
flat or sunken above, lines of interarching distinct; venation lax, distinct (leaves fre-
quently + bullate) or indistinct; petiole (0O—)2—7 by (2—)3—5 mm, usually shortly winged
by decurrent lamina; leaf bud up to 8 cm long, densely woolly-pubescent. Inflorescences
with woolly hairs 0.5—1 mm, or late glabrescent; in male: large, paniculate, many-flow-
ered, 3—5 times branched, 7-30 by 4-14 cm; female 4—10(—15) cm long; peduncle 2—7
cm, at base with a few persistent rather blunt cataphylls 2-4 mm long; bracts + con-
cave, (2—)4-8(—16) mm long, rather late caducous; flowers in male in loose clusters of
4—10, in female up to 5; perianth 2- (or 3-)lobed, often somewhat angular, glabrous or at
base glabrescent, pedicel slender, glabrescent or with hairs 0.3 mm, not articulated; flowers
before anthesis, especially in male, densely packed into subglobose or ellipsoid glomerules
4-7 mm diameter wrapped in bracts. Male flowers: pedicel 0.2—2 by 0.3 mm; buds
(narrow-)obovoid or clavate, irregularly shaped and angular by being closely packed,
1.5—2.1 by 0.5-1.3(—1.5) mm, apex obliquely obtuse, towards base + tapering into pedi-
cel, cleft 1/3—1/2, lobes 0.1—-0.3 mm thick, sometimes with a few pale dots; androecium
ellipsoid-oblong, 1-1.2 by 0.5—0.6 mm, apex broadly rounded (Plate 1: /4); thecae
8-16, completely sessile, 0.8—1.5 mm long, column solid; androphore rather broad, 0.1—
0.4 mm long. Female flowers: pedicel stout, 1.5—5.5 mm long; buds ellipsoid to broadly
ovoid, + coriaceous, 3.5—-5 by 3-4.5 mm, cleft c. 1/3; ovary broadly ovoid-ellipsoid,
2.5—3 by 2.5 mm, glabrous, stigma sessile, unlobed, 0.1 by 0.5 mm. Fruits 2-10 per in-
fructescence, ellipsoid, base and apex sometimes + acute on drying, 3.4—5.5 by 2.5—-3.5
cm, glabrous, without or with few coarse tubercles; pericarp 2—4(—5) mm thick; fruiting
pedicel 5—13 mm; perianth not persistent. — Fig. 30.

Field-notes — Striking solitary emergent tree, with pendulous branches (twigs up to
2 m), bole often very straight, in old specimens with rotten core; buttresses present or
absent, low or up to 1.5(—2.5) m high, up to 1.5 m out, up to 8(—20) cm thick, sometimes
with small stilt-roots; bark brownish, smooth or usually shallowly fissured, or slightly to
strongly peeling off in small scales; exudate pale red-brown, watery; sapwood pale yel-
lowish or straw, usually gradually passing into the slightly darker reddish heartwood.
Leaves drooping and distichous (twigs resembling compound pinnate leaves), glossy
above. Flowers bright or dark yellow, slightly fragrant or not; pollen pale yellow or
whitish. Fruits pinkish, orange, red-brown, or deep red.

Distribution — Malesia: Moluccas (Morotai to Kai Is.), Aru Is., New Guinea (not
known from Morobe and Milne Bay Prov. of Papua New Guinea).

Habitat & Ecology — Primary, degraded, and secondary forest, on alluvial soils (sandy
and clayey), especially common in the coastal plains of Bird’s Head; also in swampy
forest (with Pometia), in forest inundated by heavy rains or in stagnant water; also on
well-drained porous volcanic soils, or close to limestone outcrops, in ridge forest, or in
Castanopsis forest (at 530 m, in Bird’s Head); 0-700 m altitude; fl. & fr. throughout the
year. The trees may bear flowers and fruits simultaneously.

206 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Horsfieldia) 207

Uses — The wood is heavy, easily worked, not very durable. Fruits (sour taste) edible
(Sepik Prov.). The fruit wall is used in rodjak, and in manisan (a sweet pickle) (Moluccas).
Extract of bark is used as a drug against ‘penyakit keputihan’ by pregnant women, also
against hepatitis (Moluccas). The fruits are gathered and eaten by the Gogodala tribe
(Papua New Guinea, Western Prov.); also planted near villages. Fruits eaten by birds
(e.g., pigeons, parrots), apparently swallowed whole. The tree is recorded as beautiful,
and recommended as an ornamental. The many vernacular names indicate that the tree is
widely known by local people.

Notes — 1. Horsfieldia sylvestris is a homogeneous species, and only varies in hairi-
ness. Very hairy specimens were described as var. villosa by Warburg.

Sterile specimens may be confused with H. hellwigii, a species usually with narrower
leaves, subglobose (not clavate) male buds, and smaller and hairy fruits; in H. sylvestris
the fruits generally are larger, and always glabrous.

2. Warburg placed H. sylvestris and H. ralunensis in his section Orthanthera, which
also included H. iryaghedhi from Sri Lanka, on account of the elongate angular male
buds clustered into flower heads. The first two species, however, have the flowers clus-
tered only in immature inflorescences, and are not closely related to H. iryaghedhi.

88. Horsfieldia talaudensis W.J. de Wilde

Horsfieldia talaudensis W.J. de Wilde, Gard. Bull. Sing. 38, 1 (1985) 75. — Type: Lam 2628, Moluc-
cas, Talaud I.

Tree 15-35 m. Twigs 2—4(-7) mm diameter, early glabrescent, at first with scarce
hairs 0.1 mm or less; bark striate, not flaking; lenticels + inconspicuous. Leaves mem-
branous or chartaceous, oblong-lanceolate, 8-30 by 2.5—-10 cm, base attenuate, apex
acute-acuminate, finely pale-pustulate on both surfaces; upper surface drying greenish
to brown, lower surface bright brown, early glabrescent, hairs stellate-scaly, 0.1 mm
long or less; dots absent; midrib flat above; nerves 12—20 pairs, flat and inconspicuous
above, lines of interarching not distinct; venation thin, indistinct on both surfaces; peti-
ole 10-18 by 1.5—3 mm, leaf bud 10 by 2 mm, with hairs 0.1 mm. Jnflorescences with
dense hairs 0.1-0.2 mm long, about 3 times branched; in male and female 4-8 by 3-4
cm, rather many-flowered, peduncle 1.5—2 cm; bracts broadly ovoid-ellipsoid, pubes-
cent, 1.5—3 mm long, caducous; flowers in male in clusters of 2 or 3, perianth 2-lobed,
glabrous; pedicel pubescent, not articulated. Male flowers: pedicel slender, 1—1.5 mm,
with grey or pale brown hairs 0.1 mm; buds transversely ellipsoid, somewhat compressed
laterally, 2-2.2 by 2.5-3 mm, drying brown, not collapsing on drying, cleft 2/3—4/5,
lobes firm, 0.2—0.3 mm thick; androecium transversely ellipsoid to reniform, not much

Fig. 30. Horsfieldia sylvestris (Houtt.) Warb. a. Leafy twig apex; b & c. twigs with immature and
full-grown male inflorescences, respectively; note bracts in b; d. male flower; e. ditto, opened,
showing androecium; f. twig with female inflorescence axillary to fallen leaf; g. opened female
flower, lateral view, showing glabrous ovary; note much larger size as compared with the male
flowers; h. twig with infructescence, fruits mature [a, f, g: de Vogel 3069; b: de Vogel 3094; c—e:
Craven 739; h: de Vogel 3370]. — Scale bar for a—c, f, h = 2 cm; for d, g = 1.65 mm: for e =
0.85 mm.

208 Flora Malesiana, Ser. I, Vol. 14 (2000)

compressed, 1.5 by 2 mm (Plate 1: 9); thecae c. 36, connate for basal half and forming a
+ saucer- or cup-shaped column into which the free apical halves of the anthers are
inflexed; androphore slender, 0.2—0.3 mm long. Female flowers not seen. Fruits 3-10
per infructescence, short-ellipsoid, 1.5—1.6 by 1.3—-1.4 cm, glabrescent, at first with stellate
hairs 0.1 mm or less (hence ovary pubescent), without conspicuous tubercles; pericarp
1.5 mm thick, fruiting pedicel 3-4 mm long; perianth not persistent.

Field-notes — Tree to 35 m. Ripe fruits orange or brownish yellow.

Distribution — Malesia: Moluccas (Talaud Is.: Karakelong), possibly Sulawesi
(Minahassa, see note 2).

Habitat & Ecology — Old forest on mountain slopes; 70-200 m altitude; fl. Apr.; fr.
Apr., May.

Notes — 1. Horsfieldia talaudensis possibly is endemic on the Talaud Islands. It be-
longs to the group of species with firm, transversely ellipsoid to subglobose male buds,
the androecium with strongly inflexed anthers, connate to about halfway, with slender
hairy pedicels, and pubescent ovary (thinly pubescent young fruits).

2. Koorders 18136 (Sulawesi) is sterile but agrees in leaf colour and texture with the
specimens from the Talaud Islands.

89. Horsfieldia tenuifolia (J. Sinclair) W.J. de Wilde

Horsfieldia tenuifolia (J. Sinclair) W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 11, f. 28; Tree Fl.
Sabah & Sarawak 3 (2000) 394. — Horsfieldia polyspherula (Hook. f. emend. King) J. Sinclair
var. tenuifolia J. Sinclair, Gard. Bull. Sing. 28 (1975) 105. — Type: $ 13686 (Bujang), Sarawak.

Tree 5-15 m. Twigs 1-3 mm diameter, early glabrescent, hairs greyish rusty, (0.1—)
0.2 mm: bark striate, neither cracking nor flaking; lenticels inconspicuous or absent.
Leaves membranous or thinly chartaceous, elliptic to oblong-lanceolate, 6.5—16.5 by 3-
6.5 cm, base attenuate, apex acute-acuminate; upper surface drying olivaceous to grey-
ish brown, glabrous, lower surface dull greyish brown, glabrous; dots absent; midrib
raised above, glabrous; nerves 5—1 1 pairs, raised above, lines of interarching regular but
faint; venation lax, faint; petiole 8-16 by 1.5—2 mm; leaf bud 5—8 by 1-1.5 mm, with
dense hairs 0.2(—0.3) mm. Inflorescences with (sparse) stellate hairs 0.1—-0.2 mm long;
in male: rather many-flowered, about 3 times branched, 3-5 by 24 cm, peduncle 0.2-1
cm: in female: rather slender, few-flowered, 2-3.5 by 1 cm; bracts elliptic to oblong,
2-3 mm, pubescent, caducous; flowers in male in loose clusters of 3-8, perianth 3- (or
4-)lobed, glabrous, pedicel glabrescent or with few scattered hairs 0.2 mm in the lower
half, not or indistinctly articulated. Male flowers: pedicel 1-2 mm long, somewhat ta-
pering: buds globose to broadly obovoid, 0.8—1.3 by I-1.5 mm, apex broadly rounded,
base rounded to subattenuate, cleft c. 1/3 to nearly 1/2, lobes 0.2—0.3 mm thick, at base
to 0.4 mm thick; androecium + obovoid or ellipsoid, 0.5—0.7 by 0.5—0.8 mm, in cross
section triquetrous (Plate 3: 68); thecae 8-12, suberect, 0.3-0.4 mm long, free for at
least halfway, column largely hollow, the base continued into the somewhat tapering
androphore 0.2—0.3 mm long. Female flowers: pedicel 1.5 mm long; buds ellipsoid, 2 by
1.5 mm, cleft c. 1/3 or slightly over; ovary ellipsoid, 1.3 by 0.8—0.9 mm, glabrous, stigma
2-lobed, 0.2 mm high. Fruits 1-3 per infructescence, ellipsoid, base sometimes shortly
tapered, 1.7—2 by 1.4-1.5 cm, glabrous, without lenticel-like tubercles; pericarp 1—1.5
mm thick; fruiting pedicel 3-5 mm long; perianth not persistent. — Fig. Si.

De Wilde — Myristicaceae (Horsfieldia)

g ti

Fig. 31. Horsfieldia tenuifolia (J. Sinclair) W.J. de Wilde. a. Habit of twig with leaves and male
inflorescences; b. ditto, with female inflorescence; c. mature male flower, lateral view; d. ditto,
opened, showing androecium; e. mature female flower; f. ditto, opened, showing glabrous ovary
and minute 2-lobed stigma; g. twig with infructescence, fruits mature [a, c, d: S 24945; b, e, f:
S 34528; g: S 24914]. — Scale bar for a, b, g = 2 cm; for c—f = 0.85 mm.

210 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Slender tree, once recorded with buttresses; bark (greenish) brown,
narrowly fissured, not flaking; inner bark dark red, with red sap; sapwood whitish. Flowers
greenish yellow. Fruits yellowish.

Distribution — Malesia: Borneo (Sarawak: Ist Div.; Sabah: Beaufort Hill, Jesselton;
Brunei, E Kalimantan).

Habitat & Ecology — Understorey tree of lowland dipterocarp forest on richer soils;
on yellow (sandy) clay or loam, or brownish soil; on ridges and slopes; 0-300 m alti-
tude; fl. & fr. May—Sept.

Note — Horsfieldia tenuifolia is characterized by twigs which are glabrous to the
apex, small and slender leaf buds, with hairs only (0.2 mm long, by leaves membranous
to thinly coriaceous, not very brittle, drying to a greyish tinge, and the not markedly
different colour of upper and lower surfaces, relatively long and slender petioles, and by
comparatively small inflorescences and flowers. According to the male flowers the spe-
cies is close to H. polyspherula. Another closely related species is H. macilenta, also
with thin membranous leaves, but with much more pubescent twigs and inflorescences.

90. Horsfieldia tomentosa Warb.

Horsfieldia tomentosa Warb., Mon. Myrist. (1897) 302; J. Sinclair, Gard. Bull. Sing. 16 (1958) 403,
f. 40: 28 (1975) 149; W.J. de Wilde, Gard. Bull. Sing. 38, 2 ((1985’, 1986) 210. — Myristica
tomentosa Hook. f. & Thomson, FI. Ind. (1855) 161, nom. illeg. [non Thunb. (1782)]; A.DC.,
Prodr. 14, 1 (1856) 204: Miq., FI. Ind. Bat. 1, 2 (1858) 68; Hook. f., Fl. Brit. India 5 (1886) 105;
King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 301, pl. 129. — Type: Wallich Cat. n. 9025 (‘Myristi-
cea?’), Peninsular Malaysia.

Tree 5—20(—40) m. Twigs 2-5 mm diameter, late glabrescent, hairs dense, rusty, woolly,
1-1.5 mm: bark striate, not flaking; lenticels abundant, conspicuous. Leaves membra-
nous, elliptic or obovate to oblong-lanceolate, 9-27 by 4-10(-1 2) cm, base nearly rounded
to attenuate, apex acute-acuminate; upper surface drying brown, faintly minutely pustu-
late or not, glabrous (glabrescent), lower surface with persistent dense dendroid hairs all
of about the same size, (0.1—)0.5—0.8 mm long; dots absent; midrib + flat, glabrescent
above; nerves 7-15 pairs, flat or sunken above, lines of interarching + regular, distinct;
venation lax, faint; petiole 10-18 by 1.5—3 mm, pubescent; leaf bud ovoid-ellipsoid, 10—
15 by 2-3 mm, with dense hairs 1.5 mm. Inflorescences with dense woolly hairs 1.5—2
mm long; in male: rather many-flowered, 3 or 4 times branched, 3-12 by 1.5—7 cm,
peduncle 0.3—2.3 cm; in female: + few-flowered, 2-6 cm long; bracts elliptic, densely
woolly pubescent, 2-4 mm long, caducous; flowers (male) in small fascicles, perianth
3(—4- or 5-)lobed, glabrous; pedicel slender, in male glabrous, in female late glabrescent,
not articulated, hairs 0.2 mm. Male flowers: pedicel 1-3 mm long; buds globose, 1.4—
2.5 mm diameter, cleft nearly 1/2, lobes 0.1 mm thick; androecium much depressed-
globose, above flattish or impressed, circular to blunt-triangular in cross section, 0.6—
0.9 by 1.2-1.7 mm (Plate 2: 54); thecae 18—24(—30?), almost completely sessile, incurved
towards the apex; column broad, with apical hollow 0.2 mm deep; androphore narrow,
0.4-0.5 mm long. Female flowers: pedicel 1-2 mm long; buds broadly ellipsoid, 2.5 by
2.3 mm, cleft c. 1/3; ovary subglobose, 1.5 mm diameter, with appressed hairs 0.1—0.2
mm, style and stigma minute, faintly 2-lobed, 0.1 mm long. Fruits 1—5 per infructescence,
ellipsoid, 1.5—2 by 1.3-1.6 cm, glabrescent, usually with minute indumentum remaining

De Wilde — Myristicaceae (Horsfieldia) 211

towards the base, usually minutely pustulate; pericarp 1.5 mm thick; fruiting pedicel
2—6 mm long; perianth not (or for a short period) persistent.

Field-notes — Bark fissured, not flaky nor scaly, brown to blackish, soft; inner bark
pale reddish or pale yellowish, laminated, fibrous; exudate pink-red; sapwood whitish or
pink. Flowers yellow, with a fine perfume. Fruits yellow to orange.

Distribution — S Peninsular Thailand; in Malesia: Sumatra (E Coast, see Sinclair
1975: 149: no specimens seen), Peninsular Malaysia (Kedah, Penang, Perak, Trengganu,
Pahang, Selangor, Malacca, Johore, Kelantan), Singapore (doubtful; Sinclair 1975: 150).

Habitat & Ecology — Lowland and foothill forest, old secondary forest; 0-300 m
altitude; fl. mostly Mar.; fr. mostly July.

Notes — 1. The lobes of the male perianth in fully mature flowers are often somewhat
recurved outward.

2. Horsfieldia tomentosa belongs in the alliance with H. flocculosa, H. grandis, and
H. motleyi, all with a hairy lower leaf surface.

91. Horsfieldia triandra W.J. de Wilde

Horsfieldia triandra W.J. de Wilde, Gard. Bull. Sing. 38,2 (*1985’, 1986) 195, f. 23.—Type: Forbes
2465, Sumatra.

Low tree. Twigs sometimes faintly ridged, 1.5—3(—4) mm diameter, early glabrescent,
hairs rusty, 0.3—0.7 mm long; bark finely, lower down coarsely striate, not flaking; lenticels
small but conspicuous. Leaves membranous to thinly chartaceous, (elliptic-)oblong, 5—9
by 2-3.5 cm, base (short-)attenuate, apex acute-acuminate with acumen 8—12 mm long;
upper surface glabrous, drying dark olivaceous to dark brown, lower surface brown,
early glabrescent; dots absent; midrib raised above, glabrous; nerves 6—10 pairs, flat or
+ sunken above; venation lax, faint on both surfaces; petiole 7-13 by 1-1.5 mm; leaf
bud 6-8 by 2 mm, with dense rusty hairs 0.3-0.7 mm long. Inflorescences with dense
rusty hairs 0.3—-0.4 mm; in male: | or 2 times branched, 3-5 by 1-2 cm, peduncle 0.3—1
cm long, not many-flowered, (10-20 flowers per inflorescence and in different stages of
development), solitary or in loose clusters of 2-4; in female (from infructescences): 0.5—
1 cm long, few-flowered; bracts ellipsoid-oblong, pubescent, 2-3 mm long, caducous,
and each inflorescence usually with one (rarely two) persistent subapical enlarged bract,
resembling a small foliage leaf, 5—12 mm long (see note 1); flowers with 3-lobed peri-
anth, towards the base with fine hairs 0.2—0.4 mm, pedicel pubescent, not articulated.
Male flowers: pedicel 3-4 mm long, slender; buds + obconical-obovoid, apex broadly
rounded or blunt, base tapering, 2.5 by 2.5 mm, cleft 1/5—1/6, not collapsing on drying,
lobes at apex 0.2, towards base 0.4 mm thick, + clasping the anthers, at anthesis hardly
opening, the basal part of perianth thick-walled, 0.8—1 mm thick; androecium including
androphore + turbinate, 1.5—1.7 by 0.6—-0.8 mm, circular in cross section (Plate 2: 45);
thecae 6, acutish, erect, subsessile, 0.6—-0.7 mm long, the apical 0.3 mm free; column
solid, below proceeding into + obconical androphore, | by 0.6—0.8 mm. Female flowers
not seen. Fruits | or 2 per infructescence, broadly ellipsoid, 23—25 by 19-20 mm, drying
blackish, glabrescent with remnants of minute indumentum of scattered hairs 0.1 mm or
less towards the base; pericarp 1—1.5 mm thick; perianth not persistent. — Fig. 32.

Distribution — Malesia: C & S Sumatra.

Habitat & Ecology — Forest at 800—1000 m altitude.

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Ae tay Yi
SA Ue

Fig. 32. Horsfieldia triandra W. J. de Wilde. a. Habit of leafy twig with male inflorescences; note
leaf-like bracts: b. mature male flower; c. ditto, longitudinally opened, thick-walled perianth, and
androecium; d. androecium, longitudinal section, schematic [all Forbes 2465]. — Scale bar for a =
2 cm; for b—d = 1.7 mm.

Notes — 1. The male bud deviates from most Horsfieldias by the obconical shape
and the thick-leathery texture; it opens slightly only at the very apex, and the turbinate
androecium, with at apex only 3 anthers, is clasped by the perianth lobes before anthesis.

The flowers look as if diseased, but on opening one finds the perianth and androecium
normal. The few-flowered inflorescences, with some leaf-like enlarged bracts in the
apical part, also look aberrant. Enlarged leaf-like bracts in the inflorescences are occa-
sionally found in the inflorescences of some other species, e.g., in H. irya.

De Wilde — Myristicaceae (Horsfieldia) 213

2. Horsfieldia sterilis from Borneo, H. pulverulenta and H. crux-melitensis (to a lesser
extent), both from New Guinea, and related species from New Guinea have somewhat
resembling male flowers, with thick-leathery perianth, similarly opening only at the
apex, and a reduced number of anthers; however, all these species have a 2-lobed peri-
anth.

92. Horsfieldia tristis W.J. de Wilde

Horsfieldia tristis W.J. de Wilde, Gard. Bull. Sing. 38, 2 (1985°, 1986) 197; Tree Fl. Sabah &
Sarawak 3 (2000) 396. — Type: S 37470 (Lai Shak Teck), Sarawak.

Tree 10-15 m. Twigs 3—5(—7) mm diameter, early glabrescent, with greyish to rusty
hairs 0.1 mm long or less; bark bright brown to yellowish, coarsely striate with a ten-
dency to cracking longitudinally, or flaking; lenticels + smal, conspicuous or not. Leaves
membranous to thinly chartaceous, elliptic-oblong to lanceolate, 14-32 by 4—8.5 cm,
base attenuate, apex acute-acuminate; above drying dull with a finely wrinkled or granu-
late structure, olivaceous-brown, lower surface glabrous, almost concolorous; dots ab-
sent; midrib flat or slightly raised, glabrous above; nerves 1 1—17 pairs, sunken, flat, or
slightly raised above, lines of interarching + invisible; venation + lax, + invisible; peti-
ole 6-12 by 2-5 mm; leaf bud 10-15 by 1.5—2.5 mm, with dense greyish to dull-brown
hairs up to 0.1 mm long. Inflorescences glabrous or with few scattered hairs up to 0.1
mm; in male: about 3 times branched, rather many-flowered, 5—12 by 3.5—7 cm, pedun-
cle 0.7—2.5 cm long; in female (in fruit): 2-3 cm long, once or twice branched; bracts
not seen, caducous; flowers in male in loose clusters of 2—5, glabrous, perianth 3-
(or 4-)lobed, pedicel not articulated. Male flowers: pedicel 1.5—5 mm long, slender, well
marked off from the perianth; buds obovoid to broadly ellipsoid, 2.8-3.7 by 2-3 mm,
base rounded to short-tapering, cleft 1/5—1/4 (to nearly 1/3), lobes 0.2—0.4 mm thick,
androecium obovoid-ellipsoid, apex broadly rounded or + depressed, in cross section
subcircular or bluntly 3- or 4-angular, 2—2.7 by 1.5—2 mm (Plate 2: 46); thecae 24—40,
mutually appressed, almost completely sessile with free apices 0.1(—0.2) mm long, col-
umn broad and solid with a broad apical hollow, 0.5—0.8 mm deep, concealed by the
overcurved anthers; androphore narrow, 0.2—0.3 mm long, largely hidden by the anther
bases. Female flowers not seen. Fruits 4-8 per infructescence, ellipsoid, apex subacute,
1.5 by 1.2 cm, glabrous, drying blackish, without lenticels or tubercles; pericarp 1.5 mm
thick; fruiting pedicel 1-2 mm long; perianth not persistent.

Field-notes — Flowers yellowish, fragrant.

Distribution — Malesia: Sumatra (E Coast of Tapanuli), Lingga Arch. (Singkep L.),
Borneo (Sarawak, S Kalimantan).

Habitat & Ecology — Forest on flat land; 0-100 m altitude; fl. Aug., Nov.; fr. Sept.

Notes — 1. Horsfieldia tristis seems related to H. fulva, with similarly elongate male
flowers; in H. fulva the leaves dry brown instead of olivaceous, the twigs brown (not
pale brown), bark not cracking, inflorescences pubescent, and pedicels articulated. Dried
specimens of H. tristis have yellowish twigs and pale, olivaceous leaves; twigs, leaves
and inflorescences are almost completely glabrous.

2. Biinnemeijer 7100 (Singkep I.) differs in its thin leaves, not distinctly dull and
wrinkled on upper surface, and in the broadly ellipsoid, almost globose, male buds; its
androecium has a very broad apical cavity with a broad, almost flat base.

Flora Malesiana, Ser. I, Vol. 14 (2000)

214

Nn

De Wilde — Myristicaceae (Horsfieldia) 21

93. Horsfieldia tuberculata (K. Schum.) Warb.

Horsfieldia tuberculata (K. Schum.) Warb., Mon. Myrist. (1897) 279, t. 23. — Myristica tuberculata
K. Schum. in K. Schum. & Hollr., Fl. Kaiser Wilhelmsland (1899) 46. — Syntypes: Hollrung
848 (K, lecto), Kaiser Wilhelmsland; Kdrnbach s. n., lost, Bat I.

Tree 5—20 m. Twigs faintly ridged or not, 2—4(—6) mm diameter, early glabrescent,
hairs (rarely + woolly) 0.1—0.3 mm; bark striate, not flaking; lenticels sparse and small,
sometimes almost absent. Leaves membranous (or on higher altitudes subchartaceous),
elliptic to oblong-lanceolate, 12—25(—40) by 3—10(—16) cm, base short- to long-attenu-
ate, rarely rounded, apex acute-acuminate; upper surface drying dull (greenish) brown,
often minutely pustulate, lower surface early glabrescent, hairs 0.1(—0.3) mm; dots ab-
sent; midrib flat or + sunken above; nerves | 1—22 pairs, above thin, flattish or sunken
above, lines of interarching not very distinct; venation fine, distinct or not on both sur-
faces; petiole 8-18 by 1.5—3(—5) mm; leaf bud 10-15 by 2—3 mm, with hairs 0.1—0.3
mm long. /nflorescences with sparse stellate-dendroid hairs 0.1—0.3 mm long, some-
times + glabrescent; in male |—3 times branched, 3—15(—22) by 2-10 cm, peduncle 0.5—
2.5 cm long; female to 7 cm long; bracts 0.5—3 mm long, caducous; flowers solitary or in
loose clusters of 2—4, perianths 2- (or 3-)lobed, glabrous (early glabrescent), pedicels
glabrous or rarely sparingly pubescent, sometimes + articulated. Male flowers: pedicel
(somewhat) tapering, 2—5(—6) mm; buds laterally compressed, short-pear-shaped, about
as long as broad to slightly broader than long, (1.5—)2—3.5 by 2-4 mm, apex broadly
rounded, the lower 1/3—1/2 narrowed into the pedicel, cleft 1/2—3/4, lobes 0.2 mm thick;
androecium + flattened, apex broadly rounded to subtruncate, 1.5—2.5 by 2-3 mm (Plate
2: 33); thecae 24—40, erect, not inflexed, free apical parts up to 0.2 mm long; column
narrowly hollowed for 1/5—1/3; androphore up to 0.2 mm long. Female flowers: pedicel
1.5-2.5 mm long; buds subglobose, 2—3 by 2—3.5 mm, cleft 1/2—2/3; ovary subglobose,
glabrous, 1.5—2 by 1.5—2 mm; stigma sessile, faintly 2-lobed, 0.2 by 0.5—1 mm. Fruits
5-15 per infructescence, ellipsoid, apex rounded or slightly pointed, 1.5—3.7 by 1.1—
2.5(—3) cm, glabrous, drying blackish brown, with sparse, small to coarse, paler lenticel-
like tubercles; pericarp 1-8 mm thick; fruiting pedicel 3—10 mm; perianth not persistent.
— Fig. 33.

Distribution — Solomon & Caroline Islands; in Malesia: Papua New Guinea. Two
varieties.

KEY TO THE VARIETIES

la. Fruits 1.5—2.5 by 1-1.5 cm; dry pericarp 1—2(—3) mm thick .. a. var. tuberculata
b. Fruits 2.7—3.7 by 1.7—2.5 cm; dry pericarp 3—8 mm thick .... b. var. crassivalva

Fig. 33. Horsfieldia tuberculata (K.Schum.) Warb. var. tuberculata. a. Habit of leafy twig with in-
fructescence; b. twig with male inflorescences; c. mature male flower, lateral view; d. ditto, opened,
showing androecium; e. twig with female inflorescence; f & g. opened female flower and glabrous
ovary with shallowly 2-lobed stigma; h. twig with infructescence and mature fruits [a: BS7P 14035;
b—d: Waterhouse 820-B; e—g: BSIP 9628; h: BSIP 10611]. — Scale bar for a, b, e, h = 2 cm; for
c, d, f = 1.65 mm; for g = 0.85 mm.

216 Flora Malesiana, Ser. I, Vol. 14 (2000)

a. var. tuberculata

Horsfieldia tuberculata (K.Schum.) Warb., Mon. Myrist. (1897) 279, t. 23; K.Schum., Notizbl.
Bot. Gart. Berlin 2 (1898) 117; K. Schum. & Lauterb., Fl. Schutzgeb. Siidsee (1900) 324; Markgr.,
Bot. Jahrb. Syst. 67 (1935) 151, p.p.; A.C. Sm., J. Arnold Arbor. 22 (1941) 62; W.J. de Wilde,
Gard. Bull. Sing. 38, 1 (1985) 126, f. 19. — Myristica tuberculata K. Schum. in K. Schum. &
Hollr., Fl. Kaiser Wilhelmsland (1899) 46; Warb., Bot. Jahrb. Syst. 13 (1891) 308.

Horsfieldia novo-guineensis Warb. var. moseleyana Warb., Mon. Myrist. (1897) 273; K.Schum. &
Lauterb., Fl. Schutzgeb. Siidsee (1900) 324. — Type: Moseley s.n., Admiralty Is.

Horsfieldia solomonensis A.C. Sm., J. Arnold Arbor. 22 (1941) 64. — Type: Kajewski 1549, Solo-
mon Is.

Twigs 2—4(—6) mm diameter. Leaf blades 12—25(—40) by 3-10(—16) cm. Male flow-
ers: pedicel 2-5(-6) mm; buds (1.5—)2—3(-5) by 2-4 mm; androecium + flattened, 1.5—
2.5 by 2-3 mm, thecae 28-40, free apices up to 0.2 mm; androphore up to 0.2 mm long.
Fruits 1.5-2.5 by 1-1.5 cm; dry pericarp 1—2(—3) mm thick.

Field-notes — Exudate of bark red, watery; slash wood white or brownish white, soft;
slash bark soft, pale brown or reddish brown. Flowers yellow, sweet scented. Fruits yel-
low, or orange(-brown).

Distribution — Solomon and possibly Caroline Islands (Palau I.); in Malesia: Papua
New Guinea (Admiralty Is., Bismarck Arch., Papuan Is., Cape Vogel Peninsula).

Habitat & Ecology — Primary, degraded, and secondary forest, low mossy montane
forest: on coral rock, seashores, limestone, swamp forest; 0-700 m altitude; fl. & fr.
throughout the year.

Notes — 1. The size of flowers and fruits is variable, especially on the Solomon Is-
lands.

2. In some collections (e.g. from Cape Vogel Peninsula) the pedicels of the male
flowers are slender, and only slightly tapering; such specimens may easily key out wrongly.
They probably represent a separate taxon as yet insufficiently defined. In general, the
male buds of H. tuberculata approach those of H. laevigata.

b. var. crassivalva W.J. de Wilde

Horsfieldia tuberculata (K. Schum.) Warb. var. crassivalva W.J. de Wilde, Gard. Bull. Sing. 38, 1
(1985) 130. — Type: Brass 28352, Louisiade Arch.

Twigs (3—)4—5 mm diameter. Leaf blades 25-35 by 10—12.5 cm. Male flowers: pedi-
cel 3-6 mm; buds 3 by 3.3 mm; androecium much flattened, 1.8 by 1.8 mm, thecae c. 24,
free apices 0.1 mm long; androphore 0.1 mm long. Fruits 2.7-3.7 by 1.7—2.5(-3) cm;
pericarp 3—8 mm thick.

Field-notes — Subcanopy tree. Flowers yellow, very fragrant. Fruits to 5 cm diam-
eter, orange, ovoid or subglobose, keeled; aril pink.

Distribution — Solomon Islands (San Cristobal, doubtfull); in Malesia: Papua New
Guinea (Louisiade Arch.: Misima, Tagula, and Rossel I.).

Habitat & Ecology — Riverine rain forest at low altitudes, creek alluvial soil; 0-20 m
altitude.

Notes — 1. Specimens of var. crassivalva generally have a stout habit with coarse
twigs and large leaves, and relatively large perianths; these sizes fall, however, within
those accepted for the type variety.

De Wilde — Myristicaceae (Horsfieldia) PING

2. Specimens in fruit may be confused with H. pachycarpa, which has minutely pu-
bescent fruits (and pubescent ovaries), at least at the base towards the insertion of the
stipe.

94. Horsfieldia urceolata W.J. de Wilde

Horsfieldia urceolata W.J. de Wilde, Blumea 32 (1987) 462. — Type: LAE 77053 (Gideon), Papua
New Guinea.

Shrub or treelet, 1-5 m. Twigs 1.5 mm diameter, with dark rusty-brown hairs (0.2—)
0.3 mm, glabrescent; bark finely striate, neither cracked nor flaking; without or with few
inconspicuous lenticels. Leaves membranous, oblong to lanceolate, 6-18 by 1.5—7 cm,
base attenuate to nearly rounded, apex acute-acuminate; blade above glabrous, drying
dark brown above, beneath somewhat paler, glabrescent with some scattered scaly-stellate
hairs remaining for some time; dots absent; midrib very slender, raised above; nerves
12-15 pairs, sunken above, lines of interarching + distinct; venation lax, distinct be-
neath; petiole 6-10 by 1-2 mm; leaf bud 7 by 1.5 mm, with dense rusty woolly hairs
0.2-0.3 mm. Inflorescences (male) among the leaves, with dense rusty hairs (0.2—)0.3
mm, small, twice branched, the lowest branch 2—3 mm from the base, rather few-flow-
ered; in male: 2—3 by 2 cm, with few branches; in female (from infructescences, see
note 2): few-flowered, 2.5 cm long; bracts oblong-lanceolate, 1.5—2 mm, pubescent, ca-
ducous; flowers solitary or 2 or 3 together, perianth 2-lobed, at base with hairs 0.2—0.3
mm (the remainder of the perianth early glabrescent), drying dark brown or blackish,
pedicel pubescent, slender, not articulated. Male flowers: pedicel 1.5—3 mm long; buds
subglobose or broadly ovoid, 2.5—3 mm diameter, cleft 1/6(—1/8), lobes (perianth) 1—1.5
mm, at apex 0.2 mm thick; androecium club-shaped, 1.5—2 by 0.6—0.7 mm, the anther-
bearing apex + acute (Plate 1: /6); thecae 8, sessile, 0.3—0.4 mm long, not or hardly
projecting, column solid; androphore glabrous, with scattered tannin-dots, not or but in-
distinctly warty. Female flowers not seen. Fruits (doubtful, see note 2) 2 or 3 per
infructescence, drying dark brown to blackish, subglobose (to broadly ellipsoid), in-
cluding the minute, slender, bifid, | mm long rostrum (persistent style and stigmas) 1 cm
long, pseudostalk 1.5—2 mm; all thinly pubescent, hairs 0.1—0.2 mm; pericarp 0.5 mm
thick; fruiting pedicel slender, 7-8 mm long; perianth not persistent.

Field-notes — Shrub or low treelet. Leaves dull green above, pale or somewhat olive
below, with olive-brown midvein. Flowers yellow. Fruits red, brilliant scarlet inside.

Distribution — Malesia: Papua New Guinea (Milne Bay Prov., and, possibly, North-
em Prov.); restricted to a limited area of distribution.

Habitat & Ecology — Primary rain forest on sloping ground; Lithocarpus, Anisoptera,
Hopea-dominated forest on ridges and slopes; 25—120 m altitude; fl. Mar., Nov.; fr. May.

Notes — 1. Part of the material of H. urceolata had been included formerly in
H. squamulosa. Horsfieldia urceolata differs from that species and from the related
H. coryandra in the subglobose thick-walled male perianth.

2. The fruiting specimen NGF 31751 (Ridsdale) differs markedly from the type in
foliage, with the blades rather conspicuously tapering in the lower half, less distinct and
less projecting tertiary venation, and rather more persistent indumentum on the lower
surface. Possibly it represents a separate undescribed taxon, but more material is needed
to decide on this.

218 Flora Malesiana, Ser. I, Vol. 14 (2000)

3. Fruiting specimens of H. urceolata, and related species (H. clavata-group), may
resemble H. schlechteri and H. subtilis, but the latter two have glabrous ovaries and
fruits.

95. Horsfieldia valida (Miq.) Warb.

Horsfieldia valida (Miq.) Warb., Mon. Myrist. (1897) 349; K. Heyne, Nutt. PI. Ned. Indié (1927)
638: W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986) 25. — Myristica valida Migq., Fl. Ind. Bat. 1,
2 (1858) 67; Suppl. 1 (1860) 156. — Type: Teijsmann 479 (sterile, the fruits said to be as large as
a goose egg), Sumatra, West Coast.

Tree 10-15 m. Twigs 2.5—5 mm diameter, at insertion of inflorescences up to 10 mm
diameter, (grey-)brown, glabrescent, hairs rusty, woolly, 0.4—0.7 mm long; bark coarsely
striate, not flaking, lenticels + dense, conspicuous, pustulate. Leaves chartaceous,
(ob)ovate-oblong, 20-35 by 8-13 cm, base short- to long-attenuate, apex subobtuse to
acutish; upper surface glabrous, drying dark olivaceous-brown, lower surface bright
brown, early glabrescent but midrib late glabrescent; dots absent; midrib broad towards
the base, slightly raised above; nerves 20-25 pairs, raised, lines of interarching indis-
tinct: venation rather lax, flat, faint on both surfaces; petiole 7-12 by 3—4.5 mm; leaf
bud 15-20 by 4 mm, with dense ferruginous hairs (0.40.7 mm. Inflorescences just be-
hind the leaves, with woolly hairs 0.3-0.5 mm, subglabrescent; in male: 2 or 3 times
branched, not many-flowered, 5-6 by 3—4 cm, peduncle 1-1 5 cm long; in female (from
infructescences): 5 cm long; bracts not seen, caducous; flowers in male in loose clusters
of 3-6, glabrous, perianth 4- (or 3-)lobed, pedicel not articulated. Male flowers: pedicel
1.5 mm long; buds globose, 2.5—3 by 3-3.5 mm, cleft c. 4/5, slightly collapsing on
drying, lobes 0.3—0.4 mm thick; androecium + depressed globose, 0.8-1.2 by 1.4-1.6
mm (much smaller than the perianth), in cross section + rounded or faintly 4-angular
(Plate 3: 75); thecae 24—28, curved and largely sessile, free apices 0.2 mm; column not
or hardly hollowed; androphore narrow, 0.1—0.2 mm long. Female flowers not seen (but
see under fruits). Fruits 2-9 per infructescence, ellipsoid, 8(—9) by 5(—6) cm, glabrous,
drying brown with surface wrinkled and + warted; pericarp 15 mm thick; fruiting pedi-
cel 5 mm long; perianth persistent under young fruits in Maradjo 449, 4-lobed, 3 mm
long.

Field-notes — Erect tree, branches wide spreading and arching. Flowers brown, tinged
yellow, sweet smelling.

Distribution — Malesia: Sumatra (E & W Coast; possibly Palembang, see note I);
probably W Borneo (see note 2).

Habitat & Ecology — Primary forest, ravine forest; (2200 m, see note 1) 900-1100 m
altitude; fl. Mar., Aug.; fr. Aug.

Notes — 1. The sterile specimen Dumas 1649 (Palembang, 200 m alt.) deviates from
the other specimens (all from 900-1100 m alt.) by the shorter indumentum on the leaf
bud (hairs only 0.3 mm long).

2. Hallier 624 (with male fl., Mt Damoes, W Kalimantan) 1s obviously close to H.
valida, but markedly differs, and apparently represents a new, yet undescribed species.
The specimen is discussed further by De Wilde, l.c. Female flowering specimens which
match Hallier 624 have not been found as yet, but Afriastini (& Burley) 579 (in fruit,
from Kalimantan) is possibly identical; it has the perianth not persistent under the fruits.

De Wilde — Myristicaceae (Horsfieldia) 219

96. Horsfieldia wallichii (Hook. f. & Thomson) Warb.

Horsfieldia wallichii (Hook. f. & Thomson) Warb., Mon. Myrist. (1897) 305; J. Sinclair, Gard. Bull.
Sing. 16 (1958) 405, f. 41, pl. XI-A; 28 (1975) 156, p.p. (excl. part of the Borneo material =
Horsfieldia borneensis); W.J. de Wilde, Gard. Bull. Sing. 38, 2 (*1985’, 1986) 204; Tree FI.
Sabah & Sarawak 3 (2000) 396. — Myristica wallichii Hook. f. & Thomson, FI. Ind. (1855) 161,
p.p. (Wall., Cat. n. 6806 being a mixture, see note by Sinclair, 1975: 158); A.DC., Prodr. 14, 1
(1856) 230; Mig., Fl. Ind. Bat. 1, 2 (1858) 67; Hook. f., Fl. Brit. India 5 (1886) 105; King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 303, pl. 132 & 133, excl. syn. Myristica crassifolia Hook. f. &
Thomson. — Myristica horsfieldia auct. non Blume: Wall., Cat. (1832) n. 6806, p.p. — Lectotype:
Griffith s.n., Malacca.

Tree 10—30 m. Twigs sometimes drying somewhat flattened, usually conspicuously
hollow, 3—6 mm diameter, early to late glabrescent, hairs 0.3—0.6 mm; bark dark brown
or blackish, coarsely striate, later on fissured, sometimes flaking, lenticels inconspicu-
ous. Leaves membranous to coriaceous, ovate-oblong to oblong-lanceolate, (14—)19—40
by (4-)4.5—12 cm, base rounded to short-attenuate, apex subobtuse to acute-acuminate;
upper surface drying olivaceous to blackish brown, glabrous (with minute indumentum
sometimes remaining on midrib); lower surface late glabrescent or indumentum locally
persistent, hairs sparse or dense, dendroid, (0.3—)0.5—0.8 mm long; dark dots and/or
dashes present; midrib slightly raised above, late glabrescent; nerves (12—)15—28 pairs,
slender, flat or sunken above, lines of interarching and venation indistinct; petioles
15-35 by 2.5—4.5 mm; leaf bud 20—30 by 4—6 mm, with dense hairs 0.3—0.6 mm. Jnflo-
rescences usually behind the leaves, with sparse or dense stellate-dendroid hairs 0.5—1
mm long; in male: large, many-flowered, 3 or 4 times branched, 10—33 by 6-22 cm,
peduncle 3—7 cm; in female: rather stout, fewer-flowered, 3—7 cm long; bracts broadly
ovate, 3-10 mm long, densely woolly-pubescent, caducous; flowers in male in clusters
of 5-12, perianth 3- (or 4-)lobed, glabrous (in female glabrescent), pedicel with hairs
0.2-0.3 mm long, not articulated. Male flowers: pedicel 0.3—0.6(—1) mm; buds broadly
obovoid, 2—2.5(—3) by 2.4—2.5(—3) mm, apex broadly rounded, base + attenuate, cleft
1/3—2/3, lobes 0.2—0.3 mm thick; androecium broadly obovoid to subglobose, apex
+ depressed with a 3-radiate crack, base rounded to attenuate, faintly triangular in cross
section, |.3—2 by 1.5—2 mm (Plate 2: 50); thecae (24?—)30—46, completely connate, ses-
sile, closely appressed, at apex incurved into apical hollow; column broad with broad
hollow to c. 1/2; androphore narrow, up to 0.3 mm long. Female flowers: pedicel stout,
0.5—1.5 mm long; buds ovoid-ellipsoid, 2.5—4 by 2—3.5 mm, glabrescent, hairs 0.1 mm,
cleft c. 1/3; ovary ovoid to subglobose, 2 by 2—2.5 mm, glabrous, stigma broad, faintly
2-lobed, 0.3 by 0.8 mm. Fruits 2-9 per infructescence, ovoid-ellipsoid, 4-7 by 3—4.5
cm, glabrous, smooth or wrinkled, not or only faintly warted; pericarp 10-15 mm thick;
fruiting pedicel stout, 4-6 mm long; perianth generally persistent. — Photo 8.

Field-notes — Bole straight; crown dense; bark shallowly or deeply longitudinally
fissured, dark grey, not flaking; bark | cm thick, slash underbark bright to deep red, pink,
or reddish brown; slash wood whitish, pale or yellowish, light brown, or (red-)brown.
Flower buds blue-green or yellow at anthesis. Fruits glaucous, maturing (green-)yellow,
orange(-yellow), or red.

Distribution — Malesia: Sumatra (Aceh, Tapanuli, W & E Coast, Indragiri, Palembang,
Simeulué, Morsala, Mentawai I., Bangka), Peninsular Malaysia (all provinces except
Perlis and Negri Sembilan), Singapore, Borneo (Sarawak, Brunei, Sabah, Kalimantan).

220 Flora Malesiana, Ser. I, Vol. 14 (2000)

Habitat & Ecology — Lowland and ridge-top forest; on red soil, granitic sand, loam
soil with coral limestone; 0—470 m altitude; fl. & fr. throughout the year.

Note — Horsfieldia wallichii is vegetatively recognizable by the blackish dots and
stripes (dashes) on the lower leaf surface, and the hollow twigs. Similar dots are only
found in a few other species, including H. borneensis, with similar fruits, but differing in
general habit, and in the male flowers.

97. Horsfieldia xanthina Airy Shaw

Horsfieldia xanthina Airy Shaw, Kew Bull. 1939, n. 10 (1940) 541 (441); W.J. de Wilde, Gard.
Bull. Sing. 39, 1 (1986) 45; Blumea 41 (1996) 381; Tree FI. Sabah & Sarawak 3 (2000) 397. —
Type: Richards 1927, Sarawak, Mt Dulit.

Tree 10-30 m. Twigs sometimes faintly angular at apex, 2.5—-6 mm diameter, early
glabrescent, hairs 0.1 mm; bark coarsely striate, with a tendency to flake; lenticels con-
spicuous or not. Leaves coriaceous, elliptic to oblong-lanceolate, 8-35 by 3.5-13 cm,
base attenuate to rounded, apex subacute to acute-acuminate; upper surface drying oliva-
ceous to (dark) brown, usually with rather distinct hair scars, lower surface early glabres-
cent, drying with a reddish tinge; dots absent; midrib slightly raised above; nerves 8-20
pairs, slender, flat or slightly raised, lines of interarching faint; venation lax, faint on
both surfaces; petiole 7-12 by 2.5—3 mm; leaf bud 8-13 by 2-3.5 mm, with hairs 0.1
(—0.2) mm long. Inflorescences behind the leaves, with + dense hairs 0.1 mm; in male:
sometimes short, robust, (2—)4—20 cm long, (2 or) 3 times branched, not many-flowered,
peduncle 0.5—0.8 or 1.5—2 cm (subsp. macrophylla) long; in female (from infructescences):
1-1.5 or 5—7 cm (subsp. macrophylla) long, few-flowered; bracts not seen, caducous;
flowers in male in loose clusters of 3—5(—8), perianth 3- or 4-lobed, glabrous, pedicel
glabrescent or with few hairs 0.1 mm towards the base, not articulated. Male flowers:
pedicel thickish, somewhat tapering or not, straight or + curved (flowers reflexed),
12.5 mm long; buds broadly ellipsoid to ovoid-subglobose, 2.5—2.8 mm long, apex
rounded, base shortly rounded and somewhat tapering into the pedicel, cleft 1/2(—2/3),
slightly wrinkled but not collapsing on drying, lobes (.4—0.8(—1) mm thick; androecium
+ laterally flattened, + broadly obovoid 1—1.3 mm long (Plate 3: 86); thecae 6—16, erect,
0.9-1.1 mm long, largely sessile with free apices 0.1—0.3 mm; column broad, with api-
cal hollow 0.1—0.3 mm; androphore + tapering, broad, (0.1—)0.2—0.3 mm long. Female
inflorescences and flowers known only in subsp. macrophylla: ovary glabrous. Fruits
(1-)2-6 per infructescence, ellipsoid-obovoid or broadly ovoid, (3.5—)5 by 3—3.5 cm,
glabrous; pericarp 5(—10) mm thick, + woody towards inside; fruiting pedicel stout,
3 mm long; perianth not persistent.

Distribution — Malesia: Borneo (Sarawak, Sabah); two subspecies, both montane.

KEY TO THE SUBSPECIES

la. Twigs towards the apex 2.5—4.5 mm diam. Leaf blades 8-18 by 3.5-7 cm. Male
inflorescences 2—5 cm long, the flowers often with + reflexed pedicel; thecae 6-12
berets! ners: eet 2k a ad ek sg Pe eee a. subsp. xanthina
b. Twigs stouter, towards apex 3.5—6 mm diam. Leaf blades 22-35 by 7-13 cm. Male
inflorescences 10-20 cm, flowers erect; thecae 14 or 16... b. subsp. macrophylla

N
Nw
—

De Wilde — Myristicaceae (Horsfieldia)

a. subsp. xanthina

Tree 10-17 m. Twigs 2.5—4.5 mm diameter; bark with a tendency to flaking. Leaf
blades 8-18 by 3.5—7 cm. Male inflorescences (2—)4—5 by 2—3.5 cm, the rachis at base
2-3 mm diameter; flowers often + reflexed. Male flowers: pedicel 2—2.5 mm long; buds

laterally compressed, 1—1.2 by 0.8-1 by 0.4-0.5 mm, thecae 6-12. Female flowers not
seen.

Field-notes — Flowers yellow. Fruits (glaucous) green, glossy, maturing reddish or-
ange.

Distribution — Malesia: Borneo (Sarawak).

Habitat & Ecology — Montane, kerangas, heath forest, submontane forest; on (yel-
low) sandy soils, sandstone, or “on higher flanks of limestone mountain among huge
limestone boulders with vegetation and organic layer entwined between boulders”;
800-1800 m altitude; fl. Aug., Sept.; fr. Sept., Nov.

Note — Subspecies xanthina is characterized by the usually flaking bark of the twigs,
coriaceous and often + reddish brown tinged leaves, and coriaceous rather large male
flowers with a typical androecium of only 6—12 thecae and a marked androphore. Possi-
bly it is confined to kerangas at higher altitudes. It is close to H. majuscula, from Penin-
sular Malaysia and Sumatra, which differs by non-flaking bark of twigs, membranous
leaves, 14-18 thecae, pedicel articulated at base, and possibly by larger fruits.

b. subsp. macrophylla W.J. de Wilde

Horsfieldia xanthina Airy Shaw subsp. macrophylla W.J. de Wilde, Gard. Bull. Sing. 39, 1 (1986)
47; Blumea 41 (1996) 381; Tree Fl. Sabah & Sarawak 3 (2000) 398. — Type: Clemens 50050,
Mt Kinabalu.

Tree 25-30 m. Twigs 3.5—6 mm diameter; bark not flaking. Leaf blades 25-35 by
7-13 cm. Male inflorescences 10—20 by 7-11 cm, the rachis at base 3.5—4.5(—5) mm
diameter: flowers erect. Male flowers: pedicel 1—-1.5 mm long; buds broadly ellipsoid,
2.6-2.8 by 2.5-2.7 mm; lobes 3, cleft nearly halfway; androecium 1.3 by 0.8 mm,
subtriquetrous in cross section; thecae 14 or 16. Female inflorescences (0.5—)1—8 cm
long, once (or twice) branched. Female flowers: pedicel 1.5—2 mm, glabrous or with
some minute hairs towards the base; buds ovoid-ellipsoid, 4—4.5 by 3—3.5 mm, lobes 3,
cleft to nearly halfway, lobes 0.8(—1) mm thick; ovary ovoid, 2.2 by 2 mm, glabrous,
stigma broadly 2-lipped, 0.3 by 1 mm.

Field-notes — Poorly developed low buttresses (once); bark brown and grey, fissures
boat-shaped; exudate light red, watery. Flowers yellow or orange; ovary pale purple.
Young fruits green, when mature orange.

Distribution — Malesia: Borneo (E Sarawak; Sabah: Mt Kinabalu).

Habitat & Ecology — Mountain forest, ridge forest; on igneous derived (andesitic)
soils; 1100-1550 m altitude; fl. July, Nov.; fr. Nov.

Note — May be confused with H. subalpina subsp. kinabaluensis which has a differ-
ent androecium and generally smaller fruits.

222 Flora Malesiana, Ser. I, Vol. 14 (2000)

KNEMA

Knema Lour., Fl. Cochinch. (1790) 604; Warb., Mon. Myrist. (1897) 132, 543; J. Sinclair, Gard.
Bull. Sing. 16 (1958) 254; 18 (1961) 102; W.J. de Wilde, Blumea 25 (1979) 321; 27 (1981) 223;
32 (1987) 115: Tree Fl. Sabah & Sarawak 3 (2000) 399. — Myristica sect. Knema Blume, Rumphia
1 (1836) 187; A. DC., Prodr. 14, 1 (1856) 204; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 284.
—Type species: Knema corticosa Lour. [= Knema globularia (Lam.) Warb.}.

Trees. dioecious: stilt-roots sometimes present. Twigs without lenticels, flaky or not.
Leaves not breaking easily when dry (veins with sclerenchym sheaths), lower surface
pale, not papillose, dots present or absent; reticulation dense, distinct. Inflorescences
rarely supra-axillary; a sessile or to 5(-10) mm long pedunculate, simple or 2- (or 4-)fid,
wart-like or worm-like densely scar-covered brachyblast, 1-17 mm long, with a slow
unlimited growth; basal cataphylls absent; bracts minute, caducous; flowers in apical
subumbels (few-flowered in female). Flowers pedicellate, not fragrant, bracteole minute,
often at a distance below the perianth, usually persistent. Male flowers: perianth + ro-
tate, (thinly) carnose, inside glabrous or pubescent (rare), greenish creamy, pink or red;
buds (sub) globose or (ob)ovoid, cleft to c. 1/2 to nearly to the base, lobes 3(—5), spread-
ing at anthesis; androecium stalked, androphore cylindrical or tapering, rarely with few
hairs, androecium (or staminal disc) flat or convex (or mammillate) with at the margin
3-25 stellately attached ellipsoid anthers, (half-)sessile or shortly stiped (see also Fig.
34). Female flowers: buds (ob)ovoid or ellipsoid, ovary pubescent, style short or absent,
stigma + 2-lobed and each lobe again 2-many-lobulate. /nfructescences sessile, with
one or few fruits. Fruits + ellipsoid, 2-5 cm long, tomentose or early glabrescent (never
glabrous as in some species of Horsfieldia and Myristica); pericarp leathery; aril entire
or laciniate at the apex only. Seeds ellipsoid, not variegated; albumen ruminate with a
fixed oil and starch; cotyledons divaricate or suberect, scarcely or only very slightly
connate at the base. — Fig. 34-58.

Distribution — About 93 species in continental Southeast Asia and Malesia, from
the Deccan Peninsula (K. attenuata) to S China (Yunnan) and East to the Philippines,
Moluccas, and western New Guinea (K. tomentella in Bird’s Head Peninsula); in Malesia
75 species, the majority occurring in Borneo. — Map 4 (see p. 4).

Fig. 34. Androecia of Knema. — a. K. woodii J. Sinclair; b. K. curtisii (King) Warb. var. amoena
J. Sinclair; c. K. latifolia Warb.; d. K. elmeri Merr.; e. K. membranifolia H.J.P. Winkler [a: Sinclair
et al. 9298: b: Sinclair & Kadim bin Tassim 10442; c: Kostermans 9121; d: Sinclair et al. 9311;
e: Kostermans 4382]. — Scale bar for all = 1 mm.

De Wilde — Myristicaceae (Knema) 223

Habitat & Ecology — Usually under- and middle-storey trees of primary, often ever-
wet forest, in continental SE Asia also in seasonal forest; up to 1800(—2000) m altitude.

Notes — 1. De Wilde (1979) distinguished in the genus Knema 12 series for 83 spe-
cies accepted at that time. This division into series has been based mainly on characters
regarding the general shape of male buds, androecium and position of the anthers, and
the shape of the stigma lobes, with in addition characters derived from the indumentum
and the nature of the bark of the twigs. The series, which were fully described and dis-
cussed, at most represent possibly natural groupings of species, presumably close to
each other. However, more ample recent material and newly described species rendered
the formal distinction of the series more and more doubtful, and so, in the present Flora
Malesiana treatment, the author refrained from any formal subdivision. The series are
here indicated only in the general key to the species (Key 1), from which the main set of
characters can be inferred, and for details the reader is referred to the literature men-
tioned above. Sinclair (1958, 1961) proposed a subdivision into groups mainly based on
female flower characters. This subdivision is for the most part not in accordance with
the series distinguished by De Wilde and cannot be used satisfactorily either.

2. Besides a general key to the species (1), mainly based on male flowering speci-
mens, four separate regional keys are given, based on female flowering and/or fruiting
specimens and with emphasis on vegetative characters. These keys cover the following
areas: (2) Peninsular Malaysia, Singapore, (3) Sumatra, Java, (4) Borneo, (5) Philippines,
Sulawesi, Moluccas, Lesser Sunda Islands, and New Guinea (Bird’s Head).

(1) GENERAL KEY TO THE SERIES AND SPECIES OF KNEMA
(mainly based on male flowering specimens)

la. Buds longer than broad (in general resembling female buds: obconical, pear-shaped,
ovoid, obovoid, or ellipsoid-oblong); base rounded to attenuate, cleft 1/2—2/3;
androphore about as long as or usually longer than the diameter of the staminal disc
including anthers (these characters sometimes less obvious in K. andamanica). An-
thers 6-12, generally + erect, sometimes almost horizontal. SERIES LAURINAE .. 2

b. Buds £as broad as long, or broader than long (globose, pear-shaped, obovoid, rarely
ellipsoid), at base rounded, truncate, saccate, or + attenuate, cleft c. 2/3 to much
deeper; androphore shorter than diameter of staminal disc including anthers. An-
thers 3—25, suberect to horizontal. (In K. piriformis and K. pulchra the buds some-
times short pear-shaped and cleft 1/2—2/3, androphore comparatively long.) ... 6
2a. Lower leaf surface either early glabrescent, or with persistent sessile or mixed ses-
sile and stalked hairs, those on flowers 0.1—0.3 mm long (in K. laurina var. heteropilis
0.5—1.5 mm long). Anthers half-sessile to stiped, suberect to almost horizontal. Pedi-

eon =) ik —1 Oumimione dae ho Ae Le et RR ee ees atta ad oy -nseere ree 3

b. Lower leaf surface with conspicuous persistent indumentum of evenly spaced, more
or less equally long stalked dendroid hairs, those on flowers 0.5—1.5 mm long. An-
thers completely sessile, (sub)erect. Pedicel 2-5 mm long. — Sumatra, Peninsular
Realaysian lana, BOMmEeo 45.2 028) 9 Re 2. RR hee 29. K. laurina

3a. Twigs + flattened. Pedicel 1-3 mm long. Leaves faintly reticulate, finely pitted above
(visible with a lens!). Fruit apex acute or beaked. — Peninsular Malaysia........
MEE SRR STN es oS i Be Ly a oy eae a gy toe ys ae 44. K. oblongifolia

224

b.

4a.

a.

6a.

7a.

8a.

9a.

10a.

Flora Malesiana, Ser. I, Vol. 14 (2000)

Twigs (sub)terete. Pedicel 0.5—10 mm. Leaves faintly or distinctly reticulate above,
not finely pitted. Fruit apex blunt or subacute .............-- 002 seen eee ~
Inflorescences (partly) 1-5 mm pedunculate. Twigs 1.5—3 mm diam., with minute
hairs, glabrescent. Leaves inconspicuously tomentose beneath, glabrescent. —
N Sumatra, N Peninsular Malaysia ..................2.- 2. K. andamanica

_ Male inflorescences sessile or rarely up to | mm pedunculate. Twigs (2—)3—5 mm

diam., usually short-tomentose, rarely glabrescent. Leaves with conspicuous (sub)-
persistent indumentum beneath ..........---- 1 eee eee eee een eee 5
Hairs on twig and lower leaf surface 0.1—0.5 mm long, mainly stellate; those on
flowers 0.2 mm long. Pedicel 4-9 mm long. — Sumatra, Peninsular Malaysia . . .
ee re eet nn see citcr emen ao emareiege 52. K. pseudolaurina

. Hairs on twig and lower leaf surface 0.5—1 mm long, mainly stellate-dendroid;

those on flowers 1—-1.5 mm long. Pedicel 0.5—2.5 mm long. — Sumatra, Peninsular
Wialanstasdava oe te cca oe ees get oie tes 29b. K. laurina var. heteropilis
Lower leaf surface with persistent very short and dense indumentum, silky to the
touch, of densely interwoven scale-like hairs which are only visible with a lens.
SERIES SERICBAE 5. sobs cc fs aso oc 5+ 650 oh s cane «a> «oh A

. Lower leaf surface either glabrescent, or hairs longer, or sparse .........-. 10

Twigs 4—6 mm diam.; bark not longitudinally cracking nor flaking. Leaves 25-
GU CHU LONG on eae sek os tay cisks so yer yan ia ae i 8

. Twigs 2-4 mm diam.; bark cracking, lower down flaking. Leaves 12-32 cm long.

([Indumentum on lower leaf surface grey or pale brown. Staminal disc convex or
mammillate= anthers 10—14.|—= Bomeo™. 22)... J) oe oes 9. K. elmeri
Buds globose, at base rounded, with dense scale-like hairs 0.1 mm long. Staminal
disc convex to mammillate. [Anthers 11-15. Indumentum on lower leaf surface
pale cimmamon. | —— BORIC ng. 0. esse = earner che he Sonnet an gh 63. K. sericea

. Buds broadly obovate, at base short-attenuate. Staminal disc flat ........... 9

Leaf apex obtuse or retuse, base rounded or cordate. Indumentum on lower leaf
surface greyish. Buds short-tomentose. Anthers c. 15. — Peninsular Malaysia....
paree sed taad ete New ¢ sssmp ed A Scape Mire fe) AE: ELS 2 Ce 2 eae 56. K. retusa

. Leaf apex acuminate, base attenuate, obtuse, rounded, or rarely subcordate. Indu-

mentum on lower leaf surface cinnamon, rarely greyish. Buds with scale-like hairs
less than 0.1 mm long, seemingly glabrous. Anthers 22-24. — Borneo.........
OR nt): Oe Ue SO A Me ae es 3b. K. ashtonii var. cinnamomea
Bark of twigs at most coarsely striate, only occasionally cracking (bark of older
wood sometimes finely flaking in K. galeata, K. korthalsii, K. mandaharan, K.
membranifolia, K. pedicellata, ?K. uliginosa) .... 1... +600 eee reer eee es 22

. Bark of twigs soon longitudinally cracked, lower down flaking ........... il

Twigs (1-)1.5—4(—6) mm diam.; older bark + cracking or thinly flaking. Leaves
8—30(—40) cm long, membranous to chartaceous. Buds depressed globose or de-
pressed short-obovoid; staminal disc (flat or) convex or low-mammillate ... 20

. Twigs (3—)4—12 mm diam.; older bark cracking and flaking. Leaves 15-60 cm

long, chartaceous to coriaceous. Buds of variable shapes, if only 3—3.5 mm wide
then ellipsoid. Staminal disc + concave or flat. [Pedicel up to 20 mm long.] SERIES
PNET AR TARE os oe ticbolievefoupcetens radios nine ew be Gioe Be 6 oe ele 12

De Wilde — Myristicaceae (Knema) 225

12a.

15a.

16a.

ia

Leaf blades, petioles, and twigs glabrescent, at first with a very conspicuous lanose
indumentum consisting of hairs 3—8 mm long; hairs on flowers 1-3 mm long 13

. Leaf blades, petioles, and twigs glabrescent, at first with hairs 0.2—2.5 mm long;

Pameowerssp tolimm long 02. ieee A ee es ee. 0 RE 14

. Twigs 5 mm diam. or more; hairs to 8 mm. — Sumatra, Peninsular Malaysia... .

ePererne £9 it a) ertdpoes cytes 8 gageagt cpa peta on mate (eter gn 18. K. hookeriana

. Twigs less than 5 mm diam.; hairs 3-4 mm. — Borneo ..... 31. K. longepilosa
. Leaf base (sub)cordate, rarely rounded; basal nerves + crowded. Twigs early coarsely

erackinp and flaking: Pedicel.3=20:mm long Y<2.\eq.cc4 . 205s. ae. 15

. Leaf base attenuate, obtuse, (broadly) rounded, or occasionally cordate; basal nerves

not or but faintly crowded. Twigs striate, ridged, or cracked, lower down coarsely
Cmunelyiiakine.;Pedicel. 2-4. 5:mmvlong! vests} ck oe O22 g ee dee. 18
Buds 4-5 mm wide, glabrescent or the indumentum easily rubbed off. Pedicel 3—
15 mm long. Anthers 10-14. Disc at base of perianth absent or inconspicuous.
Bark of twigs dark brown or blackish; hairs | mm long. Female pedicel 1.5—2 mm
long; fruits (1.9—)2.5—3.5 cm long, sessile or but shortly pedicelled, hairs up to
2 mm long. — N Sumatra, Peninsular Malaysia, Singapore . . . 11. K. furfuracea

. Buds (5—)6—10 mm wide, either glabrescent or with persistent indumentum. Pedi-

cel 7-20 mm long. Anthers 14—23. Disc at base of perianth present. Bark of twigs
pale brown to blackish; hairs 0.2—2.5 mm long. Female pedicel and fruiting pedicel
8—20 mm long; fruits larger, with hairs up to5 mm long ................. 16
Buds 5—6.5 mm wide, with persistent indumentum. Pedicel 7-9 mm long. Anthers
13 or 14. Disc at base of perianth consisting of separate pads or ridges, or perianth
base thickened and coarsely transversely ridged. Bark of twigs dark brown to black-
ish; hairs 1—2.5 mm long. Female flowers not known; fruits (immature) with hairs
2—3 mm long, fruiting pedicel 8-10 mm long. — C and S Sumatra, Borneo (Anambas
Spee. 2 ATES PSL. MUI. SIGE BIE Sos: ope 26. K. lampongensis

. Buds 6.5—10 mm wide, either glabrescent or with persistent indumentum. Pedicel

12-20 mm long. Anthers (15—)17—23. Disc at base of perianth conspicuous, entire
or interrupted. Bark of twigs pale or dark brown, or blackish; hairs up to 2 mm
long. Female flowers and fruits with pedicel 14-20 mm long; hairs on fruits 1—5
EEAEHMMORE LIISA hes SAI. Ly ode a aith yond Soe sh AUR as BR EARN toe RARE IW
Buds largely glabrescent. Disc at base of perianth coarsely ridged. Bark of twigs
brown to blackish; hairs 1-2 mm long. Fruits velvety-felty with hairs 3-5 mm
fone —— Peninsular Malaysia™... 2650. 2. ee tan, 82 25. K. lamellaria

. Buds with persistent indumentum. Disc at base of perianth conspicuous, + entire or

ridged. Bark of twigs pale brown or greyish brown; hairs 0.2—1(—2) mm long. Hairs
on fruits 1-1.5(—2) mm long. — Borneo .................... 45. K. pallens

. Buds depressed globose or more or less obovoid, with persistent indumentum. An-

Set OS SArEs Le AIRSE bie: siniinss sh. deehreweg Alien dines eaves.t » 19

. Buds + ellipsoid, glabrescent in the upper part. Anthers 6 or 7.— Borneo.......

epee eae TL Eis Deets ray) Geet eel mera als | eel ee 53. K. psilantha

. Buds + obovoid, at base tapering, 3.5—5.5 by 3.5—5.5 mm; anthers curved upwards.

Leaves coriaceous, drying olivaceous above. (A variable species; the flaking of the
bark of the twigs not obvious in part of the material.) — Borneo..............
MEME ERIN, Asoc stb ete Satsw oki havrisencocteniardennvoshcbd is O08 ue wlahals ens 49. K. percoriacea

226

Flora Malesiana, Ser. I, Vol. 14 (2000)

Buds depressed globose or obovoid, 3-7 by 4-6 mm; anthers horizontal. Leaves

chartaceous to coriaceous, drying dark brown above. — Borneo.......--------
Sheth COO Ae ile 1 ek SASSI SOE SSS eA 33. K. lunduensis

_ Buds 5-8 mm diam., early glabrescent; pedicel 6-9 mm long. Twigs early glabres-

cent, often yellowish. — Borneo .......-....-++++5-- 39. K. membranifolia

_ Buds 3-6 mm diam., with persistent indumentum; pedicel (1-)2—10 mm long. Twigs

late’elabrescent, brown. 20)... sf Goes a eee 0 Rae Zl

. Buds globose, 3-4.5 mm diam., pedicel (1-)2-3 mm long, bracteole situated to-

wards the apex. Staminal disc flat or faintly convex; anthers + obtuse, opening
somewhat towards the outside. SERIES GLOMERATAE p.p. — Sumatra, Peninsular
Malaysia, Singapore, Borneo, Philippines .......---+-+-++-+5 27. K. latericia

. Buds depressed globose, 4—6 mm wide; pedicel (4~)5—10 mm long, bracteole about

median. Staminal disc mammillate; the anthers + acute, opening + downwards. —
Romeo’ ar isacd eaten: Ne 2a8d Aa. SES 21b. K. korthalsii subsp. rimosa

_ Buds (sub)globose, or broadly (ob)ovoid, base rounded, truncate, saccate, or short

attenuate: buds cleft to over 2/3. Anthers horizontal, sometimes suberect; androphore
fapeninpior cylindrical gm 2.2 tayalc iy t.\1200S * -RIER. ol. on ad 32

_ Buds obovoid or pear-shaped, in the lower half + narrowed; buds cleft 1/2—3/4 (to

c. 5/6 in K. steenisii). Anthers (half) erect; androphore tapering. SERIES OBOVOI-

DRAB ie buts lend sisaeet wp @elen 2.4%. OLS) See 23
_ Hairs on flowers more than 0.2 mm long ........------ esses eee eer eee 29
- Hairs on flowers less than 0.2 mm long (0.2 mm long in K. rigidifolia) ..... 24

_ Leaves coriaceous. Fruits subsessile. — Peninsular Malaysia........---------

I otra ash See? te ae begin: yisgasy 59b. K. rigidifolia subsp. camerona

_ Leaves chartaceous to thinly coriaceous. Fruits stalked .........--..+++-- Ds)
. Twigs 3—-6(—8) mm diam. Leaf base cordate or broadly rounded. Pedicel 4-8 mm

long; anthers 12-25. — Sumatra’, Peninsular Malaysia, Borneo 55. K. pulchra

. Twigs 1-3 mm diam. Leaf base rounded to attenuate. Pedicel 2-5 mm long . 26
_ Buds cleft c. 5/6; pedicel 2-3 mm long; anthers 4 or 5. — Lesser Sunda Islands

(Blores rani 26 etd 1G. SIE ALE TG. Heth CBI. See Aad nae 64. K. steenisii

_ Buds cleft 1/2-3/4; pedicel 2-5 mm long; anthers 8 or more. — Bomeogaa:. 27
_ Buds 2.5—4 mm long. Anthers 8-15, just stiped. [Lower leaf surface with persistent

sparse hairs 0.1—0.2(—0.3) mm.] — Lowland forest; Borneo . 69. K. subhirtella

_Buds:4=5 mm long: Anthers Wiorl2) 02.1920 0.3). AUR). Ish Ele 28
_ Anthers 11 or 12, just stiped, + horizontal. Lower leaf surface with subpersistent

sparse minute hairs. — Montane forest at (4002—)1000—2000 m; Borneo (Sabah)
BET aT, ih een ee en shee eh sees ress a Ae tbe hs ia 50. K. piriformis

_ Anthers 11, half sessile, suberect. Lower leaf surface early glabrescent, at first with

very weak indumentum. — Lowland forest up to 500 m; Borneo 68. K. stylosa

_ Leaves beneath with persistent coarse stellate and stalked hairs; old leaves par-

tially glabrescent. — Borneo .......-.---- see eee eee eees 43. K. oblongata

_ Leaves beneath either glabrescent, or hairs easily rubbed off, or very inconspicu-

ous, minutes sparse. ciht-7..2 OE eh 0 Se. | RISEN. Sateen eetel tole ote cor aa 30

. Twigs 4-8(—10) mm diam. Leaves chartaceous or coriaceous, 15—55 cm long, base

usually rounded or cordate. Pedicel 2-6 mm long. — Sumatra, Peninsular Malay-
Sirens UP: sc eek nh hha and SOS BEE Ce eee 37. K. mandaharan

De Wilde — Myristicaceae (Knema) 227

b.

31a.

b.

36a.

Twigs 1—3(—4) mm diam. Leaves membranous to thinly coriaceous, 6—30 cm long,

baseattenuate to rounded; Pedicel3=—15 mmplong!) £42 250. Oe eos ee 3]
Leaves + lanceolate, 15—30 by 1.5—4(—5.5) cm. Buds 6—7 mm long; pedicel 10—15
epIONee —ISOMICO 20. . Lk eae ee eee be eae te dom eae ess 61. K. rufa

Leaves oblong to lanceolate, 6—18(—24) cm long [often drying with a blackish me-
tallic lustre]. Buds 3-5 mm long; pedicel 3—11 mm long. — Sumatra, Peninsular
Reanstayy Java (OI ie. £5) UE AS ee eae 15. K. globularia

. Anthers 3 (or 4). Flowers subglabrescent. SERIES LATIFOLIAE. — Sumatra, Borneo

ee Resa telat aheee Gays See eee ee 28. K. latifolia

. Anthers (5 or) 6—25. Flowers with persistent indumentum or glabrescent ... 33
. Anthers (5 or) 6—23(—25), mutually free, i.e., not or at least not completely and not

tightly touching each other; anthers either half sessile or just sessile, or stiped 37

. Anthers 9-25, completely sessile, tightly set, touching, the thecae appearing as if

representing twice as many, i.e., 18—50 sessile anthers; anthers completely hidden
under the rim of the staminal disc. Twigs usually slender, often yellowish. SERIES
a. UIETUISIIATNIAN EOE a rare mapas ey er ie ma Nae ge OMS eet amen rdiie SA- ir 34

. Staminal disc circular, anthers 14 or 15. —-NE Borneo ......... 10. K. emmae
. Staminal disc (blunt-)triangular. — Sumatra, Peninsular Malaysia, Borneo .. 35
. Leaves chartaceous to thinly coriaceous, drying greenish; venation distinct at both

surfaces; base cuneate to rounded, apex acute (subacuminate) to obtuse. Buds
SHoidaneMlan with TOUNGdED angles. cs ike sla ote ae An nei anletae Ohee 36

. Leaves membranous to chartaceous, base usually cuneate to attenuate; when leaves

membranous these drying greenish, with the venation distinct or not, and apex
acute or acuminate; when leaves chartaceous these drying brown, with the vena-
tion very fine or faint, and apex acute, obtuse, or rounded. Buds (sub)triangular
with rounded or sharp angles. — Sumatra, Peninsular Malaysia, Borneo........
<= nnn 5 wo Ce SR ee er ei eee eed MAR RM Rs nan a 8. K. curtisii
Leaves up to 15 by 5 cm, lateral nerves 8—20 pairs. Buds 3.5—6 mm diam., with
short-woolly hairs 0.3—0.4 mm; lobes 0.5—1 mm thick; pedicel slender. — Borneo
cs aco be eR RRs RR ester er she tere ett A nator es ant 30. K. linguiformis

. Leaves 17-30 by 5—10.5 cm, nerves 14—23 pairs. Buds 6 mm diam., hairs 0.1 mm

long, lobes 1-2 mm thick; pedicel stout. — Borneo (Sarawak) ... 74. K. viridis

. Buds generally with persistent indumentum; sometimes seemingly glabrous be-

cause of very minute appressed scale-like hairs only (lens!). Lower leaf surface

PUDORE SCH WOR MOL 5 0 2 6512 jacsptye: © ages na Ny eS Se ee ee eee 4]
sisuds: elabrescent. Lower leaf surface early glabrescent .. 2. 4.0. 14... sae 38
. (Male) inflorescences conspicuously pedunculate. SERIES GLOMERATAE, p. p. — Phil-

ERISA (LAZO) | (open asses By te ee are Geeks Meet 58. K. ridsdaleana
minorescences: (Sub) SESSte sais tec ard ecis Se ote ees Oi oterct ects: Cea Bs 39

. Buds mitriform, with sharp angles, (8—)9-15 mm diam. Twigs 4—10 mm diam.,

grey-brown. Leaves rigidly coriaceous. SERIES GALEATAE. — Borneo...........
eer tres dad) Macao . btn wer. eyo. walt ade odisecons 12. K. galeata

. Buds (depressed-)globose, in cross section circular or obtusely triangular, 2.5—7

(—8) mm diam. Twigs 1.5—3 mm diam., often + yellowish. Leaves chartaceous to
RMAC LAC eOUSUas AMIS AG wt Fi ee ee ae ulee atu aoe pee Dae eee 40

228

“eC

Flora Malesiana, Ser. I, Vol. 14 (2000)

43a.

46a.

47a.

4. Inflorescences on the older branches, below the leaves. Buds 5—7(—8) mm diam.,

glabrescent, at first with short hairs 0.2 mm long. Staminal disc flat to convex, not
with a mammilla. Anthers 19-23. SERIES MEMBRANIFOLIAE. — Borneo..........
CR Not 1 CAG | ereenmrmenne e ies coger brbic as ee eee 39. K. membranifolia

_ Inflorescences among the leaves. Buds 2.5—4.5 mm diam., glabrescent, at first with

woolly hairs 0.5—0.7 mm long. Staminal disc with conspicuous mammilla. Anthers
9-12. SERIES MAMILLATAE p.p. — Borneo (S Kalimantan) ... 36. K. mamillata

a. Leaves on lower surface with scattered brown dots, especially on the finer nerves

(ens!) SSERIES PUNCTATAE® 22.5 oo55.0.005 2. 020s Soe eG ee: oe Se 719

_ Leaves on lower surface without dots (sometimes a few stray dots when leaves

plabrescent)) «sag sp.ths joer ose ME atten rey (PS Ae ey Or 42

_ Buds of variable consistency and size. Staminal disc concave, (+) flat, or but slightly

convex (sometimes low-mammillate in K. glomerata, K. latericia, K. rubens, K.
stenocarpa, and. K. subhirtella)......... 00. 20+. 2+ +2202 ++ 2242 -eee 46

- Buds rather robust, 4-7 mm diam. Staminal disc convex or with a mammilla (in

dry specimens the latter sometimes shrivelled and indistinct). ( Staminal disc some-
times + flat in K. korthalsii from the Philippines). SERIES MAMILLATAE p.p. . 43
Leaves rigidly coriaceous. Twig apex and young petioles with hairs 2—4 mm long;
buds with hairs 1-2 mm long. A ring-shaped disc present at base of perianth around
the androphore. — Peninsular Malaysia ...........--...-- 51. K. plumulosa

_ Leaves chartaceous to coriaceous. Twig apex and young petioles with hairs up to

1 mm long; buds with hairs up to 0.5 mm long. Perianth at base somewhat thicken-

ed or not, without a distinct ring-shaped disc ........-.----+ e+e sees: 44
_ Leaves coriaceous. Mammilla longer than broad. — Sumatra, Peninsular Malay-
sia, Singapore, W Java, Borneo ........--..-----+-+-++:: 19. K. intermedia
_ Leaves chartaceous. Mammilla about as long as broad, or shorter ......... 45

_ Lateral nerves 8-15 pairs, much raised above. Twigs 1.5—3 mm diam., (sub)terete,

smooth or finely striate. Bracteole subapical. Anthers 10-13. — Borneo........
TAYE Rs BRON come eras Ps Mn Be ee ER RC Lo 6 Siaihin. eb "el aya, 8. Og 73. K. uliginosa

_ Lateral nerves 14—40 pairs, not or but slightly raised above. Twigs 3-5 mm diam.,

rather coarsely ridged or + 2- or 3-angled. Bracteole about median. Anthers 10-20.
(Compare also K. glomerata.) — Borneo, S Philippines ..... 21. K. korthalsii
Buds small or medium-sized, rarely large, 1.5—7 mm diam. Lobes 0.3—1 mm thick.
Hairs 0.1—1 mm long. Anthers (5—)6-17. (Compare also K. pectinata from the sec-
enditcadsy) ROME See RUS Siam eee ene onto ee te se ee) 50

. Buds generally robust, (3.5—)4—8 mm diam. Lobes (0.7—)1—2 mm thick. Hairs finely

farinose or scurfy, 0.1(—0.2) mm long or less. Anthers 11-24. (Buds in K. pectinata
3.5-7 mm diam., with 11-15 anthers; in K. scortechinii 3.5—5 mm diam., with 11—
(Gtantherso)! <4 ether sxe: See WR CE See RS RT Cee ee 47
Twigs seemingly glabrous, though actually with minute appressed scale-like hairs,
chocolate, dark brown, or cinnamon. [Twigs 4-6 mm diam., angular or coarsely
striate. Leaves chartaceous to thinly coriaceous; lateral nerves 20-35 pairs. ] SERIES
SERICEAE p.p. — Borneo ..........02- +20 eee: 3a. K. ashtonii var. ashtonii

. Twigs usually early glabrescent; hairs mealy, minute, pale brown to yellowish brown;

indumentum not consisting of appressed scale-like hairs only. SERIES GLOMERATAE
PUMA oo ei ae tye ew a een Rik i mB ite Ie ite le alle nly renee eet etn 48

De Wilde — Myristicaceae (Knema) 229

48a.

49a.

50a.

S5la.

55a:

56a.

Anthers 18-23, completely sessile. Twigs + angular, dark brown or blackish, 2.5—
4 mm diam. Leaves chartaceous or thinly coriaceous; lateral nerves 20-30 pairs.
EMTS. Elsah NAY. dein. tA ee eet (Mens ARIEL AP 75. K. woodii

. Anthers 11-16, half sessile or stiped. Twigs terete, striate, greyish brown, rarely

Piehash WeavesiCOraceoust i .«. sgh ce ete 1 Lk Siem. eee eee ee 49
Leaves to 45 cm long; lateral nerves 25-50 pairs, venation coarse or fine. Buds
(3.5-)4-7 mm diam.; anthers 11—15, half sessile or nearly stiped. Perianth inside
pinkish. Twigs 3-6 mm diam. — Borneo .................. 47. K. pectinata

. Leaves to 30 cm long; lateral nerves 18-30 pairs, venation fine. Buds 3.5—5 mm

diam.; anthers 11—16, usually shortly stiped. Perianth inside greenish to yellowish.
Twigs 2—4 mm diam. — Sumatra, Peninsular Malaysia ..... 62. K. scortechinii
Indumentum of flowers and twigs apices inconspicuous, scurfy or felty, with scale-
like (in K. rigidifolia and K. rubens with stellate-dendroid) hairs 0.1—0.2 mm long
or less. SERIES GLAUCAE (excl. K. rigidifolia, K. tridactyla) .............. 58

. Indumentum of flowers and twigs apices more conspicuous, with farinose or woolly

hairs more than (0.1—)0.2 mm long (hairs sometimes short in K. glomerata from
the Philippines). SERIES GLOMERATAE p.p. .. 2... secs cee ene eee eee Si
Lower leaf surface either 1) completely glabrescent, or with sparse indumentum
persistent near midrib and nerves towards the base of the blade; hairs soft and
inconspicuous, or 2) indumentum persistent, with minute sparse soft stellate hairs.
Inflorescences sessile or pedunculate. Twigs 1-3 mm diam. .............. 54

. Lower leaf surface late glabrescent or with dense persistent indumentum. Inflores-

cences (partly) pedunculate. Twigs 1-2 mm diam. [Buds 1.5—3 mm diam.; anthers
(Sin ERAS eet 2a) sae a ei bie Orgies oale a Bok eA ee 52

. Leaves (oblong-)lanceolate; upper surface with hair scars (lens!). Buds 1.5—2 mm

See eT COGN oe das os a 4 ope ger eahovie ) tg mae oar os Pee eee 72. K. tridactyla

. Leaves ovate to oblong; upper surface without distinct hair scars. Buds 2-3 mm

AOE. a hh, Mey gL foe Gel A ea gale Ra Mme lal tg nS eR RRO Rae S AO sh DL Wet IRN 5 eS 35

. Buds 2—2.5 mm diam. Twigs 1—1.5 mm diam., not or hardly striate. — Philippines

Sulu Mindandoy ss)... oc. on oe ee eee ee oe nw eee 66. K. stenocarpa

. Buds 3 mm diam. Twigs 2—2.5 mm diam., coarsely striate or angled. — Philippines

el TECOT1 | inh Mil Rites Rap ibe i land oar gre ns Pea tanec gc aa 1. K. alvarezii

. Twigs 1.5—2.5(-3) mm diam., smooth, lower down sometimes striate. Buds 3—5

mm diam. Anthers 9—13(—15). Staminal disc + flat, convex, or low-mammillate.
Inflorescences sessile. (Compare also K. korthalsii and K. tomentella.) — Borneo
Ranawak)2 Philippines! eh 4)'S22 a0 ee ee 16. K. glomerata

. Twigs slender or rather stout, 1-3 mm diam., striate or ridged, lower down as well

as right up to the apex; if striations faint, then twigs 2-3 mm diam. (Twigs in K. to-
mentella from Sulawesi and Seram often hardly striate) ................. 50
Inflorescences all or partly up to 5 mm pedunculate. [Pedicel 3-11 mm.] Leaves 6—
18(—24) by 1.5—4(—6) cm, drying with a blackish metallic lustre above. [Twigs 1-2
(—2.5) mm diam.] SERIES OBOVOIDEAE. — Often coastal; Sumatra, Peninsular Ma-
Weaysian Widlavaliic), Lleids, 28 Ae RS ME PER ek 15. K. globularia

. Inflorescences all sessile. Leaves drying without a blackish metallic lustre. —C &

BOM Galesian, Fee sce isis seis asdeus aon: Sees LAN 7 2. eee 56
Buds 1.5—2 mm diam. — Low tree, 2—5 m; Borneo (S Brunei) .. 40. K. minima

Buds2: simimndiam: Onimore aeaere ks soe eee: Sh al Ee RE 57

Flora Malesiana, Ser. I, Vol. 14 (2000)

58a.

60a.

61a.

64a.

a. Pedicel 2-5 mm long. Buds subglobose, 2.5—3 mm diam., with hairs 0.3—0.5 mm

long; anthers (6—)8-11, each 0.3(—0.5) mm long. — Sulawesi, Moluccas, Papua
Bars (Bird?s: Headyaagccies. ce cares. DEERME 71. K. tomentella

_ Pedicel 1-2 mm long. Buds + pear-shaped, 3.5 mm diam., with coarse hairs 0.7

mm long; anthers 7-9, each 0.5 mm long. — Philippines (Palawan) (see also subsp.

ridleyi forma nana, from Borneo) .......--- 27a. K. latericia subsp. latericia
Leaf upper surface drying without blackish metallic lustre. Buds with hairs 0.1—0.2
MMMONMORIESS ..e.ecses eee ee eles eS 2S ey a9

. Leaf upper surface drying usually with a blackish metallic lustre. Buds with hairs

more than 0.2 mm long. [Pedicel 3—1 1 mm long.] (SERIES OBOVOIDEAE). — Sumatra,
Peninsular Malaysia, W Java ............ 2 ee eee eee eee 15. K. globularia

_ Leaves membranous or chartaceous, lower surface early glabrescent except for the

basal part of the midrib with minute hairs. Twigs usually flattened or blunt-triangu-
lar, at first with farinose orange-red or orange yellowish hairs 0.2 mm long. —
Sumatra, Peninsular Malaysia, Singapore .........-..-.-+++-: 60. K. rubens

_ Leaves membranous, chartaceous, or coriaceous; lower surface completely glabres-

cent or with a variable (sub)persistent indumentum. Twigs (sub)terete (in K. /uteo-
la + flattened), glabrescent, at first with hairs 0.1—0.2 mm long or less .... 60
Lower leaf surface with either 1) (sub)persistent, though usually inconspicuous,
scale-like hairs 0.1 mm long or less (lens!) or 2) (late) glabrescent, at first with
more conspicuous dense stellate-dendroid hairs 0.2 mm long. [Fruits with (sub)-
persistent indumentum.] ..........0: eee eee eee eee eect ete es 66

_ Lower leaf surface (early) glabrescent; at first with sparse or dense, weak and in-

conspicuous, minute hairs. [Venation on upper leaf surface usually distinct.] . 61
Leaves coriaceous, drying brown above; venation on upper surface distinct and
prominent. Twigs drying dark brown to blackish. Fruits with persistent very short
WMGUINENUEMMN A:-.15- ORI Ra GR Ke SNe 6 ABO MB « Ses. Saws ee ee ane 62

_ Leaves membranous or thinly coriaceous, drying greenish, brown, or blackish above,

venation distinct or not. Twigs drying grey or brown. Fruits early glabrescent, finely
warty or in K. luteola with sparse minute scale-like hairs ..........++++-- 63

. Twigs 1.5—2.5 mm diam. Leaves 9-21 by 2.5—5 cm, drying brown above. Bracteole

caducous. Perianth inside red. Anthers subsessile. — Borneo (Sabah: Mt. Kinabalu
ang. VACINILY) yan eas sR facie eerene Soe a 20. K. kinabaluensis

. Twigs 3—4 mm diam. Leaves generally larger, 14-31 by 4.5—10.5 cm, drying oliva-

ceous(-brown) above. Bracteole persistent. Perianth inside creamy-yellow. Anthers
stiped. — Central Sulawesi (Lake Matano area) .......... 38. K. matanensis

. Twigs somewhat flattened. Venation on upper leaf surface very fine, areoles less

than 0.5 mm diam. Bracteole caducous. Perianth inside yellowish 34. K. luteola

. Twigs terete. Venation on upper leaf surface with areoles 0.5 mm diam. or more.

Bracteole persistent or caducous. Perianth inside reddish (or in Java, Bali, and Borneo
réddishor yellowish). sa. 2.Guel) «- . See eee ie eee = eieleiele eae ee 64
Lateral nerves + flat or sunken above; venation in older leaves usually distinct.
Bracteole apical, caducous. [Leaves 5.5—12 cm wide, base cordate or rounded, not
attenuate. Anthers 8—10.]— Bormeo .............5+. 22. K. kostermansiana

_ Lateral nerves raised above; venation distinct. Bracteole below the apex, usually

PEUSISEEME. G.< dacireccjpese cage sete rs so euasanere tenehe = foes tei eee eee net ae ane ok ee 65

De Wilde — Myristicaceae (Knema) 231

65a. Leaves 6—20(—25) cm long. Buds 2.5—3.5 mm diam. Anthers 8-10, or in E Java and

66a.

67a.

68a.

69a.

70a.

Tia.

72a.

Bali frequently and in Borneo sometimes 11-15. Fruits 1.8—3(—4?) cm long. —
Sumatra, Peninsular Malaysia, Borneo, Java, Bali ............. 13. K. glauca

. Leaves 12—30 cm long. Buds 3-4 mm diam. Anthers 12—18. Fruits 3-4 cm long. —

Sumatra, Peninsular Malaysia: <).chpslyan? dc) cee 70. K. sumatrana
Leaves (when submature) on lower surface with + dense hairs of 0.2 mm long, of
+ mixed size; mature leaves often glabrescent. Venation on upper surface distinct,
fine, areoles 0.5 mm diam. or less. [Bracteole caducous or rudimentary.| .... 76

_ Lower leaf surface either with hairs 1) sparse, uniform, scale-like, 0.2 mm long or

less, or 2) dense, minute, scale-like, 0.1 mm only. Venation on upper surface dis-
FAMTISTROUBTROL ie, Bs doc airel guage tneg Shi yentewagies coplaconsrs cet uenars peea 4 corre haere tah cba 67
Leaves very coriaceous; venation prominent. Bracteole caducous. Buds pear-shaped,
at base tapering or not (recheck also fork 21). Fruits globose, 3—3.5 cm diam.;
fruiting pedicel 1-3 mm long. (SERIES OBOVOIDEAE). — Peninsular Malaysia... .
ae ee 59a. K. rigidifolia subsp. rigidifolia

. Leaves membranous or chartaceous; venation not so very prominent, distinct or

not. Fruits ellipsoid or obovoid, up to 2.5(—4) cm long; fruiting pedicel to 15(—20)
CETL LOT CAR aa ee eae aes spree aD Saleen Tepe ae Se oe 68
Buds 2—3.5 mm diam.; pedicel 2-7 mm long, in Borneo to 10 mm long. Anthers
5-9, in Sumatra up to 11, in Borneo up to 15. Bracteole persistent or caducous. —
PUY PES ORMCO ©, 2) Suh tee bento oe pe Menta ttn ht Sai aimed the eine eae gil

. Buds 3—5 mm diam.; pedicel 6-15 mm long. Anthers 9—-15(—17). Bracteole caducous.

MMS SRI TTBES ETN O) coe a oso oe ace oh ee eee cel AG oe EE TE ote Ae Teo vena 69
Buds truncate or + saccate at base. Lateral nerves and venation flat or sunken above,
indistinct. Perianth inside yellowish (always?). — Sumatra, Peninsular Malaysia
«ase 5 cotnieit pane ean nln de set Meade <a aren dst ln Ir rved ce 46. K. patentinervia

_ Buds rounded, subtruncate, or short attenuate at base. Lateral nerves and venation

Mie UrAMUaIStING!ADOVE™™ oC ee ae ce et ee ee ters cece meee omer 70
Twigs 1-2 mm diam., smooth. Anthers opening almost laterally. Perianth pale yel-
lowish inside (always?). Fruits often narrowed at base, with woolly hairs 0.2—0.4
mm long. — Peninsular Malaysia, Singapore .............. 35. K. malayana

. Twigs 1.5-3.5 mm diam., striate. Anthers opening + downwards. Perianth red in-

side. Fruits + rounded at base, with scurfy hairs 0.1—0.2 mm long. — Lesser Sunda
istands(E ol Balt)! Philippines... ede etre oe ee eee 5. K. cinerea
Venation on upper leaf surface very fine, distinct; areoles less than 0.5 mm diam.
Buds 2 mm diam.; pedicel 2-3 mm long. Anthers 5 or 6 (or 10—12?). Perianth
inside reddish (?). (SERIES GLOMERATAE). — Borneo (Sarawak) .............--
Pen YB NA RS eee. a 72b. K. tridactyla subsp. sublaevis

. Venation on upper leaf surface distinct or indistinct; diam. of the areoles 0.5 mm or

more. Buds 2—3.5(—4?) mm diam.; pedicel 3-10 mm long. Anthers 6—13(—15). Peri-
antminside:creamy or yellowish (always?) .<..e.45.22-4500c+2hc88 tee. We
Twigs 1 mm diam. Leaves to 15 cm long, beneath with sparse scale-like hairs less
than 0.1 mm long. Buds globose or broadly obovoid, 2.5—3 mm diameter; anthers
6-11; androphore glabrous. — Sumatra, Peninsular Malaysia, Borneo..........
MEK. So 4 Sh ol Vet dora, et eS he OE 67. K. stenophylla

. Twigs 1-3 mm diam. Leaves to 30 cm long, beneath with hairs more dense, or hairs

longer, 0.1—0.3 mm long. Androphore glabrous or finely pubescent at base .. 73

i)
eS)
bho

Flora Malesiana, Ser. I, Vol. 14 (2000)

74a.

78a.

79a.

a. Buds (broadly obovate, 2 mm diam.) cleft to the base. Anthers 9, subsessile, half-

erect; androphore + tapering, glabrous or with a few minute hairs at base. Leaves to
25 cm long, base short-attenuate or rounded, lower surface with + dense, greyish,
equal-sized hairs 0.2 mm long. — Borneo ................ 57. K. riangensis

_ Buds cleft 3/4-4/5. Anthers 6—15, subsessile or stiped, + horizontal; androphore

little tapering, glabrous or finely pubescent towards the base. Leaves variable, at
base attenuate; hairs sparse, of mixed Sizes ........-.- 0... eee eee eee eee 74
Twigs 2-3 mm diam. Leaves to 25 cm long. Buds subglobose or broadly obovoid,
3-4 mm long. [Anthers | 1-13; androphore pubescent towards the base, not seen in
var. pilocarpa. Fruits 2.5—3.5 cm long.] — Borneo (Sabah) .... 17. K. hirtella

. Twigs 1-2 mm diam. Leaves generally smaller. Buds obovoid, 2.5-3 mm long ..

RY ee ete rr. i chee eats, Sets ree elem Oe erect rr 75

_ Buds 2.5-3 mm long, anthers 8-15, androphore glabrous or with minute hairs at

base. Leaves with distinct venation above. Fruits 2 cm long. — Borneo (Sarawak,

(EYE Ua eye as 1521 1 ied et ei eG AR Ag | 1 ee a eR 69. K. subhirtella
_ Buds 2.5 mm long, anthers 6-8, androphore glabrous. Leaves with + faint venation
above. Fruits 1.5—2 cm long. — Bomeo ............-....-- 41. K. mogeana
. Twigs (1.5—)2—4 mm diam. Buds 3.5—5 mm diam. [Perianth inside greenish to yel-
FoowasinaP AC ead Goes. 3 oe es) egd thse Rete te fete ates tee tec 78
. Twigs 1-2(-3) mm diam. Buds smaller, 2.5—3.5 mm diam. ............--- V7

. Pedicel 2—5 mm long. Leaves (oblong-)lanceolate, broadest at the middle; the base

cuneate. Perianth inside reddish (?). — Peninsular Malaysia, Singapore.........
ee ee ee ECE Ae MEIN oo Mo Doo oe atric 6. K. communis

_ Pedicel 5—11 mm long. Leaves oblong to lanceolate, broadest usually below the

middle: base rounded to cuneate. Perianth inside greenish creamy or yellowish. —

Sumatra, Peninsular Malaysia, Singapore, Borneo ......... 14. K. glaucescens
Leaves chartaceous, lower surface with + sparse greyish hairs. Pedicel 3-10 mm
lones=——Sumattayss 2/52) Oe ais ye 4 AO RIBS: eke ie 32. K. losirensis

_ Leaves (sub)coriaceous, the lower surface with dense, yellowish or rusty hairs, or

glabrescent. Pedicel 7-16 mm long (in series Glomeratae). — Sumatra, Peninsular
Malaysian. 2 Aaah 6 Corea eB epee Geer 62. K. scortechinii
Leaves with persistent indumentum beneath, or hairs shed late and then leaving
hair scars (lens!); sometimes sparse minute scale-like hairs only. Twigs striate (stria-
tions sometimes not distinct in K. kunstleri subsp. coriacea and subsp. alpina 81

. Leaves early glabrescent beneath. Twigs (indumentum removed) not striate . 80
. Twigs 1.5—4 mm diam. Leaves coriaceous, 10—20 cm long, apex truncate. Bracteole

apical. Anthers long-stiped. — Sulawesi ...........+-+-++--- 4. K. celebica

. Twigs 1-1.5 mm diam. Leaves chartaceous, 5.5—15 cm long, apex acute-acumi-

nate. Bracteole about median. Anthers just stiped. — Borneo (Sarawak)........
SWS, Sey RIS oh Pe 6 wiae eS 2 ed Rae Slee ie ee 42. K. muscosa

_ Leaves 10-35 cm long; lower surface with coarse, stiff, sessile and stalked hairs.

Buds: 6554 amma diamss sc ses e565 BRE Be lS. . Bl ee 84

. Leaves generally smaller, 5-20(-27) cm long; lower surface with mainly sessile

stellate (hairss Buds: 2—3(—4) mimidiami.. .... ....s.ct. nae ln pe oo ae 82

. Perianth inside pubescent. Staminal disc convex. — Borneo . . .54. K. pubiflora

b. Perianth inside glabrous. Staminal disc flat or shallowly convex .......... 83

De Wilde — Myristicaceae (Knema) 233

83a. Lower leaf surface with stellate and/or scale-like hairs. Flowers with hairs 0.1—0.2

84a.

85a.

mm long or less. — Sumatra, Peninsular Malaysia, Borneo, Philippines.........
Re LIS Rd a SAR I, Sek eee eae tee es to fo Soa 24. K. kunstleri

. Lower leaf surface with sessile hairs mixed with dendroid hairs, the latter some-

times shed early and leaving hair scars. Flowers with dense woolly hairs 0.2—0.3
mm long, mixed with emergent dendroid hairs. — N Sulawesi, Philippines ......
ta. OS 2 os oe ae i ee eae: 65. K. stellata
Twigs 2-5 mm diam. Pedicel 7-15 mm long. Perianth 3-lobed, disc at base absent.
Staminal disc subtriangular, flat or slightly convex. Anthers 10—17. Fruits ellipsoid
to subglobose; fruiting pedicel 5-10 mm long ...................00005. 85

. Twigs 4-6 mm diam. Pedicel 20-30 mm long. Perianth 4- or 5-lobed, at base with

conspicuous disc. Staminal disc circular, distinctly mammillate. Anthers 14-18.
Fruits globose or depressed globose; fruiting pedicel 18—30 mm long. — Borneo
ne gt) ERAS LS: Brie, bots ANT Mato t Ree ae 48. K. pedicellata

Buds 2(—2.5) mm diam.; anthers 8, each 4-sporangiate. — Borneo (Central Kali-
PemaaeISD) WALES ARE SS, I, OR EAE 23. K. krusemaniana
. Buds 4—4.5 mm diam., anthers (10—)12—17, each 2-sporangiate. — Sumatra, Pe-

musular Malaysia, Singapore; Borneo) 2.2). nee a 7. K. conferta

(2) REGIONAL KEY TO THE SPECIES — PENINSULAR MALAYSIA, SINGAPORE

la.

b.

(based on female flowering and fruiting specimens)

Lower leaf surface seemingly glabrous, though actually with (sub)persistent, dense,
+ flat, tightly interwoven hairs, silky to the touch. Leaves stout .. 56. K. retusa

Lower leaf surface either glabrous (glabrescent), or with indumentum not silky to
thomuchuiteaves larseior smalls) 02 0P eee. 2 ee Se 2
. Bark of twigs longitudinally cracking, when older flaking ................ 3
ABankon twigs neithencrackineg nor flaking” §. oi002) 2.08 Oe. Pe 6
. Twigs 4-12 mm diam. Buds 6-10 mm long. Fruits 2.5-8 cm long .......... -
. Twigs 2-4 mm diam. Buds 5—8 mm long. Fruits 1.5—2.5(—3) cm long. ..........

PGR 2 ihe. ives) Sele Pr sew gree Sts. dL See AR ak hg pee 27. K. latericia

. Hairs on twigs (3—)5—8 mm long, those of flowers 1-3 mm long, of fruits 5-13 mm

ine MNase Sie nse ug led el. Wea We SAE ae 18. K. hookeriana

BlAIMESHOTEE LUGS ols sat Ae DAL OSs sc oe eee: 5
. Pedicel 1.5—2 mm long. Fruits 2.5—3.5 cm long, fruiting pedicel 2-5 mm long....

MPIOR Tan iiie OMI, cat bre fe oo See See 11. K. furfuracea

. Flowers not known. Fruits 44.5 cm long, fruiting pedicel 18-25 mm long......

reais ove eo. A ul 4a Civitte GUS EE) ee hc: 2) 25. K. lamellaria

. Lower leaf surface with dark dots, usually on the smaller veins ............ y
aeewenleafsuriace! without dots) 1.io053sr3 SoNPsee Buea ASRS 8
. Lower leaf surface with sparse scale-like hairs 0.1 mm long..................

Bes Set cd ie a RR 24a. K. kunstleri subsp. kunstleri

. Lower leaf surface with hairs (0.2—)0.3-1 mm long ............ 7. K. conferta
. Leaves 3.5—15(—20) cm long, membranous or chartaceous, drying greenish or brown.

Twigs (0.5—-)1-2 mm diam., early glabrescent, often yellowish. Fruits often
glabrescent, 2-5 cm long. Anthers 9—25(—30), closely appressed . .8. K. curtisii

234

Flora Malesiana, Ser. I, Vol. 14 (2000)

b. Leaves small or large, membranous or coriaceous, greenish or brown. Twigs slen-

9a

der or stout, drying brown or blackish. Fruits variable. Anthers not tightly appres-
eral iht SUE OR Co OO S,.120 DANES... LE ARAN iets Me) os oe 9
_ Hairs on twigs and flowers (0.1—)0.2 mm long or more, hairs often comparatively
short in K. globularia and K. pseudolaurina, both with Striate twits eee 10

b. Hairs on twigs and (usually) flowers, 0.1—0.2 mm long or less; hairs comparatively

10a.

long in K. rubens, a species with twigs + flattened...) 0006 oo 5 16
Buds 8 mm long, inside with a conspicuous disc at base; perianth persistent in
LUMLEY SL ee 51. K. plumulosa
Buds 4-10 mm long, without disc at base; perianth not persistent in froitsae 1

. Twigs striate, |-2(—2.5) mm diam. Leaves 6-1 8(—24) cm long, drying with a black-

ish metallic lustre above; lower surface late glabrescent, at first with hairs 0.1 mm
ori seeik See ie EE Ce cence = 15. K. globularia
. Twigs striate or not, 1.5—6(—10) mm diam. Leaves 9—40(—55) cm long, without a
blackish metallic lustre; lower surface glabrescent or with persistent hairs 0.2 mm
Womexor mares PbS PLE Lo i Ra eg 0s ot eT, Nie ea 12

12a. Twigs flattened or bluntly 2- or 3-angular. Leaves (lens!) pitted above (like the sur-
face of the peel of an orange). Fruits usually beaked at apex 44. K. oblongifolia

b. Twigs + terete. Leaves not pitted. Fruits not beaked!. «. 4) os oe eee 13
13a. Leaves early glabrescent beneath ..........--. esses eens sete nae: 14
b. Leaves with persistent indumentum beneath ......---- +--+ ese eeeees 15
14a. Twigs (4-)5-8 mm diam. Leaves 15-55 cm long. Buds 8-10 mm long. Fruits 3-6
cm long, with hairs (0.5—)1-2 mm long .....----------- 37. K. mandaharan

b. Twigs 1.5—4 mm diam. Leaves 9-35 cm long. Buds 5(—6) mm long. Fruits 2.5-4
cm long, rather glabrescent, hairs 0.3-0.5 mm long ......-- 19. K. intermedia

15a. Buds 6-8 mm long, with hairs 0.1—0.5 mm long. Fruits (2.5—)3—4(-5) cm long.

18a.

Hairs on lower leaf surface stellate-dendroid, of mixed sizes 52. K. pseudolaurina

_ Buds 4-6 mm long, with hairs 0.5—1(—2) mm long. Fruits 1.5—2.5 cm long. Hairs
on lower leaf surface + equal, stalked-dendroid (of mixed sizes in var. heteropilis)
MADE OSI) PRN -Ae oie 8 aT Re tence me 29. K. laurina

. Twigs flattened, with hairs 0.1—0.2 mm long. Leaves early glabrescent beneath,
except the basal part of the midrib ............ 0025s seer eee 60. K. rubens

. Twigs terete, with hairs 0.1—0.2 mm long or less. Leaves glabrescent or with per-
sistent indumentum beneath” 2.0.0. 2.0 22 ee ee oe afe Se eee 17

_ Leaf base rounded or cordate. Lower leaf surface glabrescent, or with weak hairs
0.1-0.2 mm long. Fruits ellipsoid, (3—)4—6 cm long; fruiting pedicel (2—)5—12 mm
Lonetel RES I ES Resets Ir ERE ean 55. K. pulchra

_ Leaf base rounded or attenuate (rarely (sub)cordate in K. rigidifolia with coriaceous
leaves and K. scortechinii with coriaceous leaves and with subpersistent indumentum
beneath). Lower leaf surface glabrescent or with a variable indumentum. Fruits to

A-em long: fruiting pedicel vaniable 9 eee 2. oii. 102 eats ei 18
Lower leaf surface glabrescent, at first with weak greyish hairs 0.1 mm long or less
ete PE I ie | RR |. ES SU Ma dr 19

_ Lower leaf surface late glabrescent or with (sub)persistent hairs 0.1—0.4 mm long,
sometimes remaining on and near midrib and nerves (completely glabrescent, with
distinct hair Scarsinl KY 7igidifalia) "5. PEI ae aie oes ants oP nh le Fal 20

De Wilde — Myristicaceae (Knema) 235

Ditar

22a.

26a.

. Leaves 12-30 by 4.5—12 cm. Fruits 3-4 cm long .......... 70. K. sumatrana
20a.

Leaves coriaceous. Fruits (sub)globose, 3—4 cm diam.; fruiting pedicel 1-3 mm
ln cEeerer whee amb), sure Reaper ae eae eee Behe 59. K. rigidifolia

_ Leaves membranous or chartaceous, or + coriaceous in K. scortechinii and K. pa-

tentinervia. Fruits ellipsoid, of variable sizes; fruiting pedicel (3—)4—20 mm, in K.
Eamanica. 05-7 millones Ay te oe Ae et oT 21
Lower leaf surface largely glabrescent, hairs 0.1-0.4 mm long, persistent on and
near midrib and nerves. Style slender, stigma lobes erect, together 1.5(—2) mm long.
Male buds ellipsoid or pear-shaped. Anthers 6 or7.....-..--.--05 00+ e eee
RE aos nals eg athew 2aOk 2. K. andamanica subsp. nicobarica

. Lower leaf surface with conspicuous or inconspicuous (sub)persistent indumentum.

Style shorter or absent, stigma lobes + spreading. Male buds subglobose. Anthers
618° 6—Ulin/K. stenophylla; 7-91 K communis, \): 8.2.) 180 ADE + - 22
Leaves with + conspicuous rather dense stellate-dendroid hairs 0.2 mm long be-
neath (old leaves glabrescent). Venation distinct above, with areoles 0.5 mm diam.
Omlessiee, Joscerulsls note ied. nis. 6S agra? . Aeon las eee 23

. Leaves either with sparse hairs 0.2 mm long or less, or with sparse or dense scale-

like hairs 0.1 mm long or less. Venation distinct or indistinct above; areoles 0.5 mm
Glicimomnene! ieseritidieire date er See OP RR ae eC BG 25

. Twigs 1-2(-3) mm diam., smooth, not wrinkled. Fruits 1.5—2.2 cm long .... 24
. Twigs (1.5-)2—4 mm diam., often wrinkled. Fruits 2-2.5 cmlong.............

Sern Brey ened bce le ease tent abe. Abe eae: 62. K. scortechinii

. Fruits 1.5-1.8 cm long. Buds 4 mm long; inside reddish (?). Leaves (oblong-)lan-

ceolate, broadest usually at the middle; base cuneate ......... 6. K. communis

. Fruits 1.8—2.2 cm long. Buds 4—5.5 mm long; inside greenish creamy to yellowish.

Leaves oblong to lanceolate, broadest usually below the middle; base rounded or
Stiiedte Mis: { Cue atheotaschs ais aed He ae) tS 14. K. glaucescens

. Leaves thinly coriaceous, glossy above. Lateral nerves + patent; nerves and vena-

tion on upper surface flat or sunken, indistinct. [Perianth inside yellowish (always). ]
IE oa isco keonecyervrcve seis Becca Se ee 46. K. patentinervia

. Leaves membranous or chartaceous, usually dull above. Lateral nerves less patent;

HERES and venationidistinctionnotabovers... Eee Auee tesa. seek. 26
Lateral nerves and venation sunken, flat, or raised, distinct or indistinct above.
Perianth inside creamy or yellowish (always?). Fruits at base rounded or some-
times attenuate; hairs scale-like, 0.1 mm long or less ....... 67. K. stenophylla

. Lateral nerves and venation raised and distinct above. Perianth inside creamy. Fruits

narrowed at base; hairs 0.2-0.4 mm long ................. 35. K. malayana

(3) REGIONAL KEY TO THE SPECIES — SUMATRA, JAVA
(based on female flowering and fruiting specimens)

. Bark of twigs longitudinally cracking, when older flaking ...............- 2
) Bark of twigs neither cracking nor flaking .. 0.4.65 sees ee eke. 5

Flora Malesiana, Ser. I, Vol. 14 (2000)

8a.

9a.

12a.

13a.

. Twigs 5-10 mm diam. Buds (6—)8-10 mm long ....--.... +++ +e sees 3
. Twigs 2—4 mm diam. [with hairs 0.5—-2 mm long]. Buds 5—8 mm long. [Fruits 1.5—

2.5(—3) cm long, with hairs 1-2 mm long.] — Northern Sumatra and Bangka.....
bh. doapd ieyty SeOeORR PRC SAIS TF ov IRA FE 27b. K. latericia subsp. ridleyi

a. Hairs on twigs (3—)5—8 mm long, on flowers 1-3 mm long. Fruits 4.5-8 cm long,

Wwithihales!5—= loam longo y eS. a). Bees 18. K. hookeriana

_ Hairs on twigs 1—2.5 mm long, on flowers 0.7—1.2 mm long. Fruits 3 cm long, with

hairs 0:6=2, mmilOng wc so oan Vek wee «le SUR.) CG eee 4

SBratines pedicel 1 Oman.) fh... 180). S28 eS, Lone 26. K. lampongensis
. Fruiting pedicel 2-5 mm long ........-.-. +... 26sec ees 11. K. furfuracea
_ Leaves on lower surface with dark dots, especially on the veinlets .......... 6
. Leaves on lower surface without dots ......... 0.0 e eee ee eee eee eee eee ii
_ Lower leaf surface with persistent, sparse, scale-like hairs 0.1 mm long.........

hy. ache ha, HEA RAE ee 24e. K. kunstleri subsp. macrophylla

_ Lower leaf surface with persistent stellate-dendroid hairs (0.2-)0.3-1 mm long

bial one tidbwiierd bicghiel itl lols enol: TA. PO Agee Re 7. K. conferta

. Twigs (0.5—)1—2 mm diam., early glabrescent, yellowish. Leaves olivaceous above,

early glabrescent beneath. [Fruits 2-5 cm long, often glabrescent, often + beaked
ELAS | ENR A LEY tO FO a NON eI te 8a. K. curtisii var. curtisii

. Twigs slender or stout, drying brown or blackish. Leaves olivaceous or (dark) brown

above, with a variable persistent indumentum, or glabrescent beneath ....... 8
Hairs on twigs and flowers (0.1—)0.2 mm long or more (hairs comparatively short
in K. globularia and K. pseudolaurina, both with striate twigs) .....--..--- 9

. Hairs on twigs and flowers 0.1—0.2 mm long or less (hairs comparatively long in

K. rubens, with twigs + flattened and fruits usually with acute apex) ....... 13
Twigs striate, 1-2(—2.5) mm diam. Leaves 6—18(—24) cm long, drying with a black-
ish metallic lustre; lower surface late glabrescent, hairs 0.1 mm long.........--
aries, mae edd htt. xd a ee Rates Ae RY att. D2 15. K. globularia

. Twigs striate or not, 1.5—6(—10) mm diam. Leaves 9—40(—55) cm long, without

blackish metallic lustre; on lower surface glabrescent or with persistent hairs 0.2
mmnloneorimMore Hi. GU) BT RRS Ae ES Sees 10

. Leaves early glabrescent beneath ............-- eee eee e eee teens 11
. Leaves with persistent indumentum beneath ..........-----++seeee eee 12
. Twigs (4-)5—8(—10) mm diam. Leaves 15—55 cm long. Buds 8-10 mm long. Fruits

3-6 cm long, with hairs (0.5—)1—-2 mm long ............ 37. K. mandaharan

. Twigs 1.5—4(—5) mm diam. Leaves 9-35 cm long. Buds 5(—6) mm long. Fruits

2.5—4 cm long, rather glabrescent, at first with hairs 0.3-0.5 mm long......-.--
esheets nme HGR a SARA GEER LARS IRE OY 19. K. intermedia
Buds 6-8 mm long, with hairs 0.1—0.5 mm long. Fruits (2.5—)3—4(—5) cm long.
Hairs on lower leaf surface of mixed sizes ..........-- 52. K. pseudolaurina

_ Buds 4-6 mm long, with hairs 0.5—1(—2) mm long. Fruits 1.5—2.5 cm long. Hairs on

lower leaf surface all + equally long (strongly unequal in var. heteropilis)......
idle alas nw p/p ARR SAE VAR SARMOLL RGR BERRI): 620 29. K. laurina
Leaves (obovate-)oblong, (7—) 10-30 by (2.5—)4.5—12.5 cm, drying brown or black-
ish above, often + glossy. Fruits 2-3.5 cm long, often ridged, at base somewhat
saccate, hairs minute. Anthers 3 or 4 ..........-- 0 seer eee 28. K. latifolia

De Wilde — Myristicaceae (Knema) 237

la.

_ Leaves variable in shape and size. Fruits various, not saccate at base. Anthers 6 or

TO LCM ead her ngage SOIT nT epee n= A ES 3d Ee A een Gone 14

. Twigs somewhat flattened, with hairs 0.1—0.2 mm long. Leaves early glabrescent

beneath, except for the basal part of the midrib ............... 60. K. rubens

. Twigs terete; hairs 0.1 mm long or less. Leaves glabrescent or with persistent

SAGITETTNE TAGE CTS ALIN Ley ised 5 ae cee Ae RPE eee 15

. Style slender, stigma erect, 3—8-lobed, together 1.5(—2) mm long. [Male buds

+ ellipsoid or pear-shaped; anthers 6 or 7.] — Sumatra (Aceh)................
Bea oh nin. tee ER 2. K. andamanica subsp. nicobarica

. Style up to 0.5 mm long; stigma broader, 6—12-lobulate ................. 16
. Leaves early glabrescent beneath, at first with weak hairs 0.1 mm long. Fruits

SbeeM NTIS SC CTIEN Etiam th’ Matfer. sta". Sharpest’, Seta eters 727s Ss aay eee ee 17

. Leaves with persistent indumentum or rather late glabrescent beneath, at first with

hairs 0.1—0.2 mm long, mixed with (sometimes caducous) hairs to 0.5 mm long or
ROL eruits with persistent indumentum .../2. 2. .Seeee ence =e r 18

. Leaves 12-30 by 4.5-12 cm. Fruits 3-4 cm long .......... 70. K. sumatrana
. Leaves 6—20(—25) by 2-5.5(-11) cm. Fruits 1.8-3(—-4?) cm long ............

ce AR Re ee eee eran mn ean DS wnt MeL MRR Reem LM ects 13. K. glauca

_ Lower leaf surface with stellate scale-like hairs 0.2 mm long or less. Lateral nerves

PRARVCHAIOM ADOVE GIstinet OF qOt c.ctace c o43crcuces Geesca okbercseuenees Geter “oer = 19

_ Lower leaf surface with hairs 0.1—0.2 mm long, mixed with (sometimes caducous)

hairs to 0.5 mm long. Lateral nerves and venation above distinct .......... 20

. Leaves thinly coriaceous, glossy above. Lateral nerves 12—24 pairs, rather patent,

flat, indistinct above; venation indistinct above. Male buds 3.5—5 mm diam.; an-
LRCTES 111 eae Re ee ie ee sar de SPAS cite 46. K. patentinervia

_ Leaves membranous or chartaceous, not glossy. Lateral nerves 10-21 pairs, less

patent, flat or raised, usually indistinct above; venation distinct or indistinct above.
Male buds (1.5—)2—3(—4?) mm diam.; anthers 6-11 ....... 67. K. stenophylla

. Twigs 1—2(—3) mm diam. Leaves 6-20(—23) cm long; lateral nerves 13-20 pairs.

amity less) CIMLOMS” « saecme scares AES sac one dee Ges cee 14. K. glaucescens

. Twigs 1.5—4 mm diam. Leaves 10-30 cm long; nerves 18—30 pairs. Fruits 2—2.7

erin) [OTT ES ate a mee ene npr) PoP ce iar caiy omigal ac 2M

. Twigs 1.5—2.5 mm diam., bark striate. Leaves + membranous —_ 32. K. losirensis
. Twigs 2-4 mm diam., bark usually wrinkled on drying. Leaves coriaceous ......

Re ag be on Th need UMD faut sv cere ee 62. K. scortechinii

(4) REGIONAL KEY TO THE SPECIES — BORNEO
(based on female flowering and fruiting specimens)

Lower leaf surface seemingly glabrous, but actually with persistent, dense, + flat,

imierwoven hairs, silky tothe touch 2... :..ergoeb22 4° sed oer Ge: Z
. Lower leaf surface either glabrous (glabrescent), or with variable indumentum not
silkeytoyihe fowchiys-az asa Not vse aiacys GAd ese cobs eh Sha acloer eee +
. Bark of twigs longtitudinally cracking and flaking ............. 9. K. elmeri

. Bark of twigs neither cracking nor flaking © 4)..(45: 2-2 os ese: Sek 3

Flora Malesiana, Ser. I, Vol. 14 (2000)

3a. Leaf apex + rounded or acute-acuminate. Hairs on flowers 0.1 mm long. Fruits

Aa.

subglobose to broadly ovoid, 4.5—5 cm long. [Staminal disc conspicuously con-

Vek|@ vfueemeee eee had Masia ele tease cee 63. K. sericea
_ Leaf apex acute-acuminate. Hairs on flowers inconspicuous, scale-like, 0.1 mm
long or less. Fruits (obovoid-)oblong, 5—8 cm long. [{ Venation on upper leaf surface
finer dhaninyKe S@riGe@Ga << Aasccecc oor’ me 3b. K. ashtonii var. cinnamomea
Bark of twigs longitudinally cracking, when older flaking (sometimes not apparent
inwK. PerCOMIGCeG) A). cacaae--. )\-25 a -eodine teil ck: ee DE 3)

b. Bark of twigs neither cracking nor flaking .......--.... +++ settee eens 13

. Flowers wholly or largely glabrescent ...........-- +s eee eet e eee teens 6

_ Flowers with persistent indumentum (but sometimes easily rubbed off in K. per-
COTLAGCEG)’. cc sesesciana coi foiens GOEL IRR «6s sw Dena o Hala wes lr 8

_ Fruits with persistent hairs | mm long (rarely partly glabrescent) 53. K. psilantha

Fruits slabrescemt .... 0.0.5.0 «:ssey-jey syns che shtnn 2) eee ee Ge if

_ Buds + mitriform, sharp-angled, 7-9 mm long. Twigs 4-10 mm diam., brown... .

OR EE re tot tic cero See ioe ee ope oe 12. K. galeata
_ Buds not sharp-angled, 4 mm long. Twigs 1.5—3 mm diam., often yellowish......
A POT OP IN RON eS olaraests overeat 39. K. membranifolia

. Twigs (1-)2—4 mm diam. Flowers sessile or pedicel to 1.5 mm long (see also K.

Koripalsit SUDSP. TUMOSA) a. oc 3 oo eo 27. K. latericia
. Twigs 3—-6(—12) mm diam. Pedicel 2-20 mm long (up to 2 mm in K. lunduensis;
flowers and fruits not known in K. longepilosa, but pedicel presumably compara-
LIVELY. SHOT) cgeje pg ee oe oe eg ey sce ine ea ia 9

_ Buds 6-9 mm long. Fruiting pedicel up to 3(-5) mm long ...........-+--- 10

_ Buds 9-11 mm long. Fruiting pedicel (7—)10-30 mm long .............-. Ne

_ Leaves chartaceous or thinly coriaceous, drying brown above; leaf base obtuse or

subcordate (rarely + narrowed) ..........-.....-+-+----; 33. K. lunduensis

_ Leaves coriaceous, drying greenish or brown above; leaf base attenuate to rounded
ee on a eae ere sais hig: 10 Oe) SIC ca rr 11

_ Hairs on flowers and fruits | mm long. Anthers 10-15, shortly stiped.........--.

“AERr ts ASSERT REPL SEI LN TRS RR UIE 3c Ey AO BABI 49. K. percoriacea
_ Hairs on flowers and fruits (2—)3—4 mm long. Anthers 15-18, sessile.........--
_ rca lied cea a cig aah aa re aa ee a A oN LCI ECB: 31. K. longepilosa
. Hairs on twigs 1—2.5 mm long; on flowers 0.7—1.2 mm long. Fruits 2.5 cm long. —
Sumatra and Anambasiss . Sos...) eee we eee eee 26. K. lampongensis
_ Hairs on twigs 0.2-1(—2) mm long; on flowers 0.1—0.8 mm long. Fruits 3.5—7 cm
[oteameeee ers ee tee, Sen Sore ets mine eer erste ete crane ae 45. K. pallens
_ Lower leaf surface with dark dots, predominantly on the smaller veins ..... 14
_ Lower leaf surface*withoutidots!". 25/0 001. 2702. St a ee 2 ee 19
. Twigs 1-1.5 mm diam. Leaves glabrescent beneath. Fruits obovoid-oblong, 1.7—
ih Stemilonomen set. ets A Neem as 2 a. Hitt hit are 42. K. muscosa
. Twigs 1.5-6 mm diam. Lower leaf surface with (sub)persistent indumentum. Fruits
of various shapes, 1.5=4 cmilong 950. ee ne oe 15
_ Lower leaf surface with sparse hairs 0.1 mm long, sometimes with a few larger

ones in-between. (Older leaves sometimes glabrescent in subsp. a/pina.)......--
BRD AR) 9S tot, dl ee Pe, ME Ee eR EMRE Cre 24. K. kunstleri

De Wilde — Myristicaceae (Knema) 239

16a.

b.

17a.

Leaves 4—15 cm long, drying brown above. Perianth inside hairy. [Fruits subglobose

or short-ellipsoid, 1.5—-2.5 cm long.] .....-..--.+-.0+00-05- 54. K. pubiflora
Leaves 10-36 cm long, drying olivaceous or brown above. Perianth inside gla-
Tarorrcunae (eek ill GE) tO MI, LOGS RF ek RN a Mat SENS ict BIEN | 17

Female pedicel 5—8(—10) mm long. Perianth lobes 3 or 4. No disc at base of peri-
anth. Fruits subglobose or usually ellipsoid or obovoid, (1.5—)2—4 cm long; fruiting
pedicels — 10mm long. x .)).0e Yee eh EE FREE cen ee : 18

. Female pedicel 10-12 mm long. Perianth lobes 4 or 5. Perianth with disc at base.

Fruits subglobose, usually somewhat broader than long, 2—3 by 2.5-2.8 cm; fruit-
ing pedicel 18-30 mm long ..........----.-- +s seer eee 48. K. pedicellata

_ Male buf 4—4.5 mm diam., lobes usually 3; anthers (10—)12—17, each 2-sporangiate.

— Sumatra, Peninsular Malaysia, Singapore, Borneo .......... 7. K. conferta

_ Male buds 2(—2.5) mm diam., lobes 4; anthers 8, each 4-sporangiate. — Borneo

Meomralialimantany Set oe ec ete ce ees 23. K. krusemaniana

. Fruits glabresent. Perianth glabrescent, except in K. linguiformis, K. viridis, and K.

curtisii p.p. Bark of twigs frequently smooth, glabrous, yellowish or purplish (but

ROITIIAT ORC WOO) no ego 0 vir rg oe nae a ga 20
_ Fruits pubescent or glabrescent. Perianth with persistent indumentum. Bark of twigs
FOOT IOLACKISH, ores 2 cite oocytes ig eo oho eas een nace aks Rae eae aS
. Twigs 4—8 mm diam. Buds 8—9 mm long, + mitriform, Sharp-anelede 2.0.2.5
ee cage Scares catego og) cna) pega ee ea gear 12. K. galeata
. Twigs 1-3 mm diam. Buds 4-6 mm long, not mitriform .............---- 72)
. Twigs glabrescent, at first with hairs | mmlong .............+----++++: Ups
. Twigs glabrescent, at first with hairs 0.1-0.5 mm long ........---------- DS

. Flowers glabrescent, at first with hairs 0.5—1 mm long. Fruits not seen..........

nn RRR SE er oe ee 36. K. mamillata

. Flowers glabrescent, at first with hairs 0.1 mm long. Fruiting pedicel 1-5 mm long

ca a ERR Ste en are en eee 39. K. membranifolia

_ Leaves 17-30 cm long. Pedicel stout, 15 mm long. Fruits S—6 cm long, pericarp 10

err in arr aE Ot a ores oa 74. K. viridis

_ Leaves to 20(—25) cm long. Pedicel (2—)5—10 mm long. Fruits 2-5 cm long, pericarp

A STAMINA UNL CK ge Sil A 5 ROMS 8 ete 2 wl eke ai ole oa ous BO OF Sige eis 6 eke: 24

. Twigs with hairs 0.2-0.5 mm long. Fruiting pedicel (2—)5—8(-16) mm long......

Ae ree ree 30. K. linguiformis

. Twigs with hairs 0.1 mm long or less. Fruiting pedicel 8-16 mm long........-.

(Te ee ee ee ee es Ee is 8. K. curtisii

. Twigs with hairs 0.2 mm long or more ..........--+--+-+esseeeeteeee 26
. Twigs with hairs 0.1 mm long or less ........ 052+ +15 sees sees eet ee eee 35)
. Lower leaf surface with persistent indumentum or (late) glabrescent ....... Zl,

_ Lower leaf surface glabrescent, sometimes with some indumentum persistent on

mianband nerves inK. glomeraia .......7ebi ae 2 «Year ee ees 29

_ Lower leaf surface with + uniformly long stalked dendroid hairs. Fruits 1.5—3(—4)

cm long, with hairs 1-3 mm long; fruiting pedicel up to 1(—5) mm long.........
A bieerS 4:19: perce Melbicing eaves de aiota lot: opera ase 29. K. laurina

_ Lower leaf surface with hairs of variable sizes, sessile and stellate-dendroid . 28

240

Flora Malesiana, Ser. I, Vol. 14 (2000)

28a. Buds 6—7.5 mm long. Fruits 2—3.8 cm long, with hairs 1—-1.5 mm long; fruiting

pedicel 3-10 mm long. [Female flowers and fruits not known of subsp. parviflora]
pinch ee. oo Ate wo Ao. evan gach onal hee 43. K. oblongata

. Buds 2—2.5 mm long. Fruits 1.4—2 cm long, with hairs 0.2—0.5(—1) mm long; fruit-

imeapedicel 2-3 mmmblomsyet del. Mine. 0) ererdanes Janek Ae 72. K. tridactyla

29a. Leaves drying greenish brown or blackish above. Fruits with hairs 3mm long....

det, Athe i yh Sten treba i. oes. nme teller U8 2 le peace a 61. K. rufa

. Leaves drying olivaceous brown above. Fruits with hairs 0.1-1 mm long ... 30

30a. Buds 2.5 mm long. Fruits 1.5-2 cm long. Tree 2-5 m.—S Brunei 40. K. minima

+ Buds and! fruits larcer: Trees generally larger .. ...0):).2 2-2: eee eee oul
_ Leaves coriaceous: midrib, nerves, and venation much raised and distinct above

PTR koh meron ieOSE ate pcan wae state at acacia a2

. Leaves chartaceous; midrib and nerves flat or little raised above, venation (little)

raised Wess idIStNCt ADOVE ......5....6 5 + wa. 200 oe ep ayisopehe 35

. Bracteole median on the pedicel. Staminal disc long-mammillate..............

ectnn © Sohne Sin pire rer a hey ene rant, Ramee eis 2 ot, oe ee 19. K. intermedia

. Bracteole (sub)apical. Staminal disc convex or low-mammillate 73. K. uliginosa
. Twigs 3—5 mm diam. Fruits 1.5—4 cm long, with hairs 0.3—0.5(—1) mm long; fruit-

meypedicel-2—Omnml IONS PK... oe see ne tee eee oe a yee 21. K. korthalsii

. Twigs 1.5—2.5(—3) mm diam. Fruits 1.5—2.5(—3) cm long, with hairs 0.1—0.2(—0.5)

mm long; fruiting pedicel 4-10 mmiong .......:......-.--.++eesonmme 34

. Fruiting pedicel 4—6(—10) mm long; fruits with scurfy hairs 0.1—0.2 mm. Leaves

drying green or brown, venation + distinct. —Borneo (rare) .. 16. K. glomerata

. Fruiting pedicel 7-10 mm long, more slender; fruits with mealy hairs to 0.5 mm

long. Leaves drying conspicuously green, venation distinct ..... 10. K. emmae

. Fruits largely glabrescent, somewhat flattened, often ridged, at base usually some-

what saccate. Leaves (obovate-)oblong; drying dark brown above, paler, often tinged
reddish brown or purplish beneath. Anthers 3 or4 ............ 28. K. latifolia

. Fruits either with persistent indumentum or glabrescent and then not saccate at the

base. Leaves variable in shape; drying olivaceous or brown above, greyish be-
neath “Amthers 5-25) oss ib ie we ose pe ee es, Sw eo eke oe 36

. Buds 2—2.5 mm long. Fruits 1.4—2 cm long; fruiting pedicel 2-3 mm long. Anthers

5) (Ci Caan RR Ran eee RRC RECT ots ro 72b. K. tridactyla subsp. sublaevis

. Buds 3.5-7 mm long. Fruits variable in size; fruiting pedicel usually more than

Srinimd lone sAnthers:6 Or MOre ¥.% 5 2 o: biealcas Ge 6 < ON ee 37

. Twigs 4—6 mm diam. Fruits 6-8 cm long with blunt or rounded apex. Venation of

upper surface of leaf fine, areoles 0.5 mm diam. or less (compare also K. luteola)
PER te ee eae ER Ferree TEN ich Bhatt 3a. K. ashtonii var. ashtonii

. Twigs variable, 1-6(—-8) mm diam. Fruits to 8 cm long; if over 6 cm, then the apex
acute, not rounded. Venation of upper leaf surface variable ............... 38
. Lobes of perianth 1—2 mm thick. (Female flowers not seen in K. pectinata.) . 39

» Lobes: of perianthy toil mumthicks 2:2... «23a = eee ee ee 40
. Twigs blackish brown, + angled by ridges down from the base of the petioles.

Leaves thinly coriaceous; lateral nerves 20-30 pairs ........... 75. K. woodii

. Twigs (greyish) brown, not angled. Leaves (rigidly) coriaceous; the lateral nerves

DIDO AES. | AGEs EMRE, PMS, SEALE Ya UAE TEN ES 47. K. pectinata

De Wilde — Myristicaceae (Knema) 241

40a.

b.

Ala.

50a.

Leaves 12-50 cm long, base broadly rounded to cordate. Twigs rather stout, 2-8
ARANETA ATTN § 40) 2f21 794) APNE decker tri ee ik ewe wb Oem be Eee Byes 4]
Leaves generally smaller, to 25 cm long, base attenuate to rounded (casually sub-
cordate or broadly rounded in K. glauca, K. hirtella, K. riangensis, and K. stylosa).
Mertace t—3-) iting s)y)t ). sie weet. a) adn cele oe © RD oN EC I ie 42
Twigs 3—6(—8) mm diam. Leaves (20—)30—50 by (6—)10-17 cm, nerves flat or little
raised above. Fruits subellipsoid, (3—)4—6 cm long, apex rounded. Male buds pear-
ae SCMRPS VON cries chao 6 8's 2a. Orage: eee. che ann Ae 55. K. pulchra

. Twigs 2-4 mm diam. Leaves 12-30 by 5.5—12 cm; nerves sunken to + flat above.

Fruits + slender, (5—)7—8 by 2 cm, apex long-acute. Male buds globose.........
Np Ee ented Ry op yn eG acaene Tenens 22. K. kostermansiana

. Leaves (rigidly) coriaceous, venation on upper surface raised, very distinct. — Alti-

tude 1000-2300 m; Sabah (Mt Kinabalu and vicinity) .... 20. K. kinabaluensis

. Leaves membranous to thinly coriaceous; venation on upper surface distinct or

not. — Altitude 0—2000 m; some species also on Mt Kinabalu ............ 43

. Twigs somewhat flattened, light brown. Venation on upper leaf surface distinct,

veuy, tine, jtheareoles less, than 0.5 arm diamine jo...) 22008) «ope jei2y-ceieye = 34. K. luteola

. Twigs terete or somewhat angular, drying brown or blackish. Venation on upper

featasMitace Coarser: GIStinct OF NOL ....0... +6 .<.. 4 bond qa eae oe 44

. Fruits glabrescent. Bracteole usually persistent. Leaves beneath early glabrescent

(at first with very weak greyish hairs beneath) ................ 13. K. glauca

. Fruits with persistent indumentum (hairs may be very short). Bracteole persistent

or caducous. Leaves with persistent indumentum, glabrescent beneath ..... 45

. Male buds pear-shaped, much tapered in the lower half. — A (sub)montane species

a2002=) 100022000 mvaltitide : a2... 28 ere ee. Oe Wee: 50. K. piriformis

. Male buds obovoid or globose. — Mainly in lowland forest, 0O—1000(—1200) m alti-

RRR eet SHEN ie 5) «ae eget da yaeys i221 ENE dees PETE GD oy ae Bee 46

. Lower leaf surface early glabrescent, at first with very weak indumentum. Style

5 —lanmrlong: Male buds .ebovoideyins. ies Bie S eh 68. K. stylosa

. Lower leaf surface with persistent indumentum or late glabrescent. Style to 0.5 mm

long (stigma lobes subsessile). Male buds obovoid or globose ............ 47

. Hairs on lower leaf surface dense, touching or interwoven ............... 48
Midirs sparse, notitouching each othenyc!: qua bake Pee eee 49
. Hairs brown. Lateral nerves usually raised above. Fruits 1.8—2.2 cmlong.......

a hf -sfibes iri Ge ap Os ee, ae ee 14. K. glaucescens

. Hairs yellowish or greyish white, weak and inconspicuous. Lateral nerves flat or

only partially raised above. Fruits (3—)4 cm long ........... 57. K. riangensis

. Twigs 1 mm diam. Leaves to 15 cm long, membranous, on lower surface with

sparse scale-like hairs less than 0.1 mm. [Fruits |—1.5 cm long, fruiting pedicel 10—
PO mnnvlong:)\ ot. syiead Roisin is.< 67b. K. stenophylla subsp. longipedicellata

. Twigs 1—3.5 mm diam. Leaves to 25 cm long, on lower surface with hairs of mixed

pizese |= Oe min lone, Mire 2.02 2c cecee meter pee ur een ena AWS. 50
Twigs 1—1.5(—2) mm diam. Venation on upper leaf surface rather faint. Fruits 1.5—
2 cm long, fruiting pedicel 4-10 mm long. Anthers 6-8 ...... 41. K. mogeana

. Twigs 1.5—-3 mm diam. Venation on upper leaf surface distinct. Fruits 2—3.5 cm

lenet. Asithiers Sala G15) wie oh ele Snead 2 oR RUSS BRR 51

242 Flora Malesiana, Ser. I, Vol. 14 (2000)

Sla. Twigs 1.5 mm diam. Leaves 9-20 cm long. Fruits 2—2.7 em long... aoseseee:

Tas ANE E s.0 hoe Sls oro Ee Ga Anes he eS Sono as 69. K. subhirtella
b. Twigs 3 mm diam. Leaves 16—25(—27) cm long. Fruits 2.5-3.5 cm long: baah abt.
Has Pua rls ose ad tia em ied 8. (cravat, Senet es eperaes 17. K. hirtella

(5) REGIONAL KEY TO THE SPECIES — PHILIPPINES, SULAWESI, MOLUCCAS,
LESSER SUNDA ISLANDS, NEW GUINEA (Bird’s Head)
(based on female flowering and fruiting specimens)

fa Leaves not dotted beneath (ens!) os... ee oe en 2
b. Leaves with dark dots beneath (present mainly on the nerves) .........-.. 13
2a. Twigs and lower surface of young leaves with hairs 0.2 mm long or more (usually
Short in Ke SlOMeTUIA) 0.6. etc ec ee ee ons oe ee coe s)

b. Twigs and lower leaf surface with hairs 0.1 mm long or less ......-...--+- 10
3a. Male inflorescences distinctly pedunculate. Perianth glabrescent. — Philippines
CNENEUZON) Come ee oes ee ects stom cine erery ect 58. K. ridsdaleana

b. Male inflorescences (sub)sessile. Perianth with persistent indumentum ...... 4
4a. Twigs 3-5 mm diam. [Fruits with hairs 0.5 mm long.] — S$ Philippines (incl. Pala-
NVA ee te eee Meech eas Mem | ne crear ae 21. K. korthalsii

bs Twies’3 mundiam. or less’. 2.0.5 46050 soe. es ee 5)
Sa. Twigs 1-2 mm diam. Female buds 4 mm long. Fruits 1-2 cm long A rece 6
b. Twigs (1.5—)2-3(—4) mm diam. Female buds 4.5—-6 mm long. Fruits (1—)1.5—2.5
Cai camloness (PT TI Oe EME tires sects te ere eon one eS qi

6a. Leaves on lower surface late glabrescent or with subpersistent minute sparse hairs.
Fruits 1.3—1.5 cm long, with hairs 0.1 mm long. — S Philippines (Sulu [., Mindanao)
OUT ee re UIs SME Stare Ne deere toe ney mere ene 66. K. stenocarpa

b. Leaves on lower surface early glabrescent, hairs shed as pieces of a matted indu-
mentum. Fruits 1.5—2 cm long, with hairs 0.5(-1) mm long. — S Philippines (Pala-
Wall) Montraiae, Wee Wee. Server ee 27a. K. latericia subsp. latericia var. subtilis
Ja. Twigs terete, smooth or very finely striate. Fruits with hairs 0.1—0.2 mm long. —
Enilippratcssta wey sere. Sa ea nee mene e es ote teeta nor 16. K. glomerata

b. Twigs striate or angled. Fruits with hairs 0.5-1 mm long .....-..--++----- 8
8a. Leaves on lower surface partly glabrescent or with subpersistent indumentum be-
neath. Fruits with soft hairs 0.5 mm long ..............+--+2 esses ueee 9

b. Leaves early glabrescent beneath. Fruits with rigid hairs 1 mm long. — Philippines
CBalaiaiineee 8 8 Me ot CY ete 27a. K. latericia subsp. latericia

9a. Lower leaf surface with subpersistent indumentum. — Philippines (Luzon) ......
RPV NP RAD 2s At, Es ASE PE a ig ee eee ee 1. K. alvarezii

b. Lower leaf surface largely glabrescent. — Sulawesi, Moluccas, Papua Barat (Bird’s
(G20) RUE aie eet Pee PR eB ca 71. K. tomentella
10a. Twigs 3-4 mm diam. Leaves coriaceous. [Perianth inside creamy-yellow. Fruits
with persistent short hairs. Male buds 2.5—3 mm diam.; anthers 7 or 8.] — Central
Siilawesir se BE i. OE ERE. onc Cee heme ene Beene 38. K. matanensis

b. Twigs 1-3.5 mm diam. Leaves membranous or chartaceous .........---+- 1]
lla. Lower leaf surface early glabrescent, at first with weak stellate hairs. Fruits 1.8-3
(—4) cm long, glabrescent. [Male buds globose, 2.5—3.5 mm diam.; anthers 11—15.]

—— Lesser Sunda Islands (Bali) (compare also K. glomerata) .... 13. K. glauca

De Wilde — Myristicaceae (Knema) 243

b. Lower leaf surface with persistent indumentum. Fruits 1.2-2.4 cm long, with
(sub)persistent minute indumentum ............---.e essere eerste ees 12

12a. Lower leaf surface with minute scale-like hairs. Style 0.5 mm long. Fruits 1.2—1.8
(-2) by 1-1.6 cm. Male buds globose, 3.5—4.5 mm diam.; anthers 9-14. — Lesser
Sunda Islands (except Bali), Sulawesi, Moluccas, S Philippines (Mindanao) .....
MT ee eo eM Are Die che sie tuna verieuie. estanne era Csiminars tee 5. K. cinerea

b. Lower leaf surface with mixed minute scale-like and stellate-dendroid hairs. Style

1 mm long. Fruits 2.2—2.4 by 1 cm. Male buds pear-shaped, 2 mm wide; anthers 4

or 5. — Lesser Sunda Islands (W Flores, at 780 m) ..........-- 64. K. steenisii

13a. Leaves with truncate apex, lower surface early glabrescent. — Central Sulawes! . .
A ee le, renarns we fears ie te er eee pens reas 4. K. celebica

b. Leaves not truncate at apex, lower surface with persistent indumentum or late gla-
BEE SCSTIE ete le VASAts, Aude ee SOT RINGE RILER. O). SEPA ET IS SLE We otohetene eh 14

14a. Lower leaf surface with hairs 0.1 mm long. — Philippines...............-.--
«8 dc oh oeeL RR MARS Wet aie Ua ethan eh Coat 24f. K. kunstleri subsp. parvifolia

b. Lower leaf surface with hairs of mixed sizes, 0.2—0.7 mm long. — Philippines,
RPO eS ulawesiv? / HSU PINS TORE A eRe Searls 20 32 65. K. stellata

1. Knema alvarezii Merr.

Knema alvarezii Merr., Philipp. J. Sci., Bot. 13 (1918) 288; Enum. Philipp. Flow. Pl. 2 (1923) 183;
W.J. de Wilde, Blumea 25 (1979) 437. — Lectotype: BS 22395 (Alvarez), Luzon.

Tree 10-15 m. Twigs somewhat angular, 2—2.5 mm in diameter, glabrescent, at first
with dense rusty dendroid hairs 0.5 mm long; bark striate, not tending to crack or flake.
Leaves chartaceous, oblong to lanceolate, 8-14 by 2.5-3.5 cm, base rounded or short-
attenuate, apex longly acute(-acuminate); upper surface olivaceous-brown, lower sur-
face greyish, with subpersistent grey-brown hairs 0.2—0.3 mm long, when shed leaving
minute hair scars; dots absent; midrib raised above; nerves 12-15 pairs, somewhat raised
above: venation fine and distinct above; petiole 8-15 by 1-2 mm, late glabrescent. Inflo-
rescences: peduncle 1-3 mm, the brachyblast simple or indistinctly forked, 2-4 mm in
diameter, in male 5—15-flowered; flowers with persistent rusty hairs 0.5(—0.7) mm long,
perianth 3-lobed, reddish? inside. Male flowers: pedicel 4—6 mm long, bracteole about
the middle, caducous; buds subglobose or depressed broadly obovoid, 3 by 3 mm, cleft
c. 4/5, lobes 0.6—0.7 mm thick; staminal disc flat to faintly concave, subcircular, 1.5 mm
in diameter; anthers 6 or 7, just stiped, horizontal, 0.5 mm long, not touching; androphore
slender, 1 mm long. Female flowers not seen. Fruits 1 or 2 per infructescence (+ imma-
ture), ellipsoid, 1.5—2 cm long, with dense ferruginous hairs 0.5(—1) mm long; dry pericarp
1.5 mm thick; fruiting pedicel 5 mm long, scar of bracteole median.

Distribution — Malesia: Philippines (Luzon: Prov. of Nueva Ecija & Zambales).

Habitat & Ecology — Lowland and montane forest; possibly on ultrabasic; up to
c. 850 m altitude; fl. Feb.; fr. May.

Note — Knema alvarezii is known from only a few collections. It seems related to the
East Malesian K. tomentella, in which it was included by Sinclair (Gard. Bull. Sing. 18,
1961, 277). The latter species differs in its earlier glabrescent lower leaf surface, the
somewhat smaller male flowers with 8—11 anthers, and the subapical bracteole.

244 Flora Malesiana, Ser. I, Vol. 14 (2000)

2. Knema andamanica (Warb.) W.J. de Wilde

Knema andamanica (Warb.) W.J. de Wilde, Blumea 25 (1979) 370. — Knema glauca Blume var.
andamanica Warb., Mon. Myrist. (1897) 596. — Knema cinerea (Poir.) Warb. var. andamanica
(Warb.) J. Sinclair, Gard. Bull. Sing. 18 (1961) 174, f. 4, p.p. — Lectotype: King’s coll. s.n. (K),
Andaman I.

subsp. nicobarica (Warb.) W.J. de Wilde

Knema andamanica (Warb.) W.J. de Wilde subsp. nicobarica (Warb.) W.J. de Wilde, Blumea 25
(1979) 372. — Knema glauca Blume var. nicobarica Warb., Mon. Myrist. (1897) 596. —
Lectotype: King’s coll. 536 (L), Nicobar I.

Tree 6-16 m. Twigs 1.5—3(—5) mm in diameter, early glabrescent, at first with dense
grey or yellowish brown hairs 0.1 mm long or less; bark finely striate, not tending to
crack or flake. Leaves chartaceous, oblong to lanceolate, 8-30 by 2—8.5 cm, base attenu-
ate to rounded, apex subobtuse or acute(-acuminate); upper surface greenish brown,
lower surface faintly papillate, with + dense stellate hairs 0.1—-0.4 mm (in subsp.
andamanica, not in Malesia, mixed with longer dendroid hairs or not), subpersistent or
late glabrescent, the coarser hairs early shed and leaving minute scars; dots absent; midrib
flat or slightly raised above; nerves 12-24 pairs, thin and faint; venation (very) fine,
areoles 0.5 mm diameter or less, distinct above; petiole 8—20 by 1.5—-3 mm. Inflores-
cences: sessile or pedunculate up to 5 mm, brachyblast wart-like, simple or forked, 2-8
mm in diameter, in male 3—15(—30)-flowered, in female 2—6-flowered; flowers with dull
yellowish or (grey-)brown scurfy hairs 0.1—0.2(—0.3) mm or less, perianth 3-lobed, red-
dish inside. Male flowers: pedicel 3-6 mm long, bracteole subapical, subpersistent or
caducous: buds obovoid, 2.5-4.5 by 2-3 mm, cleft 1/2—2/3, lobes 0.5—0.7 mm thick;
staminal disc + flat or convex, circular, 0.8—1.2 mm in diameter; anthers 6 or 7, half-
sessile, suberect, 0.3-0.5 mm, nearly touching; androphore slender, 1-2 mm long. Fe-
male flowers: pedicel 5—6 mm long, bracteole subapical; buds ovoid-oblong, 6 by 2.5
mm, cleft about halfway, lobes 0.4 mm thick; ovary ovoid, 2 by 1.5 mm; style including
stigma slender, 1.5 mm long, stigma + deeply 2-lobed and each lobe shallowly 2—4-
lobulate at the apex. Fruits 1 or 2 per infructescence, ellipsoid, apex rounded to faintly
pointed, 1.8—2 by 1.2—1.6 cm, with rusty or rufous hairs 0.2-0.5 mm long; dry pericarp
1.5 mm thick; fruiting pedicel 7 mm long.

Distribution — India (Nicobar I.), Peninsular Thailand; in Malesia: Sumatra (Aceh);
Peninsular Malaysia (Penang I.).

Habitat & Ecology — Lowland and lower montane forest; 0—1200 m altitude; fl. & fr.
Jan.—Dec.

Notes — 1. Besides K. andamanica subsp. nicobarica there are two more subspecies,
occurring outside Malesia: subsp. andamanica and subsp. peninsularis W.J. de Wilde,
both with somewhat larger male flower buds, 4.5—6 by 3—4.5 mm; subsp. andamanica
has 10—12 anthers, just stiped, and fruiting pedicels 0.5—4 mm long; subsp. peninsularis
has 9-11 anthers, half-sessile, its fruits unknown.

2. Perianth in S Thailand once recorded as light yellow inside, in Nicobar I. and N Su-
matra as red or reddish.

De Wilde — Myristicaceae (Knema) 245

3. Knema ashtonii J. Sinclair

Knema ashtonii J. Sinclair, Gard. Bull. Sing. 18 (1961) 162, f. 1, p.p., excl. Jaheri 6/1, f. 1A, C-E,
I, and description of male flowers; W.J. de Wilde, Blumea 25 (1979) 384; 41 (1996) 382; Tree
Fl. Sabah & Sarawak 3 (2000) 405. — Type: SAN 17386 (Smythies, Wood & Ashton), Sabah.

Tree 10-25 m. Twigs ridged or + angled, 4-6 mm in diameter, with dense chocolate,
rusty, or grey-brown scale-like hairs 0.1 mm long, glabrescent; bark striate, not tending
to crack or flake. Leaves chartaceous to thinly coriaceous, obovate-oblong to oblong-
lanceolate, 25—55 by 7-15 cm, apex acute-acuminate, base broadly or narrowly rounded,
subcordate, or (sub)attenuate; upper surface greenish brown, lower surface either gla-
brous or with sparse, minute greyish sessile stellate hairs (var. ashtonii) or with persist-
ent densely interwoven greyish, pale brown, or cinnamon stellate hairs 0.1—0.2 mm long,
silky to the touch (var. cinnamomea); dots absent; midrib slightly raised above; nerves
20-40 pairs, flat or + depressed above; venation (very) fine, often + trabeculate; petiole
late glabrescent, 10-30 by 4-7 mm. Inflorescences: sessile, brachyblast simple, up to 10
by 5 mm; in male 4—10-flowered, in female 1—S-flowered; flowers seemingly glabrous,
but with minute scale-like hairs less than 0.1 mm long (lens!); perianth 3-lobed, colour
inside not known. Male flowers: pedicel distinct, bracteole apical, caducous; buds (broad-
ly) obovoid, 5.5—8 mm long, cleft about 4/5, lobes hard-carnose, 1—1.5(—2) mm thick;
see further under the varieties. Female flower (Sinclair l.c.): pedicel S-6 mm long, brac-
teole apical; buds oblong, 5—6 by 4 mm; ovary ovoid, 3 by 2 mm; style | mm long, stig-
ma 2-lobed and each lobe again minutely 2-lobulate. Fruits 1-3 per infructescence,
obovoid-oblong or long-ellipsoid, obtuse to acute at both ends, apex often up to 5 mm
beaked, 5-8 by 3—4 cm, with dense chocolate hairs 0.2 mm long; dry pericarp 4 mm
thick; fruiting pedicel stout, (5—)10-30 mm long, bracteole scar median to apical, and in
addition a distinct collar-like scar of the perianth at the base of the fruit.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, Central E Kalimantan).

Habitat & Ecology — Primary forest; often along streams; 0—600 m altitude; fl. Mar.—
Apr.; fr. Jan.—Dec.

Note — Sinclair (1.c.) erroneously included Jaheri 611, the only male specimen known
to him when describing K. ashtonii. Since then, S 23449 (identical with the type) has
been collected, with good male flowers, quite different from those of Jaheri 611, which
is described as K. sericea.

KEY TO THE VARIETIES

la. Lower leaf surface glabrous or nearly so, and then only with sparse, weak, whitish,
stellate scale-like hairs. Fruiting pedicel 10—15(—20) mm long, the bracteole scar
above the middle. Male pedicel 9 mm long .................- a. var. ashtonii

b. Lower leaf surface densely brownish grey to cinnamon brown pubescent, silky to
the touch. Fruiting pedicel 20-30 mm long, the bracteole scar at about the middle.
ne ere CON DO 2 STM fa. ons on fo ann SA he een aa b. var. cinnamomea

a. var. ashtonii

Indumentum of twig apex, petioles, and fruits dull grey-brown, chocolate, or rusty
brown. Leaves beneath glabrous or with scattered minute, weak, grey or whitish sessile

246 Flora Malesiana, Ser. I, Vol. 14 (2000)

ea neater)

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De Wilde — Myristicaceae (Knema) QAT

stellate-scaly hairs; the lower leaf surface not silky to the touch. Male flowers: pedicel
9 mm long; buds 5.5 by 4.5—5 mm, androecium 2 mm long, somewhat tapering, staminal
disc flat or faintly convex, including anthers 3 mm in diameter, anthers 14 or 15, short-
stalked. Fruits 5—7.5(-8) by 3(—3.5) cm, apex subacute to obtuse; fruiting pedicel 10—
15(—20) mm long, the bracteole scar at or close to the apex.

Field-notes — Bark smooth, ochre. Sometimes with a few stilt-roots.

Distribution — Malesia: Borneo (Sarawak: most collections from 4th Div.; Brunei,
Sabah).

Habitat & Ecology — Preferably growing on alluvial soils along streams; 0-200 m
altitude.

Note — Certain smaller leaved specimens, if sterile or with young fruits, may be
reminiscent of K. woodii or K. luteola.

b. var. cinnamomea W.J. de Wilde

Knema ashtonii J. Sinclair var. cinnamomea W.J. de Wilde, Blumea 25 (1979) 386, f. 32 Tree El.
Sabah & Sarawak 3 (2000) 405. — Type: S 23449 (Haji Suib), Sarawak.

Indumentum on twig apex, petioles, and fruits chocolate, rusty brown, or cinnamon.
Leaves beneath with persistent indumentum silky to the touch, consisting of densely
interwoven sessile stellate-scaly hairs, brownish grey or cinnamon. Male flowers: pedi-
cel 20-25 mm long; buds 7-8 by 6 mm, androecium tapering, 1.5—2 mm long, disc sub-
triangular, flat, including anthers 3.5 mm in diameter; anthers 22-24, just stiped. Fruits
5-8 by 3—4 cm, apex obtuse; fruiting pedicel 20-30 mm long, the bracteole scar about
median. — Fig. 35.

Field-notes —Bark smooth, reddish to greyish brown, without furrows or flakes. Leaves
medium green, glossy above; cinnamon to grey-brown scaly beneath. Immature leaves
with golden indumentum. Fruits (yellowish) brown or orange; seeds (aril?) yellow.

Distribution — Malesia: Borneo (Sarawak, Central E Kalimantan).

Habitat & Ecology — Often along streams; sandy soil subject to flooding, basalt-
derived and rich clay soils; up to 600 m altitude.

4. Knema celebica W.J. de Wilde

Knema celebica W.J. de Wilde, Blumea 25 (1979) 464; 27 (1981) 233, f. 3. — Type: Boschproef-
station Cel./II-318 (L), Central Sulawesi.

Tree c. 10 m. Twigs sometimes slightly flattened, 2-4 mm in diameter, smooth, at first
with dense rusty hairs 0.5—1 mm long, glabrescent; bark smooth, not cracking or flaking.
Leaves coriaceous, oblong, 12—20 by 3—7 cm, apex truncate, base attenuate; upper sur-
face dark brown, with remnants of indumentum on the midrib, lower surface early gla-

Fig. 35. Knema ashtonii J. Sinclair var. cinnamomea W.J. de Wilde. a. Twig apex with leaves;
b. part of older twig with male inflorescences, flowers submature; c. opened mature male bud show-
ing androecium; note bracteole scar apically on pedicel; d. androecium; e. detail of lower leaf
surface showing dense indumentum; f. infructescence; note bracteole scar about halfway on the
fruiting pedicel [a—d: S 23449; e, f: Kostermans 10415]. — Scale bar for a, b, f = 2 cm; for c =
2.8 mm; for d = 1.4 mm; for e = 0.07 mm.

248 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 36. Knema celebica W.J. de Wilde. a. Habit of male flowering twig; b. male bud; c. opened
male bud showing the androecium; d. lobe of male bud seen from inside; e. androecium seen from
above; f. opened female bud showing pistil; g. twig with fruit; h. detail of lower leaf surface show-
ing minute blackish dots on the veinlets [a—e, h: van Balgooy 3847; f, g: van Balgooy 3947]. —
Scale bar for a = 2 cm; for b, c, f = 1.8; for d = 1 mm; for e = 0.5 mm; for g = 1 cm; for h = 1.65 mm.

brescent, greyish; dots present; midrib raised above; nerves c. 15 pairs, raised; venation
fine; petiole at first with dense indumentum, late glabrescent, 10-13 by 2-3 mm. Inflo-
rescences: peduncle 1—4 mm, brachyblast simple or forked, up to 10 mm long; in male
(5—)10-flowered, in female 1—4-flowered; flowers with persistent rusty hairs 0.5—0.7
(-1) mm long, the perianth 3-lobed, reddish inside. Male flowers: pedicel (0.5—)1-2 mm

De Wilde — Myristicaceae (Knema) 249

long, bracteole + persistent, apical; buds subglobose, 2.5—3 mm in diameter, cleft 2/3—
4/5, lobes 0.5 mm thick; staminal disc flat, circular, 1.2—1.5 mm in diameter; anthers
6-9, distinctly stiped, horizontal, 0.2—0.3 mm, spaced; androphore 0.5—0.7 mm long.
Female flowers: pedicel 0.5—1.5 mm long, bracteole minute, persistent, median to sub-
apical; buds ellipsoid-oblong, 5-7 by 2-3 mm, cleft slightly over half, lobes at sutures
0.4 mm thick; ovary ellipsoid, 2.5-3 by 2—2.5 mm, with dense hairs 1—1.5 mm long;
style 1 mm long, stigma 1 mm long, erect, 2-lobed and each lobe again deeply 2—5-lobu-
late. Fruits 1 (or 2) per infructescence, ellipsoid-fusiform, 2—2.2 by 1.2—1.3 cm, + acute
at both ends, with dense rusty hairs 1-1.5 mm long; dry pericarp 1.5 mm thick; fruiting
pedicel 2 mm long. — Fig. 36.

Distribution — Malesia: Sulawesi (Central: Malili-Oesoe; N part of SE Peninsula:
near Soroako).

Habitat & Ecology — Lowland forest; forest on ultrabasic rock; 100—600 m altitude;
fl. Feb.—Apr. & July; fr. July.

Notes — 1. Although a member of the group of species with dotted leaves, K. celebi-
ca is obviously also related to K. tomentella; the latter differs in having sessile inflores-
cences, striate twigs, chartaceous (not coriaceous) leaves with a less prominent reticula-
tion above, and a+ late glabrescent lower leaf surface without dots.

2. The leaf apices are very typically (variously) truncate.

5. Knema cinerea (Poir.) Warb.

Knema cinerea (Poir.) Warb., Mon. Myrist. (1897) 611, t. 25; J. Sinclair, Gard. Bull. Sing. 18 (1961)
169, f. 3A—G (p.p., for var. cinerea only); W.J. de Wilde, Blumea 25 (1979) 457; 32 (1987) 129.
— Myristica cinerea Poir., Dict. Encycl. Suppl. 4 (1816) 35; Spreng., Syst. 3 (1826) 65; A. DC.,
Prodr. 14, 1 (1856) 207; Mig., Fl. Ind. Bat. 1, 2 (1858) 72. — Type: Labillardiére s.n., Pulau
Bouton, SE off Sulawesi.

Myristica peltata Roxb., Fl. Ind. 3 (1832) 846; A. DC., Prodr. 14, 1 (1856) 207; Miq., FI. Ind. Bat.
1, 2 (1858) 72. — Knema peltata (Roxb.) Warb., Mon. Myrist. (1897) 599. — Type: Roxburgh
s.n., 1802, culta.

[Myristica caesia Zipp. ex Span., Linnaea 15 (1841) 346 (Icon. 15), nom. nud.]

Tree 4-15 m. Twigs 1—3.5 mm in diameter, at first with hairs 0.1 mm long or less, gla-
brescent; bark striate, not tending to crack or flake. Leaves membranous or chartaceous,
elliptic to oblong-lanceolate, 8-25 by 2.5—9 cm., apex acute-acuminate, base rounded to
attenuate; upper surface olivaceous, lower surface greyish, with persistent dense or sparse
greyish scale-like hairs 0.1 mm long or less (lens!); dots absent; midrib flat to faintly
raised above; nerves 10-25 pairs, raised above; venation distinct above; petiole 8-20
by 1-2.5 mm. Inflorescences: peduncle up to 3(—5) mm, brachyblast simple or + warted,
to 5 mm long; in male 5—10-flowered, in female 1—5-flowered; flowers with persistent
greyish to rusty hairs 0.1 mm long or less; perianth 3-lobed, pink inside. Male flowers:
pedicel 6-11 mm long, bracteole caducous, at or somewhat above halfway; buds glo-
bose, 3.5—4.5 mm in diameter, cleft nearly to the base, lobes 0.7—1.2 mm thick; staminal
disc flat or nearly so, circular, 1.8—2.5 mm in diameter; anthers 9-14, just to distinctly
stiped, horizontal, 0.3-0.5 mm, spaced; androphore 0.5—1.5 mm long. Female flowers:
pedicel 3-13 mm long, bracteole caducous, submedian; buds ovoid-ellipsoid to oblong,
4-6 mm long, cleft just over halfway; ovary ovoid, 2 by 1.5-2 mm; style 0.5 mm long,

250 Flora Malesiana, Ser. I, Vol. 14 (2000)

stigma suberect, 2-lobed and each lobe again (1- or) 2- or 3-lobulate. Fruits | or 2 per in-
fructescence, subglobose to + obovoid, apex obtuse, faintly pointed by style-remnant,
1.2-1.8 by 1-1.6 cm, with persistent hairs 0.1—0.2(—0.3) mm long; dry pericarp | mm
thick; fruiting pedicel 3-13 mm long.

Field-notes — Bark smooth, dark. Colour of the perianth pink inside (once recorded);
in the resembling K. glauca it is creamy.

Distribution — Malesia: Lesser Sunda Islands E of Bali (Lombok, Sumbawa, Flores,
Tanimbar), Sulawesi, Moluccas (Sula I., Banda), Philippines (one collection only, see
note 3).

Habitat & Ecology — Forest and scrub; 0—1000 m altitude; fl. & fr. Jan.—Dec.

Notes — 1. The lower leaf surface of specimens from the Lesser Sunda Islands is
densely scale-like hairy, whereas in Sulawesi the surface is more sparsely, fine-stellately
hairy.

2. Knema cinerea is related to K. stenophylla, K. patentinervia and K. glauca with
almost similar habit and flowers, the latter species mainly differing in the lower leaf
surface with very weak stellate hairs, early glabrescent, and larger glabrescent fruits.
Specimens of K. glauca from Bali have small fruits, 2.2 cm long, and may resemble
K. cinerea.

3. The only collection from the Philippines (Zamboanga Prov., Mindanao), FB 23328,
probably belongs to K. cinerea; it is in fruit, 1.8 by 1.3 cm, fruiting pedicel 12-13 mm
long, lower leaf surface with minute persistent scale-like hairs. The indumentum of the
resembling K. glomerata, common in the Philippines, consists of much softer stellate
hairs, which are either persistent or early shed; the flowers and twig apices in K. glomerata
have longer, woolly dendroid hairs, up to 0.5 mm long.

6. Knema communis J. Sinclair

Knema communis J. Sinclair, Gard. Bull. Sing. 16 (1958) 297, f. 10, pl. IX B; 18 (1961) 193, p.p..
excl. specimens from Borneo and Shah & Kadim 526 (female fl.) from Peninsular Malaysia;
W.J. de Wilde, Blumea 25 (1979) 461. — Type: SF 40522 (Sinclair), Singapore.

Tree to 20 m. Twigs often blackish, 1.5—2 mm in diameter, at first with scurfy rusty or
greyish hairs 0.1 mm long, glabrescent; bark finely striate, sometimes finely cracking,
not flaking. Leaves chartaceous or thinly coriaceous, oblong-lanceolate to lanceolate,
(6-)10—25 by (2—)2.5—5 cm, apex acute-acuminate, base cuneate, upper surface (green-
ish) brown above, lower surface grey-glaucous, with subpersistent dense rusty to grey-
ish hairs 0.1—0.2 mm long, or late glabrescent, with distinct hair scars; dots absent; midrib
raised above; nerves 15-22 pairs, slightly raised above; venation very fine; petiole 7-20
by 1-3 mm. Inflorescences: sessile, brachyblast slender, simple or 2- to 3-fid, up to 10
mm long; in male 3—10-flowered, in female 1—3-flowered; flowers with persistent rusty
hairs 0.1 mm long, perianth 3-lobed, pink or reddish inside. Male flowers: pedicel 2—5
mm long, bracteole caducous, at the middle or somewhat above; buds globose, 2.5—3
mm in diameter, cleft to over 3/4, lobes 0.5—1 mm thick; the staminal disc flat, circular,
1.5 mm in diameter, whitish; anthers 7-9, just stalked, almost horizontal, 0.3 mm long,
spaced; androphore | mm long, red. Female flowers: pedicel 3 mm long, bracteole above
the middle, + caducous; buds + obovoid, 4 by 3 mm, cleft to about halfway, lobes 0.5—1
mm thick; ovary + ovoid, nearly 2 mm long; stigma sessile, 2-lobed and each lobe

De Wilde — Myristicaceae (Knema) 251

irregularly serrate or subentire. Fruits | or 2 per infructescence, ellipsoid to obovoid,
1.5-1.8 by 1-1.3 cm, with rusty hairs 0.1 mm long; dry pericarp | mm thick; fruiting
pedicel S—10(—15) mm long.

Field-notes — Bole fluted at base; bark smooth or nearly so, or sparingly flaky; inner
bark pink or red; sapwood white or yellowish. Leaves glossy dark green above, whitish
green or glaucous with brownish midrib and veins beneath. Flowers pink inside, sweet
scented when crushed. Fruits greenish yellow or orange, with rusty or reddish indu-
mentum.

Distribution — Malesia: Peninsular Malaysia (Perak, Trengganu, Pahang, Selangor,
Johore), Singapore.

Habitat & Ecology — Lowland forest; 0-300 m altitude; fl. & fr. Jan.—Dec.

Notes — 1. Knema communis resembles K. glaucescens, a species also with subper-
sistent indumentum on the lower leaf surface. The resembling K. malayana has larger
and longer pedicelled male flowers, with 9-13 anthers, and a different indumentum on
the lower leaf surface.

2. Apparently the species is restricted to Peninsular Malaysia. The records for Borneo
mentioned by Sinclair (Gard. Bull. Sing. 18, 1968, 163) are now referred to K. glauces-
cens. Shah & Kadim 526 (Peninsular Malaysia), after which Sinclair drew the emended
description of the female flowers of K. communis, is now referred to K. glaucescens as
well.

7. Knema conferta (King) Warb.

Knema conferta (King) Warb., Mon. Myrist. (1897) 578, t. 24; Gamble, Mat. Fl. Malay Penins. 5,
23 (1912) 243; Ridl., Fl. Malay Penins. 3 (1924) 70; J. Sinclair, Gard. Bull. Sing. 16 (1958) 286,
f. 6; 18 (1961) 194; W.J. de Wilde, Blumea 25 (1979) 472; Tree Fl. Sabah & Sarawak 3 (2000)
406. — Myristica conferta King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 315, pl. 150. — Lectotype:
King’s coll. 6211, Peninsular Malaysia, Perak.

Tree 10-25 m. Twigs 2—5 mm in diameter, at first with dense rusty hairs 0.2—1 mm
long, glabrescent; bark coarsely striate, not tending to crack or flake. Leaves chartaceous
to coriaceous, oblong to lanceolate, 10-30 by 3-12 cm, base subcordate to cuneate, apex
subobtuse or acute-acuminate; above glossy (greenish) brown, with distinct hair scars
above or not; lower surface greyish or glaucous, finely papillate, and with scattered mi-
nute brownish black dots especially on the veinlets (lens!), with persistent indumentum
of mixed sessile-stellate and stalked stellate-dendroid hairs of variable size, (0.2—)0.3—-1
mm long; midrib raised above; nerves 12-28 pairs, + raised; venation + fine, distinct;
petiole 5—15 by 1.5—5 mm, late glabrescent. Inflorescences: peduncle up to 1 mm, brachy-
blast simple or 2- (to 3-)furcate, up to 15 mm long; in male (5—)10—40-flowered, in
female up to 10-flowered; flowers with dull rusty hairs (0.1—)0.2—0.3 mm long, perianth
3-lobed, greenish or yellowish inside. Male flowers: pedicel 7-15 mm long, bracteole
+ persistent or caducous, about the middle or above; buds subglobose, often + triangular,
4—4.5 mm in diameter, cleft nearly to the base, lobes 1-1.5 mm thick; staminal disc flat
to slightly convex, circular, or + triangular, 2-3 mm in diameter; anthers (10—)12-17,
(just) stiped, horizontal, 0.3-0.7 mm long, spaced; androphore 0.8—1.5 mm long. Fe-
male flowers: pedicel 5-10 mm long, bracteole about the middle or above; buds obovoid,
5 mm long, cleft about halfway, lobes 1—-1.5 mm thick; ovary subglobose to ovoid, 1.5—

252 Flora Malesiana, Ser. I, Vol. 14 (2000)

2 mm in diameter; style 0-1 mm long, stigma 3- or 2-lobed and each lobe again 2—5-
lobulate. Fruits 1-5 per infructescence, subglobose, ellipsoid or obovoid, (1.5—)2—4 by
(1—)2-3 em, hairs 0.5 mm long, later on largely shed; dry pericarp 2-5 mm thick; fruit-
ing pedicel 5-10 mm long.

Field-notes — Bole straight, with buttresses to 3 m high, extending 30 cm over the
ground, 10 cm thick (once); bark smooth, + rough, or often peeling off in narrow strips
or in isodiametric scales 1—1.5 mm thick; living bark 5-8 mm thick, light red or red-
brown inside; sapwood whitish, yellowish, or reddish brown, heartwood reddish brown.
Leaves glossy dark green above, glabrous except midrib which becomes glabrous later,
beneath whitish green with stellate indumentum to varying degree. Flowers greenish or
yellowish inside, with a brown-pink blotch at base of lobes; stigma red (?). Fruits rusty
or orange.

Distribution — Malesia: Sumatra (Tapanuli, E Coast, Indragiri, Djambi, Palembang,
Bangka, Billiton, Riau Arch.), Peninsular Malaysia (Perak, Pahang, Malacca, Johore),
Singapore, Borneo (Sarawak, Sabah, W, E, & NE Kalimantan, Nunukan Is.).

Habitat & Ecology — Forest, including inundated and swamp forest; 0-600 m alti-
tude; fl. Jan.—Dec.; fr. mainly Nov.—Mar.

Note — Knema conferta is a homogeneous species which, in the vegetative state,
may be confused with K. laurina, K. oblongata, K. scortechinii, or K. pubiflora, all dif-
ferent in details of male flowers and indumentum; K. pubiflora is similar in the dotted
lower leaf surface with almost identical indumentum, but the perianth is hairy inside.
The fruits are variable, in size as well as in shape. They are more or less ellipsoid, but
some specimens from Nunukan Is. and the Leila Forest Reserve (Sabah) differ by their
nearly globose fruits, 2.5—3.5 cm in diameter, with conspicuously thick (4-5 mm) dry

pericarp.

8. Knema curtisii (King) Warb.

Knema curtisii (King) Warb., Mon. Myrist. (1897) 567, t. 25; J. Sinclair, Gard. Bull. Sing. 16 (1958)
281. f. 4: 18 (1961) 196, f. 6, 7 (excl. var. linguiformis); W.J. de Wilde, Blumea 25 (1979) 416,
f. 9: Tree Fl. Sabah & Sarawak 3 (2000) 406. — Myristica curtisii King, Ann. Roy. Bot. Gard.
Calc. 3 (1891) 326, pl. 167. — Lectotype: Scortechini 292, Peninsular Malaysia, Perak.

For more references and synonyms see the varieties.

Shrub or tree 2—10(—35) m. Twigs sometimes faintly 2- or 3-angled, (0.5—)1—2 mm in
diameter, yellow to brownish, at first with indumentum composed of greyish hairs 0.1
mm long or less, early glabrescent; bark finely striate, not cracking nor flaking. Leaves
membranous or chartaceous, of variable shapes, elliptic, obovate, or ovate to lanceolate,
3.5—15(—20) by 1.2—4(—6.5) cm., base cuneate to attenuate, rarely + rounded, apex acute
(-acuminate), sometimes blunt or rounded; greenish or dark brown above; on lower sur-
face greyish, very finely papillate, at first with dense hairs 0.1 mm long, early glabrescent;
dots absent: midrib raised above, nerves (6—)8—14(—20) pairs, flat to slightly raised above;
venation (very) fine, distinct above or not; petiole 5-20 by 0.7—1.5 mm. Inflorescences:
peduncle up to 1 mm, brachyblast simple or forked, up to 5 mm long, in male (2—)5—15-
flowered, in female 1—3(—5)-flowered; flowers with persistent or partially caducous grey-
brown or rusty hairs 0.1—0.3 mm long, perianth 3- (or 4-)lobed, cream to pink inside.
Male flowers: pedicel 5—11 mm long, bracteole subpersistent, median, or considerably

De Wilde — Myristicaceae (Knema) 258

Fig. 37. Knema curtisii (King) Warb. — a. var. curtisii: part of branchlet with male inflorescences.
— b-d. var. paludosa J. Sinclair: b. twig with male inflorescences; c. (sub)mature male bud;
d. androecium [a: Achmad 93; b—d: SF 26155 (Corner)]. — Scale bar for: a, b = 2 cm; ¢ = 3.3 mm;
d=0.85 mm.

below or above; buds subglobose or broadly obovoid, apex often depressed, in cross
section circular, obtusely or sharply angled (see the note under the type variety), 2.5—5
(—5.5) mm in diameter, cleft 2/3—4/5, lobes 0.5(—0.8) mm thick; staminal disc flat, slightly
concave, or with a small minute mammilla, subtriangular, 1.7—3 mm in diameter; an-
thers 9—25(—30) (i.e., with 18-50 thecae), completely sessile, 0.2—0.4 mm, closely ap-
pressed, erect or horizontal, hidden under the rim of the staminal disc or partially emer-
gent; androphore slender or broad and tapering, 1-2 mm long. Female flowers: pedicel
5-10 mm long, bracteole about the middle; buds ellipsoid to obovoid or subglobose,
4-5 by 2.5—4 mm, cleft 1/2—2/3, lobes 0.7—1 mm thick; ovary ovoid, 1.5—3 mm long;
style up to 0.5 mm long, stigma 6—8-lobulate, i.e., 2-lobed and each lobe again 3—4(—5)-
lobulate. Fruits 1 or 2 per infructescence, + ellipsoid, ridged, apex acute, base obtuse, or
often up to 5 mm narrowed (beaked) at both ends, 2-5 by 1.5—3 cm, with greyish to rusty
hairs 0.1-0.2 mm long, or glabrescent; dry pericarp 1—2.5 mm thick; fruiting pedicel
8—16 mm long. — Fig. 37.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Borneo.

Note — A complex species in which four varieties are recognized. Sinclair accepted
five varieties, but his var. linguiformis is now treated as a separate species. The remain-
ing four varieties display a considerable difference in general habit, distribution area and
habitat.

254 Flora Malesiana, Ser. I, Vol. 14 (2000)

Knema curtisii and the related K. linguiformis and K. viridis can easily be distinguished
within Knema by the androecium with many tightly appressed anthers. The androecium
of K. curtisii has been described by previous authors as having 30—45 anthers; in fact
there are only 9—25(—30) anthers of which the thecae are completely sessile and closely
set and touching, and thus appearing to have twice as many anthers as there actually are.

KEY TO THE VARIETIES

1a. Leaves membranous, 5—15(—20) cm long, drying greenish, apex acute or acuminate;
nerves and venation distinct at both sides. Male buds tending to become partially
glabrescent, with blunt or sharp angles. — Lowland forest. Sumatra, Peninsular Ma-
laysia (anthers 17-25), Borneo (anthers 9-15) .........---++++5 a. var. curtisii

b. Leaves membranous or chartaceous, 3.5—10 cm long, drying + greenish brown to
dark brown above; apex acute, acuminate, obtuse, or rounded; nerves and venation
indistinct. Male buds with persistent indumentum, or partially glabrescent. Anthers

1S 25 (BON ate Sep ces: evened sigh dee ent ie a eecaiie e 2

2a. Male buds with minute mainly sessile stellate hairs, tending to be partly shed or easi-
ly rubbed off; angles sharp. Leaves drying dark brown above; apex obtuse, rounded,

or subacuminate withobtuse tip’ o......-.. Se. - ages ole > es ee 3

b. Male buds with persistent indumentum of mixed sessile-stellate and dendroid hairs;
angles rounded. Leaves drying + greenish brown above; apex acute to acuminate. —

Ridges in forest, often on sandy soils. Borneo .......-----+++ c. var. arenosa

3a. Leaves membranous or chartaceous, elliptic to oblong. — Usually in lowland marshy
forest. Sumatra, Peninsular Malaysia, Borneo ............--. d. var. paludosa

b. Leaves chartaceous, lanceolate, with nearly parallel sides. — Ridges in lowland for-
este BOMeOMBIUMEL) ooo 12: cs eee ee eee ye ae see ees enone em b. var. amoena

a. var. curtisii

Knema curtisii (King) Warb., Mon. Myrist. (1897) 567, t. 25; Gamble, Mat. FI. Malay Penins. 5, 23
(1912) 240; Ridl., Fl. Malay Penins. 3 (1924) 69; J. Sinclair, Gard. Bull. Sing. 16 (1958) 281,
f. 4: 18 (1961) 196, f. 6, 7 (excl. var. linguiformis);, W.J. de Wilde, Blumea 25 (1979) 418, f. 9a;
Tree Fl. Sabah & Sarawak 3 (2000) 407. — Myristica curtisii King, Ann. Roy. Bot. Gard. Calc.
3 (1891) 326, pl. 167.

Myristica sp. Hook. f., Fl. Brit. India 5 (1886) 113 (Maingay, Kew Distr. 1301).

Stem without stilt-roots, sometimes with buttresses. Leaves membranous, drying green-
ish, elliptic, obovate, or oblong-lanceolate, broadest at or + above the middle, in Borneo
sometimes + parallel-sided, S—15(—20) cm long, base cuneate or attenuate, apex blunt to
acute, or acuminate with blunt or acute tip; nerves (6—)8—16(—18) pairs, nerves and ve-
nation distinct at both surfaces. Male flowers: buds in cross section either with obtuse
angles (Peninsular Malaysia, some, and all specimens from Sumatra and Borneo), or
with sharp angles (most specimens from Peninsular Malaysia), 2.5—4 mm in diameter,
with mixed stellate and dendroid hairs 0.1—0.3 mm long, partly tending to be shed; an-
thers either 17-25 (in Peninsular Malaysia, the anthers + horizontal, under the rim of the
disc), or 9-15(—18) (in Borneo and Sumatra, with the anthers more laterally on the disc,
suberect). Stigma 6—8-lobulate. Fruits scurfy, 2—4.5 cm long, ridged; fruiting pedicel
8—16 mm long. — Fig. 37a.

De Wilde — Myristicaceae (Knema) 255

Field-notes — Bole sometimes with buttresses up to 50 cm high; bark smooth, with
scattered small dents, or + fissured; slash inner bark red-brown or yellow, slash wood
white; exudate pale reddish, once yellow. Leaves glossy dark green above, glaucous
with yellowish green midrib beneath. Tepals pinkish or red inside. Fruits yellow, brown,
orange-brown, or red, with light brown powdery scale-like hairs, seeds grey-white.

Distribution — Malesia: Sumatra (E Coast, Palembang, Simeulué I.), Peninsular Ma-
laysia (Perak, Kelantan, Trengganu, Pahang, Selangor, Negri Sembilan, Malacca, Johore,
Penang L., P. Tioman), Borneo (W, C, E Kalimantan, Sarawak, Brunei, Sabah).

Habitat & Ecology — Primary and degraded forest, hill slopes; on a variety of soil
types: clay, loam, sandy clay, sandstone, lime-containing soils; 0O—1000 m altitude; fl. &
fr. Jan.—Dec.

Notes — 1. The male buds (when dry) are variable in shape, size, and number of
anthers as given in the description.

2. Knema curtisii, particularly var. curtisii, resembles K. membranifolia, a species
also with yellowish twigs, but generally more distinctly angular, and with the flowers on
the older wood, below the leaves. In K. curtisii the inflorescences are found among the
leaves.

3. Sterile specimens, or material of which the androecium is not examined, may be
confused with K. globularia.

b. var. amoena J. Sinclair

Knema curtisii (King) Warb. var. amoena J. Sinclair, Gard. Bull. Sing. 18 (1961) 198, f. 7; W.J. de
Wilde, Blumea 25 (1979) 420. — Type: Sinclair (& Kadim) 10442, Brunei.

Stem without stilt-roots. Leaves chartaceous or coriaceous, drying dark brown, ob-
long to lanceolate, nearly parallel-sided, 5-8 by 1—1.8 cm, base cuneate to attenuate,
apex obtuse; nerves 10-20 pairs, nerves and venation indistinct at both surfaces. Male
flowers: buds with sharp angles, 4-5 mm in diameter, with hairs 0.1 mm long or less,
tending to become glabrescent or apparently so; creamy inside; anthers 16 or 17, almost
horizontal, just under the rim of the staminal disc. Female flowers and fruits not seen. —
Fig. 34b.

Field-notes — Tree c. 20 m; bark pale grey, smooth; sap pink, not very copious. Leaves
medium green and glossy above, paler and glaucous beneath, midrib paler. Flowers fra-
grant when crushed, cream inside when fully open.

Distribution — Malesia: Borneo (Brunei, known only from the type).

Habitat & Ecology — Ridge in lowland forest; fl. Aug.

Note — This variety is readily distinguishable by its leaf shape, and possibly de-
serves the status of a separate species.

c. var. arenosa J. Sinclair

Knema curtisii (King) Warb. var. arenosa J. Sinclair, Gard. Bull. Sing. 18 (1961) 198, f. 6D; W.J. de
Wilde, Blumea 25 (1979) 420; Tree Fl. Sabah & Sarawak 3 (2000) 408. — Type: Kostermans
8653, Nunukan Is.

Tree 6-35 m, without stilt-roots. Leaves membranous or chartaceous, drying brown
or some greenish brown, elliptic to lanceolate, broadest at or below the middle, or more
or less parallel-sided, 3.5—-13 cm long, base cuneate to attenuate, apex acute or acumi-

256 Flora Malesiana, Ser. I, Vol. 14 (2000)

nate, rarely obtuse; nerves 12-18 pairs, nerves and venation indistinct at both surfaces.
Male flowers: buds with rounded angles, 3-5 mm in diameter, with persistent minute
mixed sessile-stellate and dendroid hairs 0.1—0.3 mm long; anthers 18—23, suberect to
almost horizontal, at or just under the rim of the + triangular staminal disc. Female
flowers and fruits not seen.

Field-notes — Trunk slender; buttresses to 50 cm high; bark grey, pale brown, or
straw, smooth with scattered very small dents; inner bark 10 mm thick, (brown-)red,
with copious red latex; wood white, yellow, or brown. Flowers brown or light green, fra-
grant when crushed.

Distribution — Malesia: Borneo (S, SE, E, NE Kalimantan, Nunukan Is., Sarawak).

Habitat & Ecology — Ridges in forest; found on sandy loam and sandstone; 0-600 m
altitude; fl. Jan.—Dec.

Note — This variety is characterized by small chartaceous leaves with scarcely vis-
ible nerves and venation.

d. var. paludosa J. Sinclair

Knema curtisii (King) Warb. var. paludosa J. Sinclair, Gard. Bull. Sing. 18 (1961) 201, f. 6A, E;
W.J. de Wilde, Blumea 25 (1979) 420, f. 9b—d; Tree Fl. Sabah & Sarawak 3 (2000) 408.
Type: SF 26155 (Corner), Singapore.

Stilt-roots usually present. Leaves membranous or chartaceous, drying brown, ellip-
tic or (obovate-)oblong, broadest at or above the middle, 4-10 cm long, base cuneate or
attenuate, apex either subacute or sometimes faintly acuminate with blunt tip, obtuse or
rounded; nerves 6-16 pairs, nerves and venation faint on both surfaces. Male flowers:
buds with sharp angles (always?), 3-5 mm in diameter, with mixed sessile-stellate and
dendroid hairs 0.1—0.2 mm long, partially tending to become glabrescent; anthers 20—
30, nearly horizontally under the rim of the triangular staminal disc. Stigma 8-lobulate
(i.e., 2-lobed and each lobe again 4-lobulate, these often again with smaller incisions).
Fruits (fide Sinclair) ridged on the line of suture, 4—5 cm long; fruiting pedicel 10 mm
long, thicker than in var. curtisii. — Fig. 37b—-d.

Distribution — Malesia: Sumatra (fide Sinclair: Tapanuli, Palembang), Peninsular
Malaysia (Perak, Johore), Singapore, Borneo (Sarawak).

Habitat & Ecology — Primary lowland forest, clearings; preferably marshy forest,
and then with stilt-roots; 0-200 m altitude; fl. June—Oct.

Note — Apparently predominantly a freshwater swamp forest species. Trees usually
with stilt-roots. Furthermore characterized by leaves smaller than in the type variety,
elliptic-oblong to obovate, with obtuse apex, drying brown, and with faint nerves and
reticulation.

9, Knema elmeri Merr.

Knema elmeri Merr., Univ. Calif. Publ. Bot. 15 (1929) 75; J. Sinclair, Gard. Bull. Sing. 18 (1961)
202. f. 8: W.J. de Wilde, Blumea 25 (1979) 383; Tree Fl. Sabah & Sarawak 3 (2000) 408. —
Type: Elmer 21527, Sabah.

Tree 5-15 m, sometimes with stilt-roots. Twigs + angled, 2—4(—5) mm in diameter, at
first with rusty hairs 0.1—0.2 mm long, glabrescent; bark grey-brown, tending to crack

De Wilde — Myristicaceae (Knema) 257

longitudinally, later on flaking. Leaves chartaceous, elliptic to oblong(-oblanceolate),
12-32 by 4-13 cm, base rounded to acute, apex acute-acuminate; drying greenish or
light brown above, early glabrescent, lower surface with persistent, silvery to pale brown,
dense interwoven hairs 0.1—0.2 mm long, silky to the touch; midrib sunken to prominent
above; nerves 12—20(—25) pairs, (slightly) raised or flat above; venation distinct above;
petiole 8-18 by 24 mm. Inflorescences: sessile, brachyblast simple or irregularly warted,
up to 8 mm in diameter, in male 5—25-flowered, in female 2—7-flowered; flowers with
dense pale brown to rusty hairs 0.1 mm long, perianth 3-lobed, red or purple inside.
Male flowers: pedicel 7-14 mm long, bracteole subpersistent, about median; buds broadly
ovoid to depressed globose, often + sagged at base, 3—4 by 4—4.5 mm, cleft nearly to the
base, lobes 0.5—0.8 mm thick; staminal disc circular, distinctly convex in the centre,
22.5 mm in diameter; anthers 10-14, just sessile or stalked, horizontal, 0.3—0.6 mm,
not touching; androphore 0.5—0.8 mm, clasped by the somewhat thickened perianth base.
Female flowers: pedicel 1-2 mm long, bracteole about median; buds + ellipsoid, 6 mm
long, cleft about halfway, lobes 1 mm thick; ovary ellipsoid, 3 mm long; style 0-1 mm
long, stigma 2-lobed and each lobe again shallowly 2—3(—4)-lobulate. Fruits 1-6 per
infructescence, obovoid to ellipsoid(-oblong), apex obtuse or shortly beaked, 2—2.5 by
1.1-1.6 cm, with rusty hairs 0.1—0.2 mm; dry pericarp 2 mm thick; fruiting pedicel 1—5
mm long. — Fig. 34d.

Field-notes — Crown rounded; bole sometimes with stilt-roots, no buttresses; bark
smooth, scaly, flaky, or cracky, hard; inner bark white, reddish, or red-brown; sapwood
brown. Leaves glossy dark green above, silvery beneath.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, NE Borneo: Nunukan Is.).

Habitat & Ecology — Primary and disturbed forest, riverine or mixed dipterocarp
forest; on rich or sandy clay soils, also limestone; 0—1500 m altitude; fl. & fr. throughout
the year.

Notes — 1. Knema elmeri is characterized by the silky indumentum on the lower leaf
surface, flaking bark of the twigs, + globose male flower buds, and a distinctly convex
staminal disc.

2. The nature of the indumentum of the lower leaf surface places K. elmeri along with
K. ashtoni, K. retusa, and K. sericea, but there is also a close alliance with K. latericia,
especially subsp. rid/eyi, on account of the cracking bark of the twigs, and the convex
staminal disc.

10. Knema emmae W.J. de Wilde

Knema emmae W.J. de Wilde, Blumea 41 (1996) 384: Tree Fl. Sabah & Sarawak 3 (2000) 409. —
Type: Amdjah 144, NE Kalimantan.

Treelet c. 4 m. Twigs 1.5—2 mm in diameter, yellowish brown, at first with hairs 0.5
(—1) mm, glabrescent; bark smooth or finely striate, not cracking nor flaking; lenticels
minute. Leaves thinly chartaceous, elliptic-oblong or oblong-lanceolate, 12—20 by 3.5—
7 cm, base short-attenuate, apex + blunt or (acute-)acuminate; upper surface drying green-
olivaceous; lower surface slightly paler, glabrous, not or but little papillose; dots absent;
midrib slender above, nerves 12-15 per side, flat or slightly raised above; venation fine,
distinct at both surfaces. Petiole 10 by 2 mm. Inflorescences: sessile or pedunculate to
1.5 mm, brachyblast wart-like, 1-2 mm long, pubescent, glabrescent; in male and fe-

258 Flora Malesiana, Ser. I, Vol. 14 (2000)

male 2—5-flowered: flowers with rusty hairs (0.2-)0.5—0.7 mm, perianth 3-lobed, col-
our inside not known. Male flowers: pedicel 6-7 mm long, slender, bracteole less than
| mm, subpersistent, slightly above median; buds + depressed globose, 3 by 4 mm, cleft
3/4(—4/5), lobes 0.5 mm thick; staminal disc circular, flat, 2 mm in diameter; anthers
14 or 15. sessile, 0.3 mm, not tightly contiguous, under the rim of the disc, androphore
tapering, narrow at base, 0.5—0.8 mm long. Female flowers: pedicel 4 mm; buds obovoid,
3.5 mm long; ovary not seen. Fruits 2-4 per infructescence (slightly immature), ovoid-
oblong, 2.5—3 by 1.5—1.6 cm, base + truncate or broadly rounded, apex narrowly rounded,
with stigma remnant, | mm, dry pericarp 0.5(—1) mm thick, with rusty mealy hairs 0.5
mm; fruiting pedicel 7-10 mm long, scar of bracteole + median.

Field-notes — Low tree: bole 2 m, total height 4 m; sapwood whitish; the bark dark
brownish, inner bark brownish with red latex. Fruits yellowish.

Distribution — Malesia: Borneo (Sabah: Keningau; NE Kalimantan).

Habitat & Ecology — Hill-side forest; probably at low altitudes; fl & fr. June.

Note — Knema emmae is obviously related to K. curtisti (vat. curtisii) and K. lingui-
formis. Knema curtisii is distinguishable by more slender twigs, 1(—1.5) mm in diam-
eter, and smaller leaves with more slender lateral nerves and petiole; the fruits are
+ similar in size (although variable) but early or late glabrescent. Knema linguiformis
often has a + rounded leaf base, fruits glabrous (early glabrescent), at base + contracted
towards a short fruiting pedicel. Knema emmae is noticeable especially for its male
flowers. with a circular staminal disc, with beneath 14 or 15 not tightly contiguous small
anthers: in the other two species mentioned here there are more anthers (although often
only 9-18 in Bornean specimens of K. curtisti var. curtisii), attached to a bluntly trian-
gular staminal disc.

11. Knema furfuracea (Hook. f. & Thomson) Warb.

Knema furfuracea (Hook. f. & Thomson) Warb., Mon. Myrist. (1897) 581, t. 24; Gamble, Mat. Fl:
Malay Penins. 5, 23 (1912) 245; Ridl., Fl. Malay Penins. 3 (1924) 70; Corner, Wayside Trees
(1940) 476; J. Sinclair, Gard. Bull. Sing. 16 (1958) 275, f. 2, pl. 1B; 18 ( 1961) 209; W.J. de Wilde,
Blumea 25 (1979) 387: 27 (1981) 223. — Myristica furfuracea Hook. f. & Thomson, Fl. Ind. 1
(1855) 159; Miq.. Fl. Ind. Bat. 1, 2 (1858) 70; Hook. f., FI. Brit. India 5 (1886) 112; King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 318, p.p., pl. 155, incl. var. major. — Type: Porter or Wallich in
Hb. Hooker s.n., Peninsular Malaysia.

Myristica longifolia auct. non Blume: Hook. f. & Thomson, FI. Ind. 1 (1855) 156, p.p.

Tree 8-25 m. Twigs sometimes faintly angular, 4—8(—10) mm in diameter, at first with
dense furfuraceous hairs 1—-1.5 mm long, soon rubbed off, glabrescent; bark striate or
not. lower down brown to blackish, conspicuously longitudinally cracking and flaking.
Leaves coriaceous, oblong or (ob)lanceolate, (10-)25-50 by (3-)5—21 cm, apex acute
(-acuminate), sometimes rather blunt, base narrowly to broadly cordate; drying (green-
ish) brown, glossy or dull above, lower surface early glabrescent, finely papillate; dots
absent: midrib stout, raised above; nerves 20—40(—50) pairs, raised above; venation raised
or sunken above; petiole 5-25 by 4-8 mm, late glabrescent. Inflorescences: sessile,
brachyblast simple or coarsely warted or furcate, up to 15 mm in diameter, in male 3—30-
flowered, in female up to 20-flowered; flowers with dense furfuraceous hairs 0.5—1 mm
long, subpersistent or glabrescent, perianth 3- or 4-lobed, red inside. Male flowers: pedi-

De Wilde — Myristicaceae (Knema) 259

cel 3-15 mm long, bracteole subpersistent or caducous, about median, sometimes to-
wards the apex; buds depressed obovoid or depressed globose, somewhat triangular,
4-5 by (4-)4.5-5 mm, cleft 3/4—4/5, lobes 1—1.5 mm thick; staminal disc circular or
faintly triangular, flat or slightly concave, 2—2.5 mm in diameter; anthers 10-14, just
sessile, oblique or nearly horizontal, 0.5—0.7 mm, not touching, androphore 1-2 mm
long, striate. Female flowers: pedicel 1.5—2 mm long, bracteole median to apical; buds
ellipsoid or obovoid, 6-8 by 5-7 mm, cleft c. 2/3, lobes 1.5 mm thick; ovary ovoid or
subglobose, 2.5—4 mm long; stigma subsessile, + flat, 2-lobed, each lobe with 5-9 lobe-
lets or serrations. Fruits 1-4 per infructescence, ovoid, subglobose, or obovoid, 2.5—3.5
by 2—2.8 cm, with rusty hairs 1-2 mm long; dry pericarp 3-5 mm thick; fruiting pedicel
2—5 mm long.

Field-notes — Crown dome-shaped; trunk once recorded with buttresses; bark smooth
with longitudinal flakes or scale-like hairs, or fissured; outer bark slash red, fibrous, in-
ner bark white, laminated or fibrous, cambium red; sapwood whitish, heartwood light
red to dark brown. Flower buds yellow, pink, or deep red inside; fruits red(-brown) or
apricot.

Distribution — Southern Peninsular Thailand and Malesia: Sumatra (sterile coll.),
Peninsular Malaysia (all provinces except Perlis and Wellesley Prov.), Singapore.

Habitat & Ecology — Primary and degraded rain forest; in S Peninsular Thailand in
evergreen forest; hillsides, ridges, ridge-tops over granite, limestone; on clay and black
soil; O—900 m altitude; fl. & fr. throughout the year.

Notes — 1. Knema furfuracea is related to K. hookeriana, K. lampongensis, K. lamel-
laria, and K. pallens; the leaves also resemble large-leaved specimens of K. latericia
and K. lunduensis, species with non-cordate leaf bases. The pyramidal shape of the tree,
and the flaking bark of K. furfuracea are similar in K. hookeriana.

2. The present circumscription of K. furfuracea is much narrower than that accepted
by Sinclair (1.c.). The Indochinese specimens cited by him are now referred to K. pierrei
Warb. (not in Malesia); specimens from Thailand have been described as K. tenuinervia
(not in Malesia); specimens cited from Sumatra have been described as K. lampongensis,
those from Borneo as K. pallens; some recent specimens from Peninsular Malaysia, also
resembling K. furfuracea, have been segregated as K. lamellaria.

12. Knema galeata J. Sinclair

Knema galeata J. Sinclair, Gard. Bull. Sing. 18 (1961) 211, f. 10; W.J. de Wilde, Blumea Pa t79)
422, f. 10; Tree Fl. Sabah & Sarawak 3 (2000) 409. — Type: S 4855 (Anderson & Md. Hasan),
Brunei.

Tree 5-30 m. Twigs stout, sometimes somewhat flattened or angular, 4-10 mm in
diameter, at first with rusty hairs 1-2 mm long, glabrescent, (blackish) brown; bark finely
striate, when older tending to crack or flake. Leaves thickly coriaceous, oblong or lance-
olate, 15—50 by 2.5—14 cm, base rounded, or subcordate or subcuneate, apex obtusish or
acute-acuminate; olivaceous, glossy above; lower surface at first with dense rusty in-
dumentum, early glabrescent, greyish to pinkish; dots absent; midrib much raised above;
nerves 20-35 pairs, raised above, distinctly interarching; venation faint above; petiole
12-30(—40) by 3-8 mm. Inflorescences: sessile, brachyblast simple or warted with up to
10 protuberances, up to 20 by 10 mm, in male (5—)10—50-flowered, in female 5—25-

260 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Knema) 261

flowered: flowers at first with hairs 0.4-1 mm long, glabrescent, perianth 3-lobed, white,
pink or red inside. Male flowers: pedicel (15—)20—30 mm long, bracteole 1-3 mm long,
subpersistent, about median; buds mitriform, triquetrous in cross section, 6-10 by 9-15
mm, cleft nearly to the base, lobes broad, reniform, 0.5(—1) mm thick; staminal disc flat
or + convex, circular or + triangular, 3-5 mm in diameter; anthers 14—23, just sessile,
oblong, 1-1.5 mm long, horizontal or + oblique, not touching; androphore 0.5—1 mm.
Female flowers: pedicel 12-25 mm long, bracteole about median; buds subovoid-
mitriform, narrower than in male, 7—9 by 5—7 mm, cleft nearly to the base, lobes 0.5—1
mm thick; ovary globose or ovoid, 3—3.5 by 2.5—3 mm; stigma sessile, 2-lobed and each
lobe again (deeply) 4—8-lobulate. Fruits (1—-)2—5 per infructescence, obovoid or pear-
shaped, often contracted towards the base, 2-5 by 1.5—3 cm, sometimes ridged, apex
sometimes beaked; pericarp 2-3 mm thick, at first with indumentum of hairs 0.5 mm
long, glabrescent; fruiting pedicel 15-30 mm long. — Fig. 38.

Field-notes — Bark smooth, finely flaky or fissured; inner bark brownish; sapwood
pink-yellow, pale brown, or whitish. Leaves very glossy green with whitish green midrib
above, glaucous with yellowish green midrib and veins beneath. Flowers with rusty in-
dumentum, pure white or reddish inside; stamens yellow; fruits dark yellow, orange, or
rusty brown.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, E Kalimantan).

Habitat & Ecology — Primary mixed lowland dipterocarp forest, hill-side and ridge
forest, open kerangas, and marshy forest; on sandy soils, on yellow podzolic soils, marshy
soil, yellow clay soil, and sandy loam; at low altitudes; fl. & fr. throughout the year.

Notes — 1. The stout inflorescences are ramiflorous on the older branches of 1—2 cm
diameter

2. § 12935 (Anderson) (in L) with mainly male flowers, has one female flower with
a normal pistil, but with the perianth similar to the male flowers.

13. Knema glauca (Blume) Petermann

Knema glauca (Blume) Petermann, Pflanzenreich (1838-1845) 294; W.J. de Wilde, Blumea 25
(1979) 452; 32 (1987) 126; Tree Fl. Sabah & Sarawak 3 (2000) 410. — Myristica glauca Blume,
Cat. (1823) 111; Bijdr. 2 (1825) 576. — Type: Blume s.n., W Java.

For more references and synonyms see the varieties.

Tree 5-30 m. Twigs (1-)1.5—3.5 mm in diameter, at first with grey or brown hairs 0.1
mm long or less, early glabrescent; bark coarsely striate, not tending to crack or flake.
Leaves membranous or chartaceous, elliptic to lanceolate, 6-20(—25) by 2—5.5(-11) cm,
apex obtuse or (acute-)acuminate, base + rounded to attenuate, greenish to (blackish)

Fig. 38. Knema galeata J. Sinclair. a. Twig with leaves; b. old male inflorescence with immature
flowers, on older wood; c. older branch with male inflorescence and (sub)mature flowers; d. mature
male bud; e. androecium; f. (sub)mature female bud; g. pistil; h. apex of ovary with sessile lobed
stigma, lateral view; i. young fruit; j. apex of young fruit with sessile lobed stigma; k. mature fruit
showing seed [a, b: Kostermans 7062; c: S 1177; d, e: S 4855; f—h: Ladi anak Bikas s.n. /27-2-1961;
i, j: Ladi anak Bikas s.n./6-1-1961; k: Ladi anak Bikas s.n./23-3-1961]. — Scale bar for a—c =
2 cm; for d, f, i, k = 10 mm; for e, h = 2 mm; for g = 5 mm; for j = 3.3 mm.

262 Flora Malesiana, Ser. I, Vol. 14 (2000)

brown, dull or glossy above; lower surface grey or pale brown, at first with scattered,
very weak, greyish (or pale brown) hairs 0.1 mm long or less, early glabrescent; dots ab-
sent: midrib little raised above; nerves 12—22(—25) pairs, raised to flat above; venation
coarse (areoles 0.5 mm in diameter or more), distinct above; petiole 7-20 by 1-3 mm.
Inflorescences: sessile or (in E Java and Borneo) with peduncle up to 3 mm, brachy-
blast simple or 2- or 3-fid, up to 15 mm long; in male (2—)5—20-flowered, in female 1-8-
flowered; flowers with persistent greyish brown scale-like hairs 0.1 mm long or less,
perianth 3- (or 4-)lobed, red or in Java and Bali also yellowish (?). Male flowers: pedi-
cel 4-10 mm long, bracteole 0.5—1 mm, persistent, median or above; buds subglobose,
2.53.5 mm in diameter, cleft c. 4/5, lobes 0.5—0.8 mm thick; the staminal disc flat to
faintly convex, sometimes finely papillate, circular, 1.5-2(—2.5) mm in diameter; an-
thers 8-10, or in Borneo, E Java, and Bali frequently 11-15, subsessile to stiped, spaced,
0.3-0.5 mm long, + horizontal, the connective broad or narrow above, the thecae open-
ing nearly laterally; androphore 0.5—1 mm long. Female flowers: pedicel 1-6 mm long,
bracteole somewhat above median; buds ovoid-ellipsoid or obovoid-oblong, 4-6 mm
long, cleft to about halfway, lobes at sutures 0.8—1.5 mm thick; ovary ovoid, 1.5—2.5 by
|-2 mm; style 0—0.5 mm long, stigma + flat to erect, 2-lobed and each lobe again 2—4-
lobulate. Fruits 1—4 per infructescence, subglobose to ovoid or ellipsoid, 1.8-3(—4”) by
1.4-1.8(—2.5?) cm, at first with short scale-like hairs 0.1 mm long or less, early glabrescent,
leaving a fine-granulate surface; dry pericarp 1.5—2.5 mm thick; fruiting pedicel 3-12
mm long.

KEY TO THE VARIETIES

la. Leaves membranous or chartaceous, elliptic to lanceolate. Fruiting pedicel 3—7(-10)

mm long; fruits drying greyish brown ...........-.+++++-0+05- a. var. glauca
b. Leaves membranous, oblong to lanceolate. Fruiting pedicel 8-12 mm long; fruits
re HMODROWM §.2 2 faked oe ac uiiees is an eue aoa em vel oa) eee = ee b. var. riparia

a. var. glauca

Knema glauca (Blume) Petermann, Pflanzenreich (1838-1845) 294; Warb., Mon. Myrist. (1897)
594, t. 25 (for var. typica only); W.J. de Wilde, Blumea 25 (1979) 452; 32 (1987) 1263 dnee Fi.
Sabah & Sarawak 3 (2000) 411. — Myristica glauca Blume, Cat. (1823) 111; Bijdr. (1826) 576;
Rumphia | (1835) 187, t. 60; Miq., Pl. Jungh. (1852) 171; King, Ann. Roy. Bot. Gard. Calc. 3
(1891) 320, pl. 157, p.p.; Koord. & Valeton, Med. Lands PI. Tuin 17 (1896) 189.

Myristica intermedia Blume var. minor Migq., Fl. Ind. Bat. 1, 2 (1858) 70. — Type: Teijsmann s.n.,
Java.

Myristica corticosa (Lour.) Hook. f. & Thomson var. lanceolata Migq.., Fl. Ind. Bat., Suppl. 1 (1861)
384. — Type: Junghuhn s.n., Sumatra, Tapanuli.

Myristica palembanica Migq., Fl. Ind. Bat., Suppl. 1 (1861) 384. — Knema palembanica (Miq.)
Warb., Mon. Myrist. (1897) 592, t. 25. — Type: Teijsmann 3640, Sumatra, Palembang.

Myristica corticosa auct. non Lour.: Hook. f. & Thomson, FI. Ind. 1 (1855) 158; A. DC., Prodr. 14,
1 (1856) 205, p.p.; Miq., Fl. Ind. Bat. 1, 2 (1858) 69, p.p.

Knema glaucescens Jack var. glaucescens auct. non Jack: J. Sinclair, Gard. Bull. Sing. 16 (1958)
304, f. 12, 13 A, p.p.

Knema cinerea (Poir.) Warb. var. sumatrana auct. non (Blume) J. Sinclair: J. Sinclair, Gard. Bull.
Sing. 18 (1961) 185, p.p.; Backer & Bakh. f., Fl. Java 1 (1964) 140.

De Wilde — Myristicaceae (Knema) 263

Tree 5-30 m. Twigs 1.5—3.5 mm in diameter. Leaves membranous or chartaceous,
drying greenish to blackish brown, 6—20(—25) by 2—5.5(-11) cm, apex obtuse or acute
(-acuminate), base sometimes rounded; petiole 7-20 by 1-3 mm. Inflorescences: pedun-
cle 0-3 mm, brachyblast to 15 mm long, in male 2—20-flowered, in female 1—8-flow-
ered, flowers thinly pubescent, perianth reddish or yellow inside. Male flowers: pedicel
4-10 mm long; buds 2.5—3.5 mm in diameter, cleft 4/5; staminal disc 1.5—2.5 mm in
diameter, anthers 8-15, subsessile or stiped, horizontal, androphore 0.5—1 mm. Female
flowers: pedicel 1-6 mm; buds 4—6 mm long. Fruits 1—4 per infructescence, 1.8—3(—4?)
by 1.4—-1.8(—2.5?) cm, glabrescent; dry pericarp 1.5—2.5 mm thick; fruiting pedicel 3—7
(—10) mm long. — Photo 5.

Field-notes — Stilt-roots sometimes present, up to 1 m high; bark smooth, cracked,
flaky, scaly, or sometimes peeling off in strips 1 cm wide; bark 0.5—1 cm thick, pinkish
or red-brown; cambium brown; sapwood whitish, heartwood reddish; sap copious, pink-
ish. Leaves glossy dark green above, grey-glaucous to cinereous or whitish, with pale
yellowish or brownish midrib beneath. Flowers usually reddish inside, from Bali and
Borneo yellowish. Fruits yellow(-green), orange, or (pale) apricot; aril scarlet.

Distribution — Peninsular Thailand and Malesia: Sumatra, Peninsular Malaysia, Java,
Bali, Borneo.

Habitat & Ecology — Primary and degraded evergreen forest, including mixed dip-
terocarp forest; forest edges, riverine forest, periodically inundated forest, pole forest,
thickets; on a great variety of soils including sandstone, clay, and lateritic, granitic, and
black soil, (coral-)limestone, alluvial soils; 0O-1500 m altitude; fl. & fr. throughout the
year.

Notes — 1. Knema glauca is variable in a) the number of anthers, b) the mode of at-
tachment of the anthers, i.e. stiped or not, and with upper side of the connective broad or
narrow, hence, thecae opening + downwards or laterally, c) the colour of the flowers
inside, reddish or yellowish, and d) the size of the fruits. These features need more de-
tailed study in the field.

2. Specimens from Central and E Java, Bali, and Borneo frequently have as many as
12-15 anthers, as compared to 8-10(—12) generally in W Java and other areas; in W
Java, Sumatra, and Peninsular Malaysia 12-18 anthers is characteristic for the related
species K. sumatrana.

3. As can be seen in the keys, Knema glauca belongs to a group of related (resem-
bling) species, in which K. sumatrana is perhaps the one most closely allied; in most of
their overlapping area morphologically intergrading specimens (leaf size, fruit size) can
be found.

b. var. riparia W.J. de Wilde

Knema glauca (Blume) Petermann var. riparia W.J. de Wilde, Blumea 32 (1987) 126; Tree Fl. Sabah
& Sarawak 3 (2000) 411. — Type: S 18949 (Chai), Sarawak.

Tree 5-12 m. Twigs 1.2-2 mm in diameter. Leaves membranous, drying olivaceous
above, 11-22 by 2-6(—9) cm, apex acute or long-acuminate, base short or long attenu-
ate; petiole 10-15 by 1.5 mm. Inflorescences: peduncle up to | mm long, brachyblast
simple or forked, up to 4 mm long, in male 2-6(—10)-flowered, in female 1—2(—4)-flower-
ed: flowers sometimes subglabrescent; perianth red inside. Male flowers: pedicel 8-10

264 Flora Malesiana, Ser. I, Vol. 14 (2000)

mm long; buds subglobose, 3(—3.5) mm in diameter, cleft c. 5/6; staminal disc + flat,
subcircular, 2 mm in diameter; anthers 9—13, distinctly stiped, spaced, horizontal (open-
ing downwards), 0.4—0.5 mm long; androphore + cylindrical, slender, (1—)1.5 mm long.
Female flowers: pedicel 6 mm long; buds 4 by 2 mm. Fruits | or 2 per infructescence,
ellipsoid, apex sometimes + acute, 2—2.5 by 1.5—1.8(—2) cm, early glabrescent, + granu-
late, bright brown, dry pericarp | mm thick; fruiting pedicel 8-12 mm long.

Field-notess — Bark sometimes flaky. Flowers reddish inside, anthers yellow, disc
red. Fruits yellow to red, aril bright red, seeds whitish.

Distribution — Malesia: Borneo (Sarawak, W Kalimantan).

Habitat & Ecology — Primary riverine forest, on steep slopes; alluvial, rich clay,
yellow clay, or loamy soil; 0-500 m altitude; fl. & fr. throughout the year.

Note — This variety is distinguishable by a strikingly different general habit, with
twigs often with a yellowish tinge, comparatively narrow leaves drying greenish, and
bright brown long-stalked fruits. It superficially resembles K. luteola but that species
has a much finer venation on the upper leaf surface, larger fruits (4-7 cm), and the peri-
anth yellowish inside.

14. Knema glaucescens Jack

Knema glaucescens Jack, Mal. Misc. 7 (1821) 35; Warb., Mon. Myrist. (1897) 616 (under neglected
species); J. Sinclair, Gard. Bull. Sing. 16 (1958) 302, p.p.; W.J. de Wilde, Blumea 25 (1979)
462: Tree Fl. Sabah & Sarawak 3 (2000) 411. — Myristica glaucescens (Jack) Hook. f. & Thom-
son, Fl. Ind. 1 (1855) 157; FI. Brit. India 5 (1889) 111 (for the type only). — Type: Jack s.n.,
Sumatra (Bencoolen).

Myristica geminata Miq., Fl. Ind. Bat., Suppl. 1 (1861) 385. — Knema geminata (Miq.) Warb.,
Mon. Myrist. (1897) 604, p.p.; Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 247, p.p.; Ridl., Fl.
Malay Penins. 3 (1924) 72, p.p. — Types: Teijsmann 3620, 3924, Sumatra.

Knema cinerea (Poir.) Watb. var. sumatrana auct. non Blume: J. Sinclair, Gard. Bull. Sing. 18 (1961)
185, p.p.

Tree 8-15 m. Twigs 1—2(—3) mm in diameter, at first with very fine rusty, scurfy hairs
0.1 mm long, glabrescent; bark finely striate, not tending to crack or flake. Leaves
chartaceous or thinly coriaceous, ovate-oblong to lanceolate, 6—20(—23) by 2-6.5(-7.5)
cm, apex acute-acuminate, base rounded or subattenuate; greenish or brown above; lower
surface greyish, with subpersistent + dense stellate hairs 0.1—0.2 mm long, mixed with
larger stellate-dendroid hairs, when shed leaving minute hair scars; dots absent; midrib
flat or raised above; nerves 13-20 pairs, raised above, venation distinct above; petiole
5-15 by 1-2 mm. Inflorescences: peduncle up to 2 mm, brachyblast simple or forked, to
5 mm long; in male 5—20-flowered, in female 1—10-flowered; flowers with persistent
grey to rusty hairs 0.1—0.2 mm long, perianth 3-lobed, creamy or yellowish inside. Male
flowers: pedicel 6-11 mm long; bracteole early or late caducous, median or somewhat
higher up; buds globose, 2.5—3.5 mm in diameter, cleft c. 4/5, lobes 0.8—1 mm thick;
staminal disc flat to low-mammillate, circular or subtriangular, 1.5—2 mm in diameter;
anthers 9-13(—15?, see note 2), slightly to distinctly stiped, + horizontal, 0.2-0.4 mm
long, spaced; androphore 0.5—1 mm long. Female flowers: pedicel 3-6 mm long; bracteole
+ median; buds ellipsoid-obovoid, 4—5(—5.5) by 2.5—4 mm, cleft slightly over halfway,
lobes 1 mm thick; ovary ovoid, 2—2.5 mm long; stigma subsessile, + flat, (1- or) 2-4-
lobed and each lobe again few- to many- (up to 15-)lobulate or serrate. Fruits | or 2 per

De Wilde — Myristicaceae (Knema) 265

infructescence, ellipsoid or broadly obovoid, sometimes ridged, 1.8—2.2 by 1.2-1.8 cm,
with mealy hairs 0.1—0.3 mm long; dry pericarp 1(—2) mm thick; fruiting pedicel (3—)
4—11 mm long.

Field-notes — Bark ochre-brown, nearly smooth, not or only slightly flaking; wood
white. Leaves coriaceous, glossy dark green with whitish green midrib above, glaucous
or dirty white with brownish midrib beneath; venation not visible. Fruits with rusty
brown scurf. Aril hot, then cold in the mouth, tasting like cloves and nutmeg (Sinclair
SF 40280, 40368).

Distribution — Malesia: Sumatra (Indragiri, Benkulu, Palembang, Bangka), Penin-
sular Malaysia (Kelantan, Johore), Singapore, Borneo (Sarawak, Sabah, NW & SE Kali-
mantan).

Habitat & Ecology — Forest, including swamp forest; sand, granitic sand, and clayey
soil; 0-500 m altitude; fl. & fr. throughout the year.

Notes — 1. Knema glaucescens keys out together with K. scortechinii and K. commu-
nis on account of the subpersistent indumentum on the lower leaf surface; K. scortechinii
has a stouter habit, with larger male flowers; K. communis has smaller male flowers, and
smaller fruits.

2. Jack (1.c.) mentions in the original description 12-15 anthers; in the type, Jack s.n.
(L), 9-13 anthers were counted.

3. As compared to specimens from Sumatra and Peninsular Malaysia, those from
Borneo have more broadly obovoid or subglobose fruits (as against ellipsoid), shorter
stalked, 3-5 mm (6—11 mm in Sumatra). In general, the leaves of the Borneo specimens
are somewhat more coriaceous, and have a slightly coarser venation above.

The specimens S$ 23624 and 23644 from Sarawak 3rd Div., and S 27203 from 4th
Div., have subcoriaceous leaves with a particularly coarse venation on the upper surface,
broadly obovoid to subglobose fruits, all fairly deviating from the rest of the material.
These specimens may be found to represent a separate taxon, but because matching
male flowering material is lacking, they are tentatively included in K. glaucescens.

15. Knema globularia (Lam.) Warb.

Knema globularia (Lam.) Warb., Mon. Myrist. (1897) 601; J. Sinclair, Gard. Bull. Sing. 16 (1958)
325, f. 18; 18 (1961) 214; W.J. de Wilde, Blumea 25 (1979) 411; 32 (1987) 120, f. 2. — Myristica
globularia Lam., Mém. Ac. Paris (1788) 162. — Type: Sonnerat in Hb. Lamarck s.n., Peninsular
Malaysia (?).

Knema corticosa Lour., Fl. Cochinch. (1790) 605; ed. 2 (Willd.) (1793) 742; Warb., Mon. Myrist.
(1897) 593, t. 25 (incl. var. tonkinensis Warb.); Lecomte, Not. Syst. 1, 4 (1909) 101; FI. Indo-
Chine 5 (1914) 105, f. 10, 14; Merr., Trans. Amer. Phil. Soc. 24 (1935) 163. — Myristica corticosa
(Lour.) Hook. f. & Thomson, FI. Ind. 1 (1855) 158, p.p., only for the type; A. DC., Prodr. 14, 1
(1856) 205, p.p.; Kurz, For. Fl. Brit. Burma 2 (1877) 284, p.p. — Knema bicolor Rafin., Sylva
Tellur. (1838) 137. — Type: Loureiro s.n., Cochinchina.

Myristica sphaerula Hook. f. & Thomson, FI. Brit. India 5 (1890) 859. — Knema sphaerula (Hk. f.
& Thomson) Airy Shaw, Kew Bull. (1939) 545. — Type: Cantley 31, Peninsular Malaysia.
Myristica missionis Wall. ex King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 321, pl. 158. — Knema
missionis (King) Warb., Mon. Myrist. (1897) 602, t. 24; Gamble, Mat. Fl. Malay Penins. 5, 23
(1912) 247; Ridl., Fl. Malay Penins. 3 (1924) 71; Corner, Wayside Trees (1940) 477, f. 159, 161.

— Type: Wall. Cat. 6788, Peninsular Malaysia.

266 Flora Malesiana, Ser. I, Vol. 14 (2000)

Myristica lanceolata Wall., Cat. (1832) n. 6794.
Myristica glaucescens auct. non Jack: Hook. f. & Thomson, Fl., Ind. 1,.(1855).157,;p.p.BiaBat:
India 5 (1886) 111, p.p.

Tree 5-20 m. Twigs 1—-2(-2.5) mm in diameter, at first with yellowish brown hairs
0.1—-0.3 mm long, glabrescent; bark striate, not or but rarely tending to crack, never
flaking. Leaves membranous or thinly coriaceous, at first with indumentum mainly of
stellate golden brown or grey hairs 0.1 mm long, early or late glabrescent, oblong to
lanceolate, parallel-sided or broadest at or below the middle, 6—18(—24) by 1.5—4(-7)
cm, apex acute or acuminate, base attenuate, rarely rounded; olivaceous to dark brown,
usually with a blackish metallic lustre above; lower surface greyish, finely papillate
(lens!): dots absent; midrib sunken above; nerves 10—18(—22) pairs, faint, flat or sunken
above; venation inconspicuous; petiole 7-13 by 1—1.5(—3) mm. Inflorescences: sessile
or peduncle 1-5 mm, brachyblast simple or bifurcate, to 10 mm long; in male 5—20-flow-
ered. in female 1—10-flowered; flowers with persistent, rusty, + woolly hairs (0.05—)
0.1—0.3(—0.7) mm long, perianth 3- (or 4-)lobed, creamy or reddish (?) inside. Male
flowers: pedicel 3-5 mm long (outside Malesia to 11 mm long), bracteole above half-
way (in Malesia often apically), caducous or persistent; buds subglobose to broadly
obovoid, often triangular in the basal part, 3-5 mm in diameter, cleft c. 2/3, lobes 0.3-—
0.6 mm thick: staminal disc circular or angled, flat, 1.5-2.5 mm in diameter; anthers
8—13(—16), subsessile or just stiped, horizontal, 0.3—0.7 mm, not touching; androphore
0.5-1 mm long. Female flowers: pedicel 1.5—4 mm long, bracteole (nearly) apical; buds
ellipsoid-urceolate to ovoid, 4-5 by 2.5—3.5 mm, cleft about halfway; ovary subglobose
or conical, 1.5—2 mm long; style 0.5—1 mm long, stigma + flat, + 2-lobed and each lobe
again (2—)3—7-lobulate. Fruits 1—5 per infructescence, ellipsoid, subglobose, or some-
times + pear-shaped, 1.2—2 by 1-1.5 cm, at first with grey-brown to rusty hairs 0.1—0.2
mm long, + glabrescent, dry pericarp 2 mm thick; fruiting pedicel 3-10 mm long.

Field-notes — Bole fluted (once reported); bark flaking in small, thin scales; inner
bark pink, soft; sap red, copious. Leaves membranous or thinly coriaceous, dark green,
glossy above, glaucous, slightly pubescent beneath when young. Flowers cream or yel-
lowish inside (occasionally red in Thailand); staminal disc red, purple-red towards the
anthers. Fruits yellowish or orange, with thin orange-brown or rusty indumentum, tend-
ing to be rubbed off.

Distribution — NE India (doubtful), China (Yunnan), Burma, Vietnam, Laos, Cam-
bodia, Thailand: in Malesia: Sumatra (Aceh, Riau), most of Peninsular Malaysia, Singa-
pore, W Java.

Habitat & Ecology — Lowland primary and degraded forests on hill slopes, in South
Thailand, Peninsular Malaysia, and Java often in coastal areas: islands, rocky hill sides,
riverbanks, often along seashores; in Thailand in evergreen forest; 0-500 m altitude (in
Assam and Yunnan to c. 800 m); fl. Apr—July, in Thailand mainly Nov.—Feb.; fr. through-
out the year.

Notes — 1. Specimens of K. globularia from the Asian continent, e.g. Thailand, may
deviate by slightly larger male flowers, with buds 3.5—5 mm in diameter. Specimens
from Vietnam have large fruits, 2.5 by 2 cm.

2. Knema globularia seems related to K. attenuata from the Indian Peninsula; through
K. globulateria from Thailand it is connected with the Malesian K. latericia, the latter
differing, e.g., in flaking bark of twigs, and convex staminal disc.

De Wilde — Myristicaceae (Knema) 267

16. Knema glomerata (Blanco) Merr.

Knema glomerata (Blanco) Merr., J. Straits Branch Roy. Asiat. Soc. no 76 (1917) 81; Sp. Blanc.
(1918) 151; Enum. Philipp. Flow. Pl. 2 (1923) 183; J. Sinclair, Gard. Bull. Sing. 18 (1961) 219,
f. 11, map 9 (p.p., excl. syn. Knema stenocarpa Warb.); W.J. de Wilde, Blumea 25 (1979) 437;
32 (1987) 125; Tree Fl. Sabah & Sarawak 3 (2000) 412 g
(1837) 764: ed. 2 (1845) 525; ed. 3, 3 (1879) 164; Fern.-Vill., Noviss. Append. (1880) 27; Merr.,
Publ. Gov. Lab. Philipp. n. 27 (1905) 24, 73. — Neotype: Merrill Sp. Blanc. 504, Luzon.

Sterculia decandra Blanco, FI. Filip. (1837) 766; ed. 2 (1845) 526; ed. 3, 3 (1879) 166. — Type: not
indicated.

Myristica heterophylla Fern.-Vill., Noviss. Append. (1880) 178; Vidal, Rev. Pl. Vasc. Filip. (1886)
220. — Knema heterophylla (Fern.-Vill.) Warb., Mon. Myrist. (1897) 573, t. 25; Merr., Philipp.
J. Sci., Suppl. 1 (1906) 55; ibid., Bot., 3 (1908) 407. — Type: Vidal 507, Luzon.
Knema heterophylla (Fern.-Vill.) Warb. var. pubescens Warb., Mon. Myrist. ( 1897) 576. —[Myristica
laurina auct. non Blume: Vidal, Rev. Pl. Vasc. Filip. (1886) 221.] — Type: Vidal 509, Luzon.
Knema vidalii Warb., Mon. Myrist. (1897) 615; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 184. —
[Myristica iners auct. non Blume: Vidal, Rev. P|. Vasc. Filip. (1886) 221.] — Type: Vidal 1679,
Philippines.

Knema gitingensis Elmer, Leafl. Philipp. Bot. 3 (1911) 1065. — Type: Elmer 12200, Philippines.

Knema acuminata Merr., Philipp. J. Sci. 17 (1920) 256. — Type: Klemme 11266, Luzon.

Myristica corticosa auct. non (Lour.) Hook. f. & Thomson: Fern. -Vill., Noviss. Append. (1880)
178: A. DC., Prodr. 14, 1 (1856) 205, p.p., only for the Philipp. specimens; Vidal, Phan. Cuming.
Philipp. (1885) 139; Rev. Pl. Vasc. Filip. (1886) 220.

Myristica glaucescens auct. non (Jack) Hook. f. & Thomson: Ceron, Cat. Pl. Herb. Manila (1892)
141, p-p.

Knema glauca auct. non (Blume) Warb.: Merr., Enum. Philipp. Flow. PI. 2 (1923) 133s pip:

Tree 5—20 m. Twigs 1.5—2.5(—3) mm in diameter, at first with + woolly or scurfy rusty
hairs 0.2—0.5 mm long, glabrescent; bark finely striate or smooth, not tending to crack or
flake. Leaves chartaceous, rarely faintly lobed in the upper part, elliptic-oblong to lance-
olate, 10—-23(-35) by 3-7(-10) cm, base rounded to attenuate, apex acute(-acuminate),
rarely obtuse; greenish or brown above; lower surface at first with greyish, loose, soft-
stellate hairs 0.1—0.3 mm long, glabrescent, longer persistent on midrib and nerves; dots
absent: midrib + raised above; nerves (12—)15—20(—22) pairs, + flat above, often brown-
ish beneath; venation faint or distinct; petiole early or late glabrescent, 5-15 by 1-3 mm.
Inflorescences: sessile, brachyblast simple, sometimes forked, up to 8 by 6 mm; in male
4_20-flowered, in female 3—10-flowered; flowers with greyish brown or rusty hairs 0.2—
0.5 mm long; perianth 3-lobed, pink or red inside. Male flowers: pedicel 4-11 mm long,
bracteole caducous, + median; buds (depressed) globose, 3-5 mm in diameter, cleft nearly
to the base, lobes 0.3—0.7 mm thick; staminal disc + flat, slightly convex, or low mam-
millate, circular, 2-2.5 mm in diameter, anthers 9—13(—15), sessile to stiped, horizontal,
0.5 mm long, not touching; androphore slender, 1-2 mm long. Female flowers: pedicel
2-4 mm long, bracteole + median; buds ellipsoid-obovoid, 5 mm long, cleft about half-
way, lobes 0.5—0.7 mm thick; ovary ovoid, 2 mm long; stigma + sessile, with 2 lobes and
each lobe again 5- or 6-lobulate. Fruits 1-3 per infructescence, subglobose or globose-
ellipsoid, obtuse, 1.5—2.5(—3) by 1.2-2.3 cm, with greyish brown, scurfy hairs 0.1-0.2
mm long; dry pericarp 1.5 mm thick; fruiting pedicel (2—)4—6(—10) mm long.

Field-notes — Bark grey, smooth or slightly flaking in old trees. Leaves dull to glossy
medium green above, glaucous beneath. Flowers pink inside; staminal disc whitish; ovary
chocolate-brown tomentose, stigma sessile, green.

268 Flora Malesiana, Ser. I, Vol. 14 (2000)

Distribution — Malesia: Philippines (all over the archipelago: Mindoro, Batanes,
Babuyan, Luzon, Polillo, Marinduque, Cetanduanes, Sibuyan, Ticao, Samar, Leyte, Bili-
ran, Cebu, Negros, Panay, Guimaras, Sulu Is., Basilan, Mindanao, Palawan); one collec-
tion from Borneo (Sarawak: 4th Div., see note 2); one collection from Seram.

Habitat & Ecology — Primary and degraded (rain) forest, dipterocarp forest; 0-600
m altitude; fl. Jan.—Dec.; fr. June—Dec.

Notes — 1. Knema glomerata is the most common species of Knema in the Philip-
pines. It is close to K. korthalsii, which differs in 20-35 pairs of lateral nerves, some-
what larger flowers, in male with an average of 18 anthers, and a more pronounced
mammilla on the staminal disc. It is also close to K. tomentella (for differences see the
key), but this latter species possibly does not occur in the Philippines.

2. The specimen § 24419, from Sarawak, is highly questionable as regards the local-
ity, far away from the main area of the species in the Philippines. It has male flowers,
which agree with K. glomerata.

3. Knema glomerata is variable in the amount and length of the (deciduous) hairs on
flowers, twig apices, and petioles: in a woolly form, the hairs are relatively long, up to |
mm, and longer persistent, on the lower leaf surface it is denser; short-haired specimens
may resemble K. cinerea, which differs in a flat staminal disc, a very short indumentum
on the flowers, and the perianth being usually creamy or greenish inside, not pinkish red.

17. Knema hirtella W.J. de Wilde

Knema hirtella W.J. de Wilde, Blumea 25 (1979) 459; 32 (1987) 130; Tree Fl. Sabah & Sarawak 3
(2000) 413. — Type: SAN 30235 (Mikil), Sabah.

Tree 5-25 m. Twigs striate or not, 2-3.5 mm in diameter, at first with minute scurfy
hairs 0.1—0.2(—0.3) mm long, early glabrescent; bark finely striate, not tending to crack
or flake. Leaves chartaceous, oblong(-lanceolate), 7-25 by 2.5—10 cm, apex up to 1.5 cm
acute-acuminate or subobtuse, base subcordate, broadly rounded, or attenuate, olivaceous-
brown, with scattered hair scars above; lower surface greyish, with persistent sparse yel-
lowish hairs 0.1 mm long, or early to late glabrescent, at first with weak greyish hairs of
mixed sizes, 0.1—0.2(—0.3) mm long, in both cases mixed with scattered scars of early
shed coarser hairs; dots absent; midrib little raised above; lateral nerves 10—22 pairs, flat
or sunken above; venation + coarse (areoles 0.5 mm or more in diameter), distinct or
indistinct; petiole 8-12 by 1.5—2.5 mm. Inflorescences: sessile, brachyblast simple or
bifurcate, up to 5 mm long; in male 2—6(—10)-flowered, in female 1—6-flowered; flowers
with dense hairs 0.1 mm long or less, perianth 3-lobed, yellowish (?) inside. Male flower
(var. hirtella): pedicel 4.5-7 mm long, bracteole caducous or persistent, at or 1-2 mm
below the apex; buds globose or broadly obovoid, 3—4 mm in diameter, cleft 3/4—4/6,
lobes 0.5—0.8 mm thick; staminal disc flat or slightly convex, circular, 1.5—1.7 mm in
diameter; anthers 9—-13(—15), stiped or half-sessile, + horizontal or oblique, 0.2—0.4 mm
long, opening more or less downwards, spaced; androphore + tapering, 1-1.5 mm long,
in var. hirtella at base minutely hairy. Female flowers: pedicel 4—7.5 mm long, bracteole
caducous, at or above the middle; buds ellipsoid or long-obovoid, 4.5—6 by 3.5—4 mm,
cleft about halfway, lobes 0.7—1 mm thick; pistil 2.5—3 mm long, ovary ovoid, 1.5—2 by
|.5-2 mm, stigma subsessile, 2-lobed and each lobe again 6—8-lobulate or serrate. Fruits
1 or 2 per infructescence, ellipsoid or ovoid, 2.5—3(—3.5) by 1.7-2.6 cm, base rounded,

De Wilde — Myristicaceae (Knema) 269

apex rounded or acute, often with style remnant 0.5 mm, with hairs up to 0.5 mm long;
dry pericarp 1.5(—2) mm thick; fruiting pedicel 4—10(—15) mm long.

Distribution — Malesia: Borneo (Sarawak, Sabah, C & E Kalimantan).

Habitat & Ecology — Lowland and lower montane forest; on hill slopes and ridges;
on sandy soil, sandstone, lateritic and clay soils; 0—1000 m altitude; fl. May—July; fr.
throughout the year.

Notes — 1. Knema hirtella is a variable species which can easily be confused with
K. glauca and K. stenophylla; K. glauca differs in a very weak, early shed indumentum
on the lower leaf surface, persistent bracteole, glabrous androphore, and glabrous (gla-
brescent) fruits; K. stenophylla s.1. differs in a less stout habit, generally smaller fruits,
and glabrous androphore. Knema piriformis also resembles the present species, but has
differently shaped male buds.

2. Arbitrarily two largely sympatric varieties can be recognized, mainly based on
indumentum. Within the species there is a considerable variation in size and shape of the
fruits, but this cannot be used for a subdivision. The status of the accepted varieties is
not quite clear, moreover because male flowers of the variety pilocarpa are not known.

KEY TO THE VARIETIES

la. Twigs at apex and leaf buds with minute hairs 0.1 mm long. Fruits with stellate
(-dendroid) hairs 0.1 mm long or less. Androphore minutely pubescent at base... .
Pea he a0 008h i, beck). 2 narteds| ens eet ater eel deel ake vee ee a. var. hirtella

b. Twigs at apex and leaf buds with hairs 0.1-0.2(—0.3) mm long. Fruits with rough
dendroid hairs 0.5 mm long, especially towards the base ...... b. var. pilocarpa

a. var. hirtella

Twigs at apex with hairs 0.1 mm long or less. Leaves with persistent pale yellowish
stellate hairs 0.1 mm long, intermixed with early shed coarser hairs leaving distinct hair
scars (lens!) beneath. Androphore towards the base with minute greyish stellate hairs.
Fruits obtuse at apex; the indumentum of stellate(-dendroid) hairs 0.1 mm long or less.

Distribution — Malesia: Borneo (Sarawak: 4th Div.; Sabah).

b. var. pilocarpa W.J. de Wilde

Knema hirtella W.J. de Wilde var. pilocarpa W.J. de Wilde, Blumea 25 (1979) 461; 32 (1987) 131;
Tree Fl. Sabah & Sarawak 3 (2000) 414. — Type: SAN 85084 (Cockburn), Sabah.

Twigs at apex with rusty stellate-dendroid hairs 0.1—0.2 mm long. Leaves with per-
sistent yellowish hairs 0.1(—0.2) mm long, intermixed with early shed coarser hairs leaving
distinct hair scars beneath. Mature male and female flowers not seen. Fruits obtuse to
acute(-acuminate) at apex; the indumentum, especially towards the base of the fruit, of
stout rusty dendroid hairs 0.5 mm long.

Distribution — Malesia: Borneo (Sarawak, Sabah, E Kalimantan).

Habitat & Ecology — Periodically inundated forest.

Note — The specimen SAN 99720 (E Sabah), with immature male flowers, probably
belongs here according to its indumentum and large leaves up to 23 by 7.5 cm; the an-

270 Flora Malesiana, Ser. I, Vol. 14 (2000)

droecium (with 9 anthers) and glabrous androphore is reminiscent of K. subhirtella, a
species with smaller leaves.

18. Knema hookeriana (Wall. ex Hook. f. & Thomson) Warb.

Knema hookeriana (Wall. ex Hook. f. & Thomson) Warb., Mon. Myrist. (1897) 551, t. 24; Gamble,
Mat. Fl. Malay Penins. 5, 23 (1912) 237; Ridl., Fl. Malay Penins. 3 (1924) 67; Corner, Wayside
Trees (1940) 476: J. Sinclair, Gard. Bull. Sing. 16 (1958) 272, f. 1 & pl. I; 18 (1961) 226; W. J. de
Wilde, Blumea 25 (1979) 387. — Myristica hookeriana Wall. ex Hook. f. & Thomson, FI. Ind. 1
(1855) 156: A. DC., Prodr. 14, 1 (1856) 204; Migq., FI. Ind. Bat. 1, 2 (1858) 69, & Suppl. 1 (1861)
384: Hook. f., Fl. Brit. India 5 (1886) 109; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 325, pl.
163. — Type: Wall. Cat. 6802A, Peninsular Malaysia.

Tree 5—30 m. Twigs 5—10 mm in diameter, with dense, buff, woolly hairs (3-)5—8 mm
long, late glabrescent, the indumentum shed in compact rags; bark brown, longitudinal-
ly cracking, and flaking. Leaves coriaceous, when immature with dense hairs (2—)5—8
mm long, (oblong-)lanceolate, + narrowed towards the base, 25—60 by 5-22 cm, apex
subobtuse, acute, or acuminate, base subattenuate, rounded, or subcordate; olivaceous
to dark brown above, lower surface glabrescent, greyish, faintly papillate or not; dots
absent: midrib raised above; nerves 25-33 pairs, + raised above, reddish to yellowish
brown: venation distinct; petiole 10-30 by 8-12 mm, late glabrescent, hairs (I-)2—3 mm
long. Inflorescences: sessile, brachyblast simple or 2—4-furcate, up to 20 mm in diam-
eter, in male 5—20-flowered, in female 4—10-flowered; flowers with dense woolly hairs
1-3 mm long, perianth 3- (or 4-)lobed, striate and red inside. Male flowers: pedicel 15—
20 mm long, bracteole caducous, at or somewhat above the middle; buds depressed glo-
bose-obovoid, 7-9 by 8-10 mm, cleft c. 2/3, lobes 1-2 mm thick; staminal disc blunt-
triangular, + concave, 3-4 mm in diameter; anthers 15—25, just sessile, half-erect or
horizontal, 0.6—1 mm, not touching; androphore + tapering, 2(—3) mm long. Female
flowers: pedicel (5—)10—20 mm long, bracteole + median to subapical; buds subglobose,
or ovoid, or depressed-obovoid, 8-10 by 8-10 mm, cleft about halfway, lobes 1—2.5 mm
thick; ovary subglobose, 2-4 mm in diameter, hairs 2 mm long; style 1.5—2 mm long,
stigma + 2-lobed and each lobe again 3- or 4-lobulate, these again 2—4-lobulate, hence
stigma c. 20-lobulate-serrate. Fruits 1-3 per infructescence, ellipsoid, 4.5—7(—8) by 3—
4.5 cm, with thick, buff woolly hairs S—13 mm long, not easily rubbed off; dry pericarp
3-8 mm thick; fruiting pedicel 5S—10(—20?) mm long.

Field-notes — Bole straight, not buttressed; crown conical; bark smooth but scaly by
several layers of thin brittle blackish adherent flakes, rectangular or elongate. Branches
drooping or horizontal. Young foliage appearing periodically, pendulous, clothed in a
dense fawn down; older leaves glossy dark green above, glaucous beneath.

Distribution — Southern Peninsular Thailand and Malesia: Sumatra (East Coast,
Palembang), Peninsular Malaysia (known from all provinces except Perlis and Wellesley
Prov.), Singapore.

Habitat & Ecology — Primary and degraded forest; ridge tops, hill sides; 0-800 m
altitude; fl. Feb.—May; fr. June—Dec.

Note — Knema hookeriana is a characteristic species, noticeable for its twigs with
flaking bark, large leaves, and long, dense, woolly indumentum on twigs, petioles, flow-
ers, and fruits.

De Wilde — Myristicaceae (Knema) 271

19. Knema intermedia (Blume) Warb.

Knema intermedia (Blume) Warb., Mon. Myrist. (1897) 564, t. 25; Gamble, Mat. Fl. Malay Penins.
5, 23 (1912) 239; Koord., Exk. Fl. Java 2 (1912) 258; Ridl., Fl. Malay Penins. 3 (1924) 68;
Corner, Wayside Trees (1940) 477; Backer & Bakh. f., Fl. Java 1 (1964) 140; J. Sinclair, Gard.
Bull. Sing. 16 (1958) 315, f. 15, pl. IE B; 18 (1961) 227; W.J. de Wilde, Blumea 25 (1979) 427;
Tree Fl. Sabah & Sarawak 3 (2000) 414. — Myristica intermedia Blume, Rumphia 1 (1835)
187: Hook. f. & Thomson, FI. Ind. 1 (1855) 158; A. DC., Prodr. 14, 1 (1856) 206; Miq., FI. Ind.
Bat. 1, 2 (1858) 70: Hook. f., Fl. Brit. India 5 (1886) 112; King, Ann. Roy. Bot. Gard. Calc. 3
(1891) 317, pl. 154; Koord. & Valeton, Med. Lands PI. Tuin 17 (1896) 192. — Type: Blume s.n.,
Java.

Myristica glabra De Vriese, PI. Ind. Bat. Orient. (PI. Reinw.) (1857) 85. — Type: Reinwardt s.n.,
Java.

Myristica iteophylla Miq., Fl. Ind. Bat. 1, 2 (1858) 59. — Type: Teijsmann 478, Sumatra.

Myristica corticosa (Lour.) Hook. f. & Thomson var. decipiens Migq., Ann. Mus. Bot. Lugd.-Bat. 2
(1865) 51. — Type: Korthals s.n., Kalimantan.

Myristica glauca auct. non Blume: Zollinger msc., Moritzi, Syst. Verz. Java (1845) 38.

Tree 5—15(—30) m, sometimes with stilt-roots. Twigs slender or stout, subterete, slight-
ly angled, or flattened, 1.5—4(—5) mm in diameter, at first with rusty hairs 0.3—0.5 mm
long, glabrescent; bark smooth or finely striate, not tending to crack or flake. Leaves
chartaceous or coriaceous, oblong to lanceolate, (7—)9—35 by 1.3—8 cm, apex acute
(-acuminate), base rounded, cuneate, or attenuate; greenish brown above, lower surface
grey-brown, at first with dense indumentum, early glabrescent; dots absent; midrib raised
above; nerves 12-25 pairs, much raised; venation distinct; petiole (late) glabrescent,
10-25 by 2-3 mm. Inflorescences: peduncle up to 2 mm long, brachyblast simple or
2- or 3-furcate, up to 15 mm long, in male 5—25-flowered, in female 1-10-flowered;
flowers with woolly or mealy rusty hairs 0.3—0.5 mm long, perianth 3-lobed, creamy or
pinkish (?) inside. Male flowers: pedicel 5—13 mm long, bracteole subpersistent, median
to apical; buds subglobose, circular to subtriangular in cross section, (3—)4—5 by (3-)
45.5 mm. cleft (2/3—)3/4, lobes 0.3—0.4 mm thick; staminal disc circular to subtriangu-
lar, 2-2.5 mm in diameter, with a conspicuous mammilla 1—1.5 mm long; anthers | 1-15,
horizontal, 0.3—0.5 mm, subsessile or shortly stiped, not touching, androphore 0.5 mm
long. Female flowers: pedicel 2-8 mm long, bracteole + median; buds obovoid, 5(—6)
by 3.54 mm, cleft about halfway, lobes 0.5-0.7 mm thick; ovary subglobose or ovoid,
2.5 by 2.5—3 mm; stigma sessile, 2-lobed, each lobe again (2—)3—5-lobulate. Fruits 1—4
per infructescence, ellipsoid, 2.5—4 by 1.5—2.3 cm, often ridged, with hairs 0.3—0.5 mm
long, + easily rubbed off; dry pericarp 2-3 mm thick; fruiting pedicel (3-)5—14 mm long.

Field-notes — Trunk without buttresses, though frequently with stilt-roots; bark
smooth, scaly, or flaky; wood white or somewhat reddish. Leaves bright dark green,
glossy, the midrib much paler, yellowish above, very glaucous or light blue-green or
whitish beneath; newly emerged leaves velvety golden by the indumentum at both sides.
Flower buds yellowish brown, cream inside, stamens and disc pink; fruits often yellow
or orange with rusty or red indumentum.

Distribution — Malesia: Sumatra (W Coast, E Coast, Palembang, Lampong, Bangka,
Riau-Lingga Arch.), Peninsular Malaysia (all provinces except Perlis, Kedah, Trengganu,
Negri Sembilan), Java (W Java), Borneo (S & SE Kalimantan; Sarawak; Sabah, see
note 2; Natuna I.).

272 Flora Malesiana, Ser. I, Vol. 14 (2000)

Habitat & Ecology — Mixed lowland forest, peat forest; once from granitic sand;
0—1000 m altitude; fl. & fr. throughout the year.

Sinclair (1958) mentioned that the flower buds, observed on the tree in the Singapore
arboretum, remained closed for six weeks before opening, and that they probably had
been in the unopened stage for quite a considerable time prior to having been noticed.
The same phenomenon was seen in Knema leuserensis, in which apparently full-grown
flower buds of male and female specimens remained closed for about six weeks before
opening.

Notes — 1. Knema intermedia is characterized by the conspicuous long-mammillate
staminal disc. It is related to K. plumulosa; for differences see under that species.

2. SAN 71155, the only collection seen from Sabah, differs in a stout habit, with thick
twigs, large leaves, and short fruiting pedicels only 2-4 mm long.

20. Knema kinabaluensis J. Sinclair

Knema kinabaluensis J. Sinclair, Gard. Bull. Sing. 18 (1961) 229, f. 12; W.J. de Wilde, Blumea 25
(1979) 450; Tree Fl. Sabah & Sarawak 3 (2000) 415. — Type: Sinclair 9224, Sabah.

Tree 5-25 m. Twigs sometimes + blunt-triangular, 1.5—2.5 mm in diameter, drying
blackish brown, at first with yellowish rusty hairs 0.1 mm long, glabrescent; bark striate
or not, not cracking or flaking. Leaves (rigidly) coriaceous, (oblong-)lanceolate, 9-21
by 2.5—-5 cm, base (rounded or) attenuate, apex acute-acuminate; dark brown above;
lower surface grey-glaucous, at first with fairly dense mixed stellate and scale-like hairs
0.1 mm long or less, early glabrescent, leaving hair scars; dots absent; the midrib raised
above; nerves 12—25 pairs, raised above; venation + coarse (areoles 0.5 mm in diam-
eter), very distinct above; petiole 10—20 by 2-3 mm. Inflorescences: sessile, brachyblast
simple or bifid, up to 15 mm long; in male 4—10-flowered, in female 1—4-flowered;
flowers with mealy, rusty hairs 0.1—0.2 mm long; perianth 3-lobed, pink or reddish in-
side. Male flowers: pedicel 3-6 mm long, bracteole caducous, about median; buds sub-
globose, 2.5-3.5 mm in diameter, cleft (3/4—)4/5, lobes 0.5—1 mm thick; staminal disc
flat or slightly convex, circular, pink, 1.5(—2) mm in diameter; anthers 7 or 8(—107?), just
sessile, almost horizontal to oblique, 0.7 mm long, with broad connective, the thecae
opening downwards, spaced; androphore | mm long. Female flowers: pedicel 3-4 mm
long, bracteole caducous, about median; buds obovoid-oblong, 5 mm long, cleft about
halfway; ovary ovoid, 2 mm long; stigma + sessile, 2-lobed, lobes not clearly lobulate,
1 mm long. Fruits 1 or 2 per infructescence, ovoid or oblong, base rounded, apex acute
or short-apiculate, + ridged along the line of suture, 2—4.5 by 1.5—3 cm, with yellowish
brown to rusty hairs 0.1—0.2 mm long; dry pericarp 2-4 mm thick; fruiting pedicel 3-10
mm long. — Photo 6.

Field-notes — Bark smooth, or fine and shallowly fissured; inner bark pale brown to
reddish; sapwood whitish, cambium reddish. Leaves glossy dark green with whitish green
midrib above, glaucous with yellowish midrib beneath. Flowers pale pink or reddish
inside. Fruits orange or golden brown, or brown scurfy; aril red with pink tinge.

Distribution — Malesia: Borneo (Sabah, endemic to the Mt Kinabalu area).

Habitat & Ecology — Montane (degraded) rain forest; on ridges and slopes, along
rivers; on sandstone; also occurring in the coppermining area near Ranau; 1000-2300 m
altitude; fl. & fr. throughout the year.

De Wilde — Myristicaceae (Knema) 273

Notes — 1. Knema kinabaluensis comes close to K. cinerea and K. glauca, possibly
most closely to the latter which differs in less coriaceous leaves, persistent bracteole,
and early glabrescent fruits. By its coriaceous leaves K. kinabaluensis resembles K. rigidi-
folia from Peninsular Malaysia and also resembles certain coriaceous-leaved forms re-
ferred to K. pectinata from Sarawak, and K. kunstleri subsp. alpina from Sabah. Knema
rigidifolia differs in the shape of male buds and fruits, and the indumentum on the lower
leaf surface; K. pectinata has stouter twigs, larger male buds, and more globose fruits;
K. kunstleri can always be recognized by the blackish dots on the lower leaf surface.

2. In most of the herbarium material the mature fruits are partly ridged along the line
of suture, but in a few specimens this ridge is lacking.

21. Knema korthalsii Warb.

Knema korthalsii Warb., Mon. Myrist. (1897) 557. — Myristica korthalsii (Warb.) Boerl., Handl.
FI. Ned. Ind. 3, 1 (1900) 90. — Type: Korthals s.n., Kalimantan.

Tree 2-30 m. Twigs faintly flattened or (2- or) 3-angular, 2-5 mm diameter, at first
with rusty hairs 0.3-0.5 mm long, glabrescent; bark striate, thinly flaking or not. Leaves
chartaceous or thinly coriaceous, oblong to oblanceolate, 12—35 by 2.5—9(—12) cm, base
cuneate to subattenuate, or rounded, apex acute(-acuminate); greenish (dark) brown above;
lower surface greyish, largely glabrescent, sometimes with indumentum remaining on
midrib and nerves; dots absent; midrib slightly raised above; nerves 12—35(—40) pairs,
raised; venation distinct above; petiole tomentose or late glabrescent, 8-25 by 2-5 mm.
Inflorescences: sessile, brachyblast simple or warted, to 8 mm diameter, in male in um-
bels of (1—)5—20 flowers, female up to 10-flowered; flowers with persistent rusty hairs
0.2-0.7 mm long, perianth 3-lobed, red inside. Male flowers: pedicel 5-13 mm long,
bracteole subpersistent or caducous, 0.5—1 mm, + median; buds depressed globose, of-
ten obtusely angular in cross section, 4-6(—7) mm diameter, cleft c. 4/5, lobes 0.5 mm
thick, at base 1 mm thick; staminal disc circular, convex or mammillate, sometimes + flat
(especially in Philippines), 2-2.8 mm diameter; anthers 10—20, subsessile, horizontal,
0.5 mm long, not touching, opening + to beneath; androphore 0.7—1 mm long, + clasped
by the thickened base of the perianth. Female flowers: pedicel 1.5—2.5 mm long, bracteole
about median, caducous; buds obovoid-oblong, 5—6 mm long, cleft c. 2/3, lobes 0.5—1
mm thick; ovary ovoid, 2 mm long; stigma + sessile, 2-lobed, and each lobe again with
3-6 laciniations. Fruits 1-3 per infructescence, ellipsoid or oblong, 1.5—4 by 1-2 cm,
with rusty hairs 0.5—1 mm long; dry pericarp 3 mm thick; fruiting pedicel 1-6 mm long.

Distribution — Malesia: Borneo, S Philippines (Balabac I., Palawan, Sulu Is., Basilan,
Mindanao mainly in Zamboanga Prov.).

KEY TO THE SUBSPECIES

la. Twigs towards apex 3-5 mm diam., older bark not flaking. Lateral nerves to 40 per
side, usually clearly interarching. Fruits 2-4 cm, pedicel 2-6 mm long; persistent
style iand:.stiemato lL mmulong 222" . 22.4. ee ao. an et a. subsp. korthalsii

b. Twigs towards apex 2-3 mm diam., older bark thinly flaking. Lateral nerves 12—22
per side, lines of interarching more faint. Fruits 2 cm long, pedicel 1-2 mm long;
persistent stigma sessile (as mi latericia)....% | sven cee: b. subsp. rimosa

274 Flora Malesiana, Ser. I, Vol. 14 (2000)

a. subsp. korthalsii

Knema korthalsii Warb., Mon. Myrist. (1897) 557; J. Sinclair, Gard. Bull. Sing. 18 (1961) D2,
f. 13: W.J. de Wilde, Blumea 25 (1979) 428; 43 (1998) 242; Tree Fl. Sabah & Sarawak 3 (2000)
417. — Myristica korthalsii (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 90.

Knema insularis Merr., Philipp. J. Sci. 30 (1926) 394. — Type: BS 44288 (Ramos & Edano), Sulu
Islands.

Knema cenabrei Merr. & Quisumbing, Philipp. J. Sci. 37 (1928) 144, pl. 3. — Type: FB 29973
(Cenabre), Palawan.

Myristica mindanaensis auct. non Warb.: Merr., Bull. Bur. For. Philipp. 1 (1903) 21; Philipp. J. Sci.,
Bot. 3 (1908) 76. — Knema mindanaensis auct. non (Warb.) Merr.: Merr., Enum. Philipp. Flow.
Pl. 2 (1923) 184, p.p., excl. type.

Tree 7-30 m. Twigs 3—5 mm diameter, not flaking. Leaves chartaceous or coriaceous,
oblong to oblanceolate, 12-35 by 2.5—9(—12) cm, base (sub)attenuate; nerves (14—)20-
35(—40) per side; petiole late glabrescent, (8—)10—25 by 2-5 mm. Flowers with hairs
0.3—0.7 mm. Male flowers: pedicel 7-13 mm, bracteole + median, caducous, buds de-
pressed subglobose, obtusely trigonous, 4—6(—7) mm diameter, anthers (10—)12(—20).
Female flowers as the species. Fruits 1.5—4 cm long, with hairs 0.5—1 mm, stigma more
or less withering, fruiting pedicel 2-6 mm long.

Field-notes — Bark of trunk smooth or scaly, grey; slash of outer bark 3 mm, soft,
chocolate or blackish, inner bark 10 mm, soft, reddish; sapwood red or white; cambium
reddish or yellow; wood hard, brown. Leaves whitish or glaucous beneath.

Distribution — See the species.

Habitat & Ecology — Primary forest on level land or hill sides, mixed dipterocarp
forest; found on a variety of soil types: brown soil, (sandy) loam, clay, basalt rock,
limestone, and sandstone; 0—400 m altitude; fl. in Borneo July—Oct., in the Philippines
May—June; fr. May—Oct.

Notes — 1. Knema korthalsii belongs to the group of species with the staminal disc
distinctly convex or with a mammilla; it is also related to a group of species to which
K. glomerata, K. pectinata, and K. woodii belong. Subsp. korthalsii seems most closely
related to K. glomerata, which differs in a + flat staminal disc, more slender twigs,
leaves with fewer lateral nerves, 12—20 pairs, and fewer anthers, 9-13, as against 10-20,
for K. korthalsii. Knema glomerata and K. korthalsii overlap in area in Mindanao, Sa-
mar, and the Sulu Islands. Knema pectinata has much more lateral nerves, and K. woodii
can be distinguished by the more numerous sessile anthers, and shorter indumentum on
the twig apices and flowers. Subsp. korthalsii can be confused easily with K. membrant-
folia.

2. BS 44288 (Ramos & Edajio), the type of the synonym K. insularis, is intermediate
with K. glomerata, and might belong to this latter species.

b. subsp. rimosa W.J. de Wilde

Knema korthalsii Warb. subsp. rimosa W.J. de Wilde, Blumea 43 (1998) 242, Tree Fl. Sabah &
Sarawak 3 (2000) 417. — Type: SAN A 1996 (Wood), Sabah.

Tree 2-10(—15) m. Twigs towards apex 2-3(—5) mm diam. Bark of older twigs thinly
flaking. Leaves (thinly) chartaceous, elliptic- or obovate-oblong, 15-30 by 3.5—8.5 cm,
base acute or rounded; nerves 12—22 per side; petiole 8-12 by 2-4 mm, tomentose or

De Wilde — Myristicaceae (Knema) PAIS)

late glabrescent. Flowers with hairs 0.2—0.5 mm long; male pedicel (4—)5—10 mm long;
bracteole subpersistent, 0.5—1 mm long, buds depressed-globose, 3-4 by 4-6 mm; an-
thers 10-12. Female flowers and fruits resembling those of K. latericia subsp. ridleyi.
Fruits ellipsoid or obovoid, 2—2.3 cm, with hairs 0.5—1 mm long, persistent sessile lobed
stigma; fruiting pedicel 1—2(—3) mm long.

Field-notes — Bark smooth, blackish, (irregular) scales reveal orange-brown patches
below. No buttresses. Outer bark pale brown, soft, flaky; inner bark fibrous, pink; sapwood
off-white, no smell. The type specimen flowering abundantly throughout the crown.
Perianth brown, tomentose, inside red. Anthers yellow, disc swollen, pink. Leaves glau-
cous below.

Distribution — Malesia: Borneo (endemic to SE Sarawak and E Sabah).

Habitat & Ecology — In lowland forest, dipterocarp forest, on brown sandy soil,
leached loam over sandstone; well-drained soils; up to 900 m altitude; fl. Mar., Apr.; fr.
Apr.—June.

Note — Subsp. rimosa deceptively resembles K. latericia subsp. ridleyi in the veg-
etative state and female flowers or fruits. The male flowers are quite distinct. Both sub-
species share the flaking bark of the older twigs, a character regarded as important for
the distinction of taxa and for a proper distinction one should examine fully mature male
flowers still in buds, or just opened.

22. Knema kostermansiana W.J. de Wilde

Knema kostermansiana W.J. de Wilde, Blumea 25 (1979) 455; 32 (1987) 127; Tree Fl. Sabah &
Sarawak 3 (2000) 417. — Type: SAN 19047 (James Ah Wing), Sabah.

Tree 6-20 m. Twigs finely to coarsely striate, 2-4 mm diameter, at first with greyish
stellate hairs less than 0.1 mm long, early glabrescent; bark striate, not tending to crack
or flake. Leaves membranous or chartaceous, elliptic or oblong, 12—30(—40) by 5.5—
12(-19) cm, apex acute-acuminate, base shallowly cordate to broadly rounded; olivaceous
above; lower surface greyish, finely papillate or not, at first with inconspicuous, weak,
greyish sessile-stellate hairs 0.1 mm long or less, early glabrescent; dots absent; midrib
slightly raised above; nerves 15-22 pairs, sunken or + flat above; venation + coarse
(areoles 0.5 mm diam. or more), distinct above, faint in older leaves; petiole 10-15 by
2-3 mm. Inflorescences: sessile, brachyblast 2- or 3-furcate, up to 5 mm long; in male
5—10-flowered, female few-flowered; flowers with scale-like hairs less than 0.1 mm
long, perianth 3-lobed, reddish (?) inside. Male flowers: pedicel 2.5—7 mm long, bracteole
minute, caducous, apical; buds (depressed) globose, 3—3.2 mm diameter, cleft c. 4/5,
lobes 0.8 mm thick; staminal disc flat or faintly mammillate, circular, 1.5—2 mm diam-
eter; anthers 8-12, subsessile to just stiped, 0.5-0.6 mm long, horizontal, spaced; andro-
phore 0.6—0.7 mm long. Female flowers: pedicel 3-4 mm long, bracteole caducous,
(sub)apical; buds ellipsoid-oblong, + narrowed in the middle, (4—)5—6 by (2—-)2.5—3 mm,
cleft c. 1/2, lobes 1 mm thick; ovary long-ovoid, 2—2.5 by 1.5 mm; style slender, 1—1.5
mm long, glabrous, with 2-lobed fleshy stigma, each lobe again shallowly 3-lobulate.
Fruits solitary or 2 or 3 together, oblong(-lanceolate), + fusiform, 7 by 2 cm, base taper-
ing, apex long-tapering or beaked, with grey-brown scale-like hairs less than 0.1 mm,
appearing as if subglabrous; dry pericarp 2 mm thick; fruiting pedicel rather stout, 8-10
mm long, with the bracteole scar nearly median.

276 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Medium-sized tree to 20 m; bark chocolate- to dark-brown, fissured
dark greyish, inner bark whitish brown, sapwood white, yellowish, or brownish; exudate
red. Flowers greenish or grey-brown, turning yellowish, pink inside. Fruits greenish.

Distribution — Malesia: Borneo (Sabah, including border area of NE Kalimantan).

Habitat & Ecology — Logged-over forest, riverbank forest; on sandstone; 0-700 m
altitude; fl. June—Oct.; fr. July—Oct.

Note — Knema kostermansiana may resemble K. glauca and K. pulchra. Knema
glauca differs in its shorter fruits, and median bracteole, narrower leaves, and + raised
nerves; K. pulchra is distinguishable by pear-shaped, not globose, male buds and by
much broader ellipsoid fruits with rounded apex. The slender, + fusiform fruits of 7 cm
length of K. kostermansiana much resemble those of K. luteola, and somewhat those of
K. ashtonii var. ashtonii; both differ in a much finer venation on the upper leaf surface,
the first by narrow-rounded or attenuate leaf base and generally somewhat smaller fruits
(up to 70 mm long), the second in stouter habit and the fruits with + blunt apex, not
pointed.

23. Knema krusemaniana W.J. de Wilde

Knema krusemaniana W.J. de Wilde, Blumea 41 (1996) 385. — Type: Mogea & de Wilde 4216,
Kalimantan.

Tree 4-6 m. Twigs 4—5 mm diameter, with dense, dark brown, coarse hairs 1-2 mm
long, late glabrescent; bark blackish brown, finely striate, not cracking nor flaking, in-
conspicuously lenticellate. Leaves (thinly) chartaceous, oblong, 15—35 by 4-10 cm, base
narrowly rounded or cuneate, apex (acute-)acuminate; olivaceous or bright brown above;
lower surface pale brown or grey-brown, minutely papillose, with sparse but conspicu-
ous stellate and stellate-dendroid hairs 0.5—1.5 mm, sometimes late glabrescent; dots
present (lens!); midrib raised above, nerves 14—18 pairs, raised, venation distinct at both
surfaces; petiole 6-14 by 2.5—4 mm, late glabrescent. Inflorescences: sessile, brachyblast
wart-like or slenderly worm-like, simple or 2- (or 3-)armed, to 12 mm long, with woolly
rusty stellate hairs 0.5—1 mm, late glabrescent, in male with 10—25 clustered flowers;
flowers with dense woolly stellate and stellate-dendroid hairs 0.5—1 mm long, perianth
4-lobed, pink inside. Male flowers: pedicel 4-5 mm long, slender or thickish, bracteole
| mm long or less, 1-1.5 mm below the apex, caducous; buds (slightly immature) sub-
globose, 2 mm diameter, cleft 3/4—4/5, lobes carnose, at base enclosing the androecium;
staminal disc flat or slightly convex, circular, |-1.7 mm diameter, anthers 7 or 8, short-
stiped, spaced, each 4-sporangiate (each with 2 thecae), androphore hardly 0.5 mm long.
Female flowers not seen. Fruits 2.5—3 by 1.7—2 cm, obtuse, with hairs 0.5—1 mm, fruit-
ing pedicel 10 mm.

Field-notes — Small, slender tree, 4—6 m. Indumentum of young twigs brown. Leaves
glaucous at underside. Flower buds pale yellowish brown; flowers pink.

Distribution — Malesia: Borneo (W & Central Kalimantan, area of the upper Katingan:
Mendawai and Kahayan Rivers).

Habitat & Ecology — Old degraded forest and forest with Agathis over poor sandy
soil with thin peat layer; locally common; 200-220 m altitude; fl. Mar., Dec.

Note — Knema krusemaniana belongs, with K. conferta and K. pedicellata, to a group
of species with dots on the lower leaf surface. It is distinguishable by thinner leaves,

De Wilde — Myristicaceae (Knema) 277

twigs with longer-haired persistent conspicuous indumentum, and smaller male flowers,
with only 8 anthers. The perianth in K. krusemaniana is 4-lobed, in the related K. conferta
and K. pedicellata it is 3-lobed and 4- or 5-lobed, respectively. The pedicels are exceed-
ingly long in K. pedicellata, short in K. krusemaniana. The staminal disc is conspicu-
ously convex in K. pedicellata, + flat in K. krusemaniana and K. conferta; in K. pedicel-
lata the perianth has a raised disc at base, which is lacking in both other species. Finally
there are remarkable differences in the almost stalked anthers: 8 in K. krusemaniana,
which are uniquely 4-sporangiate; in K. conferta and K. pedicellata there are (10—)12—
18 bisporangiate anthers.

24. Knema kunstleri (King) Warb.

Knema kunstleri (King) Warb., Mon. Myrist. (1897) 568, t. 25; W.J. de Wilde, Blumea 25 (1979)
468: 43 (1998) 244: Tree Fl. Sabah & Sarawak 3 (2000) 418. — Myristica kunstleri King,
Ann. Roy. Bot. Gard. Calc. 3 (1891) 314, pl. 149. — Lectotype: King’s coll. 4216, Peninsular
Malaysia.

For more references and synonyms see the subspecies.

Tree 5-30 m. Twigs 1.5—3(—4) mm diameter, at first with rusty, chocolate, or grey-
brown hairs 0.1—0.2 mm long or less, glabrescent; bark striate or not, not tending to
crack or flake. Leaves membranous or coriaceous, variable, broadly elliptic or ovate, to
oblong-lanceolate, 5—28 by (1.5—)2—10 cm, base attenuate to broadly rounded, apex sub-
obtuse to acute(-acuminate); greenish to brown above, the lower surface greyish, finely
papillate, with (sub)persistent golden to grey-brown, mostly stellate scale-like hairs
0.1 mm long (0.3 mm in subsp. pseudostellata); + dense or remote dots, especially present
on the veinlets; midrib raised above; nerves 6—18 pairs, raised above; venation fine or
coarse, raised and distinct or not (in subsp. kunstleri raised and of a lighter colour);
petiole 5—20 by 1.5-3 mm. Inflorescences: sessile or pedunculate, up to 5 mm, some-
times supra-axillary, brachyblast simple, warted, or + forked, 2-4 mm long; in male
3-15-flowered, female 1—5-flowered; flowers with persistent scale-like hairs 0.1—0.2
mm long, perianth 3- (or 4-)lobed, reddish or cream inside (subsp. coriacea and alpina
always creamy?). Male flowers: pedicel 5-11 mm long, bracteole subpersistent, median
or somewhat above, rarely nearly apical; buds (depressed) globose, sometimes + angu-
lar, 2.5—3(—4) mm diameter, cleft nearly to the base, lobes 0.5—1 mm thick; staminal disc
flat or slightly convex, circular or somewhat angular, 1.3—2(—3) mm diameter; anthers
8-14, just to distinctly stiped, 0.3—0.6 mm, horizontal, spaced; androphore 0.5—1 mm
long. Female flowers: pedicel 2-9 mm long, bracteole about median or subapical; buds
+ obovoid, 4—5 by 2.5-3.5 mm, cleft c. 1/2, lobes 1(—1.5) mm thick; ovary ovoid, 2 mm
long; stigma subsessile, 2-lobed and each lobe again 2—S-lobulate. Fruits 1—4 per
infructescence, subglobose or ellipsoid, 1.5—4.2 by 1—2.8 cm, base rounded or narrowed,
apex rounded to subacute, with minute scale-like hairs 0.2 mm long or less, sometimes
seemingly glabrous; dry pericarp 1-2 mm thick; fruiting pedicel 3-15 mm long.

Field-notes — Bark smooth, fine scaly, rough, or sometimes fissured; grey, reddish,
or dark brown; slash pink or reddish; wood cream, sometimes yellow-brown or pink.
Leaves glossy green above, dirty white or glaucous beneath. Flowers greenish yellow
inside, or reddish in subsp. kunstleri (always?), or in subsp. coriacea and alpina cream
or pale yellow. Fruits yellow, red brown, dark (olive-)brown, or orange.

278 Flora Malesiana, Ser. I, Vol. 14 (2000)

Distribution — A polymorphous species with 7 subspecies, each with a different geo-
graphical area or habitat. Malesia: Sumatra, Peninsular Malaysia, Borneo, Philippines.

Note — Distinguishable by the minute dark brown or blackish dots, on both surfaces
of the leaves, visible with a lens.

KEY TO THE SUBSPECIES

la. Leaves 13-28 by 5.5—10 cm. Twigs at apex 4 mm diameter. Swamp forest. — Sumatra
Pee ics Eo, ent ard A eat Bs Sas lee Ray oh Slee RAEN pha e. subsp. macrophylla

b. Leaves up to 20 by 8.5 cm. Twigs at apex 1.5-3 mm diameter ............-- y
2a. Twigs 1.5—2 mm diameter, striate. Leaves chartaceous or thinly coriaceous, drying
greenish or brownish, with prominent venation above. Leaf base attenuate. Anthers
8-11. Fruits 1.5—2.5 cm; fruiting pedicel 7-15 mm long. — Lowland, but not in peat
swampiforest.: .w.. aagehiseamnelenh aeengk shiees da ae ee 3

b. Twigs 2-3 mm diameter, not or but little striate. Leaves thinly to thickly coriaceous,
drying brownish, with venation prominent or not above. Leaf base + rounded, some-
times attenuate. Anthers (7—)10—14. Fruits 1.7—4 cm; fruiting pedicel 3-9(-12) mm
long. — Lowland peat swamp forest or montane forest .........--+++++++-5 4

3a. Leaves drying greenish, with the nerves and venation paler and contrasting above.
SS sPenimsulan, Malaysia: (yc. bisce oceususysbayoee Gury ences ea a. subsp. kunstleri

b. Leaves drying more brownish, the nerves and venation not or but faintly paler, and
little contrasting above. — Philippines ..........-..-.---. f. subsp. parvifolia

4a. [Venation on upper leaf surface little prominent, distinct or indistinct. ] Fruits 1.7—2
Cun LON geatiyss cexraieero Se while do Cease SSS oat ot fog en 3

b. Fruits 3—4.5 cm long. Plant usually from montane area ........-.--+-++++-: 6
5a. Indumentum of lower leaf surface consisting of scattered scale-like hairs 0.1 mm. —
Lowland peat swamp forest or kerangas .......-...--+++++: c. subsp. coriacea

b. Indumentum of lower leaf surface consisting of mixed sessile hairs, 0.1 mm, and
few elongate hairs, to 0.3 mm. — Montane areas ...... g. subsp. pseudostellata

6a. Venation on upper leaf surface prominent and distinct; blade thinly or thickly coria-
ceous, base generally rounded. Fruits 3—4 cm, apex (subacute or) rounded. Usually

in montane forest, at (100—)900—2000 m. — Borneo .......... b. subsp. alpina

b. Venation on upper surface fine and hence indistinct; blade chartaceous (subcoria-
ceous), base attenuate. Fruits 3.5—4.5 cm long, apex (sub)acute. Hill forest at 750 m.

=, Basten iw, Kalimantan ysis). ir ota ere: eee d. subsp. leptophylla

a. subsp. kunstleri

Knema kunstleri (King) Warb., Mon. Myrist. (1897) 568, t. 25; Gamble, Mat. FI. Malay Penins. 5,
23 (1912) 241: Ridl., Fl. Malay Penins. 3 (1924) 69; J. Sinclair, Gard. Bull. Sing. 16 (1958) 291,
f. 8: 18 (1961) 236 (as var., p.p., excl. syn. Gymnacranthera cryptocaryoides Elmer and all
specimens from Borneo and the Philippines, and excl. Knema kunstleri var. surigaoensis);
W.J. de Wilde, Blumea 25 (1979) 468; 43 (1998) 244; Tree Fl. Sabah & Sarawak 3 (2000) 418.
— Myristica kunstleri King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 314, pl. 149.

Twigs 1.5—2 mm diameter, coarsely striate. Leaves chartaceous to thinly coriaceous,
elliptic(-oblong), 5—20 by 1.5—8.5 cm, base cuneate or attenuate, apex acute-acuminate;
blade drying greenish, midrib, nerves and veins paler and contrasting above; nerves

De Wilde — Myristicaceae (Knema) 279

6-14 pairs; venation prominent above. Inflorescences: sessile or all or partly with slen-
der peduncle 1-4 mm long. Male flowers: pedicel 8-11 mm long; buds 2.5 by 2.5-3 mm;
lobes reddish inside; anthers 8-11. Female flowers: pedicel 7-9 mm long. Fruits 1.5—
2.5(—3.2) by 1—-1.8(—2.5) cm; fruiting pedicel 8-15 mm long.

Distribution — Malesia: Peninsular Malaysia (Perak, Kelantan, Trengganu, Pahang,
Selangor, Negri Sembilan, Malacca, Johore).

Habitat & Ecology — Primary and degraded forest; hill sides, crests, bamboo forest;
0-900 m altitude; fl. mainly Jan.—June; fr. mainly July—Dec.

Notes — 1. Specimens may resemble subsp. parvifolia from the Philippines.

2. Shah & Noor MS 2009 and FRI 28013, from Pahang, deviate by the oblong shape
and brown drying colour of the leaves, but particularly by large fruits, 3-3.2 by 2.5—2.8
cm: MS 2007, from the same locality, is typical K. kunstleri subsp. kunstleri.

FRI 8806 (Whitmore), from N Johore, is probably of hybrid origin; it is collected at
c. 600 m altitude (with immature flowers) and deviates mainly by an oblong-lanceolate
leaf shape.

b. subsp. alpina (J. Sinclair) W.J. de Wilde

Knema kunstleri (King) Warb. subsp. alpina (J. Sinclair) W.J. de Wilde, Blumea 25 (1979) 471;
Tree Fl. Sabah & Sarawak 3 (2000) 418. — Knema cinerea (Poir.) Warb. var. alpina J. Sinclair,
Gard. Bull. Sing. 18 (1961) 287. — Type: S 45/4 (Anderson), Sarawak.

Twigs sometimes angular, (1.5—)2—3 mm diameter, faintly finely striate. Leaves (thickly)
coriaceous, elliptic-oblong to oblong-lanceolate, 4-21 by 1.5—7 cm, sometimes broadest
below the middle, base rounded to attenuate, apex acute; blade drying brown, with the
nerves not or but faintly paler, not much contrasting in colour above, lower surface
glabrescent; nerves 10-18 pairs, venation prominent above. Inflorescences sessile. Male
flowers: pedicel 5—8 mm long; buds 3 by 3—4 mm, lobes pale yellowish inside; staminal
disc flat or shallowly convex, anthers (7—)10—14. Female flowers: pedicel 2-3 mm long.
Fruits 3—4.2(—4.5) by 2-2.8 cm; fruiting pedicel (3—)5—9(—12) mm long.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, W and S Kalimantan).

Habitat & Ecology — (Sub)montane forest, mossy forest, ridges; andesite-derived
soils; (100—)900—2000 m altitude; fl. Feb., June, Oct., Nov.; fr. throughout the year.

Notes — 1. Specimens may resemble K. kinabaluensis, and can easily be noticed by
lacking minute dots on the upper and lower leaf surface.

2. § 27923 (Ilias Paie), Sarawak, differs in thinly (not thickly) coriaceous leaves and
proportionally long fruiting pedicels 7—9(—12) mm. De Vogel 851, from S Kalimantan at
c. 250 m altitude, has chartaceous leaves, non-striate twigs, with the staminal disc some-
what convex, not flat. Dransfield JD 7129 (male flowers with 7 anthers) and Prance
30585 (11 anthers), both from Brunei, have leaves glabrescent underneath; they may
represent a separate taxon.

c. subsp. coriacea (Warb.) W.J. de Wilde

Knema kunstleri (King) Warb. subsp. coriacea (Warb.) W.J. de Wilde, Blumea 25 (1979) 470; Tree
Fl. Sabah & Sarawak 3 (2000) 419. — Knema coriacea Warb., Mon. Myrist. (1897) 614. —
Myristica coriacea (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 92. — Type: Beccari 670,
Sarawak.

280 Flora Malesiana, Ser. I, Vol. 14 (2000)

=i van B96

Fig. 39. Knema kunstleri (King) Warb. subsp. leptophylla W. J. de Wilde. Habit; note the remark-

able different stages of development of the fruits on one twig [Church, Ismail, Ruskandi 2489]. —
Scale bar = 2 cm.

De Wilde — Myristicaceae (Knema) 281

Twigs 2-3 mm diameter, smooth or but faintly striate, sometimes coarsely striate or
+ angular. Leaves coriaceous, elliptic to oblong or ovate-oblong, broadest sometimes
below the middle, 6-21 by 3—7.5 cm, base rounded or attenuate, apex acute-acuminate;
blade drying light brownish, with the nerves and veins not or but faintly contrasting
above; nerves 8—18 pairs; venation prominent above. Inflorescences sessile or peduncle
stout, up to 5 mm long. Male flowers: pedicel 6—10.5 mm long; buds 2.2—2.5 by 2.5—3
mm, lobes creamy inside (always?); anthers 10-12. Female flowers not seen. Fruits
1.7—2 by 1.4-1.6 cm; fruiting pedicel 3—7(—9) mm long.

Field-notes — Flowers cream inside; fruits ochreous yellow.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah).

Habitat & Ecology — Tree with stilt-roots in primary Shorea albida peat swamp
forest; kerangas soil; 0-100 m altitude; fl. Apr.—June; fr. July—Nov.

d. subsp. leptophylla W. J. de Wilde

Knema kunstleri (King) Warb. subsp. leptophylla W.J. de Wilde, Blumea 41 (1996) 386, f. 2. —
Type: Church, Ismail, Ruskandi 2489, Kalimantan.

Twigs 1.5—2 mm diameter, early glabrescent, hairs less than 0.1 mm, faintly striate,
hardly lenticellate. Leaves chartaceous, (oblong-)lanceolate, 8-13 by 1-3 cm, base
cuneate, apex (narrowly) acute-acuminate; blade drying bright dark brown, somewhat
glossy above, lower surface with sparse brown-centred scale-like hairs less than 0.1
mm; nerves 15-17 per side, venation slightly raised, not very distinct at both surfaces.
Male inflorescences not known. Female inflorescences (immature): few-flowered, pe-
duncle 1—1.5 mm; mature buds not seen, bracteole apical, caducous. Infructescence among
the leaves. Fruits 1 per infructescence, + fusiform, with sharp edge, 3.5—4.5 by 4.5-2
cm, fruiting pedicel 7-9 mm long, broadened to above, scar of bracteole at or above
halfway. — Fig. 39.

Field-notes — Tree 25 m tall, 30 cm dbh. Outer bark reddish brown, flaking in large
patches, inner bark salmon. Sap red. Leaves glaucous beneath, dark green above, shiny.
Fruits yellowish brown.

Distribution — Malesia: Borneo, known only from the type collection, 5 km SW of
Uut Labang (W Kalimantan).

Habitat & Ecology — In sloping primary dipterocarp forest, red clay soil, semi-light
environment; seldom; 750 m altitude.

Note — The slender twigs, narrow chartaceous leaves and elongate fruits, with very
inconspicuous indumentum, give this subspecies a distinctive appearance within the com-
plex entity K. kunstleri. Also, the dots on the lower leaf surface (lens!) are finer, and less
confined to the smaller veins. Fruits on one twig are in different stages of development.
The status of subsp. /eptophylla is uncertain because it is known only from a single fruit
gathering.

e. subsp. macrophylla W. J. de Wilde

Knema kunstleri (King) Warb. subsp. macrophylla W.J. de Wilde, Blumea 25 (1979) 469. — Type:
Buwalda 6782, Sumatra.

Twigs 4 mm diameter, coarsely striate. Leaves chartaceous, elliptic-oblong, 13-28 by
5.5—10 cm, base rounded, apex acute-acuminate; blade drying (greenish) brown, with

282 Flora Malesiana, Ser. I, Vol. 14 (2000)

the nerves (not the venation) sometimes paler coloured above; nerves 8-15 pairs; vena-
tion fine, prominent above. Inflorescences sessile. Male flowers: pedicel 10-12 mm long;
buds 2—2.3 by 2.8—-3 mm; lobes reddish (?) inside; anthers 10. Female flowers and fruits
not seen.

Field-notes — Leaves white at underside. Flower buds brown-yellow.

Distribution — Possibly Thailand, and Malesia: Central Sumatra (Indragiri).

Habitat & Ecology — Lowland swamp forest; fl. May.

Note — Known only from the type collection. Closely related to subsp. kunstleri,
though distinguishable by a stout habit, with the twigs considerably thicker and the leaves
much larger, with a more brownish drying colour; the contrasting paler venation absent.
The male flowers are similar to those of subsp. kunstleri.

f. subsp. parvifolia (Merr.) W.J. de Wilde

Knema kunstleri (King) Warb. subsp. parvifolia (Merr.) W.J. de Wilde, Blumea 25 (1979) 470. —
Knema parvifolia Merr., Philipp. J. Sci., Bot. 13 (1918) 287; Enum. Philipp. Flow. Pl. 2 (1923)
184. — Type: FB 10573 (Curran), Luzon.

Twigs (1-)1.5-2 mm diameter, coarsely striate or grooved. Leaves membranous or
chartaceous, ovate-oblong to lanceolate, 4-11 by 1.5—5 cm, base cuneate to attenuate,
apex subobtuse, acute(-acuminate ); blade drying + greenish brown to bright brown above;
nerves and veins not distinctly paler and not much contrasting in colour above, nerves
8-14 pairs; venation coarse to very fine, prominent and distinct. Inflorescences sessile
or peduncle up to 4 mm. Male flowers: pedicel 7-10 mm long; buds 2.5 by 2.5—3 mm,
lobes reddish (?) inside; anthers 8-11. Female flowers not seen. Fruits 1.5—-1.7 by 1.3-
1.5 cm; fruiting pedicel 7-11 mm long.

Distribution — Malesia: Philippines (Luzon, Samar).

Habitat & Ecology — Primary dipterocarp forest; 0-300 m altitude; fl. June, Nov.—
Dec.; fr. Mar.—May.

Note — This subspecies often resembles subsp. kunstleri, from which it differs in a
number of ill-defined characters, including the thinner leaves and the lacking of much
paler contrasting venation, which is typical in subsp. kunstleri.

g. subsp. pseudostellata W.J. de Wilde

Knema kunstleri (King) Warb. subsp. pseudostellata W.J. de Wilde, Blumea 43 (1998) 244. — Type:
Church c.s. 2560, Kalimantan.

Twigs 2 mm diameter, finely striate. Leaves coriaceous, oblong, 10-15 by 2.5—4.5
cm, base subcuneate to broadly rounded; drying olivaceous-brown, lateral nerves c. 15
per side, flat and indistinct above, reticulation hardly visible; lower surface with persist-
ent open indumentum of minute stellate hairs, 0.1 mm long, mixed with dendroid hairs
0.3 mm long. Male inflorescences and flowers not known. Female inflorescences axil-
lary of leaves, sessile, wart-like, 2 mm diameter, with | (or 2) flowers. Female flowers:
pedicel 3 mm long. Fruits solitary, long-ellipsoid, 2 by 1 cm; fruiting pedicel 4-5 mm,
with bracteole scar subapical.

Field-notes — Tree 9 m tall, 10 cm dbh. Inner bark light orange. Sap red. Leaves
fleshy, glaucous beneath, dark green above. Fruits covered by rusty indumentum.

De Wilde — Myristicaceae (Knema) 283

Distribution — Malesia: Borneo (eastern W. Kalimantan, Serawai); known only from
the type specimen.

Habitat & Ecology — Primary forest; fagaceous and hill dipterocarp forest (with
Shorea, Dipterocarpua, Quercus, Lithocarpus), ridge and hillside forest; red clay soil;
not common; 1200 m altitude; fl. & fr. Oct.

Note — Subsp. pseudostellata superficially is reminiscent of K. stellata (Philippines).

25. Knema lamellaria W.J. de Wilde

Knema lamellaria W.J. de Wilde, Blumea 25 (1979) 389. — Type: FRI 3482 (Whitmore), Peninsu-
lar Malaysia.

Tree 10-20 m. Twigs 7-12 mm diameter, at first with coarse rusty hairs 1-2 mm long,
glabrescent; bark pale- or grey-brown, glabrous, not or hardly striate, coarsely flaking.
Leaves chartaceous or coriaceous, oblong(-lanceolate), 45—65 by 13—24 cm, apex bluntish
to acute-acuminate, base broadly rounded or cordate, with the basal nerves + crowded,
when immature with dense hairs 2 mm long, early glabrescent, dull greenish to brown
above; lower surface glabrous, faintly papillate or not; dots absent; midrib stout, raised
above; nerves 21-30 pairs, raised; venation distinct; petiole at first with dense rusty
hairs 1-2 mm long, late glabrescent, 15-20 by 8-10 mm. Inflorescences: sessile, brachy-
blast simple or warted, 10—15 mm diameter, in male (5—)10—25-flowered, female 2—6-
flowered; flowers with dense + woolly hairs 0.8—1 mm long, for the larger part early
glabrescent or easily rubbed off; perianth 3-lobed, inside at base ridged or coarsely stri-
ate, reddish (?). Male flowers: pedicel 16-21 mm long, bracteole subpersistent, about
median; buds + depressed globose-obovoid, 7 by 8 mm, cleft 3/4—4/5, lobes 0.5—-1 mm
thick; staminal disc circular, flat, 3.5 mm diameter; anthers c. 20, subsessile or just stiped,
nearly horizontal, 0.6 mm, not touching; the androphore tapering, 1.5 mm long, striate.
Female flowers not seen. Fruits 1-3 per infructescence, obovoid or oblong, 4—4.5 by
2-2.6 cm, with dense coarsely woolly brown hairs 3—5 mm long, caducous or easily
rubbed off; dry pericarp 3 mm thick; fruiting pedicel 18—25 mm long, with the scar of
the bracteole at or somewhat above halfway.

Field-notes — Bark black-brown, with brittle rectangular flakes; copious red sap;
slash wood white.

Distribution — Malesia: Peninsular Malaysia (Pahang, Trengganu).

Habitat & Ecology — Flat or undulating lowland forest, once at base of limestone
rock; 100—200 m altitude; fl. Mar.; fr. June—July.

Note — Knema lamellaria is related to K. lampongensis from Sumatra and K. pallens
from Borneo; it resembles also K. furfuracea, with smaller flowers.

26. Knema lampongensis W. J. de Wilde
Knema lampongensis W.J. de Wilde, Blumea 25 (1979) 388. — Type: Iboet 159, Sumatra.

Tree 10-15 m. Twigs 5—10 mm diameter, at first with dense conspicuous rusty hairs
1-2.5 mm long, glabrescent; bark hardly striate, lower down reddish brown to blackish,
conspicuously flaking. Leaves chartaceous or coriaceous, oblong(-oblanceolate), 23-36
(—56) by 5—12(—14) cm, apex acute(-acuminate), base rounded to cordate, with the basal
nerves somewhat crowded; greenish brown, and + glossy above; lower surface glabrous,

284 Flora Malesiana, Ser. I, Vol. 14 (2000)

finely papillate; dots absent; midrib distinctly raised above; nerves 17-25 pairs, raised;
venation fine or coarse, faint; petiole late glabrescent, 10—16 by 4-5 mm. Inflorescences:
sessile, brachyblast simple or up to 7-furcate or warted, 5—15 mm long, in male 2—5
(~30)-flowered, female 2—5-flowered; flowers with persistent brown hairs 0.7—1.2 mm
long; perianth 3-lobed, at base somewhat thickened and coarsely striate or grooved into
separate pads, red(?) inside. Male flowers: pedicel 7-9 mm long, bracteole caducous or
subpersistent, about median or below; buds + depressed globose-obovoid, 5—6 by 5—6.5
mm. cleft 2/3—3/4, lobes 1 mm thick; staminal disc faintly triangular, flat, 2-2.5 mm
diameter: anthers 13 or 14, subsessile, almost horizontal, 0.5—0.7 mm, not touching;
androphore tapering, striate, 1-1.5 mm long. Female flowers not seen. Fruits | or 2 per
infructescence, obovoid or subglobose, 2.5 by 2 cm, with dense dark brown hairs 0.6—1
mm long; dry pericarp 2.5 mm thick; fruiting pedicel 10 mm long; stigma remnants
many-lobulate, sessile.

Distribution — Malesia: Sumatra (E Coast, Palembang, Lampong, Riau Arch.);
Anambas Is.

Habitat & Ecology — Lowland and hill forest, 100—300 m altitude; fl. Apr., Nov.

Note — Knema lampongensis is more or less intermediate between the species K. fur-
furacea, K. lamellaria, and K. pallens, morphologically as well as geographically; for
differentiating characters see the key to the species.

27. Knema latericia Elmer

Knema latericia Elmer, Leafl. Philipp. Bot. 5 (1913) 1815; J. Sinclair, Gard. Bull. Sing. 16 (1958)
278, f. 3, pl. IIA; 18 (1961) 238, f. 15B, C, G, excl. var. lunduensis J. Sinclair; W.J. de Wilde,
Blumea 25 (1979) 395: 43 (1998) 245; Tree Fl. Sabah & Sarawak 3 (2000) 419. — Type: Elmer
12757, Philippines, Palawan.

For more references and synonyms see the subspecies and varieties.

Erect shrub or tree 2—20 m. Twigs (1—)2—4 mm diameter, sometimes + angular, at first
with rough, rusty, grey-brown, or yellowish brown hairs 0.5—2 mm long, glabrescent;
bark striate or fissured, cracking, later on coarsely or finely flaking, or not. Leaves mem-
branous or thinly coriaceous, oblong to (ob)lanceolate (broadest below the middle in
subsp. latericia), 8-30 by 1.5—9 cm, apex acute(-acuminate), base attenuate to rounded,
drying brown or greenish above; lower surface glabrous, grey(-brown) or whitish, not
obviously papillate; dots absent; midrib raised above (when dry reddish in subsp. ridleyi,
pale brown or whitish in subsp. albifolia); nerves 9—25(—30) pairs, raised; venation faint
or distinct; petiole (5—)10—18 by 1.5-3.5 mm. Inflorescences: sessile, brachyblast sim-
ple, warted or furcate, up to 10 mm long, in male 3-20-flowered, female 2—10-flowered;
flowers with pale brown to rusty hairs 0.2—0.7 mm long; perianth 3- (or 4-)lobed, red
inside. Male flowers: pedicel 2-6 mm long, bracteole caducous, median to subapical;
buds depressed obovoid (subsp. albifolia) or depressed globose, 3-3.5 by (3-)4-4.5
mm, cleft 2/3—4/5, lobes 0.5(—1) mm thick; staminal disc circular, flat, + convex, or
low-mammillate, 2 mm diameter; anthers 9-12, shortly stiped, 0.5 mm long, horizontal,
not touching; androphore | mm long. Female flowers: pedicel to 1.5 mm long, bracteole
subapical; buds much larger than in male, obovoid, 5—8 by 4-5 mm, cleft 1/3—2/3, lobes
0.5—1 mm thick; ovary subglobose to ovoid, 2.5-3 mm long; the stigma subsessile,
+ 2-lobed and each lobe again deeply (3—)4—6-serrate. Fruits 1—4 per infructescence,

Nn

De Wilde — Myristicaceae (Knema) 28

broadly ellipsoid to obovoid, sometimes subglobose, apex rounded, (1—)1.5—3 by 1-2.5
cm, with rough hairs 1-2 mm long; dry pericarp (1—)2—3 mm thick, the fruiting pedicel
2-5 mm long.

Field-notes — Slender tree, pyramidal in outline; trunk without buttresses; bark (red-
dish) brown or chocolate, scaly or flaking in fairly large elongate portions, not furrowed;
inner bark pink or reddish, sapwood white, pale yellow, or pale brown, sometimes with
streaks. Leaves dark green with whitish green midrib above, glaucous beneath. Flowers
outside yellow brown or rusty, red inside; disc pink. Fruits yellow brown or dark rusty.

Distribution — Peninsular Thailand; in Malesia: Sumatra (Bangka, no specimens seen),
Peninsular Malaysia, Singapore, Borneo, Palawan.

Habitat & Ecology — Primary and degraded forest; on hill sides, ridges, etc.,
dipterocarp forest, also in kerangas; found on a variety of soils: sand, loam, clay, black
and brown soils, limestone, sandstone, basalt hills, well-drained soils, and leached soils;
0—1500 m altitude; fl. & fr. throughout the year, in Borneo mainly May—Oct.

Note — Knema latericia is a complex species in which three subspecies are recog-
nized. Subsp. latericia (with 2 varieties), from Palawan, has twigs with non-flaking
bark, and therefore K. latericia appears twice in the general key to the species.

KEY TO THE SUBSPECIES

la. Bark of older twigs not or hardly longitudinally cracking, nor flaking. Twig apex
coarsely rusty pubescent, striate or ridged, + angled. [Male buds pear-shaped; staminal
discemilat above. |'— Palawan *.<.. «2 haesh Pact pee shee a. subsp. latericia

b. Bark of older twigs usually longitudinally cracking, later on (finely) flaking (not
flaking in subsp. ridleyi forma nana). Twig apex variably hairy, striate or not, not
PMMA cage so o's Saas Mat onent ke Re oes SE ROU ore ok. onan eee cia 2
2a. Indumentum of twig apex and flowers dark brown. Male buds obconical or globose,
often somewhat broader than long. Leaves usually large, 4-10 cm broad, drying
dark (greenish) brown above, pale brown or grey-glaucous beneath. Fruits variable

of size, with brown to rusty-reddish indumentum. — Peninsular Thailand and Ma-
laysiay Bancka, Bomeo. |i 3a ee +s -lae ae odes ie ee c. subsp. ridleyi

b. Indumentum of twig apex and flowers pale or yellowish brown. Male buds obovoid.
Leaves generally smaller, 11-22 by 2-5 cm, drying pale (greenish) brown above,
whitish beneath. Fruits 1.4—2(-3.5) cm long, with pale or yellowish brown indu-
MMCMUIM—— DOMED, <2 2 ssyarso Malas tacay es * ee eve eee b. subsp. albifolia

a. subsp. latericia

Knema latericia Elmer subsp. latericia; W.J. de Wilde, Blumea 25 (1979) 396; 39) (AIS 7) Tale stk Le
— Knema latericia Elmer var. latericia; J. Sinclair, Gard. Bull. Sing. 18 (1961) 240, p.p., for the

type only.

Twigs + angled, at apex roughly rusty tomentose; bark coarsely striate, not or hardly
finely cracking, nor flaking. Leaves 5—25 by 1-6 cm, drying dark brown with brownish
midrib, with fine prominent reticulation above, greyish beneath. Male flowers with dense
coarse dark rusty hairs 0.7 mm long, pedicel 1-2 mm long with bracteole subapical to
median, persistent; buds depressed-obovoid to pear-shaped, 3.5 by 3.5 mm, cleft c. 2/3,

286 Flora Malesiana, Ser. I, Vol. 14 (2000)

RS Uae OT
eit are } iro

wer

Jv. 86

De Wilde — Myristicaceae (Knema) 287

lobes 3, red inside, 0.5 mm thick; staminal disc faintly triangular, flat, 1.5—2 mm diam-
eter: anthers 7-9, shortly stiped, horizontal, 0.5 mm long, mutually not touching;
androphore 1—1.5 mm long. Female buds: 5.5 mm long, sessile. Fruits broadly ellip-
soid, 1.5—2 cm long, with rusty indumentum.

Note — Delicate specimens are segregated as variety subtilis.

al. var. latericia

Twigs at apex 2-4 mm diameter; bark not or hardly cracking, not flaking. Leaves
chartaceous, 7-25 by 2.5—6 cm, lateral nerves 11-18 pairs. Fruits 2. cm long, hairs 0.5—1
mm long. — Fig. 40a—d.

Distribution — Malesia: Philippines (Palawan).

Notes — 1. Resembles vegetatively (leaves) K. tomentella (Moluccas); the twig apex
is reminiscent of K. korthalsii (Palawan and Borneo).

2. In the obovoid male buds subsp. /atericia agrees with subsp. albifolia, rather than
with subsp. ridleyi which has depressed-globose male buds.

a2. var. subtilis W. J. de Wilde

Knema latericia Elmer subsp. latericia var. subtilis W.J. de Wilde, Blumea 32 (1987) INS? fe te:
41 (1996) 388. — Type: SMHI 1642 (Ridsdale c.s.), Palawan.

Twigs 1-1.5 mm diameter; bark of older twigs striate, not flaking. Leaves membra-
nous, small, elliptic to lanceolate, 5—13.5 by 1—4.5 cm, lateral nerves 8-12 pairs. Fruits
ellipsoid, 1.5—2 by 1.2 cm, indumentum of rough hairs 0.5(—1) mm long. — Fig. 40e.

Field-notes — Small tree, leaves pallid blue-green at underside, fruits brown.

Distribution — Malesia: Philippines (Palawan); possibly Sarawak and Brunei, see
note 2.

Habitat & Ecology — On primary and logged ridges in broad-leaved dipterocarp
forest, once recorded from limestone; 0-100 m altitude; fl. & fr. Apr.

Notes — 1. The occurrence of var. subtilis may be related to the nature of the bed-
rock, possibly limestone. It much resembles the delicate species K. stenocarpa from the
Sulu Is. and Mindanao.

2. Some delicate specimens from Brunei and Sarawak, discussed by De Wilde (1996),
approach var. subtilis.

b. subsp. albifolia (J. Sinclair) W.J. de Wilde

Knema latericia Elmer subsp. albifolia (J. Sinclair) W.J. de Wilde, Blumea 25 (1979) 397; 43
(1998) 246, f. la; Tree Fl. Sabah & Sarawak 3 (2000) 420. — Knema latericia Elmer var. albifolia
J. Sinclair, Gard. Bull. Sing. 18 (1961) 243, f. 15C. — Type: Sinclair 9269, Sabah.

Fig. 40. Knema latericia Elmer subsp. latericia var. latericia. a. Habit of male flowering branchlet,
note finely striate bark, not flaking; b. male bud; c. ditto, longitudinally opened, showing position
of androecium: d. androecium. — K. latericia subsp. latericia var. subtilis W.J. de Wilde. e. Fruit-
ing branchlet [a—d: SMHI 693 (Podzorski); e: SMHI 1642 (Ridsdale)|. — Scale bar for a, e = 2 cm;
for b, c = 1.6 mm; for d = 0.9 mm.

288 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Knema) 289

Twigs 1-2 mm diameter, with fine, pale grey-brown or yellowish brown scurfy
indumentum; bark coarsely striate or ridged, later on longitudinally cracking and flak-
ing. Leaves narrow, 11-22 by 2-5 cm; drying pale (greenish) brown above, midrib pale,
venation fine, distinct, whitish beneath. Male flowers: pedicel short, with pale hairs;
buds more or less depressed obovoid; staminal disc rather flat or slightly convex; an-
thers 10. Female flowers: buds 5 mm long. Fruits (sub)sessile, subglobose, ellipsoid, or
obovoid, 1.4—2(—3.5, see note) cm long, with bright (yellowish) brown indumentum. —
Fig. 41a.

Distribution — Malesia: Borneo (Sarawak; S & E Kalimantan, Nunukan Is.; Sabah).

Note — This subspecies is marked by a paler colour in diverse parts, and narrower
leaves; the shape of the male buds is depressed obovoid, not depressed (sub)globose,
and the staminal disc is rather flat, not (low) mammillate. Several specimens apparently
seem to be + intermediate with subsp. ridleyi. The collection Argent & Saridan 9333
from Central Kalimantan differs in its large fruits, 3.5 cm long.

c. subsp. ridleyi (Gand.) W.J. de Wilde

Knema latericia Elmer subsp. ridleyi (Gand.) W.J. de Wilde, Blumea 25 (1979) 396; 27 (1981)
223: 43 (1998) 246; Tree Fl. Sabah & Sarawak 3 (2000) 420. — Myristica ridleyi Gand., Bull.
Soc. Bot. Fr. 66 (1919) 226, in clavi. — Type: Ridley s.n., Singapore.

Knema conferta (King) Warb. var. borneensis Warb., Mon. Myrist. (1897) 580, p.p. — Myristica
conferta King var. borneensis (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 1. — Lectotype:
Beccari 2003, Sarawak.

Knema meridionalis J. Sinclair, Gard. Bull. Sing. 13 (1951) 297, f. 1. — Type: SF 38561 (Sinclair),
Singapore.

Knema latericia Elmer var. latericia auct.: J. Sinclair, Gard. Bull. Sing. 18 (1961) 240, f. 15B, F, G,

p.p., excl. type.
[Knema elongata Warb., nom. nud. in sched. ]

Twigs 1.5—4(—8) mm diameter, variably rusty tomentose, not angled; bark striate,
lower down longitudinally cracking and flaking (not flaking in forma nana). Leaves
4—10 cm broad; drying dark brown with brownish midrib above, venation distinct or not,
grey-brown beneath. Male flowers: pedicel 5 mm long; buds obconical or globose; sta-
minal disc flat or low-mammillate; anthers 9-12. Female flowers: buds 5—8 mm long;
pedicel up to 1.5 mm long. Fruits variable, (1—)1.5—3 cm long, with coarse rusty indu-
mentum.

Distribution — Peninsular Thailand (one deviating specimen); in Malesia: Sumatra
(N Sumatra, E Coast, Bangka), Peninsular Malaysia (Kedah, Perak, Trengganu, Johore),
Singapore, Borneo (Sarawak, S & SE, E & NE Kalimantan, Sabah).

Note — Robust sterile forms can easily be confused with small-leaved specimens of
the related K. lunduensis.

Fig. 41. Some taxa of Knema likely to be confused in the vegetative state. — a. K. latericia Elmer
subsp. albifolia (J. Sinclair) W.J. de Wilde, mature fruit. — b. K. latericia subsp. ridleyi (Gand.)
W.J. de Wilde forma nana W.J. de Wilde, mature fruit. — c. K. minima W.J. de Wilde, fruit slight-
ly immature. — d. K. tridactyla Airy Shaw aff. subsp. sublaevis W.J. de Wilde [a: SAN 38982;
b: Wong WKM 1411; c: KeBler c.s. 394; d: Wong WKM 1366]. — Scale bar for all = 2 cm.

290 Flora Malesiana, Ser. I, Vol. 14 (2000)

KEY TO THE FORMS

la. Bark of older twigs thinly flaking. Leaves (15—)20-35 by 4-9cm ..........- 2,
b. Bark not flaking. Leaves smaller, 6-15 by 1—1.5(—2) cm, lanceolate. — E Sarawak,
Brine) <:ots te Aya ee decks Oost 3e a esa he oeke c2. subsp. ridleyi forma nana

2a. Plants rather stout. Leaves drying olivaceous. — E Sabah, SE Kalimantan........
ee Re I PE Se errr ec enor c3. subsp. ridleyi forma olivacea

b. Plants of variable habit. Leaves drying (dark) brown. — S Thailand, Peninsular Ma-
laysia,; Sumatra, BOmMeO .:<.css aie gel -fee) opp) 222 cl. subsp. ridleyi forma ridleyi

cl. subsp. ridleyi forma ridleyi

Description as the subspecies, excluding the main differentiating characters as in the
key to the forms.
Distribution — S Thailand; Malesia: Sumatra, Peninsular Malaysia, Borneo.

¢2. subsp. ridleyi forma nana W. J. de Wilde

Knema latericia Elmer subsp. ridleyi (Gand.) W.J. de Wilde forma nana W.J. de Wilde, Blumea
43 (1998) 246, f. 1b. — Type: Anderson K 10, Sarawak.

Tree 7-18 m. Twigs towards apex 1.5-2 mm diameter, at first with rusty indumentum,
hairs 0.5—1 mm long; bark of older twigs finely striate, not flaky. Leaves thinly coriaceous,
(oblong-)lanceolate, 6-15 by 1-1.5(-2) cm, base (narrowly) attenuate, drying (light)
brown above, pale brown beneath. Male perianth buds 3 by 4 mm, pedicel 2—2.5 mm
long; anthers 6 or 7. Female perianth 4 mm long. Fruits obovoid, 1.3—1.8 by 1—1.2 cm,
with rusty indumentum of rough hairs 1 mm long. — Fig. 41b.

Field-notes — Bark smooth or slightly flaky. Leaves glaucous below.

Distribution — Malesia: Borneo (E Sarawak: Mt Mulu area; Brunei, Sabah).

Habitat & Ecology — Hill dipterocarp forest, ridge and hillside forest, sandy-clay
soil; 300-550 m altitude; fl. July; fr. July, Sept., Oct.

c3. subsp. ridleyi forma olivacea W.J. de Wilde

Knema latericia Elmer subsp. ridleyi (Gand.) W.J. de Wilde forma olivacea W.J. de Wilde, Blumea
43 (1998) 248. — Type: Endert 3342, Borneo, W. Kutai.

Tree 3-10 m. Twigs at apex 3—4(—6) mm diameter, with dense rusty or light brown
indumentum with hairs 0.5(-1) mm long; bark lower down longitudinally cracking and
flaking. Leaves thinly coriaceous, (15—)20-35 by 4-9 cm; upper surface drying olivaceous,
lower surface pale grey-brown; nerves 14—16(—22?) pairs, yellowish brown, contrast-
ing; petiole late glabrescent. Inflorescences, male and female flowers as in the type form
of subsp. ridleyi. Fruits ellipsoid, 2—3.5 cm long, indumentum of hairs (0.5—)1 mm long.

Field-notes — Treelet, bark smooth or scaly; living bark 13 mm, light red; leaves
blue-green below. Flowers rusty, inside pink. Fruits brown.

Distribution — Malesia: Borneo (Sabah, E & SE Kalimantan).

Habitat & Ecology — Flat and hilly country; in forest on sand- or limestone; altitude
10-700 m; fl. July, Nov.; fr. June, Sept.

De Wilde — Myristicaceae (Knema) 291

Note — Specimens of forma olivacea were previously included in subsp. ridleyi, but
differ in the distinctive pale greenish drying colour of the leaves. The fruits possibly are
slightly larger than those in forma ridleyt.

28. Knema latifolia Warb.

Knema latifolia Warb., Mon. Myrist. (1897) 610, t. 25; J. Sinclair, Gard. Bull. Sing. 18 (1961) 244,
f. 16: W.J. de Wilde, Blumea 25 (1979) 414, f. 7; Tree Fl. Sabah & Sarawak 3 (2000) 421. —
Syntypes: Forbes 2997, 3188, Sumatra; Beccari 1991, Sarawak.

Knema umbellata Warb., Mon. Myrist. (1897) 609. — Myristica umbellata (Warb.) Boerl., Handl.
Fl. Ned. Ind. 3, 1 (1900) 92, nom. inval. — Type: Beccari 1894, Sarawak.

Knema nitida Merr., J. Straits Branch Roy. Asiat. Soc. no 85 (1922) 190. — Type: Ramos 1530,
Sabah.

Knema winkleri Merr., J. Straits Branch Roy. Asiat. Soc. no 85 (1922) 189. — Type: Winkler 2390,
Kalimantan.

Tree 4-20 m. Twigs (1-)2—3 mm diameter, at first with minute greyish hairs 0.1 mm
long, early glabrescent; bark finely or coarsely striate, sometimes faintly cracking, not
flaking. Leaves (thinly) coriaceous, (elliptic-)oblong to obovate-oblong, (7—)10-—30 by
(2.5-)4.5-12.5 cm, apex broadly acute(-acuminate) or subobtuse, base attenuate to
rounded; greenish brown to blackish, often glossy above; lower surface grey or pinkish
brown, at first with sparse stellate scale-like hairs 0.1 mm long, early glabrescent; dots
absent; midrib + raised above; nerves 8—16 pairs, slightly raised above; venation distinct
above; petiole 15-30 by 2-4 mm. Inflorescences: sessile or pedunculate up to 5 mm,
brachyblast simple or 2—4-furcate, to 8 mm long; in male 5—15-flowered, female 1—5-
flowered; flowers with grey-brown stellate hairs 0.1(—0.2) mm long or less, subglabres-
cent; perianth 3- (or 4-)lobed, pale yellowish inside (always?). Male flowers: pedicel
5-10 mm long, bracteole subpersistent, subapical; buds depressed globose or broadly
(ob)ovoid, + trigonous, 2.5-3.5 by 3.5—5 mm, cleft 4/5(—5/6), lobes 0.5—1 mm thick;
staminal disc sharply 3- (or 4-)angled, flat or slightly convex, 2-2.5 mm diameter; an-
thers 3 (or 4), sessile below the angles of the disc, horizontal, 0.5—1 mm long, not touch-
ing; androphore slender, 0.5—1 mm long. Female flowers: pedicel (4-)5—10 mm long,
bracteole median or above; buds ellipsoid-obovoid, with a faint constriction below the
middle, 5—7 by 4 mm, cleft 2/3—3/4, lobes 1 mm thick; ovary ovoid, 2—2.5 mm long;
style 0.5—0.8 mm long; stigma fleshy, + 2-lobed and each lobe again (2—)3-5-lobulate.
Fruits | or 2 per infructescence, globose-ellipsoid or ellipsoid-oblong, somewhat flat-
tened, often ridged, often somewhat saccate at base, 2—3.5 by 2-3 cm; dry pericarp 2-3
mm thick, with grey to rusty hairs 0.1-0.2 mm long, subglabrescent; fruiting pedicel
7-20 mm long. — Fig. 34c, 42.

Field-notes — Bole without buttresses; crown cylindrical or pyramidal; bark nearly
smooth, peeling in narrow strips; inner bark brittle, pinkish, red, or light brown; wood
off-white, pale yellow, or pinkish. Leaves very glossy green above, grey-green or glau-
cous beneath; midrib yellow-green, drying brown to red-brown beneath. Flowers out-
side (yellowish) green, yellowish inside. Fruits yellow or (orange-)brown, the furry greyish
to rusty.

Distribution — Malesia: Sumatra (Palembang) and Borneo (Kalimantan, Nunukan
Is., Sarawak, Brunei, Sabah).

292 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 42. Knema latifolia Warb. a. Twig with male inflorescences; b. male bud; c. opened male bud
showing the androecium; d. androecium, seen from + below; e. female bud; f. opened female bud
showing the pistil; g. infructescence [a—d: SAN 51920; e, f: S 15642; g: SAN 82573]. — Scale bar
for a, g = 2 cm; for b, e, f = 2.8 mm; for c = 1.4 mm; for d = 0.66 mm.

De Wilde — Myristicaceae (Knema) 293

Habitat & Ecology — Primary or degraded lowland forest; forest on hill sides, ridges,
low undulating ground, sometimes swamp forest; mixed dipterocarp forest, Agathis for-
est; on loam-, clay-, sand-, acid-, and lime-containing soils; 0-500 m altitude; fl. through-
out the year; fr. mainly Aug.—Dec.

Notes — 1. Knema latifolia is a well-defined species characterized by only 3 (or 4)
anthers, one at each angle of the triangular staminal disc. The fruits are somewhat flat-
tened, ridged, and saccate at the base. The leaves are dry dark brown or blackish, glossy
above, with distinct coarse venation.

2. Winkler 2390, type of synonym K. winkleri, contains some 4-merous flowers, with
a 4-angular disc and 4 anthers.

29. Knema laurina (Blume) Warb.

Knema laurina (Blume) Warb., Mon. Myrist. (1897) 606, t. 24 (excl. var. malayana, p.p.; incl. var.
amboinensis, bancana, borneensis), W.J. de Wilde, Blumea 25 (1979) 379; Tree Fl. Sabah &
Sarawak 3 (2000) 421. — Myristica laurina Blume, Rumphia 1 (1835) 189, t. 61. — Type:
Blume s.n., Java.

For more references and synonyms see the varieties.

Shrub or tree 3—20(—30) m. Twigs 2-5 mm diameter, at first with dense, rough, rusty
hairs 0.5-2 mm long, late glabrescent; bark smooth or striate, not tending to crack or
flake. Leaves membranous or coriaceous, obovate-elliptic to oblong-lanceolate, 9-30 by
2.5—10.5 cm, base attenuate, rounded, obtuse, rarely subcordate, apex subobtuse to acute-
acuminate; drying greenish to brown above, lower surface finely papillate, with persist-
ent + equally long-stalked dendroid hairs 1 mm long (var. laurina), or hairs of mixed
sizes, sessile and stalked dendroid (var. heteropilis); dots absent; midrib + raised above,
late glabrescent; nerves 12-28 pairs, flat or sunken above; venation raised, clearly vis-
ible above; petiole late glabrescent, 10-20 by 1.5-3.5 mm. Inflorescences: sessile or
pedunculate up to 0.5 mm, brachyblast simple or 2-fid, to 7 mm long, in male 5—20-
flowered, female 3—8-flowered; flowers with dense rusty hairs 0.5—1.5(—2) mm long;
perianth 3-lobed, pink or red inside. Male flowers: pedicel (0.5-)2—5 mm long, bracteole
subpersistent or caduous, (nearly) apical; buds obovoid(-oblong), 3-5(—6) by 2—4 mm,
cleft c. 1/2, lobes 0.5—1 mm thick; staminal disc circular, flat or slightly concave, 1—1.5
mm diameter; anthers 6—9, sessile, (sub)erect, 0.3-1 mm long, not or little touching; an-
drophore somewhat tapering, 1.5-2.5 mm long, rarely minutely pubescent at the base.
Female flowers: pedicel to 1 mm long, bracteole apical; buds ovoid(-oblong), 4-6 by
2.5-3.5 mm, cleft c. 1/2, lobes 0.5(—1) mm thick; ovary subglobose, (1.5—)2—2.5 mm
diameter; style slender, including 2-lobed stigma (1—)1.5—2 mm long, each lobe = erect,
4-lobulate. Fruits 1-5 per infructescence, ovoid or ellipsoid(-oblong), apex obtuse to
subacute, 1.5—2.5(—3) by 1-2 cm, with rusty hairs 1-3 mm long; dry pericarp 1.5—-2 mm
thick; fruiting pedicel 1(—5) mm long.

Field-notes — Bark grey or (reddish) brown, essentially smooth but fine scaly or
slightly cracked, sometimes furrowed; inner bark reddish; sapwood white, yellowish,
red with brown rays, light brown, or yellow with brown stripes. Leaves glossy above,
glaucous with yellowish veins beneath. Flowers pink or red inside. Fruits up to 3.5 by 3
cm, yellow, golden brown, or red-brown hairy.

294 Flora Malesiana, Ser. I, Vol. 14 (2000)

Distribution — Peninsular Thailand; in Malesia: Sumatra (incl. Simeulue, Siberut,
Bangka), Peninsular Malaysia, W and C Java, Borneo (incl. Anambas, Karimata, and
Nunukan Is.).

Habitat & Ecology — Primary and degraded rain forest; mixed dipterocarp forest;
found on a variety of soils: sand, sandstone, sandy ridges, granitic sand, basalt-derived
soils, black soil, yellow clayey soil, and shales; 0O—1000(—1500) m altitude; fl. & fr.
throughout the year.

Note — Knema laurina may be confused with other species, e.g. K. pseudolaurina
and K. conferta, with persistent indumentum on the lower leaf surface. Knema pseudo-
laurina has mainly sessile stellate hairs. Knema conferta differs essentially in the shape
of the male buds, and by the dark dots on the lower leaf surface.

KEY TO THE VARIETIES

la. Lower leaf surface with predominantly stalked (stellate-)dendroid hairs, of almost

equal size and shape. Fruits 1.5—2.5(-3) cm long .............. a. var. laurina
b. Lower leaf surface with sessile stellate and stalked (stellate-)dendroid hairs of mixed
size and shape. Fruits 1:5—2/cmlomg ..<...-.----52---5-: b. var. heteropilis

a. var. laurina

Knema laurina (Blume) Warb., Mon. Myrist. (1897) 606, t. 24 (excl. var. malayana, p.p.; incl. var.
amboinensis, bancana, borneensis),; Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 248; Koord.,
Exk. Fl. Java 2 (1912) 258: Ridl., Fl. Malay Penins. 3 (1924) 72; J. Sinclair, Gard. Bull. Sing. 16
(1958) 329, f. 19, p.p.; ibid. 18 (1961) 248, p.p., excl. most synonyms; W.J. de Wilde, Blumea
25 (1979) 379: Tree Fl. Sabah & Sarawak 3 (2000) 421. — [Myristica tomentosa auct. non
Thunberg: Blume, Bijdr. 2 (1825) 577.] — Myristica laurina Blume, Rumphia 1 (1835) 189,
t. 61: A.DC., Prod. 14, 1 (1856) 206; de Vriese, Pl. Ind. Bat. Orient. (Pl. Reinw.) (1857) 96;
Mig.. Fl. Ind. Bat. 1, 2 (1858) 70, & Suppl. 1 (1861) 385; Hook. f., Fl. Brit. India 5 (1886) 112;
King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 319, pl. 156; Koord. & Valeton, Med. Lands PI. Tuin
17 (1896) 186.

Myristica laurina Blume var. longifolia Miq., Fl. Ind. Bat. 1, 2 (1858) 71, & Suppl. 1 (1861) 385. —
Syntypes: Teijsmann s.n., 484, Sumatra.

Myristica laurina Blume var. borneensis Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 51. — Type:
Korthals s.n., Kalimantan.

Myristica cantleyi Hook. f., Fl. Brit. India 5 (1886) 110, not sensu King and later authors. — Knema
cantlyi (Hook. f.) Warb., Mon. Myrist. (1897) 554, p.p., for the type only. — Type: Cantley 195,
Singapore.

Myristica laurina Blume var. bancana (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 92, nom. alt.,
inval. — Type not indicated.

Myristica laurina Blume var. amboinensis (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 92, nom.
alt., inval. — Type: Beccari s.n., introd. Hort. Bogor.

Twigs at apex with hairs 0.5-2 mm long, lower down smooth, not striate. Leaves on
lower surface with persistent hairs all or predominantly stalked (stellate-)dendroid, of
almost equal size and shape. Mature male flowers: pedicel 2-5 mm long, bracteole (sub)
apical; buds 3.5-5(—6) by 2-3.5(—4) mm, with hairs 0.5-1.5(—2) mm long; anthers 6-9.
Fruits 1.5—2.5(—3) cm long; fruiting pedicel 0—1(—5) mm long.

Distribution — As the species.

Nn

De Wilde — Myristicaceae (Knema) 29

Notes — 1. Var. malayana Warb. is excluded by its lectotype, which is K. pseudo-
laurina. Var. amboinensis Warb. is included in the synonymy; its type, Beccari s.n., was
originally collected in the botanical garden at Bogor, but cannot have originated from
Ambon, as explained by Sinclair (1961: 254).

2. Specimens from Peninsular Malaysia sometimes have relatively large fruits, up to
2.5 cm long; in Borneo fruits may reach a length of 30 mm.

b. var. heteropilis W. J. de Wilde

Knema laurina (Blume) Warb. var. heteropilis W.J. de Wilde, Blumea 25 (1979) 381. — Knema
furfurascens Gand., Bull. Soc. Bot. Fr. 66 (1919) 226, in clavi. — Type: Junghuhn s.n., 1855,
Java.

Twigs at apex with dense rusty hairs | mm long, (coarsely) striate. Leaves on lower
surface with persistent hairs of variable size and shape. Male flower: pedicel 0.5—2.5
mm long, bracteole apical; buds 3-3.5 by 2—2.5 mm, with hairs 1-1.5 mm long; anthers
7 or 8. Fruits 1.5(—2) cm long; fruiting pedicel | mm long.

Distribution — Malesia: Sumatra (Simeulué I.), Peninsular Malaysia (Kedah, Perak,
Selangor, Negri Sembilan), W Java.

Notes — 1. Specimens of var. heteropilis from Peninsular Malaysia and Java have
somewhat narrower leaves as compared to those from Simeulué I. and to those generally
found in the type variety. The specimens from Simeulué I. (Achmad 63, 1195, 1 808), are
stout, with large, broad leaves, strong twigs at apex 4 mm diameter, and fruits 2 by 1.2
cm; typical var. /aurina is known from Siberut (Iboet 178).

2. The specimen Liitjeharms 4421, from Enggano, is possibly of hybrid origin, in the
vicinity of K. glaucescens and K. losirensis.

30. Knema linguiformis (J. Sinclair) W.J. de Wilde

Knema linguiformis (J. Sinclair) W.J. de Wilde, Blumea 25 (1979) 416, f. 8; 27 (1981) 227; 32
(1987) 121; Tree Fl. Sabah & Sarawak 3 (2000) 423. — Knema curtisti (King) Warb. var.
linguiformis J. Sinclair, Gard. Bull. Sing. 18 (1961) 200, f. 6B. — Type: Sinclair (Kadim & Kapis)
9294, Sabah.

Tree 6-25 m. Twigs 1-2 mm diameter, sometimes + angled, at first with pale brown to
dark rusty woolly hairs 0.2—0.5 mm long, late glabrescent; bark pale, yellowish to brown,
+ striate, not tending to crack or flake. Leaves chartaceous or thinly coriaceous, oblong
to lanceolate, 5-15 by 1.5—5(—6) cm, base attenuate to rounded, apex acute to obtuse,
rarely subacuminate; drying greenish above; lower surface greenish grey, minutely pap-
illate, early glabrescent; dots absent; midrib raised above; nerves 8—20 pairs, raised above:
venation distinct above; petiole late glabrescent, 6-15 by I-1.5 mm. /nflorescences:
sessile, brachyblast simple or forked, 2-5 mm long; in male 5—20-flowered, female
2-10-flowered; flowers with persistent (yellowish) rusty hairs (0. 1—)0.2—0.4 mm long;
perianth 3- (or 4-)lobed, pink inside. Male flowers: pedicel 5.5-13 mm long, bracteole
persistent or late caducous, about median; buds depressed globose to broadly obovoid,
rounded or obtusely trigonous in cross section, 3-6 mm in diameter, cleft c. 3/4, lobes
0.5—1 mm thick; staminal disc subcircular or obtusely trigonous, flat, 1.5—-3 mm diam-
eter; anthers 19-25 (i.e., 40-50 closely appressed thecae), entirely sessile, suberect to

296 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 43. Knema linguiformis (J. Sinclair) W. J. de Wilde. a. Branchlet with male inflorescences;
b. male bud; c. opened male bud showing androecium; d. androecium seen from beneath; e. an-
droecium (staminal disc) seen from above (thecae of anthers + schematic); f. sample of hairs from
male bud; g. twig with female inflorescences; h. opened female bud showing the pistil; 1. in-
fructescence with submature and immature fruit [a—f: S 26978; g, h: S 32290, i: SAN 81118]. —
Scale bar for a, g, i= 2 cm; for b, c, h = 2.8 mm; for d, e = 0.66 mm; for f = 0.16 mm.

De Wilde — Myristicaceae (Knema) 297

oblique, hidden under the staminal disc, 0.2-0.3 mm; androphore tapering, 0.5—2 mm
long. Female flowers: pedicel 5-8 mm long, bracteole about median; buds depressed
globose, ellipsoid, or obovoid, 4-6 by 4—5.5 mm, cleft 1/2—2/3, lobes 0.7—1 mm thick;
ovary (broadly) ovoid, 2-3 by 1.5—2.5 mm; stigma subsessile, + flat, subcircular, faintly
2-lobed and each lobe again 4—10(—20)-lobulate. Fruits 1-3 per infructescence, ellip-
soid to ovoid, at base often contracted to 5 mm long, 3.5—4 by 2—2.5 cm, at first with
minute indumentum, glabrescent; dry pericarp 2 mm thick; fruiting pedicel 5-8 mm
long. — Fig. 43.

Field-notes — Trunk sometimes with low-footed buttresses; bark (almost) smooth, or
slightly scaly; inner bark soft, 5 mm thick, reddish or brown; sapwood dirty whitish.
Leaves medium green, glossy above, dull glaucous with yellowish green midrib be-
neath. Flowers with yellowish to rusty indumentum, pink inside; pollen pale yellow or
whitish. Fruits yellowish to deep orange.

Distribution — Malesia: Borneo (C, S and E Kalimantan, Sarawak, Sabah, Brunei).

Habitat & Ecology — Lowland and submontane primary and degraded forest; hill-
sides and ridges; sandy soil, yellow clayey soil, sandy loam, poor soil, and sandstone;
0-1500 m altitude, up to 1650 m in the Mt Kinabalu area; fl. Jan—Dec., mainly June—
July; fr. throughout the year.

Note — Knema linguiformis is related to K. curtisii, which differs in more ellipsoid,
herbaceous leaves, less thickly pubescent twigs, and longer stalked fruits.

31. Knema longepilosa (W. J. de Wilde) W.J. de Wilde

Knema longepilosa (W.J. de Wilde) W.J. de Wilde, Blumea 41 (1996) 389, f. 3; Tree Fl. Sabah &
Sarawak 3 (2000) 425. — Knema percoriacea J. Sinclair forma longepilosa W.J. de Wilde, Blumea
25 (1979) 394. — Type: S 22985 (Sibat), Sarawak.

Tree 3-10 m. Twigs 2-4 mm diameter, looking as 4-6 mm diameter because of the
dense cloth of yellow-brown or dark rusty + shaggy hairs 2-3 mm long, late glabrescent;
bark reddish brown, thinly and finely flaking. Leaves chartaceous or thinly coriaceous,
elliptic-oblong to lanceolate, 13-25 by 3—6 cm, base narrowly rounded or cuneate, apex
acute-acuminate; drying olivaceous or greenish above, lower surface greyish, early
glabrescent, with midrib and nerves brown-yellow, contrasting; dots absent; midrib nar-
row, pale, much raised above; lateral nerves 10-15 per side, raised; venation distinct at
both surfaces; petiole 10-15 by 1.5—2 mm, reddish brown, transversely wrinkled when
dry, at first with dense yellow-brown or rusty hairs 2-4 m, contrasting with the early
glabrescent leaf blades; leaf buds short, 5(—10) mm long, looking bluntish by a dense
cloth. Inflorescences: sessile, brachyblast simple or forked, to 5(—-10) mm long, late gla-
brescent, in male with clusters of 2-6 long-pubescent flowers. Male flowers: with dense,
rough, yellow-brown or rusty hairs (2—)4 mm, appearing as mops 6—8 mm in diameter;
perianth 3-lobed, colour inside not known; pedicel 3—4 mm, bracteole oblong, 0.5—1
mm long, long-pubescent like pedicel and perianth, at 0.5—1 mm below the apex; buds
subglobose, 3.5—4 mm diameter, cleft c. 1/2, lobes 0.5 mm thick; staminal disc flat,
blunt-subtriangular, 1.5(-2) mm diameter; anthers 15—18, lateral, contiguous, sessile,
suberect; androphore short, tapering, | mm long, glabrous. Female flowers and fruits
not known. — Fig. 44.

298 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Small tree, dbh c. 5 cm; bark (reddish) brown, latex red. Leaves glau-
cous beneath. Flowers and buds entirely covered by light brown or rusty indumentum.

Distribution — Malesia: Borneo (C Sarawak: 4th Div.; West-Central Kalimantan).

Habitat & Ecology — Hill-side forest; occasional, in mixed dipterocarp forest on
basaltic soil, clay soil; 100-250 m altitude; fl. Jan., Apr.

She

dn 95 i
Fig. 44. Knema longepilosa (W.J. de Wilde) W.J. de Wilde. a. Leafy twig with male inflores-
cences; b. lengthwise opened male flower; c. bracteole of male flower; d. androecium, seen from
below [a: Church & Mahyar 1613; b—d: Church & Mahyar 1466]. — Scale bar for a = 2 cm; for
b-—d=2 mm.

De Wilde — Myristicaceae (Knema) 299

Note — Knema longepilosa is readily recognized by the conspicuous dense and long
indumentum of the sterile apical leaf buds, twig apex, petioles, and flowers. It resembles
the Malayan K. plumulosa, which differs in the mammillate staminal disc and by a disc
around the base of the androphore.

32. Knema losirensis W. J. de Wilde

Knema losirensis W.J. de Wilde, Blumea 25 (1979) 463, f. 16; 27 (1981) 231. — Type: de Wilde &
de Wilde-Duyfjes 16578, Aceh, N Sumatra.

Tree 10-20 m. Twigs 1.5—2.5 mm diameter, at first with stellate hairs 0.1—0.2 mm
long, early glabrescent; bark striate, not cracking or flaking. Leaves membranous or
chartaceous, elliptic-oblong to oblong-lanceolate, 13—26 by 4—8 cm, apex up to 1.5 cm
acute-acuminate, base rounded to attenuate; greenish to brown above, the midrib late
glabrescent; lower surface greyish, at first with sparse to dense greyish hairs of mixed
size (0.1—0.2 mm long, and coarser stellate dendroid hairs 0.3—0.5 mm long), late
glabrescent, the longer hairs earlier shed and leaving hair scars (lens!); dots absent;
midrib and nerves (20-25 pairs) slightly raised above; venation fine (areoles 0.5 mm
diameter or less), distinct above; petiole 8-13 by 1.5—2.5 mm. Inflorescences: sessile or
pedunculate to 1 mm, brachyblast simple or 2- or 3-fid, up to 5 mm long; in male 5—10-
flowered, female 1—6-flowered, flowers with persistent yellowish brown hairs 0.1 mm
long or less; perianth 3-lobed, greenish or yellowish inside. Male flowers: pedicel (3—)
5—9 mm long, bracteole caducous, 1-3 mm below the apex; buds depressed globose to
broadly obovoid, 3—4 by (3—)3.5—5 mm, cleft c. 4/5, lobes | mm thick; staminal disc flat
to faintly convex, subcircular, 2—2.5(—3) mm diameter; anthers 10—15, + stiped, more or
less horizontal, 0.4—0.6 mm long, spaced, the connective broad above, thecae opening
downwards; androphore 1.2—1.5 mm long. Female flowers: pedicel 1.5—5 mm long,
bracteole caducous, above median; buds (depressed) obovoid-oblong, 4-6 by 2.5-5 mm,
cleft over 1/2, lobes 1—-1.5 mm thick; ovary ovoid, 3 by 2 mm, stigma + sessile, 2-lobed,
each lobe suberect, and again 2—4-lobulate. Fruits 1-5 per infructescence, broadly ellip-
soid, more or less ridged, base rounded, apex subacute, 2—2.7 by 1.5—2 cm, with dense
rusty hairs 0.1—0.2 mm long; dry pericarp 1—1.5 mm thick; fruiting pedicel 2-5 mm
long. — Fig. 45.

Field-notes — Bark rather smooth, slash orange-red, smelling like pine resin; wood
creamy. Leaves glaucous beneath. Male buds over 6 mm diameter, greenish brown or
very light brown; perianth greenish or pale yellow inside; staminal disc distinctly con-
vex, pale yellow or whitish; anthers sessile, yellow; stigma pale yellow. Fruits orange-
brown.

Distribution — Malesia: Sumatra (Aceh, E Coast, Palembang).

Habitat & Ecology — Forest edges, forest of alluvial terraces, hill slopes; 200-600 m
altitude; fl. & fr. throughout the year.

Note — Knema losirensis is related to K. glaucescens, particularly resembling the
Sumatran specimens of the latter. These differ in a less stout habit of the leafy twig, with
smaller male flowers and smaller fruits. The comparatively large, ridged fruits of K.
losirensis are quite different from the fruits of K. glaucescens, especially as found in
Peninsular Malaysia and Borneo. The stouter habit of K. /osirensis possibly is partly due
to more favourable conditions in forest on rich alluvial soil.

300 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 45. Knema losirensis W.J. de Wilde. a. Twig with male inflorescences; b. male bud; note scar
of fallen bracteole; c. opened male bud showing the androecium; d. androecium, almost lateral
view: e. androecium seen from below; f. hair sample of indumentum of male bud, half-schematic,
g. female bud; note scar of fallen bracteole; h. opened female bud showing pistil; i. branchlet with
infructescence; j. detail of lower leaf surface with tertiary venation and scattered small (sub)sessile
stellate hairs [a—f, j: de Wilde /Duyfjes 16578; g, h: de Wilde/Duyfjes 16577; i: de Wilde/Duyfjes
12565]. — Scale bar for a, i = 2 cm; for b, c, g, h = 2.8 mm; for d, e = 1.4 mm; for f= 0.15 mm; for
j= 0.33 mm.

De Wilde — Myristicaceae (Knema) 301

33. Knema lunduensis (J. Sinclair) W.J. de Wilde

Knema lunduensis (J. Sinclair) W.J. de Wilde, Blumea 25 (1979) 395; 41 (1996) 391; Tree Fl. Sabah
& Sarawak 3 (2000) 426. — Knema latericia Elmer var. lunduensis J. Sinclair, Gard. Bull. Sing.
18 (1961) 244, f. 15A, D, E. — Type: Sinclair (& Kadim) 10381, Sarawak.

Tree 3—10(—20) m. Twigs 4-6(—10) mm diameter, at first with rusty hairs (0.5—)1—1.5
mm long, glabrescent; bark coarsely striate, lower down finely longitudinally cracking
and flaking. Leaves chartaceous to thinly coriaceous, at first with dense indumentum,
early glabrescent, elliptic to (ob)lanceolate, 15—50 by 4.5—16(—20) cm, apex (sub)obtuse
to acute(-acuminate), base obtuse, rounded, subtruncate, or rarely subcordate; brown
above, grey(-brown), glabrous, not or but little papillate, with scattered hair scars be-
neath; dots absent; midrib raised above; nerves 12-30 pairs, raised; venation coarse,
+ fairly distinct; petiole 10-30 by 3-7 mm. Inflorescences: sessile, brachyblast simple
or forked, up to 10 mm diameter; in male S—15-flowered, female 1—6-flowered; flowers
with rusty hairs (0.4-)0.5—1.5(-2) mm long; perianth 3- (or 4-)lobed, pink-red inside.
Male flowers: pedicel 3-4 mm long; bracteole + persistent, slightly above median; buds
depressed globose, small or large (see note 2), 3-3.5 by 4-5(-6) mm, cleft c. 4/5, lobes
0.5 mm thick; staminal disc flat or slightly convex, subtriangular to circular, 1.8-2 mm
diameter; anthers 10-14, just stiped, 0.6 mm, horizontal, not touching; androphore taper-
ing, 0.6—1 mm long. Female flowers not seen; stigma remnants (on fruit): sessile, many-
lobed. Fruits 1-6 per infructescence, ellipsoid, (2—)2.5—3.5(—4.5) by 1.5—2.5(—3.5) em,
with coarse hairs 1-2 mm long, dry pericarp 2-4 mm thick; fruiting pedicel up to 3 mm
long.

Field-notes — Bark of trunk flaky, fissured, or smooth; slash light red; wood white.
Leaves dark glossy green above, with whitish green midrib. Flowers pale flesh-coloured
inside. Fruits brown, red, or chocolate; aril rich yellow, or orange.

Distribution — Malesia: Borneo (Sarawak: frequently collected in the Lundu dis-
trict, Ist Div.; Sabah).

Habitat & Ecology — Primary forest; on granitic soils, granodiorite, tertiary granitic
rock, ridges; 100-1500 m altitude; also from Quercus forest at 1500 m altitude; fl. June;
fr. June—Nov.

Notes — 1. Knema lunduensis is related to K. latericia, K. furfuracea, and K. per-
coriacea and is more or less intermediate between the first two species mentioned. Knema
latericia generally has a more slender habit, with smaller leaves, often a convex or mam-
millate staminal disc, and smaller fruits with shorter indumentum; K. furfuracea has
cordate leaf bases, though the leaf base in K. Junduensis is occasionally subcordate in
larger leaves; K. percoriacea has a differing general habit, with more coriaceous leaves.
Some specimens included in K. Junduensis come close to certain stout, large-fruited
forms of K. latericia subsp. ridleyi.

2. SAN 9957, from Sabah, has extremely large male buds, 7 by 5.5 mm, with coarse
hairs 1—5(—2) mm long; anthers 13.

34. Knema luteola W. J. de Wilde

Knema luteola W.J. de Wilde, Blumea 25 (1979) 451; Tree Fl. Sabah & Sarawak 3 (2000) 426. —
Type: Endert 2822, Central E Kalimantan.

302 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 8-20 m. Twigs somewhat flattened, sometimes blunt-triangular, pale brownish,
smooth, 1.5—2.5(—3) mm diameter, at first with greyish brown scale-like hairs, 0.1 mm
long or less, early glabrescent; bark smooth or finely striate, not tending to crack or
flake. Leaves chartaceous, oblong or lanceolate, 12-30 by 3—9.5 cm, base rounded to
cuneate, apex up to 1.5 cm acute-acuminate; greenish brown above; lower surface grey-
ish, at first with minute, weak, grey or pale brown scale-like and stellate-dendroid hairs,
early glabrescent; dots absent; midrib raised above; nerves 16—25 pairs, slender, hardly
raised above; venation fine (areoles less than 0.5 mm diameter), distinct above; petiole
10-15 by 1.5-2.5 mm. /nflorescences: sessile, brachyblast simple or 2—4-fid, to 5 mm
long; in male 5—12-flowered, flowers with persistent yellowish brown stellate scale-like
hairs 0.1 mm long or less; perianth 3-lobed, (pale) yellow inside. Male flowers: pedicel
4-7.5 mm long, bracteole caducous, 1—2.5 mm below the apex; buds globose, at base
rounded, apical part rounded or somewhat narrowed, not depressed, 2.5—4 mm diam-
eter, cleft 3/4—4/5, lobes 1 mm thick; staminal disc flat, circular, 1.7—2.3 mm diameter;
anthers 9 or 10, (just) stiped, nearly horizontal, 0.5 mm long, spaced, thecae opening
+ laterally; androphore 0.5 mm long. Female flowers not known. Fruits 1—4 per infruc-
tescence, ellipsoid to ellipsoid-oblong, 3-7 by 2-2.3 cm, usually beaked at apex, with
minute scale-like hairs less than 0.1 mm long; dry pericarp 2 mm thick; fruiting pedicel
stout, 15—20 mm long, bracteole scar + at or below median.

Field-notes — Bole straight; bark brownish, rather fissured, or strongly peeling off;
inner bark 3 mm thick, reddish or yellow inside; sapwood pale yellowish.

Distribution — Malesia: Borneo (Sarawak, W, C and E Kalimantan).

Habitat & Ecology — Hill forest, often on stream banks; 100—700 m altitude; fl. June,
Aug., Dec.; fr. Feb.—Mar.

Note — Knema luteola is one of the few species of Knema, besides K. rubens (Suma-
tra, Peninsular Malaysia) and K. oblongifolia (Peninsular Malaysia), with the twigs in
the apical part + flattened. Knema rubens differs in its longer and thicker indumentum
of twig apices, reddish in colour, hairs remaining on the lower side of the midrib, and
coarser venation of the leaves, with areoles more than 0.5 mm diameter; K. oblongifolia
has differently shaped male buds. Stout sterile or fruiting specimens may resemble
K. ashtonii var. ashtonii, which obviously is related, but differs in its larger fruits. The
fruits of K. luteola resemble those of K. kostermansiana, which are generally larger
(longer) and fusiform.

35. Knema malayana Warb.

Knema malayana Warb., Mon. Myrist. (1897) 570, t. 25; Gamble, Mat. Fl. Malay Penins. 5, 23
(1912) 242: Ridl., Fl. Malay Penins. 3 (1924) 69; J. Sinclair, Gard. Bull. Sing. 16 (1958) 293; 18
(1961) 260: W.J. de Wilde, Blumea 25 (1979) 456. — Myristica malayana (Warb.) Boerl., Handl.
FI. Ned. Ind. 3, 1 (1900) 91, nom. inval. — Syntypes: Wray 176; King’s coll. 5706, 6128, 10594;
Cantley 20; Griffith 5706; all Peninsular Malaysia.

Myristica glaucescens auct. non (Jack) Hook. f. & Thomson: Hook. f. & Thomson, Fl. Ind. 1 (1855)
157, p.p.: Hook. f., Fl. Brit. India 5 (1886) 111, p.p.; King, Ann. Roy. Bot. Gard. Calc. 3 (1891)
823; plloleip:p:

Myristica corticosa auct. non (Lour.) Hook. f. & Thomson: Hook. f. & Thomson, FI. Ind. 1 (1855)
158, p.p.; A.DC., Prod. 14, 1 (1856) 205, p.p.; Miq., Fl. Ind. Bat. 1, 2 (1858) 69, p.p.

De Wilde — Myristicaceae (Knema) 303

Tree 3-20 m. Twigs 1-2 mm diameter, at first with rusty stellate scale-like hairs 0.1
mm long or less, glabrescent; bark finely striate, not tending to crack or flake. Leaves
membranous, elliptic to oblong-lanceolate, 7-21 by 2—7 cm, base cuneate-attenuate to
rounded, apex up to 2.5 cm acute-acuminate; greenish brown above, grey-glaucous be-
neath, at first with sparse, greyish, scale-like hairs, 0.1 mm long or less, glabrescent;
dots absent: midrib raised above; nerves 14—20 pairs, raised above; venation very fine,
prominent and distinct above; petiole 7-15 by 1.5-2.5 mm. Inflorescences: sessile,
brachyblast simple or bifurcate, to 5 mm long; in male 3—15-flowered, female 1-6-flow-
ered; flowers with persistent grey-rusty hairs 0.1 mm long or less; perianth 3- (or 4-)
lobed, cream-coloured inside (always?). Male flowers: pedicel 7-15 mm long, bracteole
caducous, median or above; buds (depressed) globose, 3—4.5 mm diameter, cleft c. 4/5,
lobes 0.6—1.2 mm thick; the staminal disc flat, circular or subtriangular, 2—2.3 mm in
diameter: anthers 9-13, shortly stiped, horizontal, 0.3—0.5 mm long, spaced; androphore
0.5-1 mm long. Female flowers: pedicel 6-10 mm long, bracteole (somewhat above)
median; buds 4—5 by 2.5—3.5 mm, cleft 1/2—2/3; lobes 1—-1.5 mm thick; ovary ovoid,
1.5-2 mm long; style to 0.5 mm long; stigma erect, irregularly peltate, 5-lobed. Fruits
1-3 per infructescence, ellipsoid or obovoid, apex subobtuse, base often narrowed, |.2—
1.6 by 0.8—1 cm, with mealy hairs 0.2—0.4 mm long; dry pericarp 1—-1.5 mm thick; fruit-
ing pedicel (4—)6—15 mm long.

Field-notes — Bark not furrowed, not or but slightly flaky, nearly smooth; inner bark
pinkish, orange, or brownish, granular; wood white. Leaves slightly coriaceous, when
dry membranous. Flowers light yellow; stigma green. Ripe fruits light or dark brown, or
yellow.

Distribution — Peninsular Thailand; in Malesia: Peninsular Malaysia (most prov-
inces), Singapore.

Habitat & Ecology — Lowland and hill forest, old bamboo forest, ridges; 0-800 m
altitude; fl. throughout the year; fr. mainly May—Aug.

Note — Not in Sumatra, as was stated by Sinclair.

36. Knema mamillata W.J. de Wilde

Knema mamillata W.J. de Wilde, Blumea 25 (1979) 424, f. 12. — Type: Anderson 7 (1975), S Kali-
mantan.

Tree 5-10 m. Twigs sometimes bluntly 2—3-angled, 1.5—2.5 mm diameter, at first with
woolly soft dendroid hairs | mm long, early glabrescent; bark smooth, or finely striate,
not tending to crack or flake. Leaves chartaceous or thinly coriaceous, oblong(-lanceo-
late), (7-)11-25 by (2-)3—6.5 cm, apex acute(-acuminate), base attenuate to nearly
rounded; olivaceous to dark brown above, lower surface + reddish brown, at first with
dense hairs 0.5—1 mm long, early glabrescent; dots absent; midrib much raised above;
nerves | 1-16 pairs, raised above; venation coarse, raised and visible above; petiole 7-18
by 1.5-2.5 mm. Inflorescences: sessile, brachyblast simple or 2- or 3-fid, to 5 mm long;
in male (2-)5—10-flowered; flowers at first with dense woolly dendroid hairs 0.6—1 mm
long, early glabrescent; perianth 3- (or 4-)lobed, reddish (?) inside. Male flowers: pedi-
cel 5-10 mm long, bracteole caducous or persistent, median or below; buds (depressed)
globose, obtusely 3- (or 4-)angular in cross section, 2-3.5 by 2.5—4.5 mm, cleft 4/5,
lobes 0.30.5 mm thick; staminal disc circular, 1.5—2 mm diameter, with very distinct

Flora Malesiana, Ser. I, Vol. 14 (2000)

304

De Wilde — Myristicaceae (Knema) 305

mammilla 0.5—1 mm long; anthers 9-12, just stiped, 0.5—0.7 mm long, not touching,
horizontal, thecae opening downwards; androphore 0.5—1 mm long. Female flowers and
fruits not seen. — Fig. 46.

Distribution — Malesia: Borneo (S Kalimantan).

Habitat & Ecology — Agathis forest on sand ridges in peat forest; riparian along a
peaty river; 0-50 m altitude; fl. Mar., Aug.

Note — Knema mamillata is related to K. membranifolia, K. intermedia, and K. uligi-
nosa, especially to the latter. Knema uliginosa differs in flowers with persistent much
shorter indumentum, and staminal disc convex or only low-mammillate; in K. mamillata
the mammilla is conspicuous, up to | mm long, nearly as long as the androphore.

37. Knema mandaharan (Miq.) Warb.

Knema mandaharan (Miq.) Warb., Mon. Myrist. (1897) 553, t. 24; J. Sinclair, Gard. Bull. Sing. 16
(1958) 319, f. 16; 18 (1961) 261; W.J. de Wilde, Blumea 25 (1979) 404. — Myristica mandaharan
Mig., Fl. Ind. Bat., Suppl. 1 (1861) 384; Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 48. — Type: Diepen-
horst in Teijsmann 3091, Sumatra.

Tree 10-25 m. Twigs (4-)5—8 mm diameter, at first with rusty felty hairs (0.5—)1—1.5
mm long, + easily rubbed off, glabrescent; bark striate or not, not or only occasionally
tending to crack or flake. Leaves coriaceous, oblong to lanceolate, (15—)25—-55 by (2.5—)
5-14 cm, base rounded to cordate, or sometimes attenuate, apex (long-)acute; glossy or
bright brown above, lower surface glabrous, grey(-brown), or glaucous, not or but little
papillate; dots absent; midrib stout, much raised above; nerves 14—40 pairs, distinct,
little or much raised above; venation fine, faint or distinct; petiole late glabrescent, 15—
20 by 5-8 mm. Inflorescences: sessile, brachyblast warted, 5—15 mm diameter; in male
2-10-flowered, female 1—4(—8)-flowered; flowers with rusty persistent hairs 0.5—1 mm
long; perianth (3—)4(—5)-lobed, red (?) inside. Male flowers: pedicel 3-6 mm long, brac-
teole caducous, (sub)apical; buds obovoid, (3.5—)4—-6 by (3.5—)4—-5.5 mm, cleft 2/3
(—3/4), lobes 0.5—0.7 mm thick; the staminal disc circular, flat or slightly convex, or
concave, 1.5—2.5 mm diameter; anthers 10-17, half-sessile to just stalked, 0.5—0.7 mm,
oblique to nearly horizontal, not or but little touching; androphore tapering, 1—1.5 mm
long. Female flowers: pedicel 4-7 mm long, bracteole submedian to apical; buds ellip-
soid-obovoid, 8—10 by 5—6.5 mm, cleft c. 1/2, lobes 1—1.5 mm thick; ovary ovoid, 4mm
long; style to 1 mm long; stigma + flat, 2-lobed and each lobe again with 8-10 linear
lobelets. Fruits 1-2(—4) per inflorescence, ovoid or ellipsoid, 3—6 by 2.5—3.8 cm, with
dark brown hairs 1-2 mm long; dry pericarp 2—8 mm thick; fruiting pedicel 4-8 mm
long.

Fig. 46. Knema mamillata W.J. de Wilde. a. Twig with male inflorescences; b. male bud; note gla-
brescence and scar of bracteole about halfway the pedicel; c. opened male bud showing androecium;
d. androecium, seen almost laterally; note strongly mammillate staminal disc; e. twig with male in-
florescences; note the much smaller size of the buds as compared to a; f. male bud; note glabrescence
and bracteole towards the base of the pedicel; g. opened male bud showing mammillate androe-
cium [a—d: Anderson 7 (1975); e—g: Buwalda 7658]. — Scale bar for a, e = 2 cm; for b = 2.8 mm;
for c, f, g = 1.4 mm; for d = 0.7 mm.

306 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Bark smooth, fissured, cracked, or flaky; inner bark pinkish brown;
wood white.

Distribution — Malesia: Sumatra (Aceh, Tapanuli, W Coast, E Coast), Peninsular
Malaysia (Perak, Trengganu, Selangor, Johore).

Habitat & Ecology — Primary forest on hill sides, in Peninsular Malaysia also in
marshy forest at lower altitudes; 100—1800 m altitude; fl. Mar.; fr. throughout the year.

Notes — 1. Knema mandaharan is variable in the distinctness of the venation of the
upper leaf surface, and size of the fruits and the thickness of the pericarp. In the material
from N Central Sumatra possibly two forms can be distinguished, e.g., stout plants, with
thick twigs and large leaves with faint venation above, the fruits large, 4-6 cm long, with
short-haired pericarp 4-8 mm thick; the other form in N Sumatra has smaller leaves with
distinct venation above, with looser, more farinose hairs on immature leaves, smaller
fruits, 3-4 cm long, with longer, more lanose indumentum, especially at the base of the
fruits, and with the pericarp only 2-3 mm thick. In other material from Sumatra and
Peninsular Malaysia these differences are not apparent, as most of these specimens have
a distinct venation.

2. The aril in de Wilde 13769 is incised to nearly halfway, which is unusual in Knema.

38. Knema matanensis W. J. de Wilde
Knema matanensis W.J. de Wilde, Blumea 27 (1981) 228, f. 2. — Type: de Vogel 6060, Sulawesi.

Tree 6-25 m. Twigs sometimes faintly angled, 3-5 mm diameter, with hairs 0.1 mm
long or less, early glabrescent; bark smooth, not tending to crack. Leaves coriaceous,
oblong(-lanceolate), 14-31 by 4.5—10.5 cm, base attenuate, apex up to 1.5 cm acute-
acuminate: olivaceous to brown above; the lower surface at first with sparse stellate
(-dendroid) hairs of mixed size, 0.1 mm long or less, subglabrescent; dots absent; midrib
broad, + flat above; nerves 18-26 pairs, little raised above; venation fine, distinct above;
petiole 7-18 by 2-5 mm. Inflorescences: sessile, brachyblasts to 20 mm long, single, 2,
or 3 together, in male and female 2—5-flowered; flowers with dark rusty hairs 0.1—0.2
mm long; perianth 3-lobed, creamy to yellow inside. Male flowers: pedicel 2—3 mm,
bracteole apical, 0.5 mm long, persistent; buds (sub) globose, 2.5(—3) by 2.5—3 mm, cleft
c. 4/5, lobes 1.2 mm thick; staminal disc subtriangular, + flat, 1.5 mm diameter; anthers
7 or 8, stiped, horizontal, 0.2—-0.3 mm long, opening + downwards; androphore 0.5—0.7
mm long. Female flowers: pedicel 1 mm long; buds ovoid-oblong, 4.5 by 2.5 mm, cleft
c. 1/2, lobes 1 mm thick; ovary ovoid, 2 by 2 mm, with dense hairs 0.2 mm long; style
less than 0.5 mm, stigma 0.5 mm long, of 2 broad lobes, each 3- or 4-lobulate. Fruits
1 or 2 per infructescence, (immature) ellipsoid, 2 by 1.2 cm, with dense dark rusty den-
droid hairs 0.2—0.3(—0.5) mm; thickness of mature dry pericarp not known; fruiting pedicel
rather stout, 5 mm long. — Fig. 47.

Fig. 47. Knema matanensis W.J. de Wilde. a. Male flowering twig; b. opened male bud, showing
the androecium: c. androecium seen from beneath; d. opened female bud, showing pistil; e. twig
with submature fruit: f. detail of lower leaf surface showing scattered minute stellate hairs [a—c, f:
de Vogel 6109; d, e: de Vogel 6128]. — Scale bar for a = 2 cm; for b, d= 1.65 mm; for c = 0.4 mm;
for e = 1 cm; for f = 0.8 mm.

307

Myristicaceae (Knema)

De Wilde —

at

a eee TTT ART A
“ we Pits.

308 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 48. Knema membranifolia H.J.P. Winkler. a. Twig with leaves and infructescence; b. older
branchlet with male inflorescences in the axils of fallen leaves; c. male bud; note glabrescent peri-
anth; d. opened male bud showing androecium laterally; e. androecium seen from below, anthers
somewhat curved downwards; f. opened (sub)mature female bud, reconstructed from immature
flower and stigma persistent on fruit [a: S 19225; b—e: S 23277, f: Kostermans 9536}. — Scale bar
for a, b = 2 cm; for c, d = 2.8 mm; for e = 1.4 mm; for f = 2 mm.

De Wilde — Myristicaceae (Knema) 309

Field-notes — In both sexes the perianth is outside (ochreous) brown, yellow or cream
inside, in male with a bright dark red androphore.

Distribution — Malesia: Central Sulawesi (northern SE Peninsula, N side of Lake
Matano).

Habitat & Ecology — Forest on thin soil cover over conglomeratic rock; also on al-
luvial flatland; c. 600 m altitude.

Notes — 1. Knema matanensis seems related to species in the group of K. glauca
(with, e.g., K. cinerea, K. glauca, and K. kinabaluensis); it seems particularly related to
K. kostermansiana, a species also with small globose male buds, and the bracteole apically
on the pedicel, but differing in membranous leaves with cordate base, 8-10 anthers, and
caducous bracteole.

2. Knema matanensis (and K. stellata subsp. minahassae) was not seen in the area
with ultrabasic rock south of Lake Matano, where K. celebica exclusively was found.

39. Knema membranifolia H.J.P. Winkler

Knema membranifolia H.J.P. Winkler, Bot. Jahrb. Syst. 49 (1913) 368; J. Sinclair, Gard. Bull. Sing.
18 (1961) 262, f. 18; W.J. de Wilde, Blumea 25 (1979) 422, f. 11; Tree Fl. Sabah & Sarawak 3
(2000) 427. — Lectoype: Hubert Winkler 2460, SE Kalimantan.

Tree 6-25 m. Twigs sometimes bluntly 2- or 3-angular, 1.5-3 mm diameter, at first
with dense rusty hairs 1 mm long, glabrescent; bark pale brown to yellowish, striate,
when old tending to crack or flake. Leaves membranous or thinly coriaceous, oblong to
lanceolate, 12-28 by 3.5-8.5 cm, base cuneate-attenuate to subobtuse, apex obtuse to
acute-acuminate; drying pale brown or olivaceous above; lower surface early glabrescent,
greenish grey; dots absent; midrib stout, prominent above; nerves 12-17 pairs, hardly
raised above; venation lax or fine, distinct; petiole 15—25 by 1.5—3 mm. Inflorescences:
sessile, brachyblast simple, wart-like, or up to 2—4-tubercled, to 10 mm long; in male
5—30-flowered, female 1—4-flowered; flowers at first with dense hairs 0.5—1 mm long,
early glabrescent, or with only a few scattered hairs left; perianth 3-lobed, red (?) inside.
Male flowers: pedicel 6-9 mm long, bracteole caducous, about median; buds depressed
globose, obtusely trigonous in cross section, 3.5—5 by 5—7(—8) mm, cleft 3/4—4/5, lobes
0.7—1 mm thick; staminal disc flat to convex, not mammillate, circular, 3—-3.5 mm diam-
eter; anthers 19 or 20(—23?), + stiped, horizontal, 0.7—1 mm, not touching; androphore
tapering, 0.7—1 mm long. Female flowers: pedicel 1-4 mm long, bracteole above the
middle; buds subglobose or broadly obovoid, 3.5—4.5 mm diameter, cleft c. 3/4; ovary
subglobose to ovoid, 1.5—2 mm long: stigma + sessile, with 2 main lobes and each lobe
again deeply (2—)4—7-lobulate. Fruits 1-3 per infructescence, ellipsoid or obovoid, 2.5—
4.5 by 2-3.5 cm, at first with rusty indumentum, hairs c. 1 mm long, early glabrescent;
dry pericarp glossy, 1.5—2 mm thick; fruiting pedicel 1-5 mm long. — Fig. 34e, 48.

Field-notes — Bark thin, smooth, peeling off in thin strips, or flaky; living bark 3-5
mm, brown; wood pale brown or white; exudate of bark (yellowish) red. Flowers brown
outside. Fruits with easily detachable brown cover, becoming reddish or yellow.

Distribution — Malesia: Borneo (Sarawak, Sabah, E & SE Kalimantan).

Habitat & Ecology — Lowland dipterocarp forest, on loam, sandstone, basalt, and
lime-containing soils; 0-500 m altitude; fl. mainly Apr.—June; fr. throughout the year.

310 Flora Malesiana, Ser. I, Vol. 14 (2000)

Note — Knema membranifolia resembles typical K. curtisii by the glabrescent yel-
lowish twigs; that species, however, has terete twigs, non-flaking bark, and a different
androecium. The closest alliance of K. membranifolia is possibly with K. korthalsii,
which differs in non-flaking bark of twigs, generally larger (longer) leaves with more
veins, flowers with persistent indumentum, 12—20 smaller anthers, and smaller fruits.

40. Knema minima W. J. de Wilde

Knema minima W.J. de Wilde, Blumea 43 (1998) 249, f. lc. — Type: Dransfield c.s. JD 7071,
Brunei.

Treelet, 2-5 m. Twigs bluntly angular, 1-1.5 mm diameter, with dense rusty hairs 0.5
mm, glabrescent; bark (yellow-)brown, finely striate, not or faintly finely cracking. Leaves
chartaceous, elliptic-oblong or lanceolate, 6-18 by 1—3(—4) cm, base cuneate, rarely
nearly rounded, apex acute, early glabrescent, at first with dense woolly yellow-brown
hairs 0.5 mm; drying dull greenish above, lower surface grey-green; dots absent; midrib
raised above, nerves 10-20 per side, slightly raised, beneath yellow-brown, contrasting;
venation fine, not prominent; petiole 5-10 by 1(—1.5) mm. /nflorescences: among the
leaves, sessile, brachyblast simple (or 2- or 3-furcate), wart- or worm-like, 1-5 mm long;
in male 1—3-flowered, female 1- or 2-flowered; flowers with dense hairs, 0.5 mm; peri-
anth 3-lobed, red inside. Male flowers: pedicel 1.5(—2) by 0.5 mm, bracteole persistent,
at 1/3 below apex; buds globose, 1.5—1.8 by 2-2.2 mm, cleft c. 4/5, lobes 0.3 mm thick;
staminal disc flat, subcircular, 1—-1.5 mm diameter, anthers 7-10, shortly stiped, 0.3—0.5
mm, opening outward; androphore slender, 0.5—0.8 mm. Female flower (from remnants
under immature fruits): subsessile, 1.5—3 mm long. Fruits 1 (or 2) per infructescence,
obovoid-ellipsoid, 1.2—2 by 0.8—-1 cm, with dense bright (yellow) rusty hairs 0.5—1 mm;
dry pericarp thin, 1 mm or less; fruiting pedicel 1-1.5 mm long. — Fig. 41e.

Field-notes — Small tree, 2—5 m; branches plagiotropic, young growth rusty brown;
bark rough. Leaves dull green above, paler, glaucous beneath, midrib brownish. Inflo-
rescences racemose, axillary, few-flowered. Flowers deep red. Fruits solitary, axillary,
rusty copper-brown.

Distribution — Malesia: Borneo (S Brunei, known from a restricted area).

Habitat & Ecology — Mixed dipterocarp forest on ridges, possibly restricted to forest
on sandstone or Setap shales; loamy or podzolic soil; 50-350 m altitude; fl. Jan., Feb.,
June, July; fr. Jan., Mar., July.

Notes — 1. Knema minima resembles and may be confused with K. latericia subsp.
latericia var. subtilis or tiny forms of K. latericia subsp. ridleyi (see De Wilde, Blumea
41, 1996, 389), or with plants provisionally identified as K. tridactyla aff. subsp. sublaevis.
With K. tridactyla it has the small male flowers in common, but it differs e.g. in leaf
texture and subpersistent indumentum on the lower surface of the leaves.

2. Knema curtisii and K. minima rank among the species of the lowest stature; also
the male perianth is small, perhaps the smallest known beside those of K. curtisii, K.
mogeana, K. muscosa, and K. tridactyla.

41. Knema mogeana W.J. de Wilde

Knema mogeana W.J. de Wilde, Blumea 32 (1987) 136, f. 4; Tree Fl. Sabah & Sarawak 3 (2000)
428. — Type: Mogea (& de Wilde) 4446, C Kalimantan.

De Wilde — Myristicaceae (Knema) 311

Tree 4-15 m. Twigs 1—1.5(—2) mm diameter, with dull brown hairs 0.1—0.2 mm long,
(late) glabrescent; bark finely striate, not flaking. Leaves membranous, oblong to lance-
olate, 8-17(—24) by 1.5—4(-7) cm, base narrowly rounded to attenuate, apex acute-
acuminate, drying olivaceous or dark brown above, lower surface with persistent, sparse,
not touching, pale yellowish stellate scale-like hairs of mixed sizes, 0.1—0.2 mm; dots
absent: midrib raised above; nerves 13-20 pairs, + flat above, venation moderately fine,
not very distinct; petiole 8—12(—15) by 1—2(-3) mm. Inflorescences: sessile, brachyblast
simple or 2- or 3-forked, to 4 mm long, in male 4—8-flowered, female 2—6-flowered:
flowers with dense rusty hairs 0.1—0.2 mm long; perianth 3-lobed, greenish creamy in-
side. Male flowers: pedicel 2-4 mm long, bracteole persistent, broadly ovate, 0.5—1 mm
long, 0.5 mm below the apex; buds broadly obovoid, (2—)2.5 mm diameter, cleft 4/5—
5/6, lobes 0.5(—0.8) mm thick; staminal disc circular, flat, |-1.5 mm diameter, anthers
6—-8(—10), short-stiped, horizontal, 0.3 mm long, not touching, opening downwards;
androphore 1(—1.4) mm long. Female flowers: pedicel 3-6(—8) mm long, bracteole per-
sistent, about median or up to 2 mm below the apex; buds long-obovoid, 4 by 2.5 mm,
cleft c. 1/2, lobes 0.5 mm thick; ovary ovoid, 1.5 by 1(—1.3) mm, with hairs 0.1 mm long
or less, stigma subsessile, 1 mm long, 2-lobed and each lobe 4- or 5-lobulate. Fruits
1-6 per infructescence, ellipsoid or ellipsoid-oblong, 1.5—2 by 1—-1.2 cm, apex rounded
or short-acute, with hairs 0.1 mm long; dry pericarp | mm thick, fruiting pedicel 4-10
mm long. — Fig. 49.

Field-notes — Small tree, bark smooth, dark brown, rough when old, with some pus-
tules, not furrowed or striate; sap copious, red. Leaves glaucous beneath, with yellowish
green midrib. Perianth outside yellow-brown, greenish creamy inside, androphore whit-
ish, anthers pale yellow; fruits yellowish green.

Distribution — Malesia: Borneo (Sarawak, Sabah, C & E Kalimantan: E Kutei).

Habitat & Ecology — Primary forest, hill-slope forest, mixed dipterocarp forest, ridge
forest in kerangas; on sandy soil, sand loam soil, diorite, sandstone, limestone; 0-300 m
altitude; fl. Dec., Jan., May, June; fr. Jan., June—Sept.

Note — Knema mogeana may be confused with K. hirtella, K. stenophylla, and K.
subhirtella. Knema stenophylla has more equal-sized minute hairs on the lower leaf
surface, and larger male buds; K. hirtella and K. subhirtella both differ in stouter habit,
leaves more prominently reticulate above, androphore often minutely pubescent at base,
more anthers, 8—13(—15), and larger fruits. In K. subhirtella the bracteoles are late
caducous and the perianth is cleft less deeply.

42. Knema muscosa J. Sinclair

Knema muscosa J. Sinclair, Gard. Bull. Sing. 18 (1961) 264, f. 19; W.J. de Wilde, Blumea 25 (1979)
466; Tree Fl. Sabah & Sarawak 3 (2000) 428. — Type: Clemens 22120, Sarawak.

Tree c. 8 m. Twigs 1—-1.5 mm diameter, at first with dense rusty hairs of variable sizes,
0.2-0.7 mm long, glabrescent; bark smooth or very finely striate, not tending to crack or
flake. Leaves chartaceous, oblong(-lanceolate), 5.5-12(—15) by 1.5—3.5 cm, base (sub)
attenuate, apex acute or up to 2 cm acute-acuminate; drying brown above; lower surface
brownish grey, at first with conspicuous dense rusty mixed sessile and dendroid hairs
0.2-0.7 mm long, early or late glabrescent; dots present (lens!); midrib slender, raised
above; nerves 7-14 pairs, slender, raised; venation fine, visible above; petiole 7-10 by

Vol. 14 (2000)

Flora Malesiana, Ser. I,

AN

ee ee ee a

De Wilde — Myristicaceae (Knema) S13

1.5 mm. Inflorescences: peduncle up to 2 mm long, brachyblast simple or 2- or 3-forked,
to 3 mm long; in male (1- or) 2—8-flowered; flowers with persistent rusty woolly hairs
0.2—0.5 mm long; perianth 3-lobed, colour inside not known. Male flowers: pedicel 2—3
mm long, bracteole + persistent, + median; buds subglobose or broadly obovoid, 2—2.5
mm diameter, cleft c. 3/4, lobes 0.5 mm thick; staminal disc flat, circular, 1—-1.3 mm
diameter; anthers 6—8, + stiped, + horizontal to oblique, 0.3(—0.5) mm long, spaced;
androphore 0.7—1 mm long. Female flowers not known. Fruits solitary, obovoid-oblong,
apex shortly acute-mucronate, 1.7—1.8 by 1 cm, with hairs 0.3 mm long; dry pericarp
0.5 mm thick; fruiting pedicel 6-7 mm long, with the scar of the bracteole + median.
Distribution — Malesia: Borneo (Sarawak: Mt Majau, Gat, upper Reyang River area).
Habitat & Ecology — Montane moss forest of Mt Majau; exact altitude not known.
Note — Knema muscosa superficially resembles slenderly built specimens of K. kina-
baluensis, or also K. tomentella, K. malayana, and especially K. stenophylla, but from
all these it differs by the blackish dots on the lower leaf surface, and the leaves at first
with conspicuous dense hairs of variable sizes below, glabrescent. Knema kinabaluen-
sis has much larger fruits; K. tomentella has more striate twigs; K. malayana has larger
flowers; K. stenophylla has a much less distinct venation on the upper leaf surface.

43. Knema oblongata Merr.

Knema oblongata Merr., J. Straits Branch Roy. Asiat. Soc. no 85 (1922) 190; W.J. de Wilde, Blumea
25 (1979) 400, f. 5; Tree Fl. Sabah & Sarawak 3 (2000) 429. — Type: Ramos 1663, Sabah.

Shrub or tree 4—20 m. Twigs sometimes faintly angular, 2-5(—8) mm diameter, at first
with rusty hairs 0.2—0.6 mm long, glabrescent; bark (finely) striate, not cracking or flak-
ing. Leaves chartaceous or coriaceous, elliptic to (oblong-)lanceolate, 11-45 by 2.5-14
cm, base subattenuate, rounded, or shallowly cordate, apex acute(-acuminate):; drying
greenish brown, + glossy above, with scattered hair scars (lens!), lower surface greyish,
finely papillate, with yellow-brown to rusty hairs of variable size, persistent or partially
late glabrescent; dots absent; midrib raised above; nerves (10—)18—26 pairs, sunken or
raised above; venation fine, distinct; petiole late glabrescent, 10-25 by 2-6 mm. Jnflo-
rescences: peduncle up to 4 mm (see the subspecies), brachyblast simple or forked, to 5
mm long, in male 5—25-flowered, female |—5-flowered; flowers with rusty hairs 0.1—1(—
2) mm long; perianth 3- (or 4-)lobed, yellow (?) or red inside. Male flowers: pedicel 2—
8 mm long, bracteole caducous, median to subapical; buds depressed broadly obovoid,
tapering in the lower half, grooved when dry, 2.5—5 by 2.5-5 mm, cleft 1/2—2/3(—3/4,
see note under subsp. oblongata), lobes 0.5—1 mm thick; staminal disc circular or faintly
triangular, flat or slightly concave, 1.5—-3 mm diameter; anthers 7—9 (subsp. parviflora
and pedunculata), or 13—18(—21), + half-sessile to just stiped, + oblique or suberect,

Fig. 49. Knema mogeana W.J. de Wilde. a. Fruiting twig; b. opened fruit showing seed with aril
laciniate at apex; c. lower leaf surface with scattered small hairs; d. twig with female inflores-
cences; e. female flower; f. ditto, opened, showing pistil; g. twig with male inflorescences; h. male
flower; i. ditto, longitudinally opened, showing androecium with 7 stalked anthers [a—c: SAN 31172,
d—f: Veldkamp 8135; g—i: S 21495 (Ashton)|. — Scale bar for a, d, g = 2 cm; for b = 6.6 mm; for
C1) Anita tOmet nest — lPOsmmm:

314 Flora Malesiana, Ser. I, Vol. 14 (2000)

(0.3-)0.5—0.7 mm, not or but slightly touching; androphore tapering or not, 1-2 mm
long. Female flowers: pedicel 2.5—10 mm long, bracteole at or above the middle; buds
ellipsoid to obovoid, 6-7.5 by 4-6 mm, cleft c. 1/2, lobes 0.5—1 mm thick; ovary sub-
globose, 3-3.5 mm diameter; style |—1.5 mm; stigma 2-lobed and each lobe again 3—5-
lobulate. Fruits 1-3 per infructescence, subovoid or obovoid, apex subacute to obtuse,
base attenuate to rounded, 2—3.8 by 1.5—2 cm, with rusty hairs 1—-1.5 mm long, easily
rubbed off; dry pericarp 2-4 mm thick; fruiting pedicel 3-10 mm long.

Field-notes — Bark of trunk smooth, sometimes scaly or + fissured, once lenticellate;
inner bark red-brown, beef-red, or pinkish; slash wood pale yellowish, pale brownish, or
white. Perianth lemon-yellow to red inside. Fruits yellow or orange, reddish hairy.

Distribution — Malesia: Borneo (Sarawak, rare; Sabah; E Kalimantan).

Habitat & Ecology — Primary and degraded forest, mixed dipterocarp forest, marshy
or hill-side forest; on poor, brown, or sticky soil, and sandy ridges; 0-1200 m altitude;
fl. & fr. throughout the year.

Note — Knema oblonga is a variable species, with three subspecies. It may be con-
fused with K. tridactyla and K. laurina (the type variety), with + uniform dendroid hairs
on the lower leaf surface, absence of crateriform hair scars on the upper leaf surface, and
elongate male buds.

KEY TO THE SUBSPECIES

la. Twigs at apex with hairs 0.2(—0.3) mm long. Lateral nerves c. 10 pairs. Male buds
2.5 mm diameter, with hairs 0.1(—0.2) mm long. [Inflorescences sessile; male pedi-

cel 2 mm long; anthers 8, short-stiped.] ..............--- b. subsp. parviflora
b. Twigs at apex with hairs 0.3—0.6 mm long. Lateral nerves 18—26 pairs. Male buds
3.5—5 mm diameter, with hairs 0.4-1(-2) mm long .........-..--++++++-5- 2

2a. Leaves narrow, oblong or lanceolate, 25.5 cm wide, base attenuate to obtuse. Inflo-
rescences 1-4 mm pedunculate. Male buds 4 mm diameter; pedicel 2-3 mm long;
anthers sO. almost entirely SESSIIG jonute cy: « warepe ie 3 = 2-1 ue c. subsp. pedunculata

b. Leaves broader, elliptic to oblong-lanceolate, 4-14 cm wide, base broadly rounded
or shallowly cordate, rarely subattenuate. Inflorescences sessile or up to | mm pe-
dunculate. Male buds 3.5—-5 mm diameter; pedicel 2-8 mm long; anthers 13-21,
haliesessile TO,jUSE SUPE... 5 op. ole os estes «= ayo es oie pica a. subsp. oblongata

a. subsp. oblongata

Knema oblongata Merr. subsp. oblongata; W.J. de Wilde, Tree Fl. Sabah & Sarawak 3 (2000) 431.
— Knema obovoidea Merr., Univ. Calif. Publ. Bot. 15 (1929) 75.

Twigs stout, at apex 2.5—5(—8) mm diam., with hairs 0.3—0.6 mm long. Leaves broad,
11-45 by 4-14 cm, base broadly rounded or shallowly cordate, rarely subattenuate; lat-
eral nerves c. 20 pairs. Inflorescences: sessile or to 1 mm pedunculate; flowers with hairs
0.4-1 mm long. Male flowers: pedicel 2-8 mm long; buds broadly obovoid to obconical,
4-5 by 3.5—5 mm, cleft c. 1/2-2/3(—3/4, see note); lobes 3 (or 4), at sutures 0.5—1 mm
thick: anthers 13—18(—21), half-sessile to just stiped. Female pedicel 2.5-10 mm long.
Fruits 2-3.8 by 1.5—2 cm, apex obtuse; fruiting pedicel 3-10 mm long. — Fig. 50.

Distribution — As the species.

De Wilde — Myristicaceae (Knema) 315)

att
cnet

Hated tyiy eee
* i

a ie

|

fc

f ini

RNIN

Fig. 50. Knema oblongata Merr. subsp. oblongata. a. Branchlet with male inflorescences; b. male
inflorescence; c. male bud; d. opened male bud showing the androecium; e. androecium in half-
lateral view; f. androecium seen from below; g. opened female bud showing the pistil; h. sample of
hairs of male flower; i. detail of lower leaf surface with stellate-dendroid hairs of mixed sizes;
j. twig with infructescences, each with a single mature fruit [a, b: SAN 30740; c—f, h, i: SAN 30369;
g: Sinclair 9291; j: SAN 32857]. — Scale bar for a = 4 cm; for b = 6.6 mm; for c, d, g = 2.8 mm; for
e, f = 0.66 mm; for h = 0.16 mm; for 1 = 0.3 mm; for j = 2 cm.

316 Flora Malesiana, Ser. I, Vol. 14 (2000)

Note — Variable in the size of male buds and number of anthers. KEP FN 80161,
(Brunei) differs in its exceedingly broad male buds, 4—4.5 by 4.5—5 mm, cleft c. 3/4;
with 14—16 just stiped, rather horizontally positioned anthers.

b. subsp. parviflora W. J. de Wilde

Knema oblongata Merr. subsp. parviflora W.J. de Wilde, Blumea 25 (1979) 404; Tree Fl. Sabah &
Sarawak 3 (2000) 431. — Type: SAN 31488 (Agam), Sabah.

Twigs fairly slender, at apex 2-3 mm diameter, with hairs 0.1—0.3 mm long. Leaves
8-17 by 3-6.5 cm, base rounded; lateral nerves c. 10 pairs. Inflorescences: sessile; flow-
ers with hairs 0.1 mm long. Male flowers: pedicel 2 mm long, bracteole apical; buds
broadly obovoid, 2.5 mm diameter, cleft c. 2/3, lobes 3, at sutures 0.5 mm thick; anthers
8, just stiped, 0.3—0.4 mm long. Fruits 20 mm long, pedicel 3-4 mm long.

Distribution — Malesia: Borneo (Sabah: Lahad Datu District).

Habitat & Ecology — Primary forest on hill side; c. 150 m altitude; fl. May.

Note — Known only from the type. Its status as subspecies in K. oblongata is pro-
visional: it also seems close to K. tridactyla, a species with small flowers and only
8 anthers.

c. subsp. pedunculata W. J. de Wilde

Knema oblongata Mert. subsp. pedunculata W.J. de Wilde, Blumea 25 (1979) 403; Tree FI. Sabah
& Sarawak 3 (2000) 431. — Type: SAN 49195 (Aban Gibot), Sabah.

Twigs fairly slender, at apex 2-3.5 mm diameter, with hairs 0.5 mm long. Leaves
narrow, oblong to lanceolate, 1 1-32 by 2—5.5 cm, base attenuate to obtuse; lateral nerves
c. 20 pairs. Inflorescences: peduncle 1-4 mm long; flowers with hairs 0.4—2 mm long.
Male flowers: pedicel 2-3 mm long; buds obovoid, 3—4 by 3—4.2 mm, cleft 1/2 or slightly
over, lobes 3, variable in thickness, 0.3—1.5 mm; anthers 7—9, just sessile. Fruits (imma-
ture) 2 by 1 cm, apex acute; fruiting pedicel 6 mm long.

Field-notes — Bark chocolate; inner bark reddish; sapwood pale yellow. Flowers and
fruits reddish.

Distribution — Malesia: Borneo (W Sabah).

Habitat & Ecology — Primary forest on ridges and hill sides; yellow or brownish
soil; 0-600 m altitude; fl. Sept.

Note — This subspecies may be confused with K. tridactyla and K. laurina vat. hetero-
pilis.

44. Knema oblongifolia (King) Warb.

Knema oblongifolia (King) Warb., Mon. Myrist. (1897) 586, t. 24: J. Sinclair, Gard. Bull. Sing. 16
(1958) 321, f. 17A, B; 18 (1961) 266; W.J. de Wilde, Blumea 25 (1979) 369. — Myristica
oblongifolia King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 313, p.p., pl. 148 f. 1-7, 10, 11. — Syn-
types: King’s coll. 835, 5983 (lecto), Peninsular Malaysia.

Myristica oblongifolia King var. monticola King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 314, pl. 148,
f. 8, 9. — Knema oblongifolia (King) Warb. var. monticola (King) Warb., Mon. Myrist. (1897)
587: J. Sinclair, Gard. Bull. Sing. 16 (1958) 323, f. 17C—G; 18 (1961) 266. — Syntypes King’s
coll. 3582, 3810, 8322, 10953: Scortechini s.n.; Wray 993, 1077, 1087 (lecto); all Peninsular
Malaysia.

De Wilde — Myristicaceae (Knema) SIF

Shrub or tree (2—)5—15 m. Twigs somewhat flattened or coarsely 2- or 3-angular, 2—4
(6) mm diameter, at first with fine woolly hairs up to 1 mm long, early glabrescent; bark
smooth, finely striate, + glossy, not tending to crack or flake. Leaves chartaceous, (ellip-
tic-)oblong, 10—25(-45) by 2.5-8(—16) cm, base attenuate to rounded, rarely subcordate,
apex acute(-acuminate); drying (greenish) brown above, often with a typical pitted struc-
ture like the rind of an orange (lens!); lower surface greyish, at first with scattered grey-
ish brown stellate hairs, 0.3 mm long, persistent or glabrescent; dots absent; midrib
raised above; nerves 18-30 pairs, + flat above; venation faint; petiole 10-22 by 1.5—3
mm. Inflorescences: peduncle up to 5 mm, axillary or up to 6 mm supra-axillary, bra-
chyblast simple or forked, 2-6 mm long, in male 2—10-flowered, female 1—4-flowered;
flowers with hairs 0.2—0.5 mm long; perianth 3-lobed, reddish (?) inside. Male flowers:
pedicel (1—)2—3 mm long, bracteole persistent or caducous, almost apical; buds ellip-
soid(-oblong) to ovate, or obovate, often somewhat contracted in the middle, 3.5—5 by
2-3 mm, cleft 1/3—1/2, lobes 0.5 mm thick; staminal disc small, + circular, flat, 0.5—1
(—1.5) mm diameter; anthers 6-9, half-sessile, suberect, 0.3—0.6 mm, not tightly touch-
ing; androphore terete, slender, 1.7—3 mm long. Female flowers: pedicel 0.5—1 mm long,
bracteole (sub)apical; buds ovoid, 5—6.5 by 3.5—4.5 mm, cleft c. 1/2, lobes 0.5(-1) mm
thick; ovary subglobose, 1.7—3 mm diameter; style 1.5—2 mm long; stigma inconspicu-
ous, 2-lobed and each lobe again 2- or 3-lobulate. Fruits 1—3 per infructescence, globose
or (ob)ovoid, apex to 3 mm beaked, 2-3 by 1.5—2.3 cm, with rusty hairs 0.2—0.3 mm
long; dry pericarp 2 mm thick; fruiting pedicel 1—-3(—4) mm long.

Field-notes — Bark smooth; outer bark brittle, inside red, granular. Fruits rusty brown,
eaten by squirrels.

Distribution — Malesia: Peninsular Malaysia (Perak, Kelantan, Trengganu, Pahang,
Selangor).

Habitat & Ecology — Lowland and montane forest; 0—1700 m altitude; fl. Feb.—Mar.;
fr. Mar., July, Sept., Dec. The wide altitudinal range is unusual in Knema.

Notes — 1. Fruits are typically up to 3 mm beaked at the apex, subsessile or but
shortly stalked.

2. Especially specimens from lower altitudes have the upper leaf surface with a fine
punctate-pitted structure, under magnification reminiscent of the rind of an orange.

3. Specimens from mountain areas often have conspicuous axillary vegetative buds,
rarely found in other Knemas.

4. Knema oblongifolia is one of the few species, besides K. luteola and K. rubens,
with the younger twigs + flattened in the apical part.

45. Knema pallens W.J. de Wilde

Knema pallens W.J. de Wilde, Blumea 25 (1979) 391, f. 4; Tree Fl. Sabah & Sarawak 3 (2000) 432.
— Type: SAN 73312 (Saikeh), Sabah.

Tree 8-30 m. Twigs sometimes + angular, 5—12 mm diameter, at first with pale brown
hairs 0.2—2(—2.5) mm long, glabrescent; bark greyish brown, not striate, conspicuously
longitudinally cracking, and flaking. Leaves chartaceous or coriaceous, when immature
with dense indumentum, elliptic to lanceolate, 25-55 by (5—)6—23 cm, apex acute-acu-
minate or obtuse, base cordate, rarely rounded or subattenuate; greenish brown, + glossy
above; lower surface pale, glabrous, finely papillate; dots absent; midrib stout, raised

Flora Malesiana, Ser. I, Vol. 14 (2000)

=

.

uh

4
Duul

ise

De Wilde — Myristicaceae (Knema) 319

above; nerves 18-32 pairs, raised above; venation coarse, sunken, faint above; petiole
late glabrescent, 10-20 by 5-8 mm. /nflorescences: sessile, brachyblast simple, furcate,
or warted, 5-20 mm diameter, in male (5—)10—30-flowered, female 10—20(—40)-flow-
ered; flowers with yellowish brown hairs (0.1—)1 mm long; perianth 3- (or 4-)lobed,
reddish inside, at base thickened into a + striated disc. Male flowers: pedicel 12-20 mm
long, bracteole 2-3 mm, caducous or (sub)persistent, median to (sub)apical; buds (de-
pressed) globose or broadly obovoid, 6—10 by 6.5-10 mm, cleft 4/5 or more, lobes 1.5—
2.5(—5) mm thick; staminal disc circular, flat or slightly concave, 2.5—4.5 mm diameter;
anthers 15-23, just sessile, almost horizontal, 0.6—1 mm long, sometimes shortly apiculate,
not touching; androphore somewhat tapering, 1-1.5 mm long. Female flowers: pedicel
(7—)10-20 mm long, bracteole about median; buds obovoid, 11 by 8-9 mm, cleft c. 2/3,
lobes 2 mm thick; ovary subglobose, 3.5 mm diameter; style 1—-1.5 mm long; stigma
+ flat, 2-lobed and each lobe again 5—7-lobulate. Fruits 1-6 per infructescence, ellip-
soid, 3.5-7 by 2.5—3.5(—4.5) cm, with light brown hairs 1-2 mm long; dry pericarp 4—7
mm thick; fruiting pedicel (10—)14-30 mm long. — Fig. 51.

Field-notes — Bark scaly; inner bark light red or whitish; sapwood yellow(-ochre),
heartwood dark brown, medium hard. Leaves dark green above, glaucous beneath. Fruits
yellowish.

Distribution — Malesia: Borneo (Sarawak, Sabah, E Kalimantan).

Habitat & Ecology — Lowland forest; kerangas, forest behind mangrove, forest on
alluvial sandstone; 0—300 m altitude; fl. Jan—May, fr. throughout the year.

Note — Knema pallens is distinguishable by a stout habit, flaky bark of twigs, large
male flowers, and large, long-pedicelled female flowers; it resembles K. /amellaria from
Peninsular Malaysia, a species also with long fruiting pedicels and fruits with hairs 3-5
mm long, caducous or easily rubbed off.

46. Knema patentinervia (J. Sinclair) W.J. de Wilde

Knema patentinervia (J. Sinclair) W.J. de Wilde, Blumea 25 (1979) 456. — Knema glaucescens
Jack var. patentinervia J. Sinclair, Gard. Bull. Sing. 16 (1958) 308, f. 13C. — Knema cinerea
(Poir.) Warb. var. patentinervia J. Sinclair forma patentinervia; J. Sinclair, Gard. Bull. Sing. 18
(1961) 182, 261, 276, f. SF-H. — Type: CF 1873 (Ludin), Peninsular Malaysia.

Tree 3-16 m. Twigs 1.5—2 mm diameter, at first with minute rusty or yellowish hairs
0.1-0.2 mm long, glabrescent; bark smooth or finely striate, not tending to crack or
flake. Leaves thinly coriaceous, elliptic to lanceolate, 6-21 by 1.7—7 cm; apex obtuse,
acute, or short acute-acuminate, base attenuate to rounded; drying greenish brown,
+ glossy above; lower surface greyish, with subpersistent sparse scale-like hairs 0.1 mm
long; dots absent; midrib flat or somewhat raised above; nerves rather patent. 12-24

Fig. 51. Knema pallens W.J. de Wilde. a. Habit of twig apex with leaves; b. part of older branch
with female inflorescences; c. opened female bud showing the pistil, pedicel with bracteole scar
about halfway; d. male bud; e. opened male bud showing the androecium, somewhat shrunken by
drying; f. ditto, androecium removed, showing furrowed inner basal part of perianth; g. androe-
cium; h. sample of hairs of male bud; i. fruit, with remnants of indumentum on the fruiting pedicel
[a—c: SAN 73312; d—h: SAN 29851; i: S 32964]. — Scale bar for a = 4 cm; for b, i = 2 cm; for c-f
= 4mm; for g = 1 mm; for h = 0.16 mm.

320 Flora Malesiana, Ser. I, Vol. 14 (2000)

pairs, slender, flat, indistinct above (sometimes raised in young leaves); venation indis-
tinct; petiole 8-12 by 1-1.5 mm. Inflorescences: sessile, brachyblast simple or + warted,
up to 4 mm long; in male 5—10-flowered, female 1—6-flowered; flowers with grey-brown
or yellowish hairs 0.1 mm long or less; perianth 3-lobed, yellowish inside. Male flow-
ers: pedicel 5-16 mm long, bracteole caducous, + median; buds (depressed) globose, at
base truncate or + sagged below the lobe sutures, 2.5—4 by 3.5—5 mm, cleft c. 4/5, lobes
1—1.2 mm thick; staminal disc flat, circular or blunt-triangular, 2-3 mm diameter; an-
thers 11-17, stiped, horizontal, 0.3—0.5 mm long, not touching; androphore 0.5—1 mm
long, tapering. Female flowers: pedicel (3—)5—8 mm long, bracteole + median; buds
obovoid, apical part + triangular in cross section, 4-5 by 4-5 mm, cleft 2/3, lobes 1 mm
thick; ovary ovoid, 2—2.5 mm long; stigma + sessile, + flat, 2-lobed and each lobe again
3- or 4-lobulate. Fruits 1-3 per infructescence, subglobose or obovoid, apex rounded
and often short-acuminate, base rounded or attenuate, 1.3—2.1 by 1—1.5 cm, with yel-
lowish brown or rusty mealy hairs 0.2—0.5 mm long; dry pericarp 1.5 mm thick; fruiting
pedicel (3—)5—11 mm long.

Field-notes — Bark smooth, rugulose, finely scaly, or flaking in thin strips; inner bark
pink or red, granular, or cream and fibrous; slash wood whitish. Flowers yellowish in-
side (always?), slightly fragrant. Fruits yellow or red, aril red.

Distribution — Malesia: Sumatra (Tapanuli, Palembang, Bangka), Peninsular Ma-
laysia (Perak, Kelantan, Trengganu, Pahang, Negri Sembilan, Malacca, Johore).

Habitat & Ecology — Primary and old degraded forest; hill slopes, ridges, ridge tops,
crests; also in seasonal swamp forest; 0-600 m altitude; fl. mainly Feb—May; fr. all
records June, July.

Note — In Sumatra possibly two forms exist, one with larger, the other with smaller
male perianth.

47. Knema pectinata Warb.

Knema pectinata Warb., Mon. Myrist. (1897) 556, t. 24; J. Sinclair, Gard. Bull. Sing. 18 (1961) 266,
f. 20: W.J. de Wilde, Blumea 25 (1979) 430; 27 (1981) 227; Tree Fl. Sabah & Sarawak 3 (2000)
432. — Myristica pectinata (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 90, nom. inval. —
Type: Beccari 1607, Sarawak.

Tree 10-25 m. Twigs sometimes faintly angled, 3-6 mm diameter, at first with yel-
lowish or rusty, + scale-like stellate hairs 0.1—0.2 mm long, early glabrescent; bark
striate, not tending to crack or flake. Leaves (rigidly) coriaceous, oblong to lanceolate,
(15-)20—45 by 3.5-10 cm, base broadly truncate, obtuse, or attenuate, apex obtuse or
acute(-acuminate); drying olivaceous to dark brown, often with scattered hair scars above;
lower surface early glabrescent or with subpersistent grey-brown hairs 0.1—0.3 mm long,
especially near the nerves; dots absent; midrib little raised above; nerves 25—50 pairs,
little raised but distinct above; venation raised and distinct; petiole 15—25 by 3—7 mm.
Inflorescences: sessile, brachyblast simple or + warted, 3-5 mm diameter, in male 2-6
(—10)-flowered; flowers with yellowish brown or rusty hairs 0.1 mm long or less; peri-
anth 3-lobed, pale pink inside. Male flowers: pedicel 5-15 mm long, bracteole caducous,
about median; buds broadly obovoid to depressed globose, (3.5—)4—7 mm diameter, cleft
3/4—4/5, lobes 0.8—2 mm thick; staminal disc flat or slightly concave, circular or sub-
triangular, 2—4 mm diameter; anthers 11-15, half-sessile to nearly stiped, horizontal or

De Wilde — Myristicaceae (Knema) 321

half-erect, 0.5—0.6 mm, not or hardly touching; androphore 1—2 mm long, clasped by the
swollen base of perianth. Female flowers not seen; stigma (on immature fruits) subsessile,
1-1.5 mm diam., 2-lobed and each lobe again with 4 or 5 laciniations. Fruits | or 2 per
infructescence, ellipsoid, 3-5 by 1.7—3 cm, sometimes flanged along the line of suture;
dry pericarp 2-4 mm thick, with hairs 0.2 mm long; fruiting pedicel 2-10 mm long.

KEY TO THE SUBSPECIES

la. Leaves coriaceous. Lower leaf surface with inconspicuous, sparse, minute hairs of
variable size, subpersistent or glabrescent. Fruits flanged along the line of suture or
(Ol. . . . RRRRAR Res rae. Sateen cee rane. e ie e ee a. subsp. pectinata
b. Leaves rigidly coriaceous. Indumentum of lower leaf surface largely persistent, of
densely set minute stellate hairs, mixed with fewer emergent ones. Fruits not flanged
PS. 6 chit ok, abe ch araiae cai Rare eae eee b. subsp. vestita

a. subsp. pectinata

Twigs 3—6 mm diameter, hairs 0.1—0.2 mm. Leaves coriaceous, on lower surface with
sparse indumentum or glabrescent; leaf blade oblong or lanceolate, sometimes parallel-
sided, up to 45 cm long, lateral nerves to 50 pairs. Fruits 3-5 cm long, flanged or not,
fruiting pedicel 2-10 mm long.

Field-notes — Bark hard, with a few large flakes and dents where flakes have fallen
out, otherwise smooth. Leaves dark glossy green, with whitish midrib above, glaucous
beneath. Perianth pale reddish inside. Immature fruits tawny or orange, flanged.

Distribution — Malesia: Borneo (Sarawak, Sabah).

Habitat & Ecology — Lower montane and montane forest; (400—)500—1500 m alti-
tude; fl. & fr. June—Oct.

Notes — 1. Subsp. pectinata is related to K. korthalsii and K. glomerata in various
aspects, but these species differ in having a convex staminal disc. It is also related to K.
woodii and K. scortechinii, with which it has the thick-fleshy perianth lobes in common;
K. woodii differs in less coriaceous leaves with fewer lateral nerves, and by the androecium
with 18—23 sessile anthers; K. scortechinii has stiped anthers, and different fruits.

2. The subspecies is variable in the size of the male buds and the number of anthers.
Small-flowered specimens, and specimens with very coriaceous leaves may be confused
with K. kinabaluensis.

b. subsp. vestita W.J. de Wilde

Knema pectinata Warb. subsp. vestita W.J. de Wilde, Blumea 27 (1981) 228; Tree Fl. Sabah &
Sarawak 3 (2000) 434. — Type: S 26349 (Ilias Paie), Sarawak.

Twigs stout, at apex 3-5 mm diameter, rusty puberulous by minute stellate scale-like
hairs 0.1 mm long or less. Leaves rigidly coriaceous, on the lower surface with greyish
to yellowish rusty subpersistent densely set stellate hairs 0.1—0.2 mm long or less, mixed
with fewer slightly longer hairs; leaf blade elliptic-oblong to lanceolate, (10—)20—30 by
3-10.5 cm, base rounded to cuneate, apex obtuse to acute-acuminate; midrib and nerves
raised above, nerves 25—30 pairs; venation fine, distinct at both surfaces. Flowers not
seen. Fruits 1 per infructescence, broadly ellipsoid-obovoid, 3—4(—5) by 2.7—3(—4) cm,

Flora Malesiana, Ser. I, Vol. 14 (2000)

fn
77 alti
e4ey eel
% fi
c
©

De Wilde — Myristicaceae (Knema) 323

not flanged along the line of suture, with hairs 0.1—0.2 mm long; fruiting pedicel 3 mm
long.

Field-notes — Trees 6—25 m tall, fruits orange or brown.

Distribution — Malesia: Borneo (Sarawak).

Habitat & Ecology — Forest at 1400-1600 m altitude.

Note — The subspecies is reminiscent of K. kinabaluensis from Sabah, the latter with
a less stout habit and on the lower leaf surface scattered minute stellate scale-like hairs,
the fruits generally smaller, and ridged.

48. Knema pedicellata W.J. de Wilde

Knema pedicellata W.J. de Wilde, Blumea 25 (1979) 474, f. 18; Tree Fl. Sabah & Sarawak 3 (2000)
434. — Type: § 16626 (Ilias Paie), Sarawak.

Tree 15-30 m. Twigs 4-6 mm diameter, at first with rusty hairs 0.5-1 mm long,
glabrescent; bark striate, not or but rarely tending to crack or flake. Leaves chartaceous,
elliptic or oblong, (12—)17-36 by (5—)7—13 cm, base shallowly cordate or subcuneate,
apex acute(-acuminate); glabrescent except midrib (which remains longer pubescent in
the basal part), drying greenish to brown, with distinct hair scars above; lower surface
greenish grey, finely papillate, with subpersistent mixed sessile and stalked hairs 0.3—
1.2 mm long; dots present, predominantly on the finer nerves; midrib slightly raised
above; nerves 14—22 pairs, flat or slightly raised; venation + flat, distinct; petiole late
glabrescent, 6-18 by 3-5 mm. Inflorescences: sessile, brachyblast simple, warted or
forked, 4-8 mm diameter; in male 20—50-flowered, female 10—40-flowered; flowers
with pale to dark brown hairs 0.1—0.5(—0.8) mm long; perianth (3- or) 4- or 5-lobed,
reddish (?) inside. Male flowers: pedicel slender, 20-30 mm long; bracteole | (or 2)
1-2 mm long, persistent, about median; buds depressed globose, + sagged at base, 3-4
by 4-5 mm, cleft 3/4—4/5, lobes 0.6—1 mm thick, perianth at base with a conspicuous
disc surrounding the androphore; staminal disc appressed to perianth disc, distinctly
mammillate, circular, 2-3 mm diameter; anthers 14-18, just stiped, horizontal, 0.5—0.6
mm long, spaced; androphore 0.6—1 mm long. Female flowers: pedicel 10-12 mm long,
bracteole persistent, about median; buds ellipsoid-obovoid, 6 by 5 mm, inside with disc
at base, cleft 1/2—2/3, lobes 1 mm thick; ovary subglobose, 1.5—2 mm diameter, stigma
subsessile, about 5-lobed, each lobe again shallowly 2—8-lobulate. Fruits 1—4 per
infructescence, subglobose, somewhat broader than long, 2—3 by 2.5—2.8 cm, with dark
brown hairs, 0.3—0.5 mm long; suture grooved, dry pericarp 4-5 mm thick; fruiting
pedicel 18-30 mm long. — Fig. 52.

Fig. 52. Knema pedicellata W.J. de Wilde. a. Branchlet with male inflorescences; b. male bud:
c. opened male bud showing disc at bottom and androecium, perianth lobes 4 or 5; d. halved male
bud showing disc and androecium with convex staminal disc; e. androecium seen from above;
f. sample of hairs: indumentum of male bud, half-schematic; g. branchlet with female inflores-
cences; h. opened female flower, showing disc at bottom of perianth and pistil; i. branchlet with in-
fructescences, fruits submature; j. sample of hairs of lower leaf surface, half-schematic; k. detail of
lower leaf surface with scattered persistent stellate-dendroid hairs and minute blackish dots on the
veinlets [a—f, j, k: S 16626 (Paie); g, h: S 34477; i: S 16682]. — Scale bar for a, g = 4 cm; for
b, c, h = 2.8 mm; for d = 1.4 mm; for e = 0.7 mm; for f, j = 0.16 mm; for k = 0.33 mm; for i= 2 cm.

324 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Bole buttressed to 50 by 30 cm; bark smooth or flaky; inner bark
brownish; sapwood whitish. Indumentum of young twigs, undersurface of leaves, and
flowers (yellowish) brown. Latex pinkish or dark red.

Distribution — Malesia: Borneo (Sarawak, Sabah).

Habitat & Ecology — Mixed dipterocarp forest; forest on sandstone, sandy soil, or
sandy clay derived from granite; ridges; 0-700 m altitude; fl. May—July; fr. Nov.—Dec.

Notes — 1. Knema pedicellata resembles in habit K. conferta and K. pubiflora, but is
readily distinguishable by the rather stout twigs, broad chartaceous leaves, long flower
pedicels, mammillate staminal disc, conspicuous disc at the base of the perianth, and
broad subglobose fruits. All three species have typicallly reddish brown or blackish dots
on the lower leaf surface. The male flowers have hairs 0.1—0.3 mm long, the only female
flowers seen (S 34477) have hairs 0.2—0.5(—0.8) mm long.

2. Most flowers have 4 or 5 perianth lobes; all other species of Knema have 3 perianth
lobes. Sometimes there are 2 bracteoles.

49. Knema percoriacea J. Sinclair

Knema percoriacea J. Sinclair, Gard. Bull. Sing. 18 (1961) 268, f. 21; W.J. de Wilde, Blumea 25
(1979) 392: 41 (1996) 392; Tree Fl. Sabah & Sarawak 3 (2000) 436. — Type: Kostermans 9945,
E Kalimantan.

Tree 5-25 m. Twigs sometimes faintly angled, 3-6 mm diameter, at first with pale to
dark brown hairs 0.5-2 mm long, late glabrescent; bark smooth or striate, then usually
slightly to distincly longitudinally cracking, and flaking. Leaves chartaceous or coriaceous,
(oblong-)lanceolate, 12—40 by 2.5—9(-15) cm, apex subobtuse to acute(-acuminate), base
attenuate to rounded, drying olivaceous or brown above, lower surface glabrous, not or
but little papillate, with scattered hair scars (lens!); dots absent; midrib much raised
above; nerves 15-25 pairs, raised above; venation distinct; petiole sometimes + pulvinate,
late glabrescent, 7-25 by 3-8 mm. Inflorescences: peduncle up to 2 mm long, brachyblast
simple, warted, or 2-furcate, up to 15 mm long; in male 10—20-flowered, female up to
15-flowered; flowers with light brown or dark brown hairs 0.5—2 mm long, sometimes
partly deciduous; perianth 3-lobed, red inside. Male flowers: pedicel 3—4.5 mm long,
bracteole persistent, at or above the middle; buds depressed ( globose-)obovoid, (3.5—)
4-5.5 by (3.5-)4-5.5 mm, cleft 2/3—3/4, lobes 0.5(-1) mm thick; staminal disc circular
or subtriangular, flat, 2-2.5 mm diameter; anthers 10-18, shortly stiped or subsessile,
half-erect or curved upward, 0.3—1 mm long, not or but little touching; androphore 1—1.5
mm, finely striate. Female flowers: pedicel 2-4 mm long, bracteole subapical, buds
ovoid or ellipsoid, 6.5—8(—9) by 5—7 mm, cleft slightly over halfway, lobes 0.5—0.7 mm
thick; ovary subglobose, 3.5—4 mm diameter, stigma sessile, 2-lobed and each lobe again
8—10-lobulate or serrate. Fruits 1 or 2 per infructescence, (broadly) ellipsoid, sometimes
slightly flattened, 3—4.5(—5) by 2.5-3.5(—4.5) em, with rusty hairs 1-2 mm long; dry
pericarp 4 mm thick; fruiting pedicel 2-3 mm long.

Field-notes — Bark of trunk rather smooth, fissured, scaly, or flaking; living bark
inside red-brown or pinkish; wood usually white, sometimes reddish, or red brown. Leaves
glossy dark green with whitish midrib above, glaucous with yellowish green midrib
beneath. Flowers with yellow to rusty indumentum. Fruits ellipsoid or round, with yel-
lowish or rusty indumentum.

De Wilde — Myristicaceae (Knema) B25

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, NE and E Kalimantan).

Habitat & Ecology — Primary and degraded forest, on hill sides, ridges; sandy soil,
clay, sandy loam soil; 0-700 m altitude; fl. & fr. throughout the year, but mainly July—
Nov.

Note — A complex species, in which three forms are distinguished. The species as a
whole seems closest related to K. latericia and K. furfuracea, both with longer male
flower pedicels. Knema latericia has thinner twigs, less coriaceous leaves and smaller
male flowers, with a different androecium. Also K. /unduensis 1s related; for differences
see under that species.

KEY TO THE FORMS

la. Twigs at apex 3—4 mm diameter, with rusty red hairs 0.5 mm long. Anthers 10-12.
lecavies conaceous, —E Bomeo. ... . 2.4 a.0% ee eee a. forma percoriacea

b. Twigs at apex 4-6 mm diameter, with either pale brown hairs 0.5 mm long, or hairs
chocolate or dark brown, 1-2 mm long. Anthers 11—15. Leaves chartaceous to
BOI ASC OMS 28 hss su ogc cox Rialy ino coi cues cron al ac ty ee eae Rn eee yy
2a. Leaves chartaceous. Twig apex with pale brown or yellowish brown hairs 0.3—1 mm
long. Anthers 11-14. — Sarawak, Brunei, Sabah, Nunukan Is..................
eo AER ee eB cate BS ape ee a c. forma sarawakensis

b. Leaves coriaceous [the venation on the lower surface indistinct]. Twig apex with
chocolate or dark brown hairs 1-2 mm long. Anthers 15. [Flowers with hairs | mm
ema TUNICL, SAT AW AK. es outs oy hice. ol sigesyites wieder cht Seb oud hice amen b. forma fusca

a. forma percoriacea

Twigs usually 3-4 mm diameter, smooth, not or little striate, with dense rusty red
hairs 0.5 mm long; bark lower down inconspicuously flaking. Leaves relatively small,
coriaceous; venation distinct at both surfaces. Indumentum of flowers of hairs 0.5(—1)
mm long. Anthers 10-12.

Distribution — Malesia: Borneo (E, SE, C, and W Kalimantan); usually in kerangas.

b. forma fusca W.J. de Wilde

Knema percoriacea J. Sinclair forma fusca W.J. de Wilde, Blumea 25 (1979) 394; Tree Fl. Sabah &
Sarawak 3 (2000) 437. — Type: BRUN 5164 (Ashton), Brunei.

Twigs 6 mm diameter, with dark brown hairs 1.5 mm long; bark early flaking. Leaves
coriaceous; venation indistinct on both surfaces. Flowers with dark brown hairs | mm
long. Anthers 15, subsessile.

Distribution — Malesia: Borneo (Brunei).

Note — In habit, not in the flowers, this form resembles K. plumulosa from Peninsu-
lar Malaysia.

c. forma sarawakensis W.J. de Wilde

Knema percoriacea J. Sinclair forma sarawakensis W.J. de Wilde, Blumea 25 (1979) 393; Tree FI.
Sabah & Sarawak 3 (2000) 437. — Type: S 23889 (E. Wright), Sarawak.

326 Flora Malesiana, Ser. I, Vol. 14 (2000)

Twigs 4—6 mm diameter, coarsely striate, with pale brown or yellowish brown hairs
(.3—1 mm long; bark coarsely striate, lower down coarsely flaking. Leaves generally
relatively large, chartacous to coriaceous; venation distinct on both surfaces. Indumentum
of flowers of hairs (0.5—)1 mm long. Anthers 11—14.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah, NE & C Kalimantan,
Nunukan Is.).

Note — In this form the indumentum of the flowers is often partly deciduous or easily
rubbed off, as in K. furfuracea.

50. Knema piriformis W. J. de Wilde

Knema piriformis W.J. de Wilde, Blumea 25 (1979) 400; Tree Fl. Sabah & Sarawak 3 (2000) 437.
— Type: SAN 34619 (Meijer), Sabah.

Tree 6-12 m. Twigs sometimes + angular, 1.5—3 mm diameter, at first with hairs 0.1
(0.2) mm long, glabrescent; bark smooth or finely striate, not tending to crack or flake.
Leaves thinly coriaceous, oblong or lanceolate, 15-25 by 3-7 cm, apex acute(-acumi-
nate), base rounded to attenuate; drying olivaceous or brown, with scattered hair scars
above: lower surface at first with soft greyish stellate hairs 0.1 mm long, early or late
glabrescent, sometimes mixed with sparse larger yellow-brown hairs, early shed and
leaving hair scars; dots absent; midrib + flat above; nerves 20-25 pairs, = flat; venation
faint or distinct; petiole 8-15 by 1.5—3 mm. Inflorescences: sessile, brachyblast simple
or forked, to 6 mm long; in male 5—10-flowered; flowers with hairs 0.1—0.2 mm long, or
less; perianth 3-lobed, yellow (7?) inside. Male flowers: pedicel 2-5 mm long, bracteole
+ caducous, at or above the middle; buds obovoid, 4—5 by 3-4 mm, tapering in the lower
half, cleft 1/2(—2/3), lobes 0.4—0.6 mm thick; staminal disc circular, flat, 1.5—2 mm
diameter; anthers 11 or 12, just stiped, almost horizontal to oblique, 0.3 mm long, not
touching: androphore slender, 1.5—2.2 mm long. Female flowers not seen. Fruits | or 2
per infructescence, ovoid or ellipsoid, faintly ridged, apex subobtuse, base subtruncate,
2.42.6 by 1.5-1.7 cm, with grey hairs 0.1 mm long; dry pericarp 2 mm thick; fruiting
pedicel 6 mm long.

Field-notes — Flowers yellow.

Distribution — Malesia: Borneo (Sarawak; Sabah: mainly at Kinabalu and its vicin-
ity).

Habitat & Ecology — Montane forest, hill sides, ridges; (400?—)1000—2000 m alti-
tude; fl. Apr.—July; fr. Jan.

Notes — 1. The obovoid male buds may be quite similar to those of K. latericia
subsp. albifolia, which differs in the flaking bark, a much more conspicuous indumentum
of the twig apices, and more pronounced, not largely sunken, lateral nerves. As judged
from the shape of the male buds, K. piriformis seems most related to K. pulchra and K.
oblongata.

2. Some sterile specimens (with the lateral nerves above largely sunken) are reminis-
cent of K. patentinervia (Peninsular Malaysia); they may be confused also with K. hirtella,
K. glauca, and K. cinerea (Lesser Sunda Islands), all latter species having + globose
male buds.

Ww
i)
ay

De Wilde — Myristicaceae (Knema)

51. Knema plumulosa J. Sinclair

Knema plumulosa J. Sinclair, Gard. Bull. Sing. 16 (1958) 312, f. 14; 18 (1961) 272; W.J. de Wilde,
Blumea 25 (1979) 426. — Type: SF 40455 (Sinclair & Kiah), Peninsular Malaysia.

[Myristica glaucescens auct. non Knema glaucescens Jack: Wallich, Cat. (1832) n. 6810, nom.
nud.] — Knema intermedia (Blume) Warb. var. dubia Warb., Mon. Myrist. (1897) 567.

Myristica furfuracea auct. non Hook. f. & Thomson: A. DC., Prod. 14, 1 (1856) 206.

Myristica cantleyi auct. non Hook. f.: King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 327, pl. 168, p.p..
excl. the Cantley specimens which are Knema laurina. — Knema cantleyi auct. non (Hook. f.)
Warb.: Warb., Mon. Myrist. (1897) 554, t. 24; Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 238;
Ridl., Fl. Malay Penins. 3 (1924) 68.

Tree 5-15 m. Twigs 3—5 mm diameter, at first with dense rusty lanose hairs 2-4 mm
long, glabrescent; bark grey-brown, smooth or finely striate, not tending to crack, inci-
dentally minutely flaking. Leaves coriaceous, oblong to lanceolate, 15—35(—45) by 2.5—
8(—9) cm, apex acute(-acuminate), base rounded to cuneate; drying dull greenish brown
above; lower surface (grey-)brown, at first with dense indumentum, glabrescent; dots
absent; midrib raised above; nerves 12-23 pairs, raised; venation distinct; petiole 10—25
by 2-4 mm, late glabrescent. Inflorescences: peduncle to 3 mm long, brachyblast simple
or warted, to 10 mm long; in male (3—)5—10-flowered, female 1—5-flowered; flowers
with dense hairs 1.5—2.5 mm long; perianth 3-lobed, creamy, greenish or pinkish (?)
inside. Male flowers: pedicel 5-12 mm long, bracteole subpersistent, subapical; buds
depressed globose to broadly obovoid, 4.5—5.5 by 5—7 mm, cleft 3/4—4/5, lobes 0.3—0.4
mm thick; staminal disc convex or mammillate, circular, 2.5—3.5 mm diameter; anthers
12-15, horizontal, 0.5—0.7 mm, shortly stiped, not touching; androphore (0.5—)1—2 mm
long, clasped by an annular disc | mm high at the base of the perianth, on which the
staminal disc rests. Female flowers: pedicel 2-6 mm, bracteole towards the apex; buds
obovoid, 7-9 by 7 mm, cleft over over halfway, lobes 0.4 mm thick; ovary subglobose,
2.5 mm diameter, with a conspicuous disc around the base; stigma sessile, 2-lobed and
each lobe again (5—)6—7-lobulate or serrated. Fruits 1-3 per inflorescence, ellipsoid,
3-3.5 by 2 cm, with coarse rusty hairs 2 mm long; dry pericarp 2-3 mm thick; fruiting
pedicel 3-6 mm long; perianth persistent under the fruit.

Field-notes — Bark pale brown, once scaly; inner bark white or red; sapwood white.
Leaves glossy dark green above, pale green or glaucous beneath, the nerves at both sides
much paler. Flowers cream or pinkish inside; staminal disc pink, anthers creamy.

Distribution — Malesia: Peninsular Malaysia (Perak, Trengganu, Selangor, Johore,
P. Penang).

Habitat & Ecology — Primary and logged forest on hill sides, ridges, and in flat land;
also in marshy forest and then provided with a few stilt-roots; 0-300 m altitude; fl. & fr.
throughout the year.

Notes — 1. Knema plumulosa is related to K. intermedia, a species with which it has
the distinct mammillate staminal disc in common; K. intermedia differs in its shorter,
not lanose indumentum, thinner twigs, smaller leaves, the perianth non-persistent in
fruits, and in the absence of a disc at the base of the perianth. According to Sinclair both
species have stilt-roots if growing in wet soil.

2. By the woolly indumentum of the young twigs, and the glabrous coriaceous leaves,
the species resembles K. percoriacea from Borneo, which differs in the flat staminal
disc, absence of perianth disc, and perianth not persistent under the fruit.

oS)

28 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 53. Knema pseudolaurina W. J. de Wilde. a. Habit of male flowering branchlet; b. detail of
the lower leaf surface showing areoles of venation and stellate hairs of mixed sizes; c. male bud;
d. opened male bud showing the androecium; e. androecium; f. female inflorescence; g. opened fe-
male bud showing the pistil; h. sample of hairs of female perianth (schematic); i. infructescence
[a, c-e: KEP FN 99355; b: Soepadmo & Mahmud 1169; f-h: FRI 7832: i: FRI 4044]. — Scale bar
for a, i= 2 cm; b: 0.33 mm; c, d, g = 2.8 mm; e = 0.7 mm; f = 6.5 mm; h = 0.16 mm.

De Wilde — Myristicaceae (Knema) 329

52. Knema pseudolaurina W.J. de Wilde

Knema pseudolaurina W.J. de Wilde, Blumea 25 (1979) 377, f. 2. — Type: Curtis 1044, Peninsular
Malaysia.

Knema laurina (Blume) Warb. var. malayana Warb., Mon. Myrist. (1897) 607, p. p., for the lectotype
only: Curtis 1044, Peninsular Malaysia.

Tree 5—30 m. Twigs (2—)3—5 mm diameter, with dense yellow brown or rusty hairs
(mixed) 0.05—0.5 mm long, (late) glabrescent; bark striate, not tending to crack or flake.
Leaves chartaceous or coriaceous, oblong to (ob)lanceolate, 12-40 by 3-12 cm, base
(sub)attenuate to rounded, rarely subcordate, apex acute(-acuminate), sometimes ob-
tuse; olivaceous above, lower surface finely papillate, with (largely) persistent pale yel-
lowish to brown sparse hairs 0.1—0.2 mm long, mixed with larger stellate-dendroid hairs
(lens!); dots absent; midrib flat or slightly raised above; nerves 18—30 pairs, flat or
+ sunken above; venation (very) fine, raised, distinct above; petiole 10—22 by 1.5—3.5
mm late glabrescent. Inflorescences: sessile, brachyblast simple or warted, or 2- or 3-fid,
to 10 mm long; in male 3—25-flowered, female 3—15-flowered; flowers with short dense
yellow-brown, rusty, or golden hairs 0.1—0.5 mm long; perianth 3-lobed, reddish inside.
Male flowers: pedicel 4.5—8.5 mm long, bracteole subpersistent or caducous, at or up to
2 mm below the apex; buds obovoid(-oblong), 4—6 by 2.5—4.5 mm, cleft 1/2—2/3, lobes
1-1.5 mm thick; staminal disc flat or slightly concave, circular, 1.5—2 mm diameter;
anthers 8—11, half sessile to just stiped, half erect to nearly horizontal, 0.3—0.6 mm long,
not or only little touching, androphore tapering, 1.5—2.5 mm long, rarely with hairs 0.1
mm at base. Female flowers: pedicel to 1.5 mm long, bracteole apical; buds ovoid-
oblong, 6-8 by 3.5-4.5 mm, cleft c. 1/3, lobes 1—1.5 mm thick; ovary globose or broadly
ovoid, 3—3.5 by 2.5 mm, style and stigma 2—2.5 mm long, the latter + spreading, 2-lobed
and each lobe again indistinctly 3—5-lobulate. Fruits 1-5 per infructescence, broadly
ovoid or oblong, apex obtuse or subacute, base (broadly) obtuse, (2.5—)3—5 by (1.5-)
2-3.5 cm, with stiff rusty hairs 1-2 mm long; dry pericarp 3 mm thick; fruiting pedicel
1-6 mm long. — Fig. 53.

Field-notes — Bark rather smooth or granulose, fine and thinly scaly, or somewhat
fissured; slash of outer bark pink(-orange), slash wood whitish; sap copious, red. Leaves
medium green above, dirty white or grey-white beneath. Fruits yellowish with short,
coarse, rusty indumentum. Seed whitish, aril red.

Distribution — Peninsular Thailand; in Malesia: Sumatra (Aceh, E Coast) and the
whole of the Malay Peninsula excluding Singapore.

Habitat & Ecology — Primary and degraded evergreen forest, in flat and hilly coun-
try, on crests; over granite, sandstone, limestone, and alluvial soils; 0—1000 m altitude;
fl. mainly Jan.—Apr.; fr. mainly Apr.—Aug.

Note — Specimens of K. pseudolaurina were formerly confused with K. laurina which
differs, among others, in the indumentum of the lower leaf surface, composed of stalked
dendroid hairs, shorter male flower pedicels, fewer anthers, and smaller fruits.

53. Knema psilantha W. J. de Wilde

Knema psilantha W.J. de Wilde, Blumea 25 (1979) 391; 41 (1996) 392: Tree Fl. Sabah & Sarawak
3 (2000) 438. — Type: S 14373 (Anderson), Sarawak.

330 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 5—10 m. Twigs 3-5 mm diameter, at first with woolly bright brown to yellowish
hairs 0.5—1 mm long, soon glabrescent; bark striate or ridged, lower down cracking and
flaking. Leaves thinly coriaceous, lanceolate, 20-50 by 3.5—9.5 cm, apex (narrowly)
acute, base obtuse or (broadly) rounded; drying greenish brown above; lower surface
glabrous, inconspicuously finely papillate; dots absent; midrib raised above; nerves
30-35 pairs, raised above; venation raised, distinct or not; petiole 5-15 by 3-5 mm, late
glabrescent. Inflorescences: sessile, brachyblast simple or + warted, 2-5 mm diameter;
in male 5—25-flowered, female 1—5-flowered; flowers with pale brown hairs 0.5—0.7
mm long, early glabrescent in the upper half or more; perianth 3-lobed, red inside. Male
flowers: pedicel 2-3 mm long, bracteole subpersistent, at or above the middle; buds
ellipsoid-obovoid, 4.5—5 by 3-3.5 mm, cleft 3/4—4/5, lobes 0.5 mm thick; staminal disc
circular, + flat, 1.5 mm diameter; anthers 6—9, each 0.5 mm long, stiped for 0.5—0.7 mm,
+ horizontal, not touching; androphore cylindrical, 1-1.2 mm long. Female flowers:
pedicel 1-2 mm long, bracteole + caducous, median; buds much larger than in male
flowers, ovoid, 8-9 by 5—5.5 mm, cleft c. 1/2, lobes 0.5 mm thick; ovary subglobose, 4.5
by 5 mm, stigma subsessile, + 2-lobed and each lobe again 5- or 6-lobulate or serrate.
Fruits 1-3 per infructescence, subellipsoid, apex obtusish, base broadly rounded, 2.5-3
by 2 cm, with rough yellowish brown hairs 1 mm long, sometimes partly glabrescent;
dry pericarp 3 mm thick; fruiting pedicel 2 mm long.

Field-notes — Bark smooth, with fine lenticels, outer bark brittle, inner bark white
mottled with reddish. Flowers cream-hairy outside, scarlet inside, in the males with a
paler spot at base of stamens.

Distribution — Malesia: Borneo (Sarawak, Sabah, NW & E Kalimantan).

Habitat & Ecology — Mixed dipterocarp forest on ridges, hill sides; 50-300 m alti-
tude; fl. Aug.—Sept.; fr. June.

Note — Knema psilantha resembles K. latericia and to a lesser extent K. furfuracea;
K. latericia differs in the more depressed globose mature male flower buds, not gla-
brescent, with different androecium, and less conspicuously flaking, more slender twigs;
K. furfuracea also has differently shaped male buds, with quite different androecium,
but in that species the flowers are often glabrescent as well. Knema psilantha is well
characterized by the partly glabrescent, ellipsoid, male flower buds, and only 6 or 7 con-
spicuously long-stiped anthers.

54. Knema pubiflora W.J. de Wilde

Knema pubiflora W.J. de Wilde, Blumea 25 (1979) 466, f. 17; Tree FI. Sabah & Sarawak 3 (2000)
438. — Type: SAN 25105 (Meijer), Sabah.

Tree 10—40 m. Twigs 1.5—3 mm diameter, at first with dense rusty scurfy hairs 0. 1—-0.2
mm long, glabrescent; bark coarsely striate, not tending to crack or flake. Leaves
chartaceous or thinly coriaceous, (ovate-)oblong, 4-15 by 1.5—7 cm, base rounded to
subattenuate, apex subacute to short acute-acuminate; drying dark brown above; lower
surface grey-brown, finely papillate, at first with mixed sessile and dendroid hairs 0.2—
0.5 mm long, persistent or late glabrescent; dots present; midrib slender, + raised above;
nerves 7-14 pairs, slightly raised above; venation distinct; petiole 5—10 by 0.7—2 mm,
late glabrescent. Inflorescences: peduncle to 2 mm long, brachyblast simple, warted, or
+ forked, to 4 mm diameter; in male 5—20-flowered, female 3—10-flowered; flowers

De Wilde — Myristicaceae (Knema) 331

|
:

b

Fig. 54. Knema pubiflora W.J. de Wilde. a. Branchlet with male inflorescences; b. male bud;
c. opened male bud showing hairy inner surface and androecium; d. androecium seen from above;
e. sample of hairs of indumentum on the outer surface of male perianth, half-schematic; f. opened
female bud showing pistil; g. pistil; h. lobe of female bud showing hairy inner surface; i. branchlet
with infructescences; j. sample of hairs of lower leaf surface, half-schematic; k. detail of lower
leaf surface with scattered stellate-dendroid hairs and minute blackish dots [a—e, j, k: SAN 25105
(Meijer); f-h: Kostermans 8634; i: Kostermans 10359]. — Scale bar for a, i = 2 cm; for b, f = 2.8
mm; for c, g, h = 1.4 mm; for d = 0.66 mm; for e, j = 0.16 mm; for k = 0.33 mm.

332 Flora Malesiana, Ser. I, Vol. 14 (2000)

with dark rusty hairs 0.1 mm long; perianth 3- (or 4-)lobed, inside densely stellate-hairy,
colour not known. Male flowers: pedicel 10-15 mm long, bracteole persistent or more
or less late caducous, about median; buds (depressed) globose, 2—2.5 by 2.5—3 mm, cleft
3/4-4/5, lobes 0.5(—0.7) mm thick; staminal disc convex, circular, 1.2—1.5 mm diam-
eter; anthers 10-13, just sessile, horizontal, 0.2-0.4 mm long, not touching; androphore
0.3-0.6 mm long. Female flowers: pedicel 7-10 mm long, bracteole + median; buds
obovoid or pear-shaped, 3—4 by 3—4 mm, cleft 1/2—2/3, lobes 0.5—1 mm thick; ovary
subglobose-ovoid, 1.5 mm diameter; stigma sessile, indistinctly 2-lobed and each lobe
again deeply (3- or) 4- or 5-lobulate. Fruits 1—5 per infructescence, subglobose or short
ellipsoid, often + flattened and ridged at one side towards the base, 1.5—2.5 by 1.5-2.3
cm, with rusty hairs 0.1—0.2 mm long; dry pericarp 1.5 mm thick; fruiting pedicel 10-14
mm long. — Fig. 54.

Field-notes — Bole with buttresses to 2 m high, extending to 30 cm; bark smooth,
when old peeling off in soft irregular small scales; living bark chocolate, brown, or red-
dish; sapwood reddish white, pale brown, or pale pinkish; latex reddish. Flowers with
rusty indumentum. Fruits greenish or yellow, with rusty indumentum; aril dark red, sweet-
acid.

Distribution — Malesia: Borneo (Sarawak, Sabah, NE, E & SE Kalimantan; many
collections from Nunukan Is.).

Habitat & Ecology — Forest on sandy soil, clay, or loam, also on brown soil; hill
sides, ridges, rolling sandy areas; in Shorea laevifolia forest, scattered but locally com-
mon; 0—300 m altitude; fl. & fr. throughout the year.

Note — Knema pubiflora may be confused with K. conferta and K. kunstleri, both
also with dotted lower leaf surfaces; K. conferta differs in a stouter habit, flowers gla-
brous inside, and flat staminal disc; K. kunstleri has a different indumentum on the lower
leaf surface consisting of sparse minute scale-like hairs, flowers glabrous inside, and flat
staminal disc. Knema pubiflora is the only species of Knema with the perianth hairy
inside and the leaves have a remarkable dark brown drying colour.

55. Knema pulchra (Miq.) Warb.

Knema pulchra (Miq.) Warb., Mon. Myrist. (1897) 600, t. 24; W.J. de Wilde, Blumea 25° (1979)
399: 27 (1981) 224; Tree Fl. Sabah & Sarawak 3 (2000) 439. — Myristica pulchra Miq., Ann.
Mus. Bot. Lugd.-Bat. 2 (1865) 51. — Type: Korthals s.n., Kalimantan.

Knema glaucescens Jack var. cordata J. Sinclair, Gard. Bull. Sing. 16 (1958) 310, f. 13D. — Knema
cinerea (Poir.) Warb. var. cordata (J. Sinclair) J. Sinclair, Gard. Bull. Sing. 18 (1961) 181. —
Type: SF 39959 (Sinclair & Kiah), Peninsular Malaysia.

Tree (2.5-)6-15 m. Twigs 3-6(—8) mm diameter, at first with short grey brown to
rusty hairs 0.1 mm long, glabrescent; bark finely striate, not tending to crack or flake.
Leaves chartaceous or thinly coriaceous, elliptic-oblong to oblong-lanceolate, (20—)30—
50 by (6-)10-17 cm, base rounded or cordate, apex acute(-acuminate), drying greenish
brown, often + glossy above; lower surface with persistent or deciduous sparse pale
brown or greyish stellate hairs 0.1—0.2 mm long; dots absent; midrib raised above; nerves
18-25 pairs, + flat above; venation (in)distinct; petiole 15—40 by 3-5 mm. Inflores-
cences: sessile, brachyblast simple, warted, or 2- or 3-furcate, to 10 mm long, in male
1—10-flowered, female 1—5-flowered; flowers with greyish brown hairs 0.1 mm long or

De Wilde — Myristicaceae (Knema) 359

less; perianth 3-lobed, creamy or pale brown inside. Male flowers: pedicel 4-8 mm
long, bracteole caducous or subpersistent, median to subapical; buds obovoid, (3—)4-5.5
by 3-5 mm, cleft 1/2—3/4, lobes 0.7—1 mm thick; staminal disc circular or faintly trian-
gular, flat or concave, 1.8-3 mm diameter; anthers 12-25 (14-25 in Borneo), + half-
erect, just sessile, 0.5—-0.7 mm long, not or only little touching; androphore slender,
sometimes tapering, 0.8—1.5 mm long. Female flowers: pedicel (2-)5—8 mm long, brac-
teole about median; buds obovoid-oblong, 5—6 by 3.5—4.5 mm, cleft c. 1/2, lobes 1 mm
thick; ovary ovoid, 2.5-3 by 1.5—2 mm; style 1.5 mm long, stigma + flat, inconspicu-
ously 2-lobed and each lobe again 2—4(—5)-lobulate. Fruits variable, | or 2 per infruc-
tescence, subglobose to ellipsoid-oblong, ridged and sometimes slightly saccate at base
or not, apex obtuse to subacute, (3—)4—6 by 2—3.5 cm, with hairs (O.1—)0.2 mm long,
sometimes glabrescent; dry pericarp 3-4 mm thick; fruiting pedicel (2—)5—12 mm long.

Field-notes — Sometimes with a few stilt-roots; bark smooth with a few lenticels,
pustulate, or finely fissured; living bark orange-brown; wood pale brown or white. Leaves
dark green with whitish green midrib and nerves above, glaucous with brownish midrib
and nerves beneath. Perianth pale brown or cream inside.

Distribution — Malesia: Peninsular Malaysia (Trengganu, Johore), Borneo (Sarawak,
Brunei, Sabah, Kalimantan).

Habitat & Ecology — Primary forest, lowland dipterocarp forest, swamp forest, also
on ridges; sandy soil, loam soil with lime, swampy soil, basalt-derived soils; 0-1000 m
altitude; fl. May—Sept.; fr. Mar.—Aug.

Note — Knema pulchra is close to the more slenderly built K. piriformis and possibly
also related to K. oblongata, which differs in the conspicuous rusty indumentum on
twigs, lower leaf surfaces, and flowers. In general habit, not in the shape of the flowers,
it recalls K. kostermansiana, also a broad-leaved species.

56. Knema retusa (King) Warb.

Knema retusa (King) Warb., Mon. Myrist. (1897) 612, t. 25; J. Sinclair, Gard. Bull. Sing. 16 (1958)
318: 18 (1961) 272, f. 22; W.J. de Wilde, Blumea 25 (1979) 384. — Myristica retusa King, Ann.
Roy. Bot. Gard. Calc. 3 (1891) 330, pl. 171. — Type: King’s coll. 7690, Peninsular Malaysia.

Tree 5-20 m. Twigs 5—6 mm diameter, at first with pale rusty hairs 0.1—0.2 mm long,
glabrescent; bark (coarsely) striate, not tending to crack or flake. Leaves coriaceous,
obovate to (elliptic-)oblong, (20-)30-60 by (10—)14—25 cm, apex broadly obtuse, often
retuse, base rounded or cordate; drying brown above, lower surface with persistent dense
grey, pale brown, or rusty hairs 0.1—0.2 mm long, + silky to the touch; dots absent:
midrib stout, flat or + depressed above; nerves 15-25 pairs, widely (2—4 cm) apart, + flat
above: venation distinct; petiole 15—30 by 5-10 mm. Inflorescences: peduncle to 5 mm
long, brachyblast simple, warted, or 2- or 3-furcate, 5-20 mm long; in male 5—10-flow-
ered, female 1—5-flowered; flowers with brownish hairs 0.1-0.3 mm long; perianth
3-lobed, whitish inside. Male flowers: pedicel 10-15 mm long, the bracteole caducous
or subpersistent, 2-4 mm long, apical; buds broadly obovoid, 7-8 by 6-7 mm, cleft
about 4/5, lobes 1 mm thick; staminal disc subtriangular, flat or + concave, 4 mm diam-
eter: anthers c. 15, subsessile, + horizontal or somewhat oblique, 0.6—0.7 mm long, not
touching; androphore tapering, |—1.5 mm long. Female flowers: pedicel 3-5 mm long,
bracteole caducous, apical; buds obovoid, 9 by 6-7 mm, cleft c. 2/3, lobes | mm thick;

334 Flora Malesiana, Ser. I, Vol. 14 (2000)

ovary ovoid or subglobose, 3.5—4 by 3 mm; style 1 mm long; stigma 2-lobed and each
lobe with 2 or 3 serrations of variable depths. Fruits 1-3 per infructescence, ovoid-
ellipsoid, 3—4.5(—6?) by 2-3.5(—5?) cm, apex 3 mm apiculate, with chocolate hairs 0.2
mm; dry pericarp 3-4 mm thick; fruiting pedicel 8-10 mm long.

Field-notes — Bark of trunk dark brown, slightly rough, but not flaking nor fur-
rowed. Leaves dark green, glossy, with whitish green midrib above, greyish or glaucous
with a cover of minute pale brownish scale-like or stellate hairs beneath. Male buds 10-
12 by 7 mm, whitish inside, slightly fragrant, pollen whitish; female buds 14 by 10 mm.

Distribution — Malesia: Peninsular Malaysia (Perak).

Habitat & Ecology — Mixed lowland forest; flat land; 0-500 m altitude; fl. Oct.;
fr. May.

Notes — 1. Knema retusa is a rare species of a restricted area; no recent collections
are known. Its flowers are large for the genus, and for Myristicaceae in general.

2. Sinclair described the anthers as stalked, but I found them rather subsessile. King
described the aril as embracing only the base of the seed, but possibly this was an inci-
dental case, or due to the drying.

57. Knema riangensis W. J. de Wilde

Knema riangensis W.J. de Wilde, Blumea 32 (1987) 138: Tree Fl. Sabah & Sarawak 3 (2000) 440.
— Type: Veldkamp 7941, C Kalimantan.

Tree 3-12 m. Twigs subterete or + angular, 1.5—3(—4) mm diameter, with minute dull
brown hairs 0.1 mm long, glabrescent; bark coarsely striate, not flaking. Leaves mem-
branous, elliptic-oblong to oblong-lanceolate, 14-30 by 5—10.5 cm, base short-attenuate
or rounded, apex acute-acuminate; drying olivaceous above, paler, with (sub)persistent
dense pale yellowish to greyish almost uniform hairs 0.1(—0.2) mm long beneath; dots
absent: midrib raised above; nerves 15—23 pairs, + flat to raised above, venation distinct;
petiole 10-20 by 2-4 mm, late glabrescent. /nflorescences: peduncle to | mm long, bra-
chyblast wart-like, simple, or forked, to 5 mm long; in male 2-6-flowered, female 2—-4-
flowered: flowers with dull brown hairs 0.1(—0.2) mm; perianth 3-lobed, reddish inside.
Male flowers: pedicel 3—4 mm long, bracteole persistent, 0.5 mm long, 0.5—1 mm below
the apex; buds broadly obovoid, 2 by 1.7 mm, cleft 5/6, lobes 0.5 mm thick; staminal
disc circular or subtriangular, flat, 0.8—1 mm diameter, anthers 9, sessile, suberect, 0.3
mm long, closely set; androphore tapering (obconical), 0.8 mm long, sometimes with
few minute hairs at base. Female flowers: rather stout, pedicel S12 mm long, bracteole
slightly above median; buds + obovoid, 4-5 by 3 mm (or 7 by 4 mm, see note 2), cleft
c. 1/2, lobes 0.5—1 mm thick; ovary ovoid, 2-4 mm long, with dense hairs 0.1(—0.2) mm
long; style broad, 0.5 mm long, stigma lobes 2, each coarsely 2—4-lobulate. Fruits 1 or 2
per infructescence, ellipsoid, 2.4—4 by 1.7—2.5 cm, with persistent rust-brown hairs 0.1
mm long, dry pericarp 1-2 mm thick; fruiting pedicel 8-12 mm.

Field-notes — Young shoots cinnamon. Leaves glossy above, whitish underneath.
Flowers (pink-)red, column yellowish. Fruits orange, aril edible, somewhat sour, bright
red; pericarp edible, sweetish; outer testa white, inner black.

Distribution — Malesia: Borneo (Sarawak: possibly; C Kalimantan).

Habitat & Ecology — Primary forest, riverine forest; yellow sandy clay; 100—600 m
altitude; fl. & fr. Jan., Feb.; fr. May.

De Wilde — Myristicaceae (Knema) 335

Notes — 1. Knema riangensis is, among its relatives, characterized by a lower leaf
surface with (sub)persistent dense slightly interwoven hairs of almost uniform size,
0.1—0.2 mm long, male buds small, 2 by 1.7 mm, cleft nearly to the base, androphore
distinctly tapering, sometimes with a few minute hairs at base (which points to relation-
ship with K. hirtella), anthers 9, sessile, suberect, close to each other, style distinct, 0.5
mm long (almost as in K. stylosa), and medium-sized fruits, with short persistent indu-
mentum.

2. Veldkamp 8209 has considerably large female flowers, pedicel 12 mm long, buds 7
by 4 mm, ovary 4 by 3 mm, much larger than those in Veldkamp 7970 (a collection of
mixed male and female specimens).

58. Knema ridsdaleana W.J. de Wilde

Knema ridsdaleana W.J. de Wilde, Blumea 41 (1996) 392. — Type: Ridsdale ISU 451, Philippines,
NE Luzon.

Small tree 8 m. Twigs 3(—4) mm diameter, at first with dense rusty hairs 0.5(—1) mm,
late glabrescent; bark blackish, coarsely striate, not tending to crack or flake, lenticels
not apparent. Leaves chartaceous(-coriaceous), (ovate-)oblong, 15—25 by 5.5—8(—9) cm,
base broadly rounded, apex acute-acuminate; drying olivaceous or dark-brown above,
lower surface (ashy-grey, glabrescent but indumentum + remaining on and near lower
part of midrib; minutely punctate but (larger) dots absent; midrib raised above, nerves
15-18 per side, raised above, venation distinct at both surfaces; petiole 15(—20) by (2—)
2.5 mm, late glabrescent. Inflorescences: pubescent, among the leaves, peduncle 10-15
by 2-2.5 mm, brachyblast simple or 2- or 3-forked, to 5 mm long; in male with subumbels
of 1-3 buds of various sizes; flowers with dense dark brown woolly hairs 0.3(—0.5) mm,
as a dense mat, at anthesis (when dry) largely falling off in large flakes, leaving the peri-
anth completely glabrous; perianth 4-lobed, colour inside not known. Male flowers:
pedicel 3—4(—5) mm long, bracteole persistent, (2-)2.5 mm long, | mm below the apex,
with low ridges decurrent on the pedicel; buds depressed (sub)globose, 3 by 4—4.5 mm,
cleft 4/5(—9/10), lobes 0.4 mm thick; staminal disc flat, circular, 1.8(—2) mm diameter,
anthers + contiguous, | 1-13, half-sessile, and adnate to the tapering androphore, anthers
4-sporangiate, thecae nearly 1 mm long, androphore short (less than 0.5 mm), striate.
Female flowers and fruits not seen.

Field-notes — Small tree, 8 m; bark dark brown, flaky, inner bark red. Flowers red-
dish brown.

Distribution — Malesia: Philippines (NE Luzon, Palanan area; known only from the
type).

Habitat & Ecology — Streamside forest over ultrabasic rock; c. 50m altitude; fl. Apr.

Note — This species is superficially reminiscent of K. glomerata (especially in the
leaves) and K. tomentella (as regards the conspicuous indumentum, falling off in pieces).
Knema ridsdaleana is readily distinguishable by its grey leaf undersurface, with minute
dark punctation (not to be confused with coarser dots, strong lens!), conspicuous in-
dumentum, long-stalked (male) inflorescences, 4-merous perianth, with conspicuously
shedded indumentum, and 11-13 large (long) + contiguous subsessile 4-sporangiate
anthers.

336 Flora Malesiana, Ser. I, Vol. 14 (2000)

59. Knema rigidifolia J. Sinclair

Knema rigidifolia J. Sinclair, Gard. Bull. Sing. 16 (1958) 284, f. 5, excl. BD; 18 (1961) 274;
W.J. de Wilde, Blumea 25 (1979) 398. — Type: SF 33226 (Corner), Peninsular Malaysia.

Tree 5-15 m. Twigs 2—4(—5) mm diameter, smooth or striate, at first with hairs 0.1—
0.2 mm long, glabrescent; bark smooth or striate, not tending to crack or flake. Leaves
coriaceous, (oblong-)lanceolate, base attenuate, rounded, cordate, or subtruncate, apex
subobtuse or acute-acuminate, 12-28 by 4—8.5 cm, drying brown, with scattered hair
scars above; lower surface late glabrescent or with subpersistent scale-like hairs 0.1 mm
long or less, sometimes mixed with weak stellate-dendroid hairs 0. 1—0.2 mm long; dots
(almost) absent; midrib hardly raised above; nerves 15-28 pairs, + flat, venation raised
and distinct; petioles 15—25 by 2-4 mm. Inflorescences: sessile, brachyblast simple or
warted, up to 5 mm diameter; in male 5—20-flowered, female 1—5-flowered; flowers
with rusty hairs 0.1-0.2 mm long; perianth 3-lobed, reddish (?) inside. Male flowers:
pedicel 3-5 mm, slender or thickish, 0.5(—1) or 1.5-2 mm diameter, bracteole caducous,
about median; buds depressed broadly obovoid, variable in size and consistency (see
under the subspecies), 3.5—4 by 4—4.5 mm, cleft 3/4—4/5, lobes 0.5 or 1-1.5 mm thick;
staminal disc circular, + concave, 1.7—2.2 mm diameter; anthers 8 or (12?—)14—16 (see
the subspecies), half-sessile, 0.5—0.7 mm long, suberect, not or hardly touching;
androphore tapering, 1-1.4 mm long. Female flowers not known. Fruits 1 (or 2?) per
infructescence, almost globose, 3—4 cm diameter, with hairs 0.2 mm long; dry pericarp
2—4 mm thick; fruiting pedicel 1-3 mm long.

Field-notes — Bark smooth, slightly fissured, or faintly hooped. Flowers borne be-
hind the leaves. Leaves grey at underside.

Distribution — Malesia: Peninsular Malaysia (Pahang, Selangor).

Habitat & Ecology — Montane forest, on crests and slopes; 1000-1800 m altitude;
fl. Feb.—Apr.; fr. Apr.—Sept.

Note — Knema rigidifolia is a montane species, characterized by coriaceous leaves
and globose fruits. The male flowers of the two specimens known differ in shape, con-
sistency, and morphology, related to differences in general habit, leaf shape, and distri-
bution. By the differences in the male flowers in the two here accepted subspecies, K.
rigidifolia keyed out on two places in the general key to the species.

KEY TO THE SUBSPECIES

la. Twigs at apex slender, 2-4 mm diameter. Leaves coriaceous, at base rounded or
attenuate. Male flowers not robust; pedicel 0.5-1 mm diameter; buds cleft c. 4/5,
fobes 025 gin Eck aNUNCES O° 2 cere nace suc «koe cee ore se ae a. subsp. rigidifolia

b. Twigs stout, at apex 3-5 mm diameter. Leaves rigidly coriaceous, at base rounded or
cordate. Male flowers robust; pedicel 1.5—2 mm diameter; buds cleft 2/3—3/4; lobes
Ge CISA sgea ns cacy kee meine hase: eRe ea a b. subsp. camerona

a. subsp. rigidifolia

Twigs in apical part 2~4 mm diameter, smooth or finely striate. Leaves coriaceous, at
base rounded or attenuate. Male flowers: pedicel 3-3.5 mm long, 0.5—1 mm diameter;
buds 3.5 by 4 mm, cleft c. 4/5, lobes 0.5 mm thick. Staminal disc including anthers 1.7—

De Wilde — Myristicaceae (Knema) B5i/

1.8 mm diameter: anthers 8, half-sessile, half-erect; androphore 1.4 mm long. Fruits
subglobose, 3—3.5 cm diameter

Distribution — Malesia: Peninsular Malaysia (Pahang: Fraser’s Hill, Pine Tree Hill,
and Selangor: G. Bunga Bua).

Note — Knema rigidifolia subsp. rigidifolia resembles K. kinabaluensis from Sabah
and K. pectinata from Sarawak (see there).

b. subsp. camerona W. J. de Wilde

Knema rigidifolia J. Sinclair subsp. camerona W.J. de Wilde, Blumea 25 (1979) 398. — Type: SF
s.n. (Henderson), Peninsular Malaysia.

Twigs in apical part 3-5 mm diameter, coarsely striate. Leaves rigidly coriaceous, at
base broadly rounded or shallowly cordate. Male flowers robust; pedicels 5 mm long,
1.5-2 mm diameter; buds 4 by 4—4.5 mm, cleft 2/3—3/4, lobes 1—-1.5 mm thick. Staminal
disc including anthers 2-2.2 mm diameter; anthers 14-16, half-sessile, suberect;
androphore stout, 1-1.2 mm long. Fruits 3.84 cm diameter.

Distribution — Malesia: Peninsular Malaysia (Pahang, near the Pahang/Perak bor-
der: Cameron Highlands), at 1500-1800 m altitude.

60. Knema rubens (J. Sinclair) W.J. de Wilde

Knema rubens (J. Sinclair) W.J. de Wilde, Blumea 25 (1979) 450. — Knema glaucescens Jack var.
glaucescens forma rubens J. Sinclair, Gard. Bull. Sing. 16 (1958) 307, f. 13B. — Knema cinerea
(Poir.) Warb. var. rubens (J. Sinclair) J. Sinclair, Gard. Bull. Sing. 18 (1961) 185, p.p., excl.
specimens from Borneo. — Type: Ridley 4819, Singapore.

Tree 5—25 m. Twigs + flattened or blunt-triangular, 1.5—2 mm diameter, at first with
short farinose yellow-brown, reddish, or pale rusty hairs 0.1—0.2 mm long, glabrescent;
bark smooth, then striate, not tending to crack or flake. Leaves membranous or thinly
coriaceous, elliptic-oblong to lanceolate, (5—)8—25 by 2—5(-8.5) cm, base attenuate, rarely
rounded, apex acute-acuminate, drying greenish or brown, often with a very fine-pitted
structure above, (lens!), lower surface greyish, with a reddish tinge, glabrescent except
for the basal part of the midrib with farinose stellate scale-like hairs; dots absent; midrib
raised above; nerves (10—)15~—25 pairs, + raised above: venation faint or distinct; petiole
late glabrescent, 6-11 by 1.5—2.5 mm. Inflorescences: peduncle up to | mm long, brachy-
blast simple or 2- or 3-fid, to 5 mm long; in male 5-10-flowered, female 2—5-flowered;
flowers with rusty hairs 0.1—0.2 mm long; perianth 3-lobed, reddish inside. Male flow-
ers: pedicel 3.5—4.5 mm long, bracteole subpersistent, about median; buds (depressed)
globose, 2-3 by 2-4 mm, cleft c. 4/5, lobes 0.8 mm thick; staminal disc slightly convex
or low-mammillate, circular, 2-2.2 mm diameter; anthers 10-12, shortly stiped, hori-
zontal, 0.4 mm long, spaced; androphore 0.6—1 mm long. Female flowers: pedicel 1.5—
3 mm long, the bracteole subapical; buds ellipsoid to oblong, 3-3.5 by 22.5 mm, cleft
c. 1/2, lobes 0.5 mm thick; ovary ovoid-globose, 1.5 mm diameter, stigma subsessile,
faintly 2-lobed and each lobe again shallowly 1—3-lobulate. Fruits |— 4 per infructescence,
ellipsoid, base and apex obtuse or subacute, 1.7—2.7 by 1.2—2 cm, with mealy yellowish
brown or rusty hairs 0.2 mm long; dry pericarp 1.5—2 mm thick; fruiting pedicel 2-7 mm
long.

338 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 55. Knema rufa Warb. a. Habit of twig with male inflorescences; b. immature male bud with
the pedicel not yet fully developed; c. immature androecium; d. fruit, the bracteole not discernible,
apparently minute, attached at the base of the fruiting pedicel [a: S 13146; b, c: Sinclair 10451;
d: Sinclair 10191]. — Scale bar for a = 2 cm; for b = 4 mm; for c = 1.65 mm; for d= 1 cm.

De Wilde — Myristicaceae (Knema) 559

Field-notes — Bole straight, slender; bark smooth or slightly flaky; inner bark red,
granulate; slash wood (yellowish) white. Leaves dark green above, dirty whitish, grey-
green, or glaucous beneath. Fruits yellow(-brown) or orange, aril bright red.

Distribution — Malesia: N Sumatra (E Coast, Tapanuli, Simeulué I.), Peninsular Ma-
laysia (Kedah, Kelantan, Trengganu, Pahang, Johore, Penang I.), Singapore.

Habitat & Ecology — Lowland forest, often along streams; sandstone, red soil; alti-
tude 0-800 m; fl. Feb., Apr., Aug.; fr. in Peninsular Malaysia mainly July—Sept., in Suma-
tra Jan., June, Sept.

Note — Knema rubens is characterized by the more or less flattened twig apices,
which also occur in K. luteola and K. oblongifolia; it resembles in appearance K. latericia
subsp. ridleyi which differs in the scaly bark of the (subterete) twigs, more pronounced
convex staminal disc, and thicker indumentum on twigs and flowers.

61. Knema rufa Warb.

Knema rufa Warb., Mon. Myrist. (1897) 556, t. 24; J. Sinclair, Gard. Bull. Sing. 18 (1961) 274, 288,
f. 23: W.J. de Wilde, Blumea 25 (1979) 409, f. 6; Tree Fl. Sabah & Sarawak 3 (2000) 441. —
Myristica rufa (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 90, nom. alt. inval. — Type:
Beccari 1798, Sarawak.

Tree 5-20 m. Twigs slightly angular, 2-3 mm diameter, at first with rusty hairs 0.3
mm long, glabrescent; bark faintly striate, not or very faintly tending to crack or flake.
Leaves thinly coriaceous, oblong to lanceolate, 15-30 by 1.5—4(—5.5) cm, base attenu-
ate, apex acute or long acute-acuminate; drying olivaceous to blackish brown above,
lower surface dull grey-brown; at both surfaces at first with dense rusty hairs 0.5 mm
long, early glabrescent; dots absent; midrib raised above; nerves 15—25(—30) pairs, with
lesser side nerves in between, raised above; venation distinct; petiole 10-20 by 2-3 mm.
Inflorescences: peduncle up to 2 mm long, brachyblast simple, to 10 mm long; in male
2-10-flowered, female 1—3-flowered: flowers with rusty hairs 0.3 mm long; perianth
3- (or 4-)lobed, red inside (always?). Male flowers: pedicel 10-15 mm, bracteole cadu-
cous, above the middle; buds depressed obovoid, when dry + furrowed, 6—7 by 5-6 mm,
cleft c. 2/3, lobes inside with finely warty ridges, 0.5 mm thick; staminal disc circular,
concave, 2—2.5 mm diameter; anthers 9-15, largely sessile, suberect, not or but little
touching, the connectives extended slightly beyond the thecae; androphore tapering,
1 mm long. Female flowers not seen. Fruits 1(—3) per infructescence, ellipsoid, 2.5 by
1.5 cm, with rusty + glossy hairs 3 mm long; dry pericarp 2(—3) mm thick; fruiting pedi-
cel 10-20 mm long. — Fig. 55.

Field-notes — Bark not furrowed nor flaking, sparsely covered with lenticels. Leaves
dull dark green with whitish green midrib above, glaucous with yellowish green midrib
beneath. Male flowers red or cream inside.

Distribution — Malesia: Borneo (Sarawak, Brunei).

Habitat & Ecology — Primary mixed dipterocarp forest; sandy clay soil; 0-200 m
altitude.

Note — The shaggy indumentum of the fruits is quite unlike that of other species of
the genus Knema.

340 Flora Malesiana, Ser. I, Vol. 14 (2000)

62. Knema scortechinii (King) J. Sinclair

Knema scortechinii (King) J. Sinclair, Gard. Bull. Sing. 16 (1958) 288, f. 7; 18 (1961) 276; W.J. de
Wilde, Blumea 25 (1979) 431. — Myristica scortechinii King, Ann. Roy. Bot. Gard. Calc. 3
(1891) 317, pl. 153 (excl. female fl.). — Knema conferta (King) Warb. var. scortechinii (King)
Warb., Mon. Myrist. (1897) 580. — Lectotype: King’s coll. 5617; several syntypes, Peninsular
Malaysia.

Tree 5—20 m. Twigs 2—4(—5) mm diameter, at first with pale yellow to rusty hairs 0.1—
0.2 mm long, glabrescent; bark smooth or + wrinkled, not longitudinally striate, not
cracking or flaking. Leaves coriaceous, oblong or lanceolate, 10-32 by 2—8.5 cm, base
subcordate to cuneate, apex subobtuse or acute(-acuminate); drying greenish brown to
brown above; lower surface greyish, not distinctly papillate, late glabrescent or with
persistent golden to pale brown or greyish, usually sessile-stellate hairs 0.2—0.4 mm
long; dots absent; midrib raised above; nerves 18-30 pairs, raised above; venation
+ raised, distinct; petiole 5-20 by 1.5—4(—5) mm. Jnflorescences: sessile, brachyblast
simple or 2—4-furcate, to 10 mm long; in male 4—10-flowered, female 1—5-flowered;
flowers with rusty hairs 0.1—0.2 mm long; perianth 3- (or 4-)lobed, hard-carnose, creamy
(?) inside. Male flowers: pedicel 7-16 mm long, bracteole caducous, about median; buds
depressed globose, 3.5—5 mm diameter, cleft c. 5/6, lobes (0.8—)1—2 mm thick; staminal
disc flat, circular or subtriangular, 2-2.5 mm diameter; anthers 11—16, just sessile to
short-stiped, 0.5 mm, horizontal, not touching; androphore + slender, 1—1.5 mm long.
Female flowers: pedicel 3-11 mm long, bracteole about median; buds obovoid, 4.5—5
mm long, cleft over 1/2, lobes 1 mm thick; ovary subglobose to ovoid, 1.5—3 mm diam-
eter, style 0.5—1.5 mm long, stigma + flat, many-lobulate or + 2-lobed and each lobe
again 5—8-lobulate. Fruits 1-3 (or 4) per infructescence, subglobose or ellipsoid, 2—2.5
by 1.5—2 cm, with rusty hairs 0.1-0.3 mm long, subglabrescent; dry pericarp 1-2 mm
thick; fruiting pedicel (4—)10—12 mm long.

Field-notes — Bark dark brown with a dark reddish tinge, (nearly) smooth, adherent
scaly and probably flaky later, or closely fissured; slash inner bark pink red. Leaves dark
green, glossy above, glaucous beneath.

Distribution — Malesia: Sumatra (E Coast, Indragiri), Peninsular Malaysia (Kedah,
Perak, Kelantan, Trengganu, Pahang, Selangor, Negri Sembilan, Malacca, Johore).

Habitat & Ecology — Primary and disturbed rain forest; sandy soil; 0-500 m alti-
tude; fl. Mar.—June; fr. June, Sept., Oct.

Note — Knema scortechinii may be confused with K. conferta, which differs in the
presence of dark dots on the lower leaf surface, a coarser indumentum, and striated
twigs.

63. Knema sericea W.J. de Wilde

Knema sericea W.J. de Wilde, Blumea 25 (1979) 383; 41 (1996) 394; Tree Fl. Sabah & Sarawak
3 (2000) 441. — Type: Jaheri 6/1, NE Kalimantan.

Tree 10-20 m. Twigs 4—5 mm diameter, at first with dense grey-rusty scale-like hairs
0.1 mm long or less, glabrescent; bark finely striate, not tending to crack or flake. Leaves
membranous or chartaceous, oblong(-lanceolate), 30—45 by 9-15.5 cm, apex rounded,
retuse, or acute-acuminate, base rounded to shallowly cordate, with the basal nerves

De Wilde — Myristicaceae (Knema) 34]

somewhat crowded, drying greenish to blackish brown above, lower surface very finely
papillate or not, with persistent short, dense, greyish brown or pale cinnamon hairs 0.2
mm long, silky to the touch, mixed with sparse somewhat stouter ones; dots absent;
midrib raised above; nerves 30-35 pairs, raised above; venation very fine, distinct; peti-
ole 12-20 by 4-6 mm. Inflorescences: peduncle stout, 5—7 mm long, brachyblast + sim-
ple, 5 mm diameter; in male 4—10-flowered, female 2—5-flowered; flowers with dense
greyish brown scurfy hairs 0.1 mm long; perianth 3-lobed, colour inside not known.
Male flowers: pedicel 12-18 mm long, bracteole caducous, (subapical; buds subglobose,
6-10 mm diameter, cleft c. 4/5, lobes 0.5 mm thick; staminal disc circular, convex to
mammillate, 3 mm diameter; anthers 10—15, long-stiped, + hanging down, horizontal,
or slightly curved upwards, 0.6—0.7 mm long, not touching; androphore 0.5 mm long.
Female flowers not known. Fruits | or 2 per infructescence, subglobose to broadly ovoid,
apex 1-2 mm beaked, 3.5—5 by 3.5—4 cm, with grey-brown to rusty scale-like hairs 0.1
mm long; dry pericarp 5 mm thick; fruiting pedicel 15-20 mm, with scar of bracteole at
or somewhat above the middle.

Field-notes — Stem without buttresses; bark smooth; twigs light green, pale brown
mottled. Lower leaf surface with golden indumentum. Fruits yellowish green covered
with golden hairs.

Distribution — Malesia: Borneo (Sarawak: Ist Div.; Brunei; C & NE Kalimantan).

Habitat & Ecology — Forest; at foot of limestone mountain; 50-350 m altitude;
fr. Jan.

Note — Knema sericea is related to K. retusa, K. ashtonii (var. cinnamomea), and
K. elmeri, with which it has the dense silky indumentum on the lower leaf surface in
common. Knema elmeri has similar, though smaller male flowers, also with a convex or
mammillate staminal disc, and differs in the less stout habit of the twigs, with flaking
bark. Knema retusa (Peninsular Malaysia) differs in its stout obtuse leaves and broad,
subglobose or ovoid, + apiculate fruits, and the pedunculate inflorescences. The short
indumentum on the lower leaf surface in K. sericea is more fluffy than that of K. ashtonii
var. cinnamomea, for which it can easily be mistaken, but which has a quite different
androecium.

64. Knema steenisii W. J. de Wilde
Knema steenisii W.J. de Wilde, Blumea 27 (1981) 224, f. 1. — Type: Schmutz 4162, W Flores.

Tree. Twigs 1-2(—3) mm diameter, with stellate scale-like and dendroid hairs 0.1—0.2
mm long, glabrescent; bark smooth or finely striate, not tending to crack. Leaves membra-
nous or thinly chartaceous, (oblong-)lanceolate, 7-19 by 2—6 cm, base attenuate or nar-
rowly rounded, apex acute(-acuminate); drying olivaceous or brown above; lower sur-
face with subpersistent mixed scale-like and dendroid hairs 0.1 mm long or less, and
with few emergent ones up to 0.4 mm; dots absent; midrib slightly raised; nerves 15—25
pairs, little raised; venation + distinct; petiole 10-15 by 1—1.5 mm, late glabrescent. /n-
florescences: sessile, brachyblast simple, 1-4 mm long; in male and female 1—5-flow-
ered; flowers with rusty hairs 0.1—0.2 mm; perianth 3-lobed, pinkish(?) inside. Male
flowers: pedicel 2-3 mm, bracteole 0.5 mm long, about median, caducous; buds obovoid
or pear-shaped, at base tapering, apex + truncate, 2.8-3.5 by 2—2.2 mm, cleft c. 5/6,
lobes 0.3—0.4 mm thick, inside at apex knobby; staminal disc circular, + flat, 0.8 mm

Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Knema) 343

diameter; anthers 4 or 5, just stiped, suberect, 0.4(—0.5) mm long; androphore slender,
slightly tapering, 1.1—-1.5 mm long. Female flowers: pedicel 0.5—1.5 mm long; buds
ovoid-oblong, 5—6 by 1.5—2.5 mm, cleft slightly over 1/2, lobes 0.5 mm thick; ovary
ovoid-ellipsoid, 2 mm long, with dense hairs 0.2(—0.3) mm, style glabrous, | mm long,
stigma 1—1.5 mm long, of 2 erect lobes, |—1.5 mm long, each entire or faintly 2-lobulate
at apex. Fruits 1(—3?) per infructescence; ellipsoid-oblong, apex subacute, 2.2-2.4 by
1 cm, with brown-grey hairs 0.2—0.3(—0.5) mm long, late glabrescent; dry pericarp 0.5
mm thick; fruiting pedicel 1—-1.5 mm long. — Fig. 56.

Field-note — Ripe fruits yellow.

Distribution — Malesia: Lesser Sunda Islands (W Flores).

Habitat & Ecology — Lower montane forest; 780—900 m altitude.

Notes — 1. Knema steenisii is distinguishable by small flowers, with an unique struc-
ture: 4 or 5 anthers, androphore slender, much longer than the diameter of the staminal
disc, perianth cleft nearly to the base.

2. When sterile the species may be confused notably with K. cinerea and K. stenocarpa.
Both these species (East Malesia) have globose male buds, and differ vegetatively in the
indumentum on the lower leaf surface; in K. cinerea this consists of uniformly sized
smaller stellate scale-like hairs, in K. stenocarpa of more woolly mixed stellate and
dendroid hairs, not scale-like; in both species pistil and fruits are also different.

65. Knema stellata Merr.

Knema stellata Merr., Philipp. J. Sci., Bot. 11 (1916) 182. — Type: BS 24276 (Ramos), Philippines,
Samar I.
For more references and synonyms see the subspecies.

Tree 5-35 m. Jwigs 1.5—4 mm diameter, at first with rusty hairs 0.1—1] mm long,
glabrescent; bark coarsely striate or grooved, not tending to crack or flake. Leaves mem-
branous or chartaceous; (ovate-)elliptic or lanceolate, 6—20(—27) by 1.5—7 cm, base slight-
ly rounded or cuneate-attenuate, the apex acute or acuminate; drying (greenish) brown
above, early or late glabrescent; lower surface greyish, at first with stellate- and stellate-
dendroid hairs of mixed sizes, 0.2—0.7 mm long, late glabrescent, the dendroid hairs
sometimes more early shed and leaving distinct hair scars; dots present, especially on
the veins; midrib + flat or raised above; nerves (8—)15—24 pairs, + raised above; venation
prominent, distinct; petiole 7-13 by 1.5—2 mm. Jnflorescences: peduncle up to 2 mm
long, brachyblast simple, 2-4 mm diameter; in male 5—10-flowered, female 2—6-flow-
ered; flowers with persistent hairs 0.2—-0.3 mm long, mixed with emergent hairs 0.5 mm
long; perianth 3-lobed, reddish (?) inside. Male flowers: pedicel (subsp. cryptocaryoides)
5—9 mm long, bracteole caducous, apical; buds subglobose, 3 by 3—-3.5 mm, cleft 3/4—
4/5, lobes 0.5—0.7(—1) mm thick; the staminal disc flat, circular, 1.5—2 mm diameter;

Fig. 56. Knema steenisii W.J. de Wilde. a. Male flowering twig; b. opened male bud, showing the
androecium; c. mature androecium; d. female inflorescence; e. female flower, opened, showing the
pistil; f. habit of fruiting twig; g. detail of lower leaf surface, showing minute stellate hairs of mixed
sizes [a—c: Schmutz 4162; d, e: Schmutz 4164, f, g: Schmutz 4420). — Scale bar for a = 1 cm; for
b, e = 1.3 mm; for c = 0.4 mm; for d = 3.3 mm; for f = 2 cm; for g = 0.85 mm.

344 Flora Malesiana, Ser. I, Vol. 14 (2000)

anthers 11, just stiped, 0.4-0.5 mm long, horizontal, spaced; androphore 0.5—1 mm long.
Female flowers: pedicel 2-5 mm long, bracteole median to apical; buds obovoid-ob-
long, 5 by 3 mm, cleft c. 1/2, lobes 1 mm thick; ovary ovoid, 2 by 1.5 mm, stigma
subsessile, 2-lobed and each lobe again + 2-lobulate. Fruits 1-5 per infructescence,
subglobose or broadly obovoid, base rounded or contracted into a short stipe (pseudostalk)
up to 2 mm, 1.6—2.4 by 1.61.8 cm, with hairs either 0.1 mm or 0.5—1 mm long; dry
pericarp 1—1.5 mm thick; fruiting pedicel 3-10 mm long.

Distribution — Malesia: Philippines and N & C Sulawesi.

Note — A variable species, divided into three closely related subspecies, each with a
restricted area.

KEY TO THE SUBSPECIES

la. Hairs on fruits 0.5—1 mm long. Fruiting pedicel 6-10 mm long. — Philippines (Samar)
ee a: ASB Meare i en ae foe eee eae a. subsp. stellata

b. Hairs on fruits 0.1—0.2 mm long. Fruiting pedicel 3-10 mm long ............ ms
2a. Male pedicel S—9 mm long; buds subglobose, 3 by 3-3.5 mm; anthers 11. — Philip-
pines (Mindanao, Sibuyan, Luzon) ................ b. subsp. cryptocaryoides

b. Male pedicel 1 mm long; buds + broadly obovoid, 2.5(—3) by 2(—2.5) mm; anthers 5.
= AINE SLIME SIME < @, cles oe) eas ey ren ceo oer thar c. subsp. minahassae

a. subsp. stellata

Knema stellata Merr., Philipp. J. Sci., Bot. 11 (1916) 182; Enum. Philipp. Flow. Pl. 2 (1923) 184;
W.J. de Wilde, Blumea 25 (1979) 471; 27 (1981) 234.

Twigs 2-4 mm diameter, with woolly hairs 0.5—0.7 mm long. Leaves chartaceous,
(oblong-)lanceolate, 10-24 by 2-6 cm, on lower surface with mixed coarse sessile-stellate
and stellate-dendroid hairs, persistent or late glabrescent. Nerves 18—24 pairs. Female
inflorescences to 1 mm pedunculate, wart-like. Flowers not known. Fruits subglobose
or broadly ellipsoid, 1.8—2.4 by 1.8—2 cm, with coarse or woolly hairs 0.5—1 mm long;
fruiting pedicel 6-10 mm long, the bracteole scar somewhat above the middle.

Distribution — Malesia: Philippines (Samar).

Habitat & Ecology — Forest at low altitudes; fr. Feb.—May.

b. subsp. eryptocaryoides (Elmer) W. J. de Wilde

Knema stellata Merr. subsp. cryptocaryoides (Elmer) W.J. de Wilde, Blumea 25 (1979) 471. —
Gymnacranthera cryptocaryoides Elmer, Leafl. Philipp. Bot. 3 (1911) 1060; J. Sinclair, Gard.
Bull. Sing. 16 (1958) 115. — Type: Elmer 12262, Philippines, Sibuyan I.

Knema kunstleri (King) Warb. var. surigaoensis J. Sinclair, Gard. Bull. Sing. 18 (1961) 238, f. 14.
— Type: Wenzel 2680, Philippines, Mindanao.

Twigs 1.5—2.5 mm diameter, with scurfy hairs 0.1—0.3 mm long. Leaves membranous
or chartaceous, ovate-oblong to oblong-lanceolate, 10-17 by 2.5—6 cm, on lower sur-
face with subpersistent sparse sessile-stellate hairs 0.2-0.3 mm, mixed with early shed
longer hairs leaving hair scars, the hairs on and near midrib and nerves longer persistent.
Nerves 12-22 pairs. Inflorescences sessile to 2 mm pedunculate. Male flowers with woolly
hairs 0.2 mm long, mixed with emergent hairs 0.5 mm long. Female flowers not seen.

De Wilde — Myristicaceae (Knema) 345

Fruits subglobose or broadly ellipsoid, 1.5—1.8(—2.5, according to Elmer) by 1.4—1.7
cm, with scurfy hairs 0.1 mm long; fruiting pedicel 5-10 mm long, the bracteole scar at
or below the middle.

Distribution — Malesia: Philippines (Luzon, Sibuyan, Mindanao).

Habitat & Ecology — Forest on fertile soil; c. 250 m altitude; fl. Sept—Oct; fr. Feb.,
Apr.

c. subsp. minahassae (Warb.) W.J. de Wilde

Knema stellata Merr. subsp. minahassae (Warb.) W.J. de Wilde, Blumea 25 (1979) 472; 27 (1981)
233. — Knema laurina (Blume) Warb. var. minahassae Warb., Mon. Myrist. (1897) 620. —
Syntypes: Koorders 18167, 18169, N Sulawesi.

Twigs 1.5—3 mm diameter, with short-scurfy or woolly hairs 0.2—1 mm long. Leaves
membranous or chartaceous, elliptic to oblong-lanceolate, 6—19(—27) by 2—7 cm, on
lower surface at first with mixed scale-like stellate and dendroid hairs, late glabrescent;
nerves 8-18 pairs. Inflorescences: peduncle to 2 mm long. Male flowers: pedicel 1—1.2
mm long, bracteole late caducous, apical; buds subglobose to broadly obovoid, 2.5(—3)
by 2(-2.5) mm, cleft c. 4/5, lobes 0.2—0.3 mm thick; staminal disc + flat, subcircular,
0.8 mm diameter; anthers 5, half-erect, short-stiped, 0.3—0.4 mm long; androphore 0.6—
0.7 mm long. Female flowers: pedicel 2 mm long, bracteole subapical. Fruits obovoid-
ellipsoid, 1.8—2.5 by 1.5—1.7 cm, with scurfy scale-like hairs 0.1—0.2 mm long; fruiting
pedicel 3—6 mm long.

Field-note — Perianth yellow, with a red centre within.

Distribution — Malesia: C & N Sulawesi (Minahassa).

Habitat & Ecology — Montane forest, forest on conglomeratic rock, not on ultrabasic;
600-1000 m altitude; fl. July; fr. Mar.

Note — Knema stellata subsp. minahassae is reminiscent of K. celebica (both with
dotted leaves) in the inflorescences and the male flowers, but they differ in leaf shape
and indumentum of twigs and leaves. Vegetatively subsp. cryptocaryoides can hardly be
distinguished, but both subspecies differ considerably in the male flowers.

66. Knema stenocarpa Warb.

Knema stenocarpa Warb., Mon. Myrist. (1897) 577, p.p., only the fruiting collection, not the male
specimens; Elmer, Leafl. Philipp. Bot. 3 (1911) 1064; Merr., Enum. Philipp. Flow. Pl. 2 (1923)
184; W.J. de Wilde, Blumea 25 (1979) 436. — Myristica stenocarpa (Warb.) Boerl., Handl. Fl.
Ned. Ind. 3, 1 (1900) 91, nom. alt. inval. — Syntypes: Warburg 13303, 13303a, Philippines,
S Mindanao.

Tree or shrub 3-10 m. Twigs 1—1.5(—2) mm diameter, at first with woolly yellowish to
rusty hairs 0.5 mm long, glabrescent; bark smooth or finely striate, not tending to crack
or flake. Leaves herbaceous or chartaceous, ovate to obovate, or oblong(-lanceolate),
occasionally faintly lobed in the upper part, base attenuate to subobtuse, apex acute
(-acuminate); drying greenish above, lower surface greyish, at first with soft hairs 0.2
mm long, late glabrescent; dots absent; midrib + raised above; nerves 10—20 pairs,
+ raised above, often brownish; venation distinct; petiole 5—15 by 0.7—1.5 mm. Jnflores-
cences: peduncle up to 1 mm long, brachyblast usually simple, slender, to 10 mm long;
in male 5—25-flowered, in female 2—6-flowered; flowers with greyish to rusty hairs 0.2—

346 Flora Malesiana, Ser. I, Vol. 14 (2000)

0.4 mm long; perianth 3-lobed, reddish (?) inside. Male flowers: pedicel 2—4.5 mm long,
bracteole late caducous, median to apical; buds globose, 2—2.5 by 2—2.5(-3?) mm, cleft
over 2/3, lobes 0.3—0.5 mm thick; staminal disc flat or occasionally low-mammillate,
circular to faintly angled 1.2-1.5 mm diameter; anthers 6-9, just sessile or distinctly
stiped, 0.3—0.5 mm long, + horizontal, not touching; androphore slender, 0.7—1 mm
long. Female flowers: pedicel 0.5—2 mm long, bracteole about median; buds + ellipsoid,
4 mm long, cleft 1/2—2/3, lobes 0.4 mm thick; ovary ovoid, 1.5 by 1.2-1.5 mm; stigma
subsessile, (1-) or 2-lobed and each lobe again 3—5-lobulate. Fruits | or 2 per infructes-
cence, ellipsoid, apex subacute, 1.3—1.5 by 0.8-0.9 cm, with scurfy hairs 0.1 mm long or
less; dry pericarp | mm thick; fruiting pedicel |-3 mm long.

Distribution — Malesia: Philippines (Sulu Is.; Mindanao: mainly Zamboanga Prov.).

Habitat & Ecology — Primary and degraded forest; 300-500 m altitude; fl. & fr.
throughout the year.

Note — Knema stenocarpa is closely related to K. glomerata, which differs in its
stouter habit: thicker twigs, larger leaves with more nerves, larger male and female flowers,
and different fruits, subglobose, 1.5—2.5 cm diameter. The leaves in K. stenocarpa are
densely fine grey woolly beneath, and later glabrescent as compared to K. glomerata.

67. Knema stenophylla (Warb.) J. Sinclair

Knema stenophylla (Warb.) J. Sinclair, Gard. Bull. Sing. 16 (1958) 300, f. 11. — Gymnacranthera
stenophylla Warb., Mon. Myrist. (1897) 364, t. 20. — Type: Kehding 150, Peninsular Malaysia.
For more references and synonyms see the subspecies.

Tree 5-25 m. Twigs sometimes faintly angled, 0.5-2(-3) mm diameter, at first with
minute hairs 0.1 mm long or less, sometimes mixed with longer hairs 0.2—0.3 mm,
glabrescent; bark finely striate, not tending to crack or flake. Leaves membranous or
chartaceous; elliptic to lanceolate, (3-)5—18 by 1-5 cm, apex acute-acuminate, base at-
tenuate or cuneate, drying greenish to brown above; lower surface greyish, with sub-
persistent, scattered, mainly stellate scale-like hairs of + equal size, 0.1 mm long or less;
dots absent: midrib flat to raised above; nerves 10-21 pairs, slender, sunken or but slight-
ly raised, indistinct above; venation either raised and distinct, or sunken and indistinct;
petiole 6-16 by 0.5—2 mm. Inflorescences: peduncle up to | mm long, brachyblast sim-
ple or 2- or 3-fid, 1-3(—7) mm long; in male (1—)3—10-flowered, female 1—6-flowered;
flowers with greyish to rusty hairs 0.1 mm long, sometimes with emergent hairs up to
0.3 mm; perianth 3-lobed, creamy or yellowish inside (always?). Male flowers: pedicel
5-10 mm long, bracteole persistent or caducous, about median to nearly apical; buds
subglobose or broadly obovoid, (1.5—)2—3(—4?) mm diameter, cleft 3/4—2/3, lobes 0.5—
0.8 mm thick; staminal disc + flat, circular or subtriangular, 1-2.2 mm diameter; anthers
6-11, just to distinctly stiped, (almost) horizontal, 0.3 —0.5 mm long, spaced; androphore
1 mm long. Female flowers: pedicel 3-12 mm long, bracteole about median; buds el-
lipsoid or obovoid, 3.5—5 mm long, cleft c. 1/2; ovary ovoid, 1.5—2 by 1—1.5 mm, style
0.5 mm long, stigma 2-lobed and each lobe again (2—)3—10-lobulate or serrate. Fruits
1-5 per infructescence, subglobose or obovoid, apex rounded to subacute, base rounded,
sometimes + attenuate, (1.2—)1.5 by 0.7—1 cm, with rusty scale-like hairs 0.1 mm long or
less; dry pericarp | mm thick; fruiting pedicel 4-20 mm long.

De Wilde — Myristicaceae (Knema) 347

Field-notes — Bark smooth or rough with some pustules or flakes when old, not
striate or furrowed; slash bark reddish, granular; slash wood pale. Leaves glossy dark
green above, glaucous with yellowish green midrib beneath. Flowers creamy or yellow
inside, fragrant. Fruits greenish brown, yellow, apricot, or orange.

Distribution — Malesia: Sumatra, Peninsular Malaysia (subsp. stenophylla), Borneo
(subsp. longipedicellata).

Habitat & Ecology — Primary and degraded forest; often on hill sides, crests, ridges,
also in swamp forest, periodically inundated forest, pole forest and kerangas; on sand-
stone, loam, sandy and clayey soil, shales; O—800(—1200) m altitude; fl. & fr. throughout
the year, in Borneo fl. mainly Mar.—July; fr. mainly July—Dec., in Sumatra fl. Apr.

Notes — 1. Knema stenophylla resembles K. cinerea, K. glauca, K. hirtella, K. malaya-
na, K. patentinervia, and K. subhirtella; for differences see the key to the species.

2. The variability in K. stenophylla abounds in the venation on the upper leaf surface,
which can be either distinct, indistinct, or invisible; the length of the male and female
pedicels, the diameter of the male buds, and the number of anthers, varying from 6 to 11.
These characters show some overlap with K. malayana, which differs in the generally
thinner and broader leaves with a very distinct venation on the upper leaf surface, gener-
ally larger male flowers with longer pedicels, and the indumentum of the fruits which is
more farinose. Knema patentinervia has larger male flowers, K. subhirtella has larger
fruits.

KEY TO THE SUBSPECIES

la. Male pedicel 5—6 mm long. Fruiting pedicel 4-8 mm long. — Sumatra, Peninsular
aatawlawe de ifn Wah. e 2 esrtyad acheter SeaeRe eee a. subsp. stenophylla

b. Male pedicel 7-10 mm long. Fruiting pedicel 10-20 mm long. — Borneo........
MP Say ee yay eelek . aul ls | Le ase ee b. subsp. longipedicellata

a. subsp. stenophylla

Knema stenophylla (Warb.) J. Sinclair, Gard. Bull. Sing. 16 (1958) 300, f. 11; 18 (1961) 276; W.J.
de Wilde, Blumea 25 (1979) 457, p.p., excl. stouter forms from Borneo, now in K. subhirtella;
32 (1987) 129; Tree Fl. Sabah & Sarawak 3 (2000) 442. — Gymnacranthera stenophylla Warb.,
Mon. Myrist. (1897) 364, t. 20. — Myristica stenophylla (Warb.) Boerl., Handl. Fl. Ned. Ind. 3,
1 (1900) 88.

Myristica geminata auct. non Miq.: King, Ann. Roy. Bot. Gard. Calc. 8 (1891) 322, pl. 160, p.p.

Indumentum of twig apex and flowers + reddish brown. Leaf blades chartaceous,
usually + glossy above, with very faint nerves and reticulation; nerves faint below. Male
pedicels 5—6 mm long. Stalk of fruits 4-8 mm long.

Distribution — Malesia: Sumatra, Peninsular Malaysia.

Habitat & Ecology — Dryland or mossy forest.

b. subsp. longipedicellata (J. Sinclair) W.J. de Wilde

Knema stenophylla (Warb.) J. Sinclair subsp. longipedicellata (J. Sinclair) W.J. de Wilde, Blumea
32 (1987) 130; Tree Fl. Sabah & Sarawak 3 (2000) 442. — Knema cinerea (Poir.) Warb. var.
patentinervia J. Sinclair forma longipedicellata J. Sinclair, Gard. Bull. Sing. 18 (1961) 182,
f. SA-E. — Type: Clemens 20345, Borneo, Sarawak.

348 Flora Malesiana, Ser. I, Vol. 14 (2000)

Indumentum of twig apex and flowers greyish brown. Leaf blades membranous or
chartaceous, dull above, with nerves and reticulation clearly discernible; nerves distinct
below. Male pedicels 7-10 mm long. Stalk of fruits 10-20 mm long

Distribution — Malesia: Borneo (throughout).

Habitat & Ecology — Dryland forest on poor soils, kerangas.

68. Knema stylosa (W.J. de Wilde) W.J. de Wilde

Knema stylosa (W.J. de Wilde) W. J. de Wilde, Blumea 32 (1987) 119; Tree FI. Sabah & Sarawak 3
(2000) 443. — Knema hirtella W.J. de Wilde var. stylosa W.J. de Wilde, Blumea 25 (1979) 460.
— Type: SAN 30179 (Mikil), Sabah.

Tree 8—20(-25) m. Twigs 2-3 mm diameter, at first with pale brown or grey-brown
hairs 0.1 mm long or less, glabrescent; bark striate, not tending to crack or flake. Leaves
thinly chartaceous, (elliptic-)oblong, 12-24 by 3.5—7 cm, base (subcordate or) rounded
to short-attenuate, apex acute-acuminate; drying olivaceous to brown above, lower sur-
face glabrescent or with subpersistent sparse hairs less than 0.1 mm long, mixed with
scars of coarser caducous hairs 0.2 mm; dots absent; midrib convex, but (partly) sunken
under surface level; nerves 15—20 pairs, flat or sunken; venation usually distinct; petiole
10-20 by 2-3 mm, late glabrescent. /nflorescences: sessile, brachyblast wart-like, sim-
ple or forked, to 4 mm long; in male and female 3—6(—10)-flowered; flowers with dense
(pale) brown hairs 0.1 mm long; perianth 3-lobed, greenish (?) inside. Male flowers:
pedicel 4—5 mm long, bracteole apical, persistent; buds obovoid, 3.5—4 by 3-3.5 mm,
cleft c. 2/3, lobes 0.7 mm thick; staminal disc + flat, blunt-triangular, 1.4 mm diameter;
androphore tapering, 1.2 mm long, glabrous; anthers 11, half-sessile, suberect, 0.5 mm
long. Female flowers: pedicel 3-5 mm long, bracteole caducous, median to apical; buds
+ obovoid-ellipsoid, 4.5 by 3.5 mm, cleft c. 1/2, lobes 0.7(—1) mm thick; ovary ovoid,
1.5 by 1 mm, densely minutely pubescent, style (0.5—)1 mm long, stigma broadly
2-lobed and each lobe again 6—12-lobulate. Fruits solitary or 2 or 3 per infructescence,
ovoid, 2.5-3.5 by 1.8—2.5 cm, with dense or sparse pale brown hairs 0.1 mm or less,
sometimes glabrescent, style persistent or leaving a scar; dry pericarp 1.5—2.5 mm thick;
fruiting pedicel 3—6 mm long.

Field-notes — Bark smooth, slash pinkish with (dark) red exudate, sapwood whitish.
Flowers (pale) chocolate-brown, (greenish inside?). Fruits green or orange.

Distribution — Malesia: Borneo (NE Sarawak: 5th Div.; Sabah).

Habitat & Ecology — Primary forest, hill sides, often on blackish soil, sandstone or
sandstone-derived soils; 0-800 m altitude, fl. June; fr. throughout the year.

Note — Knema stylosa, with obovoid male buds, belongs in the group of K. obovoidea,
in the vicinity of species like K. pulchra and K. piriformis, hence species with male buds
narrowly pear-shaped, cleft c. 2/3 only. In habit it may be confused with K. glauca, a
species also with a weak indumentum on the lower leaf surface, distinct in globose male
buds, stiped anthers, and early glabrescent fruits. An additional character of K. stylosa is
the largely sunken midrib into a groove above. Knema pulchra is much stouter, always
with a cordate or broadly rounded leaf base, larger male buds, more anthers, and larger
fruits; K. piriformis is a mountain species, with more narrowly pear-shaped male buds,
stiped anthers, the indumentum on the lower leaf surface more distinct.

De Wilde — Myristicaceae (Knema) 349

69. Knema subhirtella W. J. de Wilde

Knema subhirtella W.J. de Wilde, Blumea 32 (1987) 131; Tree Fl. Sabah & Sarawak 3 (2000) 443.
— Type: S 43/32 (Yii et al.), Sarawak.

Tree 5—25 m. Twigs sometimes blunt-triangular, |.5—2(—2.5) mm diameter, with grey-
brown or pale rusty hairs 0.1—0.3 mm long, glabrescent; bark faintly or finely striate, not
flaking. Leaves membranous, elliptic-oblong or lanceolate, 9-20 by 1.5—5.5 cm, base
short- to long-cuneate, apex acute-acuminate, drying olivaceous to brown above, lower
surface with persistent sparse hairs of mixed sizes, 0.1—0.2(—0.3) mm; dots absent; the
midrib raised above; nerves 15-25 pairs, + flat or raised above, venation coarse, distinct;
petiole + late glabrescent, 8-12 by 1-2 mm. Inflorescences: peduncle up to | mm long,
brachyblast simple or forked, to 4 mm long; in male 4—7-flowered, female 1—4-flow-
ered: flowers with dense hairs 0.1—0.3 mm long; perianth 3-lobed, creamy greenish in-
side. Male flowers: pedicel 2-5 mm long, bracteole late caducous, 0.5—1.5 mm below
the apex; buds broadly obovoid, 2.5—3.5(—4) by 2.5—3.5 mm, cleft (1/2—)2/3—3/4, lobes
0.5 mm thick; staminal disc circular, flat or somewhat convex, 1.2—1.5(—2) mm diam-
eter, anthers 7—15, short-stiped, 0.3—0.4 mm long, horizontal (opening downwards), not
touching; androphore little tapering, 1-1.5 mm long, glabrous or with some minute stellate
hairs towards the base. Female flowers: pedicel 4—7(—10) mm, bracteole + median; buds
+ obovoid, 5 by 3 mm, cleft c. 1/2, lobes 0.5 mm thick; ovary 2.5 by 1.5 mm, with dense
hairs 0.1 mm long or less, stigma subsessile, 1.5 mm high, 2-lobed, each lobe + 2- or 3-
lobulate. Fruits solitary or 2 per infructescence, broadly ellipsoid, 2—2.7 by 1.5—2 cm,
with dense hairs 0.1 mm long, dry pericarp 1.5 mm thick; fruiting pedicel 4-10 mm long.

Field-notes — Understorey tree; bark usually chocolate-brown, + smooth or dippled,
not furrowed; outer bark brittle, inner bark pink-cream, fibrous; sapwood cream; exu-
date pink to blood-red. Perianth outside yellow-brown, greenish cream inside. Fruits
yellow-green, apricot, or orange; aril red. The wood is used as firewood.

Distribution — Malesia: Borneo (Sarawak, W Sabah).

Habitat & Ecology — Mixed dipterocarp forest; hill slopes, ridges; sandy clay and
loam, basalt rock, shales; 0O—800(—1200) m altitude; fl. Mar—May(—Aug.); fr. mainly
Sept, Wet.

Note — Knema subhirtella is related to K. mogeana and K. hirtella, all three with a
similar indumentum on the lower leaf surface. In general habit the species may be con-
fused with K. stenophylla, K. glaucescens, and K. piriformis. Knema mogeana differs in
its smaller male buds, and distinctly smaller fruits. Knema hirtella has stouter twigs,
larger male buds, the androphore always with hairs towards the base, and somewhat
larger fruits. In K. subhirtella the androphore is either glabrous or with a few hairs to-
wards the base only. Knema stenophylla (in Borneo subsp. longipedicellata) differs in
the subglobose (not obovoid) male buds, cleft almost to the base, and an inconspicuous
indumentum on the lower leaf surface, consisting of scattered stellate hairs of only 0.1
mm diameter. Knema glaucescens, also with a globose male buds, has much denser
hairs on the lower leaf surface. Knema piriformis (known from a limited number of
specimens from montane localities) generally has more elongate male buds, with the
androphore longer than the diameter of the disc. Through its rather variable, + pear-
shaped male buds, K. subhirtella obscures the demarcation of the group of species around
K. glauca, in which it obviously has a marginal position. On account of its ill-defined
bud shape, it keyed out twice in the general key to male flowering specimens.

350 Flora Malesiana, Ser. I, Vol. 14 (2000)

70. Knema sumatrana (Blume) W.J. de Wilde

Knema sumatrana (Blume) W.J. de Wilde, Blumea 25 (1979) 454. — Myristica sumatrana Blume,
Rumphia | (1835) 187. — Myristica glauca Blume var. sumatrana (Blume) Migq., PI. Jungh.
(1852) 171. — Myristica corticosa (Lour.) Hook. f. & Thomson var. sumatrana (Blume) Miq.,
FI. Ind. Bat., Suppl. 1 (1861) 384. — Knema glauca (Blume) Warb. var. sumatrana (Blume)
Warb., Mon. Myrist. (1897) 597. — Knema cinerea (Poir.) Warb. var. sumatrana (Blume) J. Sin-
clair, Gard. Bull. Sing. 18 (1961) 185, p.p., for the basionym only. — Type: Korthals s.n. (male
fl., lecto), Sumatra.

Myristica wrayi King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 315, pl. 151. — Knema wrayi (King)
Warb., Mon. Myrist. (1897) 572, t. 24; Gamble, Mat. Fl. Malay Penins. 5, 23 (1912) 243; Ridl.,
Fl. Malay Penins. 3 (1924) 70. — Syntypes: Wray s.n.; King’s coll. 5299, 8277, 10444 (lecto),
Peninsular Malaysia.

Knema glaucescens auct. non Jack: J. Sinclair, Gard. Bull. Sing. 16 (1958) 304 (var. glaucescens,
p-p.).

Tree 10-20 m. Twigs 2-3 mm diameter, at first with minute scale-like hairs 0.1 mm
long or less, very early glabrescent; bark finely striate, not tending to crack or flake.
Leaves membranous or chartaceous, elliptic to oblong-lanceolate, 12-30 by 4.5-12 cm,
apex obtuse to acute-acuminate, base rounded, subtruncate, or short-attenuate, drying
greenish to brown above; lower surface greyish, at first with inconspicuous scale-like
hairs 0.1 mm long or less, early glabrescent, sometimes with minute lanose hairs at
blade edge near the base, soon glabrescent; dots absent; midrib + flat or raised above,
nerves 17-22 pairs, + flat above; venation (areoles 1 mm diameter or more) distinct;
petiole 12-16 by 1.5-3 mm. Inflorescences: sessile, brachyblast simple or forked, to
8 mm long; in male 10—20-flowered, female 1—5(?)-flowered; flowers with greyish brown
hairs less than 0.1 mm long; perianth 3- (or 4-)lobed, red inside. Male flowers: pedicel
10-11 mm long, bracteole 1 mm long, at or above median, persistent or late caducous;
buds subglobose, 3-4 mm diameter, cleft c. 4/5, lobes 0.5—0.8 mm thick; staminal disc
slightly convex, finely papillate, circular, 2—2.5 mm diameter; anthers 12-18, distinctly
stiped, spaced, horizontal, 0.3—0.4 mm long (connective narrow above, thecae opening
+ laterally); androphore 1—1.2 mm long. Female flower (from aged flowers in Korthals
s.n.): pedicel 2~4 mm long, bracteole median to subapical, subpersistent; buds obovoid-
ellipsoid, 4 by 2.5 mm, cleft slightly over 1/2; ovary ovoid, 1.5 by | mm, stigma sessile,
2-lobed and each lobe again 3- or 4-lobulate. Fruits 1-3 per infructescence, ovoid-ellip-
soid or ellipsoid-oblong, apex and base subacute, 3-4 by 1.4—2 cm, at first with minute
indumentum, early glabrescent, leaving a finely granulate surface; dry pericarp 3 mm
thick; fruiting pedicel 3-6 mm long. — Photo 7.

Field-notes — Perianth bright red inside, anthers creamy white, disc dirty red.

Distribution — Peninsular Thailand; in Malesia: Sumatra (incl. Simeulué I.) and Pe-
ninsular Malaysia (Perak, Kelantan).

Habitat & Ecology — Rain forest; along rivers, Saraca streams, hill slopes; 0—1200
m altitude; fl. & fr. throughout the year.

Note — Knema sumatrana is closely related to K. glauca; several of the stouter speci-
mens included in K. glauca are difficult to separate from K. sumatrana. Possibly these
specimens are hybrids, with an intermediate habit. In most instances, however, K. glauca
can easily be distinguished by the smaller size of twigs and leaves, smaller male flowers
with fewer anthers (8-11, or in E Java up to 14), and by smaller fruits, 1.8—2.5(—4) cm

De Wilde — Myristicaceae (Knema) 351

long, as against 3—4 cm in K. sumatrana. Other characters of diagnostic value of
K. sumatrana are the early glabrescent twig apex and lower leaf surface, persistent brac-
teole, many (12-18) distinctly stiped anthers, somewhat convex, finely papillate staminal
disc, narrow connective, and almost laterally opening thecae.

71. Knema tomentella (Miq.) Warb.

Knema tomentella (Mig.) Warb., Mon. Myrist. (1897) 588, t. 25; J. Sinclair, Gard. Bull. Sing. 18
(1961) 277, f. 24, map 18, p.p., excl. syn. K. /aurina var. minahassae, K. stellata, K. alvarezit;
W.J. de Wilde, Blumea 25 (1979) 438; 41 (1996) 394. — Myristica corticosa (Lour.) Hook. f. &
Thomson var. tomentella Miq., Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 207. — Myristica tomentella
(Miq.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 91. — Type: de Fretes in Teijsmann s.n., Moluccas,
Ambon.

[Palala sexta Rumph., Herb. Amb. 2 (1750) 28.]

Myristica corticosa var. amplifolia Mig., Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 202. — Type: Riedel
in Beccari 7772, Sulawesi.

Myristica corticosa var. ceramensis Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 51.— Type: Tetjsmann
& de Vriese (12, 13), Seram.

Knema glauca (Blume) Warb. var. bancana Warb., Mon. Myrist. (1897) 597. — Myristica glauca
Blume var. bancana (Warb.) Boerl., Handl. Fl. Ned. Ind. 3, 1 (1900) 92. — Type: culta Hort.
Bog. (Beccari s.n.).

Tree 5—15(—30?) m. Twigs 2-3 mm diameter, at first with rusty hairs 0.3—0.6 mm
long, glabrescent; bark striate or in Sulawesi and Seram (partly) sometimes faintly
striate, not tending to crack or flake. Leaves chartaceous, elliptic, subobovate, or (ob)lance-
olate, 7-26 by 2—7.5 cm, apex acute(-acuminate), rarely subtruncate, base attenuate to
rounded, drying (greenish) brown above, lower surface grey-glaucous, at first with hairs
of mixed size, 0.3—0.6 mm long, largely glabrescent, with some indumentum remaining
on or near the midrib and nerves, finely papillate, with hair scars; dots absent; midrib
raised above; nerves 16-26 pairs, raised above; venation very distinct; petiole 10-20 by
1.5-3.5 mm. Inflorescences: sessile, brachyblast simple or bifurcate, to 10 mm long; in
male 2-15-flowered, female 1—5-flowered; flowers with rusty woolly hairs (0.2—)0.3-1
mm long; perianth 3-lobed, reddish (?) inside. Male flowers: pedicel 2-5 mm long, brac-
teole late caducous, (sub)apical; buds broadly obovoid or globose, 2.5—3 mm diameter,
cleft nearly to the base, lobes 0.5—0.7 mm thick; staminal disc flat, circular, 1-1.5(-2)
mm diameter; anthers 6—9(—11), stiped, horizontal or slightly oblique, 0.3—0.5 mm long,
spaced; androphore slender, 0.7-1 mm long. Female flowers: pedicel 24 mm long,
bracteole subapical; buds obovoid-oblong, 4.5—6 mm long, cleft c. 1/2, lobes 0.5 mm
thick; ovary ovoid, 1.5—2 mm long, stigma subsessile, 2-lobed and each lobe again 2—5-
lobulate. Fruits 1-3 per infructescence, obovoid, pear-shaped, or ellipsoid, apex obtuse
or shortly beaked, (1—)1.5—2.5 by (0.8—)1-1.8 cm, with harsh rusty hairs 0.5 mm long;
dry pericarp 1.5-2 mm thick; fruiting pedicel 2-5 mm long.

Field-notes — Bark smooth, brownish grey, with some dents, no flakes or fissures;
inner bark red; wood dirty white. Leaves glossy dark green with whitish midrib above,
glaucous with greenish yellow midrib beneath.

Distribution — Malesia: Philippines (Samar), Sulawesi (N, SW, & SE Peninsula),
Moluccas (Morotai, Halmahera, Bacan, Obi, Buru, Seram, Ambon), New Guinea (Bird’s
Head, one coll.).

352 Flora Malesiana, Ser. I, Vol. 14 (2000)

Habitat & Ecology — Primary and degraded forest; once from sandy loam soil; alti-
tude 0—1500 m; fl. June, Nov.—Feb.; fr. Apr., May, Aug.

Notes — 1. This is the only Knema occurring in New Guinea (and known only from
a single collection). Most specimens formerly recorded for the Philippines belong to
K. glomerata, a related species differing in the finer reticulation on the upper leaf sur-
face, bracteole median on the pedicel, and a + convex staminal disc. In the Philippines
K. tomentella is known only from Samar.

2. Specimens from Sulawesi may be reminiscent of K. celebica or K. glomerata.

72. Knema tridactyla Airy Shaw

Knema tridactyla Airy Shaw, Kew Bull. 10, 1939 (1940) 543; W.J. de Wilde, Blumea 25 (1979)
434, f. 13; 41 (1996) 395; 43 (1998) 251; Tree Fl. Sabah & Sarawak 3 (2000) 444. — Type:
Richards 1220, Sarawak.

Treelet 2-10 m. Twigs sometimes faintly 2- or 3-angular, 1.5—4 mm diameter, at first
with rusty woolly hairs 0.5—1(—1.5) mm long (subsp. tridactyla), + late glabrescent, or
with inconspicuous scale-like hairs 0.1 mm long or less (subsp. sublaevis), early glabres-
cent; bark (finely) striate, not tending to crack or flake. Leaves chartaceous or thinly
coriaceous, oblong or lanceolate, 7-28 by 2—5.5(—7.5) cm, base cuneate to rounded, apex
acute(-acuminate), drying olivaceous or brown above, often glossy and with minute hair
scars (lens!); lower surface either with subpersistent greyish stellate hairs 0.3-0.5 mm
long, intermixed with stronger dendroid hairs (subsp. tridactyla), or at first with sparse
scale-like hairs 0.1 mm long or less, early glabrescent (subsp. sublaevis), surface grey-
ish, not or hardly papillate, with scars of shed coarser hairs; dots absent; midrib much
raised above (in subsp. tridactyla towards base with appressed scale-like hairs); nerves
10-23 pairs, flat or sunken above; venation very fine, distinct above; petiole + late
glabrescent, 5—15 by 1.5—3 mm. J/nflorescences: peduncle up to 3 mm long, brachyblast
simple or 2- or 3-fid, slender or rather thick (worm-like), to 17 mm long, 1—2(-—2.5) mm
thick: in male (2—)4—10-flowered, female 1—5-flowered; flowers with hairs either 0.3—
1(—1.5) mm long (subsp. tridactyla & pachydactyla) or (scale-like) 0.1 mm long (subsp.
sublaevis); perianth 3-lobed, pinkish (?) inside. Male flowers: pedicel 1.5—3.5 mm long,
bracteole subpersistent, at or above the middle; buds globose to broadly obovoid, 1.5—2
(—2.3) by 1.5—2.5 mm, cleft 3/4—4/5, lobes 0.2—-0.5 mm thick; staminal disc flat, circu-
lar or faintly triangular, 0.6—1.2 mm diameter; anthers 5—11, partly or completely ses-
sile, 0.2-0.3 mm long, half-erect, not or hardly touching; androphore 0.3—0.7 mm long.
Female flowers: pedicel 1.5 mm long, bracteole at or above the middle; buds more or
less obovoid, 2—2.5 by 1.5 mm, cleft c. 2/3, lobes 0.2—0.3 mm thick; ovary subglobose,
1 mm diameter, style including stigmas 1—1.5 mm long, the latter 2-lobed and each lobe
again + 2-lobulate. Fruits 1 (or 2) per infructescence, ellipsoid or (ob)ovoid-oblong,
1.4—2 by 0.8-1.6 cm, with hairs either 0.2—0.5 mm long (subsp. tridactyla), or scale-
like, 0.1 mm long (subsp. sublaevis); dry pericarp 1-1.5 mm thick; fruiting pedicel 2-3
mm long.

Field-note — Bark brown, smooth.

Distribution — Malesia: Borneo (Sarawak, Brunei, Sabah).

Habitat & Ecology — Primary lowland dipterocarp forest; pole forest; clay or yellow
sandy soil; once from steep ridge; 0-1000 m altitude; fl. Jan.—Dec.; fr. Oct., Dec., Feb.

De Wilde — Myristicaceae (Knema) 353

Note — Distinguishable by the small flowers. The inflorescences are often peduncu-
late, and, as in the type Richards 1220, typically 2- or 3-fid, the ramifications worm-like,
up to 17 mm long. In other specimens the inflorescences are not or hardly branched and
much shorter.

KEY TO THE SUBSPECIES

la. Indumentum of twig apices, flowers, and fruits with hairs 0.5—1 mm long. Anthers

fe 205s caus, Ne eee RM PAS Besoin nu ot SPs oc ORE Mage OM eee ee ey

b. Indumentum consisting of hairs 0.1 mm long. Male perianth 1.5—2 mm diam.; an-
LYSIS S01 Deena, eee ot iaarnts, Sacer cca saan Rar ee an d. subsp. sublaevis

2a. Leaves narrow, 7-12 by 1.5—2.5 cm, lateral nerves 8—12 pairs. Male perianth 1.5
HTNe Alem LS OFM. 3:5. a Noo eee Mets of foils bite © ook c. subsp. salicifolia
Paeeavesmarcen lateral nerves 12-23 per side 7. 22 Bs ans ee de ew ee 3

3a. Indumentum consisting of hairs 0.5—1 mm long. Branches of male inflorescences
(1—)1.5—2 mm thick; perianth 1.5—2 mm diam.; anthers 6-8 .. a. subsp. tridactyla

b. Indumentum consisting of hairs 1(—1.5) mm long. Branches of male inflorescences
2-2.5 mm thick; perianth 2—2.3 mm diam.; anthers 10 or 11 b. subsp. pachydactyla

a. subsp. tridactyla

Twigs at apex with distinct dendroid hairs 0.5(—1) mm long. Leaves beneath with
dense stellate hairs, intermixed with coarser dendroid hairs up to 0.5 mm long.
Indumentum of flowers and fruits consisting of mixed sessile stellate, and dendroid hairs
0.5—-1 mm long. Anthers 6-8. — Fig. 57.

Distribution — Malesia: Borneo (Sarawak: 4th Div.; W Sabah).

Notes — 1. The leaves have conspicuous scattered crateriform hair scars on the upper
surface (lens!), similar to those found e.g. in K. oblongata subsp. pedunculata, the latter
differing in shape and size of the male flowers. In the indumentum on the lower leaf
surface it is somewhat reminiscent of K. scortechinii and K. conferta.

2. The midrib is prominent above, usually covered with characteristic appressed scale-
like hairs towards the base.

b. subsp. pachydactyla W. J. de Wilde

Knema tridactyla Airy Shaw subsp. pachydactyla W.J. de Wilde, Blumea 43 (1998) 251: Tree FI.
Sabah & Sarawak 3 (2000) 446. — Type: SAN 134086 (Madani et al.), male fl., Sabah.

Tree c. 7 m. Twigs at apex, lower leaf surface, and flowers with hairs 1(—1.5) mm
long. Branches of male inflorescences 2(—2.5) mm thick. Male perianth 2—2.3 mm diam.;
anthers 10 or 11. Female flowers and fruits not known.

Distribution — Malesia: Borneo (N Sabah, Penampang Distr.); known only from the
type collection; see note.

Habitat & Ecology — On ridge; c. 400 m altitude; fl. Aug.

Note — The specimen on which the present subspecies has been based represents an
exceptionally stout plant. The distinction with subsp. tridactyla may prove to be prob-
lematic, but the type of K. tridactyla, with slenderly branched male inflorescences and
small flowers (perianth 1.5 mm diam., 6 or 7 anthers), is quite different.

Flora Malesiana, Ser. I, Vol. 14 (2000)

a)
foo)

De Wilde — Myristicaceae (Knema) S55)

c. subsp. salicifolia W. J. de Wilde

Knema tridactyla Airy Shaw subsp. salicifolia W.J. de Wilde, Blumea 43 (1998) 251; Tree Fl. Sabah
& Sarawak 3 (2000) 446. — Type: SAN A 4237 (Wood), N Sabah.

Tree c. 10 m. Twigs at apex, lower leaf surface and flowers with hairs 0.5(—1) mm long.
Leaves narrow, 7-11 by 1.5—2.5 cm. Branches of male inflorescences 1.5 mm thick.
Male perianth 1.5 mm diam.; anthers 6. Female flowers and fruits not known.
Distribution — Malesia: Borneo (N Sabah, Kota Belud Distr.); known only from the
type collection.
Habitat & Ecology — On hills; c. 650 m altitude; fl. Sept.

d. subsp. sublaevis W.J. de Wilde

Knema tridactyla Airy Shaw subsp. sublaevis W.J. de Wilde, Blumea 25 (1979) 434; 41 (1996)
395: 43 (1998) 251, f. 1d; Tree Fl. Sabah & Sarawak 3 (2000) 446. — Type: S 13590 (Ilias Paie),
Sarawak.

Twigs at apex seemingly glabrous, with scale-like hairs 0.1 mm long. Leaves beneath
with sparse minute scale-like hairs, mixed with scattered somewhat coarser scale-like
hairs. Indumentum of flowers and fruits thin, consisting of scale-like hairs 0.1 mm long.
Anthers 5 or 6. — Fig. 41d.

Distribution — Malesia: Borneo (Sarawak: Ist Div.).

Note — Some slender specimens from Brunei, differing in male flowers with 10-12
anthers (instead of 5 or 6), and in a more conspicuous indumentum of the twigs, have
been discussed under subsp. sublaevis (De Wilde 1996: 395).

73. Knema uliginosa J. Sinclair

Knema uliginosa J. Sinclair, Gard. Bull. Sing. 18 (1961) 281, f. 25; W.J. de Wilde, Blumea 25 (1979)
428: Tree Fl. Sabah & Sarawak 3 (2000) 447. — Type: Main 2098 (exp. Polak), W Kalimantan.

Shrub or tree to 10 m. Twigs sometimes bluntly 2- or 3-angular, 1.5—3(—4) mm diam-
eter, at first with dense hairs 0.5—1 mm long, glabrescent; bark grey-brown, smooth or
finely striate, not tending to crack or flake. Leaves thinly coriaceous, oblong or lanceo-
late, 9-35 by 2-7.5 cm, apex subacute or acute-acuminate, base rounded to attenuate,
drying olivaceous or brown above; lower surface at first with dense bright brown hairs
0.5-1 mm long, early glabrescent, greyish to reddish brown; dots absent; midrib raised
above; nerves 8-15 pairs, much raised above; venation raised, distinct; petiole 10 by
2-3 mm. Inflorescences: sessile, brachyblast simple or forked, to 5 mm long; in male

Fig. 57. Knema tridactyla Airy Shaw subsp. tridactyla. a. Branchlet with male inflorescences;
b. piece of branchlet with male inflorescence; c. male bud; d. opened male bud showing androecium;
e. androecium, somewhat lateral view; f. ditto, seen from beneath; g. sample of hairs of male peri-
anth, half-schematic; h. piece of branchlet with infructescence; note fruit with persistent style:
i. detail of lower leaf surface showing rather dense persistent indumentum of small stellate hairs
[a—g: S§ 21378 (Othman); h, i: S 27758 (Ingah Kudi)]. — Scale bar for a = 2 cm; for b = 2.8 mm;
for c, d = 1.4 mm; for e, f, i= 0.32 mm; for g = 0.16 mm; for h = 1 cm.

Flora Malesiana, Ser. I, Vol. 14 (2000)

356

aaa arrears

De Wilde — Myristicaceae (Knema) 557

5-20-flowered, female 1—5-flowered; flowers with rusty hairs 0.3-0.4 mm long; peri-
anth 3- or 4-lobed, reddish (?) inside. Male flowers: pedicel 5-10 mm long, bracteole
subpersistent or caducous, (sub)apical; buds depressed globose, faintly angled in cross
section, 3—4 by 4—5 mm, cleft c. 4/5, lobes 0.5 mm thick; staminal disc circular, 2—2.5
mm diameter, convex or low-mammillate; anthers 10—13, almost horizontal, 0.5 mm
long, just stiped, spaced; androphore 0.5—1 mm long. Female flowers: pedicel 4-5 mm
long, bracteole subapical; buds obovoid, 5 by 3.5 mm, cleft over 1/2; ovary ovoid-glo-
bose, 2.5 mm long; stigma sessile, 2-lobed and each lobe again deeply 4- or 5-lobulate.
Fruits 1-3 per infructescence, subglobose or broadly obovoid, 1.8—2.5 by 1.5-1.7 cm,
with rusty hairs 0.3 mm long; fruiting pedicel 4-6 mm long.

Distribution — Malesia: Borneo (Sarawak, W & E Kalimantan).

Habitat & Ecology — Freshwater and peat swamp forest, lowland forest; 0-300 m
altitude; fl. Oct.

Note — Knema uliginosa is related to K. intermedia, which differs in the more pro-
nounced mammillate staminal disc, more distinctly reticulate leaves, and fruits with an
easily rubbed off scurfy indumentum, and also to K. plumulosa which differs in the
more conspicuous indumentum on twigs and leaves. Knema uliginosa may resemble
K. latericia, which differs in the cracking and flaking bark of the older twigs; compare
also K. korthalsii.

74. Knema viridis W.J. de Wilde

Knema viridis W.J. de Wilde, Blumea 32 (1987) 121, f. 3; Tree Fl. Sabah & Sarawak 3 (2000) 447.
— Type: S 37882 (Yii Puan Ching), Sarawak.
Knema linguiformis auct. non (J. Sinclair) W.J. de Wilde: W.J. de Wilde, Blumea 27 (1981) 227.

Tree 5-30 m. Twigs somewhat flattened or blunt-triangular by ridges from the base of
the petioles, (2—)2.5— 4.5 mm diameter, at first with dense pale brown hairs 0.2—0.3 mm,
early glabrescent; bark yellowish turning purple-brown, not or but little striate, neither
cracking nor flaking. Leaves chartaceous or thinly coriaceous, oblong(-lanceolate), 12—
30 by 3.5—10.5 cm, base attenuate to rounded, apex blunt to acute; drying greenish above,
lower surface early glabrescent, greyish green; dots absent; midrib raised above; nerves
1423 pairs, raised; venation distinct; petiole 12-18 by 1.5—3 mm. Inflorescences: pedun-
cle up to 2 mm long, brachyblast wart-like, simple or bifurcate, 2-5 mm diameter, in
male 8—12-flowered, female 4—7-flowered; flowers with (pale) brown scale-like hairs
0.1(—0.2) mm; perianth 3-lobed, inside colour not recorded. Male flowers: pedicel stout,
7-10 mm long, bracteole blunt-triangular, 1—1.5 mm long, caducous, about median, the
pedicel above the scar distinctly thicker than below; buds depressed-globose, 4.5—5 by
6—7 mm, faintly angled in cross section, cleft 3/4-4/5, lobes (1—)1.5—2 mm thick; staminal

Fig. 58. Knema viridis W.J. de Wilde. a. Habit of male flowering twig: b. opened male flower,
showing thick perianth and androecium with many appressed anthers; c. androecium seen from
somewhat above, staminal disc subtriangular in outline; d. part of twig with female inflorescences;
e. opened female flower, showing pistil with pubescent ovary, short style and many-lobed stigma:
f. opened fruit, pericarp glabrous, thick-lobed, aril of seed laciniate at the apex [a—c: S 37882; d, e:
S 42885; f: S 38068]. — Scale bar for a, d, f = 2 cm; for b, c, e = 1.65 mm.

358 Flora Malesiana, Ser. I, Vol. 14 (2000)

disc flat, sometimes convex in the centre, blunt-triangular in outline, 2.5—-3 mm diam-
eter, anthers 25—30 (thecae 50-60), sessile, 0.5—0.7 mm long, closely set, + hidden under
the disc below the margin; androphore tapering, 1-1.5 mm long, at base | mm diameter.
Female flower (slightly immature): pedicel stout, 15—17 mm long, bracteole + below
median; buds depressed-globose, 3.5 by 5 mm, cleft 3/4—4/5, lobes 0.5—1 mm thick;
ovary broadly ovoid, 1.5 mm diameter, with dense brown hairs 0.2—0.3 mm at apex,
narrowed into a style-like part 0.5 mm, pubescent at base, stigma 1.5 mm diameter,
broadly subpeltately 2- (or 3-)lobed and each lobe again shallowly 4—6-lobulate. Fruits
1-3 per infructescence, ellipsoid, broadly rounded at both ends, early glabrescent (hairs
not known), bright brown, 5—6 by 4—4.5 cm; dry pericarp 10 mm thick; fruiting pedicel
15 mm long. — Fig. 58.

Field-notes — Slender tree to c. 30 m, c. 95 cm girth, without or with buttresses to
one side, c. 80 cm wide; bark smooth, slightly scaly, or with lenticels, brown or blackish,
exudate red; inner bark soft. Twigs green with dark patches, dirty green or black, when
young with brown indumentum. Leaves glaucous at underside. Flowers light brown or
brown-green. Fruits elliptic or oblong, yellowish green, with slight brown indumentum.
Aril yellow to red, sweet, laciniate at apex for 1/5—1/4.

Distribution — Malesia: Borneo (Sarawak).

Habitat & Ecology — Lowland and hill mixed dipterocarp forest; on gentle slope;
wet slope near ridge; 50-600 m altitude; fl. Aug.—Nov.; fr. Sept.

Note — Knema viridis belongs in the group of K. curtisii, and seems most related to
K. linguiformis. It is markedly stouter in all parts, with larger leaves, and stouter male
buds with shorter hairs, and larger fruits with spongy dry, very thick pericarp (1 cm).
The fruits of the related K. linguiformis are 3.5—4 cm long with fruiting pedicel up to
8 mm long and pericarp 1-2 mm thick, but with seeds of about the same size as those in
K. viridis (seeds 25-30 mm long). Male buds of K. linguiformis may reach 6 mm diam-
eter, but the flower is more delicate, with woolly hairs up to 0.4 mm long. Knema viridis
also shows affinities with K. woodii, with similar twigs and thick perianth lobes, a simi-
lar obconical androecium (but with the anthers spaced, not tightly appressed), and a pis-
til with the ovary narrowed into a slender style-like part.

75. Knema woodii J. Sinclair

Knema woodii J. Sinclair, Gard. Bull. Sing. 18 (1961) 283, f. 26; W.J. de Wilde, Blumea 25 (1979)
429: 32 (1987) 125: Tree Fl. Sabah & Sarawak 3 (2000) 449. — Type: Sinclair & Kadim 9298,
Sabah.

Tree 6-30 m. Twigs + 3-angled or ridged, often blackish, 2.5—4 mm diameter, at first
with puberulous hairs 0.1—0.2 mm long, early glabrescent; the bark almost smooth or
+ striate, not tending to crack or flake. Leaves thinly coriaceous, oblong to (ob)lanceolate,
15-35 by (2-)4-9 cm, base rounded to cuneate-attenuate, apex acute(-acuminate), dry-
ing (greenish) brown above; lower surface greyish, at first with hairs 0.1—0.2 mm long,
early glabrescent; dots absent; midrib flat or raised above; nerves 22-30 pairs, sunken
above, light brown beneath; venation faint or distinct; petiole 10-20 by 3—4 mm. Inflo-
rescences: (sub)sessile, brachyblast simple or warted, 2-5(—10) mm diameter, in male
and female 1—10-flowered; flowers with pale brown or yellowish brown stellate hairs up
to 0.3 mm long, sometimes seemingly glabrous; perianth 3-lobed, red inside. Male flow-

De Wilde — Myristicaceae (Knema / Myristica) 359

ers: pedicel 8-15 mm long, bracteole + caducous, about median or lower; buds
subglobose, (5—)6—8 mm diameter, cleft c. 3/4, lobes 1.5—2 mm thick; staminal disc
subtriangular, flat, 3-5 mm diameter; anthers 18-23, sessile, + oblique, often hidden
under the disc, 0.3—0.4 mm long, not or hardly touching; androphore stout, tapering,
2(—-3) mm long, clasped by the thickened fleshy basal part of the perianth. Female flow-
ers: pedicel 3—4 mm long, bracteole about median; buds obovoid, 5—7 mm long, cleft
c. 2/3, lobes 1-2 mm thick; ovary ovoid, 1.5-2 mm long, style 1-1.5 mm long, stigma of
2 main lobes and each lobe again 4—6-lobulate. Fruits 1 or 2 per infructescence, ellip-
soid, 3—4(—5) by 2—2.5(—4) cm, thinly with mealy greyish or yellowish brown scale-like
stellate hairs 0.1 mm long or less, subglabrescent; dry pericarp 4-7 mm thick; fruiting
pedicel 7-10 mm long. — Fig. 34a.

Field-notes — Bark of trunk grey or brown, flaking in thin strips or pieces; living bark
8-10 mm thick, (pale) red; wood reddish or dirty white. Leaves glossy green above,
glaucous beneath. Flowers fleshy, red inside; male flowers fragrant, androecium pink
with yellow or brownish anthers, pollen yellow; female flowers with the pistil brown at
base, green at apex. Fruits ramiflorous, more or less solitary, with yellow pericarp; dry
seed obliquely ellipsoid, 23-25 by 16-18 mm.

Distribution — Malesia: Borneo (Sarawak, Sabah, E & NE Kalimantan).

Habitat & Ecology — Mixed dipterocarp forest, periodically flooded forest along
rivers; locally common on alluvial soil in Central Kutai; clay soil, sandstone; 0-500 m
altitude; fl. throughout the year.

Notes — 1. Knema woodii seems related to K. pectinata, K. glomerata, and K. kort-
halsii. It is distinct in the numerous, | 8—23, sessile anthers. Knema pectinata is close by
similarly thick perianth lobes, and very minute indumentum, but differs in more pairs of
lateral nerves. In K. korthalsii the staminal disc is convex or mammillate, with 11-19
(—20) anthers, the indumentum on the twig apices is more distinct, and the inflores-
cences have more flowers.

2. When sterile, K. woodii may be recognized by the pale yellow, + mealy stellate-
dendroid hairs on the twig.

MYRISTICA

Myristica Gronov., Fl. Orient. (1755) 141, nom. cons.; Warb., Mon. Myrist. (1897) 374; J. Sinclair,
Gard. Bull. Sing. 16 (1958) 333-368; 28 (1968) 1-540; W.J. de Wilde, Beitr. Biol. Pflanzen 66
(*1991°, 1992) 113; Kubitzki et al., Fam. & Gen. Vasc. Pl. 2 (1993) 466; W.J. de Wilde, Blumea
35 (1990) 233-260; 38 (1994) 349-406; 40 (1995) 237-344; 42 (1997) 111-190; Tree Fl. Sabah
& Sarawak 3 (2000) 449. — Myristica sect. Myristica Blume, Rumphia | (1836) 180; Hook. f. &
Thomson, FI. Ind. 1 (1855) 162 (sect. Eumyristica p.p.); A.DC., Prodr. 14, 1 (1856) 189, p.p.;
Mig., FI. Ind. Bat. 1, 2 (1858) 53; Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 205, p.p.; Hook. f., Fl.
Brit. India 5 (1886) 102; King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 286. — Type species: Myr-
istica fragrans Houtt.

[Comacum Adans., Fam. 2 (1763) 345. — A superfluous name for Myristica Gronov.]

Trees, dioecious (occasionally M. fragrans monoecious, M. crassa paroecious); stilt-
roots sometimes present. Twigs lenticellate, sometimes flaky, in some New Guinea spe-
cies with ridges and in places swollen, hollow, inhabited by ants; bark sometimes coal
black. Leaves brittle when dry, lower surface often pale, papillose or not, dots present or

360 Flora Malesiana, Ser. I, Vol. 14 (2000)

absent: reticulation scalariform or a lax network (never as dense as in Knema, more
abundant than in Horsfieldia). Inflorescences sometimes supra-axillary, either a pedun-
culate branched panicle with the flowers in subumbels, or a (sub)sessile woody simple
or 2—4-armed wart-like or worm-like densely scar-covered brachyblast producing flow-
ers at the apex (as in Knema), or less often a combination of the two types when the basal
part is smooth and bears 1-3 sparsely or densely scarred brachyblasts at its extremity;
basal cataphylls absent; bracts caducous. Flowers pedicellate, bracteole mostly persist-
ent, often apical and embracing the perianth. Male flowers: perianth urceolate or tubu-
lar, leathery or carnose, tubular, inside glabrous, creamy; buds variable, cleft to various
depths, lobes 3 (also 2 or 4), reflexed in anthesis. Androecium a stalked column with
8-30 completely fused linear anthers; androphore cylindrical, glabrous or pubescent;
synandrium elongate, the apex often sterile, rarely shallowly hollowed. Female flowers:
larger and more swollen than the male; ovary glabrous or hairy; style absent, stigma mi-
nutely bilobed. /nfructescences sessile or stalked. Fruits globose to oblong, or pyriform,
1-10. cm long, glabrous, tomentose, or lanose; pericarp mostly thick, fleshy but firm; aril
laciniate (nearly) to the base; seeds not variegated; albumen ruminate, with fixed oil and
starch; cotyledons connate at the base and edges, divaricate. — Fig. 59-93.

Distribution — Some 175 species, from S India and Indochina (rare) through Malesia
east to NE Australia and in the Pacific east to Fiji. Most species occur in New Guinea,
with many endemics. — Map 5 (see p. 5).

Habitat & Ecology — All species are confined to undisturbed rain forests and many
have a restricted distributional area and altitudinal range, sometimes restricted to soils
derived from specific bedrock, e.g. limestone or ultrabasic. Most species occur in low-
land forests or lower to mid-montane forests; a few species in New Guinea reach an
altitude of c. 2000 m.

Taxonomy — Sinclair, in his monograph (1968), distinguished two sections, |) sect.
Myristica, with inflorescences more or less branched panicle-like, and with a smooth,
non-scar-covered (often + flattened) peduncle, generally lasting but one flowering sea-
son, and 2) sect. Fatua, with (sub)sessile, simple or forked scar-covered, wart- or worm-
like inflorescences (as in the genus Knema), lasting several flowering seasons. Within
each section respectively 9 and 10 series are recognized, slightly less (and differently
defined) as was done by Warburg (1897). Sinclair (1958, 1968) was aware of the many
intermediate forms of inflorescences of the species, rendering the distinction of the two
sections very weak. As discussed by De Wilde (1992, 1994) the shape and architecture
of the inflorescences are useful in the keys, but a (monophyletic) division of the genus
into two sections is unwarranted. At present, no sections or series are recognized.

The nutmeg of commerce, M. fragrans, has delicate inflorescences of a variable struc-
ture, and a large part of the material actually should be grouped in the section with M.
fatua-type inflorescences. However, M. fragrans is the type species of the genus Myristica,
and hence of section Myristica.

Note to the Keys: Besides a general key to the species (1), mostly based on male flower-
ing specimens, separate regional keys (2-7) are given, mainly based on female flower-
ing and fruiting material, although vegetative characters are also used. Because most
species of New Guinea are different from those of the rest of the Malesian area, but
difficult to segregate from the others in a general key, they are keyed out separately (8).

De Wilde — Myristicaceae (Myristica) 361

(1) GENERAL KEY TO THE SPECIES OF MALESIA (EXCLUDING NEW GUINEA)

tar

Dian

3a.

)

(applicable for male flowering specimens, based on characters of male inflorescences and

flowers*:; vegetative characters and also fruit characters** have been used additionally)
g y

Male inflorescences branched (rarely unbranched), with a slightly flattened pedun-
cle, or (sub)sessile, delicate, flowers few and spaced; see further the definitions un-
der lead 2. Inflorescences lasting one or but a few flowering seasons ......... Z

. Inflorescences sessile or short-pedunculate densely scar-covered short-shoots, last-

ing several flowering seasons, each season producing new flowers at the apex; pe-
duncle smooth, terete, to 5 mm long; short shoot simple or 2—4-forked, rather thick,
Be ese mcastane ty OnfctIA= TON SNS «eT ol MICA Cy Oe VUE e chal) Cee Bey, cE et Ce AR 37.
Inflorescences with flowers in fascicles or subumbels of buds varying in size at
the end of the ramifications; occasionally unbranched, then peduncle apically with
buds in a single fascicle not growing out into a densely scar-covered short-shoot
(Gee iF ioe EA ORD rekAee aarekA ce ic ret ea a a ie a ee ee 3

. Inflorescences either simple, delicate, subsessile, with few spaced flowers, or pe-

dunculate, branched, the branches ending in spike-like + densely scar-covered short-
shoots, producing new flowers at the apex (see Fig. lo, p. 11) ............. ail
Mature leaves on lower surface glabrous or early glabrescent, or with scattered, nei-
ther touching nor interwoven weak hairs 0.1—0.3 mm or less, or with very thin cob-
Wweb-ike covering and leaves seeming glabrous =. 2°... 4: Gen. eels 55 52 -

. Mature leaves densely hairy below, but hairs may be very small (lens!); old leaves

RRNA OATES CCIM © Sree. cats RE Ae 5 hohe 5 eeu s: ee wre sae les pen ems Cae ea 26

. Leaf bud with hairs 0.1—0.2(—0.3) mm or less [Apex of young twigs and inflores-

cences with dense or sparse similar indumentum, or glabrescent.] ........... 5

. Leaf bud with hairs (0.3-)0.5 mm long or more (appressed hairs may seem shorter)

IE Ps rrersts Anata, « bays Oh are oe Gar pA ames RNa oes ARE ae «ERSTE. 21

. Buds sharply angular or winged along the lobe sutures. Leaves 10—20(—30) by 3-9

cm, nerves 8-13 per side. — S Thailand, West Malesia ........ 38. M. elliptica

meudsmounded tosubangled: Leaves:wvariables barse.c.. sity alee © ee ee 6
. Leaves 2(—3) times longer than broad, 16-44 by 8—14.5(—18) cm, apex blunt or acute,

base + rounded or subcordate; nerves (16—)20—28 per side. — Borneo...........
MN sees oc Ns sd Howe. Laces beeen Mane nits, Shah pie od ee 102. M. papyracea

. Leaves large or small, about 3 times longer than broad or more, nerves 10—33 per
side, apex acute(-acuminate), base various, rarely subcordate; leaves smaller with
apemaeute,or bluntimmost of Migicanted sc. 2v.teney aceon ae earl 7

. Plants glabrous. [Male flowers not known; infructescence partly not known.] —
Prolniccas(Seraiit).. Snccawres: es ead tee. Meare: Sere ee stent 104. M. perlaevis

alanis clabrescent, or,(parthy) pubescent ...gtt,sny.) 2 lances Arq) aera eer ee 8

. Androphore at anthesis much shorter than the synandrium ................-. 9

»Androphore almost,as-lonesas the synandniumt 2) 09) team sted as see eee 16

Male inflorescences and flowers of several species are not known, and the place in the key of
these species may be erroneous.

**) The use of fruit characters in a key mainly for male flowering specimens may be helpful when

obviously identical material of both sexes is available.

362

Flora Malesiana, Ser. I, Vol. 14 (2000)

16a.

18a.

a. Twigs 4-8 mm diam. [Buds 5—7 mm long. | Androecium (synandrium) cylindrical,

3-5 mm long. [Androphore pubescent at base.] .......--- 0.0 seer eee eees 10
. Twigs 2-5 mm diam. Androecium short, often + club-shaped or ellipsoid(-oblong),
1-5=3(3:5) mong 3.2 0 ase o,f SRE ERS. Ee Ee 11
Leaves blackish: lower leaf surface with minute + interwoven hairs, sometimes
glabrescent. — Sumatra, Peninsular Malaysia, Borneo ....... 88. M. maxima
_ Leaves drying slightly olivaceous: lower leaf surface with minute scattered hairs.
== Phillippines ene Moe ihels RUE SAS RCE aR 51. M. frugifera
. Buds 6—6.5 mm long. — Philippines ..........----2 000 ee eeee 77. M. laevis
*Buds3—4(]4:5) min longs a: sale. ele OE BORE. CI ee 12

. Twigs and inflorescences often drying yellowish, the leaves greenish yellow or

olivaceous, papillose below. [Buds cleft c. 1/2(—2/3). Androecium + ellipsoid,
androphore pubescent. Fruits with persistent scurfy short indumentum. ] — Borneo
(Ded Jets lnntuni eterna el aus mentyies MAE aeiel nelle eats: teh btea ae 12. M. borneensis

. Twigs and inflorescences drying brownish, the leaves (olivaceous-)brown .. 13
a. Androecium cylindrical or ellipsoid, androphore pubescent. Inflorescences 3-6 cm

long. — Philippines (Gincl.;Palawan)) 2.222.294) 1 14

_ Androecium + club-shaped, androphore glabrous. Inflorescences more than 6 cm

VOM Gea oo, whey svny> secesnoncysvscclcesie ines say siteehernG Me: eorda de © See Sel eae [5

_ Midrib and lateral nerves on lower leaf side yellowish. [Fruits with grey-brown in-

dumentum.] — Philippines (excl. Palawan) .........-.-.. 1. M. agusanensis

_Midrib and lateral nerves on lower leaf side dark purplish or reddish, contrasting

with greyish surface. [Fruits with rusty or orange-brown indumentum. ] — Philip-
pines (Palawan, Mindoro) ...........+-+++++e+reeeee> 116. M. rubrinervis

_ Inflorescences 7-10 cm long; buds 3—3.5 mm long, cleft 1/4—1/3; androecium with-

out sterile apex, androphore short or absent. [Fruits glabrous or largely nearly so.]
— Sumatra, Peninsular Malaysia, Borneo ...........-.-- 86. M. malaccensis

_ Inflorescences 10-15 cm long; buds 4(—4.5) mm long, cleft nearly 2/3; sterile apex

of androecium + blunt, 0.2-0.3 mm long; androphore distinct. [Fruits scurfy.] —
|EYG YG 0 Go pen Id Car rn AS Bk he aa Ba AR La BO Len SSSR. 3 1 40. M. extensa
Buds 7-8 mm long. Leaves below seemingly glabrous, but actually with scattered
minute scale-like hairs less than 0.1 mm (lens!). [Inflorescences usually unbranched.
Fruits subglobose or broadly ovoid, 3—4.5 cm long, with persistent scurf of hairs
0.1 mm or less.] — An inland species of C & E Java. (Compare also M. crassa)
MS Rs Te LIS eee er Rae, SAIS ARs NcMOMon ee ee tae 135. M. teijsmannii

_ Buds 7 mm long or less. Leaves below glabrous (early glabrescent) ........ 17
17a.

Twigs 1.5—3(—3.5) mm diam. Lateral nerves above sunken, distinct. Peduncle 2—40
mm long, the central branch present. Buds 4—6(—7) mm long, pubescent or
glabrescent. Fruits’3.5=8.5 cm long: .¥. 225. 2222 22 ees 2) ie ae ae 18

. Twigs 1-2 mm diam. Leaves papillose below (lens!), lateral nerves indistinct above.

Inflorescences almost unbranched, peduncle 4-10 mm, flowers in a subumbel. Buds
5 mm long, glabrescent. Fruits 3.5—4 cm long. — Philippines (Palawan)........
3 Bay DRO es Pa SIE ON: TY, Lea COR ice cco ntomaehegearme 141. M. umbellata
Leaves (15—)20-35 cm long, drying dull greyish (olivaceous) brown, not papillose.
Lateral nerves below flat or but slightly raised. Buds 5 mm long; androecium 3-3.5
mm long, androphore glabrous. [Fruits 5.5-8.5 cm long, pericarp (5—)10-15 mm
thick.] — Sumatra (doubtful), Peninsular Malaysia, Sabah 152.M. wyatt-smithii

De Wilde — Myristicaceae (Myristica) 363

b.

19a.

20a.

aia.

22a.

23a.

24a.

25ay

Leaves smaller, 7—24 cm long, olivaceous. Lateral nerves raised below. Androphore
esc cuededind a Noh een iran iaeia iets 9S eG ANE ae 19
Leaf apex acute-acuminate, lower surface not papillose. Buds 4-6(—7) mm long,
(late) glabrescent; androecium 3—6 mm long. Fruits 3—6(—8) cm long, pericarp
SS hihmmdthick,— ThailanditoxBomeo! minkhiinrisliiud «> ee eRRe 20

. Leaf apex blunt to acute (in leaves of sterile sapling shoots acute-acuminate), lower

surface papillose. Buds 4—-4.5 mm long, with persistent indumentum; androecium
3.5 mm long. Fruits 5-8 cm long, pericarp 15-20 mm thick. — Sumatra, Peninsu-
nm ialaysiasbormeoiin | 6s). Sterna) SiGe Se ee 56. M. gigantea
Leaves membranous, chartaceous, or subcoriaceous, 7—24 cm long. Female inflo-
rescences or infructescences with peduncle to 1 cm long. Fruits 4—6(—8) cm long,
pericarp 4-10 mm thick, late or early glabrescent. Buds 4—6(—7) mm long.......
Rent) Elo vy ore ers, Peo .8. GEL SUL GO ee 66. M. iners

. Leaves thinly membranous, 7-14 cm long. Peduncle of female inflorescences or

infructescences slender, 1—2.5 cm long. Fruits 3—4 cm long, early glabrescent; peri-
eap4—4-mm thick» Buds: 3mm long yas. se. bee. 22 oe oe 41. M. fallax
Twigs 1.5—2(—2.5) mm diam. Leaves 7—12(—15) cm long, apex subacute or rounded;
lateral nerves 10—15(—18) per side. Buds 4—4.5 mm long; synandrium without ster-
ile apex. Fruits S—8 cm long, late glabrescent, at first with scurfy hairs 0.2-0.3 mm.
= Sumatta,~cenmsular Malaysia, Bomeoy, ... Jace 2 ee 56. M. gigantea

. Twigs generally thicker; leaves larger, the apex acute-acuminate, lateral nerves 13—

18 per side or more. Synandrium with small sterile apex. Fruits pubescent or glabres-
Scien tMiialirs 0.2 Mali OGAMOTS te esMMATAE I LE ose er as ere DD
Twigs (1.5—)2—4 mm diam. Leaves membranous or thinly coriaceous. Buds 4—7
mm long. [Sterile apex of synandrium present, acute or + blunt, entire.] Fruits gla-
Brousronearhy-clabrescentria oh 8 Ue. ~~. SOSA GR ee Ds.

. Twigs 3-6 mm diam. Leaves chartaceous or coriaceous. Buds variable. Fruits gla-

brous or with persistent indumentum, rarely late glabrescent ............. 24
Twigs 1.5—3 mm diam. Leaves membranous, the lower surface not papillose. Buds
4—6(-7) mm long. Fruits (3.5—)4—6(-8) cm long. — Stout form; mainly from NE
Baie oatreriit's .-.csnepeeidale ee 2ot Carl Ns ak ee Ee 66. M. iners

. Twigs (2—)2.5—4 mm diam. Leaves chartaceous or thinly coriaceous, lower surface

distinctly papillose. Buds 6-7 mm long. Fruits 7-9 cm long, glabrescent, at first
with scurfy hairs 0.5 mm. — Peninsular Thailand, Peninsular Malaysia.........
Bree ts Herder Hi ceminads Meee els ee 85. M. maingayi
Leaves coriaceous, papillose below. Flowers pubescent with hairs 0.5—1 mm, buds
4—4.5 mm long; sterile apex of synandrium + lobed. Fruits with conspicuous shaggy
hairs 1-2 mm. — Sumatra, Peninsular Malaysia, Borneo ...... 84. M. lowiana

_ Leaves chartaceous or coriaceous. Flowers with hairs 0.3—0.5 mm; buds 5 mm

long or more; sterile apex of synandrium not lobed. Fruits glabrous or with scurfy

ins O) Samumiter! wien ee tudo, Mu Bs eee Min BORE eee: 25
Buds 5—5.5 mm long. Bark of leafy twigs early cracking. Leaves not papillose be-
low. Fruits glabrous (early glabrescent). — Borneo .......... 28. M. corticata

. Buds (5—)6—8 mm long. Bark of twigs late cracking. Leaves papillose or not. Fruits

with scurfy indumentum or late glabrescent. — Philippines.................-

364

Flora Malesiana, Ser. I, Vol. 14 (2000)

26a.

27a.

30a.

Twigs (4-)5—8 mm diam. Leaves large, 16-40 cm long; lateral nerves 23-33 per
side. [Lower leaf surface with persistent indumentum or glabrescent; hairs + scat-
tered. minute, scale-like, 0.1(—0.2) mm or less.] Inflorescences stout, 6—-12(—18)
cm long. Fruits 6-9 cm long, with scurfy hairs 0.1 mm or less, sometimes late
glabrescent. — Sumatra, Peninsular Malaysia, Borneo ....... 88. M. maxima

. Twigs 5 mm diameter or less. Leaves generally smaller with fewer pairs of lateral

nerves. Male inflorescences shorter. Fruits of various sizes ........-...--. 27
Leaf bud and young twig apex with hairs 0.1 mm. Lower leaf surface densely pu-
bescent with scale-like hairs 0.1(—0.2) mm (old leaves sometimes glabrescent).
Buds cylindrical or slightly contracted in the middle; apical part of bud + angled;
sterile apex of synandrium + blunt or subacute, usually finely pubescent (glabrous
in Sumatra). [Fruits 5-9 cm long, minutely scurfy.] — Peninsular Malaysia to
Phihippines?) ). PSY iets saa dd ad dessa cee itom tee 21. M. cinnamomea

_ Leaf bud, twig apex, and lower leaf surface with hairs short or long, 0.1 mm or

much longer. Buds + ellipsoid, apical part not angled; apex of synandrium + flat or
shallowly hollowed, the excavation concealed by the somewhat incurved apices of
thelanthersvor mots. 2210! S82 Cle RING. SR Eee 28

. Twigs 1.5—3 mm diam. Leaf bud and twig apex with hairs 0.1 mm. Buds 3—4 mm

long; androecium 1.5—2.5 mm long; anthers 6-8(—10). Fruits with hairs 0.1 mm or
less, sometimes glabrescent. — Not generally a coastal species ........... BS.

. Twigs 3—5(—10) mm diam. Leaf bud and twig apex with conspicuous hairs (0.1—)

0.2-0.5(—1) mm. Buds larger, 4-7 mm, androecium larger, anthers 8-12. Fruits
2.5—4.5 cm long, with more conspicuous persistent hairs 0.5—1(—2) mm. — Gener-
ally a coastal species; Burma, east to Bali and Philippines 59. M. guatteriifolia

_ Androecium 1.5 mm long. Lateral nerves 45—80° to the midrib. Fruits 5—7 cm long,

with dense scurfy indumentum. — Sumatra, Peninsular Malaysia, Borneo .......
LOS. ShCUY ZOE g BUREN OR TO BEM TTELOR O nk A eee Eee 35. M. depressa

_ Androecium 2.5 mm long. Lateral nerves c. 45° or less to the midrib. — Philip-

pines! ig.6{) Mek son s2e. WEN e LU. a DOSTERIONS Ue. AES ENE ee 30
Midrib and nerves on lower leaf surface (grey-)brown. Fruits 34.7 cm long, minute-
ly dull grey-brown, scurfy, sometimes partly glabrescent. — Philippines (excl. Pala-
WAI) OL, PNOR DED. GORD. SR ATEEO AIR JRE Nene 1. M. agusanensis

_ Midrib and nerves purplish or reddish brown, contrasting with grey-brown lower

leaf surface. Fruits 3-7 cm long, with short dark rusty or orange-brown indumen-
tum, sometimes partly glabrescent in var. duplex. — Philippines (Palawan, Mindoro)
BESS chal LENS BEN SA ATTA LO A OE REO De 116. M. rubrinervis

wolf be So outs, cae GR RIBN 2S LURES 5 oR Ss. 2) Jee 32

. Buds not angular, apex rounded. [Lower leaf surface generally papillose.] .. 33
. Buds ovoid-oblong, (4.5—)5—7 by 3-4 mm, hairs 0.1 mm. Leaves below grey-brown

with contrasting bright brown nerves, not or but indistinctly papillose. — Philip-
PUESHe ION LOY RIA, MAIS. LSS LOLS LIC RO es 32. M. cumingii

. Buds + elongate, (elliptic-)oblong, 4.5—8 by 2-3 mm, with hairs (0. 1-)0.2—0.5 mm.

Leaves below dull, sometimes whitish (origin of pale colour unknown), nerves not
much contrasting. — W Sarawak, W & NE Kalimantan, Sulawesi, Sulu Is., Philip-
pines, N Moluccas (Sula, Bacan) .........-..55+s eee 123. M. simiarum

De Wilde — Myristicaceae (Myristica) 365

33a.

34a.

35a.

Ala.

Buds 5—11 mm long. — Moluccas (or cultivated) ..........-..--.--005. 34
_ Buds 3—4.5(—5) mm long, rough-pubescent. Nerves on upper leaf surface flat or
Baminetivamapnessed ay: ya 7. SUPP IAM. Nin. A Aes eee ete ae 36
Buds either glabrous, early glabrescent, or short-haired. Nerves 6—12(—15) per side,
above flat or but little impressed, much raised below ...............---- a5
. Buds conspicuously pubescent. Leaves large, 25—45 cm long, lateral nerves 25—30
per side, impressed above. Bark of twigs longitudinally cracking. — N Moluccas
A tea) ATS: 2- SR Se crcccheoars louse cite eas amine mitaniolennss 47. M. fissurata

Pedicel more than | mm thick. Flowers thinly pubescent. Leaves subcoriaceous,
9-22 cm long, below late glabrescent, at first with minute indumentum. Twigs 2—
a aioamyat arn. Ni Me@liccas Mews aie Ser. ee eS 131. M. succedanea

. Pedicel less than 1 mm thick. Flowers glabrescent, at first with sparse very minute

indumentum. Leaves chartaceous, 6—13 cm long, early glabrescent. Twigs 1-2 mm
diam. — Originating from Banda (Moluccas); known only from cultivated speci-
Mier .h cs Heer ba apoio Wes Aw Lede ate ase zee 50. M. fragrans

_ Leaves membranous, whitish below, not obviously papillose. — Sulawesi.......

REED ent o> Jans ne to. een aA SE Os aoe. 63. M. impressinervia

. Leaves chartaceous, pale brownish below, distinctly papillose. — Moluccas, Papua

Ber isind. Ss IGAd) (4) eaantysra A Aer bn ise sted oencpcies Ge ee me 11. M. bifurcata
Buds elongate, (10—)13-15 by 2.5-5 mm ........-.--. 2-0 see ee eee eee 38
mends-- ovate, less than lO mm long, 2... ...5.....1J3RRD Pe eee Sere - 39

. Twigs ridged or winged, myrmecophilous. — Kai Is. (& New Guinea).........

Deweeenl If ©... esmcortlls....d wel a Jon eee Bee 129. M. subalulata

. Twigs not winged, not myrmecophilous. — Moluccas (Halmahera, Obi)......-.

me nny! or. FT Bt oe Cee re Lh ec aa 110. M. pubicarpa

. Most parts with conspicuously long hairs, 1-3 mm (lower leaf surface glabrescent).

Bark of twigs blackish, longitudinally cracked or flaking. [Fruits 4-6(—7) cm long
with hispid hairs 2—-3(—4) mm.] — Borneo .............-..-. 147. M. villosa

_ Hairs shorter, very short or up to | mm long, or indumentum (almost) absent 40
. Twigs 4—7(-10) mm diam. (in Peninsular Malaysia 3 mm diam.) .......... 4]
. Twigs generally 1—4(—5) mm diam. (sometimes 4-5 mm diam. in M. beccarii, M.

fatua, M. nivea). — Plants not from Sumatra or Peninsular Malaysia ....... 48
Leaves rather narrow, | 1—24(—30) by 3-6 cm, coriaceous. — Philippines (NE Luzon)
Wy EISLER a Hain. gob ee MARA s Le eacy e 24. M. colinridsdalei

. Leaves generally larger, comparatively broader, (15—)17—40 cm long; membranous

REEALTACCOLISY / thd. |. BCL, ates wos Shatiee | ENE See 42

_ Lower leaf surface early glabrescent or with scattered minute scale-like hairs less

fhamiosltnami(lens!'). 7.0 atn2h arsictdl. 00s he eee, eee eee = 43

_ Lower leaf surface densely tomentose (hairs may be very minute), not or not evi-

Periingnapillascw tress LAME Soo. ie eae end eee 45

. Lower leaf surface papillose (lens!). Buds 4.5—6 mm long. Fruits 3.5—5.5 cm long,

with scurfy hairs 0.1 mm. — West Malesia (Sumatra, Peninsular Malaysia, Bor-
Tee Mt Se )k- ee eT. Se a2 SA ee er ot 29. M. crassa

» Lower leaf surface not or not evidently papillose ..............+.-+-+---- 44
. Buds 8-8.5 mm long. Fruits not known. Twigs 5—10 mm diam.; leaves 30—40 by

1420 cm, nerves distinct. — N Moluccas (Bacan) .......... 114. M. robusta

366

Flora Malesiana, Ser. I, Vol. 14 (2000)

54a.

_ Male flowers not known. Fruits globose, 5 cm diam. Twigs 4-5 mm diam.; leaves

20-30 by 6-10 cm, coriaceous, nerves indistinct. — C Sulawesi............-.
sarees. SRD AG,.. SERA PEI to BE ee ry A AEs 36. M. devogelii

a. Lower leaf surface with pale scale-like hairs 0.1 mm only, thus seemingly gla-

brous. [Fruits 6 cm long, with rusty hairs 0.5—1 mm.] — Philippines...........
eB ok ae ar ee re a ee 150. M. wenzelii

_ Hairs on lower leaf surface conspicuous, brownish, 0.2—0.3(—0.4) mm. — Borneo,

BT aD aera kw sce s Chis PS ac MEL, 46

_ Flowers pedicellate. Two species, distinguishable in the indumentum of the fruits

as ere apes atat 0g oh LN: Se LR Ea 47

_ Flowers sessile. Fruits 2.5—3.5 cm long, with shaggy hairs 1—-1.5 mm. — Lesser

Sandarisiands 4:4 du 0. Re. AAS Se ee EE: 133. M. sumbawana

_ Buds 6 mm long. Fruits 5.5—8.5 cm long, pericarp 10-15 mm thick, with scurfy

hairs!0M=0.2immy—==N Moluccas) We ae eee 118. M. sangowoensis

_ Buds 4-6 mm long. Fruits 4~7 cm long, pericarp 3—12 mm thick, hairs 0.5—1 mm.

— Philippines (Mindanao), Moluccas, Sulawesi, and a deviating specimen with
male flowers from limestone in SE Kalimantan ..............-. 43. M. fatua

_ Lower leaf surface glabrous or early glabrescent, or with minute scattered hairs

De BB. orth Blk. ak Mantes SS fo hk, LE 49

_ Lower leaf surface densely hairy; sometimes hairs small and scale-like, but densely

Setior interwoven: ens!))is..\), cal coast. eRe. St ote 66

. Petiole proportionally long: leaves 10-15 cm long, petiole 20-35 mm long. [Male

inflorescences and flowers not known.] — Philippines .. 82. M. longepetiolata

. Petiole proportionally shorter ..........50. 0-000 eee eee ee 50
_ Leaves small, elliptic-oblong, thinly coriaceous, (2.5—)4—7.5 cm long; lateral nerves

faint. [Flowers not known. Fruits ovoid-ellipsoid, 2—2.5 cm long, pericarp 1-2 mm
thick, with dense scurfy hairs 0.1 mm or less.] — Central Sulawesi............
Beh Shin. 25. cn dean Lticr eae a ee 140. M. ultrabasica

eaves daraer) sui ilns 29.280) CURL 100d I Se Sil
Sla.

Buds + cylindrical, narrow; androphore about as long as the synandrium. — Moluc-
cass W. New’ Guinean). .-d ate CRU ALR oe a2

. Buds proportionally broader, + ovoid, ellipsoid(-oblong ); androphore about as long

as or shorter than the synandrium. [Androphore mostly pubescent, at least at base.
Lateral nerves not particularly closely set.] — Mainly E Malesia .......... 54

. Androphore glabrous. Lateral nerves comparatively closely set .........-- 53
. Androphore densely pubescent. [Buds 3.5—4 mm long.] Leaves pale below, lateral

nerves not particularly closely set. — Moluccas ............-+++. 2. M. alba

_ Leaves membranous; venation distinct below. Buds 2.5-3 mm long. Fruits 2.2—3.6

cm long, minutely scurfy. — NE Moluccas, Papua Barat (Bird’s Head).........
Eile genes benh Miers tah odes Set: Ge 80b. M. lepidota subsp. montanoides

_ Leaves chartaceous: venation faint below. Buds 4—6 mm long. Fruits 1.5—2.5(—3)

cm long, minutely scurfy. — Moluccas, W New Guinea ..... 78. M. lancifolia
Lower leaf surface whitish grey or glaucous, glabrescent, at first with thin indu-
mentum of which the stouter hairs leave minute scattered dark-coloured dots (lens!)
wel) vo igereh coptteel SO0kh Bert eel ly ME Oe An CA STE cai eee 3)

De Wilde — Myristicaceae (Myristica) 367

55a.

64a.

Twigs 2-4 mm diam. Leaves 20—40 cm long. Buds 5—6 mm long. — Philippines
(Mindanao), N Moluccas (Talaud Is.) ..........-2.--. eee eee 95. M. nivea
. Twigs 1.5—3 mm diam. (in M. impressa up to 5 mm diam.). Leaves up to tem
long. Buds 4(—5) mm long (only known in M. impressa, M. pilosogemma) .. 56
_ Leaf bud with hairs 1 mm. — Philippines ............ 107. M. pilosigemma
_ Hairs shorter. Three species from Sulawesi: for distinction see the regional key . . .
Pere PE NS 62. M. impressa, 74. M. kjellbergii, 75. M. koordersii
Basanore than > min long = See a eee aioe Git es oe eee 58
. Buds 2.5 mm long. — N Moluccas ......... 11b. M. bifurcata subsp. sulaica
. Buds + angular towards apex. — Philippines [incl. one fruiting collection from C
Sulawesi, Kjellberg 2990, discussed under M. impressa.| ..... 32. M. cumingii
Bemesnoraneilar 44-5): 2. ee ee ga ee ee ee 59
wiss 2-25 mm diam. or more... 27.52 eee 60
. Twigs 1—2(-2.5) mm diam. — Moluccas: W New Guinea: 4 0. c- ee 64
- Male flowers not known but, judged from the sessile female flowers, subsessile or
with short pedicel. — Lesser Sunda Islands .............-. 117. M. rumphii
. Pedicel about as long as the perianth ............-.---- see eee ee reese 61
. Bark of twigs soon cracking and/or flaking. Fruits 6 cm long, glabrescent; male
flowers not known. —Philippines (Basilan) .............--- 8. M. basilanica
_ Bark not or faintly cracking and flaking. Fruits smaller, with indumentum of minute
Hens be bP SA, MA AR Cel oct Red edn Sie ORS ot atch ible A he a elas 62
_ Leaves coriaceous. — S Taiwan, Philippines (N Luzon); mainly a coastal species
Renee mers in cet. « hig) tunel pain dient AEM, etd SAMO eee 18. M. cagayanensis
MIPeAVES Ee .ChartaceOls sure eae ee Es eae ik cae ae ae 63
_ Inflorescences much resembling a subsessile wart-like scar-covered brachyblast,

but possibly essentially paniculate. Fruits 4.5—6.5 cm long, powdery pubescent
with hairs 0.1 mm, or partly glabrescent. — Andaman and Nicobar Islands ......
perce CE PG FRY 0 OY patna M. andamanica Hook. f.

_ Fruits 2.5—4.5 cm long, with hairs 0.1-0.2 mm, sometimes partly glabrescent. —

Philippines (Mindanao), Moluccas .......-...---+++-++: 92. M. mindanaensis
Buds with hairs 0.1 mm. Pedicel about as long as the perianth. [Upper and lower
leaf surface + concolorous; lower leaf surface not papillose, venation indistinct.
Bracteole persistent. Fruits subglobose, 2.5—3 cm long, pericarp 4-8 mm thick,
hairs 0.1 mm.] — Moluccas, W New Guinea ...........----- 138. M. tristis

_ Hairs longer, 0.2-0.5 mm. Pedicel usually shorter than the Petlanthiy yeaa - 65
_ Buds cleft c. 1/3 to nearly 1/2. Inflorescences pedunculate. — N Moluccas, Papua

BatcUG bit Sead) hee eed welt acenlee cert: lla. M. bifurcata subsp. bifurcata

_ Buds cleft c. 1/4. Inflorescences all sessile. — SE Moluccas (Tanimbar), S New

rnineas Ni Australias ..<- sao! Seaeae Ooi dil de ae bere 70. M. insipida

_ Leaf bud and apex of young twigs with minute greyish hairs, glabrescent, lower

leaf surface with minute grey-silvery scale-like hairs less than 0.1 mm. [Fruits 4.5-
6.5 cm long, + glabrescent or with scurfy minute hairs 0.1 mm.] — Andaman and
micobatlslands) 24% dee oa Nane tes bak Sie te M. andamanica Hook. f.

_Indumentum more conspicuous, cinnamon or pale rufous, hairs (0.1—)0.2 mm or

niatews ee St). alduirey eee te menor bel 2a eee eee 67

Flora Malesiana, Ser. I, Vol. 14 (2000)

a. Twigs (2.5—)4-5 mm diam., bark lower down with lenticels. Leaves 20 cm long or

more.membranous or chartaceous. — E:; Malesia : ses scino4-4. .ceeebReee 68

. Twigs 14(—5) mm diam., if 4(—5) mm, then without lenticels. Leaves 20(—25) cm

long,onmless:....2..54 cnet he oy oun ioes sah agreed hyena 69

. Male flowers pedicellate. Fruits 4-7 cm long, with hairs 0.5—1 mm. — Philippines

(Mindanao), Sulawesi, Moluccas (M. fatua), Aru Is. (M. inutilis)..............
ean d chil hice PIE Be cn wares tart bh Ge 43. M. fatua, 72. M. inutilis

. Judged from female, flowers subsessile. Fruits 2.5—3.5 cm long, with shaggy hairs

1—1.5,mm.-—Lesser;Sunda Islands: ....., ..9....johs)steltevm 5 oe 133. M. sumbawana
. Twigs 1-2 mm diam. [Leaves membranous to thinly coriaceous. Buds 2—2.5 mm
long.] — W New Guinea, Aruls. .......... 80a. M. lepidota subsp. lepidota
. Bwigs 2=4(—S) mm diam. «2. 6.66 sec ee sont ss) ss sR en 70

. Twigs 2—3 mm diam. Leaves thinly coriaceous, base rounded or short-acute, nerves

above not distinct. Flowers insufficiently known. Fruits 2.5—3.5 cm long, with scurfy
hairs: 0.2—0.5,mm:— Bosneos 5 A+. tes: terete be naeeee Be 126. M. smythiesii

. Twigs 3—4(—5) mm diam. Leaves coriaceous, at base rounded or emarginate, or

—

sometimes short-acute; nerves above sunken, but distinct. Flowers with a conspicu-
ous collar of hairs at base of androecium, sterile apex minutely pubescent. Fruits
2.53.5 cm long, hairs woolly, 0.5—1 mm. — Sumatra, Borneo . 9. M. beccarii

REGIONAL KEY TO THE SPECIES — PENINSULAR MALAYSIA, SINGAPORE,
SUMATRA, JAVA (14 species)
(based on male flowering, female flowering and fruiting specimens,
with emphasis on vegetative characters)

. Inflorescences sessile, a densely scar-covered wart-like brachyblast; peduncle ab-

SENLOL Np LOL. Mim LONE: 4 «io. 2p bra oeip Seen: Cue, foam ne 2

, Inflorescences pedunculate, branched... . .:0...55 . <<< 5: + «/..- ee 4
. Leaf bud with hairs more than 0.2 mm. Mature leaves short-pubescent below. [Male

flowers: androecium + sessile, with a collar of conspicuous hairs around the base,
Slot apex, PUDESCENE |) «tesa chee ein a oie ca ane ee 9. M. beccarii

. Leaf bud with hairs 0.2 mm or less. Lower leaf surface finely papillose, early glabres-

3a.

TNE cahea cece pnren De oysnion Sooo deen east love os) oud hens ee cits See eas aioe nt eee ne 3
Leaves membranous to thinly chartaceous. Female buds (4—)5—6(—9) mm long;
lobes 0.4 mm thick. Fruits 3-4.5 cm long. Male pedicel 4-6 mm long. A species
close: tow. crassa: — CS. BE Vaviaticta.& §..4-scie see ee 135. M. teijsmannii

. Leaves chartaceous to thinly coriaceous. Female buds 4-5 mm long; lobes 0.20.3

4a.

mm thick. Fruits (3.5—)4—5.5 cm long. Male pedicel (5—)6—12 mm long. Plant gen-

erally‘stoiiter than JM: terjsmannii ts .\8 Ne oh tak 29. M. crassa
Réat bud with short hairs 0.1—0.2(—0.3) mmuoriess: 4....97). 0 4a. sae 5
Leaf budwith. hairs, more than ‘0:2 nm longs ys rs ae ee 13

Sa.

Lower leaf surface with (sub)persistent dense indumentum (hairs, even if small,
touching or interwoven; old leaves may be glabrescent). Fruits with scurfy hairs
Ou, MAMMOHA WIR Phe on eset annuus PERE em we seach yes Oe Renee 6

. Lower leaf surface glabrous (early glabrescent) or with + sparse, neither touching

nor interwoven, minute hairs. Indumentum of fruits variable, or fruits glabrous 8

De Wilde — Myristicaceae (Myristica) 369

6a. Buds (preferably to be seen in male flowers) + angular in the upper part. Twigs

9a.

10a.

lla.

ba.

bar

1.5—2.5(—4) mm diam. Leaves (8—)10—22 cm long, lateral nerves 10-17 per side.
limits 5=—9 cm. long, often subsessile:|osmi.a: eae, 21. M. cinnamomea

. Buds not angular. Twigs 2-8 mm diam., leaves 15—40 cm long, with 16—33 lateral

faeces nsidetan (Ad Ale Ohavrec leednl. daatane wa. ca eee 7

. Twigs (4-)5—8 mm diam. Leaves 16—40 cm long, drying dark brown, lateral nerves

(20—)23=33 per side. Fruits 6—9 cm long... ...02225655... 88. M. maxima

. Twigs 2-3 mm diam. Leaves 15-30 cm long, drying olivaceous, nerves 16—23 per

Se wRTtS. = /cmlong. 2 Ayaemie wisist Mateiseecals Qo Aaa. 35. M. depressa

. Fruits glabrous, or largely or almost completely glabrescent, or fruits inconspicu-

ously pubescent with remote or scattered (not densely set) minute hairs, in glabres-
cent fruits rarely some minute indumentum remaining in sheltered places near the
insertion of the pedicel. Lateral nerves c. 20 per side, or fewer ............. 9

. Fruits with persistent dense-scurfy indumentum, or fruits but partly glabrescent

and then always with distinct patches of (dense) indumentum in sheltered places of
the dried fruits, especially towards the insertion of the pedicel (hairs may be very
small). Lateral nerves (20—)23—30 per side; lines of interarching distinct. [Inflores-
cences widely paniculate. Twigs stout. Fruits (4.5—)6—9 cm long. Male buds 5—7
PPMNLONE Le aeesh ach apie) Chatboards fartdd ata ddmeerercieninveds 88. M. maxima
Twigs and leaves drying yellowish; lateral nerves 8—13(—15) per side. [Fruits dry-
ing dark brown or blackish, often with a green-yellowish exudate, 4.5—7.5 cm long,
glabrescent, at first with minute scattered hairs 0.1 mm. Buds angular in the apical
MAMA cons wpct dedauvescrtieds uailk 4 aobebse sopdetees 38. M. elliptica

. Twigs drying brown, leaves olivaceous or brown; lateral nerves 10 per side or more

Lines of interarching of the lateral nerves on lower surface of mature (older) leaves
raised and distinct. Male buds 3—3.5 mm long. Female buds 2—2.5 mm long, ovary
glabrous. [Twigs 2-4 mm diam. Lateral nerves 14—20 per side. Papillae on lower
leaf surface not apparent. Fruits 4-6 cm long, glabrous.] . .. 86. M. malaccensis

. Lines of interarching indistinct. Male and female flowers larger (female flowers

HopmieiownOLdl i 91@Gntea)).. oma! 44s Ohi bi ces a. Ss sae ee eee 11
Leaves (15—)20-35 cm long, drying a dull greyish brown; lateral nerves below
+ flat or but slightly raised; not papillose. Fruits 5.5—9 cm long, drying (dark) brown,
glabrescent, at first with hairs 0.1 mm, pericarp (5—)10—15(—20) mm thick.......
MBAS RK? coin tortie oimgh: Eig kts eho tyld ‘ttmemas+ atta weet 152. M. wyatt-smithii

. Leaves smaller, 7-24 cm long, drying olivaceous-brown; lateral nerves raised be-

low; papillose or not. Fruits 3.5—8 cm long, drying bright brown, glabrescent, hairs

woolly-scurty,0-1—O:3 MM... 6. se5 ---5 4-5 > ee ee eerie 12
Leaf apex acute-acuminate; lower surface not papillose. Fruits 3.5—6(—8) cm long,
penmcatpA—10 snmthickw.gieth.issedaeecnere anes ce cece = 66. M. iners

. Apex of leaves of fertile twigs acute or blunt; apex of leaves of sterile sapling twigs

(acute-)acuminate; lower surface distinctly papillose. Fruits 5—8 cm long, pericarp
hse Oamimthicka seiivmcusse Oo ere ere Roe - 56. M. gigantea
Leaves (12—)15—35 cm long, with persistent indumentum below; lateral nerves 15—
19(—25) per side. Leaf bud with hairs to 1(—1.5) mm. Fruits 2.5—4.5 cm long, hairs
dense, shaggy, 0.5-1 mm. — Mainly a coastal species, but sometimes inland in
destaded forest ead awoge Seas aga Saco ate ASE 59. M. guatteriifolia

370

Flora Malesiana, Ser. I, Vol. 14 (2000)

b. Leaves 7-35 cm long, glabrous (glabrescent) below; papillose; lateral nerves 10-

14a

18 per side. Leaf bud with appressed hairs 0.5—1 mm. Fruits 5 cm long or more,
glabrescent or with persistent indumentum ..... 6.66.66. eee eee eee 14
. Twigs 1.5—2.5(—3) mm diam. Leaves (of fertile twigs) small, 7—12(—15) cm long,
apex acute, blunt, or rounded; lateral nerves 10-15(—18) per side. Fruits 5-8 cm
long, (late) glabrescent, hairs scurfy, 0.2 mm. Flowers with hairs 0.2-0.5 mm....
BE AE RS nmr cic caw science ee os BATA sO AE Se cea 56. M. gigantea

b. Twigs (2—)2.5-6 mm diam. Leaves 12-35 cm long, the apex acute-acuminate; lat-

15a

eral nerves 13—22 per side. Hairs of fruits larger, 0.5 mm or more. Hairs of flowers
Ossie... ie we A ne Fee Bs HR 15
. Twigs (2-)2.5—4 mm diam. Leaves chartaceous or thinly coriaceous. Fruits 7-9 cm
long, glabrescent, hairs scurfy, 0.5 mm. Male buds 6-7 mm long 85. M. maingayi

b. Twigs 3—6 mm diam. Leaves coriaceous. Fruits (5—)6—7 cm long, with conspicuous

3a.

Aa.

Sa.

6a.

persistent shaggy hairs 1-2 mm. Male buds 44.5 long ......-. 84. M. lowiana

(3) REGIONAL KEY TO THE SPECIES — BORNEO (20 species)
(applicable for all fertile specimens; characters primarily vegetative)

_ Leaf bud with minute, appressed hairs, 0.1—0.2(—0.3) mm or less ......----- 2
. Blairs more than! 0.2mm. V2 90 SET ee a Co eee ee 14
_ Lower leaf surface with (sub)persistent dense indumentum (hairs may be very small),
old leaves sometimes glabrescent. Fruits with woolly scurfy hairs 0.1 mm. [Inflo-
rescences with distinct peduncle (often short in female).] .......-...-.++- 3
_ Lower leaf surface glabrous or early glabrescent or with scattered (not dense) minute
hairs. Fruits with variable indumentum, glabrous, or glabrescent ........... 5

Buds (best to be seen in male flowers) + angular in the upper part. Twigs 1.5-2.5
(—4) mm diam. Leaves (8—)10-22 cm long, lateral nerves 10-17 per side. [Fruits
5-9 cm long, often subsessile. Check also M. smythiesii.] . 21. M. cinnamomea
. Buds not angular. Twigs 2-8 mm diam. Leaves 15—40 cm long, with 16-33 lateral
nerves‘per side ns 220 ee. eee en ae 4
Twigs (4-)5-8 mm diam. Leaves 16-40 cm long, drying dark brown; lateral nerves
(20=)23-—33 per side. Fruits 6-9 cm long ..-...°...2...--=% 88. M. maxima
. Twigs 2-3 mm diam. Leaves 15—30 cm long, drying olivaceous; nerves 16-23 per
SIdesEnuiis is /-Cm loment-2" Ney, AIMEE 2 Oe Bere ees le eer eee 35. M. depressa
Inflorescences pedunculate; perianth largely glabrous (except in M. gigantea and
M. papyracea). Fruits glabrous, glabrescent, or with minute scattered hairs; in gla-
brescent fruits sometimes a minute indumentum remaining in sheltered places near
the msertion’of the pedicel >... 22.06 ae cee os: se: mn een ee 6
_ Inflorescences a sessile brachyblast, or pedunculate; perianth pubescent. Fruits with
persistent dense-scurfy indumentum, or partly glabrescent but with dense indumen-
tum remaining in sheltered places’... 0.000.220. 20. ee ‘15
Buds (especially in male flowers) sharply angular in the apical part. Twigs and
leaves drying yellowish; lateral nerves 8—13(—15) per side. [Fruits drying dark brown
or blackish, often with a green-yellowish exudate, 4.5—7.5 cm long, glabrescent,
hairs minute, scattered) 0.1mm: |/~ 2). 72 Ce Pee ee ee ee 38. M. elliptica
_ Buds rounded. Twigs drying (yellowish) brown, leaves olivaceous or brown, lat-
eral terves 10-30 per side. UGh ERR: OE MIN: YEAS cate verte ee eee 7

De Wilde — Myristicaceae (Myristica) 37]

Tae

b.
8a.

9a.

10a.

15a.

Lines of interarching of lateral nerves on lower leaf surface (of mature leaves)
Sms EITVCH It) eee ci: SRA AY, od bt. aed aii out dee See Bee ae 8
[eiges-of aterarching waised iandidistincty. v2 ance «-g.ee .20G) ee bet - 10
Bracteole (in male flowers) subpersistent or late caducous. Leaf apex acute-acumi-
nate; lower leaf surface not papillose. Fruits 3—6(—8) cm long, pericarp 3-10 mm
iielapeerna nya iiss iiss lst cowk iets eed ewer. cite eee 9

. Bracteole caducous. Apex of leaves of fertile twigs acute or blunt; lower leaf sur-

face papillose. Fruits 5-8 cm long, pericarp 15—20 mm thick .. 56. M. gigantea
Leaves membranous, chartaceous, or subcoriaceous, 7-24 cm long. Peduncle in
female to 1 cm long, 1-2 mm thick. Fruits 4-6(—8) cm long, pericarp 4-10 mm
thick, late or early glabrescent. Male buds 4—6(—-7) mm long ..... 66. M. iners

. Leaves thinly membranous, 7—14 cm long. Peduncle in female 1—2.5 cm long, 1.5

mm thick. Fruits 3-4 cm long, early glabrescent; pericarp 3-4 mm thick. Male
Fee OBI LONG oe cy so peye RR nrchalic nyc o Sars apes Sa hiypetic 41. M. fallax
Twigs 2-4 mm diam. Leaves about 3 times longer than broad, 10-30 by 4—9.5 cm,
apex acute-acuminate, base rounded to acute; lateral nerves 14—20 per side; lower
leaf surface papillose or not. Male buds 3—3.5 mm long. Female buds 2—2.5 mm
Mame a Eiuuits, 46, CO LONE: 3. 75 0. se dace ais Reet pence Spouse ings sac al 86. M. malaccensis

. Twigs 4-8 mm diam. Leaves large and comparatively broad, 16-44 by 8-14.5

(—18) cm, apex blunt or acute, not acute-acuminate, base + rounded or subcordate;
nerves (16—)20—28 per side; lower leaf surface papillose. Male buds 6-8 mm long.
Female buds 6 mm long. Fruits 7-9 cm long ............ 102. M. papyracea
Inflorescences (and infructescences) with a peduncle, and + branched ...... 12

_ Inflorescences sessile, a short, scar-covered sessile brachyblast. [Leaves chartaceous

Gr MITA CO RIAC SOUS oa a ek a 29. M. crassa

_ Leaves 16-40 cm long; lateral nerves 16-30 per side. Inflorescences large, widely

paniculate, 6-18 cm long in male, (1.5—)5—9 cm in female. Fruits 6-8 cm long 13

. Leaves 10-30 cm long; lateral nerves 13—20(—22) per side. Inflorescences 2-6 cm

long in male, 1.5—2.5 cm long in female. Fruits (4.5—)5—7.5 cm long. [Male buds
5 => TGS LO Tel ee ee eters eee aeons aeons eee 12. M. borneensis
Twigs (4-)5—8 mm diam. Leaves: lateral nerves (20—)23—30 per side; lines of inter-
arching + distinct; papillae usually apparent. Male buds 5-7 mm long ........-.
ee ee rrr ee a negate 88. M. maxima

. Twigs 2.5—5 mm diam. Leaves: lateral nerves 16-24 per side; lines of interarching

+ indistinct; papillae not apparent. Male buds 4(—4.5) mm long 40. M. extensa

. Twigs 5-10 mm diam. [Bark usually conspicuously cracking and flaking. ] Leaf

bud [and fruits] with very conspicuous villous hairs (1—)2—3 mm. [Inflorescences

S(CEST (EA | eet eee en ne a a 147. M. villosa
. Twigs 6 mm diam. or less. Leaf bud with hairs 1.5 mm or less ..........-- i)
Leaves pubescent below. Fruits 2.5—4.5 cm long .........--+.++-++++++-- 16
. Leaves glabrous below. Fruits 5 cm long or more ......-..-.---+++++++5 20

_ Inflorescences (best to be seen in male specimens) branched and with distinct

+ flattened main peduncle, with flowers in subumbels or spikes. Fruits 2.5—4.5 cm
Period Se PL SREP CSI MAE RIE LEE Pee oe eke My

_ Inflorescences (sub)sessile scar-covered short-shoots lasting several flowering sea-

sons. Fruits densely pubescent, 2.5—3.5 cm long. [Leaf bud with hairs up to 0.5(—1)
mm. — Inland species; not from limestone.] .......-......-..5.+0.005: 18

S72.

Flora Malesiana, Ser. I, Vol. 14 (2000)

18a.

a. Twigs 3—5(-10) mm diam. Leaves (12—)15—35 cm long; lateral nerves 15—19(—25)

per side. Leaf bud with hairs to 1(—1.5) mm. Buds rounded towards apex. Fruits
2.54.5 cm long, densely shaggy pubescent with hairs 0.5—1 mm. — Mainly a
coastal species, but sometimes inland in degraded forest .. 59. M. guatteriifolia

. Twigs 1.5-3.5 mm diam. Leaves 11-20 cm long; lateral nerves 10-18 per side.

Leaf bud with appressed hairs 0.5—1 mm. Buds sharply angular towards apex. Fruits
3.5—4 cm long, glabrescent or with thin scattered pale brown appressed hairs to
0.5 mm. — W Sarawak, NE Kalimantan; mostly from limestone..............
ee he Feet RE ee ee 123c. M. simiarum subsp. calcarea
Twigs 2-3 mm diam. Leaves thinly coriaceous, base rounded or short acute; nerves
not distinct above. Fruits 2.5—3.5 cm long, with scurfy hairs 0.2—0.3(-0.5) mm...
RS rec are acd serie te erations 126. M. smythiesii

. Twigs 3—4(—5) mm diam. Leaves coriaceous, base rounded or emarginate, some-

times short-acute; nerves distinct above. Fruits 2.5-3.5 cm long, with woolly hairs

OH lnm ee Pr eee Se oe ee Cte ce a pete als eee ae 19
a. Male flowers: androecium + sessile, with a collar of conspicuous hairs at base,
stenle apex’ pubesCelllnrtes sein tas ee hee hae ern. cee 9. M. beccarii

_ Androecium with distinct androphore, sterile apex without hairs. One collection,

SEEMNOLE S UNGEN oc ta ot te oe eps si erate cee 43a. M. fatua subsp. fatua

_ Lower leaf surface concolorous with upper surface, not papillose. Fruits glabrous

(early elabrescemt) - 2.0006 «= Gg oe coe oe cig se ih

_ Lower leaf surface distinctly papillose, generally paler than upper leaf surface. Fruits

either with persistent indumentum, late glabrescent, or early glabrescent in M. gi-
OOMILEE an spe 55 (oe ie SI gin A ne Saar paps

21a. Leaves membranous, up to 24 cm long, base attenuate. Twigs up to 4 mm diam.,

smooth or striate, the bark of the twigs lower down cracking or not; leaf bud slen-
der, 7-12 by 1-2 mm. Fruits to 8 cm long, pericarp S—10 mm thick. — Stout forms,
mainly from NEBOrmeOe oe oo acta eis weit ait eke eek ae 66. M. iners

_ Leaves chartaceous, 15-35 cm long, base broadly rounded. Twigs (4—)5—6 mm

diam., bark early conspicuously cracking; leaf bud stout, 10-15 by 4-6 mm. Fruits
7.5-9.5 cm long, pericarp 15-20 mm thick. — NE Borneo ... 28. M. corticata

. Twigs 1.5—2.5(—3) mm diam. Leaves 7—12(—15) cm long, chartaceous, apex acute,

blunt, or rounded; lateral nerves 10—15(—18) per side. Fruits 5—8 cm long, (late)
glabrescent, hairs scurty, 0.2. MT, 9322). og aerator eae me foey ne 56. M. gigantea

. Twigs 3-6 mm diam. Leaves 14-35 cm long, coriaceous, the apex acute-acumi-

nate; lateral nerves 13-22 per side. Fruits (5-)6—7 cm long, with conspicuous per-
sistentishagey Mains al 24min ©. 2... oy pee ee ee ee ee 84. M. lowiana

(4) REGIONAL KEY TO THE SPECIES — LESSER SUNDA ISLANDS (3 species)

la.

b.

ple\.

b.

Inflorescences or infructescences short peduncled. — Bali 59. M. guatteriifolia

Inflorescences:séssile}.ciechis eens ead: My eine 0 eee 2
Leaves with persistent dense indumentum below. Fruits 2.5—3.5 cm long, hairs
shaggy, (1—)1.5 mm. — Sumbawa, Flores ............. 133. M. sumbawana

Lower leaf surface glabrescent. Fruits 2.5—4 cm long, hairs 0.5 mm. — Not in Bali
GrlLombok- Wilt O08 See. ao: 1. SERGE FAR Sa 117. M. rumphii

De Wilde — Myristicaceae (Myristica) a3

la.

Sa.

6a.

Ta.

la.

(5) REGIONAL KEY TO THE SPECIES — SULAWESI (8 species)

(applicable for all fertile specimens)

Leaves (15—)20—40(—50) cm long; lower surface with persistent short dense indumen-
tum. [Inflorescence a (sub)sessile, short brachyblast. | 43b. M. fatua subsp. affinis

. Leaves generally smaller; lower surface glabrous, glabrescent, or with minute sparse

PTE CELE eee nd rn here Game). erm@dikosats a lard ee. Eee Be 2

_ Inflorescences (infructescences) with a peduncle. Buds angular in apical part. Fruits

glabrescent or with sparse greyish hairs .... 123b. M. simiarum subsp. celebica

_ Inflorescences sessile or pedunculate. Buds rounded in apical part, not angular (of

some species male flowers not known). Fruits with (minute) rusty indumentum 3

Mionesue> mint diam) «aht.te litem Ieee eke ny sale ak Bae be 4
. Twigs generally thicker, 1.5—S mm diam. Inflorescence a (sub)sessile brachyblast,

peduncle absent or to 2 mm long only ....... 2.6... ee eee eee eee eee 5)

_ Leaves membranous, 9-19 cm long. Inflorescences slenderly pedunculate, some-

times subsessile. Flowers small, conspicuously rough-hairy. Fruits not known ....
Re She VAlel nb sales Aelia See 63. M. impressinervia

_ Leaves coriaceous, (2.5—)4—7.5 cm long. Inflorescences sessile. Flowers not known.

Fruits + ellipsoid, 2-2.5 cm long. — Ultrabasic soils of Central Sulawesi........
eth idl coe. tae cries a eee Ee 140. M. ultrabasica
Twigs 4-5 mm diam. Leaves coriaceous, lower leaf surface not whitish, not minutely
dotted (lens!). [Fruits subglobose, 5 cm diam.; pericarp nearly woody, 10 mm thick. |
— Central Sulawesi; soils on conglomeratic and ultrabasic bedrock; 400 m.......
Rees Bilbao coterie one! alu nes eerie bp ae wosl..2 36. M. devogelii

. Twigs 1.5—5 mm diam. Leaves membranous, lower surface conspicuously pale,

munutely:dark-dotted: (lens!) uss .\Mewwe 2261: Sa OG A Ss ee eRe SEE 6
Twigs 4-5 mm diam. Female flowers and fruits (sub)sessile. Fruits subglobose,
2-2.5(—3) cm long; pericarp 1-3 mm thick, hairs 0.1—0.2 mm. [Male flowers not
icnenune|— NC and!SW Sulawesi :20m). a= ceo ouse toe 74. M. kjellbergii

. Twigs 1.5—4(—5) mm diam. Female flowers and fruits stipitate. Fruits + ellipsoid,

3-4 cm long; pericarp 3—-6(—8) mm thick, hairs 0.1 mm .......--...+++++- if
Twigs 1.5—3 mm diam. Fruiting pedicel 6—10 mm long, conspicuously pubescent,
hairs 1-1.5 mm. [Male flowers not known.] — NE Sulawesi (Minahasa))sise She:
PP xtra LnondgbHle wtlew! or. 8G. S eeu eee 75. M. koordersii

. Twigs 2—4(—5) mm diam. Fruiting pedicel 3-5(—7) mm long, glabrescent or with

hairs to 0.5 mm only. — Sulawesi (except Minahasa); incl. Kjellberg 2990.......
PRAg AiG He yeeros Tesco Olsb atte Mie AS Abend, Set say Somes th 62. M. impressa

(6) REGIONAL KEY TO THE SPECIES — PHILIPPINES (20 species)

(applicable for all fertile specimens; characters primarily vegetative)

Lower leaf surface with a persistent dense indumentum; whenever the hairs are small
fey are densely-interwoven) ... -iawsals shah) eee ae eee tee 2

- Lower leaf surface either with + spaced silky hairs, with a very minute cobweb-like

covering (to be seen with a lens only), glabrous, or glabrescent ........----. 7

Flora Malesiana, Ser. I, Vol. 14 (2000)

6a.

Ta.

9a.

10a.

ile.

Hairs of leaf bud (0.1—)0.5—1 mm. [Inflorescences mostly pedunculate. Fruits 2.5—
4 cm long, with conspicuous shaggy hairs 0.5—1(—2) mm.] 59. M. guatteriifolia

, Hairs of leaf bud shorter) .ccacos. saeccwinl. galls Aaa. ok ee 3
. Twigs (3—)4—8 mm diam. Leaves (15—)20 cm long or more. Inflorescences a (sub)-

sessile scar-covered wart-like brachyblast. Fruits with minute or longish hairs, to

heidi: sie Baeettiade.. tila atetiaes Soe ye 4
. Twigs 1-2.5(-4) mm diam. Leaves (5—)8—22 cm long. Inflorescences pedunculate.
Fruits with short'scurty-hairs, 0:1=0:2 mom) | 1.23 0e8 Se ee a ee 6
. Leaves coriaceous, oblong-lanceolate, (15—)20 cm long. [Petiole proportionally
long, 20-35 mm.] — NE Luzon: Palanan area .......... 24. M. colinridsdalei
. Leaves herbaceous or chartaceous, elliptic-oblong, 15-50 cm long ......... 5

. Twigs 5-8 mm diam. Leaves tapered (narrowed) in the lower half, nerves 30-35

per side; lower leaf surface with whitish dense felty hairs 0.1 mm or less, thus
seemingly glabrous (lens!). Fruits (5—)6-6.5 cm long ....... 150. M. wenzelii

. Twigs 3-4(—6) mm diam. Lower leaf surface with yellowish brown hairs 0.1—0.2(—

0:5) mame Eruits:3.5=6:5(E7 icmilonsy see eee 43. M. fatua subsp. fatua
Lower leaf surface with minute persistent hairs. Midrib below brown or yellowish.
Fruits 5—9 cm long. — Probably Mindanao ............ 21. M. cinnamomea

. Lower leaf surface late glabrescent. Midrib below purplish, contrasting with grey-

ish surface. Fruits 3-7 cm long. — Palawan, Mindoro ....116. M. rubrinervis
Buds in apical part sharp-angular. Inflorescences mostly pedunculate and branched.
Fruits rather small, 1.5—3 cm, with scattered hairs or glabrescent..............
athe OMe ls ash 7a 123. M. simiarum [subsp. simiarum and subsp. celebica]

. Apical part of buds not or not sharply angular. Inflorescences sessile or peduncu-

lafex(and! branched)? Fiitsivariabletsne em. eee, a ee ee 8

. Leaves chartaceous or coriaceous, 10—24(—30) cm long; petiole proportionally long,

20-35 mm. Inflorescences (infructescences) sessile. Fruits large, (S—)6—7 cm long
[with minute dark rusty scurfy indumentum] ...............2.-.-eeee 2

_ Leaves coriaceous, chartaceous, or membranous; petiole proportionally shorter.

Inflorescences sessile or pedunculate. Fruits large, or smaller ............. 10
Leaves coriaceous, | 1—24(—30) by 3-6 cm; lateral nerves (15—)20 per side. Twigs
4-6 mm diam: — NE Luzon: Palanan area ............. 24. M. colinridsdalei

. Leaves chartaceous, 10-15 by 2.5—6 cm; nerves c. 10 per side. Twigs 2-3 mm

diam. Petiole proportionally long — Luzon, Biliran..............-.+..+505-
We Anacapa. Baek ce dstens) A. ERM haere 82. M. longepetiolata
Twigs 3—5(—10) mm diam. Leaves large, to 45 cm long. Inflorescences or infruc-
tescences with a peduncle. Fruits large, (4—)5—8 cm long, with persistent indumen-
111. eee ao ane enw: POMPE Serr eto en roles caine bacco S.c.6 ooo 11

. Twigs 1-5 mm diam. Leaves generally smaller, to 20(—30) cm long, and inflores-

cences pedunculate or (sub)sessile brachyblasts. Whenever fruits are large, they
are\glabrescentw, siiwiaieexeriosuctoeneetage welage Hes k)- ack Geer 1,
Leaf bud with hairs 1(—2) mm. Fruits with hairs 0.3-O.5mm.................
RU REAPER, ASRS Ws LEMAR OU POND Soe e Re e ageds ree 105. M. philippensis

_ Leaf bud with hairs 0.1—0.2(—0.3) mm. Fruits with scurfy hairs 0.1 mm. [Female

flaweEs Hot koi] L AAW 08 MOO A DE I eR Re 51. M. frugifera

De Wilde — Myristicaceae (Myristica) BID

iar

a:

Dias

Twigs 1-2 mm diam. Fruits 3.5—4 cm long, indumentum minute, soon glabrescent.
Lower leaf surface distinctly papillose. Bracteole in female flowers well below the
perianth. — Palawan .......... 0.6 ee eee eee ee eee eee 141. M. umbellata

. Twigs 2-5 mm diam. Fruits pubescent or glabrescent. Lower leaf surface papillose

Graet. Bracteole at apex of pedicel 00.2... ee ne ee oie 6)
Leaves lanceolate, lateral nerves 20-30 per side, flat or slightly raised below (not
to be felt with the finger); very distinctly papillose (lens!). Pericarp 10 mm thick
one ne snc ES beg RR REAL COTS CLS Mes 3 eran Pa 77. M. laevis

_ Leaves elliptic or lanceolate, lateral nerves c. 20 per side or fewer, generally raised

below; papillae absent or present, conspicuous or not (lens!). Pericarp usually less
PEAMRUMIIIIICK ce cas State sc ons eek oe oe he oe eames eee ee 14

_ Nerves c. 45° or less to the midrib. Inflorescences (best to be seen in male speci-

mens) pedunculate, the flowers in subumbels ............----++++s5505- 15

_ Nerves c. 45° or more to the midrib. Inflorescences with few flowers at the end of

a sessile scar-covered, wart-like brachyblast lasting several flowering seasons 17

_ Nerves (45—)60° to the midrib. Lower leaf surface not papillose. — Mindanao:

Simmeao (Wenzel I93.7)\). \atiats tis Ne chi se sek Rin 05 -)s sale ee oe 66. M. cf. iners

_ Nerves less than 60° to the midrib. Lower leaf surface + papillose ......... 16
_ Fruits 3—4.7 cm long, with dull grey-brown scurfy indumentum. Midrib on lower

leaf surface (yellow-)brown when dry, not much contrasting 1. M. agusanensis

_ Fruits 3-7 cm long, with bright dark rusty or orange-brown scurfy indumentum.

Midrib on lower leaf surface drying dark purplish or reddish, much contrasting . . .
SE Bernstein es 222 )8 fel tall i AOS Shia th, 116. M. rubrinervis

_ Leaves below whitish, contrasting with upper surface; lower leaf surface minutely

dotted by the scars of fallen larger hairs ..........-.. 06s sees eee eeeeees 18

_ The two leaf surfaces + concolorous or not, not dotted ..............-.-- 19
. Twigs 1.5-2.5 mm diam. Leaves 20 cm long or less. Fruits ellipsoid, 3.5 cm long,

with conspicuous fine-woolly hairs 0.5(-1) mm. Hairs of leaf bud 1 mm. — Min-
EONS AA 2. 59, LO Acs aod on O44 oer 107. M. pilosigemma

. Twigs 2-4 mm diam. Leaves 20—40 cm long, greyish below. Fruits ellipsoid, 3.5—

4 cm long, hairs 0.1 mm or less. Hairs of leaf bud 0.1 mm. — Mindanaog...2 <i.
pee wart: bjies 0) Stee. Deion Show. oe Sonepat 95. M. nivea

. Bark of twigs soon cracking and/or flaking. Fruits 6 cm long, glabrous ( glabrescent).

[Male flowers not known.] — Basilan ...........--.-++++- 8. M. basilanica

. Bark not or only slightly cracking and flaking. Fruits shorter with minute indumen-

MIKES. ca ae sone por Rol doe §) aoekéo. - ei Bie cet shee cee 20

. Twigs 3-4 mm diam., lower down densely set with lenticels. Leaves coriaceous.

[Fruits subglobose, 3—4 cm long, hairs 0.1—0.2 mm.] — Mainly a coastal species;
S Taiwan, Philippines (N Luzon) .........--.+-++++5- 18. M. cagayanensis

. Twigs (1-)1.5—3 mm diam.; lenticels present or not. Leaves membranous or charta-

BEOIISN RUN oe tos Pir Rett o Pitan nnd CBee Se SERIES Oo See ae ys
Lenticels of twigs few and inconspicuous. Fruits subglobose or ellipsoid, 3.5—5 cm
long, hairs 0.1 mm. Leaves 8.5—23(—28) cm long ......------ 32. M. cumingii

. Lenticels present, distinct or not. Fruits ellipsoid, (2.5—)3-3.5(—4.5) cm long, hairs

0.1-0.2 mm. Leaves 14-35 cm long. — Mindanao ...... 92. M. mindanaensis

376

Flora Malesiana, Ser. I, Vol. 14 (2000)

6a.

ae

8a.

(Ja) REGIONAL KEY TO THE SPECIES — MOLUCCAS (20 species)

(based on male flowering specimens)

. Male inflorescences branched, with a smooth main peduncle sometimes subsessile,

slender, simple or branched; inflorescences lasting one or a few flowering seasons
OND es fies Shove apes tus wre ict « adakasyaert hE Ade eyes (eee o stake ae res 2

_ Inflorescences (sub)sessile densely scar-covered simple or 2—4-furcate thickish

woody short-shoots, lasting several flowering seasons, each season producing a
number of new flowers at the apex; peduncle (non scar-covered part) absent or up
G2 PITTA OIA Sc he ace ape aga Niece sy ac aelina cy 4042 peice faa 7

. Plant wholly conspicuously glabrous. [Male flowers not known.] — Seram......

ee re ene oe TE Cee ee heen me ee 104. M. perlaevis

-Indumentum obvious on terminal leaf bud 92. ss re. ics ene dye ee 3
. Buds narrow, 6-8 by 2—3 mm, the apical part markedly angular, apex (sub)acute.

—N Moluccas (Sula Is., Bacan, Buru) .... 123b. M. simiarum subsp. celebica

_ Buds variable, not. ancular, apex rounded: 2. cicc.e. 5 + ppg: cere 4
. Buds 3—4.5(—5) mm long, [rough-pubescent. Nerves on upper leaf surface flat or

impressed, lower leaf surface papillose.] — N Moluccas...........-........-
et, aerate ct ng RNa MAVEN iB LANEY 7 ran eI 11. M. bifurcata (with 2 subspecies)

Y Buds.5—lk min TOMS os. se au 527 lone G's cei ens) eos ga eee 5
. Buds conspicuously pubescent. Leaves large, 25—45 cm long, lateral nerves 25—30

per side, impressed above. Bark of older twigs longitudinally cracked. — N Moluc-
CAS SAGA ce ts <, sereatgrae ena Le hcicis ea Acat lene emememe cena cae 47. M. fissurata

. Buds either glabrous, glabrescent, or short-haired. Leaves smaller, 6-22 cm long,

lateral nerves fewer, above flat or but little impressed .................... 6
Pedicel 1 mm thick. Flowers thinly pubescent. Leaves subcoriaceous, 9-22 cm long,
lower surface with minute indumentum, late glabrescent. Twigs 2—3.5 mm diam.
mG NI@IUC CAS teen Ona tae at rere eas a eine eee eerie 131. M. succedanea

. Pedicel less than 1 mm thick. Flowers glabrescent with sparse, very minute indu-

mentum. Leaves chartaceous, 6—13 cm long, early glabrescent. Twigs 1-2 mm diam.
— Origin Banda; known only from cultivated specimens .... 50. M. fragrans
Buds l0oS]j13—15 mmlong’s. “Fa... cee eee eee es «2: 8

PBudstless than lO mimblone = A. PY MAS. eee ee ee soe 9

Twigs ridged or winged; inhabited by ants. — SE Moluccas (Kai Is.) and New
Guneae ae eee EE oy BE eee ee crate etree 129. M. subalulata

b. Twigs not winged. — N Moluccas (Obi, Halmahera) ...... 110. M. pubicarpa

Me

. Twigs 4—7(—10) mm diam. Leaves (15—)17—40 cm long ................- 10
. Twigs 1—4(—5) mm diam. (4-5 mm in some material of M. fatua and M. inutilis).

Leaves generally much shorter than 40 cm (15—40 cm in M. nivea) ........ 12

. Lower leaf surface early glabrescent or with scattered, minute, scale-like hairs less

than 0.1 mm (lens!). [Male buds 8—8.5 mm long.] — N Moluccas (Bacan).......
Ne a a oe) | Seg a eet ee ey Pe oe emery ce 114. M. robusta

. Lower leaf surface densely tomentose (hairs may be minute). [Two species, distin-

guishable:mainly by fruit characters.] .2: 00" ee I 1]
Buds @munlong.—"N Moluceas) ro ne es Se 118. M. sangowoensis

. Buds 4—6 mm long. — Moluccas (and Sulawesi, Philippines)...............--

Select be eae RC LE seas hie. eres ote ROAR Rey ee ee 43a. M. fatua subsp. fatua

De Wilde — Myristicaceae (Myristica) ST

1Qayz

Lower leaf surface densely and persistently pubescent, the hairs may be short and
Spat caieR ems pee ake. Wt a). he ok. AAR se ee Ihe)

. Lower leaf surface glabrous, early glabrescent, or with minute scattered hairs 15
. Twigs (2.5—)4-5 mm diam. Leaves 20 cm long or more. Buds 4-6 mm long . 14
. Twigs 1-2 mm diam. Leaves 15 cm long or less. Buds 2—2.5 mm long. —Aru Is.

PNIERS. or} ouria ils) vals alma wlelecloteeni se. 80a. M. lepidota subsp. lepidota

. Bracteole caducous; buds 4.5—6 mm long ........ 43a. M. fatua subsp. fatua
»Bracteole persistent; buds 3=4-5 mm long. —Aruls. .......+.2-:.22555.2-.

re Per eth ts Glee k CII R OS, eek 72b. M. inutilis subsp. papuana

_ Lower leaf surface with scattered dark brown dots and dashes (lens!). — Aru Is.

re es Wb 98 1h 6 IR ee LISS RTS ie Bilibe Diy oa 122. M. scripta
. Leaves without larger dots; small dots present or absent ................- 16
. Leaves pale below, with minute dots representing hair scars. — Known in the Moluc-
cas only from one sterile collection from Talaud Is. ............. 95. M. nivea
. Lower leaf surface not conspicuously pale, without dark dots (lens!) ....... le
. Buds cylindrical, narrow; androphore about as long as the synandrium ..... 18

. Buds ovoid or ellipsoid(-oblong); androphore about as long as or shorter than the

synandrium, mostly pubescent, at least at base. [Lateral nerves not particularly
FISTS MMOS SENT eet hhts HE MAIR. MIAESS ORL, MBAR RM Rs ae Na, f ES. SEEN 20

Pe iule@phonerelabrOUS 77.20 4.12 5 NOM. Siiette Oe Nel ae ree fee orl 1)
. Androphore densely pubescent. [Buds small, 3.5—4 mm long. Leaves pale below. ]

Sa NeVioliccas,(Bacan. Obi, Burl)).2s 2.540626 00. 28 ta cae tee 2. M. alba

_ Leaves membranous; venation distinct below. Buds 2.5—3 mm long. — NE Moluccas

TN Loch tiie a prtenionn « aoe hora 80b. M. lepidota subsp. montanoides

. Leaves chartaceous; venation faint below. Buds 4-6 mm long............-.-.

<n o aighch gh bit Aiki ME OA I BS 78. M. lancifolia (with 2 subspecies)

. Buds 2.5 mm long. — N Moluccas (Sula Is., Halmahera, Obi)..............-.

PN TS LMC bathe peAchstsenae a Styad Bids. oo Lear 11b. M. bifurcata subsp. sulaica

MEOe SMiOreltiall Ss Mind LONG 24 oe. Cae. earcle ewtnel wee rete © eee ore eRe Dill
Pi iase2—2 Sma diam, OF MOLE) sede eee er 92. M. mindanaensis
eae sal 2 (245) naman “5.00 2 EI Rie ec eee ce eee eercraes up

. Perianth with hairs 0.1 mm. Pedicel about as long as the perianth. — N Moluccas

ee ett AYER ELECT SU ACRY RA IE OBER AGI DY ORL GEL no ER Neer 138. M. tristis

. Perianth with hairs 0.2—0.5 mm. Pedicel usually shorter than the perianth ... 23
. Buds cleft about 1/3 to nearly halfway. Inflorescences sessile or pedunculate. —

NPN Ole@AS: (eee) MAL Res Oe 11a. M. bifurcata subsp. bifurcata

. Buds cleft c. 1/4. Inflorescences sessile. — SE Moluccas (Tanimbar)...........

PO 503 bE de cssonihes vo nd dese ora wodcmennatige adnan tek NT Ra 70. M. insipida

(7b) REGIONAL KEY TO THE SPECIES — MOLUCCAS (20 species)
(based on female flowering and fruiting specimens)

. Lower leaf surface with persistent dense indumentum; whenever the hairs are short

they may be densely interwoven and thus the leaf may seem glabrous ....... 2

. Lower leaf surface either glabrous, glabrescent, with sparse minute scattered hairs,

or with a very minute cobweb-like covering (only visible with a lens) ....... 5

Flora Malesiana, Ser. I, Vol. 14 (2000)

Ila.

. Twigs 1-2.5(—4) mm diam. Leaves 4.5—15 by 1.5—6 cm. Fruits 2.5-3 cm long, with

scurfy hairs 0.1-0.2 mm. —Aru Is. ......... 80a. M. lepidota subsp. lepidota

. Twigs 3-6 mm diam. Leaves 20 cm long or more. Fruits either with distinct hairs,

0:5'=liimm! or hairs seunfy 6.857! SNS SOE Ss. RIO LO a a a 3

_ Fruits 4—6.5(—7) cm long, with conspicuous rusty hairs 0.5—1 mm. [Two taxa, with

a different distribution, and distinguishable mainly by characters of the male flow-
Sree ION PE PN cb oe A oie LE, ee 4

_ Fruits 5—8.5 cm long, hairs dull cinnamon, scurfy, 0.1—0.2 mm. — N Moluccas. .

CPMEE MEOILE PAPA IE. ae Set erect Nt Oe ee eS Ste 118. M. sangowoensis

a. Male buds 4.5—6 mm long; bracteole mostly caducous .........-.--+0+-0505-

PE RN ey VE SE Ree ESS ERS 43a. M. fatua subsp. fatua

. Male buds 3—4.5 mm long; bracteole persistent. — Aruls. ..................

|, aI echt ET Reese Sete peed lt ns Gate ajar tteen ion 72b. M. inutilis subsp. papuana

. Twigs distinctly ridged or winged; myrmecophilous. [Fruits subsessile, more or

less ovoid, 2 cm long, apiculate, minutely tomentulose.] —Kails. ............
SPs tegecaes Pass dine es Eicas CRC k ene aes Or 129. M. subalulata (with 2 subspecies)

. Twigs neither ridged nor winged; not myrmecophilous ...........-++++++- 6
. Plants conspicuously glabrous. Flowers not known. — Seram.............++5

. Indumentum present on terminal leaf bud, twig, and flowers .............. 7
_ Leaves 6—17(—25) cm long, lower surface with scattered dots and dashes. — Aru

| Sunt nie 4d Rea een ten oes resets we Pinan PES ta PERE RUE Ec eee 122. M. scripta

_ Leaves of various sizes, without dots and dashes ...........-.-. +++ +eee: 8
_ Female buds much narrowed towards the acute apex, the apical part sharp-angular

im Cross section. Leaves, to 30cm long ....-..---2-se ss | | >

. Apex of female buds narrowed or not, in cross section circular or at most bluntly

angular, but not sharply angular. Leaves small or large, to 40 cm long eee 10

_ Buds 5—6 mm long. Leaves below glabrous or with minute scattered hairs. Fruits

3—3.5 cm long, with scattered hairs 0.3—0.5 mm, or glabrescent. — N Moluccas
(Sulads-sBacane Buti ic 2 a! ar tens ote 123b. M. simiarum subsp. celebica

_ Buds 10 mm long. Leaves below with dense or scattered scale-like hairs 0.1—0.2

mm. Fruits 5-6 cm long, densely dark brown short-pubescent, hairs 0.2-0.3 mm.
=— NUMOlUCCaS tals aatesste anlar eee «see tale aoe 110. M. pubicarpa

. Twigs 3—5(—10) mm diam. Leaves 25—45 cm long ........--++++++++++> 11
. Twigs 1-5 mm diam. Leaves mostly less than 30 cm long (up to 43 cm long in

M. nivea). [Fruits 2-5.5 cm long, pubescent, or fruits larger but then glabrous or
earlpelabresent. |yccia- athens thm Aloe Oh hotel oe HP eee 192
Twigs 4-7(-10) mm diam.; bark of twigs conspicuously longitudinally cracking,
+ flaking. Lower leaf surface papillose (lens!). [Fruits large, (4—)5—8 cm long, with
persistent scurfy hairs 0.1—0.2 mm.] — N Moluccas (Bacan).......---+++++--

. Twigs 5-10 mm diam.; bark of twigs striate, at most finely cracking and finely

flaking. Lower leaf surface not papillose. [Female flowers and fruits not known. ]
EDENTINIGIUCCAS (EAC AIT re ee Pare sepa eters che ie 114. M. robusta

De Wilde — Myristicaceae (Myristica) 37/2)

12a.

Fruits glabrous or largely early glabrescent, hairs 0.1 mm or less. Inflorescences
delicate, few-flowered, without or with a slender peduncle. [Leaves elliptic, with
FERVMALCLAleerVes PEkISIGes|.% 2, TORRES. WR. Pee Ree ede b3

. Fruits with persistent indumentum, hairs may be very short and inconspicuous. In-

florescences with or without a peduncle, with the flowers in a woody scar-covered
DSRESOOLNY Soa oe dieow a ee abeutedadlecwwaw AB MACRUDRL Sanate SNE eS 14

. Twigs 2—3.5 mm diam. Leaves + coriaceous, 9-22 cm long, below late glabrescent.

Fruits 4.5—8 cm long. — N Moluccas ................-. 131. M. succedanea

. Twigs 1-2 mm diam. Leaves chartaceous, 6-13 cm long, below early glabrescent.

Fruits 4—6 cm long. — Cultivated (origin Banda) ........... 50. M. fragrans

. Leaves pale below, with scattered dark-coloured small dots, representing hair scars

(lens!). — Known in the Moluccas only from a sterile collection from Talaud Is.
Re 8 96 15). ceo fi etd eecold Josh keels csepgcndb vane fh 95. M. nivea

. Leaves not conspicuously pale below, without scattered dots ............. 15
. Fruits 3—4 cm long, with conspicuous coarse hairs, (0.5—)l mm...........--..

Ud aerate Ou haat elite. aeadh 11a. M. bifurcata subsp. bifurcata

_ Fruits variable in size, hairs mealy, scurfy, or woolly, shorter, 0.1—0.5(—0.8) mm

Rea Bort Re OF), VTS TEE. NAGI GN ope NIRS, LE are ey ie 16

. Lower leaf surface greyish or whitish, glabrescent, indumentum weak, its stoutest

hairs leaving numerous regularly spaced, small, dark dot-like scars (lens!). Fruits
short-ellipsoid, 3—4.2 cm long, hairs 0.1—0.5 mm; pericarp thick and woody, 5 mm
fine eNO NI@IUTICCAS, 2 ud-205) Niatetteeet say ae O alee MID TAN case 2. M. alba

. Upper and lower leaf surface more concolorous, or, whenever greyish or whitish,

WAIN Ute STINE OLS RTRs chitosan nce Tae ERO iba bh cake re, Au che teste Ly,

. Leaves elliptic to oblong-lanceolate, 14-35 cm long. Fruits ellipsoid, (2.5—)3—-3.5

jroconlons: hairs O1—0.2.mmi wie. Je ee 92. M. mindanaensis

. Leaves generally smaller, elliptic to lanceolate, to 20 cm long. Fruits variable 18
. Lower leaf surface not papillose (lens!). Fruits subglobose, 2.5—3 cm long; pericarp

4—8 mm thick, hairs 0.1 mm or less. — N Moluccas .......... 138. M. tristis

. Lower leaf surface papillose. Fruits ellipsoid; pericarp 1-5 mm thick, hairs short or

long, 0.1—-1 mm. [Four taxa, difficult to segregate without male flowers.] ... 19

. Male inflorescences (short) pedunculate; male buds 2.5 mm long. [Fruits ovoid-

ellipsoid, 3—4 cm long, with scurfy hairs 0.2 mm.] — N Moluccas (Sula Is., Halma-
ELAN O@ DI) Jf) 5. NG LORS oR OR ee, SEL 11b. M. bifurcata subsp. sulaica

. Male inflorescences (sub)sessile; male buds 3-6 mm long ............... 20
. Leaves membranous. [ Venation on lower leaf surface distinct. Fruits 23.5 cm long,

hairs 0.1(—0.2) mm.] Male buds small, 3 mm long — NE Moluccas............
MEL AER Aish ATL ad, nie SBS 6 wy osserhe evi aces 80b. M. lepidota subsp. montanoides

. Leaves chartaceous. Male buds 4—6 mm long ...........-.-.-.+-+-+-+-: 21
. Venation on lower leaf surface generally indistinct; lateral nerves 8—20 per side,

rather dense; blade 4-17 cm long. Fruits 1.3—2.6(—3) cm long, hairs 0.1 mm.....
10. 6 RSET AD NR ee AEN. Be, TPR AINE 78. M. lancifolia (with 2 subspecies)

_ Venation distinct or not; lateral nerves more lax, 6-12 per side; blade 7-24 cm

long. Fruits 2.5—3.5(—3.8) cm long, hairs 0.3—1 mm. — SE Moluccas (Tanimbar
IGs)ineatot aii tual eis we eh aaa ees as teen nth BR 70. M. insipida

Flora Malesiana, Ser. I, Vol. 14 (2000)

3a.
. Twigs 1-2(—3) mm diameter; most leaf blades 15(-20) cm long or less ....... °
4a.

Sa.
_ Inflorescences with peduncle 5 mm long or more. Leaves membranous, chartaceous,

6a.

ee

8a.

(8) REGIONAL KEY TO THE SPECIES — NEW GUINEA (99 species)

(based on male flowering, female flowering, and fruiting specimens)

_ Lower leaf surface with scattered brown or blackish dashes or dots (lens!) .... 2
_ Lower leaf surface not dotted (including all specimens with lower leaf surface cov-

ered with dense indumentum), < os .. «cals cu wunG les we « Seperate eh. ER 22

_ Male buds before anthesis with hooded bracteole, 8-9 mm long, enveloping the

perianth, then caducous (compare also M. brassii). [Female flowers not known.] —
Papua New Guinea; (250-)800-1000 m ........---...--005- 31. M. cucullata

_ Bracteole, where known, much smaller, caducous or not, sometimes enveloping the

perianth in very immature stages only .... 6.66... 6s eee eee eee eee eee 3
Twigs (2—)3—5 mm diameter or more; most leaf blades 15 cm long or more ... . 4

Leaves with dots on both upper and lower leaf surface; this sometimes apparent only
towards the margin of the blade; lower leaf surface with subpersistent short and
dense scale-like hairs 0.1 mm or less. Inflorescences (infructescences) short pedun-
culate*. — W New Guinea, but not known from Bird’s Head; (sub)montane......
ee a nT Se Peg at) ete Bre Onoda tia ey Cee act 37. M. duplopunctata

_ Leaves with dots on lower surface only; lower leaf surface glabrous or glabrescent,

indumentum minute. Inflorescences sessile or pedunculate ..............--- 5
Inflorescences (best to be seen in male flowering specimens) without peduncle 6

OF COLMMACCOUS A. oe oak Soe cny dag ame Ce nes eee ne eae |
Leaves chartaceous or subcoriaceous; nerves 20-30 per side, faint below. Male flow-
ers: bracteole large, cucullate, caducous; androphore glabrous. — Papua New Guinea,
GSU S30 A Onin aera anon eer crta as Wren emerge LEMIRE. 31. M. cucullata

_ Leaves coriaceous; nerves c. 15 per side, distinct and contrasting in colour below.

Male flowers: bracteole not seen; androphore conspicuously pubescent in the lower
half. [Fruits not known.] — W New Guinea (SW of Bernhard Camp on Idenburg
Reeves LOO 0h aves 6 ks one <0 he cane Oe 81st aoe ee io 14. M. brassii
Twigs (2—)3 mm diam. Leaves 15-23 cm long, membranous; nerves c. 20 per side,
on lower leaf surface contrasting in colour and clearly visible. [Infructescences short-
pedunculate. Fruits 4.5-5 cm long. Flowers not known.] — Papua Barat (N Bird’s
Head, Tamrau Range); 500-800 m ............-....-.-.- 134. M. tamrauensis

. Twigs 5-7 mm diam. Leaves 20-40 cm long, chartaceous or coriaceous; nerves

+ concolorous, ZO—40 per SIS o.oo ape eee oe oe a oe eo sine oie ee 8
Leaves chartaceous or thinly coriaceous, drying dull with minutely wrinkled sur-
face; lateral nerves thin but clearly visible below. Fruits 5.5—6 cm long. Flowers not
known. — Central New Guinea; montane ............. 91. M. millepunctata

. Leaves (thinly) coriaceous, drying smooth; lateral nerves faint below. Mature fruits

not known. Male flowers: androphore completely pubescent.— Papua Barat (Bird’s
| SPECT WAM F511 Ee ae a Pea eee ar ic SSR Aen nates eke ee eS 94. M. neglecta

*) The peduncle is the non scar-covered stalk bearing a scar-covered part with flowers, the latter

sometimes much reduced and few-flowered, especially in female specimens.

De Wilde — Myristicaceae (Myristica) 381

18a.

. Specimen with female flowers and/or fruits ... 2.0.6.0... eee eee eee 10
. Specimen with male flowers (not known in M. leptophylla) ..........-+-+: 17
_ At least some inflorescences or infructescences with a peduncle* 3 mm long or
Tee ES SS SU AD a id Fe ii ie). ee eet & « 11
Inflorescences or infructescences all sessile, or with a smooth peduncle up to 3 mm
inrreamee >t) Decl! gdak dealt) 4 slanubed-l re -vneeeeaeee peele 13

_ Female buds 10 mm long. Fruits 4.5-6.5 cm long, style often persistent. — Most of

New Guinea: lowland and submontane forest; up to 1000 m 27. M. cornutiflora

_ Female buds (not seen in M. arfakensis) less than 10 mm long. Fruits 3—4.5 cm

long, style not persistent. — Montane species, at 1000 m or more ......... 12

_ Dots on lower leaf surface clearly visible only with a lens. Fruits, including pseudo-

stalk**, 3.5-4(-4.5) cm long. — Throughout New Guinea; montane...........

en Oe, ee renga cern Hoh an he eh re ee Oe 100. M. pachyphylla
_ Dots visible to the naked eye. Fruits (without pseudostalk) 3 cm long. — Papua
are bist S He AG): AMOMLANC, 5. ace epee ata Fl ny ere Ger ier 4. M. arfakensis

_ Fruits 1.5-2.5 cm long (not known in M. flavovirens). Nerves on lower leaf surface

generally faint. — Lowland; up to 700 m (in Bird’s Head to 1200m) ...... 14

_ Fruits 3.5-4.5 cm long. Nerves quite visible below. — Montane; 1400-2000 m 16
. Leaves greenish. — Papua Barat (Bird’s Head) .................---++-- 15
. Leaves (dark) brown. [Twigs 2-3 mm diam. Fruits rounded at base.] — New Guinea,

PME RMMENT A SCAG oe a eto ke eens curries Gens te ae ee ae 122. M. scripta

. Twigs 1(—2) mm diam., brown. Leaves below grey(-brown). Fruits at base contract-

eaumio a 5 mm lone pseudostalk . on. oy. < 5 acon eye cee wien cyue 26. M. conspersa

. Twigs 2-3 mm diam., blackish. Leaves below yellowish. Fruits not known. —

Papua Barat (Bird’s Head); montane ..................-- 48. M. flavovirens

_ Leaves (4-)6-14 cm long, (sub)coriaceous. Fruiting pedicel (I-)2 mm thick. —

Throughout New Guinea; montane ..............------ 100. M. pachyphylla

. Leaves 10-20 cm long, membranous or (thinly) chartaceous. Fruiting pedicel 4(—5)

mm thick. — Papua New Guinea (W Sepik Prov.); 1500 m_ 81. M. leptophylla

. Inflorescences with a smooth, often more or less flattened peduncle (1—)3 mm long

“= erent uel eee en reele ark nee. prc gyi era es tr eer wate mie pe Pes IEW saytay 2 20

_ Inflorescences (sub)sessile, simple or forked, short, scar-covered, (+ woody)

brachyblasts producing flowers at the apex, lasting several seasons; peduncle ab-
sent or up to 3 mm long (including M. leptophylla of which male inflorescences
BIPEMOWVELS Ate MOL KNOWN) cc. oe 2 = os nsec no tues 2 meee eee as 18
Twigs 1(—2) mm diam. Male buds 3.5 by 1.5 mm. [Leaves greenish. Inflorescence
sessile or 1(—2) mm pedunculate. Bracteole small, persistent.] — Papua Barat (Bird’s
ifead) lowland tp tO 900m 828 dae eee ie et ee 26. M. conspersa

. Twigs generally stouter, 2-3 mm diam. Male buds 3.5—5 mm long; if smaller, then

not occurring in Bird’s Head. — Throughout New Guinea ............... 19

*) Usually the stalk of the fruit in single-fruited infructescences is thickened and woody; it is made

up of the peduncle and the pedicel; the demarcation between the two parts may become obscure
with age.

**) The pseudostalk is a stalk-like contraction of the basal part of the fruit proper, demarcated from
the fruiting pedicel by the scar of the perianth.

382

Flora Malesiana, Ser. I, Vol. 14 (2000)

23at

a. Twigs dull or blackish brown. Leaves (dark) brown above, lower surface pale brown,

not conspicuously papillose even seen under a strong lens. Inflorescences sessile.
Flowers: pedicel 2—4 by 0.7 mm, bracteole caducous; buds 3.5—5 by 2—2.5 mm. —
Throughout New Guinea; lowland up to 600m ..........-.. 122. M. scripta

. Twigs blackish. Leaves greenish above, lower surface yellowish, densely papil-

lose. Inflorescences with peduncle 1(—2) mm long. Flowers: pedicel 4 by 1(-1.5)
mm, bracteole minute, 1 mm long, persistent; buds 5 by 2.5 mm. — Papua Barat

(Bids Headiec2 1200 mm fhe) See RR 48. M. flavovirens
_ Male buds 10 mm long or more. — Throughout New Guinea; lowland and lower
fentane arease SYP AP e Lk ee ee ce SRI 27. M. cornutiflora
“Male buds S110) mm lone-or less") 05. SL Se SD ee pa

_ Dots on lower leaf surface mostly clearly visible only with a lens. Buds elongate,

6—-7(-8?) mm long. — Most of New Guinea; montane..............-.++.+--
Petre Cy Wits GETMAIN E Sue ate or ees e, toate eae eee ee 100. M. pachyphylla

_ Dots visible to the naked eye. Buds comparatively shorter and broader, 3.5—4 mm

long. — Papua Barat (Bird’s Head); montane .............. 4. M. arfakensis

. Twigs medium-sized or stout, conspicuously hollow and + angled by distinct raised

lines or wings up to 2 mm high, at both sides of the twig connecting the bases of the
petioles; perforated ant-swellings present (but sometimes not collected). Wings
indistinct in' VM. ingrata subsp. velata <2... 20 se soo tee 23

. Twigs slender, medium-sized, or stout, generally solid or narrowly hollow, (sub)-

terete or + angled, rarely with raised lines, without wings (stem sometimes ridged
but not conspicuously hollow in M. hollrungii and M. sulcata); ant-swellings ab-
BENE rae t rte Ces tela sale eek Behind Sa dk Whee ore cas a Sin 0, cine een a 29
Twigs and leaves exceedingly stout: twig 10 mm diam., leaves to 43 by 18 cm.
Fruits 3.5 cm long with conspicuous hairs 1(-1.5) mm. — N Papua New Guinea
(SepikcProv.) low land ©% G.)2 S02 S08 |. Re eg te ee bye ete 34. M. dasycarpa

. Twigs generally narrower, leaves smaller. Fruits with shorter indumentum . . 24
_ Male buds cleft into lobes to over halfway; sizes of buds in one inflorescence vari-

able: bracteole either small, 2-3 mm long (subsp. kostermansii), or large, (S—)7-8
mm long (subsp. fissiflora). [Lines of interarching of lateral nerves and venation
distinct. Fruits not known with certainty.] — N New Guinea; lowland.........
BARE SRI MRIOES, oh re: Laat LOS DCTS tL Bt eet oes a sae 46. M. fissiflora

. Male buds cleft 1/3 or less; bracteole 2—3 mm long ..........----+++4+:: 25
_ Male buds ovoid or ellipsoid(-oblong), cleft 1/3-1/4; apex of synandrium blunt,

sterile apex absent or to 0.2 mm long. Fruits rather small, up to4 cm long .. pai |

. Male buds oblong or tubiform, cleft 1/6-1/8, lobes 1-1.5 mm long; sterile apex of

synandrium acute, 0.5—1 mm long. Fruits small or large, 1.5-7 cm long .... 26

_ Bracteole caducous at anthesis. Fruits 1.5—2.5(—3) cm long, generally with persist-

ent, short indumentum; fruiting pedicel 2-10 mm long. Venation generally faint.
— Throughout New Guinea, incl. Aru and Kai Is., not in the Bismarck Archipelago;
ECON NG U217 b:55 halter See aeatedarieae th ete geeieche te aoeisten ne BW, HRC Ra 129. M. subalulata

. Bracteole persistent at anthesis. Fruits 5-7 cm long, (late) glabrescent. Venation

faint or distinct. — Papua New Guinea (Bismarck Archipelago); lowland.......
Ghteiics 2s sonthe, sorulages mid sient ducks, eamas aitie Cemcieuenst aid taka apons cork caral keane 10. M. bialata

De Wilde — Myristicaceae (Myristica) 383

27a. Sizes of flower buds in one inflorescence generally variable; bracteole persistent.

[Female pedicel 2—5(—6) mm long. Fruits ovoid-ellipsoid, 2.5 cm long, apex nar-
rowly or broadly rounded (with minute apiculum); fruiting pedicel + stout, 3-10
mm long.] Lines of interarching of lateral nerves on lower leaf surface and vena-
tion generally faint. [Fruiting specimens may be confused with M. subalulata.| —
N & S New Guinea; lowland and montane .................- 68. M. ingrata

b. Flower buds in one inflorescence all of about the same size; bracteole caducous.

28a.

Lines of interarching of lateral nerves on lower leaf surface and venation generally
MISTING bined! n. ac.tanoeewinle fate pereeerce)-vlags aod tat es eee 28
Male buds 15 mm long, carnose; pedicel 7-8 mm long, bracteole scar below apex.
Female flowers and fruits not known. — N Papua Barat; lowland 119. M. sarcantha

b. Male buds 5 mm long; pedicel longer (10 mm) with bracteole apical. Female pedi-

cel 10 mm long; fruiting pedicel 10—15 mm long. — Papua New Guinea (W Sepik
Prov., Upper Sepik River); lowland .............-....+- 42. M. fasciculata
. Twigs lined to sharply angular. — Central New Guinea; montane; 1500-2000 m or
I i A ergo Nokon 2s cheoucren sp Me cok wie de athe eat ocr c boa beara 30

b. Twigs lined, ridged, or terete. Plants from lowland or montane areas, but in the lat-

ter case twigs (sub)terete, neither lined nor TNO UIA ee ee hs cnc ag ee 32
_ [Flowers not known.] Twigs + stout, 4-5 mm diam. Fruits subglobose, 6 cm diam.,
pericarp 20 mm thick. — Papua New Guinea (Milne Bay Proy., Mt Dayman);
=. |) SCOT arr ee re se pee eee 99. M. pachycarpidia

b. Twigs generally more slender. Fruits smaller, 2-5.5 cm long, + ellipsoid: 2. ai

_ Male buds 6 or 9 mm long (according to the subspecies); bracteole persistent. Fruits
2.5 cm long. Twigs 2—4 mm diam. — Mountains of Papua New Guinea.........
a er re rere See eee 145. M. velutina

b. Male buds 10-14 mm long; bracteole caducous. Fruits 2 cm long. Twigs 5 mm in

32a.

diam. — Papua New Guinea (Western Highlands Proy., Mt Hagen and vicinity):

2, LCT Ui er ere Sree see 129. M. subalulata var. hagensis
_ Male buds 7-9 mm long, at base verrucose inside; bracteole caducous. Fruits 3.5—
5.5 cm long. Twigs slender or medium-sized, 1.5—3 mm diam. — Throughout New
MMA NOME ANG rae Sec cae cue Rin Gee aint syns fe. Beene ciel ene eee 30. M. crassipes
Twigs 1-3 mm diam., most leaves of fertile twigs 15(-17) cm long or less (speci-
mens of M. inutilis, M. sphaerosperma, and M. subcordata may have rather small
leaves, but these species have only been entered in the alternative lead) .... 33

b. Twigs (2-)3—5 mm diam. or more, generally most leaves of a specimen 15 cm long

4° [GOI eee At rath er thai, cote PoheaPapyse enn ed A0: Bio-poatA cee s 143
. Specimen with female flowers or fruits. [Female flowers and fruits not known of
several species, incl. M. schlechteri] .........+.--.-----++-++-+2++---- 34

b. Specimen with male flowers. [Male flowers not known of several species.] . 88

_ At least some of the inflorescences or infructescences with a non scar-covered
peduncle* of 2mm long or more ........-.-+---- +e eee eee teeter eee 35

b. All inflorescences sessile: a short simple or forked scar-covered brachyblast, ses-

sile or with a smooth peduncle up to2 mmlong .............------++++- 52

*) Usually the stalk of the fruit in single-fruited infructescences 1s thickened and woody; it is made

up of the peduncle and the pedicel; the demarcation between the two parts may become obscure
with age.

Flora Malesiana, Ser. I, Vol. 14 (2000)

. Leaves ensiform, parallel-sided, 12-22 cm long. Understorey near-tree of 1.5 m

height. — Papua New Guinea (Western Prov.); c. 80 m; once found............

5 thule che Leahey ess, Sak TS a oe ER are 39. M. ensifolia
. Leaves elliptic or oblong(-lanceolate). Trees generally larger ............. 36
Hairs:ofleaf bud‘(0:2=)0!S nim. or more’ 43708 OUI A eee ou
Hairs’ of leaf bud. short, 0.1—0.2(;—0.3)'mmi or less". \/4...42)-02..0 2 42

. Lower leaf surface with persistent dense indumentum, sometimes late glabrescent.

— E Papua New Guinea (Morobe); montane; 1000-1500 m . . 125. M. sinclairii

. Lower leaf surface early (sometimes late) glabrescent or glabrous (or sometimes

with persistent indumentum in M. mediovibex) .........+-+++++-eeeeees 38
Mower leat'surtace papillose Gens!) fav DE ee ee 39
SEower leat-surtace not‘obviously papillose /2)/ 22's) he ley 40

. Fruits 3-4 cm long, with coarse hairs 0.5—0.8 mm long. — Moluccas and Papua

Bara bird's Head)... 2.55% Seek eeore 11. M. bifurcata subsp. bifurcata

. Fruits 4—4.5 cm long, hairs 0.1 mm. — Papua New Guinea .. 22. M. clemensii
a. Bracteole caducous, about halfway the pedicel. [Mature fruits (incl. pseudostalk)

4 cm long. Lower leaf surface late glabrescent or with persistent indumentum. ] —
Northern part of W New Guinea ......... 89. M. mediovibex var. mediovibex

- Bracteole at or towards the apex of the pedicel” °>. <= 3... 2. =. atte 4]
. Female peduncle slender, 20 mm long, 0.5—1 mm thick; fruiting pedicel 7-9 mm

long. Fruits (incl. pseudostalk) 5—5.5 cm long. — E Papua New Guinea.........
ed cy leering «oad ilies Nel et havea veys tape tn a phteaa ote: Riu re 8 He Sg 106. M. pilosella

. Peduncle 10 mm long, | mm thick; fruiting pedicel 14 mm long. Fruits incl. pseudo-

Stalk. 2—2.5.cn, lons.-— NE Papua -Batal 2.2 oh acces 2 mater 137. M. trianthera

. Peduncle 3—4 mm long; fruiting pedicel 2-4 mm long. Fruits (without pseudostalk)

2.5 cm long. — Papua New Guinea (Morobe Prov.); c. 100 m 112. M. pygmaea

. Lower leaf surface seemingly glabrous, but actually with dense, very short indu-

MME MEMUCLETISY) co scccs slave <yujcoe ea Fc otgm «ee, ms oe apices 2g bears ge err 43

. Leaves below glabrous or glabrescent, or with a lax indumentum, hairs + scattered,

scale-like, minute only. [Female buds generally less than 8 mm long.] ...... 45

. Lower leaf surface with silvery short cobweb-like indumentum; leaf base cuneate.

Older twigs rugose with conspicuous wart-like lenticels. Female buds 8-10 by
5—5.5 mm. Fruits early glabrescent, 7 cm long, wrinkled. — W Papua Barat; low-
Peri Pt AT 8 Re Re ee eee ne a eae 5. M. argentea

. Indumentum of lower leaf surface largely grey-brown or rust-coloured, very short.

Bark of twigs not coarsely warty-lenticellate. [Flowers not known.] Fruits shortly
rusty pubescent, not wrinkled’) Os feces cade hele tae © coe de rae 44

. Fruits 7 cm long. Petiole 2-3 cm long; blade base broadly rounded or + cordate —

Papua New Guinea (Western Prov.); lowland ............ 3. M. archboldiana

. Fruits 2.5—3.5 cm long. Petiole 1.5 cm long; blade base cuneate — N Papua Barat

Sh, AMD ARETE TUS EEL) MAURY LE, Peet eon, 89. M. mediovibex var. kosteriana

. Lower leaf surface papillose (lens!). Peduncle conspicuously flattened. Fruits (not

known in M. rosselensis) ellipsoid or oblong, 2—3.5 cm long, glabrescent except at
fheends.;, {hs space aaa ee hee is) aie ete Se ee ee 46

. Leaves not papillose below. Peduncle + flattened or terete. Fruits variable in shape,

2.7 cm long or more, with minute scurfy indumentum (hairs scale-like) .... 47

De Wilde — Myristicaceae (Myristica) 385

46a.

b.

A7Ta.

48a.

Sila

54a.

Leaves narrow, oblong-lanceolate, base (sub)acute; lateral nerves c. 20 per side;
nerves and venation faint. — Louisiade Archipelago ...... 115. M. rosselensis
Leaves proportionally broader, elliptic-oblong, base rounded; lateral nerves 8-15
per side; lateral nerves and venation + evident. — N Papua New Guinea (Bismarck
Archipelago), E to Solomon Islands; mostly coastal ...... 121. M. schleinitzii
Lower leaf surface grey-white, glabrous. Inflorescences (infructescences) conspicu-
ously supra-axillary. [Flowers not known.] Fruits without pseudostalk, ellipsoid-
oblong, 2.7 cm long. — Papua New Guinea (Madang Prov.); at low altitude.....
RY 546. 0. cheer ath, Yee Baombel a abate. 33. M. cylindrocarpa

. Lower leaf surface somewhat whitish or not. Inflorescences axillary or only shortly

supra-axillary. Fruits 4 cm long or more, including pseudostalk (mature fruits not
Laemmli isolvacem) ie. ait te: C8 3 seyret eins 1 die Jom eee a 48
Leaves drying conspicuously greenish. Inflorescences slender, the peduncle 3 cm
long; pedicel 18-20 mm, longer than the perianth. [Bracteole persistent.] — Papua

NevwaGuimes (CentralsProv:);lowland:. . 248) .).0).5 (90.0% 62 22 96. M. olivacea
. Leaves drying (olivaceous-)brown; peduncle of inflorescences (infructescences)
mnenshonter:pedicelivatiable! im. sue whee aaeae eee SAREE « 49
. [Female flowers and fruits not known.] Leaves drying conspicuously blackish brown.
Bracteole in male flowers persistent. — Papua Barat and Papua New Guinea
(Ingembit area, on the S border); lowland ................. 6. M. atrescens
. Leaves drying olivaceous or (dark) brown. Bracteole caducous. ........... 50

_ Lateral nerves on lower leaf surface slightly raised, brown, contrasting with rest of

Dingicw eeelinicl: 3.2 {de ae SOR Ee eric ee eee 51

. Lateral nerves on lower leaf surface flat or + sunken, almost concolorous. [Inflo-

rescences generally lacking a peduncle.] — Papua New Guinea; montane.......
rest 99.00,.200 ah isle Jit: tan § 8 EM ite Ses 76. M. laevifolia
Fruits (incl. pseudostalk) 4.5-6 cm long. — Papua New Guinea (Central Prov.);
local, hill rain forest at 500-700 m (a deviating fruiting specimen from montane
fonespat 1300 min Central Papua Barat). 222k eas... Sees 83. M. longipes

. Fruits 4 cm long. — Papua New Guinea (Central and Madang Prov.); montane for-

Soma 400) 12001600 arin os yatta he aes OR 52. M. fugax

. Fruits narrow, 8 by 1.5—2 cm. — SW Papua Barat; lowland or montane.........

eee At elle deonersiio..t 1.88... bashyol sean) 228 54. M. fusiformis

PEiisbtoader OL shorter ow) Aww hen sisaeRe.oL ck ta eee See 53
. Lower leaf surface early glabrescent or covered with very minute scattered pale

cobweb-like hairs only less than 0.1 mm long and papillose (lens!), the papillae
rather uniform and regularly set. — Note: some species with the lower leaf surface
very densely papillose and the papillae closely contiguous, or with the papillae
irregular in shape and size or indistinct, are included in both leads or in the alter-
native lead; specimens of M. insipida and M. lancifolia may be indistinctly papil-
loses eWesukacivihs ours 2ec ke ore eee eee ee 54

. Lower leaf surface glabrous, glabrescent, or pubescent, not or but faintly papillose,

or/papillaé not readily recognizedias such: ... 2. .2.92 222200) 225-2 ee 7
Fruiting pedicel 15-20 mm. Nerves at c. 45° to the midrib. — Papua New Guinea
OVestermProvs): lowlandwtisn.ec eee 2s Cee 101. M. papillatifolia

. Fruiting pedicel 5 mm long or less. Nerves at 60° or more to the midrib .... 55

Flora Malesiana, Ser. I, Vol. 14 (2000)

56a.

(@)

. Lateral nerves 8—20 per side, not much contrasting in colour and faint below; vena-
tion generally indistinct. [Fruits small, ellipsoid, 1.3—1.6 cm long, with hairs 0.1
mm.] — Moluccas and W & E New Guinea . 78. M. lancifolia subsp. lancifolia

. Nerves 6—14 per side, below contrasting in colour or not; venation not much con-
frasting | iIndistinctior not Maw Ts eae eee. OGL Go LS 56
Fruits ellipsoid, 1.5—2.5 cm long, including the short pseudostalk, with hairs 0.1—
0.3 mm. [Fruiting pedicel 24 mm.] — E Papua New Guinea .. 25. M. concinna

. Fruits subsessile, ellipsoid(-oblong), 2.5—3.5(—3.8) cm long, with hairs 0.3-1 mm.
— § New Guinea (also in S Moluccas and N Australia) ....... 70. M. insipida

. Fruits subsessile, + ellipsoid, 3 cm long, with hairs 0.1(—0.2) mm. — Local en-
demic of S Papua New Guinea (Central Prov.); sea level ... 13. M. brachypoda

57a. [Flowers not known.] Lateral nerves c. 25 per side, faint. Fruits egg-shaped, 7 by 5

cm, early glabrescent (a few light brown scattered appressed hairs excepted). —
Resseliinat:200misia0tt) sneha Adie ee eR eee Bee 98. M. ovicarpa

b. Lateral nerves c. 20 per side or less. Fruits generally less than 7 cm long .... 58

. Flowers glabrous or almost so. Fruits minutely pubescent or glabrescent .... 59

. Flowers pubescent (hairs may be minute and dense, the perianth seemingly gla-
brous (lens!); sometimes parts of the perianth late glabrescent). Fruits pubescent,
hairs minute or larger. [Flowers not seen in M. brevistipes, M. firmipes, or M. lasio-

carpa; fruits glabrescent in M. crassipes.| ........-+2 0020s c eee eee eees 61

a. Fruiting pedicel 2-7 mm; fruits glabrescent. — Lowland; 0-250m..........-.
SAU BOU26NDS CN DOR Ae SORT Te ee AEE 139. M. tubiflora

. Fruiting pedicel 10 mmilong ormore 2). 2222. 2a fe ee he a ie 60

. Fruiting pedicel (20-)25—30 mm by 0.5 mm; fruits minutely pubescent, with long

pseudostalk. — E Papua New Guinea (Lae subprov.); 0-15 m ... 44. M. filipes
. Fruiting pedicel stouter, 10-18 by 2 mm; fruits glabrescent, with short pseudostalk.
— Papua New Guinea; montane, 1000-1800 m ............. 76. M. laevifolia

. Leaves below either glabrescent (sometimes late), or with extremely minute indu-

mentum of scale-like, + spaced, not densely interwoven hairs ...........-. 62
_ Leaves below with persistent or subpersistent indumentum, with hairs densely set,
long or short; or late glabrescent; hairs may be extremely short and scale-like 82
. Innovations and flowers conspicuously densely clothed in rusty hairs 0.5—1 mm.
—E Papua New Guinea; lowland .. . 20. M. chrysophylla, 120. M. schlechteri
Afaiivs much: SOLE 5 ss .< os nassceso ete cere ne Foe tones amps Yom note RR: hd 63
Fruiting pedicel 2 mm long, 1-2 mm thick; fruits 2.5—3 cm long, ellipsoid-oblong.
[Flowers not/kno wang Siraerte eek SRS, SL ee 64
. Fruiting pedicel more than 2 mm long, slender or thick; fruits variable, compara-
tively broader: globose, short-ellipsoid, or + pear-shaped ................ 66
. Leaves pale below, nerves thin and faint on both surfaces. Bark of older twigs
+ corky. — Local endemic of S Central New Guinea; 150m 90. M. mediterranea
. Leaves brown-green, not pale below, nerves stronger; twigs not corky ...... 65
_ Indumentum of fruits conspicuously woolly. Lower leaf surface faintly papillose.
— S New Guinea (also in Moluccas, N Australia) ............ 70. M. insipida
_ Indumentum of fruits short-woolly. Leaves not papillose. — Local endemic of Papua
New Guinea (Central Prov.) (Check also M. concinna and M. brachypoda.)......
i dex iborn.o ann a ae, ANG a shes OE ERIE Se 15. M. brevistipes

De Wilde — Myristicaceae (Myristica) 387

Waa

73a:

Ta.

. Twigs + 2-lined or 2-angled (usually indistinctly so in M. crassipes) ....... 67
. Twigs not or only faintly angled, including M. atrocorticata, a species with some-

what angled twigs but distinguishable by the flaking bark of older twigs .... 73

. Lower leaf surface drying (pale) brownish, nerves comparatively faint, yellow or

orange-brown, not much contrasting. Fruits 2.5—4 cm long; seed attached above
the base of the pericarp and hence + transversely positioned in the fruit; fruiting
pedicel + stout, 10-25 by 2—3 mm; perianth sometimes glabrescent. — N & NE
NewaGuinea: lowland and montane’. 2. 22. 149. M. warburgii

. Lower leaf surface drying pale, greyish or not, midrib and nerves brown, and these

generally contrasting. Fruits variable; seeds basal. Perianth minutely thinly pubes-
cent, the hairs sometimes extremely small, thus the perianth seemingly glabrous
SL ee EE ah: SOR OAT [ROS te Sn oie SOS Be eee 68
Twigs 1 mm diam. Fruits in lower half contracted into a long pseudostalk ... 69

. Twigs (1.5—)2-3 mm diam. Fruits without or with pseudostalk; fruiting pedicel

SIL DIME Oop One Rae ee Pk See CR OAs oi neler Cee UO abe SEAR ee IS BESS So: 3 70

. Fruiting pedicel 20 by 0.5 mm. — E Papua New Guinea ....... 111. M. pumila
. Fruiting pedicel 3—4 by 2 mm. — Papua New Guinea (Central Provy., Sogeri area)

ss coc 2 REO ARSE AUS LE BY RUA Ese Se bY Een ge ead 127. M. sogeriensis

banana, (or UO ti-alttbden 2 ¢.2 28, 2bs SUS. aes oo cs oy eo ceeen eee qi
mi tentane. 500 mualtitude or higher \s.... bic sh. oo tae Se op) eee 12,
. Fruiting pedicel 30-35 by | mm; fruits with 10 mm long pseudostalk. — E Papua

RETA aimee crn a A hie ores anpalty 62 Fo Mie Wane ees 0G 49. M. flosculosa

. Fruiting pedicel 3-10 by (3-)4-5 mm, fruits without or with short pseudostalk.

{Leaves usually more than 15 cm long.] — Throughout New Guinea...........
PUNY AS See aes te + tte old Ma ogee weyetys 132. M. sulcata
Fruiting pedicel 4—10(—12) by 3-5(-6) mm. — Throughout New Guinea........
Rea 9) Gres syalls Peels en sere ofa Tee Ae AE ae 30. M. crassipes

. Fruiting pedicel 10-15 mm by 1.5-3 mm. — E Papua New Guinea............

PRPS NE) 1). tee are: Keel oss. Ly) Ma 149. M. warburgii subsp. hybrida

. Fruiting pedicel 25-30 by 1 mm. — Papua Barat; 500m ... 58. M. gracilipes

Small tree, 1.5—8 m tall. Fruits solitary, with slender peduncle; fruiting pedicel
8—10 by 0.5 mm, the fruits itself with distinct pseudostalk, 10 mm long. — E Papua
IWeh7 (GUTTTE ee), ae ec 93. M. nana

. Tree small or large. Fruits of variable sizes, single or 2—4 together per infructescence;

fruiting pedicel stronger and proportionally shorter; fruits without or with much
shonfampscudostalkte wan Ques) Ol meee. GhakhS See eee 74

. Leaves coriaceous or membranous. — Montane; (300?—)1600-1800m ..... TS
. Leaves subcoriaceous or membranous. [Fruits either globose, + ellipsoid, or + pear-

shaped; fruiting pedicel short or medium-sized.] — Plants generally from lowland
Bee ee OOOO USOO) a see ina a eee teeecmrsarnice ne te fo teh tere oe 76
Fruits ellipsoid or + fusiform, 3 cm long, the pericarp 3 mm thick; fruiting pedicel
10 by 3.5 mm. — Papua Barat (near Wissel Lakes); c. 1800 m (species resembling
M. laevifolia, but nerves contrasting below) ............... 148. M. vinkeana

. Fruits globose, 2.5—3 cm diam., the pericarp 7 mm thick; fruiting pedicel 5-8 mm

long. — Papuan Islands: Goodenough, possibly Tagula; 1600 m (and 300 m?; see
MOL UMGEINIMENS OCICS)» PR i AAMAS «24 342 05 eerie 108. M. polyantha

Flora Malesiana, Ser. I, Vol. 14 (2000)

TUTE.

78a.

8la

. [Male and female flowers not known.] Fruits + pear-shaped, 6—7 cm long, pericarp
thick, woody, 10 mm thick; fruiting pedicel short and thick, broader than long,

4—10 by 5-9 mm. — W Papua New Guinea ................ 45. M. firmipes
. Fruits globose or + ellipsoid, smaller, pericarp thinner; fruiting pedicel proportion-
allyamore slendeting! gas j-.€ © sii d spe dee dees sbees-le cee Hee Teh

Leaves below usually pale, grey-glaucous, with brownish contrasting nerves and
coarsely reticulate contrasting venation. [Fruits variable in size and shape, (short-)
ellipsoid or subglobose, 1.5—3.2(—4) cm long, with persistent short hairs 0.1—0.5

(0.8) mm.] — Widespread; 0O—600(—-1500) m ............... 57. M. globosa
. Both leaf surfaces generally rather concolorous; venation more faint ....... 78
Twigs 2—2.5 mm diam., somewhat angular ...........-..-.+-.+++2005- 1)

. Twigs 1-2 mm diam., subterete. [Bark of older twigs striate, + flaking or not. Peti-
ole long and slender (but proportionally shorter than in M. atrocorticata and M.
inundata), 10—20(—23) mm long. Leaves glabrous (early glabrescent) below. Fruits
Sealda plage rhace O pillt CxO kA) ict s i ni niin ki IONE EEI RC Sc cm 81

a. Bark of older twigs blackish, longitudinally cracking, or flaking. Petiole 20 mm

long. [Leaves below late glabrescent, at first with minute scale-like hairs.] .. 80
. Twigs brown, bark not flaking (but sometimes transversely cracked). Petiole 10—
13 mm long. — Northern part of New Guinea; lowland .. 16. M. buchneriana

a. Leaves oblong-lanceolate. Bracteole much shorter than the perianth. [Fruits not

known.] — W Papua Barat (Bird’s Head) and Papua New Guinea (Western Prov.);
PO Wile ete ss tea nc day er 2 tte cs TM ected emer are 7. M. atrocorticata
. Leaves (elliptic-)oblong. Bracteole in male flowers about as long as the perianth.
[Female flowers not known.] — Papua New Guinea (Western Prov.)...........
sol rou ed one Sesaiowrn saith coe a ae ie ls ene WE A a ve ar yrertb at te ey) 36 ol oh oN 71. M. inundata
Fruits subglobose or ellipsoid, 2.5—3.5 cm long. — Moluccas and West & East
INS WAGGA ee ere ete a eee oe meen en one aaa oe mete en era 138. M. tristis
. Fruits + ellipsoid-oblong, 2—3 cm long. — Moluccas and Papua Barat (Bird’s Head).
[See also M. concinna, but fruits ellipsoid, 1.5-2.5 cm long, and leaves usually
Papillosetbelawal metre tee tes eee ee 80. M. lepidota subsp. montanoides

. Fruits ellipsoid, 4.5-5 cm long. — Southern Solomon Islands ................

PEAT PESE AEBS DETER) EE CRM OS 5: M. guadalcanalensis W.J. de Wilde]
. Twigs 1.5 mm diameter. Leaves 4-14 cm long. — Moluccas and West New Guinea
aan ON ee Oe ee re 80. M. lepidota subsp. lepidota
. Twigs 2 mm diameter or more. Leaves generally 10 cm long or more ...... 83

a. Leaves below with silky, very short, (sub)persistent indumentum; venation + tra-

beculate. — Papua New Guinea (Northern Prov.); montane ... 17. M. byssacea
. Indumentum more scaly or felty, short or longer; venation reticulate, or hardly vis-
ible! Axeuu Mace seiastelstote ele Lyin ee OSB eee 84
. Fruits 2 cm long; fruiting pedicel 15-18 by 1.5—2 mm. — Papua New Guinea (Ma-
dane iProv,): lewland)srdin. ot: CR eR ee eet 103. M. pedicellata
. Fruits larger; fruiting pedicel shorter and thicker ....................5-. 85

a. Flowers, fruits, and lower leaf surface with short hairs, 0.3(—0.5) mm long or less

. Flowers, fruits, and usually lower leaf surface with longer, shaggy hairs .... 87

a. Lower leaf surface with scale-like hairs 0.1 mm long only. — Papuan Islands... .

i eG TLS eee ee Oe ee Ne CeNG OS cote care 136. M. tenuivenia

De Wilde — Myristicaceae (Myristica) 389

b. Lower leaf surface with dense, stellate hairs of mixed sizes, more than 0.1 mm
long. — Widespread (check also M. subcordata).| ...........: 72. M. inutilis
c. Lower leaf surface with dense minute scale-like hairs, less than 0.1 mm long. [Fe-
male flowers and fruits not known.] — Papua New Guinea (Western Prov.); low-
endyperiodically inundated forest Yoe.a0. 2 2.5. ee 71. M. inundata
d. Lower leaf surface densely short-felty, with hairs 0.2—0.3 mm.— Papua New Guinea
(West Sepik Prov., Telefomin subprov.; possibly Morobe Prov., Kuper Range);

femme RU Crcriner edd NE ere hive ele DEAL ERO ee ee 23. M. coacta

87a. Fruits subsessile. — E Papua New Guinea; lowland ..... 20. M. chrysophylla
b. Fruiting pedicel (5—)10 mm long. [Flowers not known.] — Papua New Guinea
(Morobe Prov., Kuper Range); 1600-2000 m ............ 79. M. lasiocarpa

88a. Inflorescences with a smooth, often + flattened peduncle (1—)2 mm long or more,
Fictmeronc OG Several SEASONS..." LONF IER) 2. f2) | ESAS 7 Meee en ee eke 89

b. Inflorescences (sub)sessile simple or forked scar-covered woody brachyblasts pro-
ducing flowers at the apex, mostly lasting several flowering seasons; in some spe-
cies with a peduncle to 2 mm long, in M. insipida the peduncle occasionally up to
4 mm long; inflorescences sometimes delicate with only a few flowers .... 109

89a. Leaves narrow, ensiform, 12—22 cm long. An understorey near-tree of 1.5 m. —
Papua New Guinea (Western Prov.);c. 80m ............... 39. M. ensifolia

b. Leaves elliptic or oblong(-lanceolate). Trees or shrubs, generally larger .... 90
90a. Leaf bud (and generally also young twig apices, inflorescences, and flowers) with
hairs of a flossy appearance, the majority (0.2—)0.5 mm long or more ...... 91
peeeoncsl—0:2(—0.3) mmoilong onless\ 2080 >.2 226,829.02 eee ie si)
91a. Male buds oblong, 7 mm long or more. [Male flowers not known in M. mediovibex,
Miemposelia Mupysmaca:| (ie vos6)5 i ofl sae ee en ei ee a2

b. Male buds ellipsoid(-oblong), 5 mm long or less ...........-.+-2-+-+5-- 95
92a. Lower leaf surface with persistent + dense indumentum, sometimes late glabrescent.
Peduncleof maleinflorescence up to 3:mm long .3..000. 220-9. 25 5ns oe 93

b. Leaves early glabrescent or glabrous below. Peduncle longer ............. 94
93a. Plants from montane area; 1000—1600 m. — Papua New Guinea (Morobe Prov.)
GUA AC hs LeNR’ 0). Ct soe AR A OS ae 125. M. sinclairii

b. Plants from lowland area; c. 100 m [Male flowers not known.] — Papua New Guinea
Mis oeweroy UL. joboels 200 ole, 2008 SUS. ee 112. M. pygmaea

94a. Bracteole in female flowers caducous, borne halfway the pedicel. [Male flowers
not known.] — N Papua Barat; lowland ...............- 89. M. mediovibex

b. Bracteole persistent, at the apex of the pedicel. — E Papua New Guinea; montane
eet) ert tone. dies thd cee. 6. a ee 106. M. pilosella

95a. Lower leaf surface papillose (lens!). Male buds + ellipsoid; anthers 6-8 .... 96

b. Lower leaf surface not papillose. Male buds ellipsoid-oblong; anthers 3. — NE

PapuaiBaratlowland QL) 4.2) Ree eee 137. M. trianthera
96a. Leaves ovate-oblong or oblong-lanceolate. Apical part of male buds rounded in
cross section. — Moluccas and Papua Barat (Bird’s Head)...................

b. Leaves oblong-lanceolate. Apical part of male buds often + triangular (sometimes
not apparent). [Dry fruits 4-5.5 cm long.] — Papua New Guinea (Morobe, North-
em. ieilands'Prov. & New Britain)... 2-222. oe. oe. 22. M. clemensii

Flora Malesiana, Ser. I, Vol. 14 (2000)

98a.

1O1a.

102a.

. Leaves below seemingly glabrous, but very short and dense indumentum evident

undervatlens. i.) saree ee ieeeeytaet eet ots ae, Bee ee 98

. Leaves below glabrous, (late) glabrescent, or with small but scattered hairs only.

Male buds either less than 12 mm long, or if 12 mm long then only 2—3 mm across
(here also included is M. cylindrocarpa, of which the flowers are not known)
wht. dren Ie Re AG, Gaciat ae eile nb ae YRS 45 eee 100
Leaves below silvery by its short cobweb-like indumentum. Older twigs rugose
with coarse wart-like lenticels. Male buds ovoid-oblong, 1 1—12 by 5—5.5 mm. —
WiPapuarbarat: lowlands eG) aie). sce) aac on ee ae 5. M. argentea

. Indumentum rust-coloured. Older twigs not with wart-like lenticels. Flowers not

kamen oe SR? ie DRE eet ee a er 99
Petiole 2—3 cm long; base of leaf blade broadly rounded or + cordate. — Papua
New Guinea (Western Prov.); lowland ................ 3. M. archboldiana

. Petiole 1.5 cm long; base of leaf blade cuneate. — Papua Barat (Jayapura Prov.);

lowlandrifvsos Qrte rOab ese SHEE VE 89. M. mediovibex var. kosteriana

. Lower leaf surface papillose (lens!). Peduncle of inflorescence conspicuously flat-

tened. Male buds ovoid to oblong, 6 mm long or less ................- 101

. Lower leaf surface not papillose. Peduncle flattened or not. Male buds oblong-

lanceolate, 6 mm long or more [Flowers not known in M. cylindrocarpa; male
buds-4- mm iniMs-atrescens.||u. 20012 ..4oLaoeatel> eel Jo. Re. Se ee 102
Leaves oblong-lanceolate, nerves and venation faint. Male inflorescences with
flattened peduncle, with sessile simple lateral flower-bearing brachyblasts; male
buds 5.5 mm long or less. — Louisiade Archipelago ..... 115. M. rosselensis

. Leaves + elliptic-oblong, lateral nerves and venation + distinct. Male inflores-

cence with flattened peduncle, with stalked simple or branched brachyblasts; male
buds 5.5—6.5 mm long. — Most frequent in coastal area of northern Papua New
Guinea, Bismarck Archipelago, E to Solomon Islands .... 121. M. schleinitzii
Leaves below glabrous, (grey-)whitish. Infructescence (inflorescence) 10 mm
supra-axillary. [Flowers not known.] — Papua New Guinea (Madang Prov.); low
altatude: pores korea ese eee, SO. Ue ceria 33. M. cylindrocarpa

. Leaves below grey or brown. Inflorescences up to 6 mm supra-axillary ... 103
. Leaves conspicuously greenish above. Inflorescences slender, with peduncle 3-5

cm long. Male buds 1 1-13 mm long; pedicel long, slender, 10-12 mm long, bracte-
ole persistent. — Papua New Guinea (Central Prov.); lowland...............

. Leaves (olivaceous-)brown. Peduncle 1.5 cm long or less. Male buds shorter;

braeteclapersistent;on caducous|xiel.ce: aes: CS ee ee 104

. Leaves conspicuously blackish brown. Male buds 4 mm long; bracteole persist-

ents SE Papua Baratslomlandit 2h) ace bit Spe 6. M. atrescens

_ Leaves olivaceous, brown, or dark brown. Male buds 6 mm long or more; bracteole

(hate yeaducons Jenner deo Ale OR eee ae..ene@i. 0G eee 105

. Male buds 10 by 3 mm [inner surface smooth; pedicel S—6 mm long.] — Local

endemic in rain forest of Papua New Guinea (Central Prov.), 5|00—700 m (possi-
blyalso CentralPapua Baraty 1300 mee eee yap seer 83. M. longipes

. Male buds 6-9 by 1.5—2(—3) mm. — Plants from lowland or montane area 106

De Wilde — Myristicaceae (Myristica) 391

106a.

109a.

112a.

sa.

Male buds inside smooth. [Lateral nerves on lower leaf surface flat or sunken,
concolorous; a species generally with sessile, non-pedunculate inflorescences. |
—N & E Papua New Guinea; montane ..............1--- 76. M. laevifolia

_ Male buds inside with verrucose ridges in the lower half. [Pedicel 3.5-6 mm

long.] (Check also M. pumila, a species from low altitudes, usually with com-
pletely sessile inflorescences) ..°/., 0; 24 tes 22 See eee 107

. Twigs 2-3 mm diam. — Throughout New Guinea; montane . 30. M. crassipes
Malo ecel—2 mami dignity! 2 .2ist. 20. eG. He ANE SA eeaere ne - 108
_ Nerves on lower leaf surface much contrasting in colour. Male buds 7 mm long,

apex acute, verrucose ridges on inside faint; bracteole persistent. — Papua New
Guinea (Central Prov., Sogeri area);c. 400m _ .......... 127. M. sogeriensis

_ Nerves less contrasting. Male buds + blunt, 4-6 mm long, verrucose ridges dis-

tinct: bracteole caducous. — Papua New Guinea (Gulf, Central, East Sepik, and
Madang Prov.); foothill and montane forest; 300-1600 m_...... 52. M. fugax
Lower leaf surface glabrous (or early glabrescent, or with scattered very minute
pale hairs less than 0.1 mm), with distinct uniform and similarly shaped papillae
(papillae may be less prominent in M. insipida and M. lancifolia) ....... 110

_ Lower leaf surface glabrous (glabrescent), or pubescent; surface not or but faintly

papillose (papillae either present but extremely small, very closely set and mutu-

ally appressed, or extremely irregular in shape and size) ............... 3
. Nerves making a comparatively sharp angle to the midrib, c. 45°. [Male flowers
nouknowmn: | — si Central; New Guinea © 2). ...1) ..6 ie 12 101. M. papillatifolia
MNeties witha wider angle tothemudnib!... 2.) Jao. S282 eee ee ee 111

_ Male buds narrow, 4—4.5 mm long, with hairs 0.1 mm; pedicel about as long as

the perianth or somewhat shorter; androphore glabrous. [Anthers 5 or 6. Lateral
nerves on lower leaf surface faint, comparatively dense.] —New Guinea: Papua
Barat (Bird’s Head, N Papua Barat); Papua New Guinea (E to Madang Prov.);
Fino ioluccasneswie sh .4-celetie Feeds, seh Oe 78. M. lancifolia

_ Male buds slender or not; pedicel considerably shorter than the perianth; andro-

phore glabrous or pubescent in the lower half ..............0..000055 12
Twigs delicate, 1—-1.5 mm diam. Lateral nerves on lower leaf surface faint, com-
paratively dense. Male buds + slender, 3.5-5 mm long, 1.5—2 mm wide, with
hairs 0.1 mm; androphore pubescent in the lower half; anthers 3 (or 4). — Papua
Nery aGuInea? 20) (nhe v1 Lexar Yate ee Sth Aae Bee eee 25. M. concinna

. Twigs 1-2 mm diam. Nerves distinct below, rather far apart. Male buds + ellip-

soid-oblong, 4.5—-6 mm long, 2—3 mm wide, with hairs (O.1—)0.2—0.4 mm; andro-
phore glabrous or pubescent towards the base; anthers 6-8. [Fruits ellipsoid, 2.5—
3.5(—3.8) cm long, hairs 0.3-1 mm.] — S New Guinea, also S Moluccas and
IVPAStraliays ssttehst Ns SR Wee Sas Ly eae ee gr tee 70. M. insipida

. Twigs delicate, 1-1.5 mm diam.; nerves distinct below, far apart. [Fruits 3 cm

long, subsessile, hairs 0.1 mm.] — Local endemic of southern Papua New Guinea

BREE Pe ote ts 34s | Seed ee Tl leer: 3. ea. ee ee 13. M. brachypoda
Leaves with c. 25 nerves per side, faint and closely set. [Male flowers not known. ]
BS ROSSELTCIISE. 25, De he. ce ee st ae SR OP PoE 98. M. ovicarpa

. Leaves with c. 10 nerves per side. [Male flowers not known.] — SW Papua Barat;

lewmlandintec Meee iw Shee I eae 54. M. fusiformis

, eaves with'en20inetves pensideyor less| 492 --reiee eee - eS eee 114

Flora Malesiana, Ser. I, Vol. 14 (2000)

. Vegetatively difficult to define, distinguishable by its fruit characters: fruits large,

+ pear-shaped, 6-7 cm long, with thick woody pericarp, 10 mm thick; fruiting
pedicel short and thick, often wider than long, 6—9(—10) mm wide. — W Papua
New'!Guinéa! ier. Sse: Oe een. ea 45. M. firmipes

. Fruits generally (much) smaller, with less thick pericarp, and proportionally more

slender peduncle. [Fruits not known in M. schlechteri.] ...........+--- 115

. Leaves below with (sub)persistent, mostly dense indumentum .......... 116
. Leaves below glabrous or glabrescent, or late glabrescent, hairs scale-like, 0.1

mm or less. [Venation (sometimes faint) reticulate. Twigs angular or not. ] 120

. Leaves 4-14 cm long. Male buds 3 mm long. —West New Guinea and Moluccas

UGH AWD Sas 8... ci. s chad loans ses tend oe ee ARE 80. M. iepidota subsp. lepidota

. Leaves 10 cm long or more. Male buds, where known, 5 mm long or more 117
_ Indumentum of lower leaf surface subpersistent, of silky, appressed hairs; vena-

tion + trabeculate. [Twigs with two sharply raised lines. Male flowers not known. ]
— E Papua New Guinea (Northern Prov.); montane forest; 1600 m.........-..
Py et ie Attala se, foie cere ae et ey Soke gma tek st) See Ae! 17. M. byssacea

_ Indumentum more scaly or felty; venation reticulate or hardly visible .... 118
118a.

Male buds 5 mm long. Indumentum of lower leaf surface felty, composed of short
+ stellate hairs of mixed sizes (check also M. chrysophylla and M. lasiocarpa). —
Widespread (small-leaved forms; for differences see lead 167 in this key)......
Pee RL Lc OO) Set ak ALS CN) 72. M. inutilis and 130. M. subcordata

_ Hairs on lower leaf surface 0.2—0.3 mm long. [Male flowers not known.] — Papua

New Guinea (West Sepik Prov., Telefomin subprov.); c. 2000 m 23. M. coacta

_ Indumentum of lower leaf surface dense, hairs very minute, scale-like. [Male

flowers not known, species only distinguishable by fruit characters.] ..... 119

. Fruits 2 cm long; fruiting pedicel comparatively slender, 1.5 cm long. — Papua

NewsGuineal(Madang Prov.) Ge: 120280) eiegink 2 ote 103. M. pedicellata

. Fruits larger; fruiting pedicel (peduncle) shorter. — Papuan Islands...........

AMM leprae iit mere wot eal, Cole Ree LATE MOA, Ae eee 136. M. tenuivenia

_ Leaves membranous, lateral nerves thin and faint on both surfaces, venation al-

most invisible; lower surface grey-white. Bark of older twigs corky, longitudi-
nally fissured. [Male flowers not known.] — Local endemic of the border area in
Céntral’S New! GuineatcehilsO) mit eke ME See 90. M. mediterranea

_ Leaves variable, nerves and venation below generally stouter, clearly visible. Bark

of older twigs either smooth, striate, or flaking, but not corky .......... 121

. Species distinguishable by its fruits; small tree, 5 m, leaves drying greenish; fruits

subsessile. [Male flowers not known.] — Local endemic of Papua New Guinea
(Central Prov.); at low altitude. (Compare also M. insipida) 15. M. brevistipes

. Fruiting pedicel longer or thicker (fruits not known in some species) ..... 122
. Male buds much elongated, oblong or + tubiform (cylindrical), slender, (6—)7 mm
long or more, at least 3 times longer than broad .............-+--0005- 123
. Male buds variable in shape, ovoid, (narrowly) ellipsoid, or nearly tubiform,
7 mm long or less, at most 3 (or 4) times longer than broad ..........-- 132
. Flowers glabrous or almost so; androphore glabrous .........---++++-- 124

. Flowers sparsely or densely pubescent, sometimes hairs very minute, thus seem-

ingly glabrous, occasionally with a few scale-like hairs towards the apex of the
perianth only; androphore wholly or partly pubescent (lens!) .........-. 28)

De Wilde — Myristicaceae (Myristica) 38)

W

124a.

126a.

Male buds 8.5—11 mm long; bracteole persistent. — Papua Barat and western part
of Papua New Guinea; lowland, up to 250m ............. 139. M. tubiflora

. Male buds 7 mm long; bracteole caducous. — Eastern Papua New Guinea (local

Sidemicon Lae subprov.); O=15 mW 284 22 44. M. filipes

. Lower leaf surface with nerves and venation (much) contrasting in colour 126
. Nerves not much contrasting in colour; venation usually inconspicuous. [Male

buds narrowed at the apex, tacute, perianth inside not warty.| NW Papua Barat;
eRe WACUINEA te es eee ee eee tee Sn ee 149. M. warburgii
Male flowers not known, but presumably perianth inside in the basal part not
minutely warty. Two taxa, pending more material at present only satisfactorily
Erotic NONMETUNTE CIALACLOLS os hee ee ae es es eo ope cas ogee 7 merece meee 27

. Male buds + blunt; perianth inside in the basal part with thickened and minutely

BPAUVASMIACE. 2 to ct cit ey AM ec cee cress Sroce ois iste ses + oho) a ee 128

. Fruiting pedicel 10-15 by 2 mm.— Papua New Guinea (Eastern Highlands Prov.);

GS. CC. Cuttir eae ae asc eae a aren oa ae ee 149. M. warburgii subsp. hybrida

. Fruiting pedicel 25-35 by 1 mm. — N Papua Barat; c.500m................

TAR eather yn Prorat ey nee peeicag: ace, ey-t ae ae 58. M. gracilipes

. Twigs 1 mm diam. Pedicel of male flower 6-10 mm long, slender ....... 129
. Twigs medium, 2—3 mm diam. Pedicel of male flower 5-7 mm long ..... 131
. Flowers with thick-woolly indumentum; synandrium subsessile. — Papua New

iGmincan MOTODE PlOVs)) fcc. castes cas eo eae Oe eee ee oe 120. M. schlechteri

. Flowers with minute indumentum; synandrium long-stalked ........... 130
. Male buds 7-10 mm long, 2 mm wide; pedicel 7-10 mm long, bracteole caducous.

= teaeapua New Guineas.) i). etl.) Aue eek eee A Pe 111. M. pumila

. Male buds 7 mm long, slender, 1(—1.5) mm wide, acute; pedicel 6 mm long, brac-

teole persistent. — E Papua New Guinea (Central Prov.) . 127. M. sogeriensis

. Plants from lowland and foothill areas, up to c. 300 m. Fruiting pedicel long. —

baeapnaNew GUINEA. 6... 5 SSP AGL an wakes oe ween 49. M. flosculosa

. Plants from montane areas; 600-2000 m. Fruits without or with a short pseudo-

stalk; fruiting pedicel short or medium-sized. — Throughout New Guinea......
3 gare bth pal aeelndms re Mapa, TaN ebro Metis ati ara i 3! 27 30. M. crassipes

male wuds 3:5—4:5 bys! mm, + tubiorl . © oe oe oe 133
. Male buds larger, 5-7 mm long, narrow or not. [Trees of variable sizes.]_ . 135
. Small tree or shrub, 1.5—8 m. [Anthers 3.] — E Papua New Guinea...........

Mee MOF) MAN eek meee, cate acc oar ae Seed be AO cae ee 134
BeATithers 5.— W. New Guinea. also.Moluccas: ..:.c...2 sees ee ea ie

a OTe ee ae eee eee 80. M. lepidota subsp. montanoides

. Anthers 6—9. [Leaves usually more than 15 cm long; twigs + angular. ] — Through-

outeNew. Guinea: lowland 66. sess on cddes cken nore ie eee 132. M. sulcata

. Leaves generally (thickly) chartaceous or coriaceous; tree of montane areas,

ClOOO mM altitud&ionmore=..c Sse sees eee eee eee 136

. Leaves generally membranous or (thinly) coriaceous; tree of lowland or sub-

montane areas, up to 1000(—1500) m. [Bracteole small, caducous or not; androphore
pubescent, often only in the lower half, or subglabrous.] ............... 138

Flora Malesiana, Ser. I, Vol. 14 (2000)

36a. Flowers short-pubescent; bracteole apical .............. 002s e ee eeee 137

137a.

138a.

142a.

143a.

144a.

. Flowers almost glabrous; bracteole small, caducous, the scar 3 mm below the

perianth. [Androphore glabrous or with extremely small hairs towards the base,
thus seemingly glabrous. Leaves pale brown below, glabrous, nerves flat or sunken,
not contrasting. Fruits ellipsoid or fusiform.] — E Papua New Guinea; montane
OME EL: Lie la EO hei i ae orale aAlotarse patie olathe 76. M. laevifolia
Bracteole (male flower) persistent; the androphore almost completely pubescent.
Leaves pale below, indumentum consisting of minute scale-like hairs, late glabres-
cent: nerves distinct, contrasting in colour.— W Papua Barat (Wissel Lake area);
TOOO SUR OO Sas cic Seer oe eat i IN cs eRe Sate Re 148. M. vinkeana

. Bracteole (male flowers) caducous; the androphore with scattered, extremely mi-

nute hairs less than 0.1 mm, or glabrous. Nerves on lower leaf surface contrasting
in colour or not. [Fruits globose, with thick pericarp.] — E Papua New Guinea;
GOO tah ea es SAAS SR Da oka: 108. M. polyantha
Leaves below pale, grey-glaucous, with darker contrasting nerves and coarsely
reticulate venation. Male buds cleft 1/4—1/2. [Fruits with persistent short hairs

O1=0:5E0:8) mm — Widespread? Ri wien eek ake ot yene eee 57. M. globosa
_ Leaves below pale or not; venation not much contrasting. Male buds cleft c. 1/6-
[ferent ABA AN LED ie INCE SSeS EN EPPO Aes Ske cer feds ae 139
. Male pedicel 1—3 mimi'lomg i. ae eee Se Shai i terete 140

. Male pedicel more than 3 mm long. [Bracteole persistent. Fruits short-haired.] . .

Mat DA ISR eck Be Lee Wicd ate, fae: COs te 141

_ Indumentum of flowers short; bracteole caducous. [Fruits rough-haired.] — NE

Papua Barat, N Papua New Guinea; lowland .......... 16. M. buchneriana

_ Indumentum of flowers conspicuously long-haired and woolly; bracteole minute,

persistent. — E Papua New Guinea; lowland .......... 20. M. chrysophylla

. Petiole proportionally long, 20 mm. Bark of older twigs blackish, conspicuously

cracking. Lower leaf surface with minute indumentum, late glabrescent. Andro-
phoreisubglabrous. .......20.00..4 ese 2 eee s Se eee 142

. Petiole usually shorter, 12-20 mm long. Bark of older twigs striate, rarely crack-

ing. Lower leaf surface glabrous (early glabrescent). Androphore almost wholly
or partially pubescent. — Moluccas; New Guinea: Papua Barat; Papua New Guinea
(E to Sepik Prov., and Louisiade Archipelago) ...........--. 138. M. tristis
Leaves oblong-lanceolate; lateral nerves faint below. Bracteole (male flower) small,
much shorter than the perianth. — Papua Barat (Bird’s Head); Papua New Guinea
(WesterntProy:): lowlands sy 3sesc Siete. car ernie een 7. M. atrocorticata

. Leaves (elliptic-)oblong; nerves distinct below. Bracteole (male flower) about as

long as the perianth, 4 mm. — Papua New Guinea (Western Prov.); swampy low-

[anid forestier UG ET. sa 8 ye ad cictene reed ter ours ebever abate 71. M. inundata
Inflorescences with a distinct, smooth peduncle, slightly flattened or not, (in male)
5 mm long or more; inflorescences branched or not ........--++++++-- 144

_ Inflorescences (sub)sessile, a wart- or worm-like, simple or forked, scar-covered

brachyblast, sessile or subsessile, or up to 5 mm pedunculate (e.g. in M. sphaero-
SDEPAIG TAs LR OE LN LA EERIE, ete osc a etebenenr 154
Leaves glabrous or glabrescent below, or with sparse minute scale-like hairs not
touching each other, not forming a dense felt-like layer (flossy hairs 1-2 mm long
remaining on lower midrib in M. uncinata) ........- 0506 eee eee eee 145

De Wilde — Myristicaceae (Myristica) 395

148a.

149a.

152a.

. Leaves with persistent indumentum below, or late glabrescent, but some (short,

dense) indumentum usually remaining on and near midrib ............. 152
mecavesipapillosebelowi(lens!) eb seeeel eis waepros ae Welsh) tae tere 146
Mifare comompapillosenh: Wii facwh tia Melee oe set... See ee ee 147

. [Flowers not known.] Peduncle of infructescence stout, 5-10 mm long, 4-6 mm

thick, not flattened; fruits 5-6 cm long, pubescent. — Papua New Guinea (West-
eqauarovt) slowland imland foOresteic ec sige. See ete es oo ew 97. M. ornata

. Peduncle slender, 10—40 by 1.5—3.5 mm, + flattened; fruits 2-3.5 cm long, gla-

brescent. [Mature male and female buds in the apical part not or but faintly angu-
iam widespread: mainly coastall...c.4-.4.-.-.4.-. 121. M. schleinitzii

. Mature male and female buds not angular in cross section. Fruits variable 148
. Apical part of mature buds angular. Fruits 4.5—6 cm long, with minute indumentum,

glabrescent. [Whole plant early glabrescent. Veins of leaf hardly visible.] — North-
ERMAN AB ARAL bot. gcste cot pass at teaeeadotuoer: ate ait: 55. M. garciniifolia
Leaves whitish below, glabrescent except lower midrib. Leaf bud often uncinate,
densely pilose, hairs 1-1.5 mm long. Fruits 6-7 cm long, apex often uncinate; in-
dumentum dark brown, persistent, with hairs 1—1.5 mm [Male flowers not known. ]
— Papua New Guinea (Central Prov., near Boridi); 1300-1400 m............
a eer re ee yes OAs Pegs ee 143. M. uncinata

. Leaves glabrous below, whitish or brownish. Leaf bud straight, its indumentum

and that of fruits shorter. — Papua New Guinea; lowland .............. 149
Leaves conspicuously pale, whitish below. [Leaves (thinly) chartaceous; nerves
flat and faint above, venation obscure on both surfaces. Hairs of leaf bud 0.5 mm.
Male buds 7 mm long, obtuse; anthers 15. Fruits + ellipsoid, S—7 by (3—)4-4.5
cm, with rusty hairs 0.5—1 mm.] — Papua New Guinea (Morobe, Northern Prov.)
se 0 a ee ee hE one 142. M. umbrosa

mlbeavies, Crey-DrOWM DOLOW 355 oes cet 2g ake ee 150
150a.

Male buds with acute apex. [Leaves membranous; lateral nerves and venation
moderately distinct above; hairs of leaf bud 0.3-0.5 mm. Male buds 8-10 mm
long, anthers 12—14. Fruits oblong, + fusiform, 4—4.5(—5) by 1.5—2 cm, with sub-
(persistent) rusty hairs (0.2—)0.5 mm.] — Papua New Guinea (SE Morobe, North-
ESTTFIM EA GOA) sess oe earet sd cans Suny Sure: Soe oes od OUR Ya eon EN eee a 19. M. carrii

mviaie buds with rounded APEX. so ciees os ee eae 151
iSyikew

Male buds ellipsoid, 9-14(—16) mm long; anthers 12—20. Leaves membranous;
lateral nerves and venation on lower leaf surface obscure; hairs of leaf bud 0.5—1
mm. Fruits 6-8 cm long, with minute brown indumentum, glabrescent. — Papua
New Guinea (Morobe, Northern, Central, Milne Bay Prov., including Normanby)
«ol ep cicbeeaaneae arene tet au Sola tit wee arta eh ae th 61. M. hooglandii

. Male buds ovoid-ellipsoid, 7 mm long, obtuse; anthers 8 or 9. Leaves thinly char-

taceous; lateral nerves and venation contrasting below. Hairs of leaf bud 0.1
(—0.3) mm. Fruits 5—5.5 cm long, glabrous, green. — Manus I. 109. M. psilocarpa
Upper surface of leaves drying blackish brown, lower surface with pale, whitish
brown, floccose indumentum. Male buds ellipsoid or obovoid, 7-9 by 5—8 mm;
pedicel 7-12 mm long. Fruits subglobose or ellipsoid, 3-5.5 cm long. [Pericarp
4-9 mm thick, with hairs (0.5—)1—1.5(—3) mm. Inflorescences sometimes grouped
into synflorescences.] — Eastern Papua New Guinea .. 87. M. markgraviana

Flora Malesiana, Ser. I, Vol. 14 (2000)

154a.

[57a:

. Leaves drying brown or olivaceous-brown, lower surface brown-pubescent or

glabrescent. Male buds ellipsoid-oblong, 8-10 mm long, narrower; pedicel 3—5
mm long. Fruits generally smaller. — Papuan Islands ..........++--+-- 153

_ Leaves subcordate at base; lower leaf surface with dense subpersistent indumen-

tum. Sterile apex of synandrium consisting of protuberances 0.1 mm long only,
equal in number to the anthers. Fruits 2—2.5 cm long, pericarp 1(—2) mm thick. —
RO SSEU Mee ek re ee te ER RH ah ee steered Say ie ts anes 65. M. incredibilis

. Leaves glabrescent, rounded at base, not cordate. Sterile apex of synandrium sim-

ple, | mm long, + acute. Immature fruits 3 by 2.5 cm, pericarp 5—6 mm thick. —
atu ale ates oy i bi ieee eaar ite ait fase nine na ae ttcts 69. M. inopinata
Leaves below either with persistent short-felty indumentum (hairs scale-like), or
with persistent indumentum of conspicuous hairs; old leaves may have become
glabrous. Fruits variously pubescent .........-- ++ +--+ eee eee tees 155

_ Leaves below glabrous or glabrescent, or with + open indumentum, hairs minute,

scattered (spaced), and seeming glabrous. Fruits pubescent or glabrescent. [Leaves
below sometimes with dense scale-like hairs, late glabrescent, in M. hollrungit.|
© Rite Mab LA Beebe dF Bubreaian ioe bahia chr ag BOURSES ACCS O32 oe 169

_ Leaves below papillose (lens!) in between the spaced, to 1.5 mm long, dark brown

ETERS erty ech tae 20% Eta ate nek Selene Mee Men Ss 0) tel ahah oc 156

_ Leaves either not or but indistinctly or irregularly papillose below; hairs contigu-

ous: on much smallem 204. ss eee se tas oe ee 157

. Twigs 4-5 mm diam. Hairs of lower leaf surface of mixed sizes, 0.5—1.5 mm

long. Bracteole on female pedicel subapical. Fruits oblong, 6-7 cm long [dry
pericarp 4-5 mm thick, hairs 1—1.5 mm.] — C & E New Guinea . 53. M. fusca

. Twigs 2.5-3 mm diam. Hairs of lower leaf surface mixed, of two types: short

ones, 0.1 mm, intermixed with slender long ones, | mm. Bracteole on female
pedicel about median. Fresh immature fruits said to be 7 by 5 cm. — Papua Barat
(Bindestte aa ee tat teh ahert ort tethee cnet cht pet cn cas eno aa 64. M. inaequalis
Indumentum of lower leaf surface felt-like, of two kinds of hairs, a dense layer of
short matted hairs 0.1—0.2 mm, intermixed with numerous emergent hairs of vari-
able sizes, the longest | mm or more, or indumentum composed of only long hairs
[MM Or IMOLe «02. fo oe whe cme pe clage > tasceue* Re he eis cho: cee er 158

_ Indumentum either dense, short, felt-like, or more open, or indumentum flocculose

by longer emergent hairs less than | mm, or without emergent hairs ..... 161

_ Fruits acorn-like, depressed at base, 2 cm diam. Leaves without sunken nerves.

(Flowers not known.] — E Papua New Guinea; lowland . 113. M. quercicarpa

_ Fruits different, not depressed at base. Leaves with nerves sunken on upper sur-

NS A ee ee ee ee OS ee Me OS Ue 159

_ Flowers and fruits with hairs 1 mm or more. — E Papua New Guinea; submontane

ot low landie. it contttichome eerie niet as Saar 20. M. chrysophylla

- Flowers and fruits with hairs 1 mm or less. Tree generally of montane areas 160
. Fruits large, globose, 6-9 cm diam.; fruiting pedicel short, 3-5 mm long. Leaves

below with long hairs, 0.1—1(—2) mm. [Male flowers not known.] — Papua New
Guinea (Eastern Highlands Proy.); montane forest at 800-2300 m........----
pee yh gi cs RT OS RE NR ar ee EE ce 3 BIE 151. M. womersleyi

De Wilde — Myristicaceae (Myristica) 397

b.

16la.

162a.

166a.

Fruits large, somewhat elongate, 6—8 cm long; fruiting pedicel longer, 7-15 mm
long. Leaves below with shorter hairs, 0.1—0.2(—1) mm. — Northern part of New
Guinea; (sub)montane forest; (300—)800-1800 m_ ... 128. M. sphaerosperma
Indumentum of lower leaf surface inconspicuous, consisting of a dense silvery-
brown layer of very short, appressed scale-like hairs, sometimes late glabrescent
(lens!). Fruits 2.5—-3.5 cm long, with very short, mealy, grey-brown or yellowish
brown hairs 0.1 mm or less, glabrescent. [Bark of older twigs often blackish,
often flaking.] — S Papua Barat, Papua New Guinea; lowland...............
a nk 2 5 NR AS RN KO EE AS Ea 144. M. undulatifolia

. Indumentum more conspicuous, hairs either minute, dense, or longer and dense

pamseairered 2!) HL ROAGE F) VOGiGoly 2200S | oR eee aes 162
Leafy twigs stout, 4-8 mm diam. Male flowers large, firmly carnose or leathery,
buds 8 by 5 mm. Fruits large, 9-12 by 7—8.5 cm, pericarp 20 mm thick; aril small,
the laciniae covering only the basal part of the seed. — Central Papua New Guinea;
Biptieane a OO SOU mits <8 em gar.) lp. ett. Fee oe ee 67. M. ingens

. Twigs 2-6 mm diam. Flowers and fruits not as above. — Lowland or lower mon-

tane; 0Q—1200(—1800) m (M. subcordata var. gigacarpa at 1200-1800 m)... 163

. Fruits large, 6-8 cm long, pericarp 5—12 mm thick. — Northern part of New

ieinea: (SUb)MOMdNe ves. ats he te oe 128. M. sphaerosperma

. Fruits generally smaller, pericarp thinner (fruits not known in M. simulans) 164
. Male buds cleft nearly halfway. [Androphore distinctly shorter than the synan-

drium; anthers 6—10. Fruits 4-7 cm long.] — Moluccas and possibly Papua Barat
en. ee cee cote: tes sh ycgGeeDR Ee oe. S : ee 43. M. fatua

mide puas clett /4—1/3 (or less) — New Guinea 2+. ..-- ee ee 165
. Leaves below with weak indumentum of minute sparse grey hairs, largely glabres-

cent; venation on lower leaf surface indistinct. [Male buds 5—6 mm long; andro-
phore much shorter than synandrium; anthers 7 or 8. Fruits 3—4.5(—6) cm long,
with rough hairs 0.5—1 mm.] — Northern New Guinea; lowland.............
ee ee | Eas Bn, 6 NS acu ate 16. M. buchneriana

. Leaves below with indumentum of densely interwoven hairs (hairs scale-like or

not); venation on lower leaf surface obscured by indumentum .......... 166
Male buds 8(—10) mm long, hairs | mm long; androphore much shorter than the
synandrium, glabrous; anthers 7 or 8. [Flowers with hairs | mm. Fruits not known. |
— E Papua New Guinea (endemic of Modewal.) ......... 124. M. simulans

. Male buds 12 mm long or less, hairs 0.1—0.5 mm; androphore about (or nearly) as

long as the synandrium, mostly (partly) pubescent ................... 167

. Leaves 12—24 cm long. Male buds ellipsoid-oblong, 34.5 mm long. [Bracteole

persistent; anthers (3—)5. Fruits variable in size, 2.5—5 cm long; fruiting pedicel
1-3 mm long.] — Widespread ............. 72. M. inutilis (subsp. papuana)

leaves 12—35,cm.long...Male. buds larger 9.5 as ewereekin ae clea cks 168
. Male buds + narrow, almost tubiform, (6.5—)7—8(—10) mm long; bracteole cadu-

cous; anthers 5 or 6; perianth towards base smooth inside. [Fruits (3.5—)4—5.5 cm
long; fruiting pedicel 1-10 mm long (fruiting specimens can be confused with
M. inutilis).| — Throughout New Guinea; lowland and lower montane........
Pe, eR nee ee ete ee ee eo ee 130. M. subcordata

398 Flora Malesiana, Ser. I, Vol. 14 (2000)

b. Male buds + oblong, 10-12 mm long; bracteole (sub)persistent; anthers 8 or 9;
perianth finely warty towards base inside. Fruits not known. — W Papua Barat

(limestone) USS aster ee PAN LOR RG 146. M. verruculosa

169a. Leaves papillose below. Bark of older twigs blackish, flaking. — Papua Barat,
Papua New Guinea; lowland, scattered ............+++--- 73. M. kalkmanii

b. Leaves not or indistinctly papillose below. Bark of older twigs brown, not or in-
conspicuously flaking. 2:08) .fisastin Soa). ee fete «1 Se 170

170a. Pedicel of male flowers 3 mm long or less. Fruits with conspicuous hairs 0.5 mm
Gismore AAA. Sta es ee as os CR oe 171

b. Pedicel of male flowers more than 3 mm long. Fruits with much shorter hairs,
persistent or glabrescent. [Twigs often lined or angled; no ant-swellings present. |

ahvels sata sees Gerloe wt eee Se els nes Ho Sek i?

171a. Pedicel of male flowers 2-3 mm long; flowers with hairs 0.2—-0.3 mm. Fruits
3—4.5(—6) cm long, hairs 0.5—1 mm. — Northern part of New Guinea; lowland

ee ee ee en 5 ate ee eee ere 16. M. buchneriana

b. Pedicel of male flowers 1-2 mm long; flowers with hairs 0.5—1.5 mm. Fruits 2.5—
4.5 cm long, hairs 1.5—3 mm. — E Papua New Guinea; up to 430) ii ets eae
Le ee I ee ee os 20. M. chrysophylla subsp. entrecasteauxensis

172a. Twigs (at least at apex) angled, 2-4 mm diam. Leaves usually dark brown. Male
buds 4 mm long. Fruits subglobose or broadly ellipsoid, (3-)3.5—4 cm diam. —
Papua Barat, Papua New Guinea; lowland .........-..---- 132. M. sulcata

b. Twigs subterete, often lined (often + hollow and somewhat lined, rarely faintly
winged), not angled, 2-5(-10) mm diam. Leaves + olivaceous. Male buds 4—6
(—6.5) mm long. Fruits subglobose or (globose-)ellipsoid, 2.5—4.5(—5) cm long,

often short pointed at the apex. — Widespread in New Guinea (incl. Bismarck
Archipelago); lowland: «is. 0 ect ts 60. M. hollrungii

c. Twigs + angled or winged, 4—8(—15) mm diam. Leaves olivaceous-brown. Male
buds 10-13 mm long. Fruits + fusiform or ellipsoid, 5~7 cm long. — Bismarck
Geli ofsl Cel Ow een Me ies Un boi eno een em Pees et 10. M. bialata

1. Myristica agusanensis Elmer

Myristica agusanensis Elmer, Leafl. Philipp. Bot. 8 (1915) 2775. — Type: Elmer 13284, Mindanao.
For more references and synonyms see the subspecies.

Tree 10-20 m. Twigs 1.5—3 mm diameter, early glabrescent, hairs pale rusty, 0.1 mm;
older twigs striate, not flaking; lenticels absent or few. Leaves membranous or thinly
chartaceous, elliptic-oblong or oblong-lanceolate, 9-29 by 2.5—7.5 cm, base short-atten-
uate, apex gradually narrowed, acute-acuminate, or bluntish; upper surface olivaceous-
brown, lower surface often greyish, pubescent or glabrescent, hairs lax or dense, 0.1
mm, with or without few scattered emergents 0.2—0.3 mm; papillose; dots absent; lateral
nerves 10-18 per side, at (30—)45° to the midrib, above flat or sunken, below with lines
of interarching and venation sometimes faint; petiole 10-30 by 1—2(—3) mm; leaf bud
10-15 by 1-1.5 mm, with hairs 0.1 mm. /nflorescences pedunculate, sometimes com-
pound (see under the subspecies), axillary to leaves or to caducous cataphylls, with hairs
0.1-0.3 mm: bracts 11.5 mm, caducous; in male: twice branched, 3-6 by 3-5 cm, pe-
duncle to 20 mm, + angled or flattened, branches usually subopposite, with clusters

De Wilde — Myristicaceae (Myristica) 399

Fig. 59. Inflorescences of some species of Myristica. — a. M. hollrungii Warb.; b. M. fragrans Houtt.,
female; c. M. cornutiflora J. Sinclair subsp. cornutiflora; d & e. M. subalulata Mig. var. subalulata,
male & female [a: BW 9399 (Schram); b: Blume s.n.; c: BW 6413 (Kalkman); d: Hoogland 3982;
e: NGF 11018]. — Scale bar for all = 2 cm. — See also Fig. 60.

of 2-7 flowers, buds of different sizes; female inflorescences: few-flowered, 1—1.5 cm
long, peduncle 0.5—1 cm; flowers with dense grey-brown hairs 0.1—0.3 mm. Male flow-
ers: pedicel slender, 2-3 mm, bracteole broadly ovate or subcircular, 1—1.5 mm, caducous;
buds ellipsoid, not angled, 3(—3.5) by 2 mm, apex rounded, base attenuate to rounded,
cleft 1/3, lobes suberect or spreading, 0.2-0.4 mm thick. Androecium 2.5 mm; androphore
1 mm, subglabrous or for the larger part with pale hairs less than 0.1 mm; synandrium
+ oblong, (1—)1.5—2 by 0.6—0.7 mm, thecae 6-10, sterile apex blunt, to 0.1 mm, glabrous
or with a few minute hairs, or central column to 0.2 mm hollowed. Female flowers:
pedicel 1.5 mm; perianth (broadly) ovoid, 2 mm; ovary ovoid, 1.8 by 1 mm, appressed
short-pubescent. Fruits single or in pairs, subglobose, ellipsoid or oblong, 3—4.7 by
2-3.5 cm, + glabrescent, hairs rusty, 0.1 mm or less; pericarp 3—4 mm thick; seeds 2.5
cm; fruiting pedicel 3—5 mm.
Distribution — Malesia: Philippines (Luzon to Mindanao).

400

Flora Malesiana, Ser. I, Vol. 14 (2000)

cu i
My

ZY, Ss
Ge NX Se == ! Ly
We \ 4
: Ly ne ”/
((
a b

Fig. 60. Inflorescences of some species of Myristica. — a & b. M. maxima Warb., female & male;
c. M. umbellata Elmer, male; d. M. iners Blume, female [a: Sinclair 10604; b: Ridley 3363, c: Elmer
12820; d: Endert 57 E IP 554]. — Scale bar for all = 2 cm. — See also Fig. 59.

KEY TO THE SUBSPECIES

la. Leaves elliptic-oblong to oblong-lanceolate, 9-29 cm; lower surface + late glabres-
cent, at first with dense woolly or arachnoid greyish stellate hairs, with a few emer-
gent pale brown hairs. Male inflorescences axillary to leaves. Flowers with hairs
OalesO: Dram. serh Bahia aera ert ET hd a. subsp. agusanensis
b. Leaves oblong-lanceolate to lanceolate, 9-13 cm; lower surface with scattered, minute
brown scale-like hairs 0.1 mm or less. Male inflorescences sometimes axillary to
reduced leaves, but then vegetative apical bud of twig present. Flowers with hairs
02=0:3) minty; . és ch Bese eA LRA Leer Oe b. subsp. squamulosa

De Wilde — Myristicaceae (Myristica) 401

== =

=
SS

=
=

SS

Fig. 61. Diverse forms of Myristica fruits. — a. M. subalulata Miq. var. subalulata. — b. M. chryso-
phylla J. Sinclair subsp. chrysophylla. — c. M. villosa Warb. — d. M. fatua Houtt. subsp. fatua. —
e. M. quercicarpa (J. Sinclair) W.J. de Wilde. — f. M. tristis Warb. subsp. tristis [a: BW 5557 (van
der Sijde); b: Hoogland 3642; c: SAN 15253; d: Kuswata & Soepadmo 14, e: NGF 1270; f: Hoogland
4393]. — Scale bar for all = 2 cm. — See also Fig. 62.

Fig. 62. Diverse forms of Myristica fruits. — a. M. womersleyi J. Sinclair; b. ditto, longitudinal sec-
tion. —c. M. maxima Warb. — d. M. fusiformis W.J. de Wilde subsp. pseudostipitata W.J. de Wilde.
—e. M. pumila W.J. de Wilde. — f. M. laevifolia W.J. de Wilde. — g. M. crassipes Warb. subsp.
crassipes [a & b: Womersley 11374; c: FMS 12132 (Strugnell); d: von Romer 304, e: Hoogland 3971;
f: Carr 13239; g: Carr 14897]. — Scale bar for all = 2 cm. — See also Fig. 61.

402 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 63. Fruits of various small-leaved Myristica species. — a. M. brachypoda W.J. de Wilde, peri-
carp glabrescent [NGF 8079]. — b. M. brevistipes W.J. de Wilde, pericarp short-pubescent [Darby-
shire 929]. —c. M. flosculosa J. Sinclair; note long slender fruiting pedicel, fruit with pseudostalk
[Guillianetti in McGregor s.n.].—d. M. fusiformis W.J. de Wilde subsp. fusiformis, pericarp largely
glabrescent [Milliken 1408]; e. subsp. pseudostipitata W.J. de Wilde, fruit largely glabrescent;
note pseudostalk [von Romer 304]. — f. M. laevifolia W.J. de Wilde, fruit glabrescent [Craven &
Schodde 1073). —g. M. longipes Warb., fruit minutely mealy pubescent; note stalked inflorescence
[Schodde 2963]. —h. M. sogeriensis W.J. de Wilde, fruit minutely pubescent, with distinct pseudo-
stalk; i. ditto, dry fruit, opened, showing seed [Forbes 396]. — Scale bar for all = 2 cm.

>
Fig. 64. Male flowers of Myristica species from the Moluccas. — a. M. alba W.J. de Wilde [de
Vogel 3975]. — b. M. bifurcata (J. Sinclair) W.J. de Wilde subsp. bifurcata |Kostermans 944]. —
c. M. fatua Houtt. subsp. fatua [Reinwardt s.n.]; d. ditto [de Vogel 3497, from Halmahera], deviat-
ing by the short androphore, completely fine-pubescent. — e. M. fissurata W.J. de Wilde [de Vogel
3836]. — f. M. fragrans Houtt. [de Vriese & Teijsmann s.n.]. — g. M. insipida R. Br. [Schodde in
Hartley 13735, from Arnhem Land, N Australia]. — h. M. lancifolia Poir. subsp. lancifolia |Teijs-
mann 7585, from Bird’s Head, Papua Barat]; i. subsp. montana (Roxb.) W. J. de Wilde [van Borssum
Waalkes 3300]. —j. M. lepidota Blume subsp. lepidota [Aet 279, from SW New Guinea]; k. subsp.
montanoides (Warb.) W.J. de Wilde [Beccari FI 7756]. — 1. M. mindanaensis Warb. |Kuswata &
Soepadmo 57). — Scale bar for all = 2 mm. — See also Fig. 65.

403

)

ristica

De Wilde — Myristicaceae (My

Flora Malesiana, Ser. I, Vol. 14 (2000)

404

SS
ANSE

aS

De Wilde — Myristicaceae (Myristica) 405

a. subsp. agusanensis

Myristica agusanensis Elmer, Leafl. Philipp. Bot. 8 (1915) 2775; Merr., Enum. Philipp. Flow. Pl. 2
(1923) 178; J. Sinclair, Gard. Bull. Sing. 23 (1968) 218, f. 17, p.p.; W.J. de Wilde, Blumea 42
(1997) 146.

Myristica laxiflora Merr., Philipp. J. Sci. 17 (Sept. 1920’, Jan. 1921) 254; Enum. Philipp. Flow. PI.
2 (1923) 179. — Type: BS 15498 (Reillo), Basilan I.

Myristica sorsogonensis Elmer ex Merr., Enum. Philipp. Flow. PI. 2 (1923) 178, in obs. sub Myristica
agusanensis Elmer, nom. nud.; Elmer, Leafl. Philipp. Bot. 10 (1939) 3809, nom. nud.; Kew
Index Suppl. 10 (1947) 149, nom. nud. — Voucher: Elmer 16010, Luzon.

Leaves elliptic-oblong to oblong-lanceolate, 9-29 by 2.5—7.5 cm, apex gradually at-
tenuate, very tip + blunt or acute; lower leaf surface late or early glabrescent, at first with
dense woolly greyish stellate hairs 0.1—0.2 mm, often mixed with a few pale brownish,
slightly emerging hairs. Male inflorescences solitary in the leaf axils, or compound, re-
sembling those of Horsfieldia or Gymnacranthera, paniculate, 3-6 cm, with peduncle to
2 cm, the central branch with up to 6 lateral branches. Flowers with hairs 0.1—0.2 mm.
Female inflorescences (from infructescences) paniculate, 1.5 cm. Female flowers not
seen.

Field-notes — Tree branched at the apex in a dense crown, bark brown, flaking slightly
in small portions; sap red. Perianth inside cream, rust-coloured outside (Sinclair, 1. c.).

Distribution — Malesia: Philippines (except for Palawan).

Habitat & Ecology — Inland forests, at low altitudes (50 m); fl. July—Aug.; fr. through-
out the year.

Note — Sinclair (1.c.) suggested that M. agusanensis and M. guatteriifolia are close,
the former being a miniature replica of the latter, but this suggestion may not be war-
ranted. Myristica guatteriifolia has larger male flowers, with a stouter androecium, more
anthers, and a more hairy androphore, and the indumentum is much longer than that of
M. angusanensis. Myristica agusanensis is not a coastal species like M. guatteriifolia.

b. subsp. squamulosa W. J. de Wilde

Myristica agusanensis Elmer subsp. squamulosa W.J. de Wilde, Blumea 42 (1997) 146. — Gym-
nacranthera lanceolata Merr., Philipp. J. Sci., Suppl. 1 (1906) 55; Elmer, Leafl. Philipp. Bot. 3
(1911) 1058 [non Myristica lanceolata Wall., Cat. (1832) n. 6794, nom. nud. = Knema globularia).
— Myristica lancifolia Merr., Enum. Philipp. Flow. Pl. 2 (1923) 178 [non Myristica lancifolia
Poir. (1816)], nom. inval. — Type: FB 3236 (Meyer), Luzon.

Leaves (oblong-)lanceolate, 9-13 cm, apex subacute or + blunt; lower leaf surface
with inconspicuous widely spaced bright brown scale-like hairs 0.1 mm long or less, late
glabrescent. Male inflorescences axillary to normal leaves or (Loher 6716) axillary to
caducous reduced leaves, and arranged in short shoot-like twigs ending in a vegetative

Fig. 65. Male flowers of Myristica species from the Moluccas. — a. M. pubicarpa W.J. de Wilde
[de Vogel 4000]. — b. M. robusta W.J. de Wilde [de Vogel 3918]. —c. M. sangowoensis (J. Sinclair)
W.J. de Wilde [Whitemore c.s. TCW 3608]. — d. M. simiarum A.DC. subsp. celebica (Miq.)
W. J. de Wilde [Koorders 18161]. — e. M. subalulata Miq. var. subalulata [Aet 16, exp. Lundquist,
SW New Guinea]. — f. M. succedanea Blume [de Vogel 3626]. — g. M. tristis Warb. subsp. tristis
[van Royen 3579, Bird’s Head, Papua Barat]. — Scale bar for all = 2 mm. — See also Fig. 64.

406 Flora Malesiana, Ser. I, Vol. 14 (2000)

bud; + paniculate, peduncle 5-20 mm, central branch short, up to 5 mm, simple or with
two lateral branches. Flowers with hairs 0.2-0.3 mm. Female inflorescences c. | mm,
few-flowered. Fruits not seen.

Distribution — Malesia: N & C Philippines (Luzon, Sibuyan).

Habitat & Ecology — Lower montane forest; fl. Apr.—June.

Note — This subspecies was originally described by Merrill in Gymnacranthera,
apparently on account of the small, ellipsoid flowers, and the inflorescences which are
often compound (subsp. agusanensis) or grouped in short-shoots.

2. Myristica alba W.J. de Wilde
Myristica alba W.J. de Wilde, Blumea 35 (1990) 238, f. 1: 1. — Type: de Vogel 3975, Moluccas.

Tree 4-20 m. Twigs (1—)1.5—2.5 mm diameter, glabrescent, hairs dull brown, 0.1 mm;
older twigs grey-brown, striate; lenticels present but inconspicuous. Leaves membra-
nous or subcoriaceous, elliptic-oblong, 14-22 by 6-8.5 cm, base shortly attenuate or
rounded, apex acute-acuminate, the very tip sometimes + blunt; upper surface glabrous,
dark or blackish brown, lower surface greyish white (possibly due to irregularly shaped
papillae), glabrescent, hairs widely spaced, stellate or scale-like, 0.1 mm long, the stouter
hairs leaving minute dark scars (lens!); dots absent; midrib above slender, slightly raised,
lateral nerves 12-15 per side, at 50—70° to the midrib, + sunken above, on the lower leaf
surface bright dark (purplish) brown, contrasting with the whitish background; lines of
interarching sometimes distinct, venation coarse, faint; petioles blackish, 7-14 by 2-3
mm; leaf bud 8-12 by 2 mm, hairs dense, dull brown, appressed, 0.1—0.3 mm. Inflores-
cences between the leaves, of the Knema-type, sessile, brachyblasts to 5 mm, slender,
with brown hairs 0.3 mm; bracts | mm, caducous; in male: with 1-4 flowers, buds of
different sizes; female inflorescences (from infructescences): similar to males; flowers
with dull brown + appressed hairs 0.2-0.3 mm. Male flowers: pedicel 0.5—1 mm, bracteole
broadly rounded or reniform, 1-1.5 mm, persistent; buds ellipsoid-oblong, 3.5—4 by 1.5
mm, apex acutish, not angled, base + rounded, cleft 1/3—-1/2, lobes 0.2—0.3 mm thick.
Androecium 3 mm; androphore 1.5 mm, hairs dense, pale brown, curved, 0.3—0.5 mm;
synandrium cylindrical, 1.5—1.7 by 0.6-0.7 mm, thecae c. 8, sterile apex minute, irregu-
larly lobed, 0.1 mm, glabrous or with a few pale hairs 0.2 mm. Female flowers not seen.
Fruits solitary, subsessile, variable in shape (see note 2), ellipsoid-oblong or subglobose
(when slightly immature), 3—4.2 by 2.5 cm, apex sometimes shortly beaked, at base
contracted into a thick, 3—5 mm long pseudostalk; hairs dense, scurfy or + woolly, 0.1 or
0.5 mm (see note 2); pericarp conspicuously woody, 5 mm thick; seeds not seen; fruiting
pedicel 3 mm. — Fig. 64a.

Field-notes — Low solitary tree, bole to 30 cm diameter. Leaves whitish glaucous
below. Flowers yellow.

Distribution — Malesia: Moluccas (Bacan, Obi, Buru).

Habitat & Ecology — Primary forest with some undergrowth on deep clayey soil
with scattered limestone boulders, forest on flat land along river, or on steep sloping
terrain, on limestone; 100—500 m altitude; fl. Nov.; fr. Aug., Dec.

Notes — 1. Resembles M. pubicarpa from the same area, M. nivea from the Philip-
pines, and M. impressa and M. impressinervia from Sulawesi, all having whitish leaves
below, later on becoming finely dark-punctate by hair scars (lens!). Myristica alba dif-

De Wilde — Myristicaceae (Myristica) 407

fers from these species in its small, nearly sessile, elongate male flowers. Both fruiting
collections of M. alba have the fruits conspicuously contracted at the base into a short
pseudostalk.

2. The specimen bb 23157 has ellipsoid fruits with dense curly woolly hairs 0.5(—1)
mm. Nooteboom 5369 (Buru) has almost globose fruits, 2.5 cm diameter, with a 3-5 mm
long pseudostalk; the scurfy hairs are very short, 0.1 mm. More collections are needed to
judge the taxonomic significance of these differences.

3. Myristica archboldiana A.C. Sm.

Myristica archboldiana A.C.Sm., J. Arnold Arbor. 22 (1941) 73; J. Sinclair, Gard. Bull. Sing. 23
(1968) 329, f. 47; Foreman, Handb. Fl. Papua New Guinea | (1978) 186; W.J. de Wilde, Blumea
40 (1995) 259. — Type: Brass 6982, Papua New Guinea.

Tree 30 m. Twigs coarsely striate, 2-3.5 mm diameter, blackish, early glabrescent,
hairs less than 1 mm, grey-white; older twigs grey-brown, 5—6 mm diameter, coarsely
striate, then cracking, fissured and flaking; lenticels not apparent. Leaves thinly
chartaceous, elliptic-oblong or obovate-oblong, 8-15 by (3—)5—7 cm, base shallowly
cordate, rounded, or short-attenuate, apex bluntish or shortly acute-acuminate; upper
surface dull blackish brown, glabrous, lower surface (greyish) rusty, largely pubescent,
hairs dense, scale-like, 0.1 mm or less; not papillose; dots absent; midrib slender, in a
shallow groove above, lateral nerves 16-21 per side, at 60—70° to the midrib, slender
and faint, sunken above, not much contrasting below, lines of interarching and venation
indistinct; petiole comparatively long, 20—30(—40) by 1.5—2 mm, blackish, glabrous;
leaf bud 15-20 by 2-3 mm, hairs dense, appressed, grey (whitish), 0.1 mm or less. Male
and female flowers not seen. Inflorescences (from infructescences) between the lower
leaves, peduncle 15-20 mm, coarsely fissured and lenticellate. Fruits (1 or) 2 or 3 per
infructescence, ovoid-oblong, 6.7—7 by 4 cm, apex rounded, tip slightly oblique, + blunt,
base narrowed (pseudostalk absent), hairs mealy, dense, rusty, 0.1 mm; pericarp smooth,
10 mm thick (towards the base nearly 20 mm thick); seeds (nearly mature) ellipsoid,
2.5—3 cm; fruiting pedicels + thick, 3(—5) mm long, bracteole scar not seen.

Field-notes — Large canopy tree with straight bole spurred at base; bark pale brown,
lenticellate, reddish when cut. Leaf blades brown below, ribbed above with impressed
nerves. Fruits rufous-brown, aril yellow.

Distribution — Malesia: Papua New Guinea (Western Prov., at Palmer River, 2 miles
below junction with Black River); known only from the type.

Habitat & Ecology — Common in forest along lower ridges; 100 m altitude; fr. June.

Note — Myristica archboldiana is easily distinguishable from other members of the
group of species with smaller leaves by its usually cordate leaves that are pubescent
below, and its relatively long petiole and the thick-walled fruits. Sinclair (1.c.: 331) sug-
gested a relationship with M. pedicellata and M. tenuivenia on account of the indumentum
on the lower leaf surface and mode of venation, but these species have sessile scar-
covered inflorescences.

4. Myristica arfakensis W.J. de Wilde

Myristica arfakensis W.J. de Wilde, Blumea 40 (1995) 259. — Type: Kostermans 2440, Irian Jaya,
Bird’s Head.

408 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 6-10 m. Twigs 1—1.5(—2) mm diameter, slightly angular, glabrescent, hairs 0.1
mm or less, grey-brown, scale-like; older twigs blackish, striate, but not cracking or
flaking; lenticels not apparent. Leaves chartaceous or subcoriaceous (brittle), + ovate-
elliptic or oblong, 5-12 by 1.5—5 cm, base (short-)attenuate, apex up to | cm acute-
acuminate; upper surface brown, lower surface bright brown, early glabrescent; not pap-
illose; dots distinct; midrib slender, raised above, lateral nerves 10-15 per side, at 60—70°
to the midrib, slender, sunken above, little raised or + flat below, generally with inter-
secondary nerves, lines of interarching and venation indistinct; petiole 8-15 by 1-1.5
mm; leaf bud acute, 5-10 by 1—1.5 mm, hairs brown-grey, less than 0.1 mm. Inflores-
cences axillary or to 4 mm supra-axillary, between the leaves or below, early glabrescent,
indumentum minute, peduncle slightly flattened, smooth, S—10 by 1—1.5 mm, ending in
a short scar-covered brachyblast; bracts minute, minutely pubescent, late caducous; male
inflorescences: brachyblast to 13 mm long, flowers 1—4, buds of different sizes; female
inflorescences: 1- or 2- (or 3-)flowered; flowers with sparse brown-grey hairs 0.1 mm or
less. Male flowers: pedicel 3-4 by 0.5 mm, bracteole broadly ovate, 0.5(-1) mm,
caducous; buds + ellipsoid-oblong, somewhat parallel-sided (or faintly narrowed in the
middle), 3.5—4 by (1.5—)2 mm, apex + blunt, not angled, base broadly rounded, cleft
1/4-1/3, lobes + membranous, 1.5 mm long, 0.3 mm thick. Androecium 3-3.5 mm;
androphore 1.5—2 by 0.5—0.6 mm, glabrous; synandrium ellipsoid-oblong, 1.5 by 0.6—
(.7(—0.8) mm, thecae 10-12, not tightly appressed, sterile apex subtruncate, 0.2 mm.
Female flowers not seen. Fruits generally solitary, ellipsoid-oblong, 3 by 1.5 cm, apex
acute, base contracted into a pseudostalk up to 5 mm long, hairs dull-brown, scurfy, less
than 0.1 mm; pericarp 1(—2) mm thick; seeds ellipsoid-oblong, 2.3 cm; stalk (including
fruiting pedicel) slender, 1—-1.5 cm, bracteole scar 2~3 mm below the apex.

Field-note — Flowers whitish.

Distribution — Malesia: Papua Barat (NE Bird’s Head, Arfak Mts).

Habitat & Ecology — Mountain forest at 1000-1800 m altitude; fl. Oct.; fr. May.

Note — Obviously related to M. pachyphylla, with which it has the brown-black dots
on the lower leaf surface and the general habit in common, but in M. pachyphylla the
dots are small, while in M. arfakensis they are readily visible with the naked eye. Myristica
pachyphylla furthermore differs in its larger, more elongate male flowers and larger
fruits (4 cm).

5. Myristica argentea Warb.

Myristica argentea Warb., Bot. Jahrb. Syst. 13 (1891) 311; Die Muskatnuss (1897) 347; Mon. Myrist.
(1897) 446; K. Heyne, Nutt. Pl. Ned. Indié (1927) 638; Burk., Dict. Econ. Prod. Malay Penins.
(1935) 1523; Markgr., Bot. Jahrb. Syst. 67 (1935) 165; J. Sinclair, Gard. Bull. Sing. 23 (1968)
235, f. 20; Foreman, Handb. Fl. Papua New Guinea | (1978) 187; W.J. de Wilde, Blumea 40
(1995) 260. — Type: Warburg 20717, West New Guinea.

Myristica finschii Warb. [Bot. Jahrb. Syst. 13 (1891) 311, as sp. nov. aff. Myristica buchneriana],
Mon. Myrist. (1897) 534, p.p.; K. Schum. & Laut., Fl. Schutzgeb. Siidsee (1900) 328, p.p. —
Type: Warburg 20715, n.v.; according to Markgraf (1.c.: 169), M. finschii consisted of a separate
fruit of M. fatua var. papuana and a leafy twig of M. argentea.

Tree 5—20(—40) m. Twigs 1.5—2 mm diameter, with minute indumentum, early
glabrescent; older twigs 3-5 mm diameter, grey-black, coarsely striate, with cracks and
conspicuous raised pustulate lenticels. Leaves membranous, elliptic-oblong to oblong

De Wilde — Myristicaceae (Myristica) 409

(-lanceolate), 9-25 by 3-10 cm, base broadly to narrowly attenuate, apex acute-acumi-
nate; upper surface olivaceous, lower surface silvery (brownish), seemingly glabrous,
with a+ rough cobweb-like texture, hairs scattered, grey or pale brown, scale-like, less
than 0.1 mm; not papillose; dots absent; midrib above slender, slightly raised above,
raised below, bright-brown, contrasting in colour with the silvery lower surface of the
blade, lateral nerves 9-17 per side, at (45—)60° to the midrib, slightly sunken above,
lines of interarching distinct or not, venation fine, + scalariform, faint on both surfaces;
petiole 10-20 by 1.5—2.5 mm; leaf bud somewhat curved, + oblique, 6-10 by | mm,
hairs 0.1 mm or less. Inflorescences axillary or to 5 mm supra-axillary, among the leaves,
sometimes axillary to reduced caducous leaves, early glabrescent, paniculate; in male:
3-5 cm, peduncle slender, + flattened, (S—)10—30 mm long, ending in one central flower
and two lateral brachyblasts to 20 mm, with scars of fallen bracts and pedicels, each
ending in a loose cluster of 2—5 flowers, buds of different sizes; female inflorescences:
peduncle 5—10(—15) mm, ending in a single stout-pedicelled flower; flowers with scat-
tered appressed dark brown hairs less than 0.1 mm, denser towards the apex. Male flow-
ers: pedicel slender, 10-17 by 0.6—1 mm, bracteole as a low collar 1 mm high, caducous,
leaving a narrow ring-shaped scar; buds ovoid-oblong or ellipsoid-oblong, 11—12 by
5—5.5 mm, apex subobtuse, not angled, base broadly to narrowly rounded, cleft 1/6,
lobes at anthesis + erect, 0.3 mm thick. Androecium slender, 8—9.5 m; androphore 3 by
0.5—0.7 mm, subglabrous or hairs pale, less than 0.1 mm, scattered towards the base;
synandrium oblong, 5—6.5 by (1—)1.5—2 mm, thecae c. 15, sterile apex 0.3 mm. Female
flowers: pedicel straight, stout, 8-12 by 1-1.5 mm, with scattered wart-like lenticels;
buds ovoid-oblong, narrowed towards the apex, 8-10 by 5—5.5 mm, cleft 1/4—1/5, lobes
2(-2.5) mm long; ovary fusiform or flask-shaped, 6 by 2—2.5 mm, gynophore | mm, all
densely minutely grey-brown pubescent. Fruits solitary, + ellipsoid-fusiform, 6.5—7.5
by 3.5—4.5 cm, apex narrowed, base somewhat contracted into a pseudostalk, 3-8 mm
long, early glabrescent, brown, usually set with conspicuous coarse pale pustules or len-
ticels; seeds 4 cm; fruiting pedicel 8—18 mm.

Field-notes — Stilt-roots sometimes present. Perianth greenish white, with scattered
brown hairs. Mature fruits yellow or brown with brown pustules, 8 by 5—5.5 cm.

Distribution — Malesia: Papua Barat (W Bird’s Head, Mt Cluer Bay area, Geelvink
Bay), locally endemic; elsewhere cultivated, e.g. in the Moluccas (Seram).

Habitat & Ecology — Primary forest, sloping forest; on sandy clay over limestone;
0-250 m altitude; fl. June; fr. May, Aug.

Uses — Cultivated in W New Guinea for its aromatic mace; the nut is less aromatic as
compared to that of M. fragrans, but is said to be used medicinally, especially in Java.

Notes — 1. Clearly related to M. fragrans, but easily distinguishable by its somewhat
larger leaves with silvery undersurface and the coarsely lenticellate-pustulate twigs.

2. Some variation has been found in BW 6346 (Kalkman),with the male buds broadly
rounded at the base and a grooved disc-like thickening around the insertion of the
androphore, features not found in e.g. Atasrip 707.

6. Myristica atrescens W.J. de Wilde

Myristica atrescens W.J. de Wilde, Blumea 40 (1995) 260. — Type: Soegeng Reksodihardjo 424,
SE Irian Jaya.

410 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 20 m. Twigs 1.5—2(—2.5) mm diameter, glabrescent, hairs (pale) brown, 0.1 mm;
older twigs blackish brown, coarsely striate, tending to crack longitudinally, not flaking;
lenticels indistinct. Leaves thinly chartaceous (but conspicuously brittle when dry), el-
liptic-oblong, 7-12 by 35.5 cm, base short-attenuate to broadly rounded, apex broadly
acute(-acuminate) with blunt tip; upper surface dark (olivaceous to) brown, glabrous,
lower surface dark brown, glabrous; not papillose; dots absent; midrib slender, raised
above, lateral nerves 10—13 per side, at c. 60° to the midrib, slender, flat and faint above,
slightly raised, concolorous below, lines of interarching and venation (a coarse network)
indistinct; petiole 12-15 by 1.5 mm; leaf bud 8-10 by 1.5 mm, hairs dense, appressed,
yellow-brown, 0.2(—0.3) mm. Inflorescences among the leaves, glabrescent, hairs 0.1-
(0.2 mm; in male: peduncle 5—7 mm, ending in a short scar-covered brachyblast | mm,
buds 2 or 3, of different sizes, hairs greyish, 0.1 mm; bracts minute, caducous. Male
flowers: pedicel 4 by 0.3—0.4 mm, bracteole ovate-ellipsoid, | mm, persistent, buds
membranous, ovoid-oblong, 4—4.2 by 2—2.2 mm, apex + blunt, not angled, base broadly
rounded, cleft 1/4, lobes | mm long, 0.2 mm thick, at anthesis + curved outwards.
Androecium 4 mm; androphore 1.5 by 0.4 mm, glabrous but drying with an irregularly
warty surface; synandrium 2 by 0.5 mm, thecae 12-14, sterile apex + blunt, 0.2—-0.3 mm.
Female flowers and fruits not seen.

Field-notes — Tree, 20 m tall, diameter 15 cm, bark smooth, dark brown, 0.3 mm,
live bark light brown, with scanty red sap. Flowers pale green.

Distribution — Malesia: SE Papua Barat (border of Western Prov. in Papua New
Guinea: at Ingembit, road to Opka), known only from the type.

Habitat & Ecology — Lowland forest; fl. June.

Note — Noticeable for a number of unique features: small leaves with faint nerves,
blades glabrous below with neither dots nor papillae, inflorescences pedunculate, peri-
anths small, 4 mm. The leaves dry conspicuously dark, to a blackish brown colour.
Superficially M. atrescens resembles M. tristis, but that species has sessile inflorescences,
larger male perianths and a (partly) pubescent androphore. The androphore in M. atres-
cens, as seen in the boiled-up flowers, has a peculiar warty-bullate surface, which may
be an artefact.

7. Myristica atrocorticata W. J. de Wilde

Myristica atrocorticata W.J. de Wilde, Blumea 40 (1995) 261. — Type: NGF 33391, Papua New
Guinea, Western Prov.

Tree 25-30 m. Twigs 1.5—3 mm diameter, drying often + 2- or 3-angled, grey or
(blackish) brown, glabrescent, hairs grey-brown, 0.1 mm or less; older twigs blackish
brown, coarsely longitudinally cracking and flaking; lenticels few, conspicuous or incon-
spicuous, or absent. Leaves chartaceous (to thinly coriaceous), oblong-lanceolate, 6-15
by 1.5-3.5 cm, gradually narrowed to the acute apex, base broadly attenuate or rounded;
upper surface olivaceous or (dark) brown, dull, lower surface grey-brown or light brown,
dull, early or late glabrescent, hairs + dense, brownish, scale-like or stellate, 0.1 mm or
less (lens!); not distinctly papillose; dots absent; midrib slender, flattish above, lateral
nerves 16-21 per side, at 60—70° to the midrib, flat and very faint above, faint, not con-
trasting below, lines of interarching and venation indistinct; petiole proportionally long,
(13—)15—22 by 1.5—2 mm; leaf bud 10-15 by 1.5—2 mm, hairs grey(-brown), appressed,

De Wilde — Myristicaceae (Myristica) 41]

0.1 mm or less. Inflorescences between and below the leaves, sometimes on the older
wood, of the Knema-type, brachyblast sessile, short, simple, to 3 mm, densely rusty
pubescent, with 5—15 flowers in a loose apical cluster, buds + equal in size; flowers with
dense dark brown hairs 0.1 mm. Male flowers (immature): pedicel slender, 3-5 mm,
bracteole ovate or rounded, | mm, (sub)apical, persistent; buds ovoid to broadly ellip-
soid, 3 by 2—-2.2 mm, rounded at apex and base, cleft 1/3, lobes 1 mm long, 0.3 mm
thick. Androecium broad-cylindrical, 2 mm; androphore 0.3 by 0.6 mm (to | mm long at
anthesis), glabrous except for a few scattered pale hairs less than 0.1 mm; synandrium
1.7 by 0.7-0.8(—1) mm, thecae c. 15, sterile apex + blunt or truncate, 0.1—0.2 mm. Fe-
male flowers (NGF 33391): pedicel straight, 4-5 by | mm, bracteole ovate, nearly | mm,
subapical; buds ovoid, 4 by 3 mm, the apical part narrowed into a + blunt apex, base
broadly rounded, cleft 1/3, lobes 1.4 mm long; ovary long-ovoid, 2.2—2.5 by 1.8 mm,
hairs dense, golden-brown, appressed, 0.2—0.3 mm. Fruits not seen, fructescences prob-
ably with several fruits, judged from the many-flowered female inflorescences.

Field-notes — Occasional or locally common tall forest tree, 30 m or more tall. Trunk
(Brass 7079) raised on short flying or prop-roots, or without buttresses (BW 122951).
Bark blackish, scaly, hard, or longitudinally fissured or not; underbark olive or light
brown; inner bark red with red sap; wood white or pink. Leaves (NGF 31774) yellow
green above, rusty below. Flowers yellow.

Distribution — Malesia: Central New Guinea (Papua New Guinea: upper Fly River
area); West New Guinea (Bird’s Head: Warsamson River valley, E of Sorong).

Habitat & Ecology — Lowland forest, on low ridges near river, forest on well-drained
flat land, on podzol (with Myrtaceae, Vatica, Campnosperma), and in valley swamp
forest; 30-160 m altitude; fl. June, July.

Note — Myristica atrocorticata is known only from a female flowering specimen
and two male flowering specimens with immature flowers; fruits are not known. It re-
sembles the variable and widespread M. tristis but is readily distinguishable by the con-
spicuously cracking blackish bark of the older twigs, the comparatively long petioles,
the faint nervation, and the late glabrescent undersurface of the leaves with inconspicu-
ous minute scale-like hairs (but especially densely haired in Brass 7079). Superficially
the present species may be confused with M. firmipes. Myristica inundata is similar as
well; for differences see under that species.

8. Myristica basilanica W. J. de Wilde

Myristica basilanica W.J. de Wilde, Blumea 42 (1997) 148. — Type: FB 18928 (Miranda), Philip-
pines.

Tree. Twigs 2.5—3 mm diameter, yellowish brown; older twigs conspicuously crack-
ing and flaking; lenticels absent or not apparent. Leaves membranous, (elliptic-)oblong,
11-22 by 5-7.5 cm, base rounded or short-cuneate, apex acute-acuminate or + blunt;
upper surface olivaceous, lower surface pale greenish yellow, early glabrescent, hairs
scattered, pale, 0.1 mm or less, hair scars pale brown; papillose; dots absent; midrib flat
above, lateral nerves (10—)13—17 per side, at (45—)60—80° to the midrib, sunken above,
yellowish, much raised below, lines of interarching and coarsely reticulate venation yel-
lowish, distinct; petiole 15-25 by 2-3 mm, yellow-brown; leaf bud 10-15 by 3-3.5 mm,
hairs appressed, dull (grey-)brown, 0.2-0.3 mm. Inflorescences (female) (sub)sessile,

412 Flora Malesiana, Ser. I, Vol. 14 (2000)

Ww

K\

us 94

Fig. 66. Male flowers of Myristica species. — a. M. bialata Warb. var. bialata [NGF 29400 (Henty)|;
b. var. brevipila W.J. de Wilde [NGF 22472 (Gillison)].—c.M. ingrata W.J. de Wilde subsp. ingrata
[Weinland s.n.|. —d. M. subalulata Migq. var. subalulata [Hartley TGF 11419]; e. var. pedunculata
W.J. de Wilde [Sands c.s. 1481]. — Scale bar for all = 2 mm. — See also Fig. 75.

De Wilde — Myristicaceae (Myristica) 413

largely as in Knema: peduncle to | mm, sometimes with a central axis to 1 mm, hairs
dull, rusty, 0.2—0.3 mm, with 3-6 flowers in a cluster, buds of about the same size, hairs
0.1-0.2 mm. Male inflorescences or male flowers not seen. Female flowers: pedicel 1-2
by 1.5 mm, bracteole broadly ovate, rounded, 4 mm, late caducous; buds broadly ovoid,
5 by 4—4.5 mm, cleft 1/5, lobes 1 mm long, 0.2—0.3 mm thick; ovary broadly ovoid or
subglobose, 4 mm diameter, hairs pale brown, of mixed sizes, (0. 1-)0.5—0.7 mm. Fruits
solitary, subsessile, broadly ovoid, 6 by 4.5—5 cm, apex and base broadly rounded, early
glabrescent; pericarp 8-10 mm thick; seeds ellipsoid, 4 cm; fruiting pedicel not seen.

Distribution — Malesia: Philippines (Basilan); known only from the type.

Habitat & Ecology — Flowers and mature fruits Aug.—Sept.

Notes — 1. Obviously close to M. iners, which is widespread in W Malesia and in the
Philippines known only from Wenzel 3537, a + deviating collection from Mindanao
(Surigao), and quite different from M. basilanica. Myristica iners has pedunculate in-
florescences and leaves of which the lower surface is not papillose. Myristica cumingit
is similar and related, but has smaller fruits, 5(—5.5) cm, with a persistent indumentum.

2. The type specimens of Gymnacranthera urdanetensis, from Mindanao (Agusan
Prov.), are intermediate between M. basilanica and M. cumingii, but G. urdanetensis has
been sunk into M. cumingit.

9. Myristica beccarii Warb.

Myristica beccarii Warb., Mon. Myrist. (1897) 518, t. 14; Merr., Enum. Born. (1921) 269; J. Sinclair,
Gard. Bull. Sing. 23 (1968) 319, f. 43; W.J. de Wilde, Blumea 42 (1997) 149; Tree Fl. Sabah &
Sarawak 3 (2000) 453. — Type: Beccari 247, Sarawak.

Tree 10-24 m. Twigs 3—4(—5) mm diameter, late glabrescent, hairs rufous, 0.5(-1)
mm; older twigs blackish, longitudinally cracking and somewhat flaking; lenticels ab-
sent. Leaves coriaceous, elliptic-oblong to oblong-lanceolate, 9—20(—25) by 3.5-6.5(—9)
cm, more or less tapering to the + blunt or acute apex, base rounded, sometimes subcordate
or short-acute; upper surface olivaceous-brown; lower surface with dense scale-like grey-
brown or pale cinnamon hairs 0.1—0.3 mm, emergent hairs scattered, caducous, to 0.4
mm; not obviously papillose; dots absent; midrib above + flat, with some indumentum
towards the base, lateral nerves (1 1—)14—19 per side, at 60—80° to the midrib, sunken but
distinct above, lines of interarching and venation indistinct on both surfaces; petiole
short, 5-10(—15) by 3—4 mm; leaf bud 7-10 by 3 mm, hairs rusty, 0.5—1 mm. Jnflo-
rescences of the Knema-type; in male: sessile, simple or forked, woody, persistent,
brachyblasts to 12 mm, hairs dull, rusty, 0.5 mm, flowers few, buds of different sizes;
female inflorescences small, one- or few-flowered, bracts + triangular, 1 mm; flowers
with dense rusty woolly hairs 0.3—0.8 mm. Male flowers: pedicel 3-4 mm, bracteole
broadly ovate-circular, 2~4.5 mm, blunt or acute, + apical, late caducous; buds ovoid-
ellipsoid, 4—5.5 by 2.5—4.5 mm, apex faintly angled or not, cleft 1/2, lobes + coriaceous,
0.2(-0.8 at base) mm thick, at anthesis erect. Androecium 2.2—3 mm; androphore (0—)
0.2-0.4 by 0.5 mm, subglabrous but at base a dense collar of stoutish rusty hairs 0.5—
0.8(—1) mm; synandrium subcylindrical, 2-3 by 0.6—0.8 mm, thecae c. 15, sterile apex
irregularly shaped, + blunt, 0.2—0.3 mm, hairs dense or sparse, pale brown, 0.1 mm or
less. Female flowers (not fully known): buds ellipsoid, 5 by 3 mm, perianth lobes = re-
curved; ovary (immature) ovoid, 2-3 mm, with bright rufous-red hairs 0.5—1 mm. Fruits

414 Flora Malesiana, Ser. I, Vol. 14 (2000)

usually solitary, subsessile, ellipsoid(-obovoid), 2.5—3.5 by 1.5—2 cm, hairs dense, ru-
fous-red, fine-woolly, 0.5—1 mm, longest towards the base of the fruit; pericarp 1-2 mm
thick; seeds ellipsoid, 2(—2.5) cm; fruiting pedicel stoutish, 2-4 mm.

Field-notes — Medium-sized tree, with low buttresses or a few stilt-roots. Bark rough,
fissured with narrow strips 1-3 mm thick, or flaky, brown-black; inner bark (orange-)
brownish, with yellowish or red sap; wood white. Flowers yellow, fragrant. Fruits yel-
low. Leaves glossy dark green above, cinnamon below.

Distribution — Malesia: Sumatra (Riau) and Borneo (Sarawak, Sabah, NE, E, & SE
Kalimantan, incl. Nunukan).

Habitat & Ecology — Lowland dipterocarp forest on (leached) sandy clay soil; roll-
ing sandy area, ridges in kerangas; 0—600 m altitude; fl. & fr. throughout the year.

Notes — 1. Myristica beccarii resembles M. cinnamomea in foliage, but often the
base of the blade is + subcordate, and the petiole shorter; the fruits are smaller and the
pericarp much thinner. Myristica beccarii also closely resembles M. smythiesii, but that
species has more coriaceous leaves, more distinct nerves, and stouter twigs. Both M.
smythiesii and M. beccarii have Knema-type inflorescences. The indumentum of the
fruits is coarser than that of both M. cinnamomea or M. smythiesit.

2. Apparently M. beccarii is a rare species, in Borneo mainly collected in Sarawak
(1st & 2nd Div.) and eastern Kalimantan; only two collections are known from Riau,
Sumatra (pointing to Corner’s Riau-pocket distribution type).

10. Myristica bialata Warb.

Myristica bialata Warb., Bot. Jahrb. Syst. 13 (1891) 308; Mon. Myrist. (1897) 483; K. Schum.,
Notizbl. Bot. Gart. Berlin 2 (1898) 117; K.Schum. & Laut., Fl. Schutzgeb. Siidsee (1900) 326:
Markgr., Bot. Jahrb. Syst. 67 (1935) 159; Peekel, Fl. Bismarck Arch. (transl. Henty) (1984) 185,
f. 302: W.J. de Wilde, Blumea 40 (1995) 264, f. 1, 2a; 43 (1998) 171. — Type: Warburg 20706,
Bismarck Archipelago.

Tree 8-30 m. Twigs subterete or slightly flattened, with two raised lines or shallow
wings up to 2 mm high joining the petioles, 4-8(-15) mm diameter, smooth or finely
striate, early glabrescent, hairs 0.1 mm or less, stem hollow and in places somewhat
swollen and with a longitudinal opening (ant-swellings) or without distinct ant-swell-
ings; older twigs coarsely striate, sometimes transversely cracked; lenticels conspicuous
or inconspicuous. Leaves membranous or (thinly) chartaceous, (obovate-)oblong or
oblong-lanceolate, 18-52 by 6-22 cm, base rounded or shallowly cordate, apex acute-
acuminate; upper surface olivaceous or dull brown, lower surface grey, glabrescent,
indumentum minute; not conspicuously papillose; dots absent; midrib faintly raised above,
lateral nerves 20—30(—35) per side, at 60—80° to the midrib, + flat above, lines of inter-
arching not very prominent, venation often scalariform, distinct or indistinct; petiole
15-20 by 2.5—5 mm; leaf bud 10-20 by 3—4 mm, hairs appressed, 0.1—0.2 mm. Inflores-

Fig. 67. Myristica bialata Warb. var. bialata. a. Lateral twig with young male inflorescence bearing
one mature male flower: b. ditto, with old male inflorescence with one mature flower; c. part of
shoot with full-grown leaf, axillary to lateral twig and a flowerless inflorescence; d. part of older
twig with infructescence and mature fruit [a, b: NGF 29400 (Henty), c, d: LAE 65435 (Croft &
Lelean)|. — Scale bar for all = 2 cm.

De Wilde — Myristicaceae (Myristica) 415

uOs 94

416 Flora Malesiana, Ser. I, Vol. 14 (2000)

cences among the leaves or below, of the Knema-type, indumentum minute, glabrescent;
in male: brachyblast simple or forked, scar-covered, wart- or worm-like, to 1.5 cm, ses-
sile or with peduncle to 1 cm (var. brevipilis), with 1—5 flowers apically, buds of differ-
ent sizes; female inflorescences: subsessile, |—3(—5)-flowered; flowers minutely pubes-
cent, + glabrescent. Male flowers: pedicel 6-15 by 0.8—2 mm, bracteole ovate(-elliptic),
3-5.5 mm, persistent; buds ovoid-lanceolate, 10—13 by 3.5-6 mm, apex narrowed, blunt,
base + rounded, cleft c. 1/8, lobes 1.5 mm long, 0.2—0.3(—0.5) mm thick. Androecium
6.5—-9 mm; androphore 1.5—2 by 0.8-1 mm, subglabrous or minutely pubescent in the
lower 2/3-3/4; synandrium narrowed to the apex, 3.5—6 by 0.8-1.5 mm, thecae 12-14,
sterile apex subacute, narrow, 0.5—1 mm. Female flowers: pedicel 3-5 by 2—2.5 mm,
bracteole broadly rounded, 2 mm; buds + leathery, ovoid-oblong, + narrowed to the
apex, (5—)7-9 by 3—4.5 mm, lobes 1.5—2 mm; ovary ovoid-oblong, finely pubescent,
including stigma, 5—6 mm. Fruits 1—6 per infructescence, (sub)sessile, + fusiform or
ellipsoid, 5—7 by 3.5—4.5 cm, minutely mealy brown pubescent, glabrescent; pericarp
10(—15) mm thick, coarsely wrinkled; seeds ellipsoid, (2.5—)3—4 cm; fruiting pedicel
(4-)5-12 by 3-5 mm, with some lenticels. — Fig. 67.

Distribution — Malesia: Papua New Guinea (lowland areas of Manus I. and Bis-
marck Archipelago); two varieties.

Notes — 1. Myristica bialata was included in M. subalulata by Sinclair [Gard. Bull.
Sing. 23 (1968) 385] and Foreman {Handb. Fl. Papua New Guinea | (1978) 210], but
this inclusion is unwarranted. Both species have winged twigs that may contain ants, but
the flower of M. subalulata has a deciduous bracteole, and fruits 1.5—2(—2.5) cm. Myristica
bialata generally is a larger tree (to 30 m tall) as compared to M. subalulata. Myristica
hollrungii is a similar species, but with twigs not or less distinctly lined, smaller flow-
ers, and smaller fruits. The collection NGF 43453 (Bagabag I.) has large fruits to 5 cm
and approaches M. bialata, but is here included in M. hollrungii. Compare also M. verru-
culosa.

2. Fresh fruits are ovoid or nearly globose (from field-notes and the figure in Peekel,
l.c.); apparently the + juicy pericarp shrinks on drying.

3. Specimens from a small area in West New Britain differ from the type species in
the extremely short hairs on perianth, ovary, and (immature) fruits, and are recognized
here as a separate variety, var. brevipila.

KEY TO THE VARIETIES

la. Flowers (incl. bracteole and immature fruits) with hairs 0.1-0.3 mm; ovary with
hairs 0.2—0.4 mm; mature fruits glabrescent. — Manus I. and Bismarck Archipelago

Se “side Neha OLS ace eh Aas ee od Nims Seite eR nov ae a. var. bialata

b. Flowers and immature fruits with hairs less than 0.1 mm, partly glabrescent; ovary
densely covered with cinnamon scale-like hairs much less than 0.1 mm; mature fruits
not seen. —W New Britain, Kandrian subdist. ............... b. var. brevipila

a. var. bialata

Twigs 4-10 mm diameter. Leaves to 40 by 16 cm. Inflorescences without peduncle.
Female flowers: buds ovoid, 5—7 by 3.5—4.5 mm, with hairs 0.1—0.3 mm; ovary ovoid,

De Wilde — Myristicaceae (Myristica) 417

with hairs 0.3—0.4 mm. Fruits 1 or 2 per infructescence, immature fruits with bright
brown or yellow brown mealy hairs (0.1—)0.2—0.3 mm; mature fruits glabrescent. —
Fig. 66a, 67.

Field-notes — Slender tree, to 30 m, branches horizontal or + drooping, often more or
less whorled. Trunk spurred at base, bark brown, shallowly fissured; blaze red-brown;
inner bark red-brown; red sap; wood straw, pale brown, or cream, strongly laced with
red channels; wood soft. Leaves grey or glaucous below. Flowers cream, brown hairy.
Fruits greenish yellow with loose orange-brown or grey-brown farinose indumentum;
seeds deep chocolate-brown.

Distribution — As the species.

Habitat & Ecology — Lowland rain forest, coastal or riverine forest; clay soil; also
in forest over limestone; found in association with Pometia, Octomeles, Eucalyptus de-
glupta, Homalium foetidum, Syzygium, Dillenia, 0-350 m altitude; fl. & fr. throughout
the year, but mostly fr. in Mar.

Note — The twigs are usually hollowed, in places thickened and with a slit-like per-
foration, and are inhabited by ants.

b. var. brevipila W.J. de Wilde

Myristica bialata Warb. var. brevipila W.J. de Wilde, Blumea 40 (1995) 264, f. 2b; 43 (1998) 171.
— Type: NGF 27188 (Henty), West New Britain.

Twigs 5—12(-15) mm diameter. Leaves to 52 by 22 cm. Inflorescences sessile or (in
male) with smooth peduncle to 10 mm. Female flowers: perianth in bud + flask-shaped,
narrowed in the upper half, 9 by 4 mm, with hairs less than 0.1 mm; ovary (ovoid-)
oblong, 6 by 2.5 mm, with dense scale-like hairs less than 0.1 mm. Fruits 2-6 per in-
fructescence, when immature with dense hairs 0.1 mm or less; mature fruits not seen. —
Fig. 66b.

Field-notes — Stilt-roots (once recorded). Branches drooping. Bark brown, shallow-
ly fissured, or grey mottled, blaze pinkish, with or without red exudate; wood straw-
coloured or white. Leaves very pale, silvery-grey below. Flowers greenish white. Imma-
ture fruits rufous-brown.

Distribution — Malesia: Papua New Guinea (West New Britain: known only from
the Kandrian subdistrict).

Habitat & Ecology — Riverside forest; red soil over limestone; 0—200 m altitude;
fl. Mar., Apr.; fr. Oct.

11. Myristica bifurcata (J. Sinclair) W.J. de Wilde

Myristica bifurcata (J. Sinclair) W.J. de Wilde, Blumea 35 (1990) 239, f. 1: 2; 40 (1995) 266; 42
(1997) 153. — Myristica lancifolia Poir. var. bifurcata J. Sinclair, Gard. Bull. Sing. 23 (1968)
460 (key p. 81), f. 80. — Type: Kostermans 944, Morotai.

Tree 10-40 m. Twigs subterete or + angular, |—-2(—2.5) mm diameter, early glabrescent,
hairs 0.1—0.5(—1) mm; older twigs sometimes becoming longitudinally cracked or slightly
flaking; lenticels not conspicuous. Leaves chartaceous, ovate-oblong to oblong-lanceo-
late, 8-20 by 2.5-7 cm, base cuneate or + rounded, apex acute-acuminate (very tip often
+ blunt); upper surface olivaceous or brown, lower surface pale, glabrous; papillose;

418 Flora Malesiana, Ser. I, Vol. 14 (2000)

dots absent: midrib somewhat raised above, lateral nerves 8-14 per side, at 60—70° to
the midrib, flat or sunken above, not much raised and sometimes hardly visible below,
lines of interarching faint, venation little raised, faint or contrasting below; petiole 8-15
by 1-2 mm; leaf bud 8-12 by 2-3 mn, hairs 0.1-0.5 mm. Inflorescences sometimes
below the leaves, glabrescent; in male: either (sub)sessile (as in Knema) or peduncle
slender to 18 mm, ending in a single or forked scar-covered brachyblast, 1-5 mm, with
clusters of 5—10 flowers, buds subequal in size; female inflorescences: similar, sessile or
peduncle to 10 mm, 1—5-flowered; flowers with short or long hairs. Male flowers: pedi-
cel shorter than the perianth, 2-3 mm, bracteole (sub)persistent or caducous, buds cleft
1/3-1/2; androphore (sub)glabrous or thinly pubescent, shorter than the synandrium;
thecae 12—16: see further under the subspecies. Female flowers: pedicel shorter or longer
than the perianth, 1.5-4 mm; buds ovoid(-ellipsoid), 3-4.5 by 2-3 mm, apex narrowly
rounded, cleft 1/3—1/2, lobes 0.3 mm thick; ovary ovoid-oblong, 2—2.5 by 1(-1.5) mm,
pubescent, hairs 0.1—0.2 or 0.5—0.8 mm. Fruits solitary or paired, ovoid-ellipsoid, 3—4
cm long, hairs + long or short, brown; pericarp 3-5 mm thick; seeds not seen; fruiting
pedicel 2-4 mm.

Distribution — Malesia: northern Moluccas and Papua Barat (Bird’s Head); two sub-
species.

Notes — 1. The variable nature of the inflorescences is noteworthy on account of its
scar-covered short-shoot of the Knema-type, and usually distinctly pedunculate.

2. Specimens with (almost) sessile inflorescences may be confused with, e.g., M. im-
pressinervia and M. lancifolia. Myristica bifurcata is distinguishable by its mostly con-
spicuously pedunculate inflorescences, papillose lower leaf surface, flat nerves and
venation of the lower leaf surface, and fairly large fruits.

KEY TO THE SUBSPECIES

la. Leaf bud, flowers including ovary, and fruits with hairs 0.5—0.8(-1) mm. Male buds
4—5 mm; androphore sparsely or densely minutely pubescent . a. subsp. bifurcata
b. Leaf bud with long or short hairs, those of flowers including ovary and fruits short,
0.10.2 mm. Male buds 2.5 mm; androphore (sub)glabrous_.. . . b. subsp. sulaica

a. subsp. bifurcata

Tree to 20 m. Leaf bud, (immature) twig apex, inflorescences and flowers with hairs
0.5(—1) mm: sometimes similar hairs remaining for some time on the midrib. Peduncles
of male inflorescences up to 18 mm. Male flowers: pedicel 3 mm, bracteole ovate, sub-
acute, 22.5 mm, + persistent; buds 4-5 by 2.5 mm. Androecium 2.5—3.5 mm; androphore
0.5—1 by 0.4 mm, with a few scattered pale brown hairs 0. 1—0.2 mm; synandrium 2—2.5
by 0.5-0.8 mm, thecae 12-16, sterile apex + acute or blunt, 0.1-0.2 mm. Female flow-
ers: pedicel 2—4 mm, bracteole minute, 0.5—1 mm, + caducous; buds (narrowly) ovoid,
4.5 by 3 mm, ovary 2-2.5 by 1.5 mm, with dense hairs 0.5-0.8 mm. Fruits (ovoid-)
ellipsoid, 3—3.5 by 2-2.5 cm, with dark rusty hairs 0.5(—1) mm. — Fig. 64b.

Field-notes — Fairly common. Flowers and fruits rusty brown.

Distribution — Malesia: N Moluccas (Morotai); Papua Barat (Bird’s Head).

Habitat & Ecology — Locally common in forest on Morotai; forest along rivulet;
30—500 m altitude; fl. Apr—May; fr. June—Aug.

De Wilde — Myristicaceae (Myristica) 419

b. subsp. sulaica W.J. de Wilde .

Myristica bifurcata (J. Sinclair) W.J. de Wilde subsp. sulaica W.J. de Wilde, Blumea 35 (1990) 241.
— Type: bb 28880, Sula Is.

Tree 10-40 m. Leaf bud and twig apex with hairs short or long, 0.1—0.5(-1) mm,
those of inflorescences and flowers only 0.1—0.2 mm. Leaves early glabrescent. Pedun-
cle of male inflorescences up to 5 mm. Male flowers: pedicel 2—2.5 mm, bracteole ovate,
1 mm, persistent; buds (slightly immature) 2.5 by 1.5 mm. Androecium | .8—2 mm; andro-
phore short, 0.5 by 0.4 mm, glabrous or almost so; synandrium 1.5 by 0.7 mm, thecae
c. 14, sterile apex minute, 0.1 mm. Female flowers: pedicel 2—4 mm, bracteole minute,
+ caducous: buds ovoid-oblong, 3—3.5 by 2(—2.5) mm; ovary 2 by | mm, with hairs
0.1(—0.2) mm. Fruits subglobose or ovoid to ellipsoid(-oblong), 3-4 by 2.5 cm, with
scurfy hairs 0.2 mm.

Field-notes — Straight tree. Bark blackish or grey, strongly fissured, not peeling off.
Trunk with or without buttresses, core solid; outer bark 0.6—1 mm thick, inner bark 6—7
mm thick, reddish or orange, turning brown, granular; not much exudate; sapwood creamy
or reddish yellow, gradually passing into the slightly more reddish heartwood. Flowers
greenish. Fruits yellow or brown(-yellow); seeds dark brown.

Distribution — Malesia: N Moluccas (E Sula Is., Halmahera, Obi).

Habitat & Ecology — Well drained ridge forest and sloping forest; on stony, vol-
canic, clayey, or serpentine-derived soils; locally abundant, 0-600 m altitude; fl. & fr.
mainly Aug.—Dec.

Uses — Bark boiled in water and exudate drunk by young women with yellow faces
(2?anaemia) (de Vogel 4455). Wood used for house construction.

Note — Subsp. sulaica is exceptionally variable in the length of the hairs on the leaf
bud, 0.1—1 mm.

12. Myristica borneensis Warb.

Myristica borneensis Warb., Mon. Myrist. (1897) 401, t. 14 [non Myristica borneensis Gandoger
(1919) = Myristica villosa Warb.],; Merr., J. Straits Branch Roy. Asiat. Soc. (1921) 269;
W.]J. de Wilde, Blumea 42 (1997) 154; Tree Fl. Sabah & Sarawak 3 (2000) 454. — Lectotype:
Beccari 1574, Sarawak.

Tree 6-20 m. Twigs 2.5—5 mm diameter, sometimes yellowish, glabrescent; hairs pale,
0.1(—0.3) mm; older twigs brownish black, longitudinally cracking, frequently flaking;
lenticels not apparent. Leaves membranous, chartaceous, or thinly coriaceous, elliptic-
oblong or oblong-lanceolate, 11-30 by 3.5-10 cm, base rounded or short-acute, apex
acute or acute-acuminate; upper surface olivaceous (greenish), lower surface grey or
pale cinnamon, early glabrescent or with (sub)persistent dispersed greyish and pale brown
hairs less than 0.1(—0.3) mm; conspicuously papillose; dots absent; midrib + flat above,
lateral nerves 13—20 per side, at 80-60° to the midrib, sunken above, lines of interarching
and venation indistinct; petiole 12-28 by 1.5—3 mm; leaf bud 8-12 by 2.5-3 mm, hairs
greyish or pale brown, appressed, 0.1(—0.3) mm or less. Inflorescences among the leaves,
pedunculate, branched; in male: slender or broad, 2-6 by 2—4.5 cm, peduncle (5—)10—20
mm, lowest branches 10—20 mm, central axis short or long, with 1—3 (sub)sessile laterals,
with clusters of 5—10 flowers, buds of different sizes; bracts 3 mm, short-pubescent,

420 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Myristica) 421

caducous: female inflorescences: as in male but smaller, 1.5—2.5 cm, few-flowered, pe-
duncle 5-10 mm; flowers drying yellow-brown or orange, hairs sparse, grey-brown,
0.1(—0.3, see note 2) mm or less. Male flowers: pedicel 3-6 by 0.5 mm, bracteoles rounded
or irregularly 3-topped, 1.5—2(—3) mm; buds (broadly) ellipsoid, 3—4 by 2.5-3.5 mm,
apex broadly rounded and slightly 3-angled, cleft 1/2-2/3, lobes 0.4-0.5 mm thick, at
anthesis + curved outwards. Androecium ellipsoid-cylindrical, 1.5—-3 mm; androphore
0.2-1 mm long, about as broad as the androecium, hairs light brown, 0.1 mm or less;
synandrium 1.5—2 by 0.8-1.2 mm, thecae 14-20, sterile apex broad and flat or irregu-
larly truncate, 0.1—0.3 mm, or absent. Female flowers: pedicel 2—3 mm; buds 4(—4.5) by
3.5—4 mm, cleft 1/2; ovary ovoid, hairs dense, minute, brown. Fruits | or 2 per infruc-
tescence, + ellipsoid, (4.5—)5—7.5 by 3—4.5 cm, indumentum persistent or late glabrescent,
hairs dark cinnamon, mealy, 0.1(—0.3, see note 2) mm; pericarp 5(—8) mm thick; seeds
(3.5-)4-4.5 cm; fruiting pedicel smooth, + tapering, 5—9 mm, bracteole scar somewhat
below the fruit. — Fig. 68.

Field-notes — Buttresses small or none. Bark dark brown or blackish, smooth or
rough, shallowly flaking and minutely (and narrowly) fissured, or longitudinally striate,
not furrowed: outer bark thin, brittle; inner bark 2.5 cm thick, soft, red-brown, exudate
pale to dark red, copious or not, watery; sapwood also exuding red sap, soft. Leaves
glossy above, dull-glaucous below. Flowers pale cream-brown or (greenish) yellow.
Fruits (greenish) yellow, with short (orange-)brown indumentum.

Distribution — Malesia: Borneo (Sarawak; Brunei; Sabah, rare; W & C Kalimantan).

Habitat & Ecology — Primary lowland dipterocarp forest, ridge or hill forest, closed
heath forest in valleys; usually on sandy clay (on igneous rock) or yellow soil, sandy
slope near stream, also kerangas; 0—500(—1000) m altitude: fl. Apr.—June; fr. mainly Aug.—
Nov.

Notes — 1. Myristica borneensis may be confused with M. malaccensis which dif-
fers in its olivaceous-brown leaves on drying, subcircular bracteoles with ciliate margin,
(subglobose-)ovoid glabrescent male buds, cleft 1/4-1/3, with thin lobes, 0.3 mm thick,
androecium consisting almost entirely of the synandrium with a sterile apex lacking or
with the apex of the synandrium hollowed, glabrous androphore, and (almost) glabrous
fruits.

2. Two forms of indumentum can be distinguished, one with very short hairs, 0.1 mm,
on apical bud, inflorescences and flowers and with the lower leaf surface early glabrescent
(most of the specimens), and another form with the indumentum more conspicuous and
persistent, with dark brown hairs, 0.1—0.3 mm. In most collections the fruits have scurfy
hairs 0.1 mm long or less, but in very immature fruits the indumentum may be longer,
0.2—0.4 mm (longer hairs later on apparently shed). The infructescence is short, 1-2
(—3.5) cm long.

Fig. 68. Myristica borneensis Warb. a. Leafy twig with male inflorescences; b. male bud; note
persistent bracteole; c. ditto, opened, showing androecium; d. twig with female inflorescences;
e. female bud; f. ditto, opened, showing pistil; g. part of branch with infructescence, bearing a
single fruit; h. dry fruit, opened, showing arillate seed; i. detail of papillose lower leaf surface
[a: S 17939; b, c: S 36582: d—f: S 36089; g, i: S 37147, h: S 35852]. — Scale bar for a, g,d,h=
2 cm; for b, c, e, f = 1.65 mm; for i = 0.4 mm.

422 Flora Malesiana, Ser. I, Vol. 14 (2000)

3. The specimen BRUN 577 (Ashton) has a not or but indistinctly papillose lower leaf
surface, but there are many minute, scattered, dark brown dots, possibly cork-warts, ab-
sent in other specimens. SAN 36839 (Beaufort Dist., Sabah) has stout twigs and broad
fruits, 5 by 4 cm; the twigs dry blackish brown; the leaves are chartaceous, with acute-
acuminate base, the lower surface not very distinctly papillose, and small scattered scale-
like hairs virtually absent; this specimen may have to be excluded from M. borneensis.

13. Myristica brachypoda W.J. de Wilde

Myristica brachypoda W.J. de Wilde, Blumea 40 (1995) 266, f. 3a.— Type: NGF 8079 (Gray, Floyd
& Middleton), Papua New Guinea, Gulf Prov.

Tree 8-10 m. Twigs 1—1.5 mm diameter, brown-yellow, early glabrescent, hairs grey-
ish, less than 0.1 mm; older twigs brown blackish, striate and fissured, and thinly flak-
ing, with small inconspicuous lenticels. Leaves membranous or thinly chartaceous, el-
liptic-oblong, 10-13 by 3.5—4 cm, base cuneate, apex acute-acuminate; upper surface
(cinnamon-)olivaceous, lower surface pale greyish cinnamon; papillose; dots absent;
midrib raised above, lateral nerves 14—16 per side, at 70—80° to the midrib, flat or sunken
above, bright brown and contrasting in colour below, lines of interarching distinct, vena-
tion fine, faint; petiole 10-14 by I(—1.5) mm; leaf bud 6—7 by 1 mm, hairs inconspicu-
ous, brown-grey, appressed, less than 0.1 mm. Male inflorescences, male and female
flowers not seen. Female inflorescences (from the infructescences) axillary to a leaf scar
just below the leaves, of the Knema-type, a minute sessile knob, bearing | (or 2) flowers.
Fruits solitary, ovoid or slightly ellipsoid-fusiform, 3 by 1.7 cm, base and apex narrow-
ly rounded or + tapered, indumentum thin, felty, inconspicuous, hairs interwoven, scale-
like, 0.1(—0.2) mm, partly glabrescent; pericarp yellowish brown, 2(?) mm thick; seeds
not seen; fruiting pedicel 1-2 by 2 mm, glabrescent, cracked and flaking. — Fig. 63a.

Field-notes — Small tree, dbh c. 25 cm. Leaves dark green above, glaucous below.
Fruits axillary, ovoid, shortly pedicellate, with a fawn close indumentum.

Distribution — Malesia: Papua New Guinea (Kikori subprov., Gulf Prov., Seribi River
near Middletown); known only from the type.

Habitat & Ecology — Logged forest at sea level; fr. July.

Note — Myristica brachypoda readily keys out beside M. insipida, but is especially
distinguishable by the leaves and the subsessile, very short-haired fruits. The seemingly
glabrous (glabrescent) fruits are reminiscent of those of M. tubiflora, and the type speci-
men on which the present species, M. brachypoda, is based was named as such by Sinclair
[Gard. Bull. Sing. 23 (1968) 342]. The leaves of M. brachypoda are very like those of
M. pumila, but that species differs in its fruits.

14. Myristica brassii A.C. Sm.

Myristica brassi A.C. Sm., J. Arnold Arbor. 22 (1941) 72 (for male fl. specimen only, excl. Brass
& Versteegh 12547 = M. sphaerosperma); J. Sinclair, Gard. Bull. Sing. 23 (1968) 245, f. 22; W.J.
de Wilde, Blumea 40 (1995) 267. — Type: Brass 12254, Irian Jaya.

Tree 6 m. Twigs 3—4 mm diameter, early glabrescent, hairs 0.1 mm or less; older
twigs coarsely striate with a few inconspicuous lenticels. Leaves thinly coriaceous, ovate-
oblong or elliptic(-oblong), 19-25 by 9-12 cm, base (broadly) rounded, apex bluntly

De Wilde — Myristicaceae (Myristica) 423

acute or short acute-acuminate; the upper surface dark brown, lower surface somewhat
paler, (greyish) brown, glabrous (or possibly early glabrescent, hairs minute, scale-like);
+ papillose (but not conspicuously so); dark (red-)brown dots present, just visible to
the naked eye; midrib moderately raised above, lateral nerves 12—-14(-17) per side, at
45—60° to the midrib, slender, sunken above, nerves below bright brown and contrasting
in colour, lines of interarching indistinct, venation coarsely reticulate, thin, impressed
above (and rendering the blade faintly bullate), hardly visible below; petiole 20 by 4
mm; leaf bud 15 by 3 mm, hairs inconspicuous, appressed, dull brown, scale-like, 0.1
mm or less. Inflorescences among the leaves, sessile, of the Knema-type; in male:
brachyblast scar-covered, 3—4 mm, glabrescent, peduncle less than 1 mm; bracts broadly
ovate, 3 mm, acute, caducous; flowers (1—)2—5 in a cluster, buds almost equal in size,
with sparse grey-brown hairs 0.1 mm or less. Male flowers: pedicel (5—)9-10 by 1.5(—2)
mm, bracteole not seen, caducous, the scar subapical; buds (ellipsoid-)oblong, 10-12 by
5 mm, apex bluntish, cleft c. 1/5, lobes + coriacious, 2—2.5 mm long, 0.5 mm thick, at
anthesis + curved outwards. Androecium (broad-)cylindrical, 8 by 1.5(—2) mm; andro-
phore 3.5 by 1.5(—2) mm, dark brown appressed-pubescent in the lower half, hairs 0.3—
0.4 mm; synandrium 4 by 1.5(-2) mm, tapering to the apex into the 0.5 mm long acute
sterile apex, thecae c. 20. Female flowers and fruits not seen.

Field-notes — Leaves brownish below. Flowers cream-coloured, fragrant.

Distribution — Malesia: Papua Barat (Idenburg River, 15 km SW of Bernhard Camp);
known only from the type.

Habitat & Ecology — Rain forest or gully; 1700 m altitude; fl. Jan.

Note — Myristica brassii was treated under M. sphaerosperma by Foreman [Contr.
Herb. Austral. no. 9 (1974) 43: Handb. Fl. Papua New Guinea | (1978) 210]. The two
may be related, but M. brassii is quite distinguishable in details. A good differentiating
character is the absence of dots on the lower leaf surface in M. sphaerosperma, a charac-
ter which holds true on the species level for all Myristicas. Myristica brassii also seems
closely related to M. cucullata (see there).

15. Myristica brevistipes W.J. de Wilde

Myristica brevistipes W.J. de Wilde, Blumea 40 (1995) 267, f. 3b. — Type: Darbyshire 929, Papua
New Guinea, Central Prov.

Treelet c. 5 m. Twigs 1.5—2 mm diameter, early glabrescent, hairs appressed, 0.1—0.2
mm, brownish yellow; older twigs brown-black, coarsely striate or finely longitudinally
fissured: lenticels not apparent. Leaves membranous, (oblong-)lanceolate, 7-17 by 2-4
(—5) cm, base short-attenuate or narrowly rounded, apex (long-)acute-acuminate; upper
surface green-olivaceous, lower surface pale greenish brown, glabrous (indumentum of
immature leaves not seen); not papillose; dots absent; midrib raised above, lateral nerves
10-15 per side, at 70—80° to the midrib, slender, slightly sunken and inconspicuous
above, not much contrasting below, lines of interarching and venation (fine) indistinct;
petiole brown-yellow, 7-10 by 1.5 mm; leaf bud 5—7 by 2 mm, hairs + rough, 0.3—0.5
mm. Male inflorescences, male and female flowers not seen. Female inflorescences (from
infructescences): between the lower leaves, of the Knema-type, a (sub)sessile, scar-cov-
ered brachyblast, 3 mm diameter, few-flowered, hairs rough, rusty, 0.5 mm; bracts not
seen. Fruits 1 or 2 per infructescence, ellipsoid-oblong, + fusiform, 2.5—3 by jeS:tema,

424 Flora Malesiana, Ser. I, Vol. 14 (2000)

apex acute, base somewhat tapered, hairs thin, woolly, light brown or rusty, 0.3 mm,
towards the base stouter, 0.5 mm; pericarp 1-2 mm thick; immature seeds ellipsoid-
oblong, 1.5 cm; fruiting pedicel 2(—3) by 1.5—2 mm, hairs rough, 0.5 mm, bracteole scar
+ median. — Fig. 63b.

Field-notes — Tree c. 5 m tall, bole c. 2 m, dbh c. 5 cm. Leaves bright green above,
pale below. Fruits green with rusty-brown bloom.

Distribution — Malesia: Papua New Guinea (Central Prov, Kairuku subproy., about
1 mile NW of Maipa village); known only from the type.

Habitat & Ecology — On summit of small hill in tall foothill forest; c. 150 m alti-
tude; fr. Sept.

Note —The fruits of M. brevistipes are very similar to those of M. pygmaea which
differs in vegetative characters and (shortly) pedunculate inflorescences. Myristica brevi-
stipes appears particularly close to certain specimens of M. concinna, a species that
differs in smaller fruits and a more or less distinctly papillose lower leaf surface. Final-
ly also some specimens of the New Guinea form of M. insipida are very similar to
M. brevistipes and generally have fruits with more conspicuous, woolly, longer hairs,
and leaves which are papillose below. In New Guinea, however, specimens linking up
with M. insipida often have leaves with a less distinct papillation. This may suggest
that the demarcation between M. brevistipes and M. insipida is indistinct, but the type of
M. brevistipes is so different in general appearance from the majority of M. insipida
specimens that its distinction seems warranted.

16. Myristica buchneriana Warb.

Myristica buchneriana Warb., Bot. Jahrb. Syst. 13 (1891) 311; Mon. Myrist. (1897) 498, t. 19;
J. Sinclair, Gard. Bull. Sing. 23 (1968) 321, f. 44 (p.p., see the note); Foreman, Handb. Fl. Papua
New Guinea | (1978) 187 (p.p., see the note); W.J. de Wilde, Blumea 40 (1995) 268. — Type:
Warburg 20714, Papua New Guinea.

Tree (5—)10-35 m. Twigs angled or subterete, 1.5—3.5(—4) mm diameter, late glabres-
cent, hairs yellowish brown or rufous, 0.3(—0.5) mm; older twigs grey-brown or black-
ish, + transversely cracked, not flaking; lenticels small, indistinct. Leaves (membranous
or) thinly chartaceous, elliptic-oblong or oblong-lanceolate, 9—22(—28) by 3.5—7(-10.5)
cm, base short attenuate or rounded, apex acute(-acuminate); upper surface (blackish)
brown, lower surface grey(-brown), hairs usually persistent on or near the midrib, espe-
cially towards the base, just or hardly touching each other, not densely interwoven, scale-
like, grey or pale brown, 0.1(—0.2) mm or smaller; not or but faintly papillose; dots ab-
sent; midrib hardly raised above, lateral nerves 10—20 per side, at 50—80° to the midrib,
flat and faint above, lines of interarching and venation indistinct; petiole 8-13 by 1.5—
2.5(-3) mm, late glabrescent; leaf bud 10-15 by 2-3 mm, hairs 0.2—0.3(—0.5) mm. Inflo-
rescences among the leaves or below, of the Knema-type: peduncle to 3 mm, ending ina
simple or forked, scar-covered, flower-bearing brachyblast to 5 mm, late glabrescent;
in male: with a cluster of 5—15 flowers, buds of + variable sizes; female inflorescences
similar, 1—3(—5)-flowered; flowers with dense rufous hairs 0.2—0.3 mm. Male flowers:
pedicel 2—3 by | mm, bracteole small, caducous; submature buds obovoid-oblong, (4-)
5-6 by 2.5—3 mm, apex rounded, cleft c. 1/4, lobes 1.5 mm long, 0.3—0.5 mm thick.
Androecium short-cylindrical, 3—4 mm; androphore 0.5—0.6 by 0.5 mm, largely gla-

De Wilde — Myristicaceae (Myristica) 425

brous, towards the base a collar of sparse or dense erect reddish brown hairs 0.4—0.5
mm; synandrium (2—)3 by | mm, thecae 14—16, sterile apex blunt, 0.2—0.5 mm. Female
flowers: pedicel 4 by 1.5—2 mm; buds ovoid, 6(—7) by 4.5(—5) mm, cleft c. 1/4, lobes 1.5
mm long; ovary ovoid, 4 by 3 mm, brown pubescent. Fruits solitary, or 2 or 3 together,
ellipsoid-oblong, 3—4.5(—6) by 2.5(—3) cm, apex rounded, base rounded or somewhat
narrowed, hairs dense, bright (dark) rusty, 0.5—1 mm, rough especially towards the base
of the fruit; pericarp 2-3 mm thick; seeds ellipsoid, 2—2.5 cm; fruiting pedicel 4-8 by
(3-)4-5 mm, late glabrescent or with persistent indumentum, fissured.

Field-notes — No buttresses. Crown deep and narrow. Branches horizontal, or some-
what drooping. Bark rather smooth, longitudinally fissured or finely tessellated, dark
(grey-)brown; outer bark reddish brown; inner bark light brown or reddish; sap watery,
red: blaze red-brown; wood white, of medium density; sapwood (cream-)pink; heart-
wood (dark) salmon. Leaves grey or glaucous below. Flowers dark brown, inside creamy-
white. Fruits rusty hairy.

Distribution — Malesia: northern and eastern New Guinea (NE Papua Barat; Papua
New Guinea: Sepik, Morobe, Madang Provinces).

Habitat & Ecology — Primary and degraded forest; levee, hill, or ridge forest; for-
ests with or dominated by Lithocarpus, Pometia, Intsia, Pandanus, Celtis, or Anisoptera;
10-350 m altitude; fl. & fr. throughout the year. Locally common.

Note — Foreman (1978), following Sinclair (1968), described the male flowers with
perianths 8-10 by 4-5 mm, possibly based on Brass 28894, a collection here placed in
M. simulans. The perianth of this collection (in L) measures 8 by 4.5 mm; the androphore
is glabrous. Sinclair noted that Brass 28894 had a deviating indumentum. The caption
under Sinclair’s figure 44 seems to be either erroneous, or the specimen drawn does not
belong to M. buchneriana.

17. Myristica byssacea W. J. de Wilde

Myristica byssacea W.J. de Wilde, Blumea 40 (1995) 269. — Type Carr 15549, Papua New Guinea,
Northern Prov.

Myristica flosculosa auct. non J. Sinclair, p.p.: J. Sinclair, Gard. Bull. Sing. 23 (1968) 359, excl.
type, for female fl. and fr. specimens only, f. 59.

Tree 6-7 m. Twigs 1.5—2(—2.5) mm diameter, + angular and with two sharp ridges,
glabrescent, hairs pale brown, scale-like, less than 0.1 mm; older twigs neither cracking
nor flaking; lenticels not apparent. Leaves subchartaceous, elliptic-oblong, 10-16 by 3—
5.5 cm, base attenuate to nearly rounded, apex acute-acuminate with conspicuous acu-
men up to 2.5 cm; upper surface olivaceous, glabrescent with some minute scale-like
hairs remaining on the midrib towards the base, lower surface pale, grey-olivaceous or
yellowish, late glabrescent or with subpersistent dense pale (grey-)brown scale-like
hairs 0.1 mm or less; not papillose; dots absent; the midrib flat or somewhat raised
above, lateral nerves 10-15 per side, at 45—60° to the midrib, curving, sunken above,
below bright brown, contrasting in colour, lines of interarching very distinct; venation
+ scalariform; petiole 10-15 by 1.5—2.5 mm; leaf bud 10-15 by 1.5—2 mm, hairs appres-
sed, 0.1 mm. Inflorescences (known only from female inflorescences and infructescence):
sessile or peduncle to 1.5 mm, among the leaves, of the Knema-type, pubescent or late
glabrescent, brachyblast 1 mm, with 1-3 flowers; bracts caducous. Male flowers not

426 Flora Malesiana, Ser. I, Vol. 14 (2000)

seen. Female flowers: pubescent, hairs 0.1 mm or less, pedicel 6-9 by 1—-1.5 mm, bracteole
caducous, its scar 2-4 mm below the apex; buds ovoid-oblong, slightly narrowed in
the upper half, 6-7 by 3.5 mm, apex narrowly rounded or subacute, base rounded, cleft
1/4—1/6, lobes 1(—1.5) mm long, 0.3 mm thick; ovary ovoid-oblong, hairs golden brown,
appressed, 0.1—0.2 mm, including stigma 3—3.5 by 2 mm. Fruits single or in pairs, broadly
ovoid-ellipsoid, 2.5-3 by 22.3 cm, apex rounded with apiculum 1.5 mm, base broadly
rounded and contracted into 3 mm long pseudostalk, hairs dull brown, scurfy, 0.1 mm or
less; pericarp 2—2.5 mm thick; seeds (immature) ellipsoid, 1.5 by 0.8 cm; fruiting pedi-
cel + stout, straight, striate, 9-10 by 3.5—4 mm, bracteole scar faint, + median.

Field-note — Fruits yellow-brown.

Distribution — Malesia: Papua New Guinea (Northern Prov. at Isuarava); known
only from two collections.

Habitat & Ecology — Montane forest at 1600-1700 m altitude; fl. & fr. Feb.

Notes — 1. Female flowering and fruiting specimens are well depicted by Sinclair
(1.c.: f. 59) under M. flosculosa.

2. The epithet alludes to the cottony appearance of especially the lower leaf surface,
which is covered by a dull, yellowish brown, very thin indumentum. Other characteristic
features are the angular twigs, and the scalariform tertiary venation of the leaves.

18. Myristica cagayanensis Merr.

Myristica cagayanensis Merr., Philipp. J. Sci. 17, 3 (Sept. 1920’, Jan. 1921) 255; Enum. Philipp.
Flow. Pl. 2 (1923) 178; Kaneh., Formos. Trees, rev. edit. (1936) 193, f. 141; Hui-Lin Li &
H. Keng, Taiwania | (1950) 112; J. Sinclair, Gard. Bull. Sing. 18 (1961) 226; Hui-Lin Li, Woody
Fl. Taiwan (1963) 193, f. 72; Fl. Taiwan 2 (1976) 396, pl. 349; Ying & Li, Fl. Reip. Pop. Sinicae
30, 2 (1979) 190, f. 86; W.J. de Wilde, Blumea 42 (1997) 154. — Myristica ceylanica A. DC.
var. cagayanensis (Merr.) J. Sinclair, Gard. Bull. Sing. 23 (1968) 442, 444; Liao, Fl. Taiwan,
ed. 2, vol. 2 (1996) 420, pl. 195, photo 160. — Type: Bernardo 24277 (PNH, lost), Luzon, Prov.
Cagayan.

Myristica laurifolia auct. non Hook. f. & Thomson: Hayata, Mat. Fl. Form., repr. from J. Coll. Sc.
Imp. Univ. Tokyo 30, 1 (1911) 236.

Myristica heterophylla auct. non Fern.-Vill.: Hayata, Gen. Index FI. Form. (1917) 61.

Myristica glomerata auct. non Blanco: Kudo & Masam., Ann. Rep. Taihoku Bot. Gard. 2 (1932) 89.

Myristica philippensis auct. non Lam.: Kaneh. & Sasaki, J. Soc. Trop. Agr. 5 (1933) 307.

Tree 8-20 m. Twigs 3—4(—6) mm diameter, early glabrescent, hairs greyish, 0.1 mm or
less; older twigs coarsely striate, tending to crack, often with scattered conspicuous
lenticels. Leaves coriaceous, (obovate-)oblong, 9-20 by 3.5—7.5 cm, base (short-)cuneate,
apex acute or + blunt; upper surface olivaceous brown, lower surface glabrescent, hairs
interwoven, scale-like, 0.1 mm; possibly papillose, papillae irregularly shaped; dots ab-
sent; midrib above somewhat raised, lateral nerves 15-20 per side, at 45—60° to the
midrib, sunken above, lines of interarching and venation distinct; petiole comparatively
long, 15-30 by 2—3(—3.5) mm; leaf bud 8-15 by 3—4 mn, hairs grey-brown, 0.1 mm.

Fig. 69. Myristica cagayanensis Merr. a. Leafy twig with male inflorescences; b. male bud; c. ditto,
opened, showing androecium; d. twig with infructescence and one fruit [a—c: Liao FN 10579,
d: Huang c. s. FN 10526]. — Scale bar for a, d = 2 cm; for b, c = 1.5 mm.

De Wilde — Myristicaceae (Myristica) 427

3
a
3

Chi

428 Flora Malesiana, Ser. I, Vol. 14 (2000)

Inflorescences among the leaves or below, of the Knema-type, in male: simple, or 2- or
3-furcate scar-covered brachyblasts to 15 mm, peduncle absent or up to 5 mm, some-
times with a central axis to 4 mm, + glabrescent, hairs sparse, 0.1—0.2 mm, with subumbels
of 5—10(—20) flowers, buds of various sizes; female inflorescences: smaller, few-flower-
ed: flowers with rusty hairs 0.1(—0.3) mm. Male flowers: pedicel + stout, 2.5—4.5 mm,
bracteole ovate, 2—-2.5 mm, caducous; buds ovoid, 4—5(-6) by 3(—4) mm, cleft c. 1/3,
lobes + pergamentaceous, 0.2—0.3 mm thick, at anthesis suberect or + curved outwards.
Androecium 3—3.5 mm; androphore 1.5(—1.7) by 0.5 mm, hairs sparse, in the lower half
pale brown, 0.3—0.4 mm; synandrium 1|.5—1.7 by 0.6 mm, thecae 10-20, when old slightly
projecting at apex, sterile apex blunt or acute, 0.1—0.3 mm, glabrous. Female flowers
(seen only in one female flower amongst male flowers in Liao Taiwan FN 10579): buds
subsessile, 3.5 by 3 mm; ovary ovoid, 2 mm diameter, with dense golden brown hairs
0.2 mm. Fruits single or 2 per infructescence, broadly ellipsoid to nearly globose, 3-4
by 2.5—4 cm, hairs dense, rust-coloured, mealy, 0.1—0.2 mm; pericarp 4-10 mm thick;
seeds 2.5 cm; fruiting pedicel 2-5 mm. — Fig. 69.

Field-note — Trunk to | m diameter.

Distribution — Southern tip of Taiwan mainland, and Lanyu & Lutao Is.; Malesia:
Philippines (islands in the extreme north, and Luzon: Cagayan Prov. ).

Habitat & Ecology — Forest of mountain slopes in coastal areas; fl. & fr. throughout
the year.

Note — Sinclair (1.c.) included this species as a variety of his broad concept of M.
ceylanica, which is according to him a species distributed in Sri Lanka as well as in the
Philippines. Myristica cagayanensis resembles M. ceylanica s.s., but the latter differs in
the more slender twigs lacking lenticels, less coriaceous leaves, androphore glabrous at
base, and ellipsoid fruits with short-haired indumentum.

19. Myristica carrii J. Sinclair

Myristica carrii J. Sinclair, Gard. Bull. Sing. 23 (1968) 160, f. 8; W.J. de Wilde, Blumea 40 (1995)
270. — Type: Hoogland 3521, Papua New Guinea.

Tree 3-7 m. Twigs 3—4 mm diameter, early glabrescent, hairs 0.1 mm long or less;
older twigs ashy-brown or grey-black, coarsely striate, tending to crack and flake; lenticels
small, sporadic, not apparent. Leaves membranous, oblong(-lanceolate) or lower half
sometimes + tapering, 25—45 by 5-13 cm, base narrowly rounded or (long-)cuneate,
apex acute-acuminate; upper surface olivaceous or brown, lower surface dull (brown-
ish) grey, glabrescent, hairs widely scattered, appressed, pale brown, less than 0.1 mm;
not papillose; dots absent; midrib raised above, lateral nerves 16-20 per side, at 45—60°
to the midrib, flat above, bright brown, contrasting in colour below, lines of interarch-
ing and (coarse) venation distinct, + flat above, obscure or invisible below; petiole dark
brown, 12-25 by 2.5—4 mm, contrasting in colour with the grey twig; leaf bud 10-15 by
3 mm, with appressed hairs (0.3-)0.5 mm. Inflorescences in male: several crowded at
the end or along lateral short-shoots 1-3 cm long (always?), with caducous cataphylls,
and with apical leaf bud, below the leaves, peduncle 5—10 by 1-2 mm, + flattened, hairs
dense, pale brown, appressed, 0.1 mm or less; flowers 2 or 3 at the end of the peduncles;
bracts caducous: female inflorescences (from infructescences): similar. Male flowers:
pedicel slender, 8—10 mm, bracteole + small, ovate, acute, ( sub)apical; buds + ellipsoid,

De Wilde — Myristicaceae (Myristica) 429

8-10 by 4 mm, apex and base (sub)acute, glabrescent, hairs dense, pale brown, appressed,
0.1 mm or less, cleft c. 1/2, lobes 4—5 mm, thin; perianth inside finely verruculose. An-
droecium 4—5 mm; androphore slender, 0.5—1 mm, hairs short, pale brown, 0.2 mm at
very base only; synandrium subcylindrical, 4 by 1.5 mm, thecae 24-28, sterile apex
absent or slender, 0.5 mm. Female flowers not seen. Fruits | or 2 together, sometimes
crowded and then infructescence appearing more-fruited, (ellipsoid-)oblong, 4-5 by
1.5—2 cm, + fusiform, apex and base narrowed into 3 mm beak and pseudostalk, hairs
dense, rusty, (0.2—)0.5 mm; pericarp 2(-5) mm thick, seeds 3 cm by 8-10 mm; fruiting
pedicel terete, 6-8 by 1.5—2.5 mm, minutely pubescent, bracteole scar subapically.

Field-notes — Low tree, stem to 10 cm diameter. Bark dark brown, longitudinally fis-
sured, with colourless exudate turning red; wood creamy-straw. Leaves glossy green
above, glaucous below. Flowers cream. Fruits pale red-brown or orange; seeds black.

Distribution — Malesia: Papua New Guinea (Morobe & Northern Prov.); a local
endemic.

Habitat & Ecology — Well drained tall lowland or open forest, forest along riverbanks;
0-350 m altitude; fl. & fr. throughout the year.

Note — May be confused with M. hooglandii from the same area, but the latter dif-
fers in its slightly thicker leaves, weaker (obscure) lateral nerves and veins, broader and
larger male flowers, rounded at the apex (not acute), and the much larger glabrescent
yellow fruits.

20. Myristica chrysophylla J. Sinclair

Myristica chrysophylla J. Sinclair, Gard. Bull. Sing. 23 (1968) 254, f. 26; Foreman, Handb. FI.
Papua New Guinea | (1978) 187; W.J. de Wilde, Blumea 40 (1995) 270. — Type: Hoogland
3642, Papua New Guinea, Northern Prov.

Tree 7-25(-30) m. Twigs 2-6 mm diameter, (late) glabrescent, hairs dense, woolly,
golden or rust-coloured, 1-2 mm; older twigs grey-brown or blackish, finely longitudi-
nally cracking, often thinly flaking; lenticels not apparent. Leaves chartaceous or (thinly)
coriaceous, ovate-elliptic (rarely obovate-elliptic) to oblong-lanceolate, 10—32(—42) by
2.5-14(—18) cm, base shortly cuneate to narrowly or broadly rounded, or shallowly cor-
date, apex acute-acuminate; upper surface yellowish-olivaceous or brown, lower sur-
face grey-brown or glaucous, indumentum persistent, or late or early glabrescent (in
subsp. entrecasteauxensis), hairs dense, light brown, 0.2-0.3 mm, with a varying amount
of scattered dendroid or almost unbranched shaggy hairs up to 1.5 mm; not or indis-
tinctly papillose; dots absent; midrib raised above, glabrescent, lateral nerves 10—23 per
side, at (50—)60-80° to the midrib, flat or sunken above (leaves in subsp. chrysophylla
+ bullate), lines of interarching generally distinct, venation reticulate, + indistinct; peti-
ole short, 5-12 by 2-5 mm, late glabrescent as the twig apices; leaf bud 5—10(-—15) by
3-4 mm, acute or blunt, hairs (1-)1.5 mm. Inflorescences as in Knema: sessile, simple or
2-fid scar-covered wart-like or vermiform brachyblasts; in male: to 10 mm, late glabrescent
or hairs persistent, woolly, 1 mm; flowers in clusters of 3-15, buds almost similar in
size: bracts (late) caducous, 2 mm, pubescent; female inflorescences shorter, few-flow-
ered: flowers with dense woolly yellow-brown hairs 0.5—1.5 mm. Male flowers: pedicel
1-2 by 1-1.5 mm, bracteole 2-3 mm, persistent; buds ovoid-oblong or nearly tubiform,
5-8 by 2.5-3 mm, apex blunt, cleft 1/4—1/3, lobes 1.5—2 mm long, 0.2 mm thick.

430 Flora Malesiana, Ser. I, Vol. 14 (2000)

Androecium slender, 4—5 mm; androphore about as long as or slightly shorter than the
synandrium, 1—3(—4) by 0.4 mm, towards the base with dense yellow-brown straight
hairs 0.5-1 mm; synandrium 2-2.2 by 0.4-0.5 mm, thecae 8-12, sterile apex 0.2—0.3
mm. Female flowers: pedicel 0.5(-1) mm; buds + ovoid, 5—6 by 4.5—5 mm, cleft c. 1/3,
lobes 1.5—2 mm, at anthesis curved outwards; ovary broadly ovoid or subglobose, 2—2.5
mm diameter, hairs dense, woolly, pale brown, 1 mm, style | mm long, stigma | mm
long, 2- or 3-lobulate. Infructescences much reduced. Fruits subsessile, solitary or 2
(or 3) together, subglobose or ovoid, 2—2.5 cm diameter, or ovoid-oblong, up to 4.5 cm,
apex rounded or narrowed, hairs conspicuous, yellow-brown, dense, woolly, 2—4(-5)
mm; pericarp 2 mm thick; seeds ellipsoid, 1.5—2.5 cm; fruiting pedicel 1-3 mm.

Distribution — In Malesia a lowland species with two closely related subspecies in
Papua New Guinea.

Notes — 1. The inflorescences with (sub)sessile densely woolly flowers look like
irregularly knobbly woolly clusters, somewhat of the same colour and general appear-
ance as immature fruits. Possibly in connection with the dense and long hairiness of the
ovary, the style (with stigma lobes) is distinct; in most species of Myristica the stigma
lobes are (almost) sessile.

2. The two subspecies have adjoining geographical distributions, with a slight over-
lap in SE Morobe Province.

KEY TO THE SUBSPECIES

la. Lower leaf surface (at least partly) with persistent indumentum. Male buds 7—8 mm.
Fruits subglobose or broadly ovoid, 2-2.5(-3) cm diameter, hairs 3-5 mm. — Morobe
Prow., Northern Prove (Kokoda))::\2). 2320, Se 4. os cane tars 2 a. subsp. chrysophylla

b. Lower leaf surface glabrescent. Male buds 5-6 mm. Fruits (ovoid-)ellipsoid, 2.5—
4.5 cm, hairs 1.5—3 mm. —SE Morobe, Central, and Milne Bay Prov............
A dite dates tle old Se LR 2 Laka ine Meee eeNen oN aete B ott Se b. subsp. entrecasteauxensis

a. subsp. chrysophylla

Twigs 2-6 mm diameter. Leaves chartaceous, rarely coriaceous, small or large, to 14
(—18) cm wide, base short-attenuate, rounded, or shallowly cordate; upper surface gener-
ally faintly or distinctly bullate, lower surface grey-brown, with (at least partly) persist-
ent indumentum. Male buds oblong-tubiform, 7-9 mm, cleft (1/5—)1/4. Fruits sub-
globose or broadly ovoid, 2—2.5(—3) cm diameter, hairs woolly, 3-5 mm. — Fig. 61b.

Field-notes — Straight-boled, horizontal branching tree. Bark grey-brown or almost
black, usually shallowly fissured; inner bark (blaze) + pink; exudate red, watery, or clear
and rapidly turning orange-brown; middle bark (dark) red; inner bark red; wood reddish
straw or pinkish, turning orange-brown, wood hard or soft. Leaves with nerves deeply
impressed above. Flowers (golden-)brown tomentose, inside creamy; male flowers in
axillary clusters. Fruits rusty or golden brown tomentose.

Distribution — Malesia: Papua New Guinea (Morobe Prov.: Lae subprov.; Northern
Prov.: Kokoda).

Habitat & Ecology — Locally frequent in lowland and lower foothill forest of slopes,
ridges, or edge of grassland; forests with Anisoptera, Quercus, and Calophyllum on ridge-
crests; 15—500 m altitude; fl. Mar., Oct.; fr. throughout the year.

De Wilde — Myristicaceae (Myristica) 431

Note — The specimen LAE 52029 (Lae subprov.) approaches subsp. entrecasteaux-
ensis in its narrow, coriaceous leaves.

b. subsp. entrecasteauxensis (J. Sinclair) W.J. de Wilde

Myristica chrysophylla J. Sinclair subsp. entrecasteauxensis (J. Sinclair) W.J. de Wilde, Blumea 40
(1995) 270. — Myristica chrysophylla J. Sinclair var. entrecasteauxensis J. Sinclair, Gard. Bull.
Sing. 23 (1968) 257, f. 27, Foreman, Contr. Herb. Austral. no. 9 (1974) 35; Handb. Fl. Papua
New Guinea 1 (1978) 189. — Type: Brass 25893, Normanby Island.

Myristica fatua Houtt. var. morobensis J. Sinclair, Gard. Bull. Sing. 23 (1968) 289, f. 35. — Type:
NGF 3142 (Womersley), Papua New Guinea, Morobe Prov.

Twigs 2-4 mm diameter. Leaves chartaceous or coriaceous, small or large, + lanceo-
late or ovate-elliptic, or (ovate-)oblong, narrower than in subsp. chrysophylla, 2.5—9.5
cm wide, base short-attenuate or narrowly or broadly rounded, not cordate; upper sur-
face not or but faintly bullate, lower leaf surface glaucous (indumentum brown under-
neath), (early) glabrescent, the hairs dense to + sparse, brown. Male buds 5—6 mm, cleft
c. 1/3. Fruits (ovoid-)oblong, (2.5—)3—4.5 by 2 cm, with woolly hairs 2-3 mm.

Field-notes — Stem with or without buttresses or spurs. Crown open; branches hori-
zontal. Bark small-flaky, brown or grey-black; bark slash (inner bark) orange-straw,
with red-brown exudate, free flowing; wood white or straw, hard. Leaves dull green,
pale green, glaucous, or brownish below. Flowers yellow or brown hairy. Fruits brown
hairy, or furry-red brown; seeds black.

Distribution — Malesia: eastern Papua New Guinea (SE Morobe, Northern, Central
& Milne Bay Provinces; also Papuan Islands).

Habitat & Ecology — Forest of slopes and ridges; once recorded from forest over
limestone: also alluvial lowland forest; advanced regrowth forest; up to 430 m altitude;
fl. May, July; fr. throughout the year.

Notes — 1. Subsp. entrecasteauxensis is variable in general habit, and possibly should
be divided. Specimens with small, almost lanceolate leaves include NGF 38578 (Henty)
and NGF 3142 (Womersley) (the type of M. fatua var. morobensis). Comparatively large
and broad-leaved specimens include Brass 25986 and Saunders 57 (from Tufi subdis-
trict); this latter specimen was discussed by Sinclair (1958: 259) as being intermediate
to var. chrysophylla.

2. The dense long hairs of flowers and fruits are somewhat reminiscent of those of
M. inopinata.

21. Myristica cinnamomea King

Myristica cinnamomea King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 292, pl. 116; Warb., Mon. Myrist.
(1897) 445; J. Sinclair, Gard. Bull. Sing. 16 (1958) 358, f. 28, pl. VILA; 28 (1968) 210; W.J. de
Wilde, Blumea 42 (1997) 156; Tree Fl. Sabah & Sarawak 3 (2000) 456. — Lectotype: King’s
coll. 7474 (male fl.), Peninsular Malaysia.

Tree (7—)10—30 m. Twigs 1.5-2.5(—4) mm diameter, glabrescent, hairs 0.1 mm; older
twigs dark grey-brown to blackish, coarsely striate, not or but rarely flaking; lenticels
absent. Leaves chartaceous or thinly coriaceous, elliptic-oblong to oblong-lanceolate,
8-22 by 2-8 cm, base broadly acute or attenuate, apex acute(-acuminate); upper surface
olivaceous, + glossy, lower surface grey- or silvery-brown, sometimes late glabrescent,

432 Flora Malesiana, Ser. I, Vol. 14 (2000)

hairs dense, inconspicuous, 0.1 mm, with scattered brown hairs to 0.2 mm; not obvi-
ously papillose; dots absent; midrib above flat or somewhat raised, in a groove, lateral
nerves 10-17 per side, at 45—60° to the midrib, + sunken, indistinct above, lines of inter-
arching and venation indistinct; petiole 12—22 by 1.5—3 mm; leaf bud 10-15 by 2-3 mm,
hairs 0.1 mm. Inflorescences among the leaves, pedunculate, with mealy hairs, 0.1—0.2
mm; bracts ovate, 2-3 mm, caducous; in male: |—2.5 cm long, peduncle 5—15 mm; later-
al branches up to 5 mm, central axis to 10 mm or absent (or reduced to one flower), with
subumbels of 5—8(—10) flowers, buds of various sizes; female inflorescence: an up to
3 mm pedunculate wart (rarely + forked), with 2—4-flowered subumbel; flowers with
hairs 0.1(—0.3) mm. Male flowers: pedicel 3-6 by 0.5 mm, bracteole 1(—2) mm, (sub)-
apical, (late) caducous; buds ellipsoid-oblong, + cylindrical or + contracted in the mid-
dle, 5-7 by 2—3 mm, apex + angled, cleft 1/3—1/2, lobes 2-3 mm long, 0.5(—1) mm thick,
at anthesis + curved outwards. Androecium 3.5—4.5 mm; androphore 1.5—2(—2.5) by
(0.6—)0.8—1 mm, hairs pale, 0.1 mm or less; synandrium ellipsoid-oblong, 1.8—2 by 0.6—
0.8 mm, thecae 12-16, sterile apex blunt, 0.2—0.3 mm, minutely pubescent (Peninsular
Malaysia, Borneo) or + acute, (sub)glabrous (Sumatra). Female flowers: pedicel 2.5—4
mm; buds ovoid-ellipsoid, angled at apex, 5—6 by 2.5—3 mm, cleft 1/3—1/2; ovary ovoid-
conical, 2.5 by 1.5—2 mm, with dense hairs 0.1 mm or less. Infructescence 0.5—2 cm.
Fruits solitary or 2 (or 3), broadly ellipsoid or oblong, or + pear-shaped, 5—9 by 2.5—5
cm, base rounded or subattenuate, hairs dense, mealy, rusty, 0.1(—0.2) mm, pericarp
(5—-)8-15 mm thick; seeds ellipsoid, 3.5—5.5 cm; fruiting pedicel rather stout, 5-10 mm.

Field-notes — Crown dense. Trunk at base often + fluted or with low buttresses, or
with a few small stilt-roots (often recorded); bark blackish brown (chocolate, dark grey),
brittle, with fine close vertical fissures, sometimes also with fine horizontal cracks, some-
times thinly flaking with appressed flakes; inner bark 5 mm thick, brown to pinkish,
soft, + laminated; exuding red watery sap slow to appear; cambium reddish; sapwood
soft, pale yellowish to brownish, often reddish speckled. Leaves dark green, glossy above,
midrib pale, in a furrow; lower leaf surface with coppery, golden, or silvery brown in-
dumentum. Flowers greenish to yellow with yellow-brown indumentum; scent not pleas-
ant; tepals bright red inside. Fruits to 10 cm, yellow-brown to apricot, with apricot or
rusty mealy indumentum; aril compact, bright or dark red; seeds blackish.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo, rare in
the Philippines (Mindanao, no material seen).

Habitat & Ecology — Locally common or scattered in various types of primary for-
est: lowland and hill mixed dipterocarp forest, ridge forest, (hilly) kerangas, forest tran-
sitional to submontane forest; on a variety of soil types: sandy soils, sandy loam, red
soil, mor soil, brownish clay-loam soil, silty sand, stony granitic soil, igneous-derived
soil, wet ground, sandstone; on hillsides, crests, undulating land; also in peat swamp
forest; 0—1000(—1400) m altitude; fl. & fr. throughout the year, but fl. predominantly
Mar.—June in all areas.

Notes — 1. Myristica cinnamomea is characterized by the persistent inconspicuous
hairs on the lower leaf surface, the short hairs on the leaf bud and twig apices, the angu-
lar apex of flower buds, and the fruits with mealy indumentum. Myristica beccarii and
M. smythiesii are superficially similar species but they have different inflorescences.

2. The pubescent sterile apex of the androecium in the material from Peninsular
Malaysia and Borneo is noteworthy.

De Wilde — Myristicaceae (Myristica) 433

22. Myristica clemensii A.C. Sm.

Myristica clemensii A.C.Sm., J. Arnold Arbor. 22 (1941) 78; W.J. de Wilde, Blumea 40 (1995)
270. — Myristica lancifolia Poir. var. clemensii (A.C. Sm.) J. Sinclair, Gard. Bull. Sing. 23 (1968)
463, f. 81; Foreman, Handb. Fl. Papua New Guinea | (1978) 204. — Type: Clemens 1668, Papua
New Guinea.

Tree 8-25 m. Twigs (1—)1.5—2 mm diameter, sometimes faintly angular, early gla-
brescent, hairs 0.1—0.2 mm; older twigs (blackish) brown; lenticels few, sometimes dis-
tinct; axillary buds sometimes conspicuous. Leaves chartaceous, (ovate-)oblong-lanceo-
late, 8-16 by 1.5—4.2 cm, base shortly attenuate, apex acutish, sometimes + rounded, or
with blunt tip; upper surface olivaceous or dark brown, lower surface brown-grey, early
glabrescent, hairs sparse, whitish, less than 0.1 mm; papillose; dots absent; midrib flat or
sunken above, lateral nerves 15—20 per side and with additional smaller intersecondary
ones often present, at 45—60° to the midrib, sunken above, indistinct below, lines of
interarching and venation indistinct; petioles 10-20 by 1-2 mm; leaf bud 10-12 by 1.5—
2 mm, with golden hairs 0.3—0.5 mm. Inflorescences among the leaves or below, with
short peduncle, approaching the Knema-type, short-pubescent, bracts small, caducous;
in male: peduncle (1—)2—4 by 1-1.5 mm, brachyblast scar-covered, simple or forked,
thickish, to 3 mm, with fascicles of (1—)2—6 flowers, buds of various sizes; female inflo-
rescence similar, 1—4-flowered: flowers with hairs 0.1—0.2 mm. Male flowers: pedicel
3-5 by 0.4-0.6 mm, bracteole + ovate, 1.5—2 mm, subpersistent; buds + carnose, ellip-
soid(-oblong), 4—5 by 2-3 mm, rounded at both ends, apical part + angled, cleft c. 1/3,
lobes 1.5-2 mm long. Androecium 3—4 mm; androphore I—1.5 by 0.8—1 mm, the lower
half with dense appressed brown hairs less than 0.1 mm; synandrium ellipsoid-oblong,
2-2.2 by 1-1.2 mm, thecae c. 16, sterile apex blunt, 0.2—0.3 mm. Female flowers not
seen. Fruits solitary, ellipsoid, 4—5.5 by 2.5—4.5 cm, apex + rounded, faintly beaked,
base + attenuate, hairs dense 0.1 mm; pericarp 5—8 mm thick, woody, often ridged at one
side, somewhat warty or not; seeds ellipsoid, 2.5 cm; fruiting pedicel woody, straight or
curved, 5-10 by 2.5—5 mm, cracked, sometimes with some lenticels.

Field-notes — Without buttresses. Bark + smooth with only shallow longitudinal
fissures, brown or grey; inner bark straw(-red), reddish brown or with red watery sap;
wood pinkish or straw; heartwood dark. Leaves glaucous below. Flower buds with golden
bloom; perianth yellow with rusty indumentum. Fruits orange or brown, up to 7 cm,
pyriform with prominent ridge, rugose or bark-like.

Distribution — Malesia: Papua New Guinea (Morobe, Northern, S Highlands Prov.,
E & W New Britain).

Habitat & Ecology — Montane forest, Castanopsis-dominated forest, on ridges and
slopes; (20—)250—900 m altitude; fl. Jan._May; fr. throughout the year. NGF 31707
(Ridsdale) (somewhat deviating by smaller, but possibly immature fruit) is from micro-
phyllous vegetation at only 20 m altitude.

Notes — 1. Myristica clemensii has distinctive smallish narrow leaves with papillate
undersurface; it superficially resembles M. lancifolia, which has sessile inflorescences,
smaller fruits, broader leaves, etc., and it has the distinct papillate undersurface of the
leaves in common with M. clemensii.

2. Specimens from New Britain differ from those of Morobe Province in having larger
fruits, 5—5.5 cm, with thick, relatively short fruiting pedicel.

434 Flora Malesiana, Ser. I, Vol. 14 (2000)

23. Myristica coacta W.J. de Wilde

Myristica coacta W.J. de Wilde, Blumea 40 (1995) 270. — Type: NGF 41553 (Henty et al.), Papua
New Guinea, West Sepik Prov.

Tree c. 10 m. Twigs 2(—3) mm diameter, early glabrescent, hairs 0.1 mm; older twigs
somewhat longitudinally cracking (not flaking), with numerous small lenticels. Leaves
(thinly) coriaceous, elliptic-oblong, 14—16 by 5—7 cm, base cuneate, apex (long-)acute-
acuminate; upper surface olivaceous(-brown), lower surface bright yellowish brown or
cinnamon by persistent dense, felty, interwoven scale-like and flocculose hairs 0.2-0.3
mm; not papillose; dots absent; midrib slightly raised above, lateral nerves 12-14 per
side, at c. 60° to the midrib, slender and flat above, little raised and faint below, lines of
interarching and venation indistinct at both surfaces; petiole 15-20 by 2-2.5 mm; leaf
bud 10-15 by 2 mn, hairs 0.2-0.3 mm. Female inflorescences (from infructescences)
(sub)sessile, + below the leaves, of the Knema-type, apparently few-flowered. Male and
female flowers not seen. Fruits (slightly immature) single, ovoid-ellipsoid, 3.5—4 by 2
cm, apex subacute, base narrowed or narrowly rounded, hairs dense, dark rust-coloured,
scurfy, 0.1(—0.2) mm; pericarp 3-4 mm thick; seeds not seen; fruiting pedicel 8—10 by
3.5-4 mm, short-pubescent, densely and conspicuously lenticellate, bracteole scar faint,
1—3 mm below the apex.

Field-notes — Leaves dark green above, brown tomentose below.

Distribution — Malesia: Papua New Guinea (West Sepik Prov. at Oksapmin; possi-
bly Takeuchi 4136 also belongs here, from Morobe Prov., Kuper Range).

Habitat & Ecology — Degraded fagaceous forest; altitude c. 2000 m; fr. Oct.

Note — Myristica coacta is distinguishable by its leaves with felty indumentum and
flocculose hairs; it keys out beside M. inutilis subsp. papuana and M. tenuivenia, spe-
cies with (partly) small leaves with persistent dense, short indumentum below.

24. Myristica colinridsdalei W.J. de Wilde

Myristica colinridsdalei W.J. de Wilde, Blumea 42 (1997) 156. — Type: Ridsdale c.s. ISU 499, NE
Luzon.

Tree 18—40 m, dbh 40-50 cm. Twigs (when fertile several pseudowhorled from near
the apex of orthotropic main stem) 4—5(—6) mm diameter, blackish, early glabrescent,
hairs pale, less than 0.1 mm, smooth or striate, cataphylls (bud scales) leaving half-
circular scars; older twigs faintly cracking, with scattered coarse lenticels. Leaves coria-
ceous, + crowded; blades (elliptic-oblong or) oblong-lanceolate, 11-30 by 3-6 cm, base
cuneate or narrowly rounded, apex acute(-acuminate) or + blunt; upper surface olivaceous-
brown, lower surface grey-brown, with subpersistent grey-brown dense interwoven hairs
0.1 mm or less; midrib slightly raised above, lateral nerves 15-22 per side, at 45—60° to
the midrib, flat or sunken above, purplish brown, contrasting in colour below, lines of
interarching and venation indistinct; petiole 20-35 by 2.5—3.5(-5) mm; leaf bud 15-20
by 3—4 mm, hairs appressed, 0.1(—0.2) mm. Inflorescences + crowded, between or be-
low the leaves, peduncle absent as in Knema, brachyblast scar-covered, simple or usu-
ally forked, to 10 mm, short-pubescent, glabrescent; bracts small, caducous; in male
with 2-10 flowers, buds of various sizes; flowers with dense, dark rust-coloured hairs,
0.1— 0.3 mm. Male flowers: pedicel 1.5—3 by 1-1.5 mm, bracteole small, 2 mm, persist-

De Wilde — Myristicaceae (Myristica) 435

ent or caducous: buds firm but thin, ovoid-ellipsoid, 4-5 by 3—3.5 mm, apex rounded,
cleft c. 1/2, lobes 2 mm long, at anthesis suberect. Androecium 4 mm, androphore
+tapering, 1—1.5 by | mm, in the lower half or almost completely with dense pale brown
hairs 0.1 mm or less; synandrium 2-2.5 by 1-1.5 mm, thecae 20, sterile apex + blunt or
broad conical, 0.2—0.3 mm. Female inflorescences and flowers not seen. Infructescences
small. Fruits (measurements of material in spirit) single (or 2) per infructescence, sub-
globose or ovoid(-ellipsoid-)oblong, 6—7 by 4—5 cm, hairs dense, short, dark brown,
0.1—0.2 mm (hence ovary pubescent); pericarp 10-12 mm thick; seeds ellipsoid, 3.5 cm;
fruiting pedicel S—10 by 8 mm.

Field-notes —Buttresses to | m, or with flying buttresses. Bark blackish, flaky or
scaly; inner bark wine red with watery sap, or streaky red with red sap. Leaves clustered
at the end of a branch, glaucous below. Flowers small, brownish. Fruits brown.

Distribution — Malesia: Philippines (NE Luzon, Isabela Prov.; endemic to the Palanan
area).

Habitat & Ecology — On ultrabasic: low stature forest with many large-girth trees,
streamside forest, or high canopy forest on low coastal hills; locally common in flat,
riverine areas; c. 50 m altitude; fl. & fr. Apr.

Note — The almost whorled lateral branching, and leaves that are somewhat cluster-
ed near the apex of the orthotropic main shoot, are possibly typical for the species. All
collections were made in areas with ultrabasic bedrock.

25. Myristica concinna J. Sinclair

Myristica concinna J. Sinclair, Gard. Bull. Sing. 23 (1968) 375, f. 62; Foreman, Handb. Fl. Papua
New Guinea 1 (1978) 189; W.J. de Wilde, Blumea 40 (1995) 271. — Type: Saunders 28, Papua
New Guinea.

Tree 3-22 m. Twigs 1-1.5 mm diameter, subangular, yellowish or reddish brown,
early glabrescent, hairs 0.1 mm or less; older twigs neither cracking nor flaking: lenticels
absent or small and indistinct. Leaves membranous or thinly coriaceous, elliptic-oblong
to oblong-lanceolate, 5—16 by 1.2—4.2(—4.5) cm, base attenuate, apex acute-acuminate;
upper surface olivaceous or brown, lower surface pale grey-brown, late glabrescent,
hairs sparse, scattered, pale, less than 0.1 mm; + papillose, but this sometimes not very
distinct, or only in old leaves (lens!); dots absent; midrib raised above, lateral nerves
7-14 per side, at 60—80° to the midrib, flat above, (little) raised below, lines of interarch-
ing and venation indistinct; petiole 7-12 by 0.7—1.2 mm; leaf bud 6-8 by | mm, hairs
0.1 mm or less. Inflorescences between and below the leaves, peduncle absent; in male:
simple or 2-armed scar-covered brachyblasts to 5 mm, early glabrescent: bracts + round,
0.5 mm, hairs coarse, caducous; flowers 1-10 in a subumbel, buds of slightly variable
sizes; female inflorescences 1—3-flowered; flowers with minute (blackish) brown hairs
0.1(—0.2) mm. Male flowers: pedicel 1—-1.5 mm, bracteole 0.5—1 mm, persistent; buds
almost tubular, 3.5—5 by 1.5—2 mm, apex rounded or subacute, base (shortly) attenuate,
cleft 1/5—1/4, lobes 1 mm long, 0.1—0.2 mm thick, at anthesis suberect. Androecium
slender, 3.2—-5 mm, included or shortly exerted; androphore 1—2 by 0.4—0.5 mm, pubes-
cent for lower 1/2-2/3, hairs pale, 0.1—0.3 mm, longest towards the base; synandrium
1.7-3 by 0.5-0.6 mm, thecae 6-8, + spaced, with a minutely 3-lobed sterile apex 0. 1—-0.3
mm. Female flowers: 3- or 4-lobed, pedicel 1-2 mm, bracteole 0.5 mm, = apical: buds

436 Flora Malesiana, Ser. I, Vol. 14 (2000)

ovoid, 4 by 2.5 mm, cleft 1/4, lobes 1 mm long, at anthesis somewhat curved outwards;
ovary ovoid, 2.5 by 1.8 mm, hairs 0.2—0.3 mm. Fruits solitary or 2 or 3 together, ellip-
soid, 1.5—2.5 by 1.2—1.5 cm, apex rounded or shortly acute, at base + contracted into an
up to 4 mm long pseudostalk; hairs mealy, 0.1—0.3 mm, longest on the pseudostalk; peri-
carp 2 mm thick; seeds broadly ellipsoid, 1.2—1.7 cm; fruiting pedicel slender, (1—) 2-4
mm, bracteole scar slightly below the apex.

Field-notes — Branches horizontal spreading. Outer bark 7 mm thick, (blackish) brown,
slightly longitudinally or finely fissured, with patches of brown and grey; blaze straw to
pale brown; sap colourless turning pink or (red-)brown; wood hard, straw, exudate turn-
ing red. Leaves glossy dark green above, dull glaucous or dull grey-green below. Flow-
ers (creamy-)yellow, orange, or brown, with brown hairs. Fruits (olive-)brown with light
brown hairs.

Distribution — Malesia: Papua New Guinea (Morobe, Northern, Central, and Milne
Bay Provinces, incl. Goodenough I.).

Habitat & Ecology — In primary, regrowth, and degraded forest, coastal, swamp, and
dry-land forest on ridges and slopes; Octomeles-forest on alluvial deposits; forest domi-
nated by Lithocarpus, Anisoptera, and Hopea; locally common; 0-500 m altitude; fl.
May—Aug.; fr. May—Nov.

Notes — 1. Myristica concinna is recorded by Foreman from the Gulf Province, but
I did not see any material from that area.

2. Sinclair, l.c., considered M. globosa to be the closest relative of M. concinna. How-
ever, it is closely related to M. insipida of which it can be regarded as a more delicate
edition. The two share short-pedicelled flowers and ellipsoid fruits (which, however, are
smaller and with shorter indumentum in M. concinna); special distinguishing features of
M. concinna are the slender twigs, small and narrow leaves, small, narrow and short-
pedicelled male flowers, and small fruits. The synandrium is made up of only 3 (possi-
bly sometimes 4) anthers, hence with 6(—8) thecae, distinctly spaced before sporulation.
The short sterile apex of the synandrium is formed by the sterile apical parts of the an-
thers, not by the central column as in most species of Myristica. The small, slender flow-
ers are reminiscent of those of M. lepidota; M. concinna may also be confused with M.
lancifolia, mainly on account of the small fruits; however, the latter species has male
flowers usually with longer pedicels, more anthers in the synandrium, a glabrous andro-
phore, and more closely set lateral nerves, fainter on the lower leaf surface.

26. Myristica conspersa W.J. de Wilde

Myristica conspersa W.J. de Wilde, Blumea 40 (1995) 272. — Type: BW 7964 (Schram), Irian Jaya,
Kebar Valley, Bird’s Head.

Tree 20-35 m. Twigs 1-2 mm diameter, early glabrescent, hairs sparse, | mm or less;
older twigs coarsely striate or finely fissured, but not cracking nor flaking; without or
with small inconspicuous lenticels. Leaves (thinly) chartaceous, (elliptic-)oblong or ob-
long-lanceolate, 5—10 by 2-2.5(—3) cm, base attenuate or cuneate, apex subacute or blunt,
or narrowly rounded; upper surface olivaceous, lower surface grey-brown or pale brown,
indumentum (sub)persistent, or early or late glabrescent, hairs dense or remote, scale-
like, 0.1 mm or less; not or faintly papillose; dots and dashes present; midrib (flat or)
raised above, lateral nerves 15—22 per side, at (70—)80° to the midrib, flat and incon-

De Wilde — Myristicaceae (Myristica) 437

spicuous above, lines of interarching faint, venation faint or invisible; petiole 7-12 by
1(-1.5) mm, leaf bud 7-10 by |.5(—2) mm, hairs | mm or less. Inflorescences among the
leaves or below, of the Knema-type: sessile or subsessile, simple (or 2-fid), scar-cov-
ered, slender, wart- or worm-like brachyblasts to 5 by 1-1.5 mm, glabrescent, peduncle
to 2 mm: in male with a subumbel of (2—)4—6 flowers, buds of various sizes; female
inflorescences 2(—4)-flowered; flowers with minute brown hairs 0.1—0.2 mm. Male
flowers: pedicel slender, 3 by 0.5 mm, bracteole | mm, persistent or caducous; buds ob-
long(-tubiform), 3.5 by 1-1.5 mm, apex narrowly rounded, base rounded, cleft c. 1/33
lobes 1(—1.5) mm long, (0.1—)0.2 mm thick. Androecium slender, 2.5 mm; androphore
1 by 0.3 mm, glabrous; synandrium 1.5 by 0.3—0.4 mm, thecae 12-14, sterile apex blunt-
ish, 0.1—0.2 mm. Female flowers: pedicel 3 by 0.6 mm, bracteole caducous; buds (sub-
mature) 3 by 2 mm; ovary not seen. Fruits solitary, subsessile, ellipsoid(-fusiform),
(2-)2.5 by 1.4 mm, apex acute with minute pointed style-remnant, base contracted into
3 mm long pseudostalk, hairs scurfy, rust-brown, 0.1 mm or less; pericarp 2 mm thick;
seeds ellipsoid, 1.5 cm; fruiting pedicel slender, 5 by | mm, indumentum minute, glabres-
cent, bracteole scar nearly | mm below the fruit.

Field-notes — Bole not buttressed. Bark with shallow fissures or not fissured, not
peeling, or strongly flaking; inner bark with red or orange milky exudate; wood pink or
reddish brown, no heartwood. Flowers (yellow-)brown. Fruits light brown.

Distribution — Malesia: Papua Barat (Bird’s Head).

Habitat & Ecology — Rather common in primary forest on sandy clay; from low
altitudes up to c. 900 m; fl. May, Nov.; fr. May.

Notes — 1. The three known specimens of M. conspersa are heterogeneous, and pro-
visionally placed here, also for convenience’s sake. None of the three fit in any other
known species, but when more material becomes available they may be found to repre-
sent more than one taxon. Brief notes on the specimens are given by De Wilde (1.c.).

2. Myristica conspersa seems closely related to M. arfakensis, from the same area,
both having small male flowers with a similar androecium; the latter differs in its stalked
inflorescences, slightly larger fruit, and dotted (not dashed) lower leaf surface. Myristica
conspersa has the faint, closely set lateral nerves in common with M. lancifolia.

Myristica scripta, occurring in most of New Guinea (except Bird’s Head), is another
closely related species; it differs in its generally stouter habit, leaves with acute apex and
usually drying darker brown, generally larger flowers, and fruits lacking a pseudostalk;
its flowers (incl. the androecium), however, are very similar.

27. Myristica cornutiflora J. Sinclair

Myristica cornutiflora J. Sinclair, Gard. Bull. Sing. 23 (1968) 348, f. 53, 54; Foreman, Handb. FI.
Papua New Guinea 1| (1978) 190, f. 87 (excl. M. gracilipes), W.J. de Wilde, Blumea 40 (1995)
273. — Type: BW 6413 (Kalkman), Irian Jaya.

Shrub or small tree, 1-20 m. Twigs 1.5—4 mm diameter, yellowish brown or dark
brown, very early glabrescent, hairs 0.1 mm or less; older twigs tending to crack lon-
gitudinally; lenticels not conspicuous. Leaves membranous or subcoriaceous, elliptic-
oblong to oblong-lanceolate, 8-24 by 3.5—10 cm, base (short-)cuneate or rounded, apex
acute-acuminate, margin + revolute on drying; upper surface olivaceous, lower surface
grey- or pale brown, early glabrescent, or extremely small whitish stellate scale-like

438 Flora Malesiana, Ser. I, Vol. 14 (2000)

hairs persistent, but leaves appearing glabrous; not papillose; sparse or dense dots and
dashes present (lens!); midrib flat or slightly raised above, lateral nerves 10—16 per side,
at 60—80° to the midrib, lines of interarching distinct or not, sunken above, venation
forming a coarse network, faint at both surfaces; petiole 12-25 by 1.5—3 mm; leaf bud
10-15 by 1.5-3 mm, hairs 0.1 mm. Inflorescences among the leaves or rarely below,
pedunculate, indumentum very minute; in male: 1.5—4 cm, with flattened peduncle 5—30
(—35) mm, with 2 or 3 up to 10 mm long scar-covered brachyblasts, with or without a
short smooth basal part, each arm with a subumbel of 3-10 flowers, buds subequal in
size, the central axis sometimes reduced to (the scar of) one flower; female inflores-
cences much smaller, peduncle 5—10(—20) mm, with (1 or) 2-4 flowers, but sometimes
along short leafless shoots suggesting larger inflorescences; bracts minute, caducous;
flowers largely glabrescent, hairs appressed, 0.1 mm or less. Male flowers: pedicel 5-15
by 0.5-1 mm, bracteole 1-1.5 mm, caducous, apical or up to 2 mm below; buds mem-
branous or horny, (ovoid-)oblong, sometimes + tubiform, much narrowed towards the
apex, 10-20 by 2-3.5 mm, the apical part + triangular in cross section, base narrowly
rounded, cleft at the apex only, lobes + thin, 0.2—-0.5 mm thick, (I—)1.5 mm long, at
anthesis not recurved. Androecium slender, 6-9 mm; androphore slender, 2.5—4.5 by
0.40.6 mm, minutely appressed-pubescent in the lower 3/4; synandrium 3.5—4.5 by
(0.6—1 mm, thecae 10-12, gradually passing into the androphore, sterile apex 0.2—0.4
mm, blunt (sometimes irregular). Female flowers: pedicel slender, 6-12 mm, bracteole
caducous, close to the apex; buds + flask-shaped, narrowed towards the apex, 10-12 by
3—4 mm, lobes 1-1.5 mm long; ovary ovoid-oblong, 4—5 by 2.5—3 mm, hairs dense, ap-
pressed, less than 0.1 mm. Fruits usually solitary, + ellipsoid(-oblong), 4.5-6.5 cm,
pubescent or glabrescent, style-remnant persistent; seeds 3 cm; fruiting pedicel straight,
slender, 1—2 cm.

Distribution — Two subspecies, one mainly in the southern, the other in the northern
part of New Guinea.

KEY TO THE SUBSPECIES

la. Male inflorescences rather stout, peduncle broadly flattened, 1.5-2 mm wide. Fruits
+ ellipsoid(-oblong), 4.5—6.5 by 2.5—3.5 cm, apex blunt, pseudostalk absent .....
SE REIN IIRL ABA LIE Ae REM: IORI UEN i SEE RUN RARE a. subsp. cornutiflora

b. Male inflorescences elegant, peduncle more slender, 1-1.5 mm wide. Fruits + fusi-
form, 4.5-5.5 by 2(—2.5) cm, apex acute, pseudostalk present .. b. subsp. elegans

a. subsp. cornutiflora

Small tree. Male inflorescences rather stout, peduncle broadly flattened, 1.5—2 mm
wide. Male flowers: pedicel 5—6 mm, bracteole scar apical; buds 10-13 by 3-3.5 mm, at
apex sharply angled, lobes horny, 0.5 mm thick; androphore 2.5—3.5 mm, as long as or
shorter than the synandrium. Fruits + ellipsoid(-oblong), 4.5—6.5 by 2.5—3.5 cm, apex
blunt, pseudostalk absent, suture deeply grooved, glabrescent, hairs minute, greyish;
pericarp 10 mm thick, woody. — Fig. 59c.

Field-notes — Buttresses up to 0.5 m, up to 3/4 m out. Outer bark shallowly fissured,
rather strongly peeling with small scales; inner bark with red opaque exudate. Flowers
greenish yellow. Fruits green-yellow or sordid yellow.

De Wilde — Myristicaceae (Myristica) 439

Distribution — Malesia: southern Central New Guinea (Papua Barat: Southern Divi-
sion); Papua New Guinea (Western & Southern Highlands Prov. ).

Habitat & Ecology — Dryland forest and marshy forest edges, also degraded forest;
limestone country; occasionally flooded; 25-800 m altitude; fl. & fr. throughout the
year.

b. subsp. elegans W.J. de Wilde

Myristica cornutiflora J. Sinclair subsp. elegans W.J. de Wilde, Blumea 40 (1995) 273. — Type:
NGF 48167 (Foreman & Kumul), Papua New Guinea.

Shrub 1—2 m, or low tree to 10 m. Male inflorescences elegant, peduncle slender, 1—
1.5 mm wide. Male flowers: pedicel slender, 8-15 mm, bracteole scar up to 2 mm below
the apex; buds elongate, 12-20 by 2-3 mm, at apex + triangular in section, but not with
sharp edges, lobes membranous, 0.2—0.3 mm thick; androphore 3—4.5 mm, as long as
synandrium. Fruits (somewhat immature) + fusiform, 4.5—5.5 by 2(—2.5) cm, apex acute,
at base with tapering pseudostalk, hairs dense, very minute, brown, scurfy, less than 0.1
mm, pericarp less than 10 mm thick.

Field-notes — Outer bark smooth, grey or dark brown; inner bark cream; sap from
bark watery, red; wood soft, straw or light brown. Leaves glossy. Flowers creamy with
waxy appearance, to 25 mm long. Fruits green- or orange-yellow, or orange-brown.

Distribution — Malesia: northern Papua New Guinea (West & East Sepik Proy.).

Habitat & Ecology — Hopea-Anisoptera forest, lowland rain forest beside rivers,
flood plains, mid-montane forest, or forest on low ridges; 200-1050 m altitude; fl. & fr.
Mar., June, Sept.

28. Myristica corticata W. J. de Wilde

Myristica corticata W.J. de Wilde, Blumea 42 (1997) 157; Tree Fl. Sabah & Sarawak 3 (2000) 457.
— Type: SAN 87293 (Krispinus), Sabah.

Tree 10-25 m. Twigs stout, angular, (4—-)5—6 mm diameter, indumentum minute, ear-
ly glabrescent, hairs | mm; older twigs pale, later on blackish, usually coarsely irregu-
larly longitudinally cracking and flaking; lenticels absent. Leaves chartaceous or thinly
coriaceous, elliptic-oblong or oblong(-lanceolate), 15—35 by 6—-11.5 cm, base broadly
rounded or subcordate, apex acute-acuminate; upper surface olivaceous brown, lower
surface dull, concolorous, glabrous; not obviously papillose; dots absent; midrib flat or
slightly raised above, lateral nerves 17—24 per side, at 60—80° to the midrib, above (flat
or) sunken, lines of interarching usually distinct, venation + indistinct on both surfaces;
petiole 15-25 by 2.5—4 mm; leaf bud 10-15 by (4—)5—6 mn, hairs rough, | mm. Jnflo-
rescences among the leaves and below, pedunculate, branched, hairs dense, woolly, rust-
coloured, 1(—1.5) mm; in male: 2—4.5 by 2-3 cm, peduncle flattened, 4—6 mm, lower
branches 5—7 mm, ending in a flower cluster, central axis 20-40 mm, with |—3 clusters,
each with 5—10 flowers, buds of various sizes; bracts caducous, pubescent; flowers with
woolly rust-coloured hairs 0.3(—0.5) mm. Male flowers: pedicel slender, 4-6 mm, brac-
teole 3.5 mm, persistent; buds ovoid, 5-5.5 by 4—4.5 mm, apex (narrowly) rounded,
base rounded, cleft c. 1/2, lobes 0.2 mm thick, at anthesis suberect. Androecium slender,
4.5 mm; androphore 1.5 by 0.7 mm, hairs conspicuous, brown, 0.2—0.3 mm; synandrium

440 Flora Malesiana, Ser. I, Vol. 14 (2000)

3 by 0.8 mm, thecae 14-16, sterile apex + blunt, 0.2 mm, glabrous. Female flowers not
seen. Infructescences little-branched, 1.5—2 cm, glabrescent. Fruits | (or 2) per infruc-
tescence, ellipsoid-oblong, 7.5-9.5 by 4-5 cm, early glabrescent, hairs dense, scurfy,
less than 0.1 mm long, glossy dark brown; pericarp 15(—20) mm thick; seeds ellipsoid-
oblong, 4.5—5.5 cm; fruiting pedicel (2—)5 mm.

Field-notes — Buttresses to 1 m or with flying buttresses, | by | m. Bole straight;
bark brown or blackish, cracked, coming off in small longitudinal thin flakes; outer bark
hard and brittle, 2.5 mm; inner bark 6 mm, hard, orange-brown, reddish, or yellowish;
sapwood pink, red, or light brown; cambium reddish. Flowers yellowish. Fruits ochre;
seed-coat black.

Distribution — Malesia: Borneo (Brunei, Sabah).

Habitat & Ecology — Primary and logged-over lowland forest, low undulating coun-
try, hillsides; sandstone ridges with sandy clay soil; 0-500 m altitude; fl. July; fr. May,
Aug., Nov.

Notes — 1. Myristica corticata is closely allied to the variable M. iners; both have the
same smooth, non-papillose structure of the lower leaf surface (lens!), which 1s of al-
most the same colour as the upper surface. Myristica corticata may be confused with
stout specimens of M. iners from northeastern Borneo. The differences between these
species have been discussed in detail by De Wilde (1.c.: 158).

2. In M. corticata the distinct marginal arches of the leaf venation are like those of
certain forms of M. extensa and M. malaccensis, and the stout twigs with conspicuously
cracking bark are like those of M. papyracea (but here the lower leaf surface is strongly
papillose).

29. Myristica crassa King

Myristica crassa King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 293, pl. 117; Warb., Mon. Myrist.
(1897) 495; J. Sinclair, Gard. Bull. Sing. 16 (1958) 366, f. 31, pl. VIIA; 23 (1968) 435; W.J. de
Wilde. Blumea 42 (1997) 159: Tree Fl. Sabah & Sarawak 3 (2000) 458. — Lectotype: King
5065, Peninsular Malaysia, Perak.

Myristica suavis King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 295, pl. 121; Warb., Mon. Myrist. (1897)
441. — Type: Cantley s.n., Peninsular Malaysia.

Myristica lowiana auct. non King, p.p.: King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 293; plnt20: 541,
5, 6, 7, flowering material (King 5537, 7258).

Tree 10-25 m. Twigs 3-7 mm diameter, dark brown or blackish, very early glabrescent,
indumentum minute; older twigs not flaking; lenticels sometimes present, indistinct.
Leaves chartaceous or thinly coriaceous, elliptic-oblong to oblong-lanceolate, (13—)17—
40 by 4.5-13 cm, base rounded or cuneate, apex acute-acuminate, margin often some-
what revolute on drying; upper surface dark olivaceous, lower surface greyish (purplish)

Fig. 70. Myristica crassa King. a. Leafy twig with male inflorescences; inset: papillate lower leaf
surface; b. male bud with persistent bracteole; c. ditto, opened, showing androecium; d. twig with
female inflorescence; e. female bud; f. ditto, opened, showing pistil; g. leafy twig with inflores-
cence bearing a single fruit; h. opened fruit showing thick pericarp and arillate seed [a: S 28098;
b, c: FRI 13537; d—f: SF 39490; g, h: FRI 8981]. — Scale bar for a, d, g, h = 2 cm; for b, c, e, f=
2 mm; for inset a = 0.4 mm.

441

De Wilde — Myristicaceae (Myristica)

442 Flora Malesiana, Ser. I, Vol. 14 (2000)

brown, glabrous or hairs scattered, scale-like, 0.1 mm or less; papillose; dots absent;
midrib flat or slightly raised above, lateral nerves 13—18(—22) per side, at 45—80° to the
midrib, sunken above, lines of interarching and venation indistinct; petiole 20-30 by
2-4 mm: leaf bud 8-14 by 3—4 mm, hairs appressed, 0.1 mm or less. Inflorescences 1-2
cm, among the leaves or below, hairs 0.1 mm or less, + of the Knema-type: sessile or
pedunculate to 6 mm, stout, with 1-3 woody brachyblasts, rarely with a lateral branch to
5 mm: in male: with 10-30 flowers, buds of various sizes; bracts |—-1.5 mm, finely pu-
bescent, late caducous; female inflorescences short, 2—5-flowered; flowers + coriaceous,
hairs less than 0.1 mm, sometimes late glabrescent. Male flowers: pedicel 5—12(-15)
mm, bracteole (sub)persistent, sometimes + lobed at apex, (2—)3—4 mm; buds ovoid or
urceolate-oblong, 4.5—6 by 2.5—5 mm, apex narrowly rounded, base broadly rounded to
subtruncate, not or but slightly angled at apex, cleft c. 1/3, lobes 0.3 mm thick. Androe-
cium slender, 3—3.5 mm; androphore 0.7—2 by 0.3—0.5 mm, minutely pubescent in the
lower 2/3 or towards the base only; synandrium ellipsoid-oblong 2 by 0.5 mm, thecae
6-10, sterile apex narrow, rather blunt or very faintly lobed, 0.3 mm. Female flowers:
pedicel + stout, 2 mm, bracteole caducous; buds + ovoid, 4—5 by 3-4 mm, cleft c. 1/3, at
anthesis curved outwards; ovary 3(—4) by 2-2.5 mm, minutely pubescent. Infructescences
short. Fruits 1 or 2 per infructescence, broadly ellipsoid, 3.5—5.5(—6) by 3-4 cm, hairs
rust-coloured, powdery (easily rubbed off), 0.1—0.3 mm; pericarp (7-)10 mm thick; seeds
ellipsoid, 3—3.5 cm; fruiting pedicel stout, 5(-10) mm. — Fig. 70.

Field-notes — Tall tree, branches spreading, a few weak stilt-roots present. Bark brown,
slightly and shallowly fissured longitudinally, or smooth, with horizontal fissures and
vertical cracks; inner bark red; wood white. Leaves dull dark green above, glaucous be-
low. Flowers pale green to yellow, with faint sweet odour. Fruits pale yellow or orange,
with orange-brown scurf; aril (light) yellow, fragrant, edible.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo. Not in
Thailand (the record by Sinclair, l.c., applies to M. maingay!).

Habitat & Ecology — Primary forest on hillsides, along rivers, (preference for) swamp
forest, rocky granite river valley; in Borneo in peat swamp and kerangas; 0-600 m alti-
tude; fl. & fr. throughout the year. Apparently not a common species.

Notes — 1. According to Sinclair (1.c.) M. crassa is sometimes monoecious, with
male and female flowers in the same inflorescence. In synonymous M. suavis, King
recorded rudimentary anthers present in female flowers.

2. Myristica crassa is related to M. teijsmannii (Java) and M. andamanica (Andaman
Is.). Sinclair (1968: 434) tabulated a comparison between these species.

30. Myristica crassipes Warb.

Myristica crassipes Warb. in K. Schum. & Laut., Fl. Schutzgeb. Siidsee (1901) 326. — Type: Bamler
50, Papua New Guinea.

Tree 4-30 m. Twigs bluntly angled or subterete, 1.5—3 mm diameter, early glabrescent,
hairs 0.1 mm or less; older twigs sometimes tending to crack longitudinally, not flaking;
lenticels often numerous, rather small but distinct. Leaves membranous, chartaceous, or
coriaceous, elliptic-oblong to (oblong-)lanceolate, 7-20 by 2.5—6.5 cm, base attenuate
or rounded, apex acute-acuminate, sometimes with blunt tip; upper surface olivaceous-

De Wilde — Myristicaceae (Myristica) 443

brown, lower surface grey-brown, with sparse or dense subpersistent pale brown scale-
like hairs 0.1 mm or less, sometimes (late) glabrescent; not papillose; dots absent; midrib
flat or raised above, lateral nerves 15—20(—24) per side, at (S0—)70—85° to the midrib,
flat or sunken above, little raised, purplish and contrasting below, lines of interarching
indistinct, venation usually + contrasting in colour below; petiole 8-16 by 1.5—3 mm;
leaf bud straight or curved, 7-16 by 1.5—2 mn, hairs 0.1 mm or less. Inflorescences
between the older leaves, essentially of the Knema-type: sessile or to 2(—4) mm pedun-
culate (in female), simple or forked, scar-covered, brachyblasts 1—5(—20) mm, minutely
pubescent, glabrescent; bracts small, caducous; in male: with 3—7 flowers in a loose
cluster, buds somewhat variable in size; female inflorescences 1- or 2- (or 3-)flowered;
flowers with minute appressed hairs 0.1 mm or less, sometimes late glabrescent. Male
flowers: pedicel 5—7 by 0.8(-1.5) mm, bracteoles small, caducous, 1-2 mm below the
apex; buds (elliptic-)oblong, sometimes narrowed in the lower half, 7-9 by 2-3 mm,
apex (narrowly) rounded or + blunt, perianth lobes thicker lower down, inside with warty-
bullate surface, possibly reddish when fresh; cleft c. 1/4, lobes thinly carnose, 2—2.5 mm
long, 0.2—0.3 mm thick. Androecium 7-9 mm; androphore 3.5—4 by 0.5—1 mm, largely
with appressed hairs 0.10.2 mm; synandrium elongate, 4—5 by 0.8—1 mm, thecae c. 16,
sterile apex + acute, 0.2-0.5 mm. Female flowers: pedicel 2.5—6 by 1-1.5 mm, bracteole
caducous, 0.5—2 mm below the apex; buds ovoid(-oblong), narrowed towards the apex,
5-6.5 by 3-4 mm, lobes 1.5 mm long; ovary ovoid-oblong, including stigma 3—4.5 by
2(-2.5) mm, hairs 0.1-0.2 mm. Fruits solitary or paired, ovoid or oblong, or + fusiform,
sometimes asymmetrical, 3.5—5.5 by 2-3(—4) cm, apex narrowly rounded, blunt, or (sub)-
acute-apiculate, base rounded, pseudostalk absent or broad, to 3 mm, hairs dense, scurfy,
0.1—0.2 mm or less, or wholly or partly (early) glabrescent; pericarp + woody, (3—-)4-8
mm thick, smooth or rugose; seeds ellipsoid, 2-3 cm; fruiting pedicel (rather) stout, 4—
17 by 3-5(—6) mm, striate or often lenticellate and rugose, bracteole scar up to 7 mm
below apex.

Field-notes — Bole straight, or bole and crown irregular, but not buttressed. Bark
dark brown-black or grey-brown, finely grooved, with small flakes; outer bark reddish,
orange, or straw but darkening rapidly; exudate slight, watery, reddish, turning brown;
wood pale brown, or pinkish, developing brown stains when cut, gummy red exudate
from bark and wood; sapwood concentrically layered; heartwood pale brown. Leaves
below grey or glaucous, once reported as scaly. Flowers cream, pale brown, or orange.
Fruits yellowish green; seeds light or black brown.

Distribution — Malesia: New Guinea, with three subspecies.

Habitat & Ecology — Canopy or subcanopy tree of (sub)montane areas; mixed mon-
tane forest, Araucaria—Anisoptera, fagaceous, or Castanopsis-dominated forest, forest
on ridges; sandy-clayey soil, or once from dark brown loam soil over limestone; (600-)
700-2100 m altitude; fl & fr. throughout the year.

Notes — 1. The holotype, Bamler 50, has been destroyed in B; an isotype in WRSL is
a poor specimen. According to Warburg (1.c.) and Markgraf (1.c.) Bamler 50 was col-
lected at c. 700 m, rather low for the species. It has comparatively thin, membranous
leaves, but generally it agrees well with the rest of the material seen. Warburg mentioned
the fruiting pedicel as conspicuously thick, 8 mm, but in the isotype in WRSL this is
only 5 mm thick.

444 Flora Malesiana, Ser. I, Vol. 14 (2000)

2. Myristica crassipes has largely the same circumscription here as that adopted by
Sinclair (1968: 353), although several specimens now in M. crassipes were named by
Sinclair M. longipes, M. gracilipes, and M. cucullata (with the leaves dotted below).
Foreman (1978: 190) included M. firmipes under M. crassipes and Sinclair (1.c.) includ-
ed some specimens with stalked inflorescences, but their views are not followed here.

3. Myristica crassipes resembles both M. tubiflora and M. laevifolia in general habit
(the latter from the same montane habitat), and also M. vinkii. Myristica crassipes also
resembles the lowland species M. flosculosa, both in general habit and in the presence of
the distinctive warty structure of the inner surface of the basal part of the male perianth,
also found in M. pumila, and they seem taxonomically related.

4. Sinclair (1.c.) cited and depicted a richly flowering male specimen, Robbins 1155
(CANB, not seen).

KEY TO THE SUBSPECIES

la. Fruits with dull rust-coloured or greyish brown mealy hairs 0.1 mm or less, or (early)

Slabrescent «oc, H-ce i -Pake--Kaperctass hax Acie eParcpepeote Gress hao Cha eee ee 2
b. Fruits with bright dark-chestnut or chocolate hairs 0.1—0.2 mm, apparently persist-
ent. —E Papua New Guinea, Milne Bay Prov. ............ c. subsp. marronia

2a. Fruits with (sub)persistent indumentum (but hairs minute! ); fruiting pedicel stout,
4-10 by 4-5(-7) mm, often longitudinally cracked with age, bracteole scar up to

3 mm below the fruit. — Papua New Guinea; (500—)700-—1500(—2000) m........
Pe ee eee me ees ores Cen a ee eee a. subsp. crassipes

b. Fruits glabrescent; fruiting pedicel more slender, 6-17 by 3-4 mm, smooth or fissured,
bracteole scar 2-7 mm below the fruit. — West and East New Guinea; (1250—)1500-
CCU a eee re ere tt oe oa oe b. subsp. altemontana

a. subsp. crassipes

Myristica crassipes Warb. in K. Schum. & Laut., Fl. Schutzgeb. Siidsee (1901) 326; Markgr., Bot.
Jahrb. Syst. 67, 2 (1935) 162; J. Sinclair, Gard. Bull. Sing. 23 (1968) 353, f. 55; Foreman, Handb.
Fl. Papua New Guinea | (1978) 190, p.p.; W.J. de Wilde, Blumea 40 (1995) 274, f. 6d.

Twigs at apex 1.5—2(—3) mm diameter. Leaves chartaceous. Fruits with (sub)persistent
indumentum (but hairs minute!); fruiting pedicel stout, 4-10 by 4—5(—6) mm, often lon-
gitudinally cracked with age, bracteole scar up to 3 mm below the fruit. — Fig. 62g,
83d.

Distribution — Malesia: Papua New Guinea.

Habitat & Ecology — Lower montane forest; (500—)700—1500(—2000) m altitude.

Notes — 1. I have seen two duplicates of Carr 15509: that in K, with male flowers,
easily keys out to M. crassipes (subsp. crassipes); that in L is vegetatively identical, but
has male inflorescences without flowers and a 1-4 mm long peduncle ending in a
worm-like brachyblast up to 20 mm. Therefore M. crassipes also keys out to species
with stalked inflorescences, in particular M. fugax and M. sogeriensis. These species
have similar male flowers, but are much more delicate in habit.

2. Jacobs 8818-B has detached fruits which may belong to M. crassipes, but they are
larger, c. 6.5 cm long, and pointed at the apex, narrowed at the base.

De Wilde — Myristicaceae (Myristica) 445

b. subsp. altemontana W. J. de Wilde

Myristica crassipes Warb. subsp. altemontana W.J. de Wilde, Blumea 40 (1995) 274. — Type:
NGF 41681 (Henty, Isgar & Galore), Papua New Guinea.

Twigs at apex (1.5—)2—3 mm diameter. Leaves generally (chartaceous-)coriaceous,
nerves impressed above, rendering the blade sometimes almost bullate. Fruits ellipsoid
or broadly fusiform, early glabrescent; fruiting pedicel more slender, 6-17 by 3-4 mm,
smooth or fissured, bracteole scar 2-7 mm below the fruit.

Distribution — Malesia: New Guinea (eastern Papua Barat and Papua New Guinea).

Habitat & Ecology — Montane forest; (1250—)1500—2000 m altitude.

c. subsp. marronia W.J. de Wilde

Myristica crassipes Warb. subsp. marronia W.J. de Wilde, Blumea 40 (1995) 275. — Type: LAE
54345 (Stevens & Veldkamp), Papua New Guinea, Milne Bay Prov., Mt Suckling.

Tree 10-15 m. Leaves membranous-chartaceous. Fruits with persistent, bright dark-
chestnut or chocolate hairs, 0.1—0.2 mm.

Field-notes — Bark brown with shallow fissures; inner bark reddish; wood pale straw
with + orange markings. Leaves glossy above, glaucous below. Flowers brownish out-
side, green inside. Fruits brown.

Distribution — Malesia: Papua New Guinea (Milne Bay Prov., Mt Suckling; known
from two collections).

Habitat & Ecology — Montane forest near river, banks of rivulet in open Araucaria
forest; 1700-1800 m altitude; fl. & fr. June, July.

31. Myristica cucullata Markgr.

Myristica cucullata Markgr., Bot. Jahrb. Syst. 67 (1935) 166; J. Sinclair, Gard. Bull. Sing. 23 (1968)
364, p.p., for the type only, f. 60G; Foreman, Handb. FI. Papua New Guinea | (1978) 192, p.p.;
W.J. de Wilde, Blumea 40 (1995) 275. — Type: Ledermann 10131, Papua New Guinea.

Tree 6—8(—30, see note 1) m. Twigs terete or faintly angular, (2—)3—4 mm diameter,
early glabrescent, hairs greyish, less than 0.1 mm; older twigs coarsely striate (but nei-
ther cracking nor flaking), with inconspicuous small brown lenticels. Leaves chartaceous
or subcoriaceous, oblong-lanceolate, 14—18(—23) by 4—6(—6.5) cm, base attenuate or
narrowly rounded, apex acute-acuminate; upper surface dark brown, lower surface grey-
brown, early glabrescent, hairs scattered, less than 0.1 mm; not papillose; dots present,
sometimes only few: midrib flat or raised above, lateral nerves 20-30 per side, at 70-80°
to the midrib, flat and inconspicuous above, hardly visible below, lines of interarching
and venation indistinct; petiole 10-20 by 2—2.5 mm; leaf bud 10 by 2 mm, hairs less than
0.1 mm. Inflorescences among the leaves and below, of the Knema-type: sessile, scar-
covered, subglabrous, brachyblasts to 5 mm diameter; bracts 1.5 mm, hairs rough, 0.2
mm, caducous; in male: a subumbel of 2—4 flowers, buds of slightly variable size; fe-
male inflorescences (from the infructescences, see note |) few-flowered, similar to male;
flowers with + scattered hairs less than 0.1 mm. Male flowers: pedicel 3—4 by 1-1.5
mm, bracteole large, deeply cucullate, 8-9 by 6 mm, with short-fimbriate margin, at first
completely enveloping the perianth, caducous; buds (slightly immature) coriaceous,

446 Flora Malesiana, Ser. I, Vol. 14 (2000)

ovoid-ellipsoid, 8 by (5-)6 mm, apex rounded or subacute, base rounded, clefticnt/3,,
lobes 2(—2.5) mm long, 0.5—0.6 mm thick. Androecium + stout, 7 mm; androphore 1|(—2)
by 2 mm, glabrous; synandrium broad-cylindrical, 6 by 2.5(—3) mm, narrowed to the
apex, thecae c. 40, narrow and tightly contiguous, sterile apex minute, 0.1 mm. Female
flowers not seen. Infructescence just below the leaves. Fruits (LAE 61173) (somewhat
immature) solitary, sessile, coarsely wrinkled, ovoid-ellipsoid, 4 by 2.5 cm, apex narrowly
rounded and with short pointed style-remnant, base narrowed into a short pseudostalk
3 mm, hairs rather pale grey-brown, scale-like, 0.1 mm or less; pericarp S—7 mm thick,
with faintly warted surface; seeds not seen; fruiting pedicel 10 by 5 mm, with few fissures
and lenticels, bracteole scar 4 mm below the apex.

Field-notes —Tree 30 m tall (LAE 6/173). Outer bark rusty brown; inner bark straw
with rusty brown exudate; wood straw. Leaves midgreen below. Fruits green brown.

Distribution — Malesia: Papua New Guinea; known only from the type (male flow-
ers), a sterile specimen from East Sepik, and a fruiting collection from Gulf Prov. (see
note 1).

Habitat & Ecology — Submontane, mossy forest with much moss in the tree crowns,
and foothill rain forest; (250—)800—1000 m altitude; fl. Dec.; fr. Mar.

Notes — 1. Markgraf described Myristica cucullata in 1912 based on two collections
from E Sepik Province, Ledermann 10131 (Lordberg, 1000 m, male fl., type) and Leder-
mann 9110 (Etappenberg, 850 m, sterile). The original specimens were lost in B, but
there are duplicates in L. Only the type is fertile. Its somewhat immature male flowers
have a very characteristic hooded bracteole, apparently caducous at anthesis. Both Sinclair
(1968) and Foreman (1978) accepted the species, and included numerous later collec-
tions, but here the species is restricted to the original specimens, with only one addi-
tional collection, LAE 61173 (Croft et al.), with somewhat immature fruit. This speci-
men agrees vegetatively in details with the isotype at L, and the fruits differ from those
of all other Myristica species from New Guinea. The only suspicious fact may be that
LAE 61173, a 30 m tall tree, was collected at a much lower altitude (240 m) in Gulf
Province, above Purari R., far from the type locality, in rain forest, on steep terrain.

2. The male flowers of M. cucullata have a coriaceous perianth, a stout androecium
acute at the apex, and a short androphore. This supports the view that M. cucullata 1s re-
lated to M. brassii, in the group of species with leaves that have dark brown dots below;
the type of M. brassii differs in general aspect, and the male flower has a hairy androphore
and fewer anthers.

32. Myristica cumingii Warb.

Myristica cumingii Warb., Mon. Myrist. (1897) 442. t. 13: Merr., Bull. Bur. For. Philipp. 1 (1903)
21; Enum. Philipp. Flow. Pl. 2 (1923) 178; W.J. de Wilde, Blumea 42 (1997) 154. — Syntypes:
Cuming 903, 1570; Vidal 854 (excl. Vidal 1679 = M. agusanensis; the FI and L duplicates =
Knema glomerata), Philippines.

?Myristica spuria Blume, Rumphia | (1837) 181, nom. nud. [For explanation see J. Sinclair, Gard.
Bull. Sing. 23 (1968) 437.]

Gymnacranthera negrosensis Elmer, Leafl. Philipp. Bot. 2 (1909) 576; (1911) 1057. — Type: Elmer
10133, Philippines.

Gymnacranthera urdanetensis Elmer, Leafl. Philipp. Bot. 8 (1915) 2773. — Syntypes: Elmer 13294,
13295, Philippines.

De Wilde — Myristicaceae (Myristica) 447

Myristica mindorensis Merr., Philipp. J. Sci., Bot. 13 (1918) 281; Enum. Philipp. Flow. Pie2 (1923)
179. — Type: Merritt 3698, Philippines.

Myristica nitida Merr., Philipp. J. Sci., Bot. 13 (1918) 282; Enum. Philipp. Flow. PI. 2 (1923) 179 =
Myristica alvarezii Merr., nom. ined. — Type: FB 22199 (Alvarez), Philippines.

Myristica ceylanica A.DC. var. ceylanica auct. non A.DC., p.p.: J. Sinclair, Gard. Bull. Sing. 23
(1968) 437, f. 77, for the Philippine material.

Tree 6-20 m. Twigs 1.5—2 mm diameter, yellowish brown, early glabrescent, in-
dumentum minute; older twigs sometimes faintly longitudinally cracking; lenticels few,
small and inconspicuous. Leaves membranous or chartaceous, elliptic-oblong to oblong-
lanceolate, 8.5—28 by 2.5—9 cm, base cuneate, apex subacute to acute-acuminate; upper
surface (olivaceous-)brown, lower surface greyish brown with contrasting bright brown
nerves, early glabrescent, hairs weak, scale-like, less than 0.1 mm; faintly papillose; dots
absent: midrib above flat or slightly raised, lateral nerves (6—)10—20 per side, at 45 —70°
to the midrib, slightly raised or sunken above, lines of interarching + distinct, venation
sometimes indistinct; petiole 10—22 by 1.5—3 mm; leaf bud 7-12 by 1.5—2 mm, hairs less
than 0.1 mm. Inflorescences among the leaves or below, + as in Knema; in male: subsessile
or with a peduncle to 15 mm, with at apex an usually bifurcate, scar-covered brachyblast
to 5 mm, with 5-10 flowers in a subumbel, buds of various sizes; female inflorescences
smaller than males, (sub)sessile, few-flowered; flowers late glabrescent, hairs sparse,
grey-brown to dark brown, 0.1 mm. Male flowers: pedicel (2—)3—6 mm, bracteole 1.5—2
mm, (sub)persistent; buds ovoid-oblong, 4.5—-7 by 3-4 mm, apex + blunt or subacute,
+ angled, base broadly rounded, cleft c. (1/3—)1/2, lobes membranous, (0.2—)0.3—0.4
mm thick, erect or reflexed at anthesis. Androecium 4—5.5 mm; androphore 1.5—2 by
0.6-1 mm, minutely pubescent (hairs 0.1 mm or less) in the lower 2/3; synandrium 2.5—
3 by 0.6-1.2 mm, thecae 10-14, not tightly contiguous, sterile apex blunt, 0.3-0.5 mm.
Female flowers: pedicel 2-3 mm, bracteole 2 mm, persistent; buds ovoid-oblong, (4—)
5-6.5 by 3—4 mm, narrowed and triangular in the upper 2/3, lobes reflexed at anthesis;
ovary ovoid, hairs dense, 0.1 mm. Fruits 1 or 2 per infructescence, broadly ovoid, sub-
globose, or ellipsoid, 3-5(—5.5) by 2.5—4.5 cm, hairs dense, rusty-brown, + mealy, 0.1—
0.2(-0.3) mm; pericarp 4—-6(—10) mm thick; seeds broad-ellipsoid, 2.5—3.8 cm; fruiting
pedicel 3—5 mm.

Field-notes — Flowers yellow. Fruits golden.

Distribution — Malesia: Philippines (Luzon, Mindoro, Sibuyan, Ticao, Samar, Leyte,
Bililar, Negros, Mindanao).

Habitat & Ecology — Common in lowland forest, rocky slope forest; 0-850 m alti-
tude; fl. mainly May to Aug.; fr. throughout the year.

Uses — Wood used for firewood and house construction. According to Sulit (PNH
17645) the bark is boiled and the decoction drunk to prevent the spitting of blood.

Notes — 1. Although much resembling M. ceylanica from Sri Lanka in appearance,
I cannot agree with Sinclair (I.c.), who considered M. cumingii as identical. Differ-
ences are, for instance, the possibly yellow aril of M. ceylanica, and the not at all angular
perianth in that species.

The widespread M. iners and the Palawan-endemic M. umbellata much resemble M.
cumingii; M. iners differs in paniculate inflorescences and early glabrescent fruits while
in M. cumingii the inflorescences are usually subsessile brachyblasts. Myristica umbellata,
with rather small glabrescent fruits, at first with minute hairs, may be very similar to M.

448 Flora Malesiana, Ser. I, Vol. 14 (2000)

cumingii. The former differs in its more delicate appearance, smaller fruits, a male inflo-
rescence presumably lasting but one flowering season, more delicately built male flow-
ers with relatively long and slender pedicel, and especially in the distinctive papillae on
the lower leaf surface, and bracteole borne well below the perianth in female flowers.

2. Myristica cumingii is variable in its fruits: shape and size, thickness of the pericarp,
and the length of the hairs; their shapes range from ellipsoid to nearly globose.

3. Most (but not all) specimens from Mindoro differ from the remainder of the mate-
rial from all over the Philippines in a more delicate habit, a thinner leaf blade, drying
more greenish above and paler below (possibly caused by the presence of scattered pa-
pillae), 6—13 pairs of lateral nerves, male inflorescences with slender peduncles (5—15
mm), and relatively small, ellipsoid fruits 3-4 cm long and a 3—4 mm thick pericarp.
Whether these collections represent a separate taxon remains as yet uncertain, but it
is noted that one of the syntypes of M. cumingii, Cuming 1570 (male, also depicted by
Warburg, |. c.: t. 3), represents this different form, and possibly the type of synonymous
M. mindorensis, Merritt 3698, as well.

33. Myristica cylindrocarpa J. Sinclair

Myristica cylindrocarpa J. Sinclair, Gard. Bull. Sing. 23 (1968) 337, f. 50, p.p.; Foreman, Handb.
Fl. Papua New Guinea | (1978) 192, p.p.; W.J. de Wilde, Blumea 40 (1995) 275. — Type: NGF
10288 (K.J. White), Papua New Guinea, ‘Josephstaal’, Madang Prov.

Tree 6-7 m. Twigs 1.5 mm diameter, + angular, reddish brown, early glabrescent,
hairs densely set, greyish, scale-like, less than 0.1 mm; older twigs blackish; lenticels
not apparent. Leaves thinly chartaceous, elliptic-oblong, 7-16 by 2.5—6 cm, base short-
attenuate, apex faintly acute-acuminate; upper surface olivaceous-brown, lower surface
silvery grey, glabrescent, hairs scattered, pale, scale-like, less than 0.1 mm; partly minutely
papillose; dots absent; midrib + flat to raised above, lateral nerves 8-10 per side, at
70—80° to the midrib, sunken above, raised, red-brown and contrasting below, lines of
interarching distinct, venation finely reticulate, not very distinct; petiole 7-13 by 1-1.5
mm; leaf bud 5 by 1 mm, hairs 0.1 mm or less. Male inflorescences, male and female
flowers not seen. Female inflorescences (from infructescences): among the leaves,
supra-axillary (see note 2), peduncle 2-6 by 1—1.5 mm, early glabrescent, with few len-
ticels, ending in a short scar-covered brachyblast 2-3 mm, with (presumably) bor 2
flowers. Fruits 1 or 2 per infructescence, (ovoid- or) ellipsoid-oblong, 2.7—2.8 by 1.2-
1.3 cm, apex acute, base rounded, pseudostalk absent, hairs dense, mealy, bright brown,
0.1 mm; pericarp 1 mm thick; seeds ellipsoid-oblong, 2.3 cm; fruiting pedicel 3-4 mm,
with fine hairs 0.1 mm, late glabrescent, the scar of the bracteole almost at the apex.

Field-notes — Fruits cylindrical, orange-brown. According to Sinclair (1. c.: 339) there
may be up to 4 fruits per infructescence, the seeds measure 20 by 8 mm, are dark brown,
glossy, the aril red and devided into four main segments.

Distribution — Malesia: Papua New Guinea (Madang Prov.); known only from the
type.

Habitat & Ecology — Common on slight rise in a seasonally inundated forest; alti-
tude c. 75 m; fr. Sept.

Notes — 1. Sinclair (1. c.) cited three specimens; besides the type also Robbins 1567
(not seen), and Darbyshire 929 (Central Prov.), recently described as M. brevistipes.

De Wilde — Myristicaceae (Myristica) 449

2. The inflorescences (infructescences) are concaulescent with the main stem for up
to 10 mm, i.e. nearly the length of an internode, so that they are inserted nearly opposite
the leaf of the following node; to check the consistency of this singular character more
material is needed.

34. Myristica dasycarpa W.J. de Wilde

Myristica dasycarpa W.J. de Wilde, Blumea 43 (1998) 171, f. 2. — Type: Regalado & Takeuchi
1520, Papua New Guinea.

Tree c. 10 m. Twigs with 2 conspicuous wings 2(—3) mm high joining the bases of the
petioles, 8-10 mm diameter, bright brown, partially somewhat swollen and hollow, ant-
hole present; older twigs becoming coarsely rectangular cracked and flaking; lenticels
slightly concolorous. Leaves membranous or thinly chartaceous, oblong, 32—47 by
13-19 cm, base (narrowly) cordate, apex broadly acute-acuminate; upper surface oliva-
ceous(-brown), lower surface with dense cinnamon hairs 0.1(—0.3) mm; not or indis-
tinctly papillose; dots absent; midrib slender, flat or slightly raised above, lateral nerves
20-25 per side, at 70—80° to the midrib, flat, lines of interarching + distinct, venation
finely scalariform, indistinct; petiole short, almost pulvinate, 15-20 by 8 mm, glabrous;
leaf bud not seen. Male inflorescences not seen (but see note 2). Female inflorescences
(from infructescences): of the Knema-type, sessile, woody brachyblasts 5 mm diameter;
female flowers not known. Fruits subsessile, 2 or 3 per sessile knotty infructescence,
on older twigs below the leaves, ovoid-ellipsoid, 3.5—4 by 2 cm, apex narrowly rounded
or (sub)acute, base broadly rounded, velutinous, with rufous-rusty hairs 0.5—1 mm; the
pericarp wrinkled and apparently much shrunken on drying, 3 mm thick; seeds oblong,
2.5 cm, grey, the aril impressed into the seed; fruiting pedicel rather stout, 2(—3) mm. —
Fig. 71.

Field-notes — Myrmecophilous. Leaves brown tomentose below. Fruits rich brown
to orange-brown; crushed dried fruit strongly smelling of nutmeg.

Distribution — Malesia: northern Papua New Guinea (East Sepik Proy., Ambunti
subprov., Waskuk Hills, area around Langu and Garuka villages); known only from the
type.

Habitat & Ecology — Subcanopy tree; on ridge; 50—70 m altitude; fr. July.

Notes — 1. Myristica dasycarpa is readily distinguishable among the New Guinean
myrmecophilous Myristicas by its thick alate twigs and large leaves, and its subsessile
fruits, 3.5—4 cm long, with velvety hairs to 1 mm, forming the most conspicuous indu-
mentum of the group.

2. Male flowers of this species are not known with certainty. Brass 13706 (Papua
Barat, Idenburg River, in flood plain forest at 850 m) possibly belongs here; in L it has
inflorescences with immature male flowers and separate fruits, the latter obviously be-
longing to M. subalulata. The male inflorescence is a wart-like brachyblast, 3—4 mm,
pubescent, glabrescent, bracts 1.5 mm, subumbel of 5 or 6 flowers, buds of variable
sizes; male flowers: pedicel slender, 6 mm, bracteole 2 mm; buds (immature) ellipsoid,
5-6 by 3.5 mm, not collapsing on drying, cleft c. 1/3?, lobes 0.5 mm thick; androecium
4 mm; androphore narrow, 1 mm, almost glabrous; synandrium 3 by | mm, acute, thecae
c. 28, sterile apex + absent.

450 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 71. Myristica dasycarpa W.J. de Wilde. a. Part of leafy twig; b. detail of indumentum of lower
leaf surface; c. lower part of older twig with ramiflorous infructescences [Regalado & Takeuchi
1520]. — Scale bar fora, c = 2 cm; for b= 0.5 mm.

De Wilde — Myristicaceae (Myristica) 45]

35. Myristica depressa W. J. de Wilde

Myristica depressa W.J. de Wilde, Blumea 42 (1997) 161; Tree Fl. Sabah & Sarawak 3 (2000) 459.
— Type: S 39018, Sarawak.

Tree 10—30(—40) m. Twigs 2-3 mm diameter, early glabrescent, hairs 0.1 mm or less,
sometimes yellowish; older twigs dark brown or blackish, coarsely striate, sometimes
flaking; lenticels absent. Leaves membranous or thinly chartaceous, oblong(-lanceolate),
15-30 by 4—9 cm, base short-cuneate or rounded, apex acute-acuminate; upper surface
olivaceous(-brown), lower surface grey-brown, with persistent dense but thin felty or
arachnoid covering of intricate greyish brown hairs 0.1 mm or less, hardly to be felt with
the finger (lens!); not obviously papillose; dots absent; midrib above flat or + raised,
lateral nerves 16-23 per side, flat or sunken above, sometimes indistinct, at 45-80° to
the midrib, lines of interarching distinct or not, venation faint on both surfaces (distinct,
scalariform in Zainal Abidih 23, from Nunukan I.); petiole 10—25 by 1.5—3 mm; leaf bud
10-15 by 1.5-3 mm, hairs 0.1—0.2 mm. Inflorescences between the lower leaves, pedun-
culate, slender, hairs woolly, 0.1—0.3 mm; in male: 2—3.5 cm, peduncle 5—10 mm, slightly
flattened, lowest branches (sub)opposite, to 5 mm, ending in a subumbel of flowers,
central axis 20-30 mm, with 2-4 sessile subumbels, each with 4-8 flowers, buds of
strongly variable sizes, hairs persistent, woolly, (0. 1-)0.2—0.3 mm; bracts broadly rounded,
1.5-2 mm, pubescent, caducous; female inflorescences as males but smaller. Male flow-
ers: pedicel 3—3.5 mm, bracteole often + 2-topped, 2 mm; buds firm, ovoid or ellipsoid,
3-4 by 2.5-3 mm, apex + rounded, not or hardly angled, cleft 3/4—4/5, lobes 0.5—0.8
mm thick. Androecium short and broad, somewhat club-shaped, 1.5 mm; androphore
0.3-0.5 by 0.8 mm, minutely pubescent (apparently glabrous in immature flowers of
bb 27505 from Sumatra); synandrium broadly ellipsoid, 1—1.3 by 1—-1.2 mm, the sterile
apex absent, the apex truncate and + hollowed for up to 0.3 mm, the edge + lobed cor-
responding with the anthers, thecae 12—16(—20), 1 mm long, not tightly contiguous.
Female flowers not seen. Infructescence 0.5—2 cm, with 1 or 2 fruits. Fruits ellipsoid
(-oblong), 5—7 by 3-4.5 cm, apex and base broadly rounded, hairs dense, scurfy, 0.1
(0.2) mm; pericarp (2—)3—6(—10) mm thick; seeds ellipsoid, 4-5 cm; fruiting pedicel
+ stout, roughly fissured or cracked, 3-5 mm.

Field-notes — With or without buttresses and stilt-roots; bole straight; outer bark brit-
tle, fissured or flaky, blackish; inner bark brownish; sapwood white or brownish. Twigs
blackish. Flowers yellowish (grey). Fruits fawn, light brown, or apricot, aril pink.

Distribution — Malesia: Sumatra (one collection, Indragiri, and one deviating speci-
men, see note 2), Peninsular Malaysia, Borneo (C Kalimantan).

Habitat & Ecology — Primary mixed dipterocarp forest, on various soil types: occa-
sionally inundated riverbanks, sandstone and diorite screes, hillsides, stony soils, yel-
low loamy soil; 0-300 m altitude; fl. & fr. throughout the year.

Notes — 1. Myristica depressa is close to M. borneensis, M. malaccensis, and M. wyatt-
smithii, but differs in its persistent indumentum on the lower leaf surface. The androecium
is distinctly short and broad, resembling that of M. borneensis and especially that of M.
malaccensis in which the sterile apex also is absent. Fruits and male flowers of M. ma-
laccensis are glabrous (early glabrescent), the latter cleft to only 1/3. Myristica borneen-
sis has fruits similar to those of the present species, but differs in lacking a dense indu-
mentum on the lower leaf surface, the male flowers which are cleft to only about halfway,

452 Flora Malesiana, Ser. I, Vol. 14 (2000)

and the androecium with a short and broad sterile apex. Other apparently related species
are M. agusanensis and M. guatteriifolia, both with a persistent indumentum on the lower
leaf surface, and both with an androecium sometimes hollowed at the apex.

2. The specimen Kostermans s.n. (Sumatra, Palembang) has leaves that dry more
brownish above, the lateral nerves are indistinct, and the blade base is conspicuously
acute-attenuate; the (almost mature) fruit is only 4.5 cm long when dry. The specimen
may represent a separate taxon.

36. Myristica devogelii W. J. de Wilde
Myristica devogelii W.J. de Wilde, Blumea 42 (1997) 163. — Type: de Vogel 6083, C Sulawesi.

Tree 20 m. Twigs 4—5 mm diameter, early glabrescent, hairs rusty or greyish, less than
0.1 mm; older twigs coarsely striate and somewhat fissured; lenticels small, not con-
trasting. Leaves coriaceous, oblong or oblong(-lanceolate), 20-30 by 6-10 cm, base
short-cuneate or nearly rounded, apex acute or subacute-acuminate, or rather blunt; up-
per surface olivaceous(-brown), lower surface glabrous (early glabrescent), brown; not
papillose; dots absent; midrib above flat, lateral nerves (12—)15—20 per side, at (45—)60°
to the midrib, slender, flat and faint above, faint below, lines of interarching and vena-
tion indistinct; petiole 15—20 by 5 mm; leaf bud 15-20 by 4—5 mm, hairs scale-like,
grey(-brown), less than 0.1 mm. /nflorescences among the leaves or just below, of the
Knema-type, a sessile scar-covered brachyblast 3-5 mm diameter, rusty-greyish short
pubescent; bracts small, caducous; in female: with 1-3 small flowers or | or 2 flowers
together with a submature fruit; flowers with short appressed brown hairs 0.1 mm. Male
inflorescences and male flowers not seen. Female flowers: pedicel short, less than 0.5
mm, bracteole + saucer-shaped, 1.5 mm, (sub)persistent; buds + ovoid, 5 by 3 mm, lobes
not seen; ovary ovoid, 3—-3.5 by 2.5 mm, hairs dense, rusty, appressed, 0.1(—0.2) mm.
Fruits usually solitary, sessile, subglobose, (4—)5 cm diameter, hairs scurfy, rust-colour-
ed, 0.1 mm or less; pericarp woody, (6—)8—10 mm thick; seeds not seen; fruiting pedicel
(almost) absent.

Field-notes — Straight tree, bole with or without buttresses, prop-roots spreading to
1.20 m. Bark fissured, not peeling; exudate red watery and gum-like.

Distribution — Malesia: Central Sulawesi (N side Lake Matano).

Habitat & Ecology — Forest on alluvial flat on conglomeratic soil, and on ultrabasic
bedrock; 400-450 m altitude; fr. June, July.

Note — Myristica devogelii is readily distinguishable by its glabrous, coriaceous leaves
and sessile, globose fruits of 5 cm diameter.

37. Myristica duplopunctata W.J. de Wilde

Myristica duplopunctata W.J. de Wilde, Blumea 40 (1995) 275. — Type: BW 9041 (Iwanggin),
West Irian.

Tree 10—25 m. Twigs subterete or + angular and grooved, 3-6 mm diameter, early
glabrescent, indumentum minute; bark smooth or fissured; with or without lenticels.
Leaves chartaceous or (thinly) coriaceous, (elliptic-)oblong, 15—23 by 5-8 cm, lower
surface with dense interwoven scale-like hairs 0.1 mm (lens!), seemingly + glabrous, or
late glabrescent; not papillose; upper and lower surface at least towards the margin with

De Wilde — Myristicaceae (Myristica) 453

scattered small dots (lens!), sometimes obscured by the indumentum; midrib raised above,
lateral nerves 20-30 per side, at 70—80° to the midrib, flat or sunken above, faint below,
venation invisible. Inflorescences between the (lower) leaves, almost as in Knema, but
pedunculate, in male peduncle (3—)10 mm, in female 5 mm, ending in a small scar-
covered part with a cluster of few flowers. Male flowers presumably resembling those of
M. brassii and M. neglecta; known at anthesis only in subsp. duplopunctata. Fruits soli-
tary, 4-5 cm, known only in subsp. versteeghii.

Distribution —Incompletely known taxon from the mountains of NW Papua
Barat and northern W Papua New Guinea; the rather heterogeneous material is placed
in two subspecies.

Notes — 1. Myristica duplopunctata is similar to M. millepunctata and M. neglecta,
but differs in smaller leaves with small scattered dots on both surfaces.

2. The male flowers of M. brassii, M. duplopunctata, and M. neglecta are all very
similar. However, the staminal column is hairy along its entire length in M. neglecta,
only in the lower half in M. brassii, and very inconspicuously hairy, with pale hairs less
than 0.1 mm, in M. duplopunctata.

KEY TO THE SUBSPECIES

la. Twigs 5 mm diameter. Leaves (thinly) coriaceous. — Papua Barat / Papua New Guinea,

northern side of Central Range, 1500-2100m ........... b. subsp. versteeghii
b. Twigs 3 mm diameter. Leaves + chartaceous. — Papua Barat, northern coastal moun-
Pam) OOM Mier hs 22nd DR I a a a. subsp. duplopunctata

a. subsp. duplopunctata

Tree c. 25 m. Twigs terete, 3 mm diameter, early or late glabrescent, hairs 0.1 mm;
older twigs (coarsely) striate; lenticels few or numerous. Leaves + chartaceous; blades
(oblong-)lanceolate, 15-23 by 5-8 cm, base (short-)cuneate, apex acute(-acuminate);
upper surface olivaceous or brown; lower surface pale brown with dense layer of inter-
woven scale-like hairs 0.1 mm or less, or late glabrescent; midrib raised above, lateral
nerves (20—)25—30 per side; petiole 15—20 by (2—)2.5 mm; leaf bud 10-15 by 2 mm,
with hairs 0.1—0.2 mm. Inflorescences in male: peduncle 5-10 mm, slightly flattened,
finely pubescent or glabrescent, ending in a minute scar-covered part with 2 or 3 flow-
ers, buds subequal in size; flowers with minute rusty hairs 0.1 mm or less. Male flowers:
pedicel 13-14 by 1.5 mm, bracteole caducous, its scar 3—-3.5 mm below the apex; buds
(ellipsoid-)oblong, 10 by 4.5 mm, apex and base narrowly rounded, cleft 1/5, lobes
2 mm long, 0.5 mm thick; androecium stout, tapered to the apex, 8 mm, synandrium 4 by
1.5(-2) mm; thecae 16-20, sterile apex acute, 0.7—1 mm, androphore stout, 3(—3.5) by
1.5-2 mm, in the lower half with minute whitish hairs less than 0.1 mm. Female flowers
and fruits not seen.

Field-notes — In young tree: bark black, slightly flaking and with some watery sap;
sapwood pinkish, heartwood brown; adult tree: outer bark grey; inner bark white within,
with little red exudate, slash light brown; sapwood white. Flowers light brown.

Distribution — Malesia: N Papua Barat (known from three collections).

Habitat & Ecology — Coastal mountains; 500-700 m altitude.

Note — May resemble M. mediovibex, which is more delicate, with leaves not dotted.

454 Flora Malesiana, Ser. I, Vol. 14 (2000)

b. subsp. versteeghii W. J. de Wilde

Myristica duplopunctata W.J. de Wilde subsp. versteeghii W.J. de Wilde, Blumea 40 (1995) 277.
— Type: Brass & Versteegh 12512, West Irian.
Myristica cucullata auct. non Markgr., p.p.: J. Sinclair, Gard. Bull. Sing. 23 (1968) 364, f. 60D-F.

Tree 10-25 m. Twigs + angular or deeply grooved, 5 mm diameter, early glabrescent,
hairs less than 0.1 mm, dark brown; older twigs coarsely striate; lenticels few. Leaves
(thinly) coriaceous, (elliptic-)oblong, 15—20 by 5—8 cm, base (broadly) rounded, apex
+ blunt or acute-acuminate; upper surface dark olivaceous, lower surface grey-brown,
with thin greyish felty interwoven scale-like hairs 0.1 mm or less; midrib flat or some-
what raised above, lateral nerves 22—30 per side (with in addition numerous intersecon-
dary ones); petiole 15—20 by 4-5 mm; leaf bud 15 by 3-4 mn, with appressed hairs less
than 0.1 mm, looking as if glabrous. Inflorescences incompletely known, finely pubes-
cent. Male and female flowers not seen (see note). Infructescence with a stout short
peduncle, 5-10 mm. Fruits solitary, + ellipsoid, 4—5 by 3.5—4 cm, hairs dense, mealy,
rust-coloured, 0.1 mm; pericarp hard, (5—)10 mm thick; seeds ellipsoid, 2.5 cm; fruiting
pedicel 6-8 mm, + stout, bracteole scar at c. 1/3 below the apex.

Field-notes — Outer bark brown or black; inner bark light brown; wood whitish brown
with brown or pink sap. Fruits orange-brown or red-brown, with yellow sap; seed brown.

Distribution — Malesia: Papua Barat (mountains on the north side of the Main Range)
and adjoining W Papua New Guinea (W Sepik).

Habitat & Ecology — Frequent in primary forest on slope of ridge; montane forest on
top of ridge with small trees, shrubs and dense moss dominating the understorey; alti-
tude 900-2100 m; fr. Feb.—Apr.

Note — Subsp. versteeghii may merit the status of a separate species; its inclusion in
M. duplopunctata is provisional, until more and especially (male) flowering material
becomes available.

38. Myristica elliptica Wall. ex Hook. f. & Thomson

Myristica elliptica Wall. ex Hook.f. & Thomson, Fl. Ind. (1855) 162; King, Ann. Roy. Bot. Gard.
Calc. 3 (1891) 295, pl. 113; Warb., Mon. Myrist. (1897) 435, t. 16; J. Sinclair, Gard. Bull. Sing.
16 (1958) 353: 23 (1968) 188; W.J. de Wilde, Blumea 42 (1997) 164; Tree FI. Sabah & Sarawak
3 (2000) 460. — Myristica elliptica var. elliptica: J. Sinclair, Gard. Bull. Sing. 16 (1958) 353,
f. 26, pl. VIA, B; 23 (1968) 188, f. 12 G, H. — Type: Porter in Wallich Cat. 6798a, Penang.

Myristica macrocarpa Wall., Cat. (1832) n. 6798b (for the fruit; herbarium sheet only), nom. nud.

Myristica calocarpa Miq., Fl. Ind. Bat. 1, 2 (1858) 71; Suppl. (1860) 156; (1861) 383; Ann. Mus.
Bot. Lugd.-Bat. 2 (1865) 48. — Type: Teijsmann 477, Sumatra.

Myristica sycocarpa Miq., Fl. Ind. Bat. 1, 2 (1858) 68; Suppl. (1860) 156. — Type: Diepenhorst &
Teijsmann 482, Sumatra.

Fig. 72. Myristica elliptica Wall. ex Hook. f. & Thomson. a. Branch with female inflorescences;
b. female bud with persistent bracteole; c. ditto, opened, showing pistil; d. branch with male inflo-
rescences: e. male flower at anthesis; f. ditto, opened, showing androecium; g. part of infructescence
with one fruit, opened, showing arillate seed; h. lower leaf surface with papilla-like structures and
scattered appressed faint hairs [a—c, h: de Wilde /Duyfjes 16544, d—f: de Wilde/Duyfjes 16452; g:
de Wilde /Duyfjes 18882]. — Scale bar for a, d, g = 2 cm; for b, c, e, f = 1.65 mm; for h = 0.4 mm.

De Wilde — Myristicaceae (Myristica) 455

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456 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 6—40 m. Twigs 2-4 mm diameter, pale, grey or yellowish, early glabrescent,
hairs greyish, 0.1—0.3 mm; older twigs straw or (brown) yellow to greenish yellow, not
longitudinally cracking; sometimes with a few pale brown lenticels. Leaves chartaceous,
elliptic or oblong, 9-21(—32) by 3—9(—12) cm, base narrowly rounded or acute, apex
acute-acuminate; upper surface yellowish or olivaceous, lower surface greyish or pale
olivaceous, + glabrous (early glabrescent), hairs scattered, appressed, greyish or pale
brown, 0.1 mm or less, surface minutely pustulate; papillae not obvious; dots sometimes
present, scattered, faint; midrib above flat or slightly raised, lateral nerves 8-13(-15)
per side, at 45—60(—70)° to the midrib, faint, flat or slightly raised above, distinct on
lower leaf surface, lines of interarching sometimes distinct, venation faint on both sur-
faces; petiole 10—20(—30) by 1.5-2.5 mm, drying the same colour as the twigs; leaf bud
(10—)15—20 by 1—1.5 mm, hairs glossy, light brown, 0.1—0.3 mm. /nflorescences among
the leaves, pedunculate, + glabrescent, hairs (sparse) pale, appressed, 0.1 mm or less;
bracts minute, subpersistent; in male: 1-2.5 by 1-1.5 cm, few-flowered, peduncle slightly
flattened, 5—12 mm, branches simple, up to 5 mm, central axis similar, sometimes absent
or with but a single flower; flowers in loose subumbels of 3—5; buds of various sizes;
female inflorescences like the male, flowers only 1—3; flowers angled, hairs sparse, ap-
pressed, pale, 0.1 mm. Male flowers: pedicel 3-5 by 1-1.5 mn, bracteole + reniform,
often + truncate or faintly + 3-topped, 0.5—1(—2) mm, persistent, apical or rarely to 1 mm
below; buds elliptic-oblong, sometimes slightly contracted below the lobes, 7-8(—9) by
2.5—3.5 mm, apex subacute, 3-angled, cleft c. 1/3, lobes 0.5—0.8 mm thick, perianth
lower down thinner. Androecium 5—7 mm; androphore slightly tapering, 2(—2.5) mm,
glabrous or at base thinly minutely pubescent; synandrium 4—4.5 by (1-)1.5 mm, thecae
10-12, + appressed, sterile apex + blunt, 0.5 mm. Female flowers: pedicel 3 mm; buds
ovoid-oblong, + urceolate, inflated, 6-8 by 3-4 mm, apex acute, angled, cleft c. 1/2,
lobes at anthesis reflexed; ovary long-conical (including style) 5 by 1 mm, hairs dense,
appressed, 0.1 mm. Infructescences often conspicuously lenticellate. Fruits | or 2, ovoid
or ellipsoid(-oblong), 4.5-7.5 by (2-)2.5—4 cm, apex rounded or bluntly narrowed or
somewhat beaked, base often narrowed into 4-12 mm long pseudostalk, early glabrescent,
hairs 0.1 mm; pericarp dull dark brown, 5(—10) mm thick; seeds ellipsoid, 4-5 cm; fruit-
ing pedicel slender, 5—10 mm. — Fig. 72.

Field-notes — Low or tall tree with dense or loose crown, sometimes branched from
the base; bole tapering or not, sometimes fluted or buttressed up to | m high, in marshy
forest with spreading stilt-roots. Bark smooth or finely fissured or scaly, sometimes
lenticellate, grey to dark brown or blackish; inner bark soft, red-brown to pink, sap
watery, pink or red; (sap) wood white to pale yellow. Twigs pale, straw-coloured. Leaves
dark glossy green above, glaucous green or very pale with yellow-green midrib below.
Flowers cream-white or (reddish) yellow, fragrant when crushed. Fruits greenish yellow
turning bright orange or apricot, sometimes with irregular swellings; seeds olive-brown.

Distribution — Peninsular Thailand; Malesia: Peninsular Malaysia, Sumatra, Borneo,
possibly also the Philippines.

Habitat & Ecology — Primarily in swamp and peat swamp forest (Swamp Nutmeg);
also in disturbed forest and belukar; alluvial forest; degraded forest, along rivers; clayey
or (poor) sandy soils, also on red soil; rocky streamsides; 0-500 m altitude; fl. & fr.
throughout the year. According to Sinclair (1.c. 1958: 214; 1968: 190) the seeds float,
a fact possibly related to the wet habitats favoured by the species.

De Wilde — Myristicaceae (Myristica) 457

Uses — Seeds once reported as combustible, containing starch and fat.

Notes — 1. A widespread species of peat swamp forest of W Malesia; not in Java.

2. Sinclair (1. c.) included the varieties celebica and simiarum in this species, both
here included in the related M. simiarum which differs mainly in its inflorescences ter-
minally with perennial brachyblasts, not few-flowered clusters lasting but one season.

3. Frequently a yellowish exudate of unknown origin has been found on dry speci-
mens, especially on the fruits.

39. Myristica ensifolia J. Sinclair

Myristica ensifolia J. Sinclair, Gard. Bull. Sing. 23 (1968) 332, f. 48; Foreman, Handb. Fl. Papua
New Guinea | (1978) 193; W.J. de Wilde, Blumea 40 (1995) 278. — Type: Brass 6857, Papua
New Guinea, Fly River.

Slender treelet, 1.5 m. Twigs bluntly 2- or 3-angular, (1—)1.5—2 mm diameter, some-
what reddish brown, early glabrescent, hairs appressed, greyish, 0.1 mm or less, smooth;
older twigs not seen. Leaves membranous (or chartaceous), lanceolate, parallel-sided,
12-22 by 1.5-3 cm, base attenuate, apex long acute(-acuminate); upper surface gla-
brous, dark brown, lower surface grey(-brown), early glabrescent, hairs minute, scat-
tered, grey, 0.1 mm or less; not clearly papillose but stomata or hair bases(?) look like
minute pustules with a dark-coloured centre (lens!); dots absent; midrib slender, raised
above, lateral nerves 16—20 per side, at 45—60° to the midrib, slender, flat or slightly
sunken above, light brown, contrasting below, lines of interarching distinct, venation
faint; petiole dark (reddish) brown, 8-10 by 1-2 mm; leaf bud 7 by (1—-)1.5 mm, hairs
dense, appressed, bright brown, 0.2(-0.3) mm. Male inflorescences, male and female
flowers not known. Female inflorescences (from imperfect broken infructescence) among
the leaves, peduncle 2 by 1.5 mm, smooth, partly glabrescent, hairs grey-brown, appressed,
0.2 mm, continuing in the fruiting pedicel (broken away in the only known specimen).
Fruits (immature) solitary, oblong-fusiform, base narrowed (pseudostalk), apex acute,
(4-)4.5 by 1.3(—1.5) cm, hairs dense, rust-coloured, scurfy, 0.1—0.2 mm; pericarp 2 mm
thick; seeds not seen; fruiting pedicel not seen, probably slender, c. 10 mm.

Distribution — Malesia: Papua New Guinea (Western Prov., Fly River, 528 mile
Camp); known only from the type.

Habitat & Ecology — In ridge forest undergrowth; 80 m altitude; nearly mature fr.
May.

Note — Despite the numerous recent collections made in New Guinea, this species
remains known only from the type specimen. The fragmentary infructescence shows
that the female inflorescence has a short, 2 mm long peduncle, and therefore it is keyed
out in the group of species with a pedunculate inflorescence; however, specimens with
male inflorescences are needed to confirm this. Myristica ensifolia is readily recogniz-
able since it is a dwarf tree with ensiform leaves which suggests it to be a rheophyte, but
actually it was collected in ridge forest undergrowth. Microscopically the fine crater-
like markings on the lower leaf surface are characteristic.

40. Myristica extensa W. J. de Wilde

Myristica extensa W.J. de Wilde, Blumea 42 (1997) 164; Tree Fl. Sabah & Sarawak 3 (2000) 461.
— Type: S 25678 (Ilias Paie), Sarawak.

458 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 15-25 m. Twigs subterete or angular, 2.5—5 mm diameter, yellowish or blackish
brown, very early glabrescent, hairs pale, 0.1 mm or less; older twigs irregularly (longi-
tudinally) cracking; lenticels few and indistinct, or absent. Leaves membranous or thinly
chartaceous, (elliptic-)oblong(-lanceolate), (16—)24—35 by (5—)8-13.5 cm, base atten-
uate or acute, or narrowly rounded, apex acute-acuminate; upper surface olivaceous
(-brown), lower surface grey-brown, glabrescent, indumentum minute, flat-lying, open
cobweb-like, silvery; not obviously papillose; dots absent; midrib above flat, lateral nerves
16-24 per side, at (45—)60-80° to the midrib, slender, indistinct and flat or + sunken
above, much raised below, lines of interarching and venation indistinct on both surfaces;
petiole 15-30 by 2.5—5 mm; leaf bud 10-15 by 2.5-3 mm, hairs appressed, 0.1 mm.
Inflorescences among the leaves or just below, pedunculate, slender, almost glabrous,
hairs scattered, 0.1 mm or less; in male: widely paniculate, 10-15 by 6-10 cm, peduncle
20-45 mm, somewhat flattened, lower branches subopposite, 10-35 mm, central axis
with 3 or 4 branches, 15 mm or less; flowers in subumbels of 5—10, buds strongly vary-
ing in size; bracts caducous, not seen, their scars dislocated upwards along the branches,
female inflorescences (from infructescences): smaller and fewer flowered, (1.5—)5-8
cm, including the 30-40 mm long peduncle. Male flowers: pedicel slender, 9-12 mm,
bracteole broadly rounded-reniform, 2—2.5 mm, persistent; buds membranous, ovoid,
4(—4.5) by 3-3.5 mm, hairs sparse, grey-brown, 0.1(—0.2) mm, cleft nearly 2/3, lobes
3 mm long, 0.3 mm thick. Androecium club-shaped, 3-3.5 mm; androphore somewhat
tapering, | by 1 mm, glabrous; synandrium ellipsoid(-oblong), 2 by 1-1.2 mm, thecae 16
(anthers rather spaced), sterile apex 0.2—0.3 mm, + blunt. Female flowers not seen. Fruits
| or 2 per infructescence, (elliptic-)oblong, 6-7 by 3-3.5 cm, rounded at both ends, with
dense minute scurf of hairs 0.1 mm or less; pericarp 3-6 mm thick, dull brown; seeds
5—5.5 cm; fruiting pedicel 6-15 mm.

Field-notes — Medium-sized tree; without buttresses, or stem fluted, with spreading
buttresses, or with stilt-roots 80 cm high, 80 cm out. Bark smooth or scaly, hard, 1 mm
thick, cracked, grey to blackish, at the base of the bole peeling off in strips, brittle, dark
brown or blackish; living bark 5(—10) mm, brown; wood brown; sap blood-red or pink,
rarely absent. Leaves below glaucous or whitish. Flower (buds) yellowish green. Fruits
cream, yellow, greenish brown, or yellowish brown, with short brown indumentum; seeds
blackish.

Distribution — Malesia: Borneo (Sarawak, Brunei, C & E Kalimantan).

Habitat & Ecology — Primary lowland forest; at top of spur; steep slope, hill forest
near summit: undulating land; rich clayey or sandy yellow loam (along rivulet), yellow
clayey soil; 0-800 m altitude; fl. June; fr. mainly Mar., Aug., Sept.

Notes — 1. Myristica extensa is similar to M. corticata and M. iners in leaf and in-
florescence features, and to M. malaccensis in the club-shaped androecium. The species
is also close to M. maxima, with widely branched paniculate inflorescences (and in-
fructescences), stouter twigs, and larger leaves and male flowers. The leaves of M. maxima
have more lateral nerves, a more distinct venation, a well-marked marginal nerve, and
a greyish lower leaf surface with minute pale hairs (glabrescent) and distinct papillae.
Its male flowers are larger, with a cylindrical, not club-shaped androecium and with
some indumentum on the androphore. For differences with the other species mentioned
above, see the keys. Fruiting specimens of M. extensa may resemble M. frugifera from
the Philippines.

De Wilde — Myristicaceae (Myristica) 459

2. Some deviating specimens from Sarawak and Sabah (see De Wilde I. c.) may rep-
resent an as yet undescribed species. They key out to fruiting specimens besides M. iners
and M. wyatt-smithii, but differ in general aspect. More material, including flowers, is
needed to solve this problem.

41. Myristica fallax Warb.

Myristica fallax Warb., Mon. Myrist. (1897) 410; Merr., Enum. Born. (1921) 269; W.J. de Wilde,
Blumea 42 (1997) 166; Tree Fl. Sabah & Sarawak 3 (2000) 461. — Type: Beccari 1590, Sarawak.

Tree 15-20 m. Twigs 1-1.5 mm diameter, (yellow-)brown, early glabrescent, hairs
scattered, less than 0.1 mm; older twigs brown, coarsely striate or faintly longitudinally
cracking; lenticels inconspicuous or absent. Leaves thinly membranous, ( oblong-)lanceo-
late, 7-14 by 2—4 cm, base cuneate, apex acute-acuminate; upper surface olivaceous or
(dark) brown, lower surface slightly paler, greyish brown, early glabrescent, hairs scat-
tered, pale grey-brown, stellate, less than 0.1 mm; not papillose; dots absent; midrib
+ flat or slightly raised above, lateral nerves 12—16 per side, at 60—80° to the midrib, flat
or sunken above, slender and little conspicuous below, lines of interarching and vena-
tion indistinct; petiole 10-20 by 1(—1.5) mm; leaf bud 10 by 1-1.5 mm, hairs 0.1 mm or
less. Male inflorescences 2 or 3 times as long as the petioles, 4 cm, with slender pedun-
cle, 10 by 1 mm, higher up once or twice forked, and with few flowers at the ends of
short branches, 2 mm long. Male flowers: pedicel slender, 8 by 0.3 mm, bracteole per-
sistent, 0.5 mm; buds urceolate, 3 mm, at the base 2 mm wide. Androecium 2.3 mm;
androphore | mm, subpuberulous; synandrium 1.2—1.3 mm, sterile apex obtuse, thecae
c. 10. Female inflorescences and flowers not seen. Infructescences in the apical part
of the twigs, among the leaves, peduncle slender, 10—25 by 1.5 mm, with or without a
branch up to 10 mm, with a few scars of fallen flowers, glabrous (early glabrescent).
Fruits solitary, ovoid-ellipsoid, 3.5—4(—4.5) by 2—2.7 cm, apex rounded, base broadly
rounded, glabrous (when immature with pale rusty scurf of scale-like hairs less than 0.1
mm, soon glabrescent), rusty brown; pericarp 3-4 mm thick; seeds ellipsoid, 2—2.5 cm;
fruiting pedicel 5—10 mm, bracteole scar subapical.

Field-notes — Erect tree; buttresses out to 1 m. Bark of trunk smooth or slightly
flaky, chocolate, with paler patches or whitish flakes; sap reddish, watery. Lower leaf
surface glaucous. Fruits light cream or yellow, with loose, thin, brown indumentum.

Distribution — Malesia: Borneo (Sarawak, E Kalimantan).

Habitat & Ecology — Ridge (hill) forest, mixed dipterocarp forest; yellow sandy
loam, tuff-plateau; 100-600 m altitude; fr. Aug., Oct., Nov.

Notes — 1. Myristica fallax is a satellite species of the variable M. iners and comes
close to the type of M. vordermannii, which goes as a delicate form under M. iners. Myr-
istica fallax is distinguishable by its slender twigs and small, thin, membranous, dark-
coloured leaves. Infructescences are long with a slender peduncle, fruits are small,
3.5—4 cm, and early glabrescent. Specimens of M. fallax are reminiscent of M. fragrans.

2. [have not seen the type of M. fallax, Beccari 1590 (Sarawak, Mt Mattang) in FI;
the description of the male inflorescences and flowers above is based on Warburg (1.c.).

3. Several tiny, slenderly built specimens (including some with immature male flow-
ers and inflorescences with comparatively long peduncles) now inserted under M. iners,
link up with M. fallax; this should be given attention during future investigations.

460 Flora Malesiana, Ser. I, Vol. 14 (2000)

42. Myristica fasciculata W.J. de Wilde

Myristica fasciculata W.J. de Wilde, Blumea 40 (1995) 278; 43 (1998) 173. — Type: Hoogland &
Craven 10400, Papua New Guinea.

Tree 5—9 m. Twigs subterete or 2-angled, with lines or wings to | mm high joining the
bases of the petioles, (4-)5—7 mm diameter, early glabrescent, hairs to 0.1 mm, hollow
and faintly swollen, in places with slit-like perforation, inhabited by ants; older twigs
not flaking, with small conspicuous lenticels. Leaves membranous, oblong, 20-35 by
7.5-14 cm, base (short-)cuneate, rounded, or shallowly cordate, apex acute-acuminate;
upper surface olivaceous, lower surface greyish; papillose or not (see note), dots absent;
midrib slightly raised towards the apex, lateral nerves 17-30 per side, at 60—80° to the
midrib, flat or impressed above, lines of interarching and venation distinct; petiole 15-
25 by 2-3 mm; leaf bud 10-15 by 2-3 mm, hairs scale-like, 0.1 mm. Inflorescences be-
tween the leaves or below, of the Knema-type, (sub)sessile, scar-covered brachyblasts,
simple or forked, to 7 by 4 mm, finely brown pubescent; bracts small, caducous; in male
with subumbels of 8—15 flowers, buds about equal in size; female inflorescences with
6-10 flowers, buds equal in size, sometimes present together with fruits of previous
season: flowers with dense or sparse brown hairs 0.1 mm or less, sometimes glabrescent.
Male flowers (immature): pedicel 10-11 by 1(—1.5) mm, bracteole sometimes + 3-topped,
3.5 mm, subpersistent; buds ellipsoid(-oblong), 4.5 by 3 mm, cleft 1/4—1/3, lobes |
(—1.5) mm long, 0.3(—0.4) mm thick. Androecium 3 mm; androphore 0.3 by 0.5 mm,
hairs scattered, pale, less than 0.1 mm long; synandrium 2.5—2.8 by 1 mm, apex rounded,
thecae 16-20, sterile apex absent. Female flowers: pedicel 7-9 mm, bracteole caducous;
buds ovoid-oblong, + tapering towards apex, 6—8(—9) by 3-5(—6) mm, lobes 2-3 mm;
ovary ovoid-oblong, 4.5-6 by 2-3 mm, hairs scale-like, less than 0.1 mm. Fruits in
fascicles of (2—)4—7, ovoid-oblong, 3—3.5 by 1(—1.5) cm, narrowed from the middle into
a narrow beak 2—3 mm (stigma remnant), base rounded, hairs dense, (red-)brown, scale-
like, less than 0.1 mm; pericarp 1 mm thick; seeds ellipsoid-oblong, 2 cm; fruiting pedi-
cel slender, 12-15 by 1(—1.5) mm, with few conspicuous wart-like mealy lenticels to-
wards the apex.

Field-notes — Slender tree. Leaves glaucous or grey-white below. Flowers cream or
whitish. Fruit orange-brown, c. 3 cm long.

Distribution — Malesia: NW Papua New Guinea (endemic to the upper Sepik River
area, Sepik Prov.).

Habitat & Ecology — Locally common myrmecophilous tree, in primary and de-
graded forest; forest of river flats; 50-150 m altitude; fl. & fr. Mar., June, July.

Note — The feature of a papillose lower leaf surface can be a character for species
delimitation, but is possibly not valid for M. fasciculata; cf. De Wilde 1.c.

43. Myristica fatua Houtt.

Myristica fatua Houtt., Nat. Hist. Pl. 2 (3) (1774) 337; Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1864)
205, p.p., incl. var. macrocarpa; Warb., Mon. Myrist. (1897) 425, t. 11; Backer & Bakh. f., FI.
Java 1 (1964) 139: J. Sinclair, Gard. Bull. Sing. 23 (1968) 268, p.p. (see note 1); W.J. de Wilde,
Blumea 35 (1990) 242, f. 1: 3; 40 (1995) 279; 42 (1997) 166. — Lectotype: Clusius, Tab. 14 (see
De Wilde 1990: 243).

For more references and synonyms see the subspecies.

De Wilde — Myristicaceae (Myristica) 461

Tree 10-25 m. Twigs 3—4(—6) mm diameter, terete or faintly 2-ridged (Halmahera,
Ternate, Bacan), glabrescent, hairs (0.1-)0.2—0.4 mm; older twigs coarsely striate or
longitudinally cracked; lenticels copious. Leaves membranous or chartaceous, elliptic
(-oblong) to (oblong-)lanceolate, 15—40(—S0) by 4.5-15(—18) cm, base cuneate to slightly
rounded, apex acute(-acuminate); upper surface olivaceous-brown, lower surface felty
with pale brown or greyish interwoven scale-like hairs 0.2—0.3 mm, in juvenile leaves
occasionally some emergent hairs to 0.4 mm (subsp. fatua), or scattered flat-lying long
hairs (subsp. affinis); papillae not obvious; dots absent; midrib slightly raised above,
lateral nerves 17-27(—30) per side, at 50—70° to the midrib, + sunken above, lines of
interarching distinct or not, venation indistinct; petiole 15-25 by 2.54 mm; leaf bud
12-20 by 3-4 mm, hairs 0.1—0.6 mm. Inflorescences between the leaves or below, of the
Knema-type; in male: sessile or to 15 mm pedunculate, wart- or worm-like, scar-cov-
ered, simple or forked brachyblasts, rarely 3-furcate (type of M. plumeriifolia), 5 to 15
(-20) mm long, with subumbel(s) of 5—10 flowers, buds of variable sizes, pedicels
+ curved upward: bracts early caducous; female inflorescences shorter: flowers with
rust-coloured hairs 0.3—0.6 mm. Male flowers: pedicel 5-8 mm, bracteole subpersistent
or caducous, 2—3 mm; buds ellipsoid or obovoid, (4—)4.5—6 by (3—)3.5—4 mm, cleft
about 1/2. lobes 0.3—0.8 mm thick. Androecium + cylindrical, 2.54.5 mm; androphore
1-1.5 by 0.6-0.7 mm, subglabrous (hairs scattered, 0.3 mm) to densely pubescent; synan-
drium 2-3 by 0.8-1.2 mm, thecae 12-20, sterile apex absent or to 0.5 mm. Female
flowers: pedicel 4 mm; buds ovoid, 4-5 by 3.5—4 mm, cleft 1/2; ovary broadly ovoid,
2.5 mm long, brown pubescent. Infructescence short. Fruits solitary or 2 together, broadly
ellipsoid or ellipsoid(-oblong), 4-7 by 3—-5.5 cm, hairs dense, velvety, rust-coloured,
0.5-1 mm; pericarp (3—)4-12 mm thick; seeds ellipsoid to oblong, 3—3.5 cm; fruiting
pedicel slender or stout, 3—10 mm long.

Distribution — Malesia: SE Borneo (occasional), Sulawesi, S Philippines, Moluccas,
and W New Guinea. There are two subspecies which resemble each other in general ap-
pearance.

Notes — 1. Sinclair accepted M. fatua in a very broad sense, including the Indian M.
magnifica as one of the many varieties, and several varieties in New Guinea. Here M.
fatua is accepted to be confined to the Moluccas, Sulawesi, and the Philippines. It is
allied to M. mindanaensis and M. sangowoensis, both with different distributional areas.

2. Myristica fatua is taxonomically close to M. inutilis and M. subcordata ( both com-
mon in New Guinea), the latter with two subspecies. Myristica inutilis differs in smaller
and narrower male buds, only shallowly cleft into lobes. Myristica subcordata var. mor-
indiifolia is similar to M. fatua, but has a narrow male bud with early falling bracteole:
the male flowers of M. subcordata var. subcordata are as yet unknown.

KEY TO THE SUBSPECIES

la. Lower leaf surface with intricate scale-like hairs 0.2-0.3 mm long, sometimes mixed
with emergent dendroid hairs 0.4 mm long. Thecae 12—16, sterile apex 0.3—0.5 mm,
or absent, androphore long or short, 1-1.5 mm, nearly glabrous or slightly pubes-
cent. Fruits with persistent indumentum; pericarp 3-5 mm thick. — Philippines
(Mindanao), Moluccas; SE Kalimantan (rare), West New Guinea (rare).........-
Meri Bl Baas Sra ft clare sauleiten: Utive O22AL eae Ee a. subsp. fatua

462 Flora Malesiana, Ser. I, Vol. 14 (2000)

b. Lower leaf surface with dense hairs 0.2—0.4 mm, and with late caducous, scattered,
flat-lying glossy hairs | mm long. Thecae c. 20, androecium truncate, sterile apex
(almost) absent, androphore short, to 0.5 mm, largely pubescent. Fruits late
glabrescent; pericarp 7-12 mm thick. — Sulawesi ............ b. subsp. affinis

a. subsp. fatua

Myristica fatua Houtt. var. fatua; J. Sinclair, Gard. Bull. Sing. 23 (1968) 269, f. 30.

Myristica tomentosa Thunb., Act. Holm. Kongl. Vet. Acad. Nya Handl. 3 (1782) 49, t. 1, f. 2, 5, 6.
— Type: probably Smith 300, from Banda (see Sinclair, l.c.: 272).

Myristica spadicea Blume, Bijdr. (1826) 277. — Type: Reinwardt s.n., culta.

Myristica macrophylla Roxb., Fl. Ind. 3 (1832) 846 (non Spruce, nec A. Gray, nec Zippelius). —
Syntypes: Smith s.n., 2640, Banda.

Myristica mascula Reinw. ex de Vriese, PI. Ind. Bat. Orient. (Pl. Reinw.) (1857) 93; nom. nud. —
Vouchers: Reinwardt s.n. (67), 1371 (L), culta.

Myristica fatua Houtt. var. macrocarpa Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1864) 205. — Type:
Teijsmann 5148, Amboina.

Myristica plumeriifolia Elmer, Leafl. Philipp. Bot. 3 (1911) 1063; Merr., Enum. Philipp. Flow. Pl. 2
(1923) 180. — Type: Elmer 11063, Mindanao.

Twigs early glabrescent. Leaf blades (elliptic-)oblong, often broadest at or above the
middle, apex mostly acute-acuminate; midrib 1-3 mm wide, lateral nerves 17-25 per
side. Indumentum of lower leaf surface dense, hairs interwoven, scale-like, 0.2—0.3 mm,
sometimes mixed with dendroid erect emergent hairs 0.4 mm long. /nflorescences to 10
mm pedunculate. Male perianth lobes rather membranous, 0.3—0.5 mm thick; thecae
12-16, sterile apex (sub)acute, 0.3—0.5 mm long, androphore slender, 1—1.5 mm, nearly
glabrous or with some short hairs towards the base. Fruits ellipsoid, 4—6.5 cm, velvety-
tomentose, hairs 0.5—1 mm long; pericarp (3—)4—5 mm thick; seeds ellipsoid. — Fig.
61d, 64c, d.

Field-notes — Bole straight, with low buttresses and prop-roots. Living bark 0.3-1
mm, rather smooth, fissured, not peeling off; inner bark 3-8 mm thick, yellowish tinged
pink, beafy red, or brown, turning chocolate on exposure; sapwood pale yellow, reddish
yellow, whitish or brown, gradually passing into slightly darker, brown(-yellow) heart-
wood; sap little, watery, pale reddish to red-brown. Fruits brown; seeds blackish brown
or chocolate.

Distribution — Malesia: SE Kalimantan, S Philippines (Mindanao), Moluccas, W New
Guinea (see note 2).

Habitat & Ecology — Hillside forest, tall forest on black porous volcanic soil or rock,
with boulders and little undergrowth; red clay soil in rugged terrain; 0-500 m altitude;
fl. throughout the year; fr. mainly Aug.—Oct.

Notes — 1. Specimens from Halmahera (Pleyte 175, de Vogel 3301, 3431, 3495, 3497,
and possibly bb 2320] from Bacan) differ from M. fatua subsp. fatua in the indumentum
of their flowers with relatively rough and long hairs, in the 2-angled twigs, in smaller
3.5—4 cm long fruits, with short 2-3 mm long fruiting pedicels, and in a short androphore
that is short-hairy (more slender and almost glabrous in the plants from the rest of the
Moluccas).

2. Two collections from western New Guinea (Papua Barat) key out to M. fatua subsp.
fatua: Docters van Leeuwen 9856 with immature male flowers, and Kloss s.n. with fruits,

De Wilde — Myristicaceae (Myristica) 463

4(—4.5) cm, fruiting pedicels 10 mm long, and short hairs 0.2—0.3 mm only. Both speci-
mens may represent a separate taxon within M. fatua. The Kloss specimen approaches
M. ingrata subsp. velata, which differs in ant-swellings in the twigs and in smaller fruits.

3. The only collection from Borneo (Kuswata 886, male fl., SE Kalimantan, from
limestone near Jaro) is far outside the main distributional area of subsp. fatua. It has a
stout habit, with + coriaceous leaves, broadly rounded at the base, and its flowers agree
with those from the Moluccas. Study of more specimens may reveal that it represents a
separate taxon.

4. Sinclair (1.c.) included in var. fatua both M. mindanaensis and M. nivea, but these
are here restored as separate species, e.g. distinguishable by their less hairy or almost
glabrous lower leaf blades and smaller fruits.

b. subsp. affinis (Warb.) W. J. de Wilde

Myristica fatua Houtt subsp. affinis (Warb.) W.J. de Wilde, Blumea 42 (1997) 166. — Myristica
affinis Warb., Mon. Myrist. (1897) 422; Burkill, Dict. Econ. Prod. Malay Penins. (1935) 1523.
— Myristica fatua Houtt. var. affinis (Warb.) J. Sinclair, Gard. Bull. Sing. 23 (1968) 275, f. 31.
— Syntypes: de Vriese s.n. (L, lecto, here designated); Beccari s.n., 1876, 7668, a; Treub s.n.,
Bogor, culta.

Myristica fatua auct. non Houtt.: Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1864) 205 (excl. var. macro-
carpa), Koord., Meded. Lands Pl. Tuin 19 (1898) 570.

Myristica celebica Gand., Bull. Soc. Bot. France 66 (1919) 225, in clavi [non Myristica celebica
Mig. (1865)]. — Type: Teijsmann s.n., Sulawesi.

Twigs late glabrescent. Leaf blades broadly elliptic to nearly lanceolate, frequently
broadest below the middle, generally tapering to above; midrib flat, broad, to 5 mm wide
towards the base, lateral nerves 20-30 per side. Indumentum of lower leaf surface in
addition to dense layer of scale-like interwoven hairs, usually also with scattered flat-
lying much longer glossy hairs, to | mm long, the latter fallen off in old leaves. Inflores-
cences subsessile or to 15 mm pedunculate. Male perianth lobes coriaceous, 0.5—0.8 mm
thick. Androecium truncate at apex; androphore short, 0.2-0.5 mm long, broad, disc-
like, densely brown pubescent towards the base; thecae c. 20, sterile apex absent or less
than 0.1 mm. Fruits broadly ellipsoid, 5—7 by 3.5—5.5 cm, hairs velvety to coarse, 0.5-1
mm long, late glabrescent (usually some hairs remaining towards insertion of the stalk);
pericarp 7-12 mm thick; seeds oblong.

Field-notes — Straight bole, many prop-roots. Outer bark 0.3 mm thick, brown or
greyish, fissured, peeling off strongly or not; inner bark 3-7 mm thick, reddish on sec-
tion, exuding much reddish sap; sapwood pale reddish yellow, gradually passing into the
slightly darker heartwood. Mature fruits brown.

Distribution — Malesia: Sulawesi.

Habitat & Ecology — Primary lowland and lower montane forest, swamp forest, pri-
mary tall dense forest with little undergrowth; deep sand-clay soil, bedrock metamor-
phic schists; 0-700 m altitude; fl. & fr. throughout the year.

44. Myristica filipes W.J. de Wilde

Myristica filipes W.J. de Wilde, Blumea 40 (1995) 280. — Type: LAE 52061 (Foreman), Papua
New Guinea, Morobe Prov.

464 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 2.5—9 m tall. Twigs 1—-1.5 mm diameter, faintly angular, very early glabrescent,
hairs less than 0.1 mm; older bark neither cracking nor flaking; lenticels inconspicuous
or absent. Leaves chartaceous, (elliptic-)oblong, 6—14(—18) by 2-4.5(—7.5) cm, base short
to long attenuate, apex acute-acuminate; upper surface dark brown, lower surface me-
dium brown, very early glabrescent, hairs less than 0.1 mm; not papillose; dots absent;
midrib (flat to) raised above, lateral nerves 11-16 per side, at 70(—80)° to the midrib,
sunken above, lateral nerves on lower surface comparatively strong, raised, yellowish
brown and contrasting, lines of interarching and venation indistinct; petiole 6—10(—16)
by 1—1.5(—2.5) mm; leaf bud 4—6 by | mm, hairs less than 0.1 mm. /nflorescences be-
tween the leaves and below, sessile, of the Knema-type; in male: a simple or sometimes
2- (or 3-)forked, scar-covered slender brachyblast to 6 mm long, glabrescent, with 1-4
flowers in a cluster, buds of slightly variable sizes; bracts minute, caducous; female in-
florescence (from infructescence): a small brachyblast, | mm long, single-flowered; flow-
ers glabrous, except for a few scattered scale-like hairs less than 0.1 mm. Male flowers:
pedicel slender, 9-11 by 0.4-0.5 mm, bracteole caducous, scar 4—4.5 mm below the
apex; buds elongate, + tubiform, somewhat narrowed below the middle, 7 by 1.5 mm,
apex + blunt (to subacute), somewhat angled, base short-attenuate, cleft c. 1/4, lobes
membranous, 2 mm long, 0.2 mm thick, at anthesis suberect, inside at base (where con-
tiguous with the androphore) thickened with irregular warty appendages. Androecium
6—6.5 mm; androphore 3.5 by 0.4 mm, glabrous; synandrium 2.8 by 0.5—0.6 mm, thecae
c. 15, sterile apex subacute, 0.2 mm. Female flowers (from remnant under the fruit,
Conn c.s. 281): perianth tubiform, 7 by (1.5—)2 mm, lobes 1.8 mm long, slightly curved
outwards. Fruits solitary, + oblong, 3.5—4.5 by 1.2-1.5 cm, apex acute (with minute
apiculum), the basal 1.5—2 cm narrowed into a slender pseudostalk, hairs dense, greyish
brown, less than 0.1 mm; pericarp | mm thick; seeds ellipsoid, 2 cm long; fruiting pedi-
cel slender, 25-30 mm long, 0.5 mm thick, bracteole scar + median.

Field-notes — Low bushy or straight tree, bole dbh to 15 cm. Bark light grey or brown,
more or less smooth; inner bark reddish or orange, exudate watery, reddish; wood white,
soft. Leaves silvery green below. Flowers yellow. Fruits yellow, orange, or rust-colour-
ed; seeds black.

Distribution — Malesia: eastern Papua New Guinea (Morobe Prov., a local endemic,
several collections from Buso River area).

Habitat & Ecology — Coastal swamp forest (with Pandanus), forest on low ridges;
also degraded forest; 0-15 m altitude; fl. Aug.; fr. May—Aug.

Note — Myristica filipes belongs with M. flosculosa and M. pumila to a coherent
group which is characterized by tubiform male flowers. The lower half of the perianth
inside (where contiguous with the androphore) is conspicuously thickened by irregular
warty ridges. As in M. pumila the fruiting pedicel of M. filipes is distinctively long and
slender.

45. Myristica firmipes J. Sinclair

Myristica firmipes J. Sinclair, Gard. Bull. Sing. 23 (1968) 355, f. 56; W.J. de Wilde, Blumea 40
(1995) 281. — Type: Brass 7181, Papua New Guinea, Western Prov.

Tree 4—30 m. Twigs 1-2 mm diameter, early glabrescent, hairs less than 0.1 mm,
older twigs striate, thinly flaking or not; lenticels distinct. Leaves chartaceous (or thinly

De Wilde — Myristicaceae (Myristica) 465

coriaceous), (elliptic-)oblong, 6-14 by 1.5—4.5 cm, base attenuate, apex acute(-acumi-
nate); upper surface olivaceous-brown, lower surface with indumentum subpersistent or
late glabrescent, hairs pale silvery brown, scale-like, less than 0.1 mm; papillae and dots
absent; midrib somewhat raised above, lateral nerves 10—15 per side, at 60(—70)° to the
midrib, flat or sunken above, lines of interarching and venation indistinct on both sur-
faces; petiole 8-14 by 1-2 mm; leaf bud 6-11 by 1.5—2 mm, hairs appressed, less than
0.1 mm. Inflorescences (known only from infructescences) of the Knema-type. Male
and female flowers not seen. Infructescences with a single fruit below the leaves; pedun-
cle absent. Fruits elliptic-oblong or + pear-shaped, 6-7 by 3—4 cm, apex rounded, with
short distinctly oblique apiculum, basal part somewhat contracted into a short thick
pseudostalk or not, hairs mealy, dull brown, 0.1 mm or less; pericarp with a marked
ridge, + woody, 10-15 mm thick; seeds (+ immature) ellipsoid, (2-)2.5 cm; fruiting
pedicel thick, 4-10 by 5—9 mm, fissured and somewhat lenticellate.

Field-notes — Small tree (NGF 41064, at c. 1300 m alt.) or tall forest canopy tree
(two records). Bole straight, spur-buttressed; crown rounded, compact. Bark brown; in-
ner bark dull pinkish brown; exudate little, reddish; wood white. Leaves greyish bronze
below. Fruits brown or brownish (green). Seeds dark brown.

Distribution — Malesia: W Papua New Guinea (possibly endemic to W Sepik, West-
ern Highlands, and Western Prov.).

Habitat & Ecology — Primary and degraded forest, ridge forest; copper-rich soil
(once); 100—1300 m altitude; fr. June, July, Nov.

Notes — 1. The specimens BW 10655 & 10698, from Wandammen Peninsula, Papua
Barat, consist of leafy twigs with short, thickened infructescences, but fruits are lacking;
the leaves, however, are very distinctly reticulate above, but the specimens still may be-
long to M. firmipes.

2. The fruits of Brass 7181 are, judged from the position of the line-shaped ridge,
situated transversely in relation to the twig; in two other specimens (NGF 41064, 42527)
the fruits are situated in a median position, turned 90°, compared to those in Brass 7181.
The leaves in the latter collection are comparatively small and I do not know whether
these differences have taxonomic consequences.

3. Sinclair described and figured the fruiting stalk as being 1-1.5 cm long, but in fact
he measured this including a part of the branches taken with the fruit; actually the fruit-
ing pedicel in the type is only some 5 mm long.

46. Myristica fissiflora W.J. de Wilde

Myristica fissiflora W.J. de Wilde, Blumea 40 (1995) 281; 43 (1998) 173. — Type: NGF 28041
(Henty), Papua New Guinea, Madang Dist.

Shrub or tree 3-10 m. Twigs subterete or somewhat flattened, (4—)5—10 mm diameter,
with distinct, raised lines or narrow wings |(—2) mm high, joining the petioles, glabrescent,
hairs 0.1—0.2 mm, hollow, and with ant-swellings similar to those in M. subalulata;
older twigs sometimes with conspicuous lenticels. Leaves membranous or chartaceous,
obovate-elliptic or oblong, 22-35 by 7-16 cm, base short-cuneate, rounded, or shallowly
cordate, apex (acute-)acuminate; upper surface olivaceous, lower surface greyish (green),
glabrous or glabrescent, hairs scale-like, 0.1 mm or less, lower surface either + papillose
(subsp. fissiflora) or not (subsp. kostermansii); dots absent; midrib flat or slightly raised

466 Flora Malesiana, Ser. I, Vol. 14 (2000)

above, lateral nerves 18-23 per side, at 45—80° to the midrib, flat or sunken above, lines
of interarching and venation distinct; petiole 15-25 by 2-4 mm; leaf bud 15 by 4 mm,
hairs 0.1 mm. Inflorescences between the leaves, of the Knema-type, a (sub)sessile scar-
covered brachyblast, minutely pubescent; in male: to 10 mm long, 4-6 mm wide, subumbel
of 2-8 flowers, buds of various sizes; bracts minute, caducous; flowers with sparse or
dense dark brown hairs 0.1—0.2 mm. Male flowers: pedicel distinct, bracteole large or
small, caducous; buds + broadly fusiform or obovate-oblong, 8—12 mm long, cleft to
over 1/2: see further under the subspecies. Female flowers and fruits not known with
certainty, see note.

Distribution — Malesia: northern New Guinea (with two subspecies).

Note — Myristica fissiflora is mainly characterized by its deeply lobed male flowers,
and vegetatively only by a strong reticulate venation of the leaves. Specimens with fe-
male flowers or fruits may have been treated under the vegetatively similar M. subalulata.
The following three fruiting collections possibly belong to M. fissiflora: Warburg 20704,
Lauterbach 2865, and NGF 46688 (Katik), all from about the distributional area of M.
fissiflora. If these specimens indeed belong to M. fissiflora then the description of the
fruits should be: small, 1.5 by 1 cm, with a4 mm long fruiting pedicel, much resembling
small-fruited M. subalulata.

KEY TO THE SUBSPECIES

la. Lower leaf surface papillose (or not, see note 1). Male buds + fusiform, 10-12 mm
long; synandrium at apex narrowed into distinct acute sterile apex (0.5—)1 mm long.
Bracteole (5—-)6-8 mm long. — Papua New Guinea (Madang Prov. and possibly
Morobe Prove es PPP PF Ae ee Cen mene meri: a. subsp. fissiflora

b. Lower leaf surface not or indistinctly papillose. Male buds + oboyoid-oblong, 8-9
mm long, apex + blunt; synandrium at apex + rounded, sterile apex absent or to 0.2
mm long. Bracteole 2—3 mm long. — NE Papua Barat (vicinity of Jayapura) ......
ES PP AN VELA! SUL SDE 5) Oe RENEE TES LAS i ae eae eee ne Oi b. subsp. kostermansii

a. subsp. fissiflora

Tree 6—8 m. Leaves (thinly) chartaceous, 25-35 by 7-16 cm, lower surface papillose.
Male flowers: pedicel 7-18 by 1—-1.5 mm, bracteole (ovate-)oblong, (5—)7-8 mm long;
buds leathery, ellipsoid-oblong, fusiform, 10—12 by 3.5—5(—6) mm, base and apex narrow-
ed, apex acute, the apical part + triangular, inner surface of lower part finely papillose,
cleft for c. 2/3, lobes 6-8 by 0.4—0.5 mm. Androecium stout, 8—-9.5 mm; androphore
2.5-3(—4) by 1(-1.5) mm, + completely pubescent, hairs bright- or yellow-brown, 0.1-
0.2 mm; synandrium + fusiform, faintly angular, 5.5—6 by 1.5(-2) mm, thecae 16-25,
sterile apex + acute, | mm.

Field-notes — Small tree. Stem hollow, full of ants. Bark brown; wood white. Leaves
light green above, glaucous below. Flowers orange, (dull) brownish cream, or yellow,
pinkish at lobe apices. Fruit brownish orange.

Distribution — Malesia: northern Papua New Guinea (Madang Proy.; see note IBY

Habitat & Ecology — Primary and old degraded (regrowth) forest, advanced regrowth
in Pometia forest, 100—400(—1000?; see note 2) m altitude; fl. Mar., May, Aug.

De Wilde — Myristicaceae (Myristica) 467

Notes — 1. Specimens of subsp. fissiflora have a papillose lower leaf surface. Takeuchi
& Soejarto 6987 (immature male fl.) and Hoogland 8944 (immature female fl. & fr; but
fruits not seen), both from Morobe Province, have an indistinct papillation, but are re-
lated to subsp. fissiflora, sharing the large bracteole. The leaves in Takeuchi & Soejarto
6987 are almost glabrous below, Hoogland 8944 has rather persistent pale cinnamon
minute scale-like hairs, and (female) inflorescences with an up 7 mm long peduncle. Fe-
male flowers: pedicel 7 mm long, bracteole hooded, 8(—9) mm long, apparently caducous
at anthesis: buds 7 mm long, cleft c. 1/3. The field-note mentions mature fruits 47 by 34
mm, but such fruits have not been observed. Pending more material the status of the
specimens mentioned above is uncertain.

NGF 49251 (Henty), from the type locality and vegetatively similar to the type, has
female inflorescences: flowers (immature) in subumbels of c. 5, pedicel and bracteole
3 mm long. Mature female flowers probably do not differ significantly from those of the
related M. subalulata.

2. Schlechter 17749 (in L, male, c. 1000 m) belongs to subsp. fissiflora. However, the
duplicate in K, also male, with the same herbarium label, is different and belongs to M.
subalulata.

b. subsp. kostermansii W.J. de Wilde
Myristica fissiflora W.J. de Wilde subsp. kostermansii W. J. de Wilde, Blumea 40 (1995) 283; 43

(1998) 173. — Myristica subalulata auct. non Migq.: J. Sinclair, Gard. Bull. Sing. 23 (1968) 385,
f. 64D. — Type: Kostermans & Soegeng 392, NE Irian Jaya, vicinity of Jayapura.

Shrub or treelet 3—10 m. Leaves membranous or subchartaceous, lower surface slightly
papillose or not. Male flowers: pedicel slender, 8-14 by 0.5(-1) mm, bracteole 2-3 mm
long; buds membranous, + obovate-oblong, narrowed towards base, 8-9 by 3.5—4(-4.5)
mm, apical part faintly triangular, apex bluntish, cleft for over 1/2, lobes 5 mm long, 0.3
mm thick. Androecium slender, 6.5—8 mm; androphore 2.5—4 by 1 mm, (almost) com-
pletely pubescent, hairs rather pale, 0.1(—0.2) mm or less; synandrium faintly triangular,
4-5 by 1-1.5 mm, the apex bluntish, thecae 14—20(—24), sterile apex absent or only 0.2
mm. Female flowers and fruit not seen.

Field-notes — Bark smooth. Flowers yellowish brown, inside of lobes white, of co-
rolla tube pinkish.

Distribution — Malesia: NE Papua Barat (vicinity of Jayapura).

Habitat & Ecology — Forest, in valley; 50—60 m altitude; fl. July, Aug.

47. Myristica fissurata W.J. de Wilde

Myristica fissurata W.J. de Wilde, Blumea 35 (1990) 243; 42 (1997) 166. — Type: de Vogel 3836,
N Moluccas.

Tree 25-50 m. Twigs 4—7(-10) mm diameter, early glabrescent, hairs 0.2 mm; older
twigs brown or blackish, coarsely longitudinally fissured or cracked, tending to flake;
lenticels absent. Leaves thinly coriaceous, oblong-lanceolate, 25—45 by 8-12 cm, base
narrowly rounded to cuneate, apex short acute(-acuminate); upper surface olivaceous,
sometimes red-brown tinged, lower surface greyish olivaceous, almost glabrous, hairs
few, scattered, scale-like, pale brown, 0.1—0.2 mm; distinctly papillose (lens!); dots ab-

468 Flora Malesiana, Ser. I, Vol. 14 (2000)

sent; midrib above slightly raised, lateral nerves 25—30 per side, at 60—80° to the midrib,
sunken above, lines of interarching and (coarsely reticulate) venation indistinct; petiole
20-35 by 3-5.5 mm; leaf bud 15-25 by 3-5 mm, hairs 0.2—0.5(-1) mm. Inflorescences
between the leaves, with shaggy woolly hairs, | mm; in male: peduncle somewhat flat-
tened, 15—25 mm long (as in panicle-like inflorescence), ending in a (Knema-like) scar-
covered brachyblast 3-6 mm long, with a subumbel of 2—5 flowers, buds strongly
varying in size; bracts minute, caducous; female (from infructescences): 5 mm long,
few-flowered: flowers with dense rusty hairs 0.5-1 mm. Male flowers: pedicel 6 mm
long, bracteole ovate-elliptic, c. 5 mm, subpersistent; buds hard-carnose, obovoid-ellip-
soid, 7 by 4.5(—5) mm, apex broadly rounded, cleft nearly halfway, lobes 0.8 mm thick.
Androecium 4—4.3 mm: androphore 0.8—1 by | mm, with hairs 0.5 mm; synandrium
faintly triangular, 3 by 2 mm, thecae c. 20, sterile apex + flat and somewhat hollowed in
the centre, with shallowly lobulate edge (the lobules corresponding with the anthers).
Female flowers not seen. Fruits solitary on short infructescences 0.5—1 cm long, broadly
ellipsoid, 6.5-7 by 5 cm, hairs + persistent, dense, scurfy, 0.1—0.2 mm, in immature
fruits somewhat longer, 0.5(—1) mm (breaking off at maturity?); pericarp 10-15 mm
thick; seeds broadly ellipsoid, 3.5 cm; fruiting pedicel thick, 2-3 mm long, 5 mm diam-
eter. — Fig. 64e.

Field-notes — Solitary tree, emergent or not, with buttresses only or with prop-roots
and buttresses, up to 1.5 m high, to 2 m out, 3-5 cm thick; clear bole to 30 m, straight,
dbh 50 cm, with hollow core or not; outer bark 1-3 mm thick, (grey-)brown, very fissured,
peeling off slightly or not; inner bark 6 mm thick, on section reddish, with little or much
watery brownish red exudate; sapwood yellowish tinged red or pale reddish cream, gradu-
ally passing into slightly darker heartwood. Flowers brown, inside pale yellow, anthers
brownish yellow. Fruits yellow, with brown indumentum.

Distribution — Malesia: N Moluccas (Bacan: Mt Tuara, near Amasing Kali).

Habitat & Ecology — Disturbed primary tall forest, c. 50 m high, with little under-
growth. Sloping dry hill ridge or hillsides, with little humus, soil loamy, with stones;
100—200 m altitude; fl & fr. Nov.

Note — Myristica fissurata seems closest related to M. philippensis, which differs in
having up to 50 mm long pedunculate male inflorescences lacking brachyblasts, a less
fleshy male perianth, a faintly triangular synandrium, and vegetatively in lacking the
cracked bark of the twigs.

48. Myristica flavovirens W. J. de Wilde

Myristica flavovirens W.J. de Wilde, Blumea 40 (1995) 283. — Type: BW 10337 (Versteegh), West
New Guinea, Bird’s Head.

Tree 12 m. Twigs 2-3 mm diameter, brown-black, glabrescent, hairs 0.1(—0.2) mm;
older twigs coarsely striate and somewhat cracking, not flaking; lenticels absent. Leaves
coriaceous, oblong-lanceolate, 9-11 by 2-3 cm, base short-attenuate or narrowly rounded,
apex blunt or narrowly rounded; upper surface dark olivaceous-green, lower surface
orange-brown, early glabrescent, hairs scattered, light brown, scale-like, less than 0.1
mm; densely papillose (lens!); dots and dashes present; midrib slightly raised above,
lateral nerves 20-22 per side, at 70—80° to the midrib, sunken above, lines of interarching
and venation faint; petiole 10-15 by 1.5-2 mm; leaf bud 10 by 1.5 mm, hairs dense,

De Wilde — Myristicaceae (Myristica) 469

golden rusty-red, appressed, 0.1(—0.3) mm. Inflorescences between the leaves and just
below, of the Knema-type, simple or bifid scar-covered brachyblasts to 5 mm long, ses-
sile or to 2 mm pedunculate; bracts small, caducous; in male: flowers in clusters of
(5-)10-15, buds of variable sizes; female inflorescences and flowers not seen. Male
flowers: with dense rusty or dark brown hairs, 0.1—0.2 mm, pedicel 3—4 by 1(—1.5) mm,
bracteole broadly rounded, | mm, persistent; buds + firm, (ellipsoid-)oblong, 5 \byr2:5
mm, apex and base rounded, cleft c. 1/3, lobes 1.5(—2) mm long, 0.3 mm thick;. An-
droecium slender, 3.5 mm; androphore 1.5 by 0.5 mm, glabrous except for a few hairs
less than 0.1 mm at base; synandrium 2 by 0.8 mm, thecae (12—)14—16, sterile apex
+ blunt, 0.2 mm. Fruits not seen.

Field-notes — Scarce; young tree c. 12 m high. Flowers yellowish.

Distribution — Malesia: W New Guinea (Bird’s Head, Kebar Valley); known only
from the type.

Habitat & Ecology — Primary forest on clay soil; 1200 m altitude; fl. Nov.

Notes — 1. Myristica flavovirescens is related to M. arfakensis, and M. conspersa,
and M. scripta, species with similarly dotted and dashed leaves and with a compara-
tively long androphore, not much shorter than the synandrium. Myristica arfakensis is
distinguishable by stalked inflorescences and smaller flowers; M. conspersa has much
smaller (male) flowers; M. scripta is rather variable in its flower size and its leaves are
generally brown on drying, with an acute apex, but the species occurs in New Guinea
outside Bird’s Head.

2. The twigs of M. flavovirens are conspicuously blackish; the leaves are coriaceous,
drying a strange green colour above, (orange-)yellow below, and are distinctly densely
papillose (lens!). The texture of its leaves and its drying colour are suggestive of a spe-
cies from ultrabasic soil.

49. Myristica flosculosa J. Sinclair

Myristica flosculosa J. Sinclair, Gard. Bull. Sing. 23 (1968) 359, f. 58 (excl. fruiting specimens,
excl. f. 59): Foreman, Handb. Fl. Papua New Guinea 1 (1978) 197, p.p.; W.J. de Wilde, Blumea
AO (1995) 284. — Type: Hoogland 3717, Papua New Guinea.

Tree 15-20 m. Twigs bluntly angled, 2-3 mm diameter, (bright) brown or reddish
brown, early glabrescent, hairs scale-like, 0.1 mm long or less; older bark striate, neither
cracking nor flaking; lenticels not apparent. Leaves chartaceous or coriaceous, elliptic-
oblong to oblanceolate, base attenuate or nearly rounded, apex acute-acuminate, 8—23
by 3-8 cm; upper surface olivaceous, lower surface pale grey-brown, late glabrescent,
hairs scattered, scale-like, 0.1 mm or less; faintly papillose; dots absent; midrib flat
above, lateral nerves 13-17 per side, at 50-80° to the midrib, sunken above, lines of
interarching indistinct, venation invisible above, faint or distinct below; petiole 12-15
by 1.5-3.5 mm; leaf bud 10 by 1.5 mm, hairs 0.1 mm or less. Inflorescences between the
leaves or below, of the Knema-type; in male: a (sub)sessile, simple, scar-covered bra-
chyblast to 5 by 3 mm, glabrescent, with (1—)2—7 flowers, buds of various sizes; bracts
broad-triangular, pubescent, 0.5(—1) mm, caducous; female inflorescence a few-flow-
ered sessile wart-like brachyblast, 1-2 mm diameter; flowers with hairs 0.1 mm. Male
flowers: pedicel slender, 5—7 mm long, bracteole small, caducous, scar thickened, apical
or to 2.5 mm below; buds firm, elongate-tubular, 8-8.5(—9) by 2 mm, apex narrowly

470 Flora Malesiana, Ser. I, Vol. 14 (2000)

rounded, base shortly attenuate, inside glabrous, smooth, but in the lower 3—-3.5 mm, the
part facing the androphore, conspicuously thickened with irregular wart-like structures,
cleft nearly 1/4, lobes 1.5—2 mm long, 0.3—0.4 mm thick, at anthesis curved outwards,
exposing the apical part of the synandrium. Androecium slender, 8-8.5 mm, androphore
slightly widened above, 3-3.5 by 0.7—1 mm, minutely pubescent in the lower 1/2-2/3,
gradually passing into the synandrium; synandrium 5—S.5 by 0.8-1.2 mm, narrowed in
the upper half, thecae 6 or 7, widely spaced (i.e. anthers 3; or 6?), sterile apex acute, 0.1-
0.4 mm. Female flowers not seen. Fruits 1 or 2 (or 3) per infructescence, fusiform or ob-
ovoid-oblong, (4—)4.5 by (1-)1.3 cm, apex subacute or narrowly rounded, base nar-
rowed into a 10 mm long pseudostalk, hairs dense, scale-like, less than 0.1 mm; pericarp
less than 1 mm thick; mature seeds not known; fruiting pedicel slender, 30-35 by 0.8
mm, bracteole scar at c. 2/3 below the apex. — Fig. 63c, 83e.

Field-notes — Medium-sized tree to 20 m. Bark pale brown or grey-brown, flaky,
shallowly longitudinally fissured, 8 mm thick; outer bark 2.5 mm; inner bark pale red-
brown; exudate colourless turning deep brown; sapwood pale brown; heartwood dark
brown or wood pinkish straw turning brown mottled. Leaves ashy green below. Flowers
yellow with fine brown dots, androphore off white, anthers pale brown.

Distribution — Malesia: eastern Papua New Guinea (Northern and Milne Bay Prov.;
known only from a few specimens).

Habitat & Ecology — Gently sloping lowland with low rain forest, at 60 m altitude,
and foothill forest at 300-350 m altitude; fl. Aug., Sept.

Notes — 1. Both Sinclair and Foreman (1.c.) included some fruiting specimens from
montane altitudes in M. flosculosa, specimens which are transferred here to different
taxa. However, the fruiting collection Giulianetti s.n. (1896; comm. W. MacGregor) in
K belongs to the present species. In M. flosculosa the fruits have characteristically a
long pseudostalk and a long and slender fruiting pedicel, suggesting a close relationship
to species like M. filipes, M. gracilipes, M. nana, and M. pumila, all species with similar
slender pedicels.

2. The basal inner surface of the perianth, facing the androphore, is conspicuously
swollen and has an irregularly warted surface. In boiled-up herbarium specimens it has
a dark, reddish brown colour, but it is possibly differently coloured when fresh.

3. Myristica flosculosa is a characteristic species with conspicuously tubiform male
flowers. Vegetatively the ashy green undersurface of the leaves is remarkable. At anthesis
the outwards curved perianth lobes expose the acute, almost acuminate apical part of the
synandrium.

50. Myristica fragrans Houtt.

Myristica fragrans Houtt., Nat. Hist. Pl. 2 (3) (1774) 333: Blume, Rumphia | (1835) 180, t. 55;
Warb., Die Muskatnuss (1897) 1-628; Mon. Myrist. (1897) 458; J. Sinclair, Gard. Bull. Sing. 16
(1958) 361, f. 29, pl. VII B; 23 (1968) 225; Flach & Tjeenk Willink in Plant Resources of
South-East Asia, A selection (1989) 192; W.J. de Wilde, Blumea 35 (1990) 244; 40 (1997) 166.
— Lectotype: J.W. Weinmann, Taalrijk Register 7 (1748) pl. 760a.

Myristica officinalis L.f., Suppl. Pl. (1781) 265.

Myristica moschata Thunb., Act. Holm. Kongl. Vet. Acad. Nya Handl. 3 (1782) 49, t. 1, f. 1.

Myristica aromatica Lam., Act. Paris (1788) 155, t. 5-7.

Myristica amboinensis Gand., Bull. Soc. Bot. France 66 (1919) 225, in clavi.

De Wilde — Myristicaceae (Myristica) 471

Myristica laurella Gand., Bull. Soc. Bot. France 66 (1919) 226, in clavi.

Myristica philippinensis Gand., Bull. Soc. Bot. France 66 (1919) 226, in clavi.

For more synonyms, references, and typifications see Sinclair, l.c. (1958) 363; (1968) 226, 231; for
pre-Linnean names and literature see Warburg, Mon. Myrist. (1897) 459-467.

Tree 4—10 m. Twigs 1-2 mm diameter, yellowish brown, early glabrescent, hairs pale,
0.1 mm or less; older twigs grey-brown, striate; lenticels inconspicuous. Leaves charta-
ceous or thinly coriaceous, (ovate-)elliptic to oblong, 6—13 by 3.5—6.5 cm, base attenu-
ate, apex acute-acuminate; upper surface olivaceous, lower surface pale, olivaceous or
greyish white, early glabrescent, hairs 0.1 mm; not conspicuously papillose; dots absent;
midrib raised above, lateral nerves 6—12 per side, at 60—80° to the midrib, flat or sunken
above, lines of interarching usually not distinct, venation faint or distinct below, con-
trasting in colour or not; petiole 7-12 by 0.7—1.5 mm; leaf bud 5—10 by | mm, hairs 0.1
mm or less. Inflorescences between the leaves, axillary or sometimes a few mm supra-
axillary, essentially paniculate; in male: peduncle slender, 5-20 mm (with an apical flower
or not) and with one or two opposite, slender (but thicker than the peduncle), scar-coy-
ered (scars spirally arranged and rather widely spaced) brachyblasts, to 20 mm long, up
to 5-flowered at apex, buds usually very variable in size, sometimes male inflorescences
reduced to 1—3 almost sessile long-pedicelled flowers; female inflorescences: peduncle

Fig. 73. Myristica fragrans Houtt. Male inflorescences [a & b: C. K. Ruffo 247, cult., Tanzania;
c: NGF 36217 (Zieck), cult., New Ireland / Papua New Guinea]. — Scale bar = 2 cm.

472 Flora Malesiana, Ser. I, Vol. 14 (2000)

slender, up to 5 mm, 1(—3)-flowered; bracts minute, caducous. Inflorescences and flow-
ers with widely spaced, appressed, pale brown hairs, 0.1 mm or less, partly glabrescent.
Male flowers: pedicel slender, 10-15 mm long; bracteole broadly ovate or semi-circular,
c. 2 mm, + caducous; buds ellipsoid-oblong, 7-8 by 4-5 mm, apex and base rounded,
cleft c. 1/5, lobes 0.3 mm thick, at anthesis somewhat spreading. Androecium slender or
broad, 5—6 mm; androphore 2—2.5 by 1 mm, hairs 0.1 mm or less, or nearly glabrous;
synandrium 3 by 1—1.2(—1.5) mm, thecae 14—20(—24), sterile apex subacute, 0.5 mm.
Female flowers: stouter than male flowers, pedicel 8-12 mm long, bracteole caducous,
apical or up to 2.5 mm below; buds + ovoid, apex narrow, base broadly rounded, 7-8 by
5 mm, cleft c. 1/5, lobes at anthesis curved outward; ovary ovoid, 5 by 3 mm, hairs less
than 0.1 mm. Fruits solitary, pendent, broadly pear-shaped or almost globose, 4-6 cm
long (6-9 cm diameter when fresh), early glabrescent, hairs widely spaced, dull brown,
0.1 mm; pericarp 10-15 mm thick; seeds broad-ellipsoid, 3 cm; fruiting pedicel 10-15
mm, + widened towards the fruit. — Fig. 59b, 64f, 73.

Field-notes — Slenderly conical trees, all parts aromatic; sometimes male and female
flowers on the same tree, but usually dioecious. Bark greyish black, slightly fissured
longitudinally in older trees. Flowers to 1 cm long, fragrant, creamy yellow, waxy. Fruits
drooping, 6—9 cm long and nearly as broad, almost globose, with a circumferential lon-
gitudinal groove and persistent remains of the stigma, with a pseudostalk, yellow, gla-
brous (early glabrescent); fruit stalk thickest towards the base of the fruit; pericarp yel-
low, succulent; aril strongly laciniate; seeds broadly ellipsoid-ovoid, purplish brown.

Distribution — Malesia: Banda Islands; widely cultivated in the Moluccas, and in
many suitable localities in the tropics all over the world; locally seemingly wild in aban-
doned or neglected plantations, or where intentionally planted in wild forest.

Supposedly all collections of M. fragrans, also those from the southern Moluccas
(including those from Ambon and Banda), are from cultivated or running wild speci-
mens. According to Rumphius (Herb. Amb. 2, 1741, 14, t. 4) the species was at the time
already amply cultivated in Banda.

Habitat & Ecology — Lowland forest, apparently preferably under rather strong sea-
sonal conditions.

Notes — 1. Sinclair (1968) listed M. fragrans with M. argentea, M. succedanea, and
M. impressinervia in his series Fragrantes (i.e., series Speciosae and Fragrans of War-
burg, l.c.: 378). Indeed, the first two species seem related to M. fragrans, but M. impressi-
nervia has far more remote affinities

2. Myristica fragrans is a homogeneous species, readily distinguishable from M. ar-
gentea and M. succedanea by its small, glabrous (early glabrescent) leaves with few
nerves, in combination with its typical slender inflorescences. Rumphius (1.c.) distin-
guished five varieties.

51. Myristica frugifera W.J. de Wilde

Myristica frugifera W.J. de Wilde, Blumea 42 (1997) 166. — Type: Ridsdale 1219, Philippines,
Mindoro.

Tree 10-20 m. Twigs 4-6 mm diameter, blackish brown, glabrous or very early gla-
brescent, hairs 0.1 mm; older twigs sometimes + cracking and flaking; lenticels small
and inconspicuous. Leaves chartaceous or thinly coriaceous, elliptic(-oblong) or oblong-

De Wilde — Myristicaceae (Myristica) 473

lanceolate, 18—30(—35) by 6—11(—13) cm, base (attenuate or) rounded, or cordate, apex
(sub)acute(-acuminate); upper surface olivaceous or brown, lower surface grey-brown
with + persistent indumentum or late glabrescent, hairs sparse, scale-like, 0.1 mm or
less; distinctly papillose; dots absent; midrib above flat, lateral nerves 20-30 per side, at
(45—)60—80° to the midrib, slender, + sunken above, faint, lines of interarching and
venation + indistinct; petiole 15—30 by 3-4 mm; leaf bud 12-18 by 4-5 mn, hairs 0.1—
0.2(-0.5) mm. Inflorescences pedunculate, between the leaves or below, with sparse
hairs 0.1 mm or less, glabrescent; bracts small, caducous; in male: 8-10 by 2—4 cm, pe-
duncle 40-50 mm, branches to 15 mm, central axis 20-30 mm long, with 2—4 subumbels,
the latter with 5—10 flowers; buds of variable sizes; in female (from infructescences):
1-8 cm, peduncle 3-50 mm, lower branches opposite, to 10 mm long, central axis to
20 mm long, sometimes several + crowded in condensed twigs, forming compound
infructescences with apical leaf bud. Male flowers: pedicel (6—)8 mm long, bracteole
broadly ovate, 3 mm, + caducous; buds ovoid, 6.5—5.7 by 5 mm, cleft 1/3 to (nearly)
1/2, lobes 0.5—1 mm thick. Androecium cylindrical, 4 mm long, + truncate; androphore
0.7 mm long, the lower 4/5 with dense hairs less than 0.1 mm; synandrium 3 by | mm,
thecae 12-15, sterile apex shaped irregularly, blunt, 0.3 mm long. Female flowers not
known. Fruits 1-3 per infructescence, broadly ellipsoid to ellipsoid-oblong, 5—7.2 by 3—
4.5 cm, hairs light to dark brown, scurfy, 0.1 mm, partially glabrescent; pericarp 5-8
mm thick; seeds (ellipsoid-)oblong, 4.5—5 cm; fruiting pedicel 3-6 mm, coarsely fissured,
scar of bracteole (sub)apical.

Field-notes — Leaves above medium green, slightly glossy with paler midrib and
veins, below pale green or glaucous with yellowish green midrib and veins. Not yet ripe
fruits glabrous, ripe fruits brown.

Distribution — Malesia: Philippines (Luzon, Leyte, Mindoro).

Habitat & Ecology — Understorey tree of primary and disturbed lowland forest;
0—500 m altitude; fr. Feb., June.

Notes — 1. The material of this species is slightly heterogeneous, according to the
islands of provenance. Sinclair & Edafio 9595 (Luzon) has glabrous immature fruits,
olivaceous leaves, drying grey below, with 25-30 pairs of lateral nerves, and links up
very well with FB 12783 (Leyte) also with immature fruits. The three collections from
Mindoro, Ridsdale 803, 1175 and 1219 (type), all in fruit, are almost identical and differ
from the Luzon and Leyte specimens in the twigs rather brown on drying instead of
blackish brown, and bright brown subcoriaceous leaves with fewer (20-25) pairs of
nerves. The infructescences are either subsessile (e.g. Ridsdale 803, in part) or up to
5 cm pedunculate (Ridsdale 1175).

2. Myristica frugifera seems related to M. extensa (Borneo), M. maxima (West Male-
sia), and M. philippensis (Philippines). Myristica extensa has an indistinct papillation
on the lower leaf surface. Myristica maxima is quite distinguishable by its generally
denser indumentum on the lower leaf surface and a blackish drying colour of the leaves.
Myristica philippensis differs in its roughly pubescent leaf bud, hairs 1(—2) mm.

52. Myristica fugax W.J. de Wilde

Myristica fugax W.J. de Wilde, Blumea 40 (1995) 284. — Type: Carr 15501, Papua New Guinea,
Central Prov.

474 Flora Malesiana, Ser. I, Vol. 14 (2000)

Treelet 4—5 m tall. Twigs (1-)1.5—2 mm diameter, glabrescent, hairs scale-like, 0.1
mm or less; older twigs not cracking nor flaking; lenticels not apparent. Leaves membra-
nous, (elliptic-)oblong, 5-12 by 2-3.5(—4.5) cm, base broadly rounded or attenuate, apex
to 1.5 cm acute-acuminate; upper surface brown (greyish), lower surface pale, glabrous
or with a few scattered scale-like hairs less than 0.1 mm; not papillose; dots absent;
midrib raised above: lateral nerves 6—12(—14) per side, at 50—70° to the midrib, flat or
+ sunken above, indistinct, on lower surface brown, + contrasting, lines of interarching
and venation indistinct; petiole 7-12 by 0.5—1(—2) mm; leaf bud 7-12 by | mm, hairs 0.1
mm or less. Inflorescences pedunculate, between the leaves, axillary or to 3 mm supra-
axillary, glabrescent, indumentum minute, grey-brown, bracts subtriangular, less than
1 mm long, caducous; in male: peduncle (1—)2-5 by 0.5-1 mm, ending in a to 1.5 cm
long scar-covered brachyblast with a subumbel of 2—5 flowers, buds of various sizes;
female inflorescences with | or 2 flowers, without or with peduncle to 5 mm long; flow-
ers thin with hairs less than 0.1 mm. Male flowers: pedicel slender, 3.5—6 by 0.4—0.5
mm, bracteole minute, caducous, scar apical or 1-2 mm below; perianth inside in the
lower half with longitudinal irregular furrows and rows of minute wart-like excrescences,
buds ellipsoid-oblong, 4.5—6.5 by 1.2-2 mm, apex + blunt or subacute, base narrowly
rounded, cleft c. 1/5, lobes membranous, 0.7—1-1.5 mm long. Androecium slender, 3.5-
6.5 mm; androphore grooved or not, 2.5—4 mm, hairs less than 0.1 mm; synandrium 1.8—
2 by 0.5-0.7 mm, thecae 8-12, sterile apex acute, 0.1—-0.7 mm. Female flowers: pedicel
2.5 mm, bracteole scar 1 mm below apex; buds ovoid-oblong, narrowed to apex, 7 by
2.5 mm, lobes 1 mm long; ovary including stigma 5 mm. Fruits solitary, ellipsoid-ob-
long, 3.7—4.5 by 1.4-1.5 cm, apical part contracted into a + curved beak 4—7 mm, base
narrowed into a 4-5 mm long pseudostalk, hairs dense, scurfy scale-like, less than 0.1
mm; pericarp | mm thick; seeds oblong, 2.8 cm; stalk (peduncle and fruiting pedicel)
slender, nearly 1 cm long, fruiting pedicel 5 mm, bracteole scar 1.5 mm below the fruit.

Distribution — Malesia: Papua New Guinea (E Sepik, Gulf, Madang, and Central
Prov.); see also under the subspecies.

Habitat & Ecology — Montane forest; (400—)1200—1600 m altitude.

Note — The irregular warty inner surface of the male perianth is also found in the
obviously related species M. pumila and M. sogeriensis.

KEY TO THE SUBSPECIES

1a. Bracteole scar in male (1—)2 mm below the perianth; androphore 4 mm long, nearly
twice as long as the synandrium. Fruits (incl. pseudostalk) 4.5 cm a. subsp. fugax

b. Bracteole scar in male apical; androphore (1.8—)2.5—3 mm long, about as long as the
synandrium. Fruits (including pseudostalk) 3.7 cm ... b. subsp. septentrionalis

a. subsp. fugax

Twigs slender, 1—1.5 mm diameter. Leaves above drying dull brown-grey; lateral nerves
8-10 pairs. Male inflorescences slender, peduncle 0.5 mm thick. Male flowers: pedicel
5-6 mm long, bracteole scar (1—-)2 mm below the apex; buds 6.5 mm long, lobes 1.5 mm
long; androphore 4 mm long; synandrium 2 mm long, sterile apex 0.4 mm long. Fruits
including pseudostalk 4.5 cm long.

Field-notes — Flowers green, cream, or cream-tipped greenish. Fruits ochre.

De Wilde — Myristicaceae (Myristica) 475

Distribution — Malesia: Papua New Guinea (Central Prov.: near Isuarava, and Lala).

Habitat & Ecology — Primary and degraded montane forest; 1200-1600 m altitude;
fl. & fr. Jan—Feb.

Note — May be confused with the related M. pumila, a species generally with non-
pedunculate male inflorescences.

b. subsp. septentrionalis W. J. de Wilde

Myristica fugax W.J. de Wilde subsp. septentrionalis W.J. de Wilde, Blumea 40 (1995) 286. —
Type: Schlechter 18670 (male fl.), Papua New Guinea.

Twigs 1.5 mm diameter. Leaves above drying dark brown; lateral nerves 8—12(-14)
pairs. Male inflorescences stouter than those in subsp. fugax, peduncle | mm thick. Male
flowers: pedicel 4 mm long, bracteole scar apical; buds 6 mm long, lobes 1 mm long;
androphore 2.5—3 mm long; synandrium 2.5 mm long, sterile apex 0.5 mm long. Fruits
including pseudostalk 3.7 cm long.

Distribution — Malesia: Papua New Guinea (E Sepik, Gulf, and Madang Prov., Bis-
marck Mts).

Habitat & Ecology — Montane forest; 300-1200 m altitude; fl. Mar., Nov.

Note — Some slightly deviating collections are discussed by De Wilde (1.c.).

53. Myristica fusca Markgr.

Myristica fusca Markgr., Bot. Jahrb. Syst. 67 (1935) 158; A.C. Sm., J. Arnold Arbor. 22 (1941) 67;
J. Sinclair, Gard. Bull. Sing. 23 (1968) 252, f. 25; Foreman, Handb. Fl. Papua New Guinea |
(1978) 198: W.J. de Wilde, Blumea 40 (1995) 286. — Type: Schlechter 16848, Papua New Guinea,
West Sepik.

Tree 20-30 m. Twigs 2.5-6 mm diameter, late glabrescent, hairs red-brown, coarse,
+ erect, 1(—1.5) mm; older twigs dark grey-brown, not cracking; not or inconspicuously
lenticellate. Leaves chartaceous or subcoriaceous, ovate-oblong or oblong-lanceolate,
14-33 by 5-12 cm, base cuneate, rounded, or subcordate, apex acute( -acuminate); upper
surface olivaceous-brown, lower surface greyish (brown), hairs (especially on the veins)
persistent, conspicuous, spaced, 0.1—1.5 mm, with view on papillose lower leaf surface;
dots absent; midrib slightly raised above, lateral nerves 20—25(—30) per side, at 60—90°
to the midrib, + sunken above, lines of interarching distinct, venation coarsely netted,
sunken above and rendering the leaf bullate; petiole pubescent or late glabrescent, 10—
17 by 3-5 mm; leaf bud 10-15 by 4-5 mm, hairs dense, + rough, rust-coloured, 1—1.5
mm long. Inflorescences between the leaves or below, of the Knema-type, sessile or
pedunculate; in male a to 6 (by 4) mm pedunculate, + woody, scar-covered brachyblast,
simple or bifid, to 1 cm long; flowers in a loose cluster of (2—)4—6; buds of almost equal
size; female inflorescence not seen, presumably 1—3-flowered only. Inflorescence and
flowers with persistent rough dense rusty hairs 1-1.5 mm; bracts 2 mm, late caducous.
Male flowers: stout, pedicel 10-14 by 2-2.5 mm, bracteole broadly rounded, | mm
long, (late) caducous; buds (ovoid-)oblong or (long-)fusiform, 14—16(-17) by 5-6 mm,
apex subacute, cleft 1/8-1/10, lobes leathery, 1.5(—2) mm long, 0.5—0.6 mm thick.
Androecium 12-14 mm; androphore 3—4 by (1.5—)2 mm, in the lower half with stiff
rufous hairs, 1-1.5 mm; synandrium 7-8 by 1.5-2 mm, thecae c. 20, sterile apex broad
or narrow, conical, subacute, 2 mm long. Female flowers (Markgraf, Sinclair, Foreman,

476 Flora Malesiana, Ser. I, Vol. 14 (2000)

l.c.): pedicel 3-8 by 2 mm; buds broadly ovoid, 10 by 5—7 mm, cleft 1/4; ovary nearly as
long as the perianth, ovoid-conical, densely setose, stigma substipitate. Fruits single,
(ellipsoid-)oblong, 6-7 by 3.5—4 cm, apex narrowed, (sub)acute or shortly beaked, base
+ narrowed, hairs dense, rust-coloured, 1—1.5 mm long; pericarp 4—5 mm thick; seeds
ellipsoid-oblong, 2.5—3.5 cm, dark brown; fruiting pedicel stout, 10-12 by 5-6 mm, late
glabrescent, bracteole scar + apical.

Field-notes — Branching horizontally. Buttresses to 1 m out. Bark brown or black,
scaly; inner bark reddish; exudate red; wood red-brown, heartwood pink. Leaves pale or
glaucous below. Flowers brown hairy, or yellow. Fruits rust-coloured.

Distribution — Malesia: New Guinea (a limited number of collections known from a
restricted area in NE Papua Barat and Papua New Guinea: W Sepik & Madang Prov.).

Habitat & Ecology — Subcanopy tree of hillside or foothill forest; also on copper-
rich soil; 100—850 m altitude; fl. & fr. throughout the year.

Notes — 1. Myristica fusca is close to M. sphaerosperma and M. womersleyi; for dif-
ferences, see the key to the species.

2. Myristica fusca is readily recognizable by its somewhat bullate leaves with coarse
dark rust-coloured indumentum below and its comparatively large flowers.

54. Myristica fusiformis W. J. de Wilde

Myristica fusiformis W.J. de Wilde, Blumea 40 (1995) 287, f. 3d. — Type: Milliken 1408, W Irian
Jaya.

Treelet c. 3 m tall. Twigs 1-1.5 mm diameter, early glabrescent, hairs very minute;
older twigs coarsely striate or slightly flaking, with few inconspicuous lenticels. Leaves
membranous or chartaceous, elliptic-oblong, 9.5—15(—17) by 3.5—6 cm, base attenuate,
apex acute-acuminate; upper surface olivaceous or brown, lower surface brown-grey,
glabrous, with some scattered whitish scale-like hairs less than 0.1 mm; inconspicu-
ously densely fine-papillose; dots absent; midrib slightly raised above, lateral nerves
10-15 per side, at 60—70° to the midrib, flat or sunken above, on lower surface brownish
and + contrasting, lines of interarching + distinct, venation fine, faint or distinct below;
petiole 8-15 by 0.8—-1.5 mm; leaf bud 10 by 1-1.5 mm, hairs 0.1 mm or less. Male in-
florescences, male and female flowers not seen. Female inflorescences (from infruc-
tescences) sessile, of the Knema-type, small, consisting of a single axillary flower. Fruits
solitary, + narrowly fusiform, 7—-8.5 cm, apex acute, with or without pseudostalk, hairs
inconspicuous, pale rust-coloured, mealy, interwoven, scale-like, 0.1 mm or less, (partly)
glabrescent; pericarp (1—)2 mm thick; seeds oblong, 3 cm; fruiting pedicel slender or
thickish, 9-10 mm, fissured and transversely cracked, glabrous (hence flowers possibly
glabrescent), bracteole scar obscure, 2.5 mm below the apex.

Distribution — Malesia: SW Papua Barat (known only from the type specimens of
the two subspecies).

KEY TO THE SUBSPECIES

la. Fruits 7.5 cm long, tapering at base, but without pseudostalk. — Montane, at 1700 m
eyelet aawolow, mae dash Oh, Ree ee a. subsp. fusiformis

b. Fruits 7.5—8 cm long, tapering at base into a narrow pseudostalk 15-20 mm long. —
owlandiicen’,.Ameishes ey eee ee b. subsp. pseudostipitata

De Wilde — Myristicaceae (Myristica) ATT

a. subsp. fusiformis

Treelet 3 m. Twigs 1-1.5 mm diameter. Leaves chartaceous, 10-17 by 4.5—6 cm, apex
narrowly acuminate; upper surface dark brown, lower surface grey-brown, venation faint,
lateral nerves 10-15 per side; petiole (10—)15 by 1-1.5 mm. Fruits + fusiform, 7.5 by 2
cm, the basal part gradually narrowed but pseudostalk absent; pericarp largely glabrescent;
fruiting pedicel 10 by 3-4 mn, fissured, with a few lenticels. — Fig. 63d.

Field-notes — Leaf underside pale. Fruits ochre.

Distribution — Malesia: W Papua Barat (Kurima, close to Ilamik village).

Habitat & Ecology — Nothofagus-dominated forest; 1700 m altitude; fr. Oct.

Note — Related to M. tubiflora (a lowland species with smaller fruits) on account of
the sessile, elongate fruits.

b. subsp. pseudostipitata W.J. de Wilde

Myristica fusiformis W.J. de Wilde subsp. pseudostipitata W.J. de Wilde, Blumea 40 (1995) 287,
f. 3e. — Type: von Rémer 304, West Irian.

Possibly a small tree, size not known. Twigs 1 mm diameter. Leaves thinly membra-
nous, 9-10 by 3.5 cm, apex shortly acute-acuminate; upper surface rather pale brown-
olivaceous, lower surface grey, venation distinct, lateral nerves c. 10 per side; petiole 8
by 0.8 mm. Fruits long-fusiform, 8—8.5 by 1.5 cm including the basal part contracted
into a narrow pseudostalk 15-20 mm long; pericarp largely minutely pubescent, partly
glabrescent; fruiting pedicel 9 by 1.5 mm. — Fig. 62d, 63e.

Distribution — SW Papua Barat (lower Lorentz River, “Noord River’).

Habitat & Ecology — Presumably collected in forest at low altitude; fr. Sept.

Notes — 1. Known only by its fragmentary type specimen, which is readily distin-
guishable from the type subspecies and all known Myristicas in New Guinea by its elon-
gate solitary fruit with long pseudostalk, and long fruiting pedicel, in the axil of a leaf
scar. Vegetatively it is similar to M. brachypoda from S Papua New Guinea, but in that
species the papillation on the lower leaf surface is more pronounced, and the fruits,
although with a similar indumentum, are smaller, almost sessile, and lack a pseudostalk.

2. The distinct reticulation of the lower leaf surface is reminiscent of M. globosa, a
species differing in its (sub)globose fruits.

55. Myristica garciniifolia Warb.

Myristica garciniifolia Warb., Mon. Myrist. (1897) 525, t. 19; Markgr., Bot. Jahrb. Syst. 67 (1935)
169; J. Sinclair, Gard. Bull. Sing. 23 (1968) 196, f. 13; Foreman, Handb. Fl. Papua New Guinea
1 (1978) 198: W.J. de Wilde, Blumea 40 (1995) 288. — Type: Beccari FI acc. nos. 7735, a,
northern West Irian, Humboldt Bay.

Tree 10-30 m. Twigs 2.5—4(—5) mm diameter, yellowish or reddish brown, early
glabrescent, hairs grey, 0.1 mm or less; older twigs sometimes + fissured or flaking,
sometimes with few lenticels. Leaves chartaceous or subcoriaceous, elliptic-oblong, 12—
30 by 5—11 cm, base + cuneate or (narrowly) rounded, apex acute-acuminate; upper sur-
face olivaceous or brown, lower surface almost concolorous, glabrous; not papillose;
dots absent; midrib flat or somewhat raised above, lateral nerves 17—25 per side, at 45—
80° to the midrib, flat, faint above, faint and little raised below, lines of interarching and

478 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 74. Myristica garciniifolia Warb. a. Leafy twig with male inflorescences; b. male bud; c. male
flower at anthesis; d. androecium; e. female inflorescence; f. female flower at anthesis; g. pistil;
h. fruit [a: Atasrip 79, b—d: BW 3326 (Runtuboy); e—h: BW 4320 (Koster)]. — Scale bar for a, e, h
='2.cm; for'b, c, f =5 mm; for d, ¢ = 2 mm.

De Wilde — Myristicaceae (Myristica) 479

venation indistinct; petiole 15—25 by 2.5—4 mm; leaf bud 15-20 by 3-4 mm, hairs less
than 0.1 mm. Inflorescence between the leaves or below, pedunculate, glabrescent,
indumentum minute; bracts small, caducous; in male: 3—5 by 2—3.5 cm, peduncle flat-
tened, 10-23 by 2-4 mm, ending in a single (somewhat adaxial) flower and with (1) 2
(or 3) branches, (1—)2—5(-15) mm, + flattened (occasionally the middle branch once
more forked), with to 10 mm long scar-covered brachyblasts with subumbels of (5—)10—
15 flowers; buds of almost equal sizes; female like male, but much reduced, peduncle
10-15 mm, generally 3-flowered; flowers + coriaceous, appearing as if glabrous, but
hairs widely spaced, 0.1 mm or less, or glabrescent. Male flowers: pedicel + flattened,
5-8 by 1.5 mm, bracteole 2 mm, persistent, (sub)apical; buds ovate-ellipsoid, 7—8.5 by
(3-)4-5 mm, base + attenuate or rounded, + narrowed to the subacute apex, angled, cleft
2/3—3/4, lobes (4—)5—6 mm long, 0.3(—0.5) mm thick. Androecium slender, 5—6.5 mm;
androphore 1—1.5 by 0.8 mm, in the lower half with brownish hairs, 0.2 mm; synandrium
sometimes blunt triangular, 4—5 by | mm, thecae 14-20, sterile apex absent or irregular-
ly shaped, to 0.2 mm. Female flowers: pedicel 6-8 by 2 mm, bracteole subapical; buds
ovoid-oblong, narrowed to the apex, angled, 8—9 by (4—)5 mm, lobes 4 mm; ovary ovoid,
4 by (3-)3.5 mm, hairs 0.2—0.3 mm. Fruits 1-3 per infructescence, (ovoid-)oblong, 4.5—
5.5(-6) by (2.5—)3 cm, apex narrowly rounded with stigma remnant, base rounded or
+ attenuate, hairs brown powdery, 0.5 mm long, partly glabrescent; pericarp 3 mm thick;
seeds ellipsoid-oblong, 3—3.5 cm; fruiting pedicel 10-17 by 3 mm, glabrous, bracteole
scar 3-4(-10) mm below the apex. — Fig. 74.

Field-notes — Medium tree with small buttresses. Bark smooth, rarely scaly, brown
or blackish: slash red; exudate orange or red; sapwood pale, no heartwood. Leaves light
green below. Flowers yellow or creamy yellow. Fruits brown hairy.

Distribution — Malesia: NW Papua Barat (in a rather restricted area in the Jayapura
region, also Japen I.); not yet recorded from NW Papua New Guinea where it can be
expected in the adjoining NW Sepik Province.

Habitat & Ecology — Coastal rain forest; not rare in the lower Cyclops Mts; clayey
soils; up to c. 350 m altitude; fl. & fr. throughout the year.

Notes — 1. Myristica garciniifolia seems related to the West Malesian M. elliptica,
with similar inflorescences and conspicuously angled buds. It also resembles the Papuan
M. schleinitzii which differs in its non-angular buds and smaller glabrescent fruits.

2. By its very short, inconspicuous indumentum of leaf bud and innovations, and its
early glabrescence, M. garciniifolia makes a completely glabrous impression; it is fur-
ther distinguishable by its faint leaf venation.

56. Myristica gigantea King

Myristica gigantea King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 288, pl. 110; Warb., Mon. Myrist.
(1897) 400; J. Sinclair, Gard. Bull. Sing. 16 (1958) 343, f. 22; 28 (1968) 163; W.J. de Wilde,
Blumea 42 (1997) 167; Tree Fl. Sabah & Sarawak 3 (2000) 462. — Lectotype: King’s coll. 6050,
Peninsular Malaysia.

Myristica motleyi Warb., Mon. Myrist. (1897) 400 & 684, t. 14, f. 1, nom. nud., p.p. quoad Motley
145 (Labuan).

Tree 17-40 m. Twigs 1.5—2.5(—3) mm diameter, late glabrescent, hairs either short,
0.1—0.2 mm, or hairs conspicuous, rust-coloured, 0.5—1 mm mixed with emergent hairs

480 Flora Malesiana, Ser. I, Vol. 14 (2000)

1-1.5 mm (see note 2); older twigs (coarsely) flaking; lenticels inconspicuous or absent.
Leaves chartaceous or thinly coriaceous, elliptic-oblong or + lanceolate, 7-15 by 2—4
cm (to 22 by 8 cm of sapling shoots), base rounded or attenuate, apex rounded, rather
blunt, or acute; upper surface olivaceous or brown, lower surface greyish brown,
glabrescent, hairs scattered, 0.1 mm or less; distinctly papillose; dots absent; midrib
above slightly raised, lateral nerves 10—15(—18) per side, at 45—70° to the midrib, lines
of interarching and venation indistinct on both surfaces; petiole 12-25 by 1-2 mm; leaf
bud 10-15 by 1.5—2.5 mm, hairs 0.1—0.5 mm. /nflorescences generally below the leaves,
pedunculate, with shaggy hairs (0.2—)0.5—1 mm, sometimes glabrescent; in male: 2.5—
3.5 by 2.5—3 cm, peduncle 6-10 mm, lower branches 10—15 mm, central axis 10-20 mm
long, with | or 2 subsessile subumbels, the latter with 6-12 flowers, buds of variable
sizes; bracts caducous, 4 mm; in female: 1.5 cm long, + branched, few-flowered; flow-
ers and inflorescences with hairs (0.2—-)0.5 mm. Male flowers: pedicel 4-8 by 0.5 mm,
bracteole 2 mm, caducous; buds ovoid-ellipsoid, 4-4.5 by 3(—3.5) mm, cleft 1/5—1/4,
lobes membranous, (0.2—)0.3 mm thick. Androecium slender, 3.5 mm; androphore 1.5
by 1 mm, with hairs 0.3 mm in the lower 2/3; synandrium 2 by 0.8—1 mm, apex = acute,
thecae 14—16(?), sterile apex absent. Female flowers not seen (young fruits densely pu-
bescent). Infructescence 1—1.5 cm long. Fruits 1 or 2, broadly ovoid, 5—8 by 4.5—5 cm,
early or late glabrescent, hairs mealy, 0.2-0.3 mm, persistent at the base; pericarp 15-20
mm thick; seeds ellipsoid, 3—3.5 cm; fruiting pedicel + stout, 5-9 mm.

Field-notes — Frequently with (low) buttresses or (few) stilt-roots, up to 2 m high,
slender or thick, round. Bark generally blackish (charcoal), longitudinally fissured, or
often flaking in strips or pieces, 1 cm wide, brittle; inner bark 10—12 mm thick, brown or
red; sap red, sometimes sticky; sapwood white-yellow; heartwood brown. Leaves dull
pale green below. Flower buds brown. Fruits yellow or orange, with minute brown hairs.
Torquebiau c.s. ET 751 suggested the beautiful tree to be tried in plantations.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Singapore, Borneo (Sarawak,
Sabah, W, E & NE Kalimantan, Labuan, Nunukan).

Habitat & Ecology — Lowland dipterocarp forest; sandy (loam) soil; 0—100(—450) m
altitude; fl. & fr. possibly throughout the year.

Notes — 1. Strange enough, of this apparently fairly common tree, only Scortechini
1949, and one recent collection (Borneo), bear well-developed male flowers. Female
flowers have never been collected.

2. Variable in length of the hairs on the sterile leaf bud, either 0.1—0.2 mm only or up
to 2 mm long; because of this, M. gigantea appears twice in the keys. Whether this dif-
ference has taxonomic implications is not known.

3. Related to M. maingayi, but with smaller inflorescences, flowers and fruits. It reaches
a large size as a tree, and is known as the tallest Malayan Myristica. Myristica maingayi
has a lower stature, stouter twigs, larger leaves with acute apex, larger male inflorescen-
ces and flowers, androecium with sterile apex, and slightly longer glabrescent fruits.

57. Myristica globosa Warb.

Myristica globosa Warb., Mon. Myrist. (1897) 540, t. 19 — Type: Forbes 212 (Chalmers 10 in B,
lost), Papua New Guinea.
For more references and synonyms see the subspecies.

De Wilde — Myristicaceae (Myristica) 481

Tree 5-30 m. Twigs subterete or angular, |—2(—2.5) mm diameter, early glabrescent,
hairs 0.1 mm or less; older twigs not or but finely cracking or flaking; small lenticels
usually present. Leaves membranous or thinly coriaceous, elliptic-oblong to oblong-
lanceolate, 7-16(—21) by 2—7(—8) cm, base attenuate, apex acute-acuminate (the very tip
sometimes rather blunt); upper surface olivaceous or brown, lower surface greyish, early
glabrescent, hairs sparse, less than 0.1 mm; not or indistinctly papillose; dots absent;
midrib slightly raised or sunken above, lateral nerves 8—14(—17) per side, at 45—80° to
the midrib, flat or sunken (and sometimes very faint) above, lines of interarching not
distinct, venation reddish brown, mostly conspicuously contrasting with the lower leaf
surface; petiole (S—)8—12(—15) by 0.8—2(-2.5) mm; leaf bud 5—10(-13) by (0.5—)1-2
mm, hairs 0.1 mm or less. Inflorescences between the leaves and below, of the Knema-
type, sessile; in male: a simple or 2- or 3-armed scar-covered brachyblast, to 4 mm long,
with a subumbel of up to 15(—30) flowers, buds of various sizes, short pubescent; in
female: to | mm long, with 1—5 flowers (rarely more); flowers with appressed (dark)
brown hairs 0.1—0.2 mm, sometimes glabrescent, bracteole 1(—2) mm, caducous, (sub)-
apical. Male flowers: pedicel 4—6(—7) by 0.40.6 mm; buds membranous, ovoid-oblong
or ellipsoid-oblong, (4—)5—6.5 by 2-3(-4) mm, apex rather blunt or subacute, cleft 1/3
to (nearly) 1/2, the lobes 1.5—2.5(-3) mm long. Androecium 3—4(—5.5) mm; andro-
phore 1.5—2 by 0.4-0.6 mm, hairs 0.1—-0.3 mm in the lower 2/3—1/3, rarely glabrous;
synandrium (2—)2.5—4 by 0.6—0.8(—1) mm, thecae 10—16(—20?), sterile apex more or
less acute to flat, 0.2-0.3 mm or less, minutely lobed. Female flowers: pedicel 2—2.5
mm; buds ovoid, 4 by 3—3.5 mm, the lobes 1.5(—2) mm; ovary ovoid, 2.5—3 by 2-2.5
mm, hairs 0.2 mm. Fruits 1—3(—5) per infructescence, (sub)globose, | .5—3.2(—4) by 1.5—
2.5(-3) cm, with scurfy hairs, (0.1—)0.3 mm; the pericarp (1—)2—5 mm thick; seeds sub-
globose or broadly ellipsoid, 1-2(—2.2) cm; fruiting pedicel slender or rather thick, 2—5
(—7) mm.

Field-note — Pericarp + juicy.

Distribution — Solomon Islands; Malesia: NE Papua Barat (Jayapura region) and the
whole of Papua New Guinea (incl. New Britain). Two subspecies.

Habitat & Ecology — Usually in lowland, more rarely in submontane everwet or
rather seasonal forest.

Notes — 1. Myristica globosa is a widespread variable species. The two subspecies,
both also variable, are arbitrary. Frequently specimens are difficult to place in a subspe-
cies, especially those from the Solomon Islands. A few specimens obviously belonging
to M. globosa s.1., but not readily fitting in one of the subspecies, are separately dis-
cussed in the notes to the subspecies.

2. Myristica globosa s.1. is closely related to M. muelleri (S Solomon Is., NE Queens-
land) and M. tristis (NW Papua Barat, including Bird’s Head, and the Moluccas). The
latter species differs in a brown, not pale greyish lower leaf surface, without papillae, a
persistent bracteole, and subglobose fruits with thick pericarp. Although M. muelleri is
not known from Papua New Guinea, it may be found there as well, and therefore it
appears in the key to the subspecies. It is distinguishable by the male flower buds, cleft
1/4 to slightly over 1/3, broadly ellipsoid fruits with hairs up to 0.8 mm long [see W.J. de
Wilde, Blumea 36 (1991) 185; 42 (1997) 122, 132].

482 Flora Malesiana, Ser. I, Vol. 14 (2000)

KEY TO THE SUBSPECIES
(based on fruiting material, and also including M. muelleri)

la. Fruits 1.5-2.2(—2.4) cm long, (sub)globose, hairs 0.1—0.3 mm; pericarp (1—)2-3 mm
thick. — New Guinea (incl. New Britain), Solomon Islands .b. subsp. chalmersii

b. Fruits (2-)2.5—3(—4) cm long (if 2-2.5 cm, then fruits rather ellipsoid) ....... 2
2a. Fruits (broadly) ellipsoid, 2-3 cm long, hairs (0.1—)0.2—0.5(—0.8) mm; pericarp 2-3
mm thick. — S Solomon Islands, Australia (Queensland) .... M. muelleri Warb.

b. Fruits (sub)globose, 2.5—3(—4) cm long, hairs 0.1—0.3 mm; pericarp 3-5 mm thick.
— New Guineatincl New Brita) i csc scr spss Se 2). cyseetiees op sane a. subsp. globosa

a. subsp. globosa

Myristica globosa Warb., Mon. Myrist. (1897) 540, t. 19; J. Sinclair, Gard. Bull. Sing. 23 (1968)
378, p.p., f. 63, p.p. (excl. van Royen 3579), Foreman, Handb. Fl. Papua New Guinea 1! (1978)
199, p.p.; W.J. de Wilde, Blumea 38 (1994) 364, f. 5; 40 (1995) 288.

Male buds at anthesis cleft 1/3 to nearly halfway. Fruits (sub)globose, 2.5—3(—4) cm
long, hairs 0.1—0.3 mm; pericarp 3-5 mm thick; fruiting pedicel mostly short and thick,
up to 4 mm thick, coarsely fissured.

Field-notes — Tree to 30 m tall, rarely with small stilt-roots. Bark smooth or shallowly
fissured: outer bark (red-)brown; inner bark pink; wood white or yellowish with a red
tinge; exudate red. Flowers cream, yellow, golden, or brown; scented or not. Seeds brown.

Distribution — Malesia: Papua New Guinea (incl. New Britain).

Habitat & Ecology — Lowland, (degraded) rain forest on volcanic ash soil, forest
over limestone, Castanopsis-forest, and montane forest; 30-1220 m altitude; fl. & fr.
throughout the year.

Note — Carr 12890 has a stout habit and large male flowers, buds 6.5 by 4 mm; LAE
58538 (Stevens & Lelean) (New Britain) has conspicuously rich-flowered male inflores-
cences (c. 30 flowers per cluster), and further deviates in the warty inner surface of the
perianth; NGF 33284 (Leach), and NGF 46929 (Katik) from the foothills of Mt Suckling
(Milne Bay Prov., 350-400 m), have exceptionally large fruits, 4 cm long.

b. subsp. chalmersii (Warb.) W. J. de Wilde

Myristica globosa Warb. subsp. chalmersii W.J. de Wilde, Blumea 38 (1994) 366; 40 (1995) 288.

Myristica chalmersii Warb., Mon. Myrist. (1897) 519, t. 19. — Type: Chalmers 5 (B, lost; iso MEL,
n.v.), Papua New Guinea.

Myristica bduerlenii Warb., Mon. Myrist. (1897) 541, t. 19; Markgr., Bot. Jahrb. Syst. 67 (1935)
168. — Type: Bduerlen 1 (B, lost; iso MEL, n.v.), Papua New Guinea.

Myristica schumanniana Warb. in K.Schum. & Lauterb., Fl. Schutzgeb. Siidsee (1900) 328. —
Type: Rodatz & Klink (Tappenbeck) 70 (B, lost; iso BRSL, n.v.), Papua New Guinea.

Male buds at anthesis cleft 1/3 to nearly halfway. Fruits (sub)globose, 1.5—2.2(-2.4)
cm long, hairs 0.1—0.3 mm; pericarp (1-)2-3 mm thick; fruiting pedicel either rather
long and slender or short and thickish with fissured surface.

Field-notes — Small or medium-sized tree to c. 33 m; bole straight, sometimes with
small buttresses; branches often + horizontal, + whorled. Bark brown, 5 mm thick, smooth
or mostly finely longitudinally fissured or furrowed, often scaly (flaky); inner bark pink,

De Wilde — Myristicaceae (Myristica) 483

straw, or orange-brown, wood soft, cream or straw, staining reddish or orange-brown on
exposure; exudate red, watery, odourless, sometimes appearing in separate drops, rarely
sticky. Leaves mostly glaucous, greyish, or pale green below. Flowers (golden) yellow
or brown. Fruits (orange-)brown or red-brown; pericarp firm but fleshy, with a strong
spicy smell; seed brown, glossy.

Distribution — Solomon Islands; Malesia: NE Papua Barat (Jayapura area) and the
whole of Papua New Guinea (incl. New Britain).

Habitat & Ecology — Usually in (degraded) dry-land forest of flat land, slopes and
ridges, also in alluvial and bogged forest; red clay soil, volcanic soil, limestone, and
well-weathered soil; associated with Dracontomelum, Buchanania, Alstonia, with Celtis,
Vitex, or with Pometia, Spondias, and Canarium; 10—650(—1500) m altitude; fl. & fr.
throughout the year.

58. Myristica gracilipes J. Sinclair

Myristica gracilipes J. Sinclair, Gard. Bull. Sing. 23 (1968) 334, p.p. (excl. Schodde 2291 = M.
warburgii), £. 49; W.J. de Wilde, Blumea 40 (1995) 288. — Type: Kanehira & Hatusima 12277,
Irian Jaya.

Tree 6 m tall. Twigs faintly blunt-angular, 2-2.5 mm diameter, early glabrescent, hairs
minute, greyish; older twigs incidentally thinly flaking, with some small lenticels. Leaves
(thinly) chartaceous, + obovate-oblong, 16-19 by 5.5—6.5 cm, base short-attenuate or
nearly rounded, apex acute-acuminate; upper surface blackish brown, lower surface
brown-grey, glabrous (or glabrescent, hairs few, minute, scattered, whitish, appressed);
not papillose; dots absent; midrib flat above, lateral nerves 12-16 per side, at 45—60° to
the midrib, sunken above, bright reddish brown below, contrasting, lines of interarching
and venation distinct; petiole 8-15 by (2—)3-3.5 mm; leaf bud 7-8 by 2 mm, hairs 0.2—
0.3 mm. Inflorescences (from infructescences) between the apical leaves, presumably of
the Knema-type: 1- (or 2-)flowered, peduncle 1.5—2 mm long (see note 2). Male and
female flowers not seen. Fruits solitary, somewhat immature: elliptic(-oblong), 2.5 by
1.5 cm, apex 2 mm beaked, base contracted into + broad 4-5 mm long pseudostalk, hairs
thinly set, mealy, dull rust-coloured, 0.1 mm or less; pericarp 2 mm thick; seeds not
seen; fruiting pedicel 2.7—3.5 cm by 1 mm, bracteole scar 10 mm below the apex.

Distribution — Malesia: N Papua Barat (Dalman, 45 km inland from Nabire); known
only from the type.

Habitat & Ecology — In fringing rain forest; 500 m altitude; fr. Mar.

Notes — 1. Myristica gracilipes is close to M. warburgii subsp. hybrida, but the latter
has smaller leaves, fruits lower on the twigs, and shorter fruiting pedicels.

2. If male inflorescences of M. gracilipes appear to be pedunculate, the species keys
out in the vicinity of M. cylindrocarpa.

59. Myristica guatteriifolia A. DC.

Myristica guatteriifolia A.DC., Ann. Sc. Nat. 4, 4 (1855) 20, t. 4 (guatteriaefolia); Prodr. 14, 1
(1856) 193: Warb., Mon. Myrist. (1897) 412, t. 13; J. Sinclair, Gard. Bull. Sing. 16 (1958) 350,
f. 25: 23 (1968) 213; Backer & Bakh. f., Fl. Java 1 (1964) 139; W.J. de Wilde, Blumea 42 (1997)
168: Tree Fl. Sabah & Sarawak 3 (2000) 463. — Type: Cuming 1582, Luzon, Prov. Batangas.

484 Flora Malesiana, Ser. I, Vol. 14 (2000)

Myristica littoralis Miq., Fl. Ind. Bat. 1, 2 (1858) 57; Koord. & Valeton, Bijdr. Booms. Java 4
(1896) 173; Warb., Mon. Myrist. (1897) 418; Koord., Exk. Fl. Java 2 (1912) 257; K. Heyne,
Nutt. Pl. Ned. Indié (1927) 647. — Type: Teijsmann s.n., Java.

Myristica cookii Warb., Mon. Myrist. (1897) 414, t. 15; Lecomte, Not. Syst. 1, 4 (1909) 98; FI.
Gén. Indo-Chine 5 (1914) 97. — Syntypes: Phillips s.n.; Hb. Pierre 5433, S Vietnam, Pulau
Condor.

Myristica riedelii Warb., Mon. Myrist. (1897) 417, t. 15. — Types: Beccari Fl acc. no. 7749 (Riedel),
cultivated in Hort. Bogor., orig. Billiton, and Beccari FI acc. no. 7750, orig. Singapore.

Myristica palawanensis Merr., Philipp. J. Sci., Bot. 13 (1918) 283; Enum. Philipp. Flow. Pl. 2
(1923) 179. — Type: Merrill 9253, Philippines, Palawan.

Tree 10-25 m. Twigs 3—5(—10) mm diameter, hairs rust-coloured, 0.2—0.5 mm; older
twigs glabrescent, smooth or rarely flaking, usually without lenticels. Leaves chartaceous,
elliptic-oblong to oblong-lanceolate, (12—)15—35 by 3-12 cm, base rounded or attenu-
ate, apex acute-acuminate; upper surface olivaceous(-brown), lower surface grey-brown
or cinnamon, pubescent or late glabrescent, hairs dense, 0.2—-0.3 mm, with scattered
emergents ().5(—2) mm; not papillose; dots absent; midrib + raised above, lateral nerves
15—19(—25) per side, at about 45° to the midrib, flat or slightly sunken, lines of interarching
and venation distinct or faint; petiole 15-30 by 2—4 mm; leaf bud 10-20 by 2.5—4 mm,
hairs 0.2—1 mm. Inflorescences paniculate, between the leaves or sometimes axillary to
reduced leaves in short-shoots, with dense hairs 0.2—0.6 mm; bracts caducous, 2 mm
long; in male: 2-8 cm long, once or twice branched, peduncle (0—)4—20 mm, flowers in
clusters of (1-)2-10, buds of variable sizes; female inflorescences similar but shorter
and fewer flowered, simple or little-branched, 0.5—2 by 0.5—1 cm, peduncle to 7 mm;
flowers with hairs 0.3—0.6 mm. Male flowers: pedicel 3-6 mm, bracteole (2—)2.5—-3 mm,
caducous; buds ovoid or ellipsoid, (4—)5—7 by 3.5—5 mm, cleft 1/4—1/3, lobes 0.3-0.5
mm thick. Androecium 3—5 mm; androphore 1.5—2 mm long, the basal (1/2—)2/3—3/4
with pale hairs 0.40.7 mm; synandrium 2-3 by 0.8—1.8 mm, thecae 16-24, sterile apex
up to 0.2 mm, glabrous or rarely with few hairs 0.2—0.3 mm, or apex of synandrium
+ truncate, rounded, or to 0.3 mm hollowed. Female flowers: pedicel 1-2(—4) mm long;
buds broadly ovoid, 4—6 by 3.5—6 mm; ovary broadly ovoid, 2.5—3.5 mm diameter, hairs
dense, 0.5 mm long, stigma lobes oblique, faintly lobulate or not. Fruits up to 3 per
infructescence (sometimes seemingly more-fruited when infructescences are + clustered),
short ellipsoid or subglobose, 2.5—4(—4.5) by (2—)2.5—3 cm, hairs dense, velvety or shaggy,
0.5—1(—2) mm; pericarp 3—5(—8) mm thick (not woody); seeds 2—2.5(—3) cm long; fruit-
ing pedicel 1.5—5 mm long.

Field-notes — Sometimes buttresses 20(—200) cm high, 30(—150) cm out. Outer bark
chocolate, 2-4 mm thick, smooth or + rough, rarely fissured, usually cracked or scaly,
with irregular to + rectangular flakes, 2 by 4 cm, adherent; cambium white to yellow or
red: inner bark 5—10 mm thick, soft, fibrous, pinkish to beefy red; exudate watery, (pale)
red; sapwood usually pale, whitish to yellow, streaked with red; heartwood reddish.
Leaves below cinnamon, aureous, or rusty, which can be seen from a distance. Flowers
outside yellow-brown or rusty tomentose, perianth inside smooth, yellowish or some-
times partly purple-red. Fruits to 5 cm long; seeds to 3 cm.

Distribution — Burma, possibly Thailand (no specimens known), Indochina (P. Con-
dor); Malesia: Peninsular Malaysia (East Coast, according to Sinclair never found on
the West Coast), through Malesia east to Bali and the Philippines; not yet in Sulawesi.

De Wilde — Myristicaceae (Myristica) 485

Habitat & Ecology — Mainly but not always coastal; in primary conditions on coastal
rock, forest behind beach, e.g. in Oncosperma filamentosa forest near the sea (Nunukan),
or behind the Casuarina equisetifolia belt (especially in Brunei); also more inland at
riversides, hill slopes; mainly on sandy or rocky soils, white or granitic sand, etc., but
sometimes red earth, or black soil over sandstone, limestone, ultrabasic, rarely in de-
graded forest or belukar, once from (disturbed) peat swamp forest in Brunei; 0—200 m
altitude; fl. & fr. throughout the year.

Notes — 1. Specimens from black soil over sandstone in Sabah have conspicuously
small flowers. In several specimens from Brunei and Sabah the inflorescences are small
(reduced) and grouped in short-shoots, axillary to fallen leaves, giving the impression of
one compound inflorescence, mostly with an apical vegetative bud. PNH 117198 ( Samar)
deviates in having rather thick blackish twigs with distinct contrasting lenticels; its fruits
have a thick pericarp, 6-8 mm.

2. Myristica guatteriifolia is mainly a coastal species, well characterized by essen-
tially paniculate inflorescences, and leaves pubescent below (hairs of mixed sizes). It is
related to M. agusanensis from the Philippines, which is a smaller and more delicate
version of the present species. Small-leaved inland specimens from Borneo may resem-
ble M. beccarii.

60. Myristica hollrungii Warb.

Myristica hollrungii Warb., Mon. Myrist. (1897) 490, t. 19; K. Schum. & Lauterb., Fl. Schutzgeb.
Siidsee (1900) 328, p.p. (excl. syn. Myristica heterophylla, p.p.); A.C. Sm., J. Arnold Arbor. 22
(1941) 67; J. Sinclair, Gard. Bull. Sing. 23 (1968) 405, f. 68, 69; Foreman, Handb. Fl. Papua
New Guinea 1 (1978) 199: W.J. de Wilde, Blumea 40 (1995) 289; 43 (1998) 180. — Type:
Hollrung 701, Papua New Guinea.

Myristica heterophylla K. Schum. in K. Schum. & Hollrung, Fl. Kaiser Wilhelmsland (1889) 45,
p.p. — Type: Hollrung 648 (B, lost; iso BO, n.v.) (see Sinclair, 1.c.: 410), Papua New Guinea.

Myristica albertisii Warb., Mon. Myrist. (1897) 532, t. 19. — Type: d’Albertis 9 (B, lost; iso FI,
MEL, n.v.), Papua New Guinea.

Myristica euryocarpa Warb. in K. Schum. & Lauterb., Fl. Schutzgeb. Siidsee (1900) 327. — Type:
Rodatz & Klink (Tappenbeck) 33 (B, lost; BRSL, n.v.), Papua New Guinea.

Tree 6-36 m. Twigs 2—5(—10) mm diameter, usually subangular with two raised lines
joining the petioles, but not winged, frequently + hollow, but without ant-swellings;
(early) glabrescent, hairs palish, less than 0.1 mm; older twigs with small, + concolorous
lenticels. Leaves chartaceous, elliptic-oblong or oblong-lanceolate, 15—35(—40) by 4-12
(-15) cm, base cuneate, rounded, or shallowly cordate, apex acute-acuminate;, upper
surface olivaceous, lower surface olivaceous or pale brownish, early or late glabrescent,
hairs weak, dense or scattered, stellate, 0.1 mm or less; not or + papillose; dots absent;
midrib flat above, lateral nerves 15-25 per side, flat, at 45—60° to the midrib, lines of
interarching and venation indistinct; petiole 15-25 by 2-5 mm; leaf bud 10-15 by 2-4
mm, hairs 0.1(—0.2) mm. Inflorescences between the lower leaves or below, as in Knema;
in male: a simple or up to 4-branched, to 1 cm (rarely to 3 cm) long, scar-covered, gla-
brescent brachyblast, rarely to 5 mm pedunculate, with subumbel(s) of 5—10 flowers,
buds of various sizes; in female similar, short, (sub)sessile, with a 5—10-flowered clus-
ter; flowers with mealy hairs 0.1(—0.2) mm. Male flowers: pedicel 4—6 by 1 mm, bracteole
rounded, 2(-3) mm, (sub)persistent; buds thinly leathery, (subglobose), broadly ellip-

486

soid, or ovoid, 4-6.5 by 4-5 mm,
cleft 1/4(-1/3), lobes 1.5—2 mm
long, (0.3—)0.5 mm thick. Androe-
cium 3—4(—5) mm; androphore
(0.5—)1(—1.5) by 0.5—0.6 mm, hairs
minute, in lower 1/4 or lower; sy-
nandrium ovoid-oblong, 2—4 by |
mm, thecae c. 16, the sterile apex
+ acute, 0.1—0.3 mm. Female flow-
ers: pedicel 2—3(—4) by 2 mm, brac-
teole 2 mm; buds ovoid, 5—6 by
4-5 mm, lobes | mm long, ovary
ovoid-oblong, 4 by 3 mm, minutely
pale brown pubescent, stigma 2
(-4)-lobed, | mm. Fruits subsessile,
1—4(—5) per infructescence, (sub-
globose-)ellipsoid, 2.5—4.5(—5, see
note 2) by 2-3 cm, the apex often
shortly pointed, hairs thinly set,
mealy, brown, 0.1(—0.3) mm, + gla-
brescent, with some indumentum
remaining near the base and in de-
pressions; pericarp 3—5(—7) mm
thick; seeds (broadly) ellipsoid, 2—
3 cm; fruiting pedicel 3—7(—10) mm,
smooth or with few lenticels, bracte-
ole scar subapical. — Fig. 59a, 75a.

Field-notes — Usually a tall tree,
locally common; branches + whorl-
ed, often + horizontal; crown nar-
row; buttresses, prop-roots, or usu-
ally stilt-roots present. Bark smooth,
or mostly finely longitudinally fis-
sured or flaky (scaly), or faintly
peeling, dark (grey-)brown or black-
ish, or bark with raised horizontal
lenticels; inner bark reddish; exu-
date + colourless turning reddish;

Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 75. a. Myristica hollrungii Warb. Opened male bud
and schematic longitudinal section of androecium [De-
moulin 5958]. — b. M. hooglandii J. Sinclair. Opened
male bud (note scar of bracteole) and schematic longitu-
dinal section of androecium [Brass 25471]. — Scale bar
= 2mm.

blaze dark chestnut; sapwood straw, dark brown on exposure; heartwood absent or but
little differentiated in colour. Leaves grey-green or glaucous below. Flowers sometimes
on the older wood; perianth pale yellow or cream, the tips flushed pink, inside yellow-
ish; anthers straw. Fruits yellow(-brown) or pale brown, aril thick, fatty.

Distribution — Malesia: throughout New Guinea, including the Bismarck Archipelago

(New Ireland, W New Britain).

Habitat & Ecology — Lowland marshy forest; less frequently in dry-land forest or
lower montane and ridge forest; alluvial forest (with Barringtonia), periodically inun-

De Wilde — Myristicaceae (Myristica) A87

dated or marshy forest, close to mangrove; occasionally in sago-swamp woodland, or
forest on (deep) volcanic soil; also from regrowth forest. According to Brass 8008 char-
acteristic of tall forest of low flood plains of the Fly River; in some places overhanging
the rivers; 0-500(—1000) m altitude; fl. & fr. throughout the year.

Notes — 1. A homogeneous species, distinct in the (seemingly) glabrous lower leaf
surface, the often lined twigs, the seemingly (very short-pubescent) glabrous flowers,
and sessile infructescences with short-stalked glabrescent fruit.

2. Larger fruits, 4.5—5 cm long, are found among smaller-fruited specimens on Bagabag
and Umboi Is. (LAE 66001, NGF 43453); the latter specimen is somewhat approaching
M. bialata from the Bismarck Archipelago.

3. Myristica rumphii, based on a sterile collection originally described in Lauraceae,
does not belong to M. hollrungii as suggested by Kostermans; M. rumphii (see there) 1S
an older name of M. spanogheana.

A. Sometimes the lower leaves of a flush are retarded in growth and are cataphyll-
like, as is normally found in M. subcordata var. morindiifolia.

5. The lined twigs are often narrowly hollow inside, and sometimes a small hole 1S
present, obviously related to inhabitation by ants; however, conspicuous ant-swellings
in the twigs, as characteristic for, e.g., M. subalulata, are never present.

61. Myristica hooglandii J. Sinclair

Myristica hooglandii J. Sinclair, Gard. Bull. Sing. 23 (1968) 156, f. 7; Foreman, Contr. Herb. Aus-
tral. no. 9 (1974) 39; Handb. Fl. Papua New Guinea 1 (1978) 200, f. 89 (excl. syn. M. carrit);
W.J. de Wilde, Blumea 39 (1994) 347, f. le, e’; 40 (1995) 289. — Type: Hoogland 4206, Papua
New Guinea.

Tree 8-27 m tall. Twigs (2—)3—5 mm diameter, early glabrescent, hairs appressed,
glossy, 0.1 mm or less; older twigs blackish, cracking, indistinctly lenticellate. Leaves
(thickly) membranous, (elliptic-)oblong, (12—)15—35 by 3-10 cm, base attenuate or nar-
rowly rounded, apex acute(-acuminate); upper surface brown, lower surface (brownish)
grey, early glabrescent, hairs 0.1 mm or less; not papillose; dots absent; midrib (flat or)
raised above, lateral nerves (10—)13-17 per side, at 45—70° to the midrib, + flat, incon-
spicuous above, more or less raised below, often with intersecondary nerves, lines of
interarching and venation indistinct on both surfaces; petiole 15-20 by 2-4 mm; leaf
bud 10-15 by 2-3 mm, hairs 0.5—0.8 mm long. /nflorescences between the leaves or (as
in the type) below the leaves, arranged in I—1.5 cm long brachyblasts ending in a small
sterile leaf bud, pedunculate, glabrescent, indumentum minute; bracts caducous; in male:
peduncle of partial inflorescence 10-30 by 2-3 mm, with a subumbel of 4—7 flowers,
buds of almost equal sizes; in female: peduncle 5—10(—15) mm, one- or few-flowered;
flowers coriaceous, glabrescent, hairs appressed, silky, 0.1 mm. Male flowers: pedicel
10-16 by 1.5—2 mm, bracteole (?broadly rounded), caducous, scar conspicuous; buds
(broadly) ellipsoid, 9-16 by 6-9 mm, hardly collapsing on drying, cleft c. 1/2, lobes
5-7 mm long, | mm thick. Androecium (broadly) obconical or cylindrical, 5—10 mm
long; androphore 1-2 by 0.8-1 mm, (sub)glabrous (hairs 0.1—0.2 mm), synandrium 4—8
by 3-4 mm, apex rounded or + truncate, sometimes + triangular, thecae 30—40, sterile
apex absent, or apex sometimes shallowly hollowed. Female flowers: pedicel 5 mm
long; buds ovoid, 8 by 6 mm; ovary ovoid, 6 by 4.5-5 mm, hairs (0.5—)1(—2) mm.

488 Flora Malesiana, Ser. I, Vol. 14 (2000)

Infructescences: peduncle smooth or flaky. Fruits generally single, (broadly) ellipsoid
or oblong, 6-8 by 4-5 cm, apex (sub)obtuse, (late) glabrescent, hairs mealy, rust-col-
oured, 0.5—1(—2) mm long; pericarp 5(—10) mm thick; seeds ellipsoid or oblong, 3.5 cm;
fruiting pedicel 5—8 by 3—5 mm, striate, glabrescent. — Fig. 75b.

Field-notes — Small or medium-sized straight-boled tree. Bark grey-brown or black-
ish, longitudinally fissured and flaky; inner bark red-brown, or red on exposure; exudate
red or colourless, sticky or resinous; blaze red; wood cream or straw, tinged reddish
brown, of medium hardness. Leaves pale, dull, whitish or glaucous below, shiny dark
green above. Flowers (cream-)yellow, fragrant. Fruits ovoid or almond-shaped, lemon-
green, or (orange-)yellow, with brown indumentum; seeds grey.

Distribution — Malesia: a local endemic species of Papua New Guinea, known from
Morobe, Northern, Central and Milne Bay Provinces (also Normanby Is.).

Habitat & Ecology — Lowland rain forest, often fairly wet or swampy, also on lower
slopes of hills on skeletal soils, coastal forest; found associated with Mastixiodendron,
Maniltoa, and Tristaniopsis, or in Eucalyptopsis-dominated forest (at c. 450 m); altitude
0-450 m; fl. Apr.—July; fr. May—Dec.

Note — LAE 62150 (Katik) from Central Province has exceedingly small leaves, 12—
15 cm long only.

62. Myristica impressa Warb.

Myristica impressa Warb., Mon. Myrist. (1897) 537, t. 15; J. Sinclair, Gard. Bull. Sing. 23 (1968)
235, 481 (as doubtful species; see note 1); W.J. de Wilde, Blumea 42 (1997) 168. — Type:
Warburg 16716 (B, lost), Sulawesi, southwestern arm.

Myristica koordersii auct. non Warb.: J. Sinclair, Gard. Bull. Sing. 23 (1968) 262, f. 28, p.p., excl.
Kjellberg 2962 (= Myristica kjellbergii), excl. specimens from Moluccas and NE arm of Sula-
wesi (= Myristica koordersii).

Tree 10-25 m. Twigs 2-4(—5) mm diameter, early glabrescent, hairs 0.1 mm or less;
older twigs with or without few inconspicuous lenticels. Leaves thinly chartaceous or
subcoriaceous, elliptic-oblong or oblong(-lanceolate), 14-27 by 4-9 cm, base cuneate
(or nearly rounded), apex acute(-acuminate); upper surface olivaceous or dark brown,
lower surface conspicuously grey-brown or whitish, (late) glabrescent, with scale-like
widely spaced hairs 0.1 mm or less, and with scattered contrasting points (possibly scars
of fallen stouter hairs) (lens!); papillation not apparent; dots absent; midrib above little
raised, lateral nerves (13—)15—20 per side, at 40—50° to the midrib, + sunken (and faint)
above, nerves below purple or brown, sometimes contrasting in colour, lines of inter-
arching and venation indistinct; petiole 12-20 by 2.5—3 mm; leaf bud 10—15 by 2-3 mm,
with (appressed) hairs 0.1(—0.2) mm. Inflorescences between the leaves or below, of the
Knema-type, a (sub)sessile, simple or bifurcate scar-covered brachyblast to 7 mm, pu-
bescent or glabrescent; in male: with a cluster of 4-8 flowers, buds of variable sizes; in

Fig. 76. Myristica impressinervia J. Sinclair. a. Leafy twig with male inflorescences; b. male in-
florescence; c. male bud; d. ditto, opened, showing androecium. — M. impressa Warb. e. Twig with
female inflorescences; f. female bud, opened, showing pistil; g. male bud; h. ditto, opened, showing
androecium; i. fruit [a—d: Kjellberg 2427; e, f: bb Cel. I-403; g, h: Elbert 3025; i: Meijer 10061). —
Scale bar for a, i = 2 cm; for b = 3.3 mm; for c, d= 1.5 mm; for e = 6.5 mm; for f—h = 2 mm.

De Wilde — Myristicaceae (Myristica)

489

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490) Flora Malesiana, Ser. I, Vol. 14 (2000)

female similar, fewer-flowered; flowers with hairs 0.2—0.4 mm. Male flowers (submature):
pedicel 4—5 mm long, bracteole ovate or + triangular, 3.5 mm, late caducous; buds ellip-
soid, (4-)4.5—5 by 3—3.5 mm, cleft c. 1/3, lobes 0.3 mm thick. Androecium (3—)3.5 mm;
androphore | by 0.8 mm, hairs less than 0.1 mm; synandrium 2 by | mm, thecae 16-20,
sterile apex hemiglobose, 0.5 mm, glabrous. Female flowers: pedicel 4(—5) mm, bracteole
3 mm, caducous; buds (immature) ovoid, 3.5(—5) by 3(—4) mm, cleft c. 1/3; ovary ovoid,
2.5 by 2 mm, hairs less than 0.1 mm. Fruits solitary or 2 per infructescence, broadly
ellipsoid, 3-4.2 by 2-3 cm, hairs dense, mealy, 0.1 mm; pericarp 2-4(—5) mm thick;
seeds subglobose or ellipsoid, 2-3 cm; fruiting pedicel slender or + stout, 2-4(—7) mm
long. — Fig. 76e-i.

Field-notes — Sap deep red. Fruit with golden or yellow indumentum; seed reddish
brown.

Distribution — Malesia: Sulawesi (Minahasa?, N Central, Central, SW, SE).

Habitat & Ecology — Hill or montane forest, moderately dry forest; also on ultrabasic;
200-1500 m altitude; fl. July, Sept.; fr. throughout the year.

Notes — 1. Myristica impressa (type lost) was regarded by Sinclair (1.c.) as a doubt-
ful species, possibly based on an atypical specimen of M. koordersii, which was accept-
ed by him as a very variable entity. However, M. koordersii can best be regarded as a
related local species of NE Sulawesi (Minahasa). The type of M. impressa was collected
in the southwestern arm; preferably a neotype should be chosen from that area. Myristica
koordersii differs in its more slender twigs, smaller leaves, longer and more coarsely
hairy fruiting pedicel (possibly the indumentum of the flowers is longer as well), and
fruits with a thicker pericarp. Other species with which M. impressa may be confused
are M. agusanensis (Philippines; resembling in the leaves but differing in paniculate in-
florescences and more conspicuously hairy fruits), M. kjellbergii (Sulawesi; see De Wilde,
l.c.: 170), and M. pilosigemma (Philippines; with conspicuous long-haired indumentum
on terminal leaf bud and fruits).

2. Milliken 989 (male) and 1091 (sterile) (N Sulawesi, 1500 m) probably belong to M.
impressa, but deviate in the 2 mm long pedunculate male inflorescence, the slightly
larger, 5 mm long male perianth, the shorter pedicel, 1.5 mm only, and the shorter brac-
teole, 2 mm; the androecium is + club-shaped, the androphore fine-pubescent, the synan-
drium about as long as the androphore, both c. 1.5 mm long, anthers (?)10, sterile apex
absent. The collection was annotated as a tall tree, 24 cm in diameter, with conspicuous
stilt-roots. More fertile material of M. impressa is needed to understand its variability.
Kjellberg 2990, a fruiting collection from C(—SW) Sulawesi, may represent an undescribed
species. It resembles M. impressa, but differs in a more tiny habit, with smaller leaves
which are not whitish below.

63. Myristica impressinervia J. Sinclair

Myristica impressinervia J. Sinclair, Gard. Bull. Sing. 23 (1968) 232, f. 19; W.J. de Wilde, Blumea
42 (1997) 168. — Type: Kjellberg 2427, Sulawesi.

Tree 6-15 m. Twigs 0.7—1.5(—3) mm diameter, early glabrescent, hairs 0.2—0.3(0.5)
mm; older twigs not flaking, without or with few indistinct lenticels. Leaves membra-
nous or thinly chartaceous, 9-19 by 46.5 cm, elliptic(-oblong), base (short) cuneate,
apex acute(-acuminate); upper surface dark brown, below greyish, seemingly glabrous,

De Wilde — Myristicaceae (Myristica) 49]

but hairs grey-brown, scale-like, scattered, 0.1 mm or les; not papillose; dots absent;
midrib above sunken, lateral nerves 6—13 per side, at 45(—60)° to the midrib, slender,
and sunken above, nerves below brown, contrasting in colour, lines of interarching and
venation indistinct; petiole 5—12 by 1(-1.5) mm; leaf bud 6-9 by 1—-1.5(—2) mm, hairs
rough, 0.3(—0.5) mm. Inflorescences between the leaves or below, approaching the Knema-
type; in male: peduncle slender, (3—)5—10 mm, pubescent, ending in 1—3 slender scar-
bearing brachyblasts to 5 mm long, each with a loose subumbel of 2—4 flowers, buds of
various sizes; female inflorescence not known; flowers with rough hairs 0.2—0.3 mm.
Male flowers: pedicel slender, 4-5 mm long, bracteole broadly reniform, 2 mm long,
persistent; buds ovoid(-oblong), 3.5—4 by 2.5 mm, cleft nearly 1/2, lobes 0.2(-0.3) mm
thick. Androecium 2.5 mm long; androphore well-demarcated from synandrium, 0.5—
0.6 by 0.3—-0.4 mm, with hairs 0.2 mm in the lower half; synandrium ovoid-ellipsoid,
2 by 1-1.2 mm, thecae c. 16, 1.5 mm long, the sterile apex comparatively large, blunt,
0.5 mm. Female flowers and fruits not seen. — Fig. 76a—d.

Field-notes — Richly branched small tree, stem dbh up to 25 cm. Leaf undersurface
silvery white. Flowers outside brown or yellow, inside yellowish white.

Distribution — Malesia: S Central and SE Sulawesi.

Habitat & Ecology — Primary forest, forest on limestone: 0-100 m altitude; fl. Oct.

Note — Myristica impressinervia is characterized by its delicate slender habit, slen-
der pedunculate (male) inflorescences, and rather small thin leaves with a whitish lower
surface and nerves sunken above.

64. Myristica inaequalis W.J. de Wilde

Myristica inaequalis W.J. de Wilde, Blumea 43 (1998) 252, f. 2. — Type: R. J. Johns 7699, Irian
Jaya, Bird’s Head.

Tree to 20 m. Twigs 2.5-3 mm diameter, late glabrescent, hairs rusty, 0.5(-1) mm;
older twigs blackish brown, finely striate, neither cracking nor flaking, with scattered
inconspicuous lenticels. Leaves chartaceous or coriaceous, elliptic-oblong, (13—)15-19
by 3-5.5 cm, base cuneate, apex acute-acuminate; upper surface dark brown, lower sur-
face light brown, hairs (sub)persistent, widely spaced, mixed: 0.1 mm and | mm long;
papillose surface clearly visible (lens!); dots absent; midrib raised above, lateral nerves
16-20 per side, at 70—80° to the midrib, flat and inconspicuous above, lines of interarching
thin though distinct below, venation indistinct; petiole 8-15 by 2-2.5 mm, late glabrescent;
leaf bud 15 by 2—2.5 mm, hairs 0.5—1 mm. Male inflorescences unknown; female inflo-
rescences (known from a single specimen) between the leaves, almost of the Knema-
type, a scar-covered brachyblast 3 mm diameter, 1- or 2-flowered, peduncle subterete,
1-4 by 2 mm, hairs 0.5(—1) mm; bracts minute, caducous; flowers with bright brown
hairs, 0.5-1 mm. Male flowers unknown. Female flowers: pedicel 10 by (1.5—)2 mm,
bracteole about median, caducous, scar inconspicuous; buds ovoid-oblong, 12-13 by
5—6 mm, much narrowed in the upper half, cleft c. 1/5, lobes 2.5—3 mm long, 0.5 mm
thick, at anthesis curved outwards; ovary c. 7 mm long, the hairs dense, golden brown,
1 mm. Fruits not seen, but see field-notes. — Fig. 77.

Field-notes — Young stems densely covered with ferruginous hairs. Leaves borne
horizontally to the stem, to 18 by 3.5 cm, petiole to 1.5 cm. Fruits (immature) to 7 by
5 cm, ferruginous.

492 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 77. Myristica inaequalis W.J. de Wilde. Habit of leafy twig with one-flowered female inflores-
cence [R. J. Johns 7699]. — Scale bar = 2 cm.

Distribution — Malesia: Papua Barat (NE Bird’s Head, Arfak Mts, track from Wamare
to Mokwam); known only from the type.

Habitat & Ecology — Ridge forest; 800-850 m altitude; fl. & fr. Mar.

Note — Myristica inaequalis is provisionally reckoned to the group with larger leaves
(blade length 15 cm or more) and subsessile or short-stalked inflorescences (as in Knema).
It keys out next to M. fusca (to which it is closely related). The caducous bracteole,
situated about halfway on the pedicel, is reminiscent of that of M. mediovibex from
about the same area.

De Wilde — Myristicaceae (Myristica) 493

65. Myristica incredibilis W.J. de Wilde

Myristica incredibilis W.J. de Wilde, Blumea 40 (1995) 289. — Type: LAE 70971 (Katik) (male
flowering element), Papua New Guinea, Rossel I.

Tree 8 m. Twigs 3-4 mm diameter, late glabrescent, hairs conspicuous, brown, 0.5
mm long; older twigs and lenticels not seen. Leaves chartaceous, (ovate-)elliptic, 12-17
by 6-8.5 cm, base (shallowly) cordate, the blade + narrowed towards the blunt apex;
upper and lower surface of recently full grown leaf with dense brown hairs of mixed
sizes, 0.1—0.6 mm, upper surface glabrescent, olivaceous, lower surface with persistent
conspicuous indumentum, partly late glabrescent; not papillose; dots absent; midrib raised
above, lateral nerves 9-13 per side, at 45—70° to the midrib, flat or + sunken above, lines
of interaching and venation indistinct; petiole late glabrescent, 12-15 by 3 mm; leaf bud
12 by 3 mm, hairs 0.5(-1) mm. Inflorescences between the leaves, densely pubescent, as
in Knema but pedunculate; in male: peduncle 5—10 by 2-3 mm, ending in 2 (or 3) scar-
covered brachyblasts to 5 mm long, flowers in loose clusters of 5—10, buds not much
variable in size: female inflorescences not seen; flowers with hairs 0.5 mm. Male flow-
ers: pedicel 2—2.5 by 1—-1.5 mm, bracteole rounded, 1 mm, caducous; buds ellipsoid-
oblong, 8 by 4 mm, cleft c. 1/3, lobes 2.5—3 mm long, 0.3 mm thick. Androecium 6.5—7
mm; androphore 3 by | mm, at base with dense rufous hairs 0.5 mm long; synandrium
3.5(—4) by (1-)1.5 mm, thecae 14-16, sterile apex consisting of minute excrescences
0.1 mm corresponding with the anthers. Female flowers not seen. Fruits (doubtful, see
note 1) ovoid, 2 by 1.5 cm, apex rounded, base (broadly) rounded, hairs dense, | mm,
pericarp 1(-1.5) mm thick; seeds ellipsoid, 1.5 cm; fruiting pedicel not known.

Field-notes — Medium-sized tree; bole 5 m, dbh 14 cm. Bark grey-brown; underbark
brown; inner bark red; sap red; wood creamy straw. Leaves dark glossy green above,
light green and brown hairy below. Flowers creamy yellow. Fruits orange.

Distribution — Malesia: E Papua New Guinea (Louisiade Archipelago: Rossel L.);
known only from the type.

Habitat & Ecology — Hill forest; c. 10 m altitude; fl. & fr. Mar.

Notes — 1. The species is described from a single male flowering collection. Accord-
ing to the herbarium label the fruit is orange. The specimen in L has a separate single
mature fruit which, by necessity, was collected from another tree and hence may belong
to a different species. It cannot be matched with fruits of a known species; it approaches
that of M. chrysophylla, but is smaller and has shorter hairs.

2. Myristica incredibilis is, particularly in habit, reminiscent of M. schleinitzii, mainly
on account of the resembling leaves with cordate base and rounded apex. It has the gen-
eral features of male inflorescences and flowers in common with M. inopinata, but the
latter differs markedly in the distinct, acute, sterile apex of the synandrium. Myristica
schleinitzii differs in its (nearly) glabrous papillose leaves, smaller flowers, and the gla-
brous or finely stellate-hairy androphore. Possibly M. incredibilis 1s of a hybrid origin,
both species occurring in the same region.

66. Myristica iners Blume

Myristica iners Blume, Bijdr. (1826) 575; Rumphia | (1837) 184, t. 58; J. Sinclair, Gard. Bull. Sing.
16 (1958) 363, f. 30; 23 (1968) 177; Backer & Bakh. f., Fl. Java 1 (1964) 139; W.J. de Wilde,
Blumea 42 (1997) 168; Tree Fl. Sabah & Sarawak 3 (2000) 464. — Type: Blume s.n., W Java.

494 Flora Malesiana, Ser. I, Vol. 14 (2000)

Myristica sublanceolata Miq., Fl. Ind. Bat. 1, 2 (1858) 58. — Type: Horsfield s.n. (K sheet 683),
C Java.

Myristica vordermanii Warb., Mon. Myrist. (1897) 525, t. 14, ‘vordermanni’. — Type: Vorderman
s.n., Pulau Mendanau, near Belitung.

Myristica heritieriifolia Pierre ex Lecomte, Not. Syst. 1, 4 (1909) 99; Fl. Gén. Indo-Chine 5 (1914)
98. — Type: Pierre 5435, Indochina.

Myristica cumingii Warb. var. floribunda Airy Shaw, Kew Bull. 1939, no. 10 (1940) 539. — Type:
Richards 1615, Sarawak.

Tree 5—40 m. Twigs 1.5—3 mm diameter, early glabrescent, hairs greyish, less than 0.1
mm (in NE Borneo to 0.5 mm); older twigs (blackish) brown, + flaking, sporadically
with a few lenticels. Leaves membranous or chartaceous, rarely (at c. 1000 m) sub-
coriaceous, elliptic-oblong to oblong-lanceolate, 7-24 by 1.5—8(—10) cm, base rounded
or attenuate, apex acute-acuminate; upper surface olivaceous(-brown), lower surface
early glabrescent, hairs scale-like, less than 0.1 mm; not papillose; dots absent; midrib
slightly raised above, lateral nerves 1 1—16 per side, at (30—)45(—80)° to the midrib, above
flat to sunken, below rather flat or raised, lines of interarching and venation indistinct;
petiole 11—25 by (0.5—)1-2 mm; leaf bud 7-12 by 1-2 mm, hairs 0.2(—0.5) mm. Inflores-
cences between the leaves or below, paniculate, pedunculate, glabrescent; in male: 2-8
cm long, peduncle slender or stout, 2-30(—40, see note 1) mm long, branches to 20 mm,
central axis absent or up to 40 mm, with | or 2 subumbels of 5—15 flowers, buds of
various sizes; bracts 3-4 mm, caducous; female inflorescences: reduced, sometimes
+ sessile, 1-6-flowered; flowers glabrescent, hairs + woolly, 0.1—0.3 mm. Male flowers:
pedicel slender, 4-12 mm, bracteole 1.5—3 mm, subpersistent or caducous; buds ovoid
(-ellipsoid), 4-6(-7) by 2—4(-6) mm, cleft (1/4—)1/3-1/2, lobes membranous or perga-
mentaceous, 0.2—0.3 mm thick. Androecium 3(—6) mm; androphore cylindrical or some-
what swollen, about as broad as the synandrium, |.2—2(—3) mm long, with hairs 0.1—0.2
mm in the lower half; synandrium ellipsoid-lanceolate, 2-3 by 0.6—1 mm, thecae 10—16,
+ contiguous, sterile apex 0.1—0.3 mm, entire or shallowly lobed, glabrous. Female flow-
ers: pedicel 1—2 or 3—-7 mm long (see note 1); buds ovoid, 4—4.5 by 3.5—4 mm (in galled
flowers larger); ovary long-ovoid, 3 by (1—)1.5 mm, hairs dense 0.1 mm or less. Fruits
usually single, ellipsoid or ovoid-ellipsoid, 3.5—6(—8) by 2.5—3.5(—4.5) cm, glabrescent,
hairs dense, mealy, grey-brown, 0.2 mm; pericarp (4-)5—10 mm thick; seeds ellipsoid,
2—4(-5) by 1—2(—2.3) cm; fruiting pedicel either short and thick, 1-3 mm long (Java), or
slender, 5—15 mm long (Borneo) (see note 1). — Fig. 60d.

Field-notes — Tree with stilt-roots or buttresses, or without stilt-roots. Bark rough,
(deeply) fissured (strips 10 cm long, wavy, 3 mm thick) and flaky, dark brown, grey, and
blackish: outer bark 3—5 mm thick; inner bark 10 mm thick, soft, laminated, pinkish or
brown; slash wood white or pale, sometimes red streaked; exudate clear, reddish, some-
times copious. Flowers yellow, sometimes recorded as scented. Fruits subglobose or
ellipsoid, 4—8 cm long, green- or pale yellow, + glossy, early glabrescent, hairs minute,
brown; aril orange to red; seeds (glossy) brown or black.

Distribution — Cambodia, Thailand; Malesia: Sumatra, Peninsular Malaysia, Java,
Borneo, possibly Philippines (see note 4).

Habitat & Ecology — Forest on flats, slopes and ridges; evergreen forest; peat swamp
forest (Thailand); on sandstone and sandy-loam soils or tuff soil; O-1200 (in Aceh to
1800) m altitude; fl. & fr. throughout the year.

De Wilde — Myristicaceae (Myristica) 495

Notes — 1. Myristica iners is variable especially in the shape and texture of leaves,
the flower size, and the fruits. Leaves may be thinly chartaceous or subcoriaceous, espe-
cially in Java, with relatively distinct venation on the upper surface. Also in Java the
inflorescences are generally compact and with a short peduncle, occasionally subsessile.
In Borneo slender elongate and delicate (from sandy soils?) as well as stout, much-
branched inflorescences exist, e.g. in Hose 392 (Sarawak), SAN 1954] (Sabah), or
Kostermans 6789 (E Kalimantan), sometimes with + woolly hairs 0.3-0.5 mm long
(e.g. SAN 21323). Delicate specimens link up with the much related M. fallax from
Borneo. Some specimens from Sumatra (Palembang) are distinguishable by exceeding-
ly long peduncles of the male inflorescences, c. 40 mm, and these also link up with the
related M. fallax.

2. Fruits are variable in size; smallest in Java (c. 4 cm long), largest in Peninsular
Malaysia (7-8 cm long). Specimens from Java and the islands west of Sumatra gener-
ally have short fruiting pedicels (I-3 mm long), in most specimens from Borneo the
fruiting pedicels are long and slender, to 15 mm long.

3. Superficially resembling species are M. malaccensis, M. umbellata, and M. wyatt-
smithii; taxonomically related are M. corticata, M. fallax, and M. umbellata.

4. Wenzel 3537 (K, immature fr.), from the Philippines (Mindanao: Surigao), with a
comparatively long and slender fruiting pedicel, and long hairs on the leaf bud, agrees
with specimens from NE Borneo, but its locality is remarkable, far from the main distri-
butional area of the species.

67. Myristica ingens (Foreman) W.J. de Wilde

Myristica ingens (Foreman) W.J. de Wilde, Blumea 40 (1995) 290. — Myristica fatua Houtt. var.
ingens Foreman, Contr. Herb. Austral. no. 9 (1974) 37, f. 1; Handb. Fl. Papua New Guinea |
(1978) 193. — Type: Brass 31999, Papua New Guinea.

Tree 10-30 m. Twigs 4-8 mm diameter, glabrescent, hairs 0.1(—0.2) mm or less; older
twigs not cracking nor flaking; finely lenticellate. Leaves chartaceous or coriaceous,
(elliptic-)oblong, (22—)25—40 by 5—15 cm, base (short-cuneate or) broadly rounded, apex
acute-acuminate; upper surface olivaceous or brown, lower surface cinnamon, hairs in-
terwoven, scale-like, 0.1 mm; not papillose; dots absent; midrib flat above, lateral nerves
20-25(—30) per side, at (60—)70—80° to the midrib, lines of interarching and venation
indistinct; petiole 20-35 by 4-6 mm; leaf bud 20-30 by 3-4 mm, hairs 0.1 mm or less.
Inflorescences between the lower leaves, or below; of the Knema-type: a + woody scar-
covered simple (not forked) brachyblast to 5 mm long, sessile or to 5 mm pedunculate,
glabrescent; bracts 2-3 mm long, caducous; in male: with a cluster of 2-7 flowers, buds
of variable sizes: in female: 1—2(—4)-flowered; flowers with flocculose appressed dense
hairs 0.1—0.2(—0.3) mm. Male flowers (LAE 51941): hard leathery, stout, pedicel 10-12
by 2—3 mm, bracteole + boat-shaped, 6—7 mm long, caducous; buds (almost mature)
+ ellipsoid-obovoid, 8 by 5 mm, cleft c. 1/4, the lobes | mm thick. Androecium broad,
+ fusiform, 6 mm long; androphore | by | mm, almost glabrous (hairs scattered, pale,
less than 0.1 mm); synandrium 5 by 1.5(—2) mm, apex acute, thecae c. 20, sterile apex
0.1 mm or absent. Female flowers (NGF 38940, slightly immature): pedicel 5 by 4 mm,
bracteole broadly ovate, 4 by 5 mm; buds subglobose, 5 mm; ovary not seen. Fruit soli-
tary, subglobose or broadly ovoid(-ellipsoid), 7-10 by 6-8 cm, apex rounded or rather

496 Flora Malesiana, Ser. I, Vol. 14 (2000)

acute, hairs dense, mealy, yellow-brown or rust-coloured; pericarp 10-20 mm thick (with
thin woody endocarp 0.1—0.2 mm); seeds broadly ovoid(-subglobose), 3.5—5 cm, aril-
lobes reaching only to 1/5-1/3 of the seed; fruiting pedicel stout, 15-20 by 6-10 mm,
tomentose, bracteole scar at c. 1/3 below the apex.

Field-notes — Straight boled tree, sometimes emergent from the canopy; crown small
or narrow; branches horizontal; no or but slight buttresses. Bark grey-brown or dark
brown, vertically shallowly fissured, or peeling with small flakes; underbark orange-
brown, wood pale, rapidly staining on exposure; slash bark exuding copious (pale) red
sticky or watery sap. Leaves with brown or rufous indumentum below. Flowers brown-
ish, stamens cream. Fruits 9-11 by 7—8.5 cm, reddish brown or rust-coloured; aril small;
seeds glossy dark brown or black.

Distribution — Malesia: mountains of Central Papua New Guinea (Western, South-
ern, and Eastern Highlands Prov.).

Habitat & Ecology — Locally frequent in montane forest; sometimes emergent from
the canopy; degraded forest, forest remnants, and advanced regrowth; gullies in Lithocar-
pus—Castanopsis-forest; forest flats, near gullies in limestone area; 1200-1800 m alti-
tude; fl. Sept.—Oct.; fr. July—Jan.

Notes — 1. According to Foreman (1978) this is the Myristica with the largest fruits
in New Guinea. It is distinguishable by the stout habit of all parts, the large leathery
flowers, the large fruits with thick pericarp, and the aril-lobes only covering the basal
part of the seed.

2. Myristica ingens is the only species in the genus Myristica with a reduced aril; in
all other species the laciniae of the aril cover the seed over its entire length.

68. Myristica ingrata W.J. de Wilde

Myristica ingrata W.J. de Wilde, Blumea 40 (1995) 290, f. 2c; 43 (1998) 175. — Type: bb 30596,
Irian Jaya, Jayapura.

Tree 5-20 m. Twigs subterete or + flattened, 4-10 mm diameter, with wings 1-2 mm
high, early glabrescent, hairs 0.1 mm, hollow, and usually with ant-swellings with longi-
tudinal slit-like opening; older twigs with scattered lenticels. Leaves (sub)chartaceous
or subcoriaceous, (ob)ovate-oblong or oblong(-lanceolate), 15—40 by 5-14 cm, base
cuneate, rounded, or subcordate, apex acute-acuminate; upper surface olivaceous or
brown, lower surface grey(-brown), glabrescent, hairs scale-like, scattered, less than 0.1
mm, or indumentum dense, short (subsp. velata); conspicuously irregularly papillose;
dots absent; midrib faintly raised above, lateral nerves 20-25 per side, at 50—70° to the
midrib, flat or sunken, lines of interarching and venation not prominent; petiole 15 by
3—4 mm; leaf bud 10-15 by 2-3 mm, hairs 0.1 mm. /nflorescences of the Knema-type,
(sub)sessile scar-covered brachyblasts, subglabrous, to 10 mm long, simple or 2-branched;
in male: with subumbels of (5—)10—15 flowers, buds unequal in size; in female: 5—8-
flowered, sometimes bearing some fruits of a previous season; flowers with hairs 0.1
mm or less. Male flowers: pedicel (7—)10-12 by (0.6—)1 mm, bracteole + 3-topped, 2-3
by 2—3 mm, persistent; buds ovoid-oblong, 6.5—7.5 by 3.5—4 mm, apex blunt and faintly
triangular or not, base (broadly) rounded, cleft 1/4—1/3, lobes 1.5—2.5(-3) mm long,
0.2—0.3 mm thick. Androecium 5.5—6.5 mm; androphore (1—)1.5—2 by 0.5—0.8 mm, hairs
0.1 or less + in the lower half, sometimes only at the base; synandrium 4—5 by 0.8-1

De Wilde — Myristicaceae (Myristica) 497

mm, rather acute, thecae 12-14, sterile apex 0.2—0.3 mm. Female flowers: pedicel 3-5
mm, bracteole 2 mm, buds + ovoid, 5 by 3.5 mm, lobes 1.5 mm; ovary + ovoid, 3 mm,
finely pubescent. Fruits solitary or 2—4 per infructescence; (broadly) ovoid(-ellipsoid),
2.5-3.5 by (1.5—)2(-2.5) cm, apex + rounded and short-apiculate, hairs rust-coloured,
0.1(—0.2) mm; pericarp 2 mm thick; seeds ellipsoid, 2 cm; fruiting pedicel + stout, short
or long, 3-10 by 3-4 mm, with a few lenticels.

Distribution — Malesia: lowland and lower montane forest of Papua Barat and Papua
New Guinea; with two subspecies.

KEY TO THE SUBSPECIES

la. Leaves thinly chartaceous; lower surface (sub)glabrescent, hairs sparse, greyish, scale-
like, less than 0.1 mm. Fruits broadly ovoid(ellipsoid), 2.5 cmlong.............
PPPS Stes tI LE A) ARID hice gah ie, PSL gaan SL MES a. subsp. ingrata

b. Leaves subcoriaceous; lower surface with dense pale brown scale-like hairs, 0.1
iments: (long-)ellipsoid, 3: cmdJong... .2..6- ess eso b. subsp. velata

a. subsp. ingrata

Tree 5-20 m. Twigs brown, distinctly 2-winged, + fissured and lenticellate. Leaves
thinly chartaceous; lower surface (sub)glabrescent, hairs sparse, greyish, scale-like, less
than 0.1 mm. Fruits broadly (ovoid-)ellipsoid, 2.5 cm long. — Fig. 66c.

Field-notes — Slender tree. Bark brown, smooth or with sparse fine pustules or finely
fissured; blaze with watery clear sap turning brown or (chestnut-)red; wood cream, straw,
or orange; heartwood dark brown. Leaves grey, glaucous or silvery below. Flowers cream
or (yellow-)brown. Fruits brown, globose.

Distribution — Malesia: northern E Papua Barat, northern and southern Papua New
Guinea.

Habitat & Ecology — Lower montane forest, ridge forest, and lowland (alluvial)
forest; 20-1250 m altitude; fl. & fr. throughout the year. Twigs myrmecophilous, with
small black ants; sometimes with galls, resembling fruits.

Notes — 1. NGF 48095 (Foreman) has mature fruits among flowers in the same
inflorescence.

2. Myristica ingrata can be confused with M. subalulata but is clearly distinguishable
by the ovoid male perianth, persistent bracteole, and generally larger fruits with stout
fruiting pedicel. Male flowers of M. ingrata are rather similar to those of the related
M. hollrungit.

b. subsp. velata W. J. de Wilde

Myristica ingrata W.J. de Wilde subsp. velata W.J. de Wilde, Blumea 40 (1995) 292; 43 (1998)
175. — Type: Jacobs 9127, Papua New Guinea.

Tree 10-20 m. Twigs dark brown or blackish, faintly ridged, coarsely fissured,
lenticellate. Leaves subcoriaceous, greyish or grey-brown below, hairs persistent, inter-
woven, scale-like, 0.1 mm. Flowers not seen. Fruits 1-4 per infructescence, (ovoid-)
oblong, (2.5—)3-3.5 by 2 cm, with minute scurfy rust-coloured indumentum; fruiting
pedicel + stout, 3—5(—10?) by 3-5 mm.

498 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Branches spreading, inhabited by ants. Bark with sticky red exudate.
Fruits cinnamon or brownish orange.

Distribution — Malesia: submontane areas of Central Papua New Guinea (West Sepik
and Eastern Highlands Provy.); possibly also in West New Guinea (De Wilde l.c.).

Habitat & Ecology — Ridge forest and forest over limestone; 500-800 m altitude;
fr. June, Sept., Oct.

Notes — 1. Myristica ingrata subsp. velata may merit the status of a separate species;
its inclusion in M. ingrata is provisional, until more and (male) flowering material be-
comes available.

2. This subspecies may be close to M. velutina, a species without ant-swellings.

69. Myristica inopinata J. Sinclair

Myristica inopinata J. Sinclair, Gard. Bull. Sing. 23 (1968) 199, f. 14; Foreman, Handb. Fl. Papua
New Guinea 1 (1978) 200; W.J. de Wilde, Blumea 40 (1995) 292. — Type: Brass 28055, Papua
New Guinea, Tagula I.

Tree 8-20 m. Twigs 4—5 mm diameter, late glabrescent, hairs copious, 0.5—1 mm;
older twigs not seen. Leaves (thinly) chartaceous, (elliptic-)oblong, 12-24(-27) by 5-8
(10) cm, base (broadly) rounded, apex (subacute or) blunt; upper surface bright (oliva-
ceous-)brown, glabrescent, hairs dense, mealy, pale brown, 0.3 mm, lower surface late
glabrescent, hairs dense, much-branched, of mixed sizes, up to | mm long, persisting on
and near the base of the midrib; not papillose; dots absent; midrib raised above; lateral
nerves 12-18 per side, at 50-80° to the midrib, flat, lines of interarching and venation
indistinct; petiole 15-20 by 3(—4) mm, late glabrescent; leaf bud 10-15 by 3-4 mm,
hairs (0.5—)1 mm. Inflorescences as in Knema but pedunculate, between the leaves, pe-
duncle with persistent hairs 0.5—1 mm, or late glabrescent; in male: peduncle 5—10(—20)
mm long, ending in 1-3 short scar-covered flower-bearing brachyblasts, each inflores-
cence with 10—15 flowers, buds of various sizes; in female: peduncle up to 5 mm long,
|—3-flowered: flowers with dense hairs 0.5(—1) mm. Male flowers: pedicel 3-5 by 1-1.5
mm, bracteole | mm long, caducous, (sub)apical; buds ellipsoid-oblong, 9-10 by (4—)
4.5—5 mm, apex rounded, not angled (see note 1), cleft 1/4-1/3, lobes 2.5—3 mm long,
0.3 mm thick. Androecium slender, 8 mm; androphore 3 by 1(—1.5) mm, in basal (1/2—)
1/3 with dense setose yellow-brown hairs | mm; synandrium 3.5—4 by 1.5 mm, thecae
c. 12, sterile apex rather acute, | mm. Female flowers: pedicel 3 mm long, bracteole ca-
ducous, subapical; buds ovoid, 7(—9) by 5(—7) mm, lobes 3 mm long; ovary ovoid, 4-5
mm, hairs | mm. Fruits (submature) solitary or paired, subsessile, ovoid(-ellipsoid), 3
by 2-2.5 cm, apex (narrowly) rounded, hairs dense, rust-coloured, 2(—3) mm; pericarp
5—6 mm thick; seeds not known; fruiting pedicel 4-5 by 3 mm, pubescent.

Field-note — Subcanopy tree c. 20 m tall, stem 25 cm in diameter.

Distribution — Malesia: Papua New Guinea (Papuan Islands: known from 2 collec-
tions from Tagula I., at Rambuso).

Habitat & Ecology — Ridges in rain forest; 30-150 m altitude; fl. & fr. Sept.

Notes — 1. Sinclair (1.c.: 202) mentioned that the male perianth towards the apex is
triangular in cross section, to be seen when the dense indumentum is removed, but I
think that this observation rests on an artefact caused by drying; in fact, the bud is not or
only faintly angled at the apex.

De Wilde — Myristicaceae (Myristica) 499

2. Myristica inopinata is related to M. incredibilis and M. schleinitzii. It may be con-
fused with M. chrysophylla by the conspicuous indumentum of flowers and fruits, but
M. chrysophylla has (sub)sessile inflorescences, short pedicelled smaller flowers, and
fruits with a thin pericarp.

70. Myristica insipida R. Br.

Myristica insipida R.Br., Prod. Fl. Nov. Holl. ed. 1 (1810) 400; ed. 2 (1827) 256; Benth., Fl. Aus-
tral. 5 (1870) 281; Blake, Austral. J. Bot. 2, 1 (1954) 124; J. Sinclair, Gard. Bull. Sing. 23 (1968)
369, f. 61 (excl. syn. M. muelleri); Foreman, Handb. Fl. Papua New Guinea | (1978) 200; W.J.
de Wilde, Blumea 35 (1990) 245, f. 1: 6; 36 (1991) 187, f. 2; 40 (1995) 292; 42 (1997) 168. —
Myristica cimicifera R. Br. var. insipida (R. Br.) Warb., Mon. Myrist. (1897) 501. — Type: Brown
25, N Australia.

Myristica cimicifera Soland. ex R.Br., Prod. Fl. Nov. Holl. ed. 1 (1810) 400; ed. 2 (1827) 256;
Warb., Mon. Myrist. (1897) 499, t. 18 f. 1-6 (incl. var. typica). — Type: Banks & Solander s.n.,
1770, Queensland.

Myristica cimicifera R. Br. var. acutifolia Warb., Mon. Myrist. (1897) 502 (= var. kingti Warb., msc.,
nom. nud. in sched.). — Type: King s.n., W Australia.

Myristica macgregorii Warb., Mon. Myrist. (1897) 479. — Type: MacGregor 12, Papua New Guinea,
Barawara, Milne Bay Prov.

Shrub or tree (4—)6-25 m. Twigs 1-2 mm diameter, early glabrescent, hairs woolly,
0.2-0.5 mm; older twigs sometimes blackish, longitudinally cracked; lenticels incon-
spicuous or absent. Leaves membranous or chartaceous, elliptic-oblong or oblong-lan-
ceolate, 7—20(—24) by 2-6(—8) cm, base rounded or cuneate, apex acute( -acuminate), the
very tip often + blunt; upper surface olivaceous, lower surface greyish, indumentum
subpersistent or late glabrescent, hairs inconspicuous, scattered, 0.1 mm or less; papillae
distinct (specimens from Tanimbar Is.) or inconspicuous (part of New Guinea material);
dots absent; midrib above flat or sunken, lateral nerves 6—12 per side, at 45—80° to the
midrib, far apart, flat or sunken, (much) raised below, lines of interarching not distinct,
venation (not always) raised below; petiole 9-14 by 1-2 mm; leaf bud 7-12 by 1-2 mm,
hairs woolly, or appressed, 0.2—0.5 mm. Inflorescences between the leaves, axillary or
up to 2 mm supra-axillary, as in Knema: sessile or up to 2-4 mm pedunculate, 1—3-
branched scar-covered brachyblasts, 2—5(—7) mm long, short-pubescent, glabrescent; in
male: with subumbels of 5—15 flowers, buds of various sizes; bracts minute, caducous;
in female: sessile, few-flowered; flowers with woolly hairs (0.1—)0.2—0.4 mm, towards
the apex often darker coloured. Male flowers: pedicel 2—3(—4) mm (shorter than peri-
anth), bracteole (0.5—)1—1.5 mm, persistent or late caducous; buds ellipsoid-oblong, 4.5—
6 by 2-3 mm, cleft c. 1/4, lobes 0.2-0.3 mm thick. Androecium 3.5—4.5 mm; androphore
(1.5—)1.8—2 by 0.4—0.8(-1) mm, glabrous or (New Guinea) in the lower half with scat-
tered hairs 0.1—0.2 mm: synandrium 2-2.5 by 0.6-1.2 mm, thecae 12-16, sterile apex
+ absent or irregularly lobulate by short processes from the anthers, or rather blunt to
subacute, (0.1—)0.2-0.3 mm. Female flowers (from Specht 666): pedicel 0.5(—1) mm;
buds ovoid, 4 by 3 mm, cleft c. 1/4; ovary ovoid, 3 by 2 mm, hairs 0.3—0.4 mm. Fruits |
or 2 per infructescence, subsessile, ovoid-ellipsoid or ellipsoid-oblong, 2.5—3.8 by 1.5—
2 cm, apex acute or with short beak 1-2.5 mm, base rounded or cuneate or + narrowed,
hairs dense, pale or dark brown, woolly, 0.3—1 mm, glabrescent, but hairs remaining in

500 Flora Malesiana, Ser. I, Vol. 14 (2000)

3

Me
De
oe

NIV EAA OC

4

Erratum: Flora Malesiana vol. 14 — The drawing on page 500 (Myristica markgraviana) should
be replaced by the figure below (= Fig. 78. Myristica insipida R. Br.).

De Wilde — Myristicaceae (Myristica) 501

the depressions left after drying and towards the base of the fruit; pericarp 2-3 mm
thick, seeds ellipsoid-oblong, 2—2.5 cm, bright brown; fruiting pedicel 1-3 mm. — Fig.
64g, 78.

Field-notes — Shrub or medium-sized (understorey) tree. Bark (grey-)brown or black-
ish, usually shallowly fissured or finely scaly; inner bark 4-12 mm thick, pinkish, sap
reddish; sapwood straw or light brown, undefined; true wood light brown or pale red-
brown, rather soft. Leaves pale green or glaucous below. Flowers yellow green, pale
yellow, or brown-yellow, androphore white, anthers light brown, pollen cream. Fruits
(greenish) brown, golden, or rust-coloured; seeds brown.

Distribution — N Australia (N Western Australia, E to N Queensland); Malesia: SE
Moluccas (Tanimbar Is.), S New Guinea.

Habitat & Ecology — Coastal or cliff forest, rain forest behind dunes, deciduous vine
thickets, monsoon forest and scrub, riparian, gallery, or gully forest (in gorges in sand-
stone, Australia), fringing forest of savanna woodland; generally not too far from the
coast; sandstone, sandy soil, lateritic or coral sand; locally common; 0—200 m altitude;
fl. & fr. throughout the year.

Notes — 1. In M. insipida three groups can be recognized: 1) Material from the
Tanimbar Islands, characterized by well-developed papillae on the lower leaf surface
(lens!), glabrous androphore, and rather elongate fruits with dark rust-coloured com-
paratively short hairs, up to 0.5 mm long only. 2) Material from New Guinea with scat-
tered pale brown hairs on the lower half of the androphore; some collections with the
venation on the lower leaf surface either raised and distinct or + flat and indistinct (e.g.
Brass 6430), and the bracteoles persistent or late caducous (e.g. Brass 6505). 3) Mate-
rial from northern Australia, containing the type specimens of M. insipida and most of
its synonyms, characterized by a distinct venation of the lower leaf surface, persistent
bracteole, glabrous androphore, and fruits with pale or bright brown hairs to | mm long.

2. Branderhorst 294 (L) (two separate branches, one with male flowers, one with
fruits) from S Papua Barat, agrees with M. insipida vegetatively and also in the shape of
the male flowers with thickish woolly indumentum, but differs markedly in its androphore
with dense conspicuous brown hairs 0.2—0.4 mm. UPNG 645A ( Frodin c.s.) differs in
male inflorescences with the peduncles up to 4 mm long.

3. Myristica insipida is characterized by subpersistent + scattered hairs and usually
distinct papillation on the lower leaf surface, widely spaced lateral nerves, woolly-haired
flowers, usually glabrous androphore (partly pubescent in part of the material from New
Guinea), and fruits with woolly hairs to 1 mm. The species is related to M. globosa
subsp. muelleri from the Pacific (Solomon Islands); the basionym M. muelleri was in-
cluded by Sinclair (1.c.) in M. insipida.

Fig. 78. Myristica insipida R. Br. a. Leafy twig with male inflorescences; b. male bud with persist-
ent bracteole; c. ditto, opened, showing androecium; d. twig with female inflorescences; e. female
bud; f. ditto, opened, showing pistil; g. twig with infructescences, each bearing one or two fruits;
h. dry fruit, opened, showing arillate seed (a—c: Telford & Wringly 7825 (Australia); d—f: Specht
666 (Australia); g, h: Darbyshire 710 (Papua New Guinea)]. — Scale bar for a, d, g = 2 cm; for b, c,
es Ly inm: tor h'=" lem.

502 Flora Malesiana, Ser. I, Vol. 14 (2000)

71. Myristica inundata W. J. de Wilde

Myristica inundata W.J. de Wilde, Blumea 40 (1995) 292. — Type: NGF 18331] Streimann & Lelean,
Papua New Guinea, Western Prov.

Tree 25—30 m. Twigs faintly angular, 2(—3) mm diameter, sometimes blackish, with
small, scattered, pale lenticels, early glabrescent, hairs less than 0.1 mm; older twigs
(blackish) brown, + fissured, densely lenticellate. Leaves (thickly) chartaceous, (ellip-
tic-)oblong, 10-16 by 4—6 cm, base short-cuneate or narrowly rounded, apex (short)
acute-acuminate; upper surface brown, lower surface brown-grey, late glabrescent, hairs
dense, scale-like, less than 0.1 mm; indistinctly papillose; dots absent; midrib + flat
above, lateral nerves 10-15 per side, at (SO0—)60—70° to the midrib, flat and inconspicu-
ous above, little raised and + contrasting in colour below, lines of interarching and vena-
tion faint; petiole 20-25 by 2.5 mm; leaf bud 12 by 2.5 mm, hairs greyish, scale-like,
less than 0.1 mm. Jnflorescences between the lower leaves and below, of the Knema-
type; in male: a pubescent scar-covered brachyblast, 1-2 mm, sessile or to 2 mm pedun-
culate, with a cluster of 10-15 flowers, buds of variable sizes; bracts not seen; flowers
with bright yellowish brown hairs 0.3 mm. Male flowers: pedicel 8-10 by 0.5(—1) mm,
bracteole 4 by 4 mm, (sub)persistent; buds ovoid-ellipsoid, 4.5 by 3.5 mm, cleft c. 1/3,
lobes 1.5 mm long, 0.2—0.3 mm thick. Androecium 4 mm; androphore I—1.5 by | mm,
with few hairs at the base, pale, 0.1 mm; synandrium 3 by 1—1.2 mm, thecae c. 16,
subcontiguous, sterile apex 0.2 mm. Female inflorescences and flowers, and fruits not
seen.

Field-notes —Tree c. 25 m tall, bole 20 m, straight, dbh 40 cm, with stilt-roots; crown
open, spreading. Leaves dull dark green above, bronze below. Flower buds brownish
yellow.

Distribution — Malesia: Papua New Guinea (Western Prov., Kiunga subprov., Tuide-
masuk Road); known only from the type.

Habitat & Ecology — Seasonally inundated swamp forest; at about 25 m altitude;
fl. Sept.

Notes — |. Myristica inundata is similar to M. atrocorticata from the same area; the
latter is distinguishable by narrower leaves, a male bracteole that is much shorter than
the perianth, shorter indumentum, and prop-roots. Myristica inundata was collected in
seasonally inundated swamp forest, and recorded as having stilt-roots. Prop-roots have
been recorded for M. atrocorticata.

2. Because of the (sub)persistent, short and dense indumentum of the lower leaf sur-
face, this species may be confused with M. inutilis. The male flowers have some remi-
niscence with those of M. hollrungii, a species not closely related.

72. Myristica inutilis Rich. ex A. Gray

Myristica inutilis Rich. ex A. Gray in Wilkes, U.S. Explor. Exped. 1 (1854) 34; Warb., Monogr.
Myrist. (1987) 481; W.J. de Wilde, Blumea 38 (1994) 357, f. 3, 4; 40 (1995) 293.

This species reaches far into the Pacific; there are three subspecies of which one,
subsp. papuana, occurs in the Malesian area.

De Wilde — Myristicaceae (Myristica) 503

subsp. papuana (Markgr.) W.J. de Wilde

Myristica inutilis Rich. ex A. Gray subsp. papuana (Markgr.) W.J. de Wilde, Blumea 40 (1995)
293: 42 (1997) 169: 43 (1998) 180. — Myristica fatua Houtt. var. papuana Markgr., J. Arnold
Arbor. 10 (1929) 77, 214; Bot. Jahrb. Syst. 67 (1935) 160, p.p.; J. Sinclair, Gard. Bull. Sing. 23
(1968) 294, p.p., excl. f. 37 which depicts M. subcordata var. morindiifolia; Foreman, Contr.
Herb. Austral. no. 9 (1974) 39; Handb. Fl. Papua New Guinea | (1978) 194, p.p. — Type: Brass
1070, Papua New Guinea.

Myristica finschii Warb., Mon. Myrist. (1897) 534, t. 19, p.p. — Type: Warburg 20715 (B, lost),
Papua New Guinea, ‘Sattelberg’.

Myristica sericea Warb., Mon. Myrist. (1897) 521. — Type: Warburg 20722 (B, lost), New Ireland.

Myristica wallacea Warb., Mon. Myrist. (1897) 530, t. 19. — Syntypes: Beccari FI acc. nos. 7707,
A, B, n.v.; Warburg 20721 (B, lost), Aru Islands.

Tree 7—-25(—40) m. Twigs (1.5—)2—3(—3.5) mm (in var. foremaniana Cc. 5 mm) diam-
eter, faintly angled, glabrescent, hairs 0.1 mm or less; older twigs not flaking, finely len-
ticellate. Leaves (thickly) chartaceous, elliptic-oblong to oblong-lanceolate, 12-30 by
3.5-7.5(—9) cm, base attenuate or narrowly rounded, apex acute-acuminate; upper sur-
face olivaceous or brown, indumentum of lower surface persistent or sometimes late gla-
brescent, hairs pale (yellowish) brown or greyish, densely interwoven, scale-like, 0.1
(—0.2) mm; not papillose nor dotted; midrib above + flat, lateral nerves 18-25 per side,
at 40-75° to the midrib, lines of interarching and venation indistinct; petiole 10-20 by
1.5-3 mm; leaf bud 10-15 by 1.5—2.5 mm, hairs woolly, 0.1(—0.2) mm. Inflorescences
between the leaves or below, (supra-)axillary, of the Knema-type: simple or 2- (or 3-)
branched, woody, wart-like or vermiform, scar-covered brachyblasts to 15 mm long, ses-
sile or to 3(—5?) mm pedunculate, short pubescent, glabrescent; in male: with subumbels
of 5—10(—20) flowers, buds of various sizes; bracts minute; in female fewer flowered;
flowers with hairs 0.1-0.5 mm. Male flowers: pedicel 2-5 by 0.5—0.7 mm, bracteole
acute or blunt, (1—)1.5—2(—2.5) mm, persistent; buds ellipsoid-oblong, 34.5 by 1.5—2
mm, cleft (1/4—)1/3(—c. 1/2), lobes 1.5—2 mm long, 0.2—0.3 mm thick. Androecium 3—
4.2 mm; androphore slender, 1.5—2 mm long, glabrous, or with few hairs at base or
almost completely minutely hairy; synandrium 2—2.4 by 0.4-0.6 mm, thecae ( 6—)10,
+ contiguous, sterile apex 0.1-0.2 mm, blunt. Female flowers: pedicel 1-2 mm long,
bracteole 2 mm long; buds ovoid, 2.5 mm long; ovary ovoid, 2 mm, densely fine pubes-
cent. Infructescence with 1-5 fruits in a cluster. Fruits broadly obovoid(-ellipsoid), apex
rounded or pointed, variable in size, 2.5-5(—6) by 1.8-3.5 cm, hairs grey-brown or rusty,
0.1—0.5(—1) mm; pericarp (1—)2—5 mm (8-10 mm in var. foremaniana) thick; seeds el-
lipsoid, 2—2.5 cm; fruiting pedicel 1-3 mm long, glabrescent, bracteole scar + apical.

Field-notes — Slender tree, bole straight, with or without buttresses, sometimes with
(flattened) stilt-roots. Bark smooth or rough, fissured, rarely flaking, brown, grey, or
black: slash bark (blaze) red; wood white, pinkish, or orange; no heartwood, central rot
10 cm diameter. Leaves bronze, cinnamon, or orange (tinged) below. Flowers golden,
rust-coloured, cream, or greenish inside. Fruits usually (orange-)brown or ferruginous.
Seeds brown to almost black.

Distribution — Malesia: Moluccas (Aru Islands); throughout New Guinea, including
Papuan Islands, Bismarck Archipelago.

Habitat & Ecology — Flat, colline, or ridge forest on a variety of soil types; 0-800 m
altitude; fl. & fr. throughout the year.

504 Flora Malesiana, Ser. I, Vol. 14 (2000)

Note — 1. Subspecies papuana is the westernmost subspecies of the widespread M.
inutilis; two more subspecies occur in the Pacific, subsp. inutilis (Samoan Islands) and
subsp. platyphylla (Samoan Islands and New Hebrides), the latter erroneously men-
tioned for New Guinea by De Wilde (1994). The three subspecies can be separated on
characters of the male flowers. Fruits are strongly variable in size, shape and indumentum,
and cannot be used to corroborate the subspecies. Variety foremaniana is accepted within
subsp. papuana on account of a remarkably distinct collection, with large leaves and
large globose fruit.

2. Myristica inutilis is related to M. fatua and M. subcordata;, for delimitation see the
key to the species.

KEY TO THE VARIETIES

la. Leaves 12-24 by 3.5—7 cm. Fruits obovoid-ellipsoid, 2.5—5(—6) cm long, hairs 0.1—

05C1)immy pencarp (I=) 2—s mim thick. 2% a ae et: Se tee a. var. papuana
b. Leaves larger, up to 30 by 9 cm. Fruits subglobose, 5—5.5 by 5 cm, hairs minute,
meaty. O02) mm-iperearp 10mm thick "Yo si rs a b. var. foremaniana

a. Var. papuana

Distribution — Widespread, as the subspecies.

Note — Highly variable in leaf size, some flower characters, and fruits. The androphore
is usually hairy towards the base, but in some (not all) specimens from the Bismarck
Archipelago (almost) glabrous. Fruits vary from 2.5 to 5 cm in length, with 0.1—0.5 mm
long hairs; Ridsdale NGF 33918 is one of the extremes, with fruits 5 by 3.5 cm, rough
hairs, 0.5 mm long, and pericarp 3—5 mm thick.

b. var. foremaniana W.J. de Wilde

Myristica inutilis Rich. ex A. Gray subsp. papuana (Markgr.) W.J. de Wilde var. foremaniana W.J.
de Wilde, Blumea 40 (1995) 294. — Type: LAE 52230 (Foreman), Papua New Guinea.

Tree 18 m. Twigs 5 mm diameter. Leaves up to 30 by 9 cm, densely grey-cinnamon
felty pubescent below. Fruits subsessile, subglobose, not wrinkled, 5—5.5 by 5 cm, hairs
light dull brown, mealy, 0.1(—0.2) mm; pericarp 10 mm thick; fruiting pedicel 2-3 mm
long.

Field-notes —Bark slightly fissured; underbark reddish; sap slight, reddish; wood
light brown. Leaves dark green above, covered with cinnamon-brown scale-like hairs
below. Fruits brown.

Distribution — Malesia: Papua New Guinea (Madang Prov., S of Dumpu); known
only from the type.

Habitat & Ecology — Forest of saddle between two ridges; 730 m altitude; fr. Mar.

Note — Known only in fruit. When flowers become available this taxon may appear
to represent a species of its own, characterized by the conspicuous (sub)globose sessile
fruits. It can be confused with M. sphaerosperma, which differs among others in a much
longer fruiting pedicel, and also with M. pachycarpidia which has a much longer fruit-
ing pedicel and a (sub)glabrous lower leaf surface.

De Wilde — Myristicaceae (Myristica) 505

73. Myristica kalkmanii W. J. de Wilde

Myristica kalkmanii W.J. de Wilde, Blumea 40 (1995) 294. — Type: BW 8522 (Kalkman), SE Irian
Jaya.

Tree 15(-20) m. Twigs sometimes + angular, 2.5—5 mm diameter, early glabrescent,
hairs 0.1(—0.2) mm or less; older twigs blackish, longitudinally cracking and flaking;
lenticels absent. Leaves (thinly) chartaceous, oblong-lanceolate, 15-32 by 5—11 cm, base
cuneate or (narrowly) rounded, apex acute-acuminate; upper surface brown, lower sur-
face grey(-brown), glabrous; papillose; dots absent; midrib nearly flat above, lateral
nerves 14-20 per side, at 50—70° to the midrib, flat or sunken, lines of interarching
distinct or not, venation indistinct; petioles 18-38 by 2.5—4.5 mm, leaf bud 10(—15) by
4(—5) mm, + longitudinally cracked on drying, hairs 0.1—0.2 mm. Inflorescences be-
tween the lower leaves or (just) below, of the Knema-type: sessile, simple or 2- or 3-
branched scar-covered brachyblasts; bracts not seen; in female (from infructescences):
branches 5—8 mm long, glabrescent, hairs yellow-rufous, 0.2—0.3 mm. Male and female
flowers not seen. Fruits 1(-3?) per infructescence, + ellipsoid-fusiform, 3.5—4 by (2-)
2.5-3 cm, apex narrowly rounded or somewhat acute, base rounded or slightly narrowed,
glabrescent, hairs mealy or powdery, rust-coloured, 0.3(—0.5) mm; pericarp 5 mm thick,
light (yellowish or orange-)brown; seeds ellipsoid, 2 by 1.2-1.3 cm, the aril deeply im-
pressed into the seed; fruiting pedicel + slender, 5—8 by 2-3 mm, yellow-brown pubes-
cent, glabrescent, bracteole apparently small, scar apical, or up to 1.5 mm below the
apex.

Field-notes — Small, straight, slender tree, bole 10-15 m, dbh 10-18 cm, without or
with stilt-roots up to 20 cm high; branches few; crown small, open. Leaves light blue-
green below. Fruits 4 by 3 cm; pericarp and pulp yellowish white; seeds brown; imma-
ture fruits red brown or light rusty brown.

Distribution — Malesia: southern (SE Papua Barat; Papua New Guinea: Western
Prov.) and possibly (see note 2) northern New Guinea (W Sepik Prov.).

Habitat & Ecology — Lowland forest on clayey soil, on slight ridge, or on sloping
ground; 25-120 m altitude; fr. Mar., Sept.

Notes — 1. Myristica kalkmanii is close to M. sulcata, with similar dark brown dry-
ing colour of the leaves. Myristica sulcata differs in its non-papillose lower leaf surface,
thinly ridged (lined) or angular twigs, and subglobose fruits. Flowers of M. kalkmaniti
are not known. The species somewhat resembles M. ornata, with which it has the mark-
edly papillose lower leaf surface in common, but M. ornata differs in its fainter nerves,
stalked infructescences, and larger fruits.

2. NGF 3931 (Womersley) (K, in fr.) from W Sepik probably belongs here; the speci-
men agrees with the material from southern New Guinea in twig structure, leaf shape,
and fruits, but it differs in the non-papillose lower leaf surface.

74. Myristica kjellbergii W. J. de Wilde
Myristica kjellbergii W.J. de Wilde, Blumea 42 (1997) 169. — Type: Kjellberg 2962, Sulawesi.

Tree 4-15 m. Twigs 2.5—3(—5) mm diameter, late glabrescent, hairs 0.1 mm; older
twigs coarsely striate, with scattered not contrasting lenticels. Leaves chartaceous or
subcoriaceous, broadly ellipsoid-oblong, 15-24 by 5—10 cm, base attenuate, apex short

506 Flora Malesiana, Ser. I, Vol. 14 (2000)

acute-acuminate; upper surface brown, lower surface greyish, glabrescent, hairs inter-
woven, soft, stellate, 0.1-0.2 mm, the larger hairs leaving point-like scars; distinctly
papillose or not; dots absent; midrib almost flat above, lateral nerves 10—16 per side, at
45—60° to the midrib, flat or sunken, indistinct above, on lower surface purple-brown
contrasting, lines of interarching and venation indistinct; petiole 10-15 by 3—4 mm; leaf
bud 15 by 3 mm, hairs woolly, 0.2-0.5 mm. Female inflorescences between the leaves
and below, of the Knema-type: sessile wart-like brachyblasts, 2-3 mm long, finely woolly
pubescent, with 2—5 flowers, buds almost equal in size. Male flowers not known. Fe-
male flowers (Meijer 9363) with dense bright golden-rusty hairs 0.2 mm, pedicel 0.5
mm, bracteole ovate, 3 mm, persistent; buds ovoid, narrowed towards the blunt or sub-
acute apex, 5.5 by 3.5(—4) mm, cleft into 3 or 4 lobes for c. 1/3 to nearly 1/2, lobes 0.3
mm thick; ovary ovoid, 3 by 2 mm, hairs golden brown 0.2(—0.3) mm. Fruits in sessile
clusters of 2 per infructescence, subsessile, (sub)globose, 2—2.2 by 2 cm, apex some-
times with beak 1.5(—2) mm, hairs dense, 0.1—0.2 mm; pericarp woody-pergamentaceous,
1-2 mm thick; seeds ellipsoid, 16-18 mm, with thick aril; fruiting pedicel 1-1.5 mm.

Field-notes — Low tree, branches horizontal, 3-5 from the same spot. Bark dark
grey-brown, finely flaking. Leaves silvery white below. Fruits globose, 2.5 cm diameter;
yellow or brown felty or rusty brown.

Distribution — Malesia: N, C and SW Sulawesi, known from a few collections. De-
viating collections from SW Peninsula, with rather small subglobose fruits, are discussed
by De Wilde (I. c.: 170).

Habitat & Ecology — Primary rain forest, ridges at 500-800 m altitude; female fl.
Apr., Aug.; fr. Aug.—Dec., Mar.

Note — Myristica kjellbergii is characterized by its pale, greyish or whitish lower
leaf surfaces, with conspicuously contrasting brown nerves, a feature shared with M.
koordersii, M. impressa, and M. impressinervia; it differs from the first two species in
the extremely faint nerves on the upper leaf surface, the papillae on the lower surface
(always?), and smaller fruits with thin pericarp. Myristica impressinervia, known only
from male flowers, has a much more tiny habit, smaller and thinner leaves, lower sur-
face apparently not papillose, and distinctly pedunculate inflorescences.

75. Myristica koordersii Warb.

Myristica koordersii Warb., Mon. Myrist. (1897) 619; Koord., Meded. Lands Pl. Tuin 19 (1898)
572: J. Sinclair, Gard. Bull. Sing. 23 (1968) 262, p.p., for the specimens from NE Sulawesi
(Minahasa) only, excl. f. 28 (= Myristica impressa); W.J. de Wilde, Blumea 42 (1997) 170. —
Syntypes: Koorders 18128 (L, lecto, here designated), 18129 (immature fr.), 18/44 (BO, n.v.),
all NE Sulawesi.

Tree 20-50 m. Twigs 1.5—3 mm diameter, early glabrescent, hairs 0.2 mm; older twigs
sometimes cracking; lenticels present but inconspicuous. Leaves (thinly) chartaceous,
(elliptic-)oblong or lanceolate, 13—20 by 3—5(—6) cm, apex acute or shortly acute-acu-
minate, base attenuate, upper surface olivaceous, lower surface pale greyish, glabrescent,
hairs + widely spaced, appressed, of mixed sizes, 0.1(—0.3) mm, the stouter ones leaving
dark points; papillae not obvious but possibly present (causing the whitish surface); dots
absent: midrib above slightly raised, lateral nerves 13-16 per side, at 45—60° to the

De Wilde — Myristicaceae (Myristica) 507

midrib, faint and flat above, lines of interarching and venation indistinct (the latter some-
times distinct towards the margin); petiole 15-30 by 2-3 mm; leaf bud 12-15 by 1.5-3
mm, hairs (0.1—)0.2-0.3 mm. Inflorescences, male and mature female flowers not seen,
infructescences of the Knema-type: peduncle up to 2 mm long, hairs dense, shaggy,
yellowish brown 0.5—1 mm. Female flowers (Foreman 262, immature) with dense shaggy
hairs 0.5—1 mm, pedicel at least 3 mm long, bracteole caducous. Fruits 1 or 2 per
infructescence, broadly ovoid-ellipsoid, 3—-3.5 by 2.5—3 cm, apex with | mm long style
remnant, hairs mealy, 0.1 mm, and with longer sparse hairs towards the base of the fruit;
pericarp 6-8 mm thick; seeds not seen; fruiting pedicel 6-10 mm long, slender,
subglabrescent or with persistent shaggy hairs (0.5—)1 mm, bracteole scar 1-2 mm be-
low the apex.

Field-notes — Solitary emergent tree. Prop-roots few, covering an area of 3 m in
diameter. Clear bole very straight, 20 m. Outer bark 0.3 mm thick, dark brown, rather
fissured, peeling off strongly; inner bark 8 mm thick, on section yellowish, out + red-
dish, with little watery red exudate. Sapwood pale yellowish tinged red. Leaves glau-
cous below. Fruits green (somewhat immature), with golden brown indumentum.

Distribution — Malesia: Sulawesi (eastern part of NE arm, Minahasa).

Habitat & Ecology — Hillside forest; deep sandy-clayey volcanic soil; 500-1000 m
altitude; fr. Jan., Feb., June, Oct.

Notes — 1. Sinclair accepted M. koordersii in a wider sense than is done here, includ-
ing specimens now transferred to M. impressa (type lost). The latter occurs almost through-
out Sulawesi (except the Minahasa), but was by Sinclair (1.c.: 235, 481) considered to
be a dubious species based on a possible atypical specimen of M. koordersii.

2. Myristica koordersii resembles M. impressa and M. kjellbergit; for differences see
the keys.

76. Myristica laevifolia W. J. de Wilde

Myristica laevifolia W.J. de Wilde, Blumea 40 (1995) 295, f. 3f. — Type: NGF 23957 (Streimann),
Papua New Guinea.

Tree 6-30 m. Twigs 1-2 mm diameter, glabrescent, hairs greyish, less than 0.1 mm;
older twigs smooth with indistinct lenticels. Leaves (thinly) coriaceous, (ovate-)elliptic
or oblong-lanceolate, 6-15 by 2—6 cm, base + attenuate, apex acute-acuminate; upper
surface olivaceous or brown, lower surface light brown, glabrescent, hairs sparse, grey-
ish, less than 0.1 mm; not papillose; dots absent; midrib raised above; lateral nerves 8—
14 per side, at 45—70° to the midrib, flat or sunken and indistinct at both surfaces, not
really contrasting in colour, lines of interarching and venation faint on both surfaces;
petioles 10-15 by 1.5-2 mm; leaf bud 8-12 by 1.5—2 mm, hairs less than 0.1 mm. Jnflo-
rescences between the leaves or below, largely of the Knema-type: a sessile sometimes
up to 2.5 mm pedunculate, scar-covered brachyblast to 7 mm long, glabrescent; in male:
with 2-6 (rarely more) flowers, buds somewhat variable in size; in female: with 1-3
buds about equal in size; flowers partly glabrescent, hairs scattered, 0.1 mm or less, and
appearing glabrous. Male flowers: pedicel slender, (4-)5—8 mm, bracteole 0.5 mm or
less, caducous, apical or up to 2 mm below; buds elliptic-oblong, 6(—6.5) by 2-2.2(-3)
mm, apex broadly rounded, base rounded or attenuate, cleft c. 1/3, lobes 2 mm long,

508 Flora Malesiana, Ser. I, Vol. 14 (2000)

0.2—0.3 mm thick. Androecium 5—6.5 mm; androphore 2—2.3 mm long, glabrous or hairs
pale, less than 0.1 mm at the base; synandrium 2.2—3 by 0.7—1.5 mm, thecae 12-14,
sterile apex 0.3-0.5 mm. Female flowers rather stout, pedicel 9-15 by 1 mm, bracteole
scar 2-3 mm below the apex; buds ovoid-ellipsoid, S—5.5 by 3 mm, cleft 1/4(-1/3),
lobes 1.5-1.7 mm; ovary ovoid-oblong, including stigma 4—4.5 by 2 mm, hairs less than
0.1 mm, or partly glabrous. Fruits solitary or 2 or 3, ellipsoid-oblong, + fusiform, 4—5.5
by (1.7—)2-2.5 cm, apex narrowly rounded or subacute, base narrowed into a pseudostalk
up to 3 mm, glabrescent, hairs scurfy, 0.1 mm or less; pericarp 2-3 mm thick; seeds
ellipsoid, 2—2.5 cm; fruiting pedicel + stout, 10-15(—18) mm long, 2mm thick, bracteole
scar 2—5 mm below the apex. — Fig. 62f, 63f.

Field-notes — Tree without buttresses, straight-boled or crooked. Bark smooth, rough,
or fissured, not flaking off, grey-brown or dark brown; underbark pinkish or salmon;
exudate watery or sticky, red; wood cream or straw, staining brown on exposure. Leaves
dull green below. Flowers cream, yellowish, or whitish. Fruits brown, orange, or green
with brown tinge.

Distribution — Malesia: Papua New Guinea (Sepik, Morobe, Eastern Highlands,
Central, and Milne Bay Prov.).

Habitat & Ecology — Mid-mountain mixed rain forest; oak-forest, Castanopsis-domi-
nated forest, mixed forest with Kibara, Lithocarpus (Pasania), Elaeocarpus, Elmerrillia;
1000-1800 m altitude; fl. & fr. throughout the year.

Note — This species is close to M. crassipes subsp. altemontana and M. tubiflora (for
discussion, see De Wilde I.c.: 296).

77. Myristica laevis W.J. de Wilde

Myristica laevis W.J. de Wilde, Blumea 42 (1997) 170. — Type: PNH 118091 (Gutierrez), Philip-
pines, Samar.

Tree 4-20 m. Twigs 2-3 mm diameter, early glabrescent, hairs 0.1 mm or less; older
twigs brown or rather pale; lenticels indistinct or absent. Leaves thinly chartaceous,
lanceolate, 9-22 by 2—4.5 cm, base cuneate or (sub)rounded, apex acute(-acuminate) or
+ blunt; upper surface olivaceous or brown, lower surface pale grey-brown, early gla-
brescent; densely papillose; dots absent; midrib above flat, much raised below, yellow-
brown, lateral nerves 20-30 per side, at 60-80° to the midrib, flat, hardly visible above,
slightly raised below, lines of interarching and venation indistinct; petiole 10-20 by 1.5-
2.5 mm; leaf bud 7-16 by 3-4 mm, hairs 0.1—0.5 mm. Inflorescences between the leaves
and below, pedunculate, ending in a subumbel of flowers (but brachyblast absent); bracts
minute, caducous: in male (PNH 1/809], + immature): peduncle 5-8 mm, subumbel
with 3—6(—10) flowers, buds of various sizes; female similar, peduncle up to 25 mm,
fewer flowered; inflorescences and flowers with rusty hairs 0.2—0.3(—0.5) mm. Male
flowers: pedicel rather stout, 5-7 mm, bracteole 4 mm long, caducous; buds woody-
coriaceous, ovoid or ellipsoid(-oblong), apex narrowly rounded, 6—-6.5 by 4(—4.5) mm,
cleft to over 1/2 (possibly nearly to the base), lobes 0.8 mm thick. Androecium broadly
cylindrical, 2.5-3 mm; androphore 0.2 mm long, and broad, like a disc at the base of the
synandrium, with sparse erect hairs 0.1—0.2 mm; synandrium 2-2.3 by | mm, thecae
16-18, sterile apex blunt, 0.2—0.4(-0.5) mm. Female flowers not seen; pedicel variable
in length (see the subspecies). Infructescences: peduncle conspicuously longitudinally

De Wilde — Myristicaceae (Myristica) 509

cracked. Fruits single or up to 3, subglobose or ellipsoid, (3.6—)4—5.5 cm long, hairs
brown, scurfy, 0.1—0.2 mm; pericarp 7-10 mm thick, seeds not seen; fruiting pedicel
glabrous.

Distribution — Malesia: E Philippines, with two subspecies.

Note — Myristica laevis is distinguishable from all other Philippine Myristica spe-
cies by the glabrous, lanceolate, faintly but many-nerved leaves, large + coriaceous male
flowers with almost sessile synandrium, and fruits with thick pericarp. Dry leaves are
exceedingly brittle. With M. umbellata it has the distinct papillose lower leaf surface
and the pedunculate, somewhat umbellate non-branched inflorescences in common.

KEY TO THE SUBSPECIES

la. Peduncle of infructescence 3-10 mm long; fruiting pedicel 2-3 mm long; fruits
subglobose, 3.8—4.2 cm long, with cinnamon-rusty scurfy hairs 0.1 mm, easily rubbed
Ht cc ek. Uovad taba Oped ae ances ved: Cet aes a. subsp. laevis

b. Peduncle of infructescence 15—25(—30) mm long; fruiting pedicel more slender, 13—
18 mm long, [bracteole scar 2-4 mm below the fruit;] fruits (slightly immature)
ellipsoid-oblong, 5—5.5 cm long, with persistent dark chocolate-coloured hairs 0.1—
WNMPMITNINTO MN tenes sens bec; teecene Bones ie eon soa ce Oana ase b. subsp. badia

a. subsp. laevis

Tree 4-20 m. Male inflorescences and flowers as the species. Female flowers: pedicel
short. Fruits solitary, subglobose, 3.6—4.2 by 3.34 cm, indumentum cinnamon or rust-
coloured, rather easily rubbed off, hairs scurfy, 0.1 mm; pericarp 7-9 mm thick; pedun-
cle of infructescence 3-10 mm long; fruiting pedicel 2-3 mm long.

Field-notes — Small tree, dbh 8-60 cm. Flowers yellow-green, fruits (slightly imma-
ture) brown.

Distribution — Malesia: E Philippines (Samar).

Habitat & Ecology — Forest of Mt Sohoton, altitude not recorded; fl. & fr. Apr.

b. subsp. badia W. J. de Wilde

Myristica laevis W.J. de Wilde subsp. badia W. J. de Wilde, Blumea 42 (1997) 172. — Type: Rojo
159, NE Mindanao.

Tree c. 5 m. Male and female flowers not seen. Fruits (slightly immature) 1-3 per
infructescence, ellipsoid-oblong, 5—5.5 by 3 cm, apex rounded with short apiculum, base
rounded, hairs persistent, dark chocolate, scurfy, 0.1—0.2 mm; pericarp hard, 10 mm
thick; peduncle of infructescence 15—25(—30) mm long; fruiting pedicel rather slender,
glabrous, 13-18 mm long, bracteole scar 2-4 mm below the fruit.

Field-notes — Low tree, dbh 10 cm, twigs and leaves rather brittle. Bark blackish;
wood soft. Infructescences among the leaves, with one or usually three fruits per pedun-
cle: fruits ovate, ferruginous to chocolate brown hairy, with mucro at apex and two shal-
low longitudinal depressions.

Distribution — Malesia: E Philippines (NE Mindanao, S Surigao), known only from
the type.

Habitat & Ecology — Hillside forest; 400 m altitude; fr. Nov.

510 Flora Malesiana, Ser. I, Vol. 14 (2000)

78. Myristica lancifolia Poir.

Myristica lancifolia Poir. in Lam., Encycl. Méth. Bot., Suppl. 4, 1 (= 12) (1816) 35 [non Myristica
lancifolia Poepp. ex Warb. (1897), nec Myristica lancifolia Merr. (1923)]; W.J. de Wilde, Blumea
35 (1990) 246; Jessup & W.J. de Wilde, Blumea 38 (1993) 39; W.J. de Wilde, Blumea 40 (1995)
297; 42 (1997) 172.

For more references and synonyms see the subspecies.

Tree 4—20(—30) m. Twigs subterete or + angular, 1-2 mm diameter, early glabrescent,
hairs 0.1(—0.3) mm; older twigs with small lenticels. Leaves chartaceous or thinly coria-
ceous, ovate-elliptic or lanceolate, (4—)7—17(—21) by (1.5—)2—6(—9) cm, base cuneate,
apex acute-acuminate, the very apex sometimes + blunt; upper surface olivaceous or
brown, lower surface pale greyish brown, glabrescent, hairs 0.1—0.2 mm or less, or per-
sistent, sparse (lens!); usually distinctly papillose; dots absent; midrib above somewhat
raised, lateral nerves 8—20 per side, at 45—80° to the midrib, rather closely set, flat and
faint above, little raised, sometimes hardly visible below, lines of interarching and vena-
tion indistinct; petioles S—15 by 1—2(—2.5) mm; leaf bud 6-12 by 1—2 mm, hairs (0.1—)
0.2(—0.3) mm. Inflorescences between the leaves and/or below, + pubescent, of the
Knema-type; in male: (sub)sessile (exceptionally peduncle up to 2 mm long), simple or
2—4-branched tubercle-like, scar-covered brachyblasts to 4(—10) mm, flowers in sub-
umbels of (2—)5—10(—15), buds usually of variable sizes; bracts minute, early caducous;
in female: brachyblasts 1(—2) mm long, few-flowered; flowers with (sparse) hairs 0.1
mm. Male flowers: pedicel slender, 1-7 mm long, bracteole 1—1.5 mm, persistent; buds
long-ellipsoid to oblong, 4-6 by 1.5—2.5 mm, somewhat narrowed towards the blunt
apex, cleft c. 1/4-1/3, lobes 0.2(—0.3) mm thick. Androecium slender, 3.5-5 mm;
androphore |.5—2 by 0.4—0.5 mm, glabrous; synandrium 1|.8—2.5(—0.3) by 0.5—0.8 mm,
thecae 10-14, (sub)contiguous, sterile apex + blunt, 0.1—0.2 mm. Female flowers: pedi-
cel 1-2 mm long; buds ovoid or conical, narrowed towards the apex, 3—4.5 by 2-3 mm,
cleft c. 1/3; ovary narrowly ovoid, 2.5—3.5 by 1.5—2 mm, hairs less than 0.1 mm, stigma
1 mm, deeply and broadly 2-lobed. Fruits 1-3 per infructescence, short- or long-ellip-
soid, 1.3—-2.6(—3) by 0.9-1.3 cm, apex rounded or subacute, usually to 1 mm beaked,
hairs scurfy, c. 0.2 mm or less (largely glabrescent in subsp. australiana); pericarp 1—2
(—3) mm thick; seeds 11-20 mm long; fruiting pedicel slender, 1-5 mm long.

Distribution — Malesia: Moluccas, W Papua Barat (Waigeo, Bird’s Head), and Papua
New Guinea (east to Madang Prov.), with three subspecies; a fourth, subsp. australiana
Jessup & W.J. de Wilde, occurs in N Australia (Jessup & De Wilde 1993: 41).

KEY TO THE SUBSPECIES

la. Leaves elliptic-oblong to lanceolate. Male buds 4—4.5 by 1.5—1.8 mm, pedicel 1—3.5
mm long. Fruits small, 1.3—1.6(—2) cm long. — Papua Barat (Obi, Waigeo, Bird’s
Head), Papua New Guinea (E to Madang Prov.) ............ a. subsp. lancifolia

b. Leaves variable but not lanceolate. Male buds somewhat larger, (4—)4.5—6 by 1.8-
2.5 mm, pedicel longer. Fruits 2.2—2.6(—3) cm long. — Moluccas and W Papua Barat
(Adtls, “sée note) ieee Ue ae ee A ae ae c. subsp. montana

c. Leaves ellipsoid-oblong, small, 4-10 cm long. Male flowers not known. Fruits ovoid-
ellipsoid, 2 by 1.5 cm. — Papua New Guinea (Southern Highlands Prov., 500—
COO iex fhe ath oR) sie ee ee b. subsp. kutubuensis

De Wilde — Myristicaceae (Myristica) 511

a. subsp. lancifolia

Myristica lancifolia Poir. in Lam., Encycl. Méth. Bot., Suppl. 4, 1 (= 12) (1816) 35; Warb., Mon.
Myrist. (1897) 519, t. 19; J. Sinclair, Gard. Bull. Sing. 23 (1968) 456, f. 79 (var. lancifolia, see
note), p.p.; W.J. de Wilde, Blumea 35 (1990) 246; Jessup & W.J. de Wilde, Blumea 38 (1993)
39. — Type: Labillardiére s.n., W Irian Jaya, Pulau Waigeo.

Myristica papuana Scheff., Ann. Jard. Bot. Buitenzorg | (1876) 47; F. Muell., Descr. Notes Papuan
Pl. 1, 5 (1877) 96. — Myristica montana Roxb. var. papuana (Scheff.) Warb., Mon. Myrist.
(1897) 514. — Type: Teijsmann 7585, Irian Jaya, Bird’s Head.

Tree 15 m. Leaves variable in shape, elliptic-oblong to lanceolate, lateral nerves 8-14
per side, comparatively rather far apart, sometimes very faint below. Male flowers: pedi-
cel 1-3.5 mm long; buds 4—4.5 by 1.5—-1.8 mm. Fruits (short-)ellipsoid, small, 1.3—
1.6(—2) by 0.9-1.1 cm, indumentum persistent; fruiting pedicel 1-3 mm. — Fig. 64h.

Field-notes — Small to rather large and straight tree, stem somewhat fluted. Bark
brown with numerous longitudinal fissures, not peeling off, 0.4 mm thick; inner bark 5
mm thick, pale ochre-red; exudate little, watery, dull brown or red; sapwood cream,
faintly reddish, gradually passing into the slightly more reddish heartwood. Flowers yel-
lowish. Fruits light brown or ochre; aril (immature?) yellow.

Distribution — Malesia: NE Moluccas (Obi), NW Papua Barat (Waigeo, Bird’s Head);
Papua New Guinea (E to Madang Prov.).

Habitat & Ecology — Hillside and riverine forest; humus soil over limestone, sandy
clay on limestone; in Waigeo I. common along many creeks; 0-650 m altitude; fl. Oct.—
Jan.; fr. May, Nov., Jan.

Note — Myristica lancifolia var. lancifolia, as accepted by Sinclair, l.c., has a wider
concept than the present subsp. /ancifolia. Most of the material cited by Sinclair for New
Guinea (including the Aru Islands), has now been transferred to other taxa.

b. subsp. kutubuensis W. J. de Wilde

Myristica lancifolia Poir. subsp. kutubuensis W. J. de Wilde, Blumea 40 (1995) 297. — Type: Jacobs
9280, Papua New Guinea.

Tree 25 m. Innovations, leaf bud, lower surface of immature leaves and flowers with
minute rust-coloured stellate hairs 0.1-0.3 mm. Leaves thinly chartaceous, elliptic-ob-
long, 4-10 by 1.53.5 cm, papillose below, lateral nerves 10-15 per side, faint, venation
almost invisible. Male flowers not seen. Female flowers: pedicel 3 mm, bracteole broadly
rounded, caducous, apical; buds ovoid, 3.5 mm long. Fruits solitary or paired, ovoid-el-
lipsoid, 2 by 1.5 cm, hairs bright rust-coloured, mealy, 0.1—0.2 mm; pericarp + woody,
2-3 mm thick: seeds 1.5 cm long; fruiting pedicel 5 by 3 mm, bracteole scar subapical.

Field-notes — Innovations olive-green. Flowers white. Fruits dull brown, rather hard.

Distribution — Malesia: Papua New Guinea (Southern Highlands Prov., near Waro
Airstrip, 20 km SSW of Kutubu, known only from the type).

Habitat & Ecology — Primary forest of limestone area; 500—600 m altitude; fl. & fr.
Oct.

Note — The status of subsp. kutubuensis is uncertain as it is as yet known only from
one female flowering and fruiting specimen. It is tentatively described as a subspecies
under M. lancifolia, as it keys out to that species. Its distinctness is possibly related to its
habitat in forest over limestone at quite a distance from the area of the type subspecies.

512 Flora Malesiana, Ser. I, Vol. 14 (2000)

c. subsp. montana (Roxb.) W.J. de Wilde

Myristica lancifolia Poir. subsp. montana (Roxb.) W.J. de Wilde, Blumea 35 (1990) 247, f. 1: 7b;
40 (1995) 298. — Myristica montana Roxb., [Hort. Beng. (1814) 105, nom. nud.] FI. Ind. 3
(1832) 846; Warb., Mon. Myrist. (1897) 512, t. 15. — Myristica lancifolia Poir. var. montana
(Roxb.) J. Sinclair, Gard. Bull. Sing. 23 (1968) 467, f. 82. — Syntypes: Wallich Cat. 6792, Smith
s.n. in Herb. Roxb., Banda I.

Myristica diversifolia Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1864) 205. — Syntypes: Teijsmann 1951
(Seram), 1964, 5057 (Ambon); de Vriese s.n. (Ambon).

Medium-sized or low tree. Leaves variable in shape, ovate-elliptic to oblong, not lan-
ceolate, lateral nerves 12-20 per side, generally rather closely parallel, faint or rather
distinct below. Male flowers: pedicel 2.5—7 mm long; buds (broader than in subsp. lanci-
folia) 4.5—6 by 1.8—2.5 mm. Fruits ellipsoid-oblong, 2.2—2.6(—3) by 1—1.3 cm, apex
narrowly rounded or subacute, indumentum persistent; fruiting pedicel 1—3(—5) mm. —
Fig. 641.

Field-notes — Bole straight, without or with only a few buttresses (0.5 by 1 m). Bark
(blackish) brown, either smooth or slightly longitudinally cracked, fissured, flaky, or
(strongly) peeling off or not; outer bark 0.5 mm thick; inner bark 4-8 mm thick, on sec-
tion outside reddish, inside (reddish) ochre or brown; exudate little, (yellow- or red-)
brown; sapwood yellowish (tinged red), the transition to brown heartwood gradual or
well marked off. Flowers cream, or (greenish) yellow, fragrant (once recorded). Fruits
(brownish) yellow; seeds dark or (blackish) brown.

Distribution — Moluccas (Halmahera, Bacan, Obi, Buru, Seram, Ambon, Banda: no
material seen, Tanimbar I.), and W Papua Barat (Adi L.).

Habitat & Ecology — Forest on clay and loam soil, with or without humus layer, over
grey schist or limestone; alluvial or porous, stony or rocky soils; 0-900 m altitude; fl. &
fr. mainly Nov.—Jan.

Note — The only specimens seen from W Papua Barat are from lowland forest, ster-
ile or with very immature flowers. Vegetatively they agree except for comparatively
indistinct papillae on the lower leaf surface (BW 9560, 11532).

79. Myristica lasiocarpa W.J. de Wilde

Myristica lasiocarpa W.J. de Wilde, Blumea 40 (1995) 298. — Type: Kairo 401, Papua New Gui-
nea.

Tree 10-12 m. Twigs 1.5—2(-3) mm diameter, late glabrescent, hairs + rough, 0.2—
0.3(—0.5) mm; older twigs + longitudinally cracking, with scattered faint lenticels. Leaves
chartaceous or coriaceous, oblong-lanceolate, 10-16 by 3—5 cm, base attenuate, apex
acute-acuminate; upper surface dark-olivaceous or brown, lower surface brown-grey
with (sub)persistent dense felty pale brown scale-like hairs 0.1—0.2 mm and scattered
emergent bright brown dendroid hairs 0.3—0.5 mm; indistinctly papillose; dots absent;
midrib (flat or) raised above, lateral nerves 13—17 per side, at 45—60(—70)° to the midrib,
(flat or) sunken above, lines of interarching and venation faint but visible in old leaves;
petiole 13-20 by 1.5—2 mm; leaf bud 15 by 2—2.5 mm, hairs 0.2-0.3 mm long. /nflores-
cences (from infructescences) between the leaves, of the Knema-type; in female: a few-
flowered, sessile, scar-covered wart-like brachyblast, 2-3 mm diameter, short-pubes-

1S)

De Wilde — Myristicaceae (Myristica) 51

cent, glabrescent. Male inflorescences, male and female flowers not seen. Fruits solitary,
ellipsoid, 4.5 by 3.5 cm, base slightly attenuate, hairs conspicuous, + thick and shaggy,
rust-coloured 2(—3) mm; pericarp 3 mm thick; seeds ellipsoid, 2.5 cm; fruiting pedicel
10(—12) by 4(—5) mm, late glabrescent, + cracking, bracteole scar 2 mm below the apex.

Field-notes — Fruits reddish brown or orange.

Distribution — Malesia: Papua New Guinea (Morobe Prov., Kuper Range area); known
only from the type.

Habitat & Ecology — Subcanopy layer of (mossy) Nothofagus-forest, with multiple-
tiered and congested canopy, understorey open, moist, dimly lit, in broad gully; 1600—
2000 m altitude; fr. Oct.—Nov.

Note — Myristica lasiocarpa is similar to M. chrysophylla mainly in its conspicu-
ously shaggy-pubescent fruits, but it differs markedly from the latter in its long fruiting
pedicel. Myristica chrysophylla is a lowland species with a wider distribution in about
the same area.

80. Myristica lepidota Blume

Myristica lepidota Blume, Rumphia | (1837) 183, t. 57; W.J. de Wilde, Blumea 35 (1990) 247. —
Type: Zippelius 159a, SW Irian Jaya.
For more references and synonyms see the subspecies.

Tree 10-35 m. Twigs 1-2 mm diameter, glabrescent, hairs 0.1 mm or less; older twigs
with numerous small pale lenticels, contrasting or not. Leaves membranous or thinly
coriaceous, elliptic-oblong, 4.5—15 by 1.5—5(—6) cm, base narrowly rounded or cuneate,
apex up to 20 mm acute-acuminate; upper surface olivaceous or blackish brown, lower
surface early glabrescent (subsp. montanoides) or with persistent indumentum (subsp.
lepidota), hairs interwoven, pale cinnamon or greyish, 0.1—0.2 mm; papillation indis-
tinct; dots absent; midrib slightly raised above, lateral nerves 6-14 per side, at 45(—60)°
to the midrib, flat or sunken above, brown and contrasting below, lines of interarching
distinct or not, venation clearly visible below; petiole (S—)10—17 by 1—1.5 mm; leaf bud
7-10 by 1 mm, hairs 0.1(—0.2) mm. Inflorescences between the leaves or below; of the
Knema-type: sessile or up to 1(—2) mm pedunculate, simple or branched scar-covered
brachyblasts to 4 mm long, 1.5 mm wide, minutely woolly brown pubescent; in male:
with clusters of 4-8 flowers, buds of strongly variable sizes; bracts minute; in female:
smaller, 2—5-flowered; flowers with hairs 0.1—0.2(—0.3) mm. Male flowers: pedicel 1—2
mm long, bracteole 0.5(—1) mm, (late) caducous; buds ellipsoid-oblong, apex and base
truncately broadly rounded, 2.5—3(—4) by 1-1.4 mm, cleft 1/3-1/4, lobes 0.2 mm thick.
Androecium slender, 2—2.6 mm long; androphore 1—1.2 by 0.4—0.5 mm, glabrous or
with hairs less than 0.1 mm in the lower half; synandrium 1.2—1.5 by 0.4—0.5 mm, the-
cae 10-12, sterile apex absent or + acute, to 0.2 mm. Female flowers: pedicel 1.5 mm;
buds ovoid, 2.5 by (1.5—)2 mm; ovary ovoid-oblong, 1.5—2 mm, hairs 0.1 mm, style and
stigma 0.3 mm. Fruits solitary or paired, subsessile, + ellipsoid-oblong, 2—3 by 1.2-1 8
cm, apex blunt or (sub)acute, or 1(—2) mm beaked, base + rounded or narrowed into the
stalk, hairs scurfy, 0.1(—0.2) mm; pericarp 2-3 mm thick; seeds ellipsoid(-oblong),
2 cm; fruiting pedicel stout, 3-4 mm long and broad.

Distribution — Malesia: E Moluccas and W & SW New Guinea; with two sub-
species.

514 Flora Malesiana, Ser. I, Vol. 14 (2000)

KEY TO THE SUBSPECIES

la. Lower leaf surface with dense persistent indumentum. Androphore minutely pubes-

centinthelower Nal faethe sc ceh werd ace a beers somal ae a. subsp. lepidota
b. Lower leaf surface early glabrescent, indumentum scarce, minute. Androphore gla-
brous or at base with few pale hairs less than 0.1 mm_....b. subsp. montanoides

a. subsp. lepidota

Myristica lepidota Blume, Rumphia | (1837) 183, t. 57; Warb., Mon. Myrist. (1897) 434, t. 19, p.p.;

Markgr., Bot. Jahrb. Syst. 67 (1935) 161, p.p.; J. Sinclair, Gard. Bull. Sing. 23 (1968) 265, f. 29;
W._J. de Wilde, Blumea 35 (1990) 248, f. 1: 8a. — [Myristica microcarpa Zipp., nom. nud. in
sched. ]

Leaves below rather woolly-pubescent, hairs pale cinnamon or grey-brown, inter-
woven, 0.1—0.2 mm. Male flowers: buds 2—2.5 mm long; thecae 10, lower half of andro-
phore with minute grey-white hairs 0.1 mm or less. — Fig. 64j.

Field-notes — Tall tree, to 35 m, dbh c. 50 cm. Flowers yellow. Fruits yellow.

Distribution — Malesia: Moluccas (Aru Islands), SW Papua Barat (also Bird’s Head).

Habitat & Ecology — Lowland rain forest; flat country; sandy and clayey alluvial
soils; 0-150 m altitude; fl. May—June, Dec.; fr. May—June.

Note — Distinguishable by slender twigs and small leaves with short but dense per-
sistent indumentum on lower surface. Both male and female flowers are small, 2—2.5
mm long, with short pedicels. The fruits are subsessile, ellipsoid-oblong, pear-shaped,
or obovoid, with comparatively short thick stalks.

b. subsp. montanoides (Warb.) W. J. de Wilde

Myristica lepidota Blume subsp. montanoides (Watb.) W.J. de Wilde, Blumea 35 (1990) 248, f. 1:
8b: 40 (1995) 299; 42 (1997) 173. — Myristica montanoides Warb., Mon. Myrist. (1897) 514.
— Myristica globosa auct. non Warb.: J. Sinclair, Gard. Bull. Sing. 23 (1968) 378, p.p., as for
the synonym M. montanoides only. — Type: Beccari Fl 7756, NE Moluccas.

Leaves below early glabrescent, hairs dispersed, pale, less than 0.1 mm. Male flow-
ers: buds 3(—4) mm long; thecae 10 or 12, androphore somewhat grooved corresponding
to the anthers, glabrous or nearly so, with at base a few pale hairs less than 0.1 mm. —
Fig. 64k.

Field-notes — Buttresses few, to 50 cm high and out. Bark of trunk fissured, not
peeling off, or scaly; outer bark 0.3 mm thick, dark grey to light brown; inner bark 4 mm
thick, pinkish; sap reddish; slash wood pink or pale yellowish tinged reddish, gradually
passing into slightly darker heartwood, or heartwood absent. Flowers yellowish. Fruits
orange, light brown, or pale brown, with abundant ochre or clear exudate.

Distribution — Malesia: NE Moluccas (Ternate, Obi, Bacan), Papua Barat (Bird’s
Head).

Habitat & Ecology — Primary and disturbed forest, hill ridge forest; loamy soil with
stones; 50—600 m altitude; fl. Feb.; fr. Sept.—Nov.

Note — Resembles M. tristis from the same area, which differs in its larger and dif-
ferently shaped male flowers and larger, almost globose fruits with thick pericarp.

De Wilde — Myristicaceae (Myristica) SIS)

81. Myristica leptophylla W.J. de Wilde

Myristica leptophylla W.J. de Wilde, Blumea 40 (1995) 299. — Type: LAE 67023 (Vinas), Papua
New Guinea.

Tree 10-12 m. Twigs (1.5—)2.5 mm diameter, early glabrescent, hairs less than 0.1
mm; older twigs with few inconspicuous lenticels. Leaves membranous or (thinly) char-
taceous, oblong-lanceolate, 14~20 by 3.5—5 cm, base short-attenuate or narrowly rounded,
apex acute-acuminate; upper surface olivaceous-brown, lower surface grey-brown, early
glabrescent, hairs less than 0.1 mm; not papillose; dots present (lens!); midrib raised
above, lateral nerves 20(—25) per side, at 60—80° to the midrib, thin and sunken above,
brown and contrasting in colour below, lines of interarching and venation inconspicu-
ous; petiole 10-20 by 2—2.5 mm; leaf bud 10-12 by 2 mm, hairs less than 0.1 mm.
Inflorescences (from incomplete female and fruiting specimens): of the Knema-type: a
sessile or 2(—3) mm pedunculate, scar-covered brachyblast 2 mm long, with 2 or 3 flow-
ers (according to their scars), minutely pubescent; bracts not seen. Male inflorescences,
male and female flowers not seen. Infructescence below the leaves, sessile or with pe-
duncle to 3 mm long. Fruits (somewhat immature) solitary, broadly fusiform, 4.5 by 2.5
cm, apex acute with 3 mm long beak formed by style-remnant, base + narrowed into a
3 mm long pseudostalk, hairs dense, rust-coloured, scurfy, 0.1 mm; pericarp 3-4 mm
thick; seeds not seen; fruiting pedicel 10-12 by 4(—5) mm, bright brown, finely fissured
and with small contrasting lenticels, bracteole scar + median.

Field-notes — Tree 10-12 m, bole 8 m, dbh 10-20 cm. Bark finely fissured, outer
bark grey or brown, inner bark orange-straw, with red sap; wood straw or whitish brown,
turning orange on exposure, or with red sap. Leaves glossy dark green above, light green
below. Fruit with orange-brown scurf; aril of immature seeds with watery liquid.

Distribution — Malesia: Papua New Guinea (W Sepik Prov., Telefomin subprov.,
near Busilmin); known only from the type area.

Habitat & Ecology — Secondary regrowth of submontane forest; 1500 m altitude;
fr. Mar.

Note — This species keys out beside M. pachyphylla to which it is similar and closely
related. The two specimens now known of M. leptophylla differ in larger and thinner
leaf blades, and in slightly larger fruits with stouter fruiting pedicel, sessile or on a short
peduncle. Myristica tamrauensis, from Bird’s Head, is similar too.

82. Myristica longepetiolata W.J. de Wilde

Myristica longepetiolata W.J. de Wilde, Blumea 42 (1997) 173. — Type: PNH 21548 (Sulit), Phil-
ippines, Biliran Is., northern slope of Mt Suiro.

Tree 17 m. Twigs 2-4 mm diameter, faintly angular, early glabrescent, hairs scatter-
ed, greyish, less than 0.1 mm; older twigs with scattered conspicuous lenticels. Leaves
(thinly) chartaceous, (ellipsoid-)oblong, sometimes narrowed in the upper half, 10-15
by 2.5—6 cm, base cuneate, apex blunt to acute-acuminate; upper surface glossy, olivaceous
or bright brown, lower surface grey-brown, glabrous; not or indistinctly papillose; dots
absent; midrib flat or slightly raised above, lateral nerves (10—)12—15 per side, at 45
(—60)° to the midrib, flat and faint above, lines of interarching indistinct, venation coarsely
reticulate, not very distinct; petiole comparatively long, bright reddish or yellowish brown

516 Flora Malesiana, Ser. I, Vol. 14 (2000)

(the same colour as the twigs), 20-35 by 1.5—2 mm; leaf bud 10 by 1.5—2 mn, hairs in-
conspicuous, appressed, greyish or brown, 0.1 mm or less. Inflorescences (known only
from infructescences) subsessile, peduncle 1-2 mm long, one-fruited, with a few scars
of abortive flowers. Male and female flowers not seen. Fruits subglobose or broadly
(ovoid-)ellipsoid, 5—6 by 4—4.5 cm, hairs dense, dark rusty, 0.1(—0.2) mm; pericarp 10
mm thick; seeds ellipsoid, 3.5—4 cm; fruiting pedicel short (see note).

Field-notes — Tree, dbh 50 cm. Fruits large, brown.

Distribution — Malesia: Philippines (Luzon: Sorsogon and Camarines Prov.; Biliran).

Habitat & Ecology — Sloping forests; c. 800 m altitude; fr. Apr—May, July—Aug.,
Dee:

Note — The subsessile infructescences with rather stout, short, 1-2 mm long pedun-
cle suggest that male inflorescences may be (short) paniculate, with a more or less dis-
tinct peduncle, pointing to a relationship with M. agusanensis. The latter species differs
in the minute indumentum on its lower leaf surface (+ glabrescent), and in its smaller
fruits.

83. Myristica longipes Warb.

Myristica longipes Warb., Mon. Myrist. (1897) 535; Markgr., Bot. Jahrb. Syst. 67 (1935) 163 (p.p.,
for the type only); J. Sinclair, Gard. Bull. Sing. 23 (1968) 343, f. 52C, D, F (p. p.. for part of the
specimens from Central Proyv., PNG, only); Foreman, Handb. Fl. Papua New Guinea | (1978)
205, p.p.; W.J. de Wilde, Blumea 40 (1995) 300. — Type: MacGregor s.n. (B, lost), Papua New
Guinea, Central Prov., Mt Yule. Neotype: Forbes 647 (L, iso K), Papua New Guinea.

Tree 7-20 m. Twigs 1.5—2.5 mm diameter, somewhat angular, glabrescent, hairs 0.1—
0.2 mm; older twigs with inconspicuous scattered lenticels. Leaves + membranous, el-
liptic-oblong, 9-17 by 4—6.5 cm, base narrowly rounded or cuneate, apex acute(-acum1-
nate), upper surface dark brown, lower surface silvery-grey, early or late glabrescent,
hairs sparse, scale-like, 0.1 mm or less; indistinctly papillose; dots absent; midrib slightly
raised above, below chestnut-coloured, contrasting in colour, lateral nerves 10-15 per
side, at 40—50° to the midrib, flat or sunken, indistinct above and below, lines of
interarching and venation faint; petiole 10-20 by 2 mm, leaf bud 8-15 by 2 mm, hairs
0.1-0.2 mm. Inflorescences pedunculate, between the leaves or just below, with two
short branches. and one central flower (or this abortive), flowers in small subumbels,
hairs sparse, shaggy, 0.1 mm; bracts minute, pubescent, caducous; in male: peduncle
5—15 mm, lateral branches 15 mm with up to 2 mm long sterile basal part proceeding in
a scar-covered slender brachyblast; flowers 2 or 3, buds of various sizes; female inflo-
rescences: similar, peduncle 5—10 mm, few-flowered. Male flowers: pedicel 5—6 by 0.6—
0.7 mm, bracteole caducous, minute; buds subcoriaceous, oblong-lanceolate, 10 by 3
mm, apex acute, base subacute, cleft c. 1/10, lobes 1 mm long, 0.7 mm thick. Androecium
slender, 9-10 mm; androphore cylindrical, 4 by 0.7 mm, hairs sparse, less than 0.1 mm;
synandrium 4 by 0.7—0.8 mm, thecae c. 14, sterile apex slenderly acute, 1.5 mm, gla-
brous. Female flowers: pedicel 14-15 mm, bracteole caducous, 4 mm below the apex;
buds ovoid, apex acute, 5.5 by 3 mm, cleft c. 1/3; ovary 3.5 mm, minutely pubescent.
Infructescences: peduncle + slender, 10-15 mm long. Fruits single, 4.5—6 by 2-2.5 cm,
fusiform, apex acute, beaked by style-remnant, base narrowed into a 5—7 mm long pseudo-
stalk, hairs dense, mealy, dark-brown, 0.1 mm; pericarp + woody, (3—)4-5 mm thick, the

De Wilde — Myristicaceae (Myristica) Sly

suture distinctly sunken at one side; seeds (somewhat immature) elliptic-oblong, 3 cm;
fruiting pedicel 15-17 by 2-3 mm, glabrescent, bracteole scar 5 mm below the apex. —
Fig. 63g.

Field-notes — Bark smooth, mottled grey outside, inside straw turning brown on ex-
posure. Leaves dark green glossy above, pale dull grey-green or glaucous below; dry
leaves extremely brittle. Fruits rufescent orange, brown, or brownish ochre; seeds black.

Distribution — Malesia: Papua New Guinea (Central Prov., Sogeri & Kotaki region);
possibly C Papua Barat (but see note 2); a locally endemic species.

Habitat & Ecology — Mixed hill rain forest, foothill forest; (300—)500—750 m alti-
tude (but see note 2); fl. June; fr. June, Sept.

Notes — 1. Myristica longipes was described after a single fruiting collection, later
on lost in Berlin. When revising the Papuasian Myristicas Markgraf (1936) added a few
collections, but Ledermann 9728, on which he described the male flowers, was (is) con-
sidered by Sinclair and me to belong to a different species. Myristica longipes, more-
over, contains in Sinclair’s (1968) and Foreman’s (1978) treatments many specimens
which at present are transferred to various other species. The present species is now
restricted to five collections and Forbes 647 (with almost mature fruit) is herewith des-
ignated as neotype.

2. Brass & Versteegh 12574 (Central Papua Barat, 1300 m altitude), with dark orange
fruits, keys out to M. longipes. It is possible that it is a species of its own. The specimen
differs from M. longipes (Papua New Guinea) in shorter and less slender fruiting pedi-
cel, 10 by 4—5 mm, with the bracteole scar 4 mm below the apex; the fruit is compara-
tively slightly broader, 4—4.5 by 2 cm, with short and broad pseudostalk, but is some-
what immature. The collecting locality is far apart from that of M. longipes, and at a
higher altitude. More (preferably male flowering) material from Papua Barat should
elucidate its status. The specimen is also close to M. laevifolia and M. vinkii, both (gen-
erally) with sessile inflorescences.

84. Myristica lowiana King

Myristica lowiana King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 293, p.p., pl. 120 2-44 S-7=
Myristica crassa); Warb., Mon. Myrist. (1897) 496, p.p.; J. Sinclair, Gard. Bull. Sing. 16 (1958)
345, f. 23, pl. IV B; 23 (1968) 164; W.J. de Wilde, Blumea 42 (1997) 174; Tree FI. Sabah &
Sarawak 3 (2000) 465. — Lectotype (Sinclair 1958): Scortechini 185], Peninsular Malaysia.

Myristica hackenbergii Diels, Bot. Jahrb. Syst. 60 (1926) 308. — Syntypes: Hackenberg 86 (male),
86a (female) (B, both lost), S Kalimantan.

Tree 7-25 m. Twigs 3-6 mm diameter, late glabrescent, hairs rough-woolly, rufous,
1-1.5 mm; older twigs blackish, cracking and flaking; lenticels indistinct or absent. Leaves
coriaceous, oblong(-lanceolate), 14-35 by (3.5-)9-11.5 cm, base rounded or short-
cuneate, apex acute-acuminate; upper surface glossy, olivaceous or brown, lower sur-
face grey-brown, early glabrescent, hairs mealy, rust-coloured, 0.1—0.2 mm; distinctly
papillose; dots absent; midrib raised above, glabrescent, lateral nerves 15-22 per side, at
(45—)60-70° to the midrib, usually (not always, see note 1) sunken above, lines of inter-
arching indistinct, venation sunken above, rendering the blade somewhat bullate; peti-
ole 25-50 by 2.5-5 mm, late glabrescent; leaf bud 10-15 by 4-5 mn, hairs 1—-1.5 mm.
Inflorescences between the lower leaves or below, pedunculate, branched, with rough-

518 Flora Malesiana, Ser. I, Vol. 14 (2000)

woolly hairs 1-1.5 mm; bracts 3-5 mm, late caducous; in male: 2.5-5.5 by 2-3 cm,
peduncle (10—)15—30 mm, basal branches 5-10 mm long, central axis 5-25 mm, each
branch with 1 or 2 (or 3) (sub)sessile brachyblasts, each with a subumbel of 5—10 flow-
ers, buds of various sizes; in female: 1—1.5 cm long, hardly branched, rather thick, 3-5-
flowered: flowers with hairs 0.5-1 mm long. Male flowers: pedicel 2—3(—5) by 1 mm,
bracteole 3 mm diameter (apex sometimes subtruncate), late caducous; buds ellipsoid
(-oblong), 4—4.5 by 3 mm, apex narrowly rounded, cleft 1/4(—1/3), lobes 0).3(—0.4) mm
thick. Androecium (3—)3.5—4 mm long; androphore 1.5—2 by 0.8—1 mm, with dense
hairs 0.5 mm in the lower 4/5; synandrium oblong, 2—2.2 by 0.8 mm, thecae 10-14,
sometimes at apex loosely coalescent and proceeding into a (0.1—)0.2—0.3 mm long,
lobed, sterile apex. Female flowers: pedicel short; buds ovoid, 5 by 4 mm, cleft c. 1/4;
ovary rusty tomentose. Infructescence 1-1.5 cm. Fruits generally solitary, ellipsoid,
(5-)6-7 by 3.5—4 cm, hairs velvety, 1-2.5 mm; pericarp (5S—)10(-15) mm thick; seeds
ellipsoid, 4-5 cm; fruiting pedicel rather thick, 2-5 mm long.

Field-notes — Usually with stilt-roots or with (flying) buttresses (to 4 ft tall); also in
drier situations. Bark chocolate or blackish, hard, brittle, longitudinally fissured (or
ridged), or scaly, 6 mm deep; outer bark blackish; inner bark slightly laminated, (pink
or) brown; exudate watery, red; cambium white; sapwood white turning brownish. Flowers
fragrant, cream, rusty tomentose on outside; once (Lorzing 14260) pink, fragrant like
Mimusops elengi; perianth teeth at anthesis slightly reflexed. Fruits with brown furry
indumentum; seeds glossy dark brown.

Distribution — Malesia: Sumatra (incl. Bangka), Peninsular Malaysia, Singapore,
Borneo.

Habitat & Ecology — Fresh water peat swamp forest and kerangas; also riverine
forest, Dryobalanops rappa-forest, ridge forest (on yellow soil) and undulating hills (on
yellow sandy soils); 0—200(—800) m altitude; fl. & fr. throughout the year.

Notes — |. Usually the nerves are clearly sunken above, but in some specimens from
the East Coast of Sumatra, Bangka, and many from Borneo, (in older leaves) they may
be raised above.

2. Myristica lowiana is related to M. maingayi which differs in less stout twigs, less
coriaceous leaves, slightly larger flowers with less thickly pubescent androphore, and
less thickly tomentose fruits; apparently M. lowiana is largely confined to forest on poorer
soils, like swamp forest, kerangas, and ridge forest.

3. Variation mainly concerns the thickness of the indumentum of the fruits, especially
in Borneo, where it may vary between | and 3 mm; in S 16509 the hairs are dense and
shaggy, 2-3 mm long. In Sumatran and Malayan specimens the hairs are shorter, 1-2
mm. FRI 7734 (Cockburn) from Peninsular Malaysia (G. Panti) deviates from other
Malayan material by its stout habit, rather small fruits (6 cm) with very thick fur-like
indumentum, hairs 3-4 mm long; possibly this specimen represents a separate variety.

85. Myristica maingayi Hook. f.

Myristica maingayi Hook. f., Fl. Brit. India 5 (1886) 104; King, Ann. Roy. Bot. Gard. Calc. 3 (1891)
294, pl. 114 (excl. Curtis 2455); Warb., Mon. Myrist. (1897) 398; J. Sinclair, Gard. Bull. Sing.
16 (1958) 348, f. 24, plate V B; 23 (1968) 166; W. J. de Wilde, Blumea 42 (1997) 175. — Type:
Maingay 1289, Peninsular Malaysia.

De Wilde — Myristicaceae (Myristica) og

Tree 10—40 m. Twigs (2—)2.5—4 mm diameter, (late) glabrescent, hairs rusty, woolly,
0.2—0.5 mm; older twigs blackish, longitudinally and transversely cracking, or coarsely
flaking; lenticels inconspicuous. Leaves chartaceous or (thinly) coriaceous, elliptic-ob-
long or oblong-lanceolate (rarely linear-oblong), 12-28 by 48.5 cm, base rounded or
short-attenuate, apex acute-acuminate; upper surface olivaceous-brown, glossy, lower
surface grey-brown, early glabrescent, hairs 0.1—0.3 mm, often with hair scars; distinctly
papillose; dots absent; midrib + raised above; lateral nerves 13-20 per side, at 45—80° to
the midrib, sunken, lines of interarching indistinct, venation sometimes visible; petiole
15-30 by 1.5—3 mm, glabrescent; leaf bud 8-15 by 2.5—4 mm, hairs shaggy or appressed
0.5—1(-1.5) mm. Inflorescences between the leaves or below, pedunculate, with hairs
1(-1.5) mm; in male: 1.5—2 by 1.5—2 cm, peduncle 5—12 mm long, branches to 4 mm,
central axis absent or to 6 mm, simple (or bifid), apical subumbel(s) of 5—15(—20) flow-
ers, buds of various sizes; bracts 4 mm, caducous; female inflorescences like the males,
but shorter and stouter, 1—4-flowered; flowers with shaggy hairs 0.5—1 mm. Male flow-
ers: pedicel 5—6(-9) mm, bracteole 4-5 mm, rounded, or truncate or faintly angular
(3-topped), subpersistent; buds ovoid-ellipsoid, 6-7 by 4.5—5 mm, cleft 1/5—1/4, lobes
0.3-0.4 mm thick. Androecium 5—5.5 mm; androphore 1.2-2 by 1—-1.2 mm, hairs 0.5
mm in the lower 3/4; synandrium 2.8-3.5 by 1 mm, thecae 12-16, sterile apex + acute,
0.5(-1) mm. Female flowers: pedicel 3 mm, bracteole 5 mm, caducous; buds ovoid
(-ellipsoid), 6 by 4.5 mm, cleft c. 1/4; ovary 3.5(—4) by 3 mm, hairs 0.5 mm, stigma
0.5(-1) mm long. Fruits | or 2 per infructescence, broadly ovoid-ellipsoid, 7-9 by 5-6
cm (but see note 2), glabrescent, hairs rust-coloured, mealy, 0.5 mm; pericarp 15-20
mm thick; seeds ellipsoid, 4—5 cm; fruiting pedicel short, stout, 4-11 by 5-7 mm.

Field-notes — Bark much fissured longitudinally or with adherent scales, greyish or
blackish, brittle; inner bark reddish; sap pink, watery, copious; wood white or reddish.
Flowers rusty-tomentose, perianth inside (female) creamy-yellow. Fruits 10.5 by 6—6.5
cm, pale yellow with some reddish brown scurf, soon becoming entirely glabrous; aril
scarlet; seeds 6 cm long, blackish brown, glossy, smooth.

Distribution — Thailand; Malesia: Peninsular Malaysia, Singapore; possibly Sumatra
(see note 2).

Habitat & Ecology — Hillside forest (near river), ridge forest; sandstone ridge; alti-
tude 0-700 m; fl. & fr. throughout the year.

Notes — 1. Related to M. gigantea and M. lowiana (series Maingayae of Sinclair,
l.c.); for differences see the keys.

2. This species is variable especially in the size of the leaves. FRI 17603 (Chan) and
KEP 77818 (Jabil), both from Peninsular Malaysia, differ from the rest of the material
of M. maingayi in stouter twigs with coarsely striate older bark, neither cracking nor
flaking, in slightly broader leaves, and mainly in the smaller fruits, 4.5—5 by 3.5 cm,
with subpersistent mealy light brown indumentum. Possibly these two specimens repre-
sent a separate taxon, but more fruiting and flowering material is needed to make a
decision. Krukoff 4384 (Sumatra, East Coast) differs in its smaller fruits, 6.5 by 4 cm,
with a spongy-woody pericarp, so that it did not dry coarsely wrinkled as in the Malaysian
specimens. It is the only collection of M. maingayi known from Sumatra. This specimen
was also discussed separately by Sinclair (1968: 166).

520 Flora Malesiana, Ser. I, Vol. 14 (2000)

86. Myristica malaccensis Hook. f.

Myristica malaccensis Hook. f., Fl. Brit. India 5 ( 1886) 104: W.J. de Wilde, Blumea 42 (1997) 177;
Tree Fl. Sabah & Sarawak 3 (2000) 466. — Type: Maingay 1305, Peninsular Malaysia.
For more references see under the subspecies.

Tree 7-35 m. Twigs 2-4 mm diameter, + angular, early glabrescent, hairs less than 0.1
mm; older twigs often blackish, longitudinally cracking and moderately flaking; lenticels
absent. Leaves thinly chartaceous, (elliptic-)oblong, sometimes contracted below the
middle, 10-30 by 4-10 cm, base rounded or cuneate, apex acute-acuminate; upper sur-
face olivaceous-brown, lower surface greyish brown, early glabrescent; papillation ei-
ther not obvious or distinct (subsp. papillosa); dots absent; midrib raised above, lateral
nerves 14—20 per side, at 60—80° to the midrib, flat or sunken above, contrasting below,
lines of interarching and venation distinct; petiole 12-24 by 2-3 mm, leaf bud 7-12 by
1.5—3 mm, hairs 0.1 mm or less. Inflorescences between the leaves or below, early gla-
brescent, hairs less than 0.1 mm; in male: narrowly paniculate, many-flowered, 7-10 by
3-4 cm, peduncle 20-30 mm, basal branches 10-20 mm, + branched, central axis with
up to 13 side branches 10 mm long or less, subumbels of 5—15 flowers, buds of various
sizes: bracts 2-3 mm, caducous; female inflorescences smaller than male, 1.5—2.5 cm
long; flowers early glabrescent, hairs 0.1 mm or less. Male flowers: pedicel 3-5 mm,
bracteole 1.5 mm, subcircular, glabrescent, margin ciliate, caducous or persistent; buds
(subglobose-)ovoid, 3—3.5 by 2—3(—3.5) mm, cleft 1/4—1/3, lobes 0.3 mm thick.
Androecium 1.5(—2) by 0.8(-1) mm, consisting almost entirely of the synandrium;
androphore 0.2—0.5 mm long, glabrous; thecae 12—16, anthers 1.5 mm long, with free
apices 0.1—0.2 mm, sterile apex absent or apex of synandrium shallowly hollowed. Fe-
male flowers: pedicel 2 mm long, bracteole (1—)1.5 mm, margin ciliate, persistent; buds
broadly ellipsoid, 2-2.5 by 1.5 mm, cleft 1/4—1/3; ovary conical-ovoid, 1.3 by 1 mm,
almost glabrous. Infructescence 1-2 cm long, with few flower scars, 1- or 2-fruited.
Fruits ellipsoid-oblong, 4-6 by 2.5—4 cm, glabrous, but with minute indumentum in
grooves near the fruiting pedicel; pericarp S—10 mm thick; seeds 3.5—4.5 cm; fruiting
pedicel 5—8 mm long.

Field-notes — Tree with buttresses 1-3 m high. Bark smooth, greenish grey to dark
brown, with narrow furrows, or with narrow square sections, fissures and ridges with
small flat adherent scales; outer bark thin, soft; inner bark pale yellow, soft; exudate
clear, watery, faintly red, slow to appear; sapwood white or brown. Flowers yellowish.
Fruits ellipsoid or oblong and slightly tapered, large, pale green, yellow-cream, or pink.

Distribution — Malesia: Sumatra, Peninsular Malaysia, Borneo.

Habitat & Ecology — Primary forest on hillsides; also disturbed forest; stony soil,
undulating land near stream with rich clay soil; 0-700 m altitude; fl. June; fr. Nov.— Mar.

Note — After flowering, the female perianth breaks off, leaving a singular, persistent,
small disc-like collar around the base of the young fruit.

KEY TO THE SUBSPECIES

la. Lower leaf surface distinctly and regularly fine-papillose (lens!). — Sarawak .....
1 LA ee Nis na eee cd arene: CEE MU NE te no ata. Ones) NE b. subsp. papillosa

b. Lower leaf surface not, or very irregularly and faintly papillose. — Sumatra, Penin-
sulan Malaysia, OMEO Gas. ection ee a. subsp. malaccensis

De Wilde — Myristicaceae (Myristica) 37

a. subsp. malaccensis

Myristica malaccensis Hook. f., Fl. Brit. India 5 (1886) 104; King, Ann. Roy. Bot. Gard. Calc. 3
(1891) 287, pl. 107 bis; Warb., Mon. Myrist. (1897) 411; J. Sinclair, Gard. Bull. Sing. 16 (1958)
356, f. 27; 23 (1968) 184, p.p., excl. most of Borneo [non Myristica malaccensis Gand. (1919)
= Ardisia teijsmanniana Scheff.]; W. J. de Wilde, Blumea 42 (1997) 177; Tree Fl. Sabah & Sarawak
3 (2000) 467.

Myristica pandurifolia H. J.P. Winkler, Bot. Jahrb. Syst. 49 (1913) 367; Merr., Enum. Born. (1921)
269. — Type: Winkler 2405, Kalimantan.

Leaf blades large or small; venation faint or distinct; lower surface not or very irregu-
larly and faintly papillose.
Distribution — Malesia: Sumatra, Peninsular Malaysia, Borneo

b. subsp. papillosa W. J. de Wilde

Myristica malaccensis Hook.f. subsp. papillosa W.J. de Wilde, Blumea 42 (1997) 7. ire.
Sabah & Sarawak 3 (2000) 467. — Type: S 36528 (Ilias Paie), Sarawak.

Leaf blades comparatively large, 15-28 by 7-10 cm, veins strongly reticulate; lower
surface distinctly and regularly fine-papillose (lens!).

Distribution — Malesia: Borneo (Sarawak).

Note — The presence of papillae on the lower leaf surface is an important taxonomic
character in Myristica. Specimens of subsp. papillosa are generally stouter than those of
subsp. malaccensis. Possibly the male flowers are identical in both subspecies. To judge
the definite status of subspecies papillosa more material (with fruits), is needed.

87. Myristica markgraviana A.C. Sm.

Myristica markgraviana A.C.Sm., J. Arnold Arbor. 22 (1941) 66; J. Sinclair, Gard. Bull. Sing. 23
(1968) 221, f. 18; W.J. de Wilde, Blumea 40 (1995) 300, f. 4. — Type: Clemens 1142, Papua
New Guinea.

Myristica philippensis auct. non Lam.: Markgr., Bot. Jahrb. Syst. 67 (1935) 158 (based on Schlech-
ter 16789).

Tree 5-30 m. Twigs 1.5—3(-5) mm diameter, faintly triangular, + late glabrescent,
hairs rusty to blackish brown, mealy, 0.5—1 mm; older twigs sometimes flaking; lenticels
absent. Leaves membranous or chartaceous, (elliptic-)oblong, 8-23 by 3-10 cm, base
attenuate or (broadly) rounded, apex acute-acuminate (or + blunt); upper surface green-
ish or (dark) brown, lower surface grey, late glabrescent or hairs (sub)persistent, dense,
interwoven, brown or grey, stellate, 0.1—0.2 mm, mixed with scattered brown dendroid
hairs 0.5(—0.7) mm rendering the surface with a powdery appearance; not papillose;
dots absent; midrib flat, glabrescent above, lateral nerves (6—)10—15 per side, at c. 60° to
the midrib, sunken below, lines of interarching + distinct, venation faint or invisible;
petiole 10-22 by 2-3 mm; leaf bud 6-16 by 2-5 mm, hairs shaggy, 0.5—1 mm. Jnflores-
cences with rust-coloured hairs 1 mm, pedunculate, in male 1—7, each axillary to caducous
cataphylls along short-shoots 1-5 cm long, in female fewer; short-shoots to 1 cm long
and ending in a reduced leaf bud, axillary to foliage leaves; bracts 3-4 mm, caducous;
male: peduncle 8—20 mm, with (sub)opposite wart-like lateral branches, each with a
fascicle of 1-3 flowers, and with or without an apical central flower, buds + equal in

522 Flora Malesiana, Ser. I, Vol. 14 (2000)

— — —s

Th Met yh oe BASRA
Mere ee eee lived b

MAMA Stee cae

bray atdaebere Tyeibed
les Mtg oe, t -<

previa iiuara Cantey Me ee nketzea

X SCANS
Saas

eis
es

BK

Ags.

De Wilde — Myristicaceae (Myristica) 23,

size; in female: smaller, peduncle 5—10 mm, apically 1—3-flowered; flowers easily dis-
articulating, with shaggy hairs 0.5—-1.5 mm. Male flowers: pedicel 7-12 mm long, brac-
teole subcircular to broadly ovate, 3—4 mm; buds ellipsoid or obovoid, inside towards
base verrucose, 7-9 by 5-8 mm, not collapsing on drying, cleft 1/5—1/3, lobes 0.5 mm
thick. Androecium (3—)4—5 mm; synandrium 3—4.5 by 1-1.5(—2) mm, at apex shallowly
hollowed to 0.5 mm, thecae (14—)20-26, at apex incurved, at base contracted into slen-
der androphore, 0.5—1 mm, glabrous, but usually with frill of dense hairs 0.5—1 mm at
the bottom of the perianth. Female flowers: pedicel thickened towards apex, 6-10 mm
long, bracteole caducous, 1-4 mm below apex; buds ovoid, 7 by 7 mm; ovary ovoid,
4 mm long, with lanose hairs 2(-3) mm; style including stigmas 1.5—2 mm, lobes shallowly
lobulate. Fruits 1(—3) per (partial) infructescence, subglobose or ellipsoid, apex usually
+ hooked, 3-5.5 by 2.5—3 cm, hairs dense, shaggy, (red-)brown, (0.5—)1—1.5(—3) mm;
pericarp 4—9 mm thick; seeds ellipsoid, 2 cm; fruiting pedicel 6—12 mm long. — Fig. 79.
Field-notes — Medium-sized understorey tree; bole straight, when old fluted to 2 m
or with small buttresses up to 1 by 0.3 m; branches nearly horizontal. Bark 6 mm thick,
dark grey brown, smooth or usually longitudinally fissured or with longitudinal cracks,
when old flaking in nearly rectangular strips; inner bark concentrically layered, red-
brown or brown; exudate watery or thickish, pink to red (also in the wood); blaze
(sapwood) white or straw, cream, or pinkish; heartwood pink. Twig apices, young leaves
and flower buds golden-brown hairy. Leaves below silvery or ashy, with brown scat-
tered hairs emerging from paler dense layer of scale-like hairs. Immature aril white.
Distribution — Malesia: Papua New Guinea (Eastern, Milne Bay, Morobe, Madang,
Northern Provinces).
Habitat & Ecology — Lowland and submontane rain forest, (oak) forest on ridges;
clay soil (once recorded); to c. 950 m altitude; fl. mainly June—Dec.; fr. Jan., Mar.—June.
Note — The inner surface of the tubular part of the male perianth is conspicuously
verrucose, i. e., with (densely set) irregular wart-like appendages, irregularly scattered
or sometimes arranged in + horizontal structures, the warts 0.2—0.4 mm high, possibly
purplish or reddish when fresh; inner surface of the perianth lobes completely glabrous.

88. Myristica maxima Warb.

Myristica maxima Warb., Mon. Myrist. (1897) 385; J. Sinclair, Gard. Bull. Sing. 16 (1958) 339,
f. 20, 21, pl. I-IV A; 23 (1968) 131; W.J. de Wilde, Blumea 42 (1997) 178; Tree Fl. Sabah &
Sarawak 3 (2000) 467. — Lectotype: King’s coll. 5513, Peninsular Malaysia.

Myristica bracteata auct. non A. DC.: King, Ann. Roy. Bot. Gard. Calc. 3 (1891) 286, pl. 107 (excl.
pl. 106 = Myristica philippensis Lam.).

Fig. 79. Myristica markgraviana A.C. Sm. a. Apical part of leafy twig; b. detail of indumentum of
lower leaf surface; c. short and long dendroid hair of indumentum of lower leaf surface; d. twig with
male inflorescences arranged along axillary short-shoots each ending in a vegetative bud; e. male
bud, opened, showing androecium; note finely warty inner surface of perianth; f. androecium in
section, schematic; note shallowly impressed apex; g. part of twig with lateral short-shoot with
one female inflorescence; h. female bud, opened, showing pistil; note bracteole scar, perianth minute-
ly warty-bullate inside; i. older twig with one-fruited infructescence at the end of lateral short-shoot
[a—c: NGF 46566; d—f: NGF 14828; g, h: Saunders 426; 1: NGF 45898]. — Scale bar for a, d, g, 1
= 2 cm: for b = 0.83 mm; for c = 0.2 mm; for e, h = 3.3 mm; for f = 1.65 mm.

524 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 15-35 m. Twigs (4—)5—8 mm diameter, early glabrescent, indumentum minute;
older twigs becoming rough and scaly, not longitudinally cracking; lenticels often present
but indistinct. Leaves membranous, chartaceous or (thinly) coriaceous, elliptic or ob-
long-lanceolate, 16—40 by (6-)10—20 cm, base rounded or shallowly cordate, sometimes
attenuate, apex acute-acuminate; upper surface (blackish) brown, sometimes bullate,
lower surface (brownish) grey, seemingly glabrous, but actually minutely pubescent, or
glabrescent, hairs interwoven or sparse, grey-brown, scale-like, 0.1(—0.2) mm or less;
usually papillose; dots absent; midrib flat or raised above, lateral nerves 23-33 per side,
at 60—70° to the midrib, flat or impressed above, lines of interarching and venation dis-
tinct (or indistinct) on both surfaces; petiole 25-35 by 2.5—4 mm; leaf bud 15-20 by
(3-)4-5 mm, hairs 0.1 mm or less. Inflorescences usually between the leaves, subglabres-
cent, indumentum minute, grey-villous, essentially paniculate, stout, the peduncle dis-
tinctly flattened, sometimes lenticellate; in male: 6—12(—18) by 3-8 cm, peduncle 20—45
by 4-6 mm; lower branches up to 3 cm, central axis 4-8 cm long, with up to 6 side
branches, subumbels shortly stalked or sessile, each with S—10 flowers, buds of various
sizes; bracts caducous; female inflorescences smaller, 5-9 cm long, subumbels 1—5-
flowered; flowers coriaceous, thinly with brown-grey hairs less than 0.1 mm. Male flow-
ers: pedicel 3-10 mm long, bracteole broadly reniform, angular, or 2- to 3-topped, sub-
umbels 1.5—2 mm, with ciliate margin, persistent; buds (ovoid-)ellipsoid, 5-7 by 4—5
mm, cleft c. 1/2 or slightly over, lobes 0.7 mm thick. Androecium 3—4.5(—5) mm; andro-
phore 0.5—0.8(—1) by 0.8—1 mm, finely pubescent in the lower half; synandrium = cylin-
drical, 2-2.5 by 1-1.2 mm, thecae 16-24, sterile apex + blunt, 0.3-0.5 mm. Female
flowers: pedicel stout, 1-3(—4) mm long, bracteole broadly rounded, 2(—3) mm, persist-
ent; buds ovoid, 5—7 by 4—6 mm, cleft 1/3 to halfway; ovary ovoid, 3-4 mm, hairs ap-
pressed, 0.2—0.4 mm, stigma lobes (0.5—)1 mm long. Fruits 1—4 per infructescence, el-
lipsoid-oblong, (4.5—)6—9 by 3.5-5 cm, base sometimes narrowed, hairs mealy, dark
brown, 0.1 mm or less, sometimes late glabrescent; pericarp 10-15 mm thick; seeds
ellipsoid-oblong, (4—)4.5—6 cm, aril in lower 1/4 sometimes undivided; fruiting pedicel
stout, 3-10 mm long. — Fig. 60a, b, 62c.

Field-notes — Large tree, with or without buttresses, sometimes with stilt-roots or
with stilt-roots and buttresses; buttresses variable, up to 3 m out and high, + narrow;
crown deep and dense, pyramidal in outline, with numerous horizontal and descending
branches; twigs deep green. Bark smooth or rather rough, hard, shallowly scaly, dippled,
or shallowly fissured, grey-brown or chocolate; living bark 10-20 mm, usually reddish
brown, laminated, soft, with clear light to dark red exudate, copious or not; cambium
whitish; sapwood pale brown to whitish, also with red exudate; heartwood brown. Leaves
glossy dark green above, glaucous-grey or whitish below; Sinclair (1.c.: 133) measured
in a cultivated female tree leaves up to 54 cm long. Flowers (pale) yellow, in male pale
flesh-coloured (once), male may reach 8 mm, female 8—9 mm, ovary 5 mm long, fra-
grant or not. Fruits 9 cm long, (yellow-)brown or rusty brown; aril first yellow, then red.

Distribution — S Peninsular Thailand; Malesia: Sumatra, Peninsular Malaysia, Sin-
gapore, whole of Borneo.

Habitat & Ecology — Primary forest; seasonal swamp forest, forest along rocky Saraca
streams (granite rock); ridges; sandy loam, yellow clay loam, sandy and black soils,
shales, loam soil containing limestone, volcanic rock; 0—700 m altitude; fl. & fr. through-
out the year.

De Wilde — Myristicaceae (Myristica) 525

Note — Herbarium specimens generally have a markedly dark drying colour. The
leaves seem glabrous below but actually have a very minute indumentum of + scattered
very small scale-like grey or pale brown hairs, glabrescent or not.

89. Myristica mediovibex W. J. de Wilde

Myristica mediovibex W.J. de Wilde, Blumea 40 (1995) 301. — Type: Brass & Versteegh 13573,
Irian Jaya.

Tree 18-30 m. Twigs 1.5—3 mm diameter, somewhat angular, late glabrescent, hairs
shaggy, up to 0.5 mm long; older twigs sometimes blistering or flaking; lenticels small,
many or few. Leaves chartaceous, elliptic-oblong or oblong-lanceolate, (7-)9-17 by 2.5-
7.5 cm, base rounded or attenuate, apex acute-acuminate; upper surface dark brown,
lower surface brown-greyish, indumentum persistent (or late glabrescent), hairs mealy,
stellate scale-like, 0.2 mm or less; not papillose; dots absent; midrib raised above, lateral
nerves 13-17 per side, at c. 60° to the midrib, flat or sunken above, little raised below,
lines of interarching and venation not distinct; petiole 12-15 by 2-3 mm; leaf bud 8-18
by 1.5—3 mm, hairs rufous, 1—-1.5 mm long, + erect or appressed, brownish grey, 0.1 mm
(var. kosteriana) or longer. Inflorescences pedunculate, between the leaves or just be-
low, hairs 0.1—0.2 mm, male not seen; in female: peduncle 4-8 by 2 mm (to 4 mm wide
in infructescences), ending in | or 2 knob-like scar-covered brachyblasts 2 mm long,
each with 1-3 flowers, buds of slightly various sizes; bracts small, pubescent, caducous;
flowers subcoriaceous, with shaggy hairs of mixed sizes, (0.1—)0.2—0.3 mm. Male flow-
ers not seen. Female flowers (slightly immature): pedicel 7-8 by 1.5—2 mm, broadening
distally and + gradually passing into the perianth; bracteole | mm or less, caducous, scar
nearly ring-shaped, + median; buds subellipsoid, broadest below the middle, 4—4.5 by
2.5(—3) mm, cleft c. 1/6, lobes 0.5(—1) mm long, 0.5 mm thick; ovary ovoid-oblong, 325
by 1.5 mm, hairs 1-1.5 mm long. Fruits solitary or 2 or 3, ellipsoid-oblong, (2.5—)4 by
2 cm, apex + acute, base contracted into a rather stout pseudostalk or not, hairs dense,
mealy, brown, 0.1(—1) mm; pericarp + woody, 4 mm thick; seeds ellipsoid-oblong, 2.2
cm (var. mediovibex); stipe (peduncle and fruiting pedicel) + stout, 1.5 cm long, nearly
5 mm wide at base, striate, + lenticellate, bracteole scar slender, ring-shaped, 5 mm
below the fruit.

Distribution — Malesia: northern Papua Barat, locally with two varieties.

Note — The almost ring-shaped bracteole scar at about halfway female pedicel and
fruiting pedicel is characteristic.

KEY TO THE VARIETIES

la. Leaves below + late glabrescent ...~.. 02+ + 5204s se0++--5:> a. var. mediovibex
b. Leaves below with persistent indumentum ............-.--- b. var. kosteriana

a. var. mediovibex

Tree 20-27 m. Leaves below + late glabrescent; leaf bud with hairs up to 0.5 mm
long. Fruits 4 by 2 cm, apex acute and narrowly, 1.5 mm beaked, basal part contracted
into a rather stout pseudostalk 5 mm long; indumentum short, with the strong conspicu-
ous hairs as found on the ovary remaining on the apex and the base (pseudostalk).

526 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Medium-sized tree, dbh of bole c. 58 cm; buttresses to 1 m high, 20 cm
out, 3 cm thick, or not buttressed. Bark 14 mm thick, black, scaly, or not peeling or
fissured; exudate little, light red; slash dark red; wood red-brown; sapwood white. Fruits
(green-)brown.

Distribution — Malesia: Papua Barat (Idenburg River; Japen I.).

Habitat & Ecology — Locally frequent in primary forest on sandy soil; forest on
ridges; 130-550 m altitude; fl. & fr. Apr., Sept.—Oct.

Note — May be confounded with M. longipes, a local endemic of SE Papua New
Guinea, differing in short-haired indumentum of the leaf bud. Myristica mediovibex vat.
mediovibex is characterized by long-haired leaf bud, conspicuous short-felty subpersistent
indumentum on the lower leaf surface, coriaceous (female) flowers and hard, woody
fruits.

b. var. kosteriana W. J. de Wilde

Myristica mediovibex W. J. de Wilde var. kosteriana W. J. de Wilde, Blumea 40 (1995) 303. — Type:
BW 8084 (Ch. Koster), Irian Jaya.

Tree 18 m. Leaves below with persistant indumentum; leaf bud with hairs 0.1 mm or
less. Fruits (somewhat immature) + ovoid, 2.5 by 2 cm, apex + blunt, base without
pseudostalk, hairs at base shorter than in the typical variety.

Field-notes — Small to medium-sized tree, bole 15 m, dbh 45 cm, buttresses up to 1.5
m high, | m wide, thick 10 cm. Outer bark dark brown, not fissured, faintly peeling with
large scales; inner bark with much scarlet milky exudate, brown within; slash dark brown;
heartwood dark brown. Young fruits brown.

Distribution — Malesia: Papua Barat (Jayapura, 60 km SE of Sarmi, Bodem River),
Papua New Guinea (E Sepik).

Habitat & Ecology — Locally common in primary forest on clayey soil; 70-200 m
altitude; fr. Feb.

90. Myristica mediterranea W. J. de Wilde

Myristica mediterranea W.J. de Wilde, Blumea 40 (1995) 304. — Type: Soegeng Reksodihardjo
328, SW Irian Jaya.

Tree 5-8 m. Twigs (1.5—)2—3 mm diameter, early glabrescent, hairs grey-brown, scale-
like, less than 0.1 mm; older twigs corky, dark grey-brown, with longitudinal and trans-
verse cracks, but not much flaking; lenticels inconspicuous. Leaves membranous, (ellip-
tic-)oblong, 9-18(—20) by 2.5—6(—7) cm, base attenuate, apex acute-acuminate; upper
surface (blackish) brown, lower surface conspicuously pale, greyish, sometimes with a
yellow-brown discoloration, early glabrescent, indumentum minute; papillae present,
dense, but inconspicuous, unequal in size and irregularly spaced; dots absent; midrib
raised above, lateral nerves 13—20 per side, at c. 60° to the midrib, flat and faint above
and below, lines of interarching and venation indistinct; petiole short, 5-10 by 1.5—3
mm; leaf bud 5-10 by 1.5—2 mm, hairs grey-brown, less than 0.1 mm. Inflorescences
(from female inflorescences and infructescences): short-pubescent, between the leaves,
of the Knema-type: sessile, tubercle-like brachyblast, 1—-1.5 mm, 1—3-flowered; flowers
with dark brown hairs 0.1(—0.2) mm. Male flowers not seen. Female flowers (somewhat

De Wilde — Myristicaceae (Myristica) 527

immature): pedicel 1.5 by | mm, bracteole broadly rounded, | mm, (sub)apical, (sub)-
persistent; buds membranous, ovoid(-oblong), 3 by 1.8—2 mm, lobes 0.8 mm; ovary
ovoid, (including stigma) 2 by 1.5 mm, hairs less than 0.1 mm. Fruits single (or paired),
ellipsoid-oblong, 2.2-2.5 by 1—1.4 cm, apex short-acute, somewhat oblique, base (nar-
rowly) rounded, hairs dense, mealy (scurfy), rust-coloured, less than 0.1 mm; pericarp
1 mm thick; seeds ellipsoid-oblong, 2 cm; fruiting pedicel short, striate, 1.5—2 by 2 mm.

Field-notes — Small tree, dbh c. 5 cm; branches horizontal, verticillate. Bark smooth,
dark brown: inner bark aromatic. Leaves very pale green below. Fruits orange(-brown).

Distribution — Malesia: New Guinea; known only from three collections from the
southern border area between Papua Barat and Papua New Guinea, at Ingambit.

Habitat & Ecology — Disturbed forest on flat well-drained land, valley forest with
semi-swamp near Sagu patches; c. 150 m altitude; fl. & fr. June, July.

Notes — 1. Myristica mediterranea is characterized by the corky, deeply fissured
bark of the older twigs, the membranous leaves with very pale, whitish lower surface,
and the ellipsoid-oblong, short-stiped fruits.

2. Myristica mediterranea seems close to M. lancifolia, a species also with rather
closely set, faint, lateral nerves. Myristica lancifolia has smaller fruits, and the lower
leaf surface is more distinctly and regularly papillose, with the papillae equal in size.

91. Myristica millepunctata W. J. de Wilde

Myristica millepunctata W.J. de Wilde, Blumea 40 (1995) 305, f. 5. — Type: Soegeng Reksodi-
hardjo 545, E Irian Jaya.

Tree 30—40 m. Twigs sometimes subangular, (4-)5—7 mm diameter, early glabrescent,
hairs 0.1 mm or less; older twigs somewhat cracking, coarsely lenticellate. Leaves (thinly)
coriaceous (but brittle), (oblong-)lanceolate, 25-40 by 7-11 cm, base short-cuneate or
narrowly rounded, apex acute(-acuminate); upper surface olivaceous or blackish brown,
finely wrinkled, lower surface pale, grey-brown, late glabrescent, hairs dense, greyish,
scale-like, less than 0.1 mm; not papillose; dots present; midrib raised above, lateral
nerves 28-40 per side, at 70—80° to the midrib, usually interspaced by intersecondary
nerves, lines of interarching and venation indistinct or invisible; petiole 25—40 by 4-7
mm; leaf bud 20-25 by 4-5 mm, the hairs 0.1 mm or less. Inflorescences incompletely
known (from infructescences): peduncle short; male and female flowers not seen. In-
fructescence below the leaves, short, + woody, nearly 1 cm long, peduncle 5 by 8 mm,
cracked and lenticellate, and with some old flower scars at apex. Fruits solitary, rather
stout, broadly ovoid or subglobose, 5.5—6 by 4.5—5 cm, shortly narrowed at apex and
base, hairs fine, brown, mealy, 0.1 mm; pericarp hard, woody, 10-12 mm thick, outer
layer with scattered stone-cell-like concretions visible on section; seeds broadly ovoid-
ellipsoid, 2 cm; fruiting pedicel not discernible. — Fig. 80.

Field-notes — Tree with narrow crown. Bark fissured, 2 mm thick, dark brown; inner
(live) bark 15 mm, light brown or pinkish; wood pinkish straw; exudate red, sticky.
Leaves dull green above, pale bronze-green or glaucous below. Fruits brown.

Distribution — Malesia: central part of New Guinea (E Papua Barat, in the head-
water area of Ok Denim River and Oksibul River).

Habitat & Ecology — Montane and ridge forest; 1400-1600 m altitude; fr. June.

528 Flora Malesiana, Ser. I, Vol. 14 (2000)

Bee re 9
ora aaa
2 Ke AY

Fig. 80. Myristica millepunctata W.J. de Wilde. a & b. Leafy twig and part of apex with sterile
terminal leaf buds and axillary buds; c. leaf seen from below; d. detail of lower leaf surface with
scattered dots; e & f. fruit from outside and longitudinally opened [all NGF 33212]. — Scale bar for

a—c, e, f=2 cm; ford=0.4 mm.

De Wilde — Myristicaceae (Myristica) 529

Note — Related to M. neglecta. Mature fruits of M. neglecta are not known, the
immature fruit closely resembles that of M. millepunctata. Differences are found in the
appearance of the dried leaves and the quite different habitat, M. neglecta being a spe-
cies of lowland areas in Bird’s Head.

92. Myristica mindanaensis Warb.

Myristica mindanaensis Warb., Mon. Myrist. (1897) 497, t. 13; W.J. de Wilde, Blumea 35 (1990)
249, f. 1: 9; 42 (1997) 178. — Type: Warburg 13300, Mindanao.

Myristica fatua Houtt. var. morotaiensis J. Sinclair, Gard. Bull. Sing. 23 (1968) 292, f. 36A. —
Type: Kostermans 771, Morotai.

Tree 10—20 m. Twigs angular and 2-ridged, 2-2.5(—3) mm diameter, early glabrescent,
hairs less than 0.1 mm; older twigs sometimes faintly cracking; lenticels scattered. Leaves
membranous or chartaceous, (obovate-)elliptic or oblong-lanceolate, 14-35 by 6.5—-16
cm, base narrowly rounded or cuneate, apex acute-acuminate; upper surface olivaceous
or brown, lower surface greyish, glabrescent, hairs scattered or touching, scale-like, 0.1
mm or less, but leaving the surface visible; distinctly papillose or not; dots absent; midrib
slightly raised above, lateral lateral nerves 17—27 per side, at 45—70° to the midrib, flat
or sunken, lines of interarching and venation visible; petiole 15—25 by 2-4 mm; leaf bud
10-16 by 2-3 mm, hairs 0.1 mm. Inflorescences between the leaves or below, of the
Knema-type: sessile, simple or forked, woody, tubercle-like scar-covered brachyblast
up to 12 mm long, minutely pubescent; in male: with 6—20 flowers in subumbels, buds
unequal in size; in female: smaller, with 1 or 2(—4) flowers; flowers with hairs 0.2—0.3
mm long. Male flowers: pedicel 4-6 mm long, bracteole + acute, 2.5—3.5 mm,
subpersistent; buds ovoid(-oblong), apex subacute, 5—5.5 by 3-4 mm, cleft c. 1/4 to
nearly halfway (see note 2), lobes 0.2—0.3 mm thick. Androecitum 44.5 mm; androphore
1(—1.5) mm, glabrous or with few minute hairs towards base; synandrium 2.5 by 0.6—0.7
mm, thecae 12-14, sterile apex 0.2(—0.3) mm, irregular in shape, rather blunt. Female
flowers (Binnendijk s.n., Buru): pedicel 1-1.5 mm long, bracteole caducous; buds ovoid,
4 by 3-3.5 mm, cleft 1/4-1/3; ovary ovoid, 2.5 mm. Fruits solitary or paired, ovoid-
ellipsoid, (2.5—)3—3.5(—4.5) by 2-2.5(-3) cm, apex often shortly beaked, hairs scurfy,
pale or dark brown, 0.1—0.2 mm, partly (late) glabrescent; pericarp 2—3(-5) mm thick;
seeds ellipsoid, 2-2.5 cm; fruiting pedicel glabrescent, 2-6 mm long. — Fig. 641.

Field-notes — Erect tree; buttresses inconspicuous, prop-roots terete, c. 70 cm high
(once recorded). Bark blackish brown; outer bark 0.5—1.5 mm thick, longitudinally
fissured or cracked, or very fissured, not peeling off; inner bark 2.5—5 mm thick, beefy
red, turning brown(-red) on exposure; exudate clear, reddish; sapwood cream or yellow-
ish, gradually passing into slightly darker heartwood. Flowers fascicled, rust-coloured.
Fruits pale ochre or brown; seeds brown.

Distribution — Malesia: Philippines (Mindanao); Moluccas.

Habitat & Ecology — Lowland primary and degraded forest, tall forest with little
undergrowth, hill slopes; alluvial soil (inundated after heavy rains), sandy clay or deep
clayey soil (bedrock grey schists); 0-250(—700) m altitude; fl. & fr. throughout the year.

Notes — 1. A lowland species, differing from M. fatua in usually distinctly ridged
twig apices, much less developed indumentum on flowers and lower leaf surface (old
leaves glabrescent), short indumentum on leaf bud (hairs 0.1 mm long or less), smaller

530 Flora Malesiana, Ser. I, Vol. 14 (2000)

fruits with very short indumentum, and somewhat smaller flowers, opening at anthesis
to only c. 1/3 (but see note 2).

2. The male buds in the type and in other specimens from Mindanao are cleft as deep
as nearly halfway, in other material (e.g. Kuswata & Soepadmo 57, Seram) the buds are
more slender, opening at anthesis only to 1/4—1/3.

3. The papillose lower surface of the leaves is especially apparent in Moluccan
specimens from Bacan, Obi, Halmahera, and Seram.

93. Myristica nana W. J. de Wilde

Myristica nana W.J. de Wilde, Blumea 40 (1995) 307. — Type: LAE 50348 (P.F. Stevens), Papua
New Guinea.

Treelet, 1.5—8 m. Twigs 1-1.5 mm diameter, early glabrescent, hairs less than 0.1 mm;
older twigs neither cracking nor flaking; lenticels inconspicuous or absent. Leaves mem-
branous, (elliptic-)oblong or lanceolate, 6—14 by 1.5—4 cm, base attenuate, apex acute-
acuminate, the very tip often rather blunt; upper surface olivaceous or brown, lower
surface grey-brown, (late) glabrescent, hairs scattered, less than 0.1 mm, not distinctly
papillose; dots absent; midrib slightly raised above, lateral nerves 11-17 per side, at
c. 70° to the midrib, flat, indistinct above, below reddish brown, indistinct, lines of
interarching and venation indistinct; petiole 7-12 by 1(-1.5) mm; leaf bud 5-6 by
(0.5—)1 mm, hairs 0.1 mm or less. Inflorescences between the leaves, of the Knema-
type: sessile or to 1.5 mm pedunculate, slender, simple or forked scar-covered brachyblasts
to 6 mm long, glabrescent; in male: ending in | or 2 loose clusters each with |—3 flow-
ers, buds of various sizes; bracts 0.5 mm, short-pubescent, caducous; female inflores-
cences consisting of merely one axillary flower; flowers with hairs 0.1 mm or less. Male
flowers: pedicel 3—4.5 by 0.4 mm, bracteole | mm, persistent, 1-2 mm below apex;
buds (submature) ovoid-oblong, 3.5(—4) by | mm, apex narrowly rounded, cleft c. 1/4,
lobes membranous, | mm long, 0.2 mm thick. Androecium 3.5 mm; androphore 1.7 by
0.3 mm, with hairs 0.1 mm all over; synandrium 1.8 by 0.4 mm, thecae 6, sterile apex
0.1 mm. Female flowers: pedicel slender, 5 mm long, bracteole scar about halfway; buds
ovoid-oblong, 3.5 by 1.5(—2) mm, lobes 0.7 mm; ovary 2.5 by 1(-1.5) mm, minutely
pubescent. Fruits solitary, + ellipsoid-oblong, 3-3.5 by 1-1.3 cm, apex subacute with
minute apiculum (stigma remnant), the basal part contracted into a narrow pseudostalk,
10 mm long, hairs dull brown, 0.1 mm or less; pericarp | mm thick or less; seeds ellip-
soid, 1.7-2 cm; fruiting pedicel slender, 8-12 by 0.5 mm, bracteole scar at or below the
middle.

Field-notes — Treelet, dbh c. 4 cm. Bark smooth, finely fissured, dark brown, grey, or
blackish; inner bark straw; sap dark red; wood whitish. Leaves glaucous below. Flowers
cream or greenish white; female flower buds cream, pale green at tip. Fruits (mature)
greenish with brown scurf or olive-green; seeds brown.

Distribution — Malesia: eastern Papua New Guinea (Central and Milne Bay Prov.);
locally endemic.

Habitat & Ecology — Flat-land and sloping forest, ridge or level forest; 400-750 m
altitude; fl. & fr. July, Aug.

Notes — 1. Myristica nana resembles M. pumila, but is distinguished by smaller
male flowers and a shorter fruiting pedicel.

De Wilde — Myristicaceae (Myristica) 531

2. Myristica nana is also superficially similar and possibly related to M. ensifolia
(from Western Province) in its narrow leaves. The latter, known only from one fruiting
specimen, differs in the still more elongate leaves, the venation and lower leaf surface
texture (lens!), and in the infructescence with stouter stalk, including a short peduncle.

94. Myristica neglecta Warb.

Myristica neglecta Warb., Mon. Myrist. (1897) 542, t. 17; Markgr., Bot. Jahrb. Syst. 67 (1935) 170;
J. Sinclair, Gard. Bull. Sing. 23 (1968) 154, f. 6; Foreman, Contr. Herb. Austral. no. 9 (1974) 40;
Handb. Fl. Papua New Guinea | (1978) 206, p.p., excl. fruiting material; W. J. de Wilde, Blumea
40 (1995) 309. — Syntypes: Beccari FI acc. nos. 7702, 7703 (female), 7704 (male), Bird’s
Head.

Tree 25 m. Twigs stout, 5-7 mm diameter, early glabrescent, hairs less than 0.1 mm;
older twigs incompletely known, possibly somewhat fissured but not distinctly flaking,
with a few lenticels. Leaves (thinly) coriaceous (but very brittle), 21-40 by 6-12 cm,
base narrowly rounded, apex acute; upper surface brown, smooth, lower surface slightly
paler, glabrous; not papillose; dots present (lens!); midrib slightly raised above, lateral
nerves 20—40 per side, at 60—70° to the midrib, flat and faint above, hardly visible be-
low, lines of interarching and venation invisible; petiole 30 by 4—5 mm; leaf bud 10 by 4
mm, hairs appressed, grey-brown, less than 0.1 mm. Inflorescences between the leaves,
minutely brown pubescent, late glabrescent, with pedunculate (surface may be fissured)
brachyblast; bracts 2—2.5 mm, caducous; in male (van Royen 3498): peduncle S—15 mm,
with transverse lenticels, scar-covered brachyblast to 10 mm, with subumbel of 3-6
flowers, buds of various sizes; female inflorescences (incompletely known, Beccari 7703):
peduncle 5 mm long, brachyblast to 10 mm long, ending in a subumbel of 2 or 3(-5)
flowers; flowers leathery with hairs 0.1-0.2 mm. Male flowers: pedicel 10-15 by 1.5—
2.5 mm, bracteole scar conspicuous, 2-3 mm below the apex; buds ellipsoid-oblong,
9-11 by 4-5 mm (according to Warburg and Sinclair 13-14 by 6-8 mm), cleft c. 1/5,
lobes 2—2.5 mm long, 0.5 mm thick. Androecium + stout, broad-cylindrical, 7 mm long;
androphore 2—2.5 by 1.5—2(—2.5) mm, entirely with hairs 0.3—-0.5 mm; synandrium 2.5—
2.5 by 1.5-2 mm, gradually narrowed into acute sterile apex, 1-1.5 mm, thecae c. 20:
Female flowers: pedicel stout, 5-6 mm long, bracteole scar 1-2 mm below the apex;
buds ovoid(-oblong), narrowed towards the apex, 9 by 5 mm, lobes 2(—2.5) mm; ovary
ovoid-oblong, tapering, 7 by 3.5 mm, hairs 0.5 mm. Fruits (immature, see note 2) broadly
fusiform, 3 by 2 cm, hairs mealy, 0.3(-0.5) mm; pericarp hard, 8(—10) mm thick; seeds
not seen; fruiting pedicel (immature) 7 by 3 mm.

Field-notes — Bark brown, with long flakes; wood reddish brown, with a soft acid
smell. Leaves dark green above, greyish green below. Flower buds yellow-brown.

Distribution — Malesia: Papua Barat (Bird’s Head).

Habitat & Ecology — Primary forest dominated by Quercus and Canarium; c. 15 m
altitude; fl. Apr.

Notes — 1. The name M. neglecta should be restricted to plants from the lowland
area of Bird’s Head, i.e., the syntypes and one additional collection, van Royen 3498.
This means that the description of the fruit by Foreman (1974: 40; 1978: 206) is ex-
cluded; this fruiting material is transferred to M. millepunctata. Other related species are
M. brassii, M. duplopunctata, and M. tamrauensis.

582 Flora Malesiana, Ser. I, Vol. 14 (2000)

2. Van Royen 3498 (L), with male flowers, has mounted on the sheet an envelope con-
taining a loose female inflorescence and three immature fruits, 3 cm long, which must
have been taken from a different tree.

95. Myristica nivea Merr.

Myristica nivea Merr., Philipp. J. Sci. 1, Suppl. (1906) 191; Enum. Philipp. Flow. Pl 21923) 179;
W.J. de Wilde, Blumea 42 (1997) 178. — Type: FB 3454 (Hutchinson), Philippines, Basilan.

Tree 10-15 m. Twigs 2—3(—4) mm diameter, glabrescent, hairs greyish, 0.1(—0.2) mm;
older twigs (coarsely) striate; lenticels few, inconspicuous. Leaves chartaceous, elliptic-
oblong or oblong-lanceolate, 15—43 by 4.5—13 cm, base narrowly to broadly rounded,
apex acute or acuminate; upper surface olivaceous or (blackish) brown, lower surface
grey-white (possibly by dense irregular papillae), glabrescent, hairs scale-like, 0.1(—0.2)
mm, stouter hairs leaving point-like scars; (larger) dots absent; midrib above slightly
raised, lateral nerves 13-20 per side, at 45—70° to the midrib, sunken above, dark brown,
much contrasting in colour below, lines of interarching and venation faint; petiole 10-20
by 2-4 mm; leaf bud 10-15 by 1.5-3 mm, hairs appressed, 0.1 mm. Inflorescences
+ below the leaves, of the Knema-type: sessile, simple or forked, scar-covered, woody,
brachyblasts up to 4 mm, with short brown hairs; in male: with 2-5 flowers, buds of
various sizes; in female (from infructescences, BS 37070): similar to male, few-flow-
ered: flowers with hairs 0.1-0.2 mm. Male flowers: pedicel 2.5—3 mm, bracteole 2—2.5
mm, persistent; buds ovoid-ellipsoid, 6 by 4 mm, apex narrowly rounded or subacute,
cleft (1/4—)1/3, lobes (0.2-)0.3 mm thick. Androecium broad, 4—4.5 mm, androphore
1.5 by 0.6—-0.7 mm, with few pale brown hairs 0.1—0.2 mm towards the base; synandrium
2.5-3 by 1.3-1.5 mm, thecae c. 20, sterile apex + blunt, glabrous, 0.3 mm. Female flow-
ers not seen. Fruits solitary, subsessile, ellipsoid, 3.5—4 by 2.5—3 cm, hairs dense, dark
rust-coloured, scurfy, 0.1 mm or less; pericarp 4 mm thick; submature seeds ellipsoid,
2.5—3 cm; fruiting pedicel rather stout, 4 mm long.

Distribution — Malesia: Philippines (Mindanao: Zamboanga Prov.; Basilan I.);
Moluccas (Talaud Is.).

Habitat & Ecology — Forest at low altitude; fl. Dec.; fr. Oct —Nov., Mar.

Note — Sinclair [Gard. Bull. Sing. 23 (1968) 268, 273] included M. nivea in M. fatua,
the latter from the same area in the Philippines. However, M. nivea differs in the struc-
ture of the lower leaf surface (in M. fatua with dense and more woolly persistent
indumentum, not leaving scattered dark point-like hair scars), and the indumentum of
the fruits (in M. fatua dense, velvety, hairs 0.5—1 mm). Myristica nivea seems also close
to M. impressa and M. koordersii, two species with the leaves below whitish as well.

96. Myristica olivacea W. J. de Wilde

Myristica olivacea W.J. de Wilde, Blumea 40 (1995) 309. — Type: Pullen 7663, Papua New Guinea,
Central Prov.

Small undergrowth tree, 2.5—4.5 m. Twigs 1-2 mm diameter, early glabrescent, smooth,
hairs 0.2 mm; older twigs yellowish brown, not or only faintly cracking or flaking; lenticels
not seen. Leaves membranous, oblong-lanceolate, 10-30 by 2.5—7.5 cm, base attenuate,
apex acute-acuminate or + blunt; upper surface conspicuously olive-green, lower sur-

De Wilde — Myristicaceae (Myristica) 3)3)3)

face early glabrescent, hairs minute, scale-like, leaving minute crater-like scars; not pap-
illose; dots absent; midrib yellowish, slightly raised above, much raised below, lateral
nerves 10-15 per side, at 60—-80° to the midrib, faint above, lines of interarching and
venation faint or absent; petiole 20-30 by 1.5-2.5 mm; leaf bud 5—10 by 1 mm, hairs
appressed, 0.2(—0.3) mm. Inflorescences between the leaves, axillary or to 7 mm supra-
axillary, + paniculate, with slender peduncle, early glabrescent; in male 6-10 cm long,
peduncle 3—5 cm, lower branches 1—2.5 cm, central axis 1-3 cm, with 1—3 side branches
to 1 cm long, all branches ending in scar-covered brachyblasts to 5 mm, each with a
subumbel of 4—10 flowers, buds of various sizes; bracts small, 1-2 mm, late caducous;
in female: peduncle 2.5—3 cm, shorter and less branched than the male, subumbels 2—4-
flowered; flowers with sparse, dark-brown, appressed hairs, partly glabrescent. Male
flowers: pedicel 10-12 by 0.4-0.6 mm, bracteole 0.5—1 mm, half-circular, persistent,
subapical; buds membranous, elongate, almost tubiform, narrowed to both ends, especi-
ally towards the apex, 12-13 by 2.5(-3) mm, cleft c. 1/10, lobes 1-1.5 mm, erect, 0.2
mm thick. Androecium slender, 8—9 mm long; androphore 3—3.5 by 0.5 mm, hairs less
than 0.1 mm; synandrium 4.5-5.5 by 0.6-0.8 mm, thecae 10-12, sterile apex 0.2—0.3
mm, with 5 or 6 lobules corresponding with the anthers. Female flowers: pedicel rather
stout, 18-20 mm, bracteole 2-5 mm below the apex; buds ovoid-oblong, narrowed to-
wards the apex, 9 by 3-3.5 mm, lobes | mm long; ovary ovoid-oblong, 6—7 by 2-3 mm
(including the 1 mm long deeply 2-lobed stigma), hairs dense, pale brown, less than 0.1
mm. Fruits single, oblong, 3—3.5 by 1-1.3 cm, top acute, base + narrowed into 3-5 mm
long pseudostalk, hairs scale-like, 0.1 mm; pericarp 1 mm thick; seeds 2—2.5 cm.

Field-notes — Undergrowth tree, dbh c. 4 cm. Bark shallowly fissured, dark grey or
black; blaze with red exudate. Leaves green, midrib yellowish. Inflorescence + pendent;
flowers cream.

Distribution — Malesia: Papua New Guinea (Central Prov.); known from three col-
lections.

Habitat & Ecology — Rain forest on low hills; clayey soil; 75-120 m altitude; fl.
June.

Notes — 1. Myristica olivacea is close to M. cornutiflora from which it differs chiefly
by its more slender habit. The twigs dry yellowish, the leaves conspicuously olivaceous-
green. The flowers are similar to those of M. cornutiflora subsp. elegans, but differ in
the persistent bracteole and longer androecium in proportion to the perianth. The leaves
below apparently lack the small dark brown dashes and dots, but possibly the latter are
represented as very small scattered points, only visible here and there (lens!).

2. The stomata on the lower leaf surface are deeply sunken in the surrounding tissue,
which apparently is not papillose.

97. Myristica ornata W. J. de Wilde

Myristica ornata W.J. de Wilde, Blumea 40 (1995) 310. — Type: LAE 51752 (Streimann & Katik),
Papua New Guinea.

Tree 25-30 m. Twigs almost angled, + yellowish brown, 2.5(—3) mm diameter, very
early glabrescent, hairs scattered, scale-like, less than 0.1 mm; older twigs tending to
flake, with few brownish lenticels. Leaves subcoriaceous, (elliptic-)oblong, 15—20 by
3.5-6.5 cm, base cuneate, apex acute; upper surface olivaceous or yellowish brown,

534 Flora Malesiana, Ser. I, Vol. 14 (2000)

lower surface slightly paler, glabrous; papillose (lens!); dots absent; midrib slightly raised
above, lateral nerves 15-20 per side, at 45—60° to the midrib, slender, flat and faint
above, faint below, lines of interarching and venation inconspicuous; petiole 15-20
(—25) by 2.5-3 mm, early glabrescent; leaf bud 15 by 3-4 mm, longitudinally cracked
on drying, hairs dense, yellow-brown, appressed, (0.1—)0.3 mm. Inflorescences known
only from the infructescences. Male and female flowers not seen. Infructescences below
the leaves, peduncle 5-10 by (4-)5—6 mm, glabrescent, with coarsely flaking bark, end-
ing in one or two fruiting pedicel(s). Fruits (immature) solitary or paired, broadly fusi-
form, 5—6 by 3—3.5 cm, base narrowed into 5(—7) mm long pseudostalk, apex + gradu-
ally tapered, acute, hairs light brown, mealy, 0.3(—0.4) mm; pericarp 10 mm thick; seeds
not seen; fruiting pedicel 10 mm, fissured and with lenticels or coarsely flaking, bracteole
scar about median.

Field-notes — Bole straight, slender, 23 m, dbh 50 cm. Bark dark grey, slightly verti-
cally cracked; middle bark deep red; inner reddish; exudate red; wood of moderate weight
and hardness; sapwood straw; heartwood reddish, stained by sap. Leaves with thick
margin slightly curved inwards, dull dark green above, green below. Fruits rusty brown.

Distribution — Malesia: Papua New Guinea (Western Prov., Kiunga area); known
only from the type.

Habitat & Ecology — Lowland rain forest flats, c. 50 m altitude; fr. Aug.

Notes — |. Keys out beside M. schleinitzii on account of pedunculate inflorescences
and papillate lower leaf surface, but otherwise quite different.

2. Myristica ornata is reminiscent of M. mediovibex (also with the scar of the bracte-
ole + halfway the fruiting pedicel), and especially of M. millepunctata and M. neglecta;
these latter, however, with the lower leaf surface non-papillose and with scattered dots.

98. Myristica ovicarpa W. J. de Wilde

Myristica ovicarpa W.J. de Wilde, Blumea 40 (1995) 311. — Type: LAE 74587 (Damas & Katik),
Papua New Guinea, Rossel I.

Tree 25 m. Twigs 1—2(—2.5) mm diameter, early glabrescent, hairs greyish, less than
0.1 mm; older twigs neither cracking nor flaking; lenticels inconspicuous or absent.
Leaves membranous, (elliptic-)oblong, 6—10.5 by 2.5—4 cm, base broadly attenuate, apex
subacute or acute-acuminate; upper surface blackish brown, lower surface brown, early
glabrescent, hairs less than 0.1 mm; not papillose; dots absent; midrib almost flat above,
lateral nerves 25-30 per side, at 70—80° to the midrib, slender, flat and faint above, faint
below, lines of interarching and venation indistinct; petiole 10-18 by I-1.5 mm; leaf
bud 6-10 by 1 mm, hairs less than 0.1 mm. Inflorescences (from infructescence): ses-
sile, short, of the Knema-type, possibly a single axillary flower only, with the pedicel
4 mm long or less. Male and female flowers not seen. Fruits (apparently of mature size,
but seeds not yet fully developed, i.e. shrunken within the aril): axillary to a lower leaf,
solitary, ovoid-ellipsoid, 7 by 5 cm, apex and base rounded, glabrous, except for some
minute appressed brownish hairs less than 0.1 mm (hence ovary minutely pubescent)
towards the insertion of the fruiting pedicel; pericarp 2—2.5 cm thick; seeds (immature)
ellipsoid, 2 cm; fruiting pedicel short, rather thick, 4 mm long, 5 mm thick, brown, lon-
gitudinally cracked.

De Wilde — Myristicaceae (Myristica) 535

Field-notes — Large tree, bole 20 m, dbh 50 cm. Outer bark greenish black; inner
reddish, fibrous or slightly corky, exudate red on exposure; wood cream. Leaves gla-
brous, dark green above, light green below, growing tips rusty brown. Fruits axial, soli-
tary, large, egg-shaped, light green-yellow.

Distribution — Malesia: E Papua New Guinea (Rossel I.: Mt Dom, Pambwa); known
only from the type.

Habitat & Ecology — Lowland hill forest; c. 200 m altitude; fr. Mar.

Note — This species can be distinguished from other small-leaved Myristicas in the
area by the glabrous nature of all parts, the many-nerved leaves, and the large short-
stiped glabrescent fruits with very thick pericarp.

99. Myristica pachycarpidia W. J. de Wilde

Myristica pachycarpidia W. J. de Wilde, Blumea 40 (1995) 312; 43 (1998) 180. — Type: Brass
23293, Papua New Guinea, Mt Dayman.

Tree c. 25 m. Twigs with two low wings or sharp ridges between the petioles, 4-5 mm
diameter, early glabrescent, hairs less than 0.1 mm; older twigs tending to crack and
flake; lenticels inconspicuous. Leaves thinly chartaceous, elliptic-oblong or oblong-lan-
ceolate, 12-25 by 4.5-10 cm, base rounded, apex + blunt or subacute; upper surface
(blackish) brown, lower surface grey-brown, seemingly glabrous, hairs scattered, whit-
ish, scale-like, less than 0.1 mm; not papillose, dots absent; midrib slightly raised above,
lateral nerves 20-30 per side, faint and flat above, lines of interarching and venation
indistinct, somewhat impressed above, and rendering the blade slightly bullate; petiole
15-20 by 2.5—4 mm; leaf bud 10 by 2 mm, seemingly glabrous, but hairs appressed,
grey-white, less than 0.1 mm. /nflorescences (from infructescences): just below the leaves,
of the Knema-type: sessile, small, glabrous. Male and female flowers not known. Fruits
(somewhat immature) solitary, subglobose, 5(—6) by 4—5 cm, strongly wrinkled, apex
+ acute, hairs + sparse, dull brown, less than 0.1 mm, with a tendency to become gla-
brous; pericarp 2 cm thick; seeds (immature) subglobose or broadly ellipsoid, 2 cm;
fruiting pedicel stout, straight, 10-20 by 4—6 mm, glabrescent, hairs minute, scattered,
and scale-like, + warty or lenticellate, bracteole scar at 1/3 to halfway below the apex,
perianth scar conspicuous, apical.

Field-notes — Leaves grey below. Unripe fruits subglobose, 5—6 cm diameter.

Distribution — Malesia: Papua New Guinea (Milne Bay Prov., Mt Dayman); known
only from the type.

Habitat & Ecology — Occasional in oak forest of north slopes; 1550 m altitude; fr.
July.

Note — Ignoring its provenance from 1550 m altitude, M. pachycarpidia may key out
beside M. sulcata, a lowland species with smaller fruits. Similar also is M. bialata from
lowland areas in the Bismarck Archipelago but that species has twigs with ant-swell-
ings.

The (immature) fruits of M. pachycarpidia may resemble those of M. sphaerosperma,
also a montane species, with terete twigs and a much more conspicuous indumentum on
the lower leaf surface.

536 Flora Malesiana, Ser. I, Vol. 14 (2000)

100. Myristica pachyphylla A.C. Sm.

Myristica pachyphylla A.C. Sm., J. Arnold Arbor. 22 (1941) 69; W. J. de Wilde, Blumea 40 (1995)
313. — Type: Brass 12173, Papua New Guinea.

Tree 5-22 m. Twigs (1-)1.5-2.5 mm diameter, glabrescent, hairs 0.1 mm or less;
older twigs neither cracking nor flaking; lenticels small, indistinct. Leaves chartaceous
or subcoriaceous, ovate- or elliptic-oblong, (4-)6-14 by (1.5—)2-5.5 cm, base (short)
attenuate, apex to 2 cm acute-acuminate; upper surface olivaceous or brown, lower sur-
face somewhat paler, glabrescent, hairs sparse, cobweb-like, greyish, less than 0.1 mm;
not papillose; dots present, of various sizes; midrib raised above, lateral nerves 10-16
per side, at 60—80° to the midrib, interspaced by intersecondary nerves, fine, sunken
above, + flat and faint below, lines of interarching and venation thin, sunken above,
indistinct; petiole 7-15 by 1-2 mm; leaf bud 10 by 1.5 mm, hairs 0.1 mm or less. Inflo-
rescences between the leaves, minutely pubescent, pedunculate (sometimes subsessile
in female), ending in a simple or forked scar-covered brachyblast; in male: peduncle
3-8 by 1 mm, brachyblast up to 5 mm with a subumbel of 2-5 flowers, buds of various
sizes; bracts | mm, rusty-pubescent, late caducous; in female: sessile or to 10 mm pe-
dunculate, ending in 1 (or 2) flower(s); flowers with sparse hairs 0.1 mm or less. Male
flowers: pedicel 2—4 by 0.5-0.7 mm, bracteole ovate, 0.5—1 mm, caducous, subapical;
buds carnose, oblong-cylindrical, 6-8 by 2(-2.5) mm, apex (sub)acute, base shortly nar-
rowed, cleft 1/4—1/5, lobes 1.5 mm long, 0.3 mm thick. Androecium slender, 4—4.5 mm;
androphore 2.5 by 0.6(—0.7) mm, with few hairs less than 0.1 mm at base; synandrium
1.5 by 0.7 mm, thecae 10-12, sterile apex 0.1—0.2 mm long, blunt. Female flowers:
pedicel 7-10 mm, bracteole caducous, scar distinct, 2-3 mm below the apex, the part
above the scar thickened; buds elongate, larger than in male, 7-10 by 2.5—3 mm, nar-
rowed towards the apex, cleft 1/6(-1/8), lobes 1-1.5 mm long; ovary elongate, 6 by
2 mm, with silky hairs 0.3 mm. Fruits single (rarely 2), + ellipsoid, 3.5-4(—4.5) by 2
(—2.2) cm, apex + acute or somewhat beaked, base narrowed into a broad pseudostalk
5 mm long, hairs rust-coloured, scurfy, 0.1 mm or less; pericarp 2-3 mm thick; seeds
ellipsoid(-oblong), 2.5 cm; stipe (including fruiting pedicel) straight, 20-25 by 1-2 mm,
with scars of bract, bracteole, and perianth visible.

Field-notes — Slender trees. Outer bark black; inner bark pale, cream; wood white.
Leaves glossy green above. Flowers cream-green. Fruits distinctly beaked or with acute
apex, orange(-red) or fawn.

Distribution — Malesia: montane areas of Central West and East New Guinea.

Habitat & Ecology — Montane forest, on slopes and ridges; laurel ridge or oak forest;
steep limestone hillsides with Nothofagus forest, mossy forest with Myrtaceae,
Papuacedrus, Podocarpaceae; 1400-2000 m altitude; fl. & fr. July—Nov., Jan.

Notes — 1. Myristica pachyphylla is similar to M. arfakensis, both with characteris-
tic scattered dots on the lower leaf surface. May also be confused with M. longipes.

2. A deviating specimen is BW 13780 (Koster) (with fruits, in L broken off and lack-
ing) from Wandammen Peninsula, at 920 m altitude. In the absence of fruits it keys out
near M. pachyphylla or M. arfakensis and is characterized by: twigs slender, leaves nar-
row, 10 cm, long-acuminate, conspicuously greenish, dotted below, and peduncle of
infructescence slender, 15—20 mm long. It may represent an as yet undescribed species,
but more material is needed.

De Wilde — Myristicaceae (Myristica) Soy

101. Myristica papillatifolia W. J. de Wilde

Myristica papillatifolia W.J. de Wilde, Blumea 40 (1995) 313. — Type: NGF 33366 (Ridsdale &
Galore), Papua New Guinea, Western Prov.

Small tree c. 4 m. Twigs triangular, 1.5—2(—3) mm diameter, early glabrescent, hairs
dense, less than 0.1 mm; older twigs neither cracking nor flaking; lenticels inconspicu-
ous or absent. Leaves membranous, elliptic-oblong, 6-11 by 1.5—3 cm, base long-atten-
uate, apex acute(-acuminate); upper surface (blackish) brown, lower suface light brown
or greyish, very early glabrescent, hairs few, scattered, whitish, less than 0.1 mm; papil-
lose (lens!); dots absent; midrib slightly raised above, lateral nerves 10—13 per side, at
c. 45° to the midrib, fine, + sunken above, on lower surface brownish, lines of interarching
indistinct, nerves and venation in old leaves contrasting in colour; petiole 7-10 by 1—1.5
mm; leaf bud 6-10 by 1.5(—2) mm, hairs less than 0.1 mm. Inflorescences (from in-
fructescence) of the Knema-type: a sessile, simple, short tubercle-like brachyblast. Male
and female flowers not seen. Fruits axillary to leaf scars below the leaves, solitary, ellip-
soid-fusiform, 3—3.3 by 1.5—1.7 cm, apex narrowly 2(—3) mm beaked, base narrowed
into a slender 5(—7) mm long pseudostalk, hairs dull, mealy, grey-brown, interwoven,
less than 0.1 mm, appearing as if glabrous; pericarp 1(—2) mm thick; seeds broad-ellip-
soid, blackish brown, 1.5—1.7 cm; fruiting pedicel slender, 15—17 by 1.5 mm, bracteole
scar at (4—)5 mm below the apex, part above the scar smooth, below coarsely striate or
cracked.

Field-notes — Small tree. Mature fruit orange-brown.

Distribution — Malesia: Central Papua New Guinea (Western Province, Kiunga sub-
prov., near Ingembit (Ingambet, border area); known only from the type.

Habitat & Ecology — Valley forest, semi-swamp with sago patches; c. 130 m alti-
tude; mature fr. July.

Note — Distinguishable among the smaller-leaved New Guinean Myristicas with
Knema-type inflorescences by the leaves with lateral nerves steeply running from the
midrib (c. 45°), the distinctly papillose lower surface, and the delicately built, broadly
fusiform fruits, conspicuously narrowed at both ends, with a thin pericarp bearing a very
thin indumentum, almost as if it is glabrous.

102. Myristica papyracea J. Sinclair

Myristica papyracea J. Sinclair, Gard. Bull. Sing. 23 (1968) 133, f. 1; W.J. de Wilde, Blumea 42
(1997) 178: Tree Fl. Sabah & Sarawak 3 (2000) 468. — Type: SAN A 4775 (Wood), Sabah.

Tree 20-40 m. Twigs somewhat angular, 4-8 mm diameter, early to late glabrescent,
hairs 0.1 mm or less; older twigs coarsely flaking; with or without lenticels. Leaves
(thinly) coriaceous, elliptic-oblong or oblong-oblanceolate, 16-44 by 8—14.5(-18) cm,
base narrowly rounded or subcordate, apex + blunt (rounded) or acute; upper surface
olivaceous(-brown); lower surface yellowish, glabrous; papillose; dots absent; midrib
above slightly raised, lateral nerves (16—)20—28 per side, at 45—60° to the midrib, sunken
or flat above, lines of interarching sometimes distinct, venation fine-scalariform, dis-
tinct or not; petiole 20-50 by 3.5-5 mm; leaf bud 15—25 by 3.5—5 mm, hairs 0.1 mm or
less. Inflorescences between the leaves or below, paniculate, pedunculate, hairs mealy,
0.1—0.2(—0.3) mm; in male: rather stout, 2-5 by 2—4 cm, peduncle 8—20 mm, lower

aw We
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Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 81. Myristica papyracea J. Sinclair. a. Leafy twig; b. twig with male inflorescence; c. male
bud; note bracteole scar; d. male flower at anthesis; e. androecium (sterile apex not drawn); f. twig
with female inflorescence; g. female flower at anthesis; note bracteole scar; h. pistil; i. fruit; j. long-
ellipsoid seed with aril (a, i: § 18329 (Ashton); b-e: SAN 31080 (J. Singh); f-h: Cult. Hort. Bog. XF
55 (n.v.); j: Kostermans 5732}. — Scale bar for a, b, f, i,j =2cm; force, d,g=5 mm; fore, h=2 mm.

De Wilde — Myristicaceae (Myristica) 539

branches to 15 mm, central axis similar, unbranched, each branch ending in a short
brachyblast, with few flowers, size of buds strongly variable, in older inflorescences
with dispersed scars of fallen flowers; bracts 2~3(—4) mm, caducous; female inflores-
cences like male but smaller; flowers coriaceous, with hairs 0.1 mm, sometimes late
glabrescent. Male flowers: pedicel 5-8 mm, bracteole broadly reniform, 1.5—2 mm,
caducous, apical or to 2 mm below; buds ellipsoid, 6-8 by 4-5 mm, cleft 2/3—3/4, lobes
0.5—1 mm thick. Androecium (4—)5—6 mm; androphore somewhat tapering, 1—2 by 1.5
(—2) mm, hairs dense, 0.1 mm; synandrium oblong, (2—)2.5—3 by 1.5 mm, thecae 18-24,
sterile apex + blunt or subacute, 0.3—0.6 mm. Female flowers: pedicel 2-4 mm, bracte-
ole scar apical; buds ovoid, 6 by 4 mm, cleft 2/3—3/4; ovary (3—)4.5 by 3-3.5 mm, hairs
0.1 mm or less, stigma lobes sessile. Infructescences 1-3 cm long. Fruits | or 2, ellip-
soid, 7-9 by 4.5—6 cm, with the style remnant + lateral (always?), glabrescent, indumen-
tum mealy, dark brown, hairs 0.1 mm (persistent at the very base); pericarp glossy blackish
brown, 10-20 mm thick: seeds long-ellipsoid, 5—6 cm; fruiting pedicel stout, short,
4-10 mm long. — Fig. 81.

Field-notes — Large tree, often flanged and/or with conspicuous (laterally compres-
sed) stilt-roots to 2 m high. Bark dark grey or blackish brown, + cracked, in old trees
flaking in thin papery strips; sap pink, not copious; living bark 8-13 mm thick, red-
brown; wood pale (red-)brown, or white streaked with red. Leaves dark green glossy
above, conspicuously pale, yellowish green below. Flowers (pale) yellow. Fruits on twigs
and branches, yellow or pale apricot-coloured, glabrescent from minute powdery indu-
mentum.

Distribution — Malesia: Borneo (Sarawak, E & NE Kalimantan). No recent collec-
tions known.

Habitat & Ecology — A species of tall mixed dipterocarp forest on better soils; yel-
low clayey soil, loamsoil with coral limestone; along rivulet, wooded hillside, foot of
hill; 0-300 m altitude; fl. & fr. throughout the year.

Note —A lofty tree, apparently related to M. maxima, the latter differing in the lower
leaf surfaces which are (brownish) grey on drying, with subpersistent very thin indu-
mentum, not obviously papillose, larger inflorescences, somewhat more delicate flow-
ers, the androphore pubescent only in the lower half, and indumentum of the fruits longer
persistent.

103. Myristica pedicellata J. Sinclair

Myristica pedicellata J. Sinclair, Gard. Bull. Sing. 23 (1968) 324, f. 45; Foreman, Handb. Fl. Papua
New Guinea 1 (1978) 206; W. J. de Wilde, Blumea 40 (1995) 314. — Type: Pullen 944, Papua
New Guinea.

Tree c. 30 mtall. Twigs 2-3(—5) mm diameter, glabrescent, hairs 0.1 mm or less; older
twigs brown, coarsely striate, with small paler lenticels. Leaves chartaceous, oblong-
lanceolate, 12-21 by 3—6 cm, base short-attenuate, apex acute-acuminate; upper surface
brown, lower surface dull cinnamon or light brown, hairs persistent, dense, pale brown
(greyish when older), scale-like, 0.1 mm or less; not papillose; dots absent; midrib flat
above, lateral nerves 20-28 per side, at 50—70° to the midrib, slightly sunken above,
faint below, lines of interarching and venation indistinct; petiole 20-25 by 2-2.5 mm;
leaf bud 12—15 by 2—2.5 mm, hairs appressed, 0.1 mm. /nflorescences (from infructes-

540 Flora Malesiana, Ser. I, Vol. 14 (2000)

cences): between the leaves and below, of the Knema-type: sessile knob-like woody
scar-covered 1—3 topped brachyblasts 2-3(-5) mm long, minutely brown pubescent.
Male and female flowers not known. Fruits | or 2 (or 3) per infructescence; (broadly)
ovoid-ellipsoid, 22.3 by 1.5-1.7 cm, beaked for 1 mm by style-remnant, hairs fine,
mealy, rust-coloured, 0.1 mm; pericarp 1(—2) mm thick, seeds (not fully mature) ellip-
soid, 1.5 cm; fruiting pedicel slender, (13—)15—18 by 1.5—2 mm, bracteole scar 1(—1.5)
mm below the apex, perianth scar clearly visible.

Field-notes — Bole 25 m, with low buttresses, the whole base on short stilt-roots.
Bark short flaky, with slight acrid smell; blaze pink turning brown; sapwood cream, on
exposure quickly deepening in colour.

Distribution — Malesia: Papua New Guinea (Madang Prov.); known only from the
type.

Habitat & Ecology — On river terrace, in Pometia—Celtis forest; at c. 115 m altitude;
fr. Aug.

104. Myristica perlaevis W. J. de Wilde

Myristica perlaevis W.J. de Wilde, Blumea 42 (1997) 178. — Type: Burley, Tukirin & Ismail 4330,
Seram.

Tree 20 m. Twigs 3(—4) mm diameter, slightly angular, glabrous; older twigs pale
grey-brown, coarsely longitudinally cracking; lenticels not apparent. Leaves thinly mem-
branous, elliptic-oblong, 16-23 by 6—9 cm, base cuneate or nearly rounded, apex abruptly
acute-acuminate; upper surface dark olivaceous, glossy, lower surface bright or light
brown or olivaceous, glabrous; not papillose; dots absent; midrib flat above, lateral nerves
13-15 per side, at (S0—)60—70° to the midrib, faint, flat above, not contrasting in colour
below, lines of interarching and venation (fine) indistinct; petiole 20-25 by 1.5-2 mm,
brown blackish, rather contrasting in colour with the twig; leaf bud 10 by 1—-1.5 mm,
hairs powdery, 0.1 mm. Male inflorescences and flowers not seen. Female inflorescen-
ces glabrous, peduncle 6(-10) mm long, ending in one flower. Female flowers: gla-
brous, pedicel 12 mm, bracteole scar 3 mm below the apex; buds carnose, 9 by 7 mm,
cleft c. 1/2, lobes 4—4.5 mm long; ovary 7 by 4.5 mm, minutely pubescent. Infructescence
short pedunculate. Fruits solitary, ellipsoid-oblong, 8-8.5 by 3-3.5 cm, base and apex
+ blunt or narrowly rounded, glabrous; pericarp 8—10(—15) mm thick, finely wrinkled,
brown, glabrous; seeds elongate-ellipsoid, 4.5—5 cm; fruiting pedicel not seen.

Field-notes — Tree 20 m, dbh 20 cm. Bark grey, deeply grooved and longitudinally
fissured; inner bark orange-red; sap red, watery. Flowers waxy, yellow. Fruits greenish
yellow.

Distribution — Malesia: Moluccas (Seram, c. 40 km E of Masohi); known only from
the type.

Habitat & Ecology — Natural forest with Myristicaceae, Fagaceae, Myrtaceae and
Dipterocarpaceae; on steeply dissected ridge; reddish clay soil on grey shale and quartzite
bedrock; 275 m altitude; fl. & fr. Dec.

Note — This species is glabrous in all parts, except for a fine indumentum on leaf bud
and ovaries: some minute stellate scattered hairs may be seen at the base of the fruit
(lens!).

De Wilde — Myristicaceae (Myristica) 541

105. Myristica philippensis Lam.

Myristica philippensis Lam., Hist. Acad. Roy. Sc. Paris (for 1788) (1791) 161; Encycl. Méth. Bot.
4 (1797) 387; Warb., Mon. Myrist. (1897) 386, t. 12; Merr., Philipp. J. Sci., Suppl. 1 (1906) 55:
Sp. Blanc. (1918) 151; Enum. Philipp. Flow. Pl. 2 (1923) 179: J. Sinclair, Gard. Bull. Sing.
23 (1968) 138, f. 3, 4; W.J. de Wilde, Blumea 42 (1997) 179. — Type: Sonnerat s.n., Philip-
pines.

Myristica commersonii Blume, Rumphia | (1837) 181; A.DC., Prodr. 14, 1 (1856) 193; Miq., Fl.
Ind. Bat. 1, 2 (1858) 61. — Type: Commerson s.n., Mauritius, culta.

Myristica macrocarpa Blume, Rumphia | (1837) (= Nux moschata quarta ... in Ray, Hist. Pl. 3,
App. 58, sine descr.); A.DC., Prodr. 14, 1 (1856) 207; Fern.-Vill., Nov. App. (1880) 178; Vidal,
FI. For. Fil. Atlas (1883) (excl. pl. 57A 1, 2 & 3 = Knema glomerata) — Type: not preserved
(Sinclair, l.c.: 144).

Myristica luzonica Blanco, Fl. Filip. (1837) 664; ed. 2 (1845) 462, 463; ed. 3, 3 (1879) 69, 70:
A.DC., Prodr. 14, 1 (1856) 207; Migq., Fl. Ind. Bat. 1, 2 (1858) 72. — Type: no specimens pre-
served (Sinclair, l.c.: 144).

Myristica bracteata A.DC., Prodr. 14, 1 (1856) 192 (incl. var. longifolia ADE. Vice 195)2 Berm
Vill., Nov. App. (1880) 177; Vidal, Phan. Cuming. (1885) 139; Rev. Pl. Vasc. Filip. (1886) 221.
— Syntypes: Bojer s.n., 1833; Callery 33; Cuming 1481; Wallich Cat. n. 6800; DuPetit Thouars
s.n.; Sieber 365; etc. — Type of var. longifolia: Cuming 829 (see Sinclair, |.c.).

Myristica grandiflora Wall., Cat. (1832) n. 6800b, nomen (K, herb. sheet, not in the Wallich Cata-
logue).

Myristica madagascariensis auct. non Lam.: Bojer, Hort. Maurit. (1837) 275; Vent. ex A. DC.,
Prodr. 14, 1 (1856) 192 (in syn. Myristica bracteata).

Myristica sylvestris auct. non Houtt.: Sieber ex A.DC., Prodr. 14, 1 (1856) 192 (in syn. Myristica
bracteata).

Tree 6-15 m. Twigs 3-5(-7) mm diameter, early glabrescent, hairs 0.5—1 mm; older
twigs sometimes fissured; lenticels many, few, or absent. Leaves chartaceous or thinly
coriaceous, (obovate-oblong or) elliptic-oblong, or oblong-lanceolate, 15—45(—50) by
5—14(—15) cm, base cuneate or (broadly) rounded, apex acute (with + blunt tip) or acute-
acuminate; upper surface olivaceous-brown, lower surface grey-brown, glabrescent,
indumentum cobweb-like, hairs scale-like, 0.1 mm, mixed with few dendroid hairs 0.2
mm, in old leaves a faint cobweb-like covering remaining; distinctly or faintly papil-
lose; dots absent; midrib above flat, lateral nerves 18-30 per side, at 40—80° to the midrib,
flat above, brown and contrasting in colour below, lines of interarching and venation
distinct above, indistinct below; petiole 20-40 by 2-5 mm; leaf bud 6-10 by 3 mm,
hairs 1-2 mm long. Inflorescences pedunculate, between the leaves or below, or fre-
quently axillary to (fallen) reduced leaves at the twig apex, with shaggy rusty hairs 1-2
mm; bracts S—10(—20) mm, pubescent, caducous; in male: 4-9 by 3—4(—5) cm, peduncle
10-50 mm, branches to 20 mm, central axis (10—)20-50 mm long, with | or 2\(or3)
dense clusters of (3—-)5—10 flowers, buds of various sizes; in female: smaller, 2-3 cm
long, fewer flowered; flowers + coriaceous, with woolly hairs 0.2—0.5 mm, sometimes
subglabrescent. Male flowers: pedicel 5-8 mm, bracteole 4-6 by 6-8 mm, apex sub-
acute or + 3-topped, late caducous; buds ovoid, 5-8 by 5—7 mm, apex + blunt, base
broadly rounded, cleft 1/3—1/2, lobes 3 or 4, at sutures 0.5—1 mm thick. Androecium
cylindrical or + club-shaped, 4-6 mm, subtruncate; androphore broadly cylindrical,
(1-)1.5-2 by 1-1.2 mm, with hairs 0.2—0.4 mm long in the lower 2/3; synandrium 3-4
(-5) by 1.2-1.5 mm, thecae 12-16, sterile apex to 0.2—0.3 mm. Female flowers: pedicel

542 Flora Malesiana, Ser. I, Vol. 14 (2000)

1-2 mm long, bracteole caducous; buds ovoid, apex subacute, 7—8(—9) by 6-7 mm, cleft
nearly 1/2; ovary conical-ovoid, (3—)4 by 3-3.5 mm, hairs 0.3(—0.4) mm, stigma | mm.
Fruits usually solitary (seemingly more-fruited in aggregated inflorescences), broadly
ellipsoid, (4-)5-8 by 3—4.5 cm, hairs dense, rust-coloured, + mealy, 0.3—0.5 mm, some-
times + glabrescent; pericarp 7-10 mm thick; seeds not seen; fruiting pedicel 3-5 mm.

Field-notes — Bark blackish brown, longitudinally fissured. Seeds narrowly oblong,
conform to the fruits, dark brown and slightly glossy when dry. Sinclair (1.c.: 140) re-
ported a fresh fruit, cultivated in Madagascar, as large as 12 cm long.

Distribution — Cultivated in Mauritius, Madagascar, Martinique, and India (Calcutta);
Malesia: throughout the Philippines (except Palawan); no recent collections known.

Habitat & Ecology — Formerly a common understorey tree in primary lowland for-
est, up to c. 400 m altitude; fl. & fr. possibly throughout the year.

Note — Inflorescences of M. philippinensis appear frequently before the leaves, prob-
ably in relation to the seasonal climate in the Philippines, and are often axillary to re-
duced leaves and somewhat crowded at the apex of the twigs, with a vegetative bud at
the end, reminiscent of a compound apical inflorescence. The species is distinguishable
by its stout habit, paniculate inflorescences, and glabrescent leaves.

106. Myristica pilosella W. J. de Wilde

Myristica pilosella W. J. de Wilde, Blumea 40 (1995) 314. — Type: NGF 28986 (Streimann & Katik),
Papua New Guinea.

Treelet 5 m. Twigs 1-1.5 mm diameter, + angular, late glabrescent, hairs shaggy, 0.5
(-1) mm; older twigs yellowish brown, longitudinal cracks and lenticels not apparent.
Leaves thinly chartaceous, + oblong-lanceolate, 8-13 by 1.5—3.5 cm, base short-atten-
uate, apical part narrowed into long acute-acuminate apex; upper surface dark brown,
lower surface grey-brown, glabrescent, hairs scattered, less than 0.1 mm (lens!), subpersis-
tent and stouter on the midrib; not papillose; dots absent; midrib raised above (but in a
groove), lateral nerves 14-16 per side, at 60—70° to the midrib, sunken above, lines of
interarching and venation distinct on both surfaces; petiole 7-15 by 1-1.5 mm, glabrescent,
hairs pilose, to 1 mm; leaf bud 6-12 by 1.5—2 mm, hairs 1 mm. Inflorescences between
the leaves, with slender peduncle, paniculate, late glabrescent, hairs woolly, 0.5 mm
long or less; male inflorescences not known, but likely resembling those of M. cornutiflora
subsp. elegans and M. olivacea; female inflorescences: peduncle 15-25 by 0.5-1 mm,
ending in a scar-covered brachyblast up to 5 mm long, with one flower (bud), with or
without the initiation of a next bud; flowers with woolly light brown hairs 0.5 mm or
less. Female flowers: pedicel 6 by 0.7(-1) mm, bracteole ovate, 0.5—1 by 1 mm, sub-
persistent, apical; buds (submature) ovoid-oblong, narrowed in the apical half, 8 by 3
(3.5) mm, apex subacute, cleft c. 1/8, lobes 0.7(-1) mm long; ovary (from the perianth
and immature fruit) ovoid-oblong, 6—6.5 by 2.5 mm, hairs 0.5—1 mm. Fruits solitary, on
long slender stalk, presumably pendulous, fusiform, 5—5.5 by 1.5(-2) cm, apex + beaked,
base narrowed into a 5-10 mm long pseudostalk, hairs dense, yellow-rusty, pilose, 0.2—
0.4 mm and stouter straight hairs towards the apex and at the base of the pseudostalk;
pericarp 1 mm thick; seeds not seen but presumably (ellipsoid-)oblong, 3-3.5 cm; fruit-
ing pedicel 7-9 mm long, pubescent, with the (sub)persistent bracteole 1-2 mm below
the apex.

De Wilde — Myristicaceae (Myristica) 543

Field-notes — Small tree, bole 3 m, dbh 5 cm. Leaves dull dark green above, green
below: soft reddish scale-like hairs on midrib and new growth. Flower buds reddish
brown. Fruits reddish brown.

Distribution — Malesia: Papua New Guinea (Milne Bay Prov., Ruba Ruba subprov.,
at the junction of Ugat and Mayu Rivers, near Mayu I.); known only from the type.

Habitat & Ecology — In Castanopsis-dominated forest; 1000 m altitude; fl. & fr.
July.

Note — According to the inflorescences M. pilosella links up with M. cornutiflora
and M. olivacea. It is similar to M. cylindrocarpa, a lowland species also known only
from a single fruiting specimen which differs in the shorter indumentum, leaf venation,
shorter infructescences, and smaller fruits.

107. Myristica pilosigemma W. J. de Wilde

Myristica pilosigemma W.J. de Wilde, Blumea 42 (1997) 179. — Type: PNH 118078 (Gutierrez),
Philippines, Samar I.

Tree 12 m. Twigs 1.5—2(—2.5) mm diameter, early glabrescent, inconspicuously,
minutely pubescent; older twigs finely fissured or thinly flaky; lenticels small. Leaves
chartaceous or thinly coriaceous, elliptic-oblong or oblong-lanceolate, 10-24 by 3-6
cm, base + cuneate or (narrowly) rounded, apex acute(-acuminate), the very tip acute or
+ blunt; upper surface olivaceous, lower surface + grey, late glabrescent, hairs pale brown,
dense, felty, appressed, of mixed sizes, 0.1—0.4 mm, the larger hairs leaving minute dark
point-like scars; somewhat papillose; dots absent; midrib above slightly raised, lateral
nerves 14-17 per side, at 40-60° to the midrib, brown below, contrasting in colour, lines
of interarching and venation indistinct; petiole 12-18 by 2-2.5 mm; leaf bud 15-20 by
2-3.5 mm, hairs rough, bright brown, + appressed, | mm. /nflorescences minutely rusty
pubescent, glabrescent, between the leaves, as in Knema; in male: a simple subsessile
(peduncle | mm or less), worm-like scar-covered brachyblast to 13 mm long, with a
subumbel of 1—5 flowers, buds of various sizes; bracts small, caducous; female inflores-
cences not known: flowers with dense red-brown hairs 0.2 mm. Male flowers: pedicel
slender, 3 mm, bracteole ovoid-oblong, 3 mm, late caducous or subpersistent; buds el-
lipsoid-oblong, 4—4.5 by 2(—2.5) mm, cleft c. 1/2, lobes 1.5—-2 mm long, 0.2 mm thick.
Androecium 3 mm; androphore 1.5 by 0.4 mm, with hairs 0.2—0.3 mm in lower 1/4 only;
synandrium 1.5 by 0.5 mm, thecae 14, sterile apex shallowly lobulate at apex, (0.2—)0.3
mm. Female flowers not known. Infructescences sessile, wart-like, 5 mm long. Fruits
solitary, subsessile, ellipsoid, (3—)3.5 by 2.5-3 cm, hairs woolly, 0.5—1 mm; pericarp
4-5 mm thick; seeds ellipsoid, (2.5—)3 cm; fruiting pedicel short, 2—3 mm long.

Field-notes — Tree dbh c. 15 cm. Flowers yellow-brown.

Distribution — Malesia: Philippines (Samar, Mindanao).

Habitat & Ecology — Along trail; c. 600 m altitude; fl. Apr.; fr. June—July.

Notes — 1. Species of similar general habit in the Philippines are e.g. M. agusanensis
and M. cumingii (with + pedunculate inflorescences), but the present species differs in
the more conspicuous indumentum (hairs to 1 mm long) of leaf bud and fruits.

2. Myristica pilosigemma has the whitish lower leaf surface with minute punctation
caused by scars of fallen larger hairs in common with several species outside the Philip-
pines, e.g. with M. impressa from Sulawesi.

544 Flora Malesiana, Ser. I, Vol. 14 (2000)

108. Myristica polyantha W. J. de Wilde

Myristica polyantha W.J. de Wilde, Blumea 40 (1995) 315. — Type: Brass 24772, Papuan Islands
(Goodenough I.).

Tree 25-30 m. Twigs (1.5—)2—3 mm diameter, early glabrescent, hairs less than 0.1
mm; older twigs + longitudinally fissured, not flaking; lenticels scarce. Leaves coriaceous,
ellipsoid-oblong or obovoid-oblong, 7—17 by 3—9 cm, base (short-)attenuate, apex acute-
acuminate, the very tip rather blunt; upper surface brown, lower surface brown-grey,
glabrescent, hairs widely spaced, scale-like, less than 0.1 mm; not papillose; dots ab-
sent; midrib slightly raised above, lateral nerves 12—15 per side, at 45—60° to the midrib,
flat or sunken above, lines of interarching indistinct, venation distinct or not; petioles
12-15 by 2-3.5 mm; leaf bud 10-15 by 2-3 mm, hairs 0.1 mm or less. /nflorescences of
the Knema-type: sessile wart-like simple or somewhat forked woody scar-covered bra-
chyblasts to 5 mm diameter, rusty short-pubescent; in male: flowers in dense clusters of
10-20, buds subequal in size; female inflorescences not seen; flowers with hairs 0.1—0.2
mm. Male flowers: pedicel 4—6 by (0.6—)0.8 mm; bracteole sometimes + 3-topped, 2.5—
3 mm long, caducous, (sub)apical; buds (ellipsoid-)oblong, 4-6 by 2—2.5(—3) mm, base
+ rounded or tapering, cleft c. 1/4, lobes (1-)1.5 mm long, (0.1—)0.2 mm thick. An-
droecium broad-cylindrical, (3.5—)4—4.5 mm; androphore 2 by 0.8—1.2 mm, glabrous or
with scattered pale hairs less than 0.1 mm only; synandrium oblong, + truncate, 1.8—2.5
by 1-1.5 mm, thecae 10-12, not tightly contiguous, sterile apex + absent. Female flow-
ers not seen. Infructescences sessile, minute. Fruits (immature) | or 2 per infructescence,
globose, 2.5(—3) cm diameter, hairs (grey-)rusty, mealy, 0.1 mm; pericarp 7 mm thick;
seeds not seen; fruiting pedicel 5-8 by 3.5-5 mm, with coarse lenticels, bracteole scar
apical.

Field-notes — Diameter of trunk to 35 cm. Leaves grey below.

Distribution — Malesia: Papua New Guinea (Milne Bay Prov., Goodenough I. and
possibly Tagula I., but see note 2).

Habitat & Ecology — Canopy or subcanopy tree of montane oak forest and foothill
forest, heavily mossed forest of ravine; c. 1600 m altitude (but see note 2); fl. & fr. Aug.,
Oct.

Notes — 1. Related to the variable M. globosa and to M. laevifolia. The former is
mainly a lowland species, with thinner leaves and flowers with thinner perianth, nar-
rower androecium, smaller bracteole, and fruits with thinner pericarp; M. /aevifolia has
very similar male flowers, with extremely thin indumentum, but the fruits differ in their
elongate shape.

2. Brass 27918 (male, Tagula I., 300 m altitude) differs in its non-contrasting veins.

3. Myristica polyantha may be related to M. tenuivenia subsp. lignosa, with similar
globose, thick-walled fruits, but the latter differs in the (+) persistent indumentum of the
lower leaf surface and a more conspicuous fruit coat, with hairs (0.2—)0.5 mm. The
taxonomic status of M. polyantha needs confirmation by more field study.

109. Myristica psilocarpa W. J. de Wilde

Myristica psilocarpa W.J. de Wilde, Blumea 40 (1995) 317. — Type: LAE 52394 (Foreman), Papua
New Guinea, Manus I.

De Wilde — Myristicaceae (Myristica) 545

Tree 18—22 m. Twigs 3 mm diameter, early glabrescent, hairs less than 0.1 mm; older
twigs not seen. Leaves thinly chartaceous, oblong-lanceolate, 14—24 by 4—7 cm, base
subcuneate or narrowly rounded, apex blunt or subacute; upper surface olivaceous or
brown, lower surface grey- or pale greenish brown, glabrescent, hairs widely scattered,
scale-like, less than 0.1 mm; not papillose; dots absent; midrib + flat above, lateral nerves
15-20 per side, at 45—70° to the midrib, flat or sunken, little raised below, reddish brown,
contrasting, lines of interarching indistinct, venation sunken and faint above, reddish
brown, contrasting below; petiole 15—25 by 2—2.5 mm; leaf bud 15 by 2.5—3 mm, hairs
0.1(—0.3) mm. Inflorescences pedunculate, between the leaves, (early) glabrescent, hairs
scattered, pale, 0.1 mm, like on the flowers; bracts 2 mm, caducous; in male: peduncle
(15—)20 mm long, ending in one or two (sub)sessile scar-covered brachyblasts to 5 mm
long, each with a subumbel of (2—)3—5 flowers, buds of various sizes; in female (from
infructescences): 5-12 mm pedunculate, 1-flowered. Male flowers: pedicel 6—7 by 1.5—
2 mm, bracteole 3—3.5 mm, caducous, scar conspicuous; buds leathery, ovoid-ellipsoid,
7-7.5 by 6 mm, cleft c. 1/2, lobes 3.5 mm long, 0.5 mm thick. Androecium 5.5 mm;
androphore (1—)1.5 by 0.4 mm, glabrous; synandrium 4 by 1.2—1.4 mm, thecae 16-18,
sterile apex + cushion-shaped, impressed in the centre, 0.3(—0.4) mm high. Female flow-
ers not known. Fruits single, obovoid-oblong, 5—5.5 by 3 cm, apex + blunt or subacute,
basal part narrowed, early glabrescent, hairs + scattered, appressed, rather pale, less than
0.1 mm; pericarp hard, 5(—7) mm thick; seeds ellipsoid-oblong, 2.5(—3) cm, with hard,
bony, yellowish brown testa; peduncle of infructescence 5—12 by 3-4 mm, fissured, with
scattered small lenticels, passing into fruiting pedicel, 5—7 by 3 mm, striate.

Field-notes — Small buttresses, or with stilt-roots at base. Bark dark brown, scaly;
sap reddish; wood white or brown flecked, with red sap. Leaves shiny, dark green above,
whitish below. Flowers white. Fruits (mature) green, + obovoid; seeds brown.

Distribution — Malesia: Papua New Guinea (endemic to Manus I.).

Habitat & Ecology — Lowland rain forest, at 150-180 m altitude; fl. June; fr. July.

Note — Myristica psilocarpa is similar and related to M. hooglandii, but the latter is
distinguishable by a stouter habit, leaves with faint or invisible venation, much larger
male flowers (synandrium without sterile apex) and fruits with less woody and less hard
pericarp, at first with a thin mealy indumentum, glabrescent.

110. Myristica pubicarpa W. J. de Wilde

Myristica pubicarpa W.J. de Wilde, Blumea 35 (1990) 251, f. 2 (10) & 3a-e; 42 (1997) 180. —
Type: Pleyte 310, Halmahera.

Tree 10-25 m. Twigs 1.5—2(—3) mm diameter, early glabrescent, hairs 0.5—1 mm long;
older twigs distinctly lenticellate. Leaves membranous or chartaceous, (elliptic-)oblong,
15-26 by 5—11.5 cm, base short-attenuate or narrowly rounded, apex acute-acuminate;
upper surface brown, lower surface grey-brown, seemingly glabrous but late glabrescent.
hairs thin, widely spaced, scale-like, 0.1 mm, mixed with few pale brown larger ones,
0.2 mm, the larger hairs leaving point-like scars; not obviously papillose; dots absent;
midrib slightly raised above, lateral nerves 12-16 per side, at 45(—60)° to the midrib, flat
or sunken above, on lower surface bright brown, contrasting in colour, lines of inter-
arching distinct, venation faint or hardly visible; petioles 10-22 by 1.5—4 mm; leaf bud

546 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Myristica) 547

10—15(—20) by 2-3 mm, hairs silky, yellow-brown, (0.5—)1 mm long. Inflorescences be-
tween the leaves and below, of the Knema-type: sessile, simple or forked + woody scar-
covered brachyblasts, with rusty hairs 1 mm; bracts caducous; in male: brachyblasts up
to 5 mm, at apex with up to 5 flowers, buds not much variable in size; in female: brachy-
blasts smaller, 2 mm, 1—3-flowered; flowers + glabrescent except at apex, hairs (0.2—)
0.3 mm. Male flowers: pedicel 8-10 mm, bracteole ovoid, subacute, 2—3 mm, caducous;
buds subcylindrical, somewhat angled, (10—)13—15 by 2.5—3(—3.5) mm, apex (sub)acute,
base narrowly rounded, cleft 1/15—1/20, the lobes 0.5 mm long, 0.3—0.4 mm thick.
Androecium elongate, 9-12 mm; androphore slender, (3—)3.5—4 mm, with dense hairs
0.5 mm; synandrium 6.5—7.5 by 0.4-0.5 mm, thecae 14-16, + contiguous, sterile apex
+ acute, (0.5—)1 mm, glabrous. Female flowers: pedicel 4-6 by 1.5(—2) mm, bracteole
1.5-2 mm, caducous, scar about median; buds ovoid-oblong, 8-10 by 4-5 mm, nar-
rowed towards subacute apex, apical part + angled, base + rounded, cleft 1/8, lobes 0.5
mm thick; ovary ovoid-oblong, 4 by 2—2.3 mm, with dense hairs 0.5—1 mm, style and
stigma slender, 1.5-2 mm long, deeply 2-lobed. Fruits solitary, sometimes several to-
gether (on short-shoots with several inflorescences), subglobose or broadly ellipsoid,
5-6 by 3.5—4.5 cm, apex + rounded or shortly beaked, base narrowly rounded and nar-
rowed into a pseudostalk up to 5 mm, indumentum coarse-mealy, hairs dense, dark rust-
coloured, 0.2—0.3 mm; pericarp hard-leathery or woody, often wrinkled or bullate on
drying, 5-10 mm thick; seeds broadly or narrowly ovoid-ellipsoid, 2.5—3 cm, aril thick,
fatty, deeply sunken into the seed leaving deep grooves when removed; fruiting pedicel
stout, 10 mm long. — Fig. 65a, 82a-e.

Field-notes — Bole very straight, up to 25 cm dbh, buttresses to 40 cm high and out,
2 cm thick, or with low stilt-roots, c. 50 cm high, covering an area of about one square
metre. Outer bark 0.2—0.3 mm thick, (grey-)brown, not fissured, slightly peeling off or
not; inner bark 2.5—5 mm thick, pale red on section; exudate little, watery, (orange-)
brown; sapwood cream turning reddish (cream). Flowers yellow. Fruits with dark brown
indumentum.

Distribution — Malesia: N Moluccas (endemic to Halmahera and Obi).

Habitat & Ecology — Solitary, locally common tree. Primary open or rather dense
foothill forest, with little or dense undergrowth; clayey soil; rather level terrain or hill-
side; along small stream; 500—600 m altitude; fl. & fr. Nov.

Note — This species, readily distinguishable by its large, elongate, male flowers and
sessile inflorescences, is related to M. succedanea from nearby Ternate, Tidore and Bacan.
The latter differs in its dense (but low) subpersistent indumentum on the lower leaf sur-
face, usually distinctly pedunculate inflorescences, persistent or late-falling apical brac-
teole, shorter indumentum of twig apex, leaf bud and flowers, larger glabrescent fruits,
and a thinner aril.

Fig. 82. Myristica pubicarpa W.J. de Wilde. a. Leafy twig with female inflorescence; b. twig with
male inflorescences; c. twig with female inflorescences; d. female bud, opened, cleft at anthesis to
the depth as indicated at the apex; e. twig with two infructescences, each with one fruit, one opened,
showing subglobose seed with thick aril. — M. succedanea Blume. f. Fruit, opened, showing el-
lipsoid seed with thin dried aril [a: de Vogel 3979; b: de Vogel 4000; c-f: de Vogel 4092]. — Scale
bar for a—c, e, f = 2 cm; for d = 2 mm.

548 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Myristica) 549

111. Myristica pumila W. J. de Wilde

Myristica pumila W. J. de Wilde, Blumea 40 (1995) 319, f. 6a—c. — Type: Hoogland 3935, East
Papua New Guinea.

Shrub or treelet, 1.5—7 m tall. Twigs (faintly) angular, (0.5—)1—1.5 mm diameter, early
glabrescent, hairs less than 0.1 mm; older twigs neither cracking nor flaking; lenticels
inconspicuous. Leaves membranous, (elliptic-)oblong, 6—15.5 by 2—5 cm, base attenu-
ate, apex acute-acuminate; upper surface olivaceous-brown, lower surface + pale grey-
ish brown (rather concolorous with upper surface), early glabrescent, hairs less than 0.1
mm; not papillose; dots absent; midrib raised above, lateral nerves 9-16 per side, at 60—
80° to the midrib, sunken (and indistinct) above, brown and contrasting in colour below,
lines of interarching distinct, venation distinct or not; petiole 7-15 by 1—1.5 mm; leaf
bud 5 by 0.5—1 mm, hairs less than 0.1 mm. /nflorescences between the leaves, of the
Knema-type: a sessile or up to | mm pedunculate, scar-covered brachyblast; in male:
simple or forked, up to 5 mm long, ending in a cluster of 1—4 flowers, buds of various
sizes; female with a single flower, sessile;. flowers thinly or scarcely with hairs less than
0.1 mm, looking as if glabrous. Male flowers: pedicel 6.5—10 by 0.4(—0.5) mm, bracteole
+ ovate, caducous, scar 2-4 mm below the apex; buds elongate, tubiform, (6—)7—9.5 by
1.8-2 mm, slightly narrowed or not in the lower half, apex + blunt, base + rounded-
attenuate, inner surface in basal part minutely verrucose; cleft c. 1/6, lobes membranous,
1-1.7 mm long, 0.2 mm thick. Androecium slender, 6-9 mm; androphore 3—4.5 by
(0.3—-)0.4—0.5 mm, with hairs 0.1 mm or less in the lower 1/2—2/3; synandrium 2.4—4.2
by 0.6—0.7 mm, thecae c. 12, subacute, sterile apex 0.3(—0.4) mm. Female flowers (sub-
mature) solitary, pedicel thread-like, at least 14 mm long, bracteole scar about median;
buds (narrowly) ellipsoid-oblong, at least 5 by 1.5—2 mm, lobes | mm; ovary not seen.
Fruits solitary, oblong, 5 by 1.5 cm, apex + blunt, base gradually contracted into a nar-
row pseudostalk for nearly half of the total length of the fruit, hairs sparse, greyish
brown, appressed scale-like, less than 0.1 mm; pericarp 1 mm thick; seeds ellipsoid, c. 2
cm long; fruiting pedicel filiform, 20-25 by 0.5 mm, bracteole scar below the middle. —
Fig. 62e, 83a-c.

Field-notes — Stem to 10 cm diameter. Bark (grey-)brown, faintly longitudinally fis-
sured; inner bark pale brown; exudate colourless turning red; wood straw(-brown). Leaves
slightly glaucous below. Flowers pale yellow or greenish. Fruits pale rusty brown; seeds
dark brown.

Distribution — Malesia: Papua New Guinea (Northern Prov., Milne Bay Prov.:
Fergusson I.).

Habitat & Ecology — Primary and degraded forest; rather open tall forest, lowland
forest by river, fairly dense c. 10 m tall forest; 170-400 m altitude; fl. Mar., Sept., Nov.;
fr. Sept.

Fig. 83. Three related small-leaved Myristica species with the perianth finely warty inside. —
M. pumila W.J. de Wilde. a. Male flowering twig; b. male bud, opened, bracteole caducous; c. apex
of leafy twig with one-fruited infructescence. — M. crassipes Warb. subsp. crassipes. d. Male bud,
opened, with caducous bracteole. — M. flosculosa J. Sinclair. e. Male flower, opened [a, b: Hoog-
land 3935; c: Hoogland 3971, d: Carr 16091; e: Saunders 175\. = Scale-bar for a; .¢ = 2'ems for
bydie= 15 mm.

550 Flora Malesiana, Ser. I, Vol. 14 (2000)

Notes — 1. Obviously close to M. tubiflora, and especially to M. filipes and M. flos-
culosa: with the latter two it shares the warty-fleshy inner surface of the lower half of the
perianth. Myristica pumila can be regarded as a delicate edition of M. flosculosa. An-
other much resembling species is M. sogeriensis.

2. Myristica pumila keys out beside species with a sessile, non-pedunculate inflores-
cence: however, some of the inflorescences have a short peduncle, to 1 mm long, usually
in specimens in which the majority of the inflorescences is completely sessile.

112. Myristica pygmaea W. J. de Wilde

Myristica pygmaea W.J. de Wilde, Blumea 40 (1995) 320. — Type: LAE 66758 (Barker & Katik),
Papua New Guinea.

Treelet 2-4 m. Twigs subangular, 1.5(—2) mm diameter, early or rather late glabrescent,
hairs 0.3—0.5 mm; older twigs striate; lenticels small and inconspicuous. Leaves mem-
branous or thinly chartaceous, oblong, 8—15(-17.5) by 3-5(—6.5) cm, base short-attenu-
ate or rounded, apex acute-acuminate, often with + blunt tip; upper surface olivaceous-
brown, lower surface grey-brown or cinnamon, late glabrescent, hairs weak, flossy, 0.1
(0.2) mm; indistinctly papillose; dots absent; midrib slightly raised above, lateral nerves
(4-)7-14 per side, at 45—60° to the midrib, the lower nerves sometimes steeper, (flat or)
sunken above, (light) brown and contrasting in colour below, lines of interarching and
venation indistinct; petioles 10-12 by 1.5—2(-2.5) mm; leaf bud 10 by 2.5 mm, hairs
+ rough, 0.3—0.5 mm. /nflorescences (from infructescences) between the (lower) leaves,
axillary or up to 3 mm supra-axillary, + paniculate, peduncle 3-4 mm long, ending ina
small, scar-covered brachyblast with few flowers, partly late glabrescent, hairs dark brown
(rusty), 0.3—0.5 mm; male inflorescences, male and female flowers not seen. Infructes-
cences: peduncle 3—4 by 2 mm, + pubescent. Fruits (slightly immature) solitary, + ellip-
soid, 2.5—2.7 by 1.5 cm, apex subacute, narrowed towards the base, hairs powdery, rust-
coloured, (.3-0.4 mm; pericarp + 2 mm thick; seeds not seen; fruiting pedicel 2-4 by
2-2.5 mm, hairs woolly, 0.2 mm, bracteole scar near the apex.

Field-notes — Outer bark grey; inner bark reddish brown; wood whitish brown. Leaves
dull dark green or + shiny light green above, hairy below. Fruits (orange-)yellow, with
brown indumentum.

Distribution — Malesia: Papua New Guinea (Morobe Prov., Mumeng & Lae sub-
prov.); locally endemic to that area.

Habitat & Ecology — Lowland rain forest, logged forest; c. 120 m altitude; fr. Nov.

Note — Distinguishable by its small stature, 2-4 m. Superficially resembling other
small-statured species of Papua New Guinea and possibly most related to M. cylindrocarpa
or to the montane M. sinclairii, all with pedunculate inflorescences, partly supra-axil-
lary to the leaves, but M. pygmaea is different in indumentum and fruits. The species is
known only with fruits; female flowers apparently have a comparatively short pedicel.

113. Myristica quercicarpa (J. Sinclair) W. J. de Wilde
Myristica quercicarpa (J. Sinclair) W. J. de Wilde, Blumea 40 (1995) 321. — Myristica fatua Houtt.

"var. quercicarpa J. Sinclair, Gard. Bull. Sing. 23 (1968) 302, f. 38D-F; Foreman, Contr. Herb.
Austral. no. 9 (1974) 37 (key); Handb. Fl. Papua New Guinea | (1978) 196. — Type: NGF 1270

(L.S. Smith), Papua New Guinea.

De Wilde — Myristicaceae (Myristica) 551

Tree 20—40 m. Twigs 2-4 mm diameter, glabrescent, hairs 0.5(—1) mm long; older
twigs somewhat longitudinally cracking, probably later on + flaking; lenticels absent.
Leaves thinly chartaceous, elliptic- or obovate-oblong, 15—21 by 4.5-10 cm, base cuneate,
rounded, or subcordate, apex acute-acuminate; upper surface olivaceous(-brown), lower
surface yellowish brown, hairs persistent, dense, pale, floccose, interwoven, 0.1—-0.2 mm
with numerous emergent brown or yellowish hairs of varying length, up to 1 mm, not
papillose; dots absent; midrib slightly raised above, with some indumentum remaining
in younger leaves, lateral nerves 13—23 per side, at 60—80° to the midrib, lines of inter-
arching distinct, venation distinct or faint; petiole late glabrescent, 12-20 by 2.5-3.5
mm; leaf bud 10-15 by 2-3 mm, hairs 0.5—1 mm long. Inflorescences (from infructes-
cences) of the Knema-type: in female a sessile, wart-like brachyblast, few-flowered,
between the leaves (or below), or rarely a few in a lateral short-shoot. Male and female
flowers not known. Infructescences very short. Fruits: subsessile, single or 2—4 ina
cluster, acorn-shaped, about as broad as long or broader than long, + broadly ovoid, 1.3-
2.3 by 1.8—2.7 cm, apex (broadly) rounded with short pointed tip, base broadly truncate,
flat, or depressed, hairs dark (yellow-)brown, floccose, 0.5—1 mm long; pericarp 2(—3)
mm thick; seeds broadly ellipsoid, broadly rounded at both ends, 1.5 cm; fruiting pedi-
cel slender, 5—10 by 2-3 mm, pubescent, bracteole scar apical. — Fig. 61e.

Field-notes — Bole straight, sometimes channelled at base, not buttressed. Branches
horizontal. Bark blackish, with fine longitudinal fissures or wrinkled and with less dis-
tinct horizontal grooves, or peeling off in rectangular flakes; inner bark red-brown, con-
centrically layered; wood pale, red-brown, staining on exposure; exudate red, watery.
Leaves brown-green below, or with brown hairs. Fruits (pale) reddish brown.

Distribution — Malesia: Papua New Guinea (endemic to Northern and Milne Bay
Prov.).

Habitat & Ecology — Lowland forest on flats; 50-100 m altitude; fr. Mar., June.

Note — Known only from fruiting specimens. The indumentum of the lower leaf sur-
face with long, emergent hairs, rather than only a thin felt-like covering, makes the spe-
cies quite distinguishable from the M. fatua-species group, even in the sterile state.

114. Myristica robusta W. J. de Wilde

Myristica robusta W. J. de Wilde, Blumea 35 (1990) 253, f. 2: 11; 42 (1997) 180. — Type: de Vogel
3918, Moluccas (Bacan).

Tree c. 15 m. Twigs 5-10 mm diameter, + hollow, early glabrescent, hairs scurfy, less
than 0.1 mm; older twigs soon finely longitudinally cracking, thinly scaly; lenticels grey-
brown, abundant, but not conspicuous. Leaves thinly coriaceous, elliptic-oblong, 30—40
by 14-20 cm, base rounded or somewhat cordate, apex subacute; upper surface olivaceous-
brown, lower surface pale brown, + glabrescent, hairs scurfy, less than 0.1 mm, not pap-
illose; dots absent; midrib slightly raised above, lateral nerves 18-23 per side, at 60—80°
to the midrib, flat or sunken, lines of interarching distinct, venation + trabeculate, faint
at both surfaces; petiole 26-40 by 4-6 mm (in vigorous sapling shoots + alate); leaf bud
25 mm long, hairs 0.1(—0.2) mm or less. Inflorescences between the leaves and below, of
the Knema-type, in male: peduncle up to 6 mm long, with a simple or a 2—4-fid woody,
scar-covered brachyblast (5—)10-20 by 4-7 mm, with + woolly hairs 0.2—0.3 mm, end-
ing with clusters of 3-6 (or more) flowers, buds of various sizes; bracts minute, caducous;

552) Flora Malesiana, Ser. I, Vol. 14 (2000)

flowers with woolly hairs 0.2-0.3 mm. Male flowers: pedicel (10—)14 by 1.5(—2) mm,
bracteole (sub)persistent, 4.5—5 mm long; buds ellipsoid(-oblong), 8-8.5 by 4—4.5 mm,
apex narrowly rounded, cleft c. 1/5, lobes (0.4-)0.5 mm thick. Androecium 7 mm;
androphore 2.5 by (1—)1.5 mm, with a few scattered pale hairs 0.1 mm; synandrium 4.5
by 1.5(-2) mm, thecae c. 20, sterile apex 0.1 mm, rather flat with lobed ege. Female
flowers and fruits not seen. — Fig. 65b.

Field-notes — Solitary tree, clear bole 8 m, dbh 13 cm, straight with solid core. Outer
bark 0.4 mm thick, fissured, not peeling off; inner bark 2.5 mm thick, on section pale
red-brown; exudate little, watery, pale reddish brown; sapwood light yellowish, gradu-
ally passing into the slightly red-tinged heartwood. Flowers yellowish, brown by indu-
mentum.

Distribution — Malesia: N Moluccas (Bacan); known only from the type.

Habitat & Ecology — Disturbed, tall, 50 m high forest with little undergrowth; allu-
vial flat, with in places stagnant water at c. 5 m altitude; fl. Nov.

Note — Myristica robusta is distinguishable by its stout twigs and leaves, stout inflo-
rescences of the Knema-type, and the comparatively large male flowers, 8 mm long. The
species seems closely related to the widespread M. fatua which readily differs in leaves
that are pubescent below, and smaller male flowers. Also close to M. frugifera from the
Philippines, a species known only in fruit, noticeable for its paniculate, pedunculate in-
florescences.

115. Myristica rosselensis J. Sinclair

Myristica rosselensis J. Sinclair, Gard. Bull. Sing. 23 (1968) 205, f. 16; Foreman, Handb. Fl. Papua
New Guinea 1 (1978) 207; W. J. de Wilde, Blumea 40 (1995) 32. — Type: Brass 27447, Papua
New Guinea, Misima I.

Tree 7-25 m. Twigs faintly angular, 1.5—2.5 mm diameter, early glabrescent, indu-
mentum minute; older twigs finely longtidudinally cracking; lenticels not obvious. Leaves
membranous or (thinly) chartaceous, oblong or lanceolate, 6-16 by 2-3.5(—4) cm, base
narrowly rounded or acute, apex subacute or blunt; upper surface olivaceous brown,
lower surface grey-brown, seemingly glabrous, but hairs (sub)persistent, scattered, less
than 0.1 mm; papillose (lens!); dots absent; midrib above flat or slightly raised, on lower
surface yellowish or reddish brown, contrasting in colour, lateral nerves 15—22 per side,
at 60—70° to the midrib, above and below flat or sunken and faint, lines of interarching
and venation faint or invisible; petiole 10-15 by 1.5—2 mm; leaf bud 5-10 by 1-1.5 mm,
hairs 0.1 mm. Inflorescences between the leaves or below, pedunculate, early glabrescent,
the lateral branches with sessile, slender, scar-covered brachyblasts, central axis short
or reduced to one flower; bracts + acute, caducous, 1—1.5 mm long; male inflorescence:
4-7 cm long, peduncle flattened, 25-40 by 2—2.5 mm, lateral brachyblasts to 20 mm
(subsp. minutiflora), flowers in subumbels of 2—4(-5), buds subequal in size; female
like male, shorter (20-30 mm), 2—4-flowered, the central axis short or absent, each sub-
umbel reduced to a single flower; flowers membranous, hairs 0.1 mm (densest at apex
and base). Male flowers variable in size; pedicel slender, bracteole apical; buds cleft
1/5—1/4: androecium slender, the apical half fertile with 8-14 thecae, sterile apex absent
or to 0.2 mm. Female flowers: buds urceolate, 4.5 mm long; ovary pubescent (see subsp.
minutiflora). Fruits not seen.

De Wilde — Myristicaceae (Myristica) 355

Distribution — Malesia: Papua New Guinea (Louisiade Archipelago). With two sub-
species.

Note — Closely allied to M. schleinitzii, the latter with a widespread distributional
range; M. rosselensis differs in the more narrow leaves with acute base, and faint vena-
tion, and inflorescences with sessile lateral slender brachyblasts. Sinclair (1968) men-
tioned the glabrous androphore for M. rosselensis, but in M. schleinitzii the androphore
may be + glabrous as well. The leaves of M. rosselensis resemble those of M. clemensii.

KEY TO THE SUBSPECIES

la. Male buds 5.5 mm long, pedicel 5 mm long, about as long as the bud............
Paes! Gaeikycs cause dew oda wdde meee $2) sere a. subsp. rosselensis

b. Male buds 3.5 mm long, pedicel | mm long, much shorter than the bud..........
IE A Foes sys ad coys'd rath anyougs odd scien gaaenide Ss SA: b. subsp. minutiflora

a. subsp. rosselensis

Tree c. 25 m tall. Leaves 6-16 cm long. Male inflorescences with distinct central axis
with flower clusters on 2 (or 3?) levels, the clusters subsessile (short, but possibly later
on growing out into scar-covered brachyblasts). Male flowers: pedicel 5 mm long; buds
ovoid-oblong, 5—5.5 by 2.5 mm, cleft c. 1/5, lobes 1 mm long. Androecium 5 mm; andro-
phore about as wide as the synandrium, 2.5 mm long, glabrous; synandrium 2.5 by 1.2
mm, thecae 8—10, with broad connectives, sterile apex almost absent. Female flowers
and fruits not seen.

Field-notes — Leaves glaucous below. Flowers yellow, fragrant.

Distribution — Malesia: Papua New Guinea (Louisiade Archipelago: Misima I., Mt
Sisa); known only from the type.

Habitat & Ecology — In forest associated with oaks, on north slopes at c. 300 m
altitude; fl. Sept.

Note — The male flowers are almost identical with those of M. schleinitzii, including
the androecium with 8-10 thecae (with broad connectives) as against 12-16 in M.
schleinitzii.

b. subsp. minutiflora W. J. de Wilde

Myristica rosselensis J. Sinclair subsp. minutiflora W. J. de Wilde, Blumea 43 (1998) 254. — Type:
Brass 28245, male fl., Rossel I.

Tree 7-25 m tall. Leaves 6-10 cm long. Male inflorescences with two opposite ses-
sile scar-covered slender brachyblasts 10-20 mm long, the central axis reduced to one
flower or absent. Male flowers: pedicel 1 mm long; buds (ellipsoid-)oblong, 3.5 by 1.5
mm, cleft c. 1/4, lobes 0.8 mm long. Androecium 2.8—3 mm; androphore about as wide
as the synandrium, 1.5 mm long, glabrous; synandrium 1.5 by 0.7 mm, thecae 10-12,
sterile apex blunt, 0.2 mm. Female flowers: pedicel 1-2 mm long, bracteole scar subapi-
cal; buds ovoid-oblong, 4.5 by 2.5-3 mm, somewhat narrowed to the apex, cleft c. 1/5,
lobes 1 mm long; ovary ovoid-oblong, 3.5 by 2 mm, narrowed into 0.4 mm long stigma,
hairs dense, 0.1 mm. Fruits not seen.

554 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Substage tree or lesser canopy tree. Leaves grey below. Flowers yel-
low.

Distribution — Malesia: Papua New Guinea (Louisiade Archipelago: Rossel and
Tagula I.); known from two collections.

Habitat & Ecology — Rain forest of ridge crests at 100—300 m altitude; fl. Sept.

116. Myristica rubrinervis W. J. de Wilde

Myristica rubrinervis W.J. de Wilde, Blumea 42 (1997) 180. — Type: Ridsdale SMHI 258, Philip-
pines, Palawan.

Tree 8-30 m. Twigs somewhat angular or not, 2-3 mm diameter, early glabrescent,
hairs grey or orange-brown, 0.1 mm; older twigs with inconspicuous lenticels. Leaves
chartaceous or (sub)coriaceous, elliptic-oblong or (obong-)lanceolate, 8-16 by 1.8-4.5
(6) cm, base cuneate, apex acute-acuminate, the very tip (subacute or) + blunt; upper
surface olivaceous or brown, lower suface greyish, subpersistent or glabrescent, hairs
dense, woolly, 0.1—0.2(—0.3) mm; papillose (lens!); dots absent; midrib above some-
what raised, purplish brown, lateral nerves 1 1—15 per side, at c. 45° to the midrib, above
flat or sunken, thin, below distinct, conspicuously purplish brown (or yellow when fresh),
contrasting in colour, lines of interarching and venation distinct; petiole 12—30(—40) by
1.5-2.5 mm: leaf bud 8-15 by 2—4 mm, hairs 0.1-0.2 mm. Inflorescences pedunculate,
between the leaves, with rufous hairs 0.1—0.3 mm; in male: 1.5—2.5 cm long, peduncle
10 mm, flattened, proceeding into a branch 5—10(—20) mm, with 1—4 (sub)sessile or
short-stalked lateral clusters each of 5(—10) flowers, buds of various sizes; bracts ovate,
3 mm long, caducous; female inflorescences (from infructescences): glabrescent, (1—)
1.5-3 cm long; flowers densely pubescent. Male flowers (Soejarto c.s. 8079): pedicel
2-4 mm, slender, bracteole small, caducous; buds oblong, 3—4.5 by 2—2.5 mm, cleft
c. 1/3, lobes 1-1.5 mm long, c. 0.3 mm thick. Androecium 3.5 mm; androphore 2 mm,
for the larger part minutely pubescent; synandrium 1.5 mm, thecae c. 12, sterile apex
(almost absent) consisting of 5 or 6 protrusions of the anthers, 0.1 mm long or less.
Female flowers not seen. Fruits 1 or 2 (or 3) per infructescence, ovoid-ellipsoid to ellip-
soid(-oblong), 3-7 by 2.5—3.5 cm, apex and base (narrowly) rounded, hairs scurfy,
orange-brown or rusty, 0.1—0.2 mm; pericarp 2-6 mm thick; seeds ellipsoid, 2—4.5 cm;
fruiting pedicel 2-4 mm.

Distribution — Malesia: Philippines (Palawan, Mindoro, Leyte). There are two vari-
eties.

Note — Known mostly from fruiting specimens, all quite distinct from the related
M. agusanensis in more coriaceous leaves, with contrasting purplish or reddish brown
midrib below. Myristica rubrinervis may be confused with M. laevis.

KEY TO THE VARIETIES

la. Lower leaf surface with rather persistent dense indumentum. Fruits 3-4 cm long,
pencarp/2=3. mm jthiieks:....9, 29 ee PS ae See a. var. rubrinervis

b. Lower leaf surface glabrescent. Fruits 5-7 cm long, pericarp S—6 mm thick.......
UE DTS be STATE ANE CUMS oe he Gary ar al es Tale he b. var. duplex

De Wilde — Myristicaceae (Myristica) 555

a. var. rubrinervis

Tree 8-15 m. Lower leaf surface with subpersistent, rather dense pale brownish or
greyish stellate hairs 0.1—0.3 mm, sometimes partially glabrescent. Petiole 1.5—2.5 cm
long. Fruits ovoid(-ellipsoid), 3-4 by 2.5—3 cm, hairs persistent, dense, scurfy, dark
rusty, 0.1—0.2 mm; pericarp 2-3 mm; seeds 2—3 cm long.

Field-notes — Bark blackish, shallowly fissured; inner bark reddish purple; exudate
red. Fruits brown.

Distribution — Malesia: Philippines (Palawan).

Habitat & Ecology — Stunted montane rain forest with many epiphytes but little
moss: closed broad-leaved rain forest on ultrabasic rock; 450-900 m altitude; fr. Mar.,
May.

b. var. duplex W. J. de Wilde

Myristica rubrinervis W.J. de Wilde var. duplex W. J. de Wilde, Blumea 42 (1997) 182. — Type:
Ridsdale SMHI 1528, Philippines, Palawan.

Tree 20-30 m. Lower leaf surface (late) glabrescent, hairs rather sparse, 0.1—0.2 mm.
Petiole 2-4 cm long. Fruits ovoid-oblong or ellipsoid-oblong, 5—7 by 3—3.5 cm, apex
+ narrowly rounded, hairs persistent, scurfy, bright rust-coloured, 0.1—0.2 mm; pericarp
5-6 mm: seeds 3.5—4.5 cm long.

Field-notes — Large canopy trees. Bark blackish brown, pustular, flaky, or vertically
finely cracked; inner bark red or pinkish brown, sap reddish, watery; wood pale. Leaves
very pale or glaucous below with yellow nerves. Fruits and seeds brown.

Distribution — Malesia: Philippines (Palawan, Mindoro, Leyte); on Leyte one slightly
deviating specimen occurs (Soejarto c. s. 8079), in leaves approaching M. agusanensis.

Habitat & Ecology — Rain forest (with Dipterocarpus), montane and ridge forest
(with A gathis); 40-900 m altitude; fl. (immature) & fr. Feb—Apr.

Note — The species is named after the purplish red contrasting colour of midrib and
nerves on the lower leaf surface; for fresh leaves the nerves were mentioned as being
yellow on the otherwise glaucous lower leaf surface.

117. Myristica rumphii (Blume) Kosterm.

Myristica rumphii (Blume) Kosterm., Gard. Bull. Sing. 22 (1968) 446; W.J. de Wilde, Blumea 42
(1997) 182. — Tetranthera rumphii Blume, Mus. Bot. Lugd. Bat. 1 (1851) 382. — Litsea rumphii
(Blume) Fern.-Vill. in Blanco, Fl. Filip. Nov. App. ed. 3 (1880) 180 (cf. Kosterm., Bibl. Laur.
874, no. 402). — Type: Spanoghe s.n. (L, HLB 905234-28), Timor, annotated as originating
from the Moluccas.

Tree 12—25(—30) m. Twigs terete or faintly 2-angled, 2.5—4(—-6) mm diameter, yellow-
ish or reddish brown, or blackish, early glabrescent, hairs 0.1 mm or less; older twigs
with scattered concolorous lenticels. Leaves chartaceous or (sub)coriaceous, (elliptic-)
oblong, base cuneate, rounded, or (sub)cordate, apex acute (with + blunt tip) or acute-
acuminate, 17-30 by 5-10 cm; upper surface olivaceous or brown, lower surface grey-
ish, (sub)glabrescent, indumentum variable, hairs scale-like, + touching, but surface vis-
ible, 0.1 mm long or less, sometimes mixed with few scattered emergents; + papillose;
dots absent: midrib above slightly raised, lateral nerves 16-28 per side, at 45-—60° to the

556 Flora Malesiana, Ser. I, Vol. 14 (2000)

midrib, sunken above, lines of interarching and venation faint or distinct; petiole 12—25
by 2-4 mm; leaf bud 10-15 by 2-3 mm, hairs appressed, 0.2—0.5 mm. Inflorescences
between the leaves or below, of the Knema-type; male inflorescences not seen; female:
(sub)sessile, simple or 2- to 3-armed, or irregularly shaped, scar-covered, brachyblasts
to 8 mm long, short-woolly rust pubescent, with clusters of 24 flowers, buds of various
sizes. Male flowers not seen. Female flowers: with hairs 0.2—0.4 mm, pedicel 0.5—1 mm,
bracteole ovate, 2—2.5 mm, subacute, late caducous, apical; buds ovoid, apex subacute,
base broadly rounded, 5 by 3.5—4 mm, cleft c. 1/3; ovary ovoid, 3 mm long, pubescent.
Fruits usually single, subsessile, ovoid-ellipsoid, 2.5—4 by 2—2.5 cm, hairs dense, vel-
vety, (dark) rust-coloured, 0.3-1 mm; pericarp 2—4.5 mm thick; seeds 2.5 cm; fruiting
pedicel 1—3 mm long.

Distribution — Malesia: Lesser Sunda Islands (except Bali and Lombok). There are
two varieties.

Note — A homogeneous species, characterized by the weak minute indumentum on
the lower leaf surface, the rather small fruits (2.5—4 cm long) with persistent short indu-
mentum, and the + sessile inflorescences. The species resembles M. fatua, which has a
densely interwoven indumentum on the lower leaf surface.

KEY TO THE VARIETIES

la. Twigs 2.5—3.5 mm diameter, yellowish or reddish brown. Leaves (thinly) chartaceous,
base cuneate or (narrowly) rounded. Fruits 2.5—3.5(—4) cm long, hairs bright brown,
0.5=1 mm-_— Lesser, sunda islands nce atin hseie doe ie a. var. rumphii

b. Twigs 3—4 mm diameter, blackish brown. Leaves coriaceous, base broadly rounded
or subcordate, sometimes short-cuneate. Fruits 4 cm long, hairs (very) dark brown,
Or3:mim., Flores, G00 = 1400 iccees Hy cetesg) Seer ices anu, Chee b. var. florentis

a. var. rumphii

Myristica rumphii (Blume) Kosterm., Gard. Bull. Sing. 22 (1968) 446; W. J. de Wilde, Blumea 42
(1997) 182. — Tetranthera rumphii Blume, Mus. Bot. Lugd. Bat. 1 (1851) 382. — Litsea rumphii
(Blume) Fern.-Vill. in Blanco, Fl. Filip. Nov. App. ed. 3 (1880) 180 (cf. Kosterm., Bibl. Laur.
874, no. 402).

Myristica spanogheana Migq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 47; Warb., Mon. Myrist. (1897)
531, t. 15. — Myristica glauca auct. non Blume: Span., Linnaea 15 (1841) 346. — Myristica
fatua Houtt. var. spanogheana (Migq.) J. Sinclair, Gard. Bull. Sing. 23 (1968) 304, f. 39, p.p.
(excl. M. sumbawana). — Type: Spanoghe s.n., Timor.

Twigs 2.5—3.5 mm diameter, yellowish to reddish brown. Leaves (thinly) chartaceous,
(elliptic-)oblong, base rounded or cuneate, lateral nerves 22—28 pairs; lower leaf surface
usually greyish, (sub)glabrescent, hairs scattered, pale, greyish brown, scale-like, 0.1 mm.
Leaf bud with appressed hairs 0.3—0.5 mm. Fruits 2.5—4 cm long, hairs bright brown,
0.5—1 mm; pericarp 2—4.5 mm thick.

Field-notes — Tree c. 10 m tall (may be up to 30 m). Bark black. Fruits rusty.

Distribution — Malesia: Lesser Sunda Islands (Sumbawa, Sumba, Flores, Timor,
Wetar, Damar).

Habitat & Ecology — Lowland and montane forest; scattered in wet forest; altitude
100—800 m; fl. & fr. throughout the year.

De Wilde — Myristicaceae (Myristica) 7)

Notes — 1. Var. rumphii is identical with Sinclair’s Myristica fatua var. spanogheana,
one of the 14 varieties recognized by Sinclair (1968) in his large concept of M. fatua;,
however, the East Malesian varieties are treated now as four related species, each with
its own distributional area, viz. M. fatua, M. mindanaensis, M. rumphii, and M. sango-
woensis.

2. Schmutz 4013 (W Flores, at 300 m alt.), with female flowers, keys out to Myristica
rumphii on account of its almost glabrous lower leaf surface, with only a very faint and
fine arachnoid covering. It differs from the remainder of the material of var. rumphii in
the more glossy upper leaf surface, and particularly in the larger female flowers (6 by
4.5—5 mm), with large late caducous bracteole, 5 mm long, and distinctly longer hairs,
0.5—-1 mm. More complete material from the area is needed to assess the status of this
deviating specimen.

b. var. florentis W. J. de Wilde

Myristica rumphii (Blume) Kosterm. var. florentis W. J. de Wilde, Blumea 42 (1997) 183. — Type:
Kostermans & Wirawan 728, Lesser Sunda Islands, Flores.

Twigs 3—4 mm diameter, blackish brown. Leaves coriaceous, elliptic-oblong, broadly
rounded or subcordate at base, rarely narrowly rounded, lateral nerves 15—22 pairs; lower
leaf surface pale greyish brown, late glabrescent, hairs dense or scattered, brownish,
scale-like, 0.1 mm, sometimes with a few scattered longer appressed hairs. Leaf bud
with dense hairs 0.2 mm. Fruits 4 cm long, hairs dense, dark brown, 0.2—0.3 mm; pericarp
4—4.5 mm thick.

Field-notes — Bark + rough, 2 mm, dark brown, peeling off in small particles. Living
bark light brown. Sap red. Fruits rusty.

Distribution — Malesia: Lesser Sunda Islands (Flores).

Habitat & Ecology — Montane forest, 600-1400 m altitude; fr. Apr.—May, Nov.

118. Myristica sangowoensis (J. Sinclair) W. J. de Wilde

Myristica sangowoensis (J. Sinclair) W.J. de Wilde, Blumea 35 (1990) 254, f.2 (12); 42 (1997)
183. — Myristica fatua Houtt. var. sangowoensis J. Sinclair, Gard. Bull. Sing. 23 (1968) 304,
f. 36B, C. — Type: Kostermans 1039, N Moluccas.

Tree 10-15 m. Twigs sometimes faintly ridged, 3.5—4(—5) mm diameter, early glabres-
cent, hairs 0.1 mm; older twigs conspicuously lenticellate. Leaves chartaceous or thinly
coriaceous, oblong or (ob)lanceolate, 20-35 by 7-12 cm, base rounded or short-attenu-
ate, apex acute-acuminate; upper surface olivaceous or brown, lower surface cinnamon
by dense persistent indumentum of stellate(-dendroid) hairs 0.1—0.2(—0.3) mm, mixed
with few appressed longer hairs; not papillose; dots absent; midrib above slightly raised,
lateral nerves 20-25 per side, at 50—70° to the midrib, flat or sunken, lines of interarching
and venation indistinct or invisible; petiole 18-25 by 3.5—5 mm; leaf bud 15 by 3.5 mm,
hairs 0.1(—0.2) mm. Inflorescences between the leaves or below, of the Knema-type:
sessile, single or forked, woody, scar-covered brachyblasts to 5 mm long, with hairs 0.1—
(0.2 mm: bracts caducous; in male: with subumbels of 3-7 flowers, buds of various sizes;
female similar, fewer-flowered; flowers with dense hairs (0.3—)0.5 mm long. Male flow-

558 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 84. Myristica sarcantha W.J. de Wilde. a. Leafy twig; b. male flower, opened; note scar of
caducous bracteole [both: McDonald & Ismail 3800]. — Scale bar for a = 2 cm, for b = 1.65 mm.

De Wilde — Myristicaceae (Myristica) 559

ers (Whitmore c.s. TCW 3608): pedicel (upcurved when young) 4-5 mm, bracteole
+ obovate-elliptic, 5 mm, (sub)persistent; buds (ob)ovoid-ellipsoid, 6 by 3—4 mm, cleft
c. 1/3, lobes 0.5 mm thick, perianth inside somewhat callose and bullate (always?).
Androecium small, 4 mm; androphore | by 0.7 mm, at base with few hairs, dark brown,
0.2—0.3 mm; synandrium 2.5 by 0.8(—1) mm, thecae 16-20, sterile apex more or less
acute, 0.5—0.7 mm long, glabrous. Female flowers not seen. Fruits solitary, subsessile,
(broadly) ellipsoid (immature fruit subglobose in Kostermans 1039), 5.5-8.5 by 4-5
cm, base rounded or + attenuate, hairs cinnamon, scurfy, 0.1—0.2 mm; pericarp 10-15
mm thick; seeds ellipsoid(-oblong), 4.5 cm; fruiting peduncle including pedicel thick,
8 by 8 mm. — Fig. 65c.

Field-notes — Small tree. Bark dark brown; inner bark red brown, fibrous. Leaves
golden brown below, becoming grey. Flowers golden yellow in bud. Fruits red.

Distribution — Malesia: Moluccas (Morotai, Halmahera, Bacan).

Habitat & Ecology — Submontane and lower montane forest; forest on ridge with
abundant Fagaceae; volcanic soil; on G. Sango(a)wo (Morotai) many specimens seen;
400-800 m altitude; fl. Sept.; fr. May, Aug.

Note — May be confused with M. fatua which has smaller fruits with thinner pericarp,
and longer hairs, 0.5—1 mm long; the pericarp of M. fatua subsp. affinis (Sulawesi) has
also a thick pericarp but differs in the much more conspicuous indumentum, male flow-
ers of subsp. fatua generally are smaller and those of subsp. affinis lack a sterile apex on
the synandrium.

119. Myristica sarcantha W. J. de Wilde

Myristica sarcantha W. J. de Wilde, Blumea 43 ( 1998) 175, f. 3. —Type: McDonald & Ismail 3800,
male fl., northern Irian Jaya.

Treelet 4 m. Twigs 5—6 mm diameter, smooth, hardly striate, bright brown, subterete
with shallow lines from petiole to petiole, partially hollow (see field-notes) but ant-holes
not seen, glabrous (glabrescent, but twig apex not seen); older twigs not seen; lenticels
small, scattered, inconspicuous. Leaves membranous, obovate-oblong, 22-32 by 8-15
cm, base cuneate, apex broadly rounded with acute-acuminate tip; upper surface green-
olivaceous, lower surface pale, subglabrous, hairs extremely minute, scattered, whitish,
appressed; minutely irregularly papillose; dots absent; midrib almost flat above, lateral
nerves 20-22 per side, at 70—80° to the midrib, flat, lines of interarching and venation
slender, distinct at both surfaces; petiole 25-30 by 3 mm, glabrous; leaf bud not seen.
Male inflorescences between the leaves, of the Knema-type: sessile, scar-covered
brachyblasts to 5 mm diameter, the flowers in subumbels of 2—5, buds slightly variable
in size, with + scattered mealy stellate hairs (less than) 0.1—0.2 mm. Male flowers: pedi-
cel 7-8.5 by 2.5(—3) mm, bracteole scar at about 1/3—1/2 from the apex; buds firm,
oblong-fusiform, 15-16 by (4-)5—6 mm, broadest above the middle, apex acute, base
tapered, apical part triangular with lines of lobe sutures clearly marked, cleft c. 1/3,
lobes (4—)5 mm long, 0.6—0.8 mm thick. Androecium 12-13 mm; androphore 4 by 1.5—
2 mm, appearing to be glabrous but with very minute (less than 0.1 mm) pale stellate
hairs in the lower 2/3; synandrium oblong, acute, 8 by 3 mm, thecae 24-26, sterile apex
0.5—-1 mm long. Female flowers and fruits not seen. — Fig. 84.

560 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Understorey tree, trunk straight, cylindrical, 3 cm diameter, branching
above and below; lateral branches whorled and covered with muricate bodies that attract
aggressive ants. Flowers pale yellow and green, lobes reflexed.

Distribution — Malesia: northern Papua Barat (Sarmi); known only from the type.

Habitat & Ecology — Coastal plain, mixed lowland forest, canopy 25—45 m high;
10-20 m altitude; fl. June.

Notes — 1. This species, belonging to the group of ant-inhabited species, is readily
distinguishable by the large, robust, carnose flowers. In general habit M. sarcantha is
close to M. fissiflora (with deeper lobed less thick-fleshy male perianth, and the bracteole
apical) and to M. fasciculata (mature male flowers not known, but immature flowers
very different). It also resembles M. verruculosa, which has larger flowers with a longer
pedicel.

2. The nature of the muricate bodies mentioned in the field-notes is unknown.

120. Myristica schlechterii W. J. de Wilde

Myristica schlechteri W.J. de Wilde, Blumea 40 (1995) 322. — Type: Schlechter 17461, Papua
New Guinea.

Small tree. Twigs 1(—1.5) mm diameter, glabrescent; hairs 0.5—1 mm long; older twigs
without lenticels. Leaves membranous, oblong-lanceolate, 6—10 by 2—3 cm, base attenu-
ate, apex acute-acuminate, with + blunt tip; upper surface (blackish) brown, lower sur-
face grey, early glabrescent, hairs 0.5—-1 mm; irregularly densely grey-white papillose,
appearing non-papillose; dots absent; midrib flat or sunken above; lateral nerves 6—10
per side, at 70—80° to the midrib, far apart, brown and contrasting in colour below, lines
of interarching and venation faint; petiole 6—8 by 0.8(—1) mm; leaf bud rather blunt,
4—5 by 1.5—2 mm, hairs dense, rough-woolly, 0.5—1 mm long. Inflorescences between
the leaves, in the basal part of innovations, of the Knema-type; in male: consisting of
2 or 3 flowers at the end of a 1.5 mm long peduncle; bracts not seen. Whole inflores-
cence, including flowers, densely clothed in reddish brown rough-woolly hairs 0.5—-1
mm. Female inflorescences not seen. Male flowers: pedicel 7 by 0.7—0.8 mm, bracteole
less than 1 mm, caducous, scar 0.5(—1) mm below the apex; buds (ovoid-)oblong, apex
subacute or + blunt, base + rounded, (8—)9 by 3 mm, cleft c. 1/4, lobes 2(—2.5) mm long,
0.1—0.2 mm thick, perianth inside fine-warty. Androecium tapering towards the apex,
largely consisting of the synandrium; androphore (0.1—)0.2 mm long, 1 mm wide, at very
base with a collar of minute reddish brown hairs 0.1—0.2 mm; synandrium (5.5—)6 by
1.2(-1.5) mm, thecae 12-14, + contiguous, sterile apex narrow, + blunt, (0.3—)0.4 mm.
Female flowers and fruits not seen.

Distribution — Malesia: Papua New Guinea (Morobe Prov., Waria area, near Pema);
known only from the type, in K.

Habitat & Ecology — Understorey treelet; c. 350 m altitude; fl. Mar.

Note — This species is characterized by a small stature, with slender twigs and small
herbaceous leaves, the innovations, inflorescences, and flowers densely clothed in ru-
fous rusty rough-woolly indumentum with hairs 0.5—1 mm long, and by the unique al-
most sessile synandrium. By the small stature and the finely verrucose inner surface of
the perianth M. schlechteri seems related to species like M. crassipes, M. filipes, M. flos-
culosa, M. pumila, and M. sogeriensis.

De Wilde — Myristicaceae (Myristica) 561

121. Myristica schleinitzii Eng].

Myristica schleinitzii Engl., Bot. Jahrb. Syst. 7 (1886) 455; Warb., Mon. Myrist. (1897) 392, t. 19;
A.C.Sm., J. Arnold Arbor. 22 (1941) 74; J. Sinclair, Gard. Bull. Sing. 23 (1968) 202, f. 15;
Foreman, Handb. Fl. Papua New Guinea 1 (1978) 207, f. 91; W.J. de Wilde, Blumea 38 (1994)
367, f. 6; 40 (1995) 322. — Type: Naumann s.n., 20 July 1875 (B, lost), New Hanover.

Myristica mas (non Rumph.) Labill., Rélat. du Voy. ala Rech. de la Pérouse 1 (1799) 237. — Myris-
tica spanogheana auct. non Miq.: K.Schum. Bot. Jahrb. Syst. 9 (1887) 200. — Myristica faro-
ensis Hemsl., Ann. Bot. 5 (1891) 506. — Type: Guppy 209, Santa Isabel I.

Tree 5—15(—20) m. Twigs 1.5—3 mm diameter, (pale) reddish or yellowish brown,
glabrescent, hairs 0.1 mm; older twigs + fissured or cracking; lenticels inconspicuous.
Leaves membranous or thinly chartaceous, elliptic(-oblong), 7-27 by 2.5—-12 cm, base
rounded or shallowly cordate, sometimes subacute, apex narrowed with blunt tip; upper
surface olivaceous, lower surface grey-olivaceous, seemingly glabrous, but hairs sparse,
0.1 mm or less; distinctly papillose; dots absent; midrib slightly raised above; lateral
nerves 8—13(—20) per side, at 50—70(—80)° to the midrib, flat or slightly sunken above,
lines of interarching and venation (usually) indistinct; petiole 6-20 by 1.5—3 mm,
glabrescent; leaf bud 5—12(—15) by 1.5—3 mm, hairs 0.1—0.3 mm. Inflorescences be-
tween the leaves and below, glabrescent, paniculate, with flattened peduncle; in male:
(1-)2-7 cm long, peduncle 10—40 by 1.5—3.5 mm, lower lateral branches to 20(—25)
mm, central axis absent or to 30 mm long, with 1—4 lateral branches, each branch with a
scar-covered brachyblast to 15 mm, with a subumbel of 3—10 flowers, buds of various
sizes; female as male but shorter, subumbels of 2—6 flowers; bracts 1—-1.5 mm, pubes-
cent, caducous, the scars sometimes higher up on the branches; flowers with thick peri-
anth, hairs 0.1 mm, partly glabrescent. Male flowers: pedicel 3.5—5 by 0.5—0.7 mm,
bracteole ovate, 1—1.5 mm, persistent, (sub)apical; buds ovoid-oblong, 5.5—6.5 by 2.5—
3.5 mm, sometimes faintly angular, apex subacute or + blunt, cleft 1/5—1/3, lobes 1.5—2
mm long. Androecium 5—6 mm; androphore about as wide as the synandrium, 2—2.5 mm
long, glabrous or hairs sparse, 0.1 mm or less; synandrium 2.5—4 by 0.7—1 mm, apex
blunt, thecae 10-15, often with short sterile apical appendages 0.1 mm, sterile apex 0.2
mm or absent. Female flowers: pedicel slender, 2-5 mm; buds ovoid-oblong, 4.5—5.5 by
2.5-3 mm, apex + acute, cleft c. 1/3; ovary ovoid-ellipsoid, 3.5 by 2 mm, hairs less than
0.1 mm. Fruits solitary or 2—6 per infructescence, ellipsoid or oblong, 2—3.5(—4) by
1-1.7 cm, apex (sub)acute, base (sub)acute or narrowed into an up to 5 mm long pseudo-
stalk, hairs mealy, pale brown, 0.1—0.2 mm, glabrescent except very base and apex;
pericarp 0.5—1 mm thick, brown, wrinkled or + longitudinally striate; seeds ellipsoid,
2-2.5(—3) cm, (blackish or) dark brown; aril deeply slenderly laciniate; fruiting pedicel
3-7 mm. — Fig. 85.

Field-notes — Erect or bushy trees, largely glabrous, but growing twigs with devel-
oping leaves and inflorescences with dense short brown-yellow indumentum. Stem
crooked or not, mostly without buttresses, stilt-roots occasionally present; branches some-
times whorled, horizontal (Terminalia-branching). Bark soft, brown-red, when old grey-
brown to blackish, longitudinally fissured and/or flaking (scaly); slash bark banded or
not, straw turning pink(-brown); exudate pinkish, watery or opaque; wood rather hard,
pale, straw or orange, discolouring to brown; heartwood reddish. Leaves glossy (blue-)
green above, pale green to glaucous below, midrib + yellow. Flowers cream or (green-)
yellow, scented or not, sometimes not opening. Fruits green-yellow or orange, partly

562 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 85. Myristica schleinitzii Engl. a. Twig with male inflorescences; b. ditto, aged, axillary to leaf
scar, with distinct scar-covered flower-bearing brachyblasts; c. female inflorescence; d. male bud,
opened, showing androecium; e. opened female bud just before anthesis, showing pistil; f. branch
with infructescences and mature fruits [a, d: Goetghebeur 3379; b: Brass 3288; c, e: Lauterbach
1492: f: NGF 15512 (Croft & Katik)]. — Scale bar for a—c, f = 2 cm, for d, e = 1.65 mm.

De Wilde — Myristicaceae (Myristica) 563

with thin golden indumentum; seeds dark brown to black, glossy, slightly aromatic, in
mature fruit white with yellow-green aril (once recorded)

Distribution — Malesia: northern part of Papua New Guinea, (W & E Sepik, Madang,
Morobe, Northern?, Milne Bay, and Central Proy.: one collection); Papuan Islands
(d’Entrecasteaux); Bismarck Archipelago (New Hanover, New Ireland, New Britain);
Solomon Islands (Bougainville, Santa Isabel, Novatana, New Georgia Group).

Habitat & Ecology — Coastal. Forest on cliffs and crests, well-drained; sand dunes,
on coral beach, forest in and behind mangrove; also inland but not too far from the sea;
in New Britain common in the Calophyllum, Intsia, and Terminalia association; also on
limestone. In Milne Bay Prov., Dugumunu I., in forest on coral debris with Eugenia;
0-200(—350) m altitude; fl. & fr. throughout the year.

Uses — Aril used for nail varnish.

Note — Related to M. rosselensis, M. garciniifolia (habitat similar, Papua Barat), and
M. inopinata, the latter two with stouter leaves. Myristica schleinitzii is a homogeneous
species, characterized by paniculate inflorescences with flattened peduncle, elegant yel-
lowish twigs with small- or medium-sized leaves, often with subcordate base, greenish
on drying, papillose on lower surface (lens!), and by rather small subglabrous fruits in
pedunculate infructescences.

122. Myristica scripta W. J. de Wilde

Myristica scripta W.J. de Wilde, Blumea 40 (1995) 323; 42 (1997) 183. — Type: Pullen 7185,
Papua New Guinea.

Tree 4-30 m. Twigs terete or faintly angular, (1.5—)2—3 mm diameter, glabrescent,
hairs 0.1(—0.2) mm; older twigs faintly fissured but neither cracking nor flaking; lenticels
not conspicuous. Leaves thinly coriaceous (but brittle), ovate-elliptic or ovate-lanceo-
late, 6-17(—25) by 2.5—6(—9) cm, base short-attenuate or broadly rounded, apex + blunt
or short-acuminate, often with blunt tip; upper surface dark brown, lower surface pale
brown, glabrescent, hairs sparse, 0.1 mm or less; inconspicuously papillose; dots and
dashes present, best to be seen towards the margin (lens!); midrib usually raised above:
lateral nerves 12—20(—30) per side, at 50—80° to the midrib, closely set, flat and hardly
visible above, almost flat and faint below, lines of interarching and venation faint or
invisible; petiole 8-15 by 1.5—3 mm; leaf bud 8-15 by 2—2.5 mm, hairs 0.1—0.2 mm.
Inflorescences between the leaves and below, of the Knema-type: up to 1 mm peduncu-
late, woody, simple or 2- to 3-armed, scar-covered brachyblasts to 5 mm long, ending
in subumbels of flowers, brown pubescent or glabrescent; in male: with (5—)10—40
flowers, buds of various sizes; in female: with (1—)2—5 flowers, buds about equal in size:
flowers with hairs 0.1—0.2 mm, sometimes partly glabrescent. Male flowers: pedicel
2-4 by 0.6-0.8 mm, bracteole rounded, 0.5—1 mm, caducous, (sub)apical; buds (ellip-
soid-)oblong, 3.5—5 by 2—2.5 mm, faintly triangular, cleft c. 1/3, lobes 1.51 .7 mm long,
0.3 mm thick. Androecium subcylindrical, 2.5—4.2 mm; androphore shallowly grooved,
1-1.7 by 0.4-0.6 mm, glabrous or with few minute hairs in the lower half; synandrium
narrowly oblong, 1.5—2.5 by 0.7-0.8 mm, thecae c. 12, contiguous or not, sterile apex
irregularly rounded, 0.2—0.4 mm. Female flowers: pedicel 4-8 by 1-1.5 mm, bracteole
scar 0.5—2 mm below the apex: buds ovoid(-oblong), narrowed to the apex, 4.5—5.5 by
3-4 mm, lobes 1.5—2 mm long; ovary ovoid(-oblong), 3.5—4 by 2—2.5 mm, hairs 0.2

564 Flora Malesiana, Ser. I, Vol. 14 (2000)

mm. Fruits solitary or 2—5 in a cluster, ovoid, (broadly) ellipsoid, or subglobose, (1.5—)
22.5 by 1.5-2 cm, apex rounded or slightly narrowed, sometimes minutely beaked,
hairs rusty, mealy, 0.1(—0.2) mm; pericarp 2(—3) mm thick; seeds ellipsoid, 1.5 cm; fruit-
ing pedicel slender or + thick, 5—10 mm long, bracteole scar 1-3 mm below the apex.

Distribution — Malesia: Moluccas (Aru Is., type variety); W, C, & E New Guinea.
There are two varieties consisting of specimens with conspicuously different fruiting
pedicels, but similar in vegetative and flower characters.

Habitat & Ecology — Lowland forest up to 700 m altitude.

Note — Vegetatively M. scripta resembles M. lancifolia from the Moluccas and the
western part of New Guinea in the similar leaves (same size and venation). Myristica
scripta is readily distinguishable by the conspicuous dots on the lower leaf surface.

KEY TO THE VARIETIES

la, Fruitine pedicel.2=sanmy thick, 24 .0.0-ehat eagnee oe See a. var. scripta
b; Fruiting pedicel stout, 5—7 mm thick 2.) .).cu: 2 «as-sem-9- ee: b. var. incrassata

a. var. scripta

Tree up to 30 m. Flowers minutely appressed-pubescent, hairs 0.1—0.2 mm. Fruit-
ing pedicel + slender, 2-3 mm thick, smooth or finely striate, sometimes with minute
lenticels.

Field-notes — Bark without or (rarely) with buttresses to 1.5 m high and out, smooth,
or finely, irregularly or deeply fissured, often peeling or flaking; outer bark thin, dark
brown; inner bark (slash bark) reddish, with wavy appearance, exudate reddish, watery
or sticky; wood white or (pale) brown, sometimes mottled; heartwood rich red. Leaves
grey-green below. Flowers cream, yellow, or brown; ovary pale green. Fruits yellow,
orange, or brown.

Distribution — Malesia: Moluccas (Aru Is.); Papua Barat (Biak, Japen I., Jayapura
area); Papua New Guinea (W Sepik, Western, Gulf Prov.).

Habitat & Ecology — Primary or disturbed dryland forest, on clay or volcanic soil,
limestone; also from forest along dry creeks, streamside forest, stony riverbanks; alti-
tude 0-500 m: fl. & fr. mainly June—Dec. Locally common.

b. var. incrassata W. J. de Wilde

Myristica scripta W.J. de Wilde var. incrassata W.J. de Wilde, Blumea 40 (1995) 324. — Type:
NGF 18306 (Streimann & Lelean), Papua New Guinea.

Tree up to 30 m. Flowers rather roughly tomentose, hairs 0.2 mm. Fruiting pedicel
stout, 5-7 mm thick, conspicuously lengthwise fissured, sometimes with coarse lenticels.

Field-notes — Bole slender, without buttresses; branches verticillate, horizontal. Bark
smooth or finely fissured; live bark light brown; underbark reddish; wood white; exu-
date red, sticky. Leaves greyish green below. Flowers yellow. Fruits yellow-brown.

Distribution — Malesia: S Central New Guinea: SE Papua Barat; Papua New Guinea
(Western Prov., W Sepik Prov.).

Habitat & Ecology — Primary and degraded dryland forest (on volcanic soil) and
swamp forest; 25-600 m altitude; fl. & fr. June—Sept.

De Wilde — Myristicaceae (Myristica) 565

123. Myristica simiarum A. DC.

Myristica simiarum A.DC., Ann. Sc. Nat. 4, 4 (1855) 29; Prodr. 14, 1 (1856) 192; W.J. de Wilde,
Blumea 35 (1990) 254; 42 (1997) 184: Tree Fl. Sabah & Sarawak 3 (2000) 469. — Myristica
elliptica Wall. ex Hook.f. & Thomson var. simiarum (A. DC.) J. Sinclair, Gard. Bull. Sing. 16
(1958) 356; 23 (1968) 62 (key), 190, f. 12. — Type: Callery 34, Philippines, Luzon, Manilla.

For more references and synonyms see the subspecies.

Tree 5—36 m. Twigs 1.5—3(—3.5) mm diameter, whitish or pale brown (or dark brown),
early glabrescent, hairs 0.1—0.2(—0.4) mm; older twigs without or with few to many
small paler coloured lenticels. Leaves membranous or thinly chartaceous, very brittle,
ovate-elliptic or oblong(-lanceolate), 11—18(—32) by 3—7(—12) cm, base acute or nearly
rounded, apex acute-acuminate; upper surface light or blackish brown, sometimes scab-
rous, lower surface grey(-brown), pubescent or (late) glabrescent, hairs appressed, silky,
(0.1—)0.3-0.5 mm; papillation present, absent, or indistinct; dots absent; midrib slightly
raised above; lateral nerves 10—14(—18) per side, at 45—60° to the midrib, flat and faint
above, lines of interarching distinct or not, venation faint; petiole 14—25(—35) by 1-2
(—2.5) mm; leaf bud 12—25 by 1—2.5 mm, hairs appressed, pale or (reddish) brown, 0.5—
1 mm. Inflorescences pedunculate, between the leaves (rarely + below), late glabrescent,
hairs scattered, 0.5(—1) mm; in male: + slender, 1.5—4 by 1.5—3.5 cm, peduncle 7—15
(—17) by 1-2 mm, basal branches rectangular to the peduncle, to 7 mm long, central axis
simple, or with 2 or 3 sessile or short-stalked, scar-covered brachyblasts to 10 mm long
(rarely absent), flowers in clusters of 3-12, buds of + variable sizes; bracts + triangular,
1-1.5 mm,+ pubescent, subpersistent or caducous; female as male, but smaller and fewer
flowered: flowers with hairs 0.2—0.3(—0.5) mm. Male flowers: pedicel (2—)3—5 mm,
slender, bracteole + triangular, 1-2 mm, pubescent, persistent; buds + ellipsoid-oblong,
+ triangular, 4.5—8 by 2-3 mm, apex + acute, cleft c. 1/2, the sutures thicker than rest of
perianth. Androecium slender, 4—5.5 mm; androphore (0.5—)1—1.5 mm , hairs sparse or
dense, 0.3—1 mm; synandrium (2.5—)3—3.5 by 0.8—1 mm, thecae 12—16(?), sterile apex
irregularly blunt, 0.3-0.5 mm long, or shallowly hollowed. Female flowers: pedicel 1.5—
2(-2.5) mm long, bracteole 1—1.5 mm long, persistent; buds ovoid, 5—6 by 3-3.5 mm,
+ angled, cleft c. 1/3 (to nearly 1/2); ovary (narrowly) ovoid, 3 by 1.5—2 mm, hairs 0.5
mm, stigma broadly 2-lipped, 0.3-0.5 mm long. Fruits (variable according to the sub-
species), solitary or 2 or 3 per infructescence, subglobose or broadly ellipsoid or ovoid-
ellipsoid, 2—4 by 2-3 cm, apex rounded or shortly beaked, base rounded or shortly con-
tracted, glabrescent or thinly pubescent, hairs inconspicuous, grey-brown, silky, (0.3—)0.5
mm; pericarp 2-5 mm thick, (blackish) brown, seeds broad-ellipsoid; fruiting pedicel
slender or stout, 3-6 mm long. — Fig. 86.

Distribution — Malesia: Borneo (W & NE Kalimantan, W Sarawak), Philippines,
Sulawesi, Moluccas (Bacan, Sula Is.).

Note — A variable species in which, according to the area of distribution and based
on rather weak differences in the fruits, three subspecies are recognized. Further varia-
tion is noteworthy in subsp. simiarum and briefly discussed there. Sinclair (1968) in-
cluded the species in the related M. elliptica, but M. simiarum differs in its inflores-
cences and its smaller fruits. Both species have the markedly triangular upper part of the
buds in common. Myristica elliptica, the swamp nutmeg, is a species mostly from marshy
forest, frequently with stilt-roots which have never been recorded for M. sumiarum.

566

Flora Malesiana, Ser. I, Vol. 14 (2000)

ye

Ys

On
Mi}

IW
WH)

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SS RSG

Scene

De Wilde — Myristicaceae (Myristica) 567

a

—

Fig. 86. Myristica simiarum A.DC. subsp. simiarum. a. Male flowering twig; b. part of male flow-
ering twig, inflorescences in an advanced stage, with scar-covered brachyblasts; c. male bud;
d. ditto, opened, showing androecium:; perianth triangular in apical part; e. twig with infructescence
with one fruit. — M. simiarum subsp. celebica (Miq.) W.J. de Wilde. f. Twig with male inflores-
cence; g. twig with female inflorescence; h. female flower at anthesis; i. ditto, opened, showing
pistil; j. part of older twig with two infructescences, each with one fruit. — M. simiarum subsp.
calcarea W.J. de Wilde. k. Part of twig with male inflorescence; 1. male bud, opened, showing an-
droecium; m. twig with infructescence with two fruits [a: BS 80173 (Ramos): b—d: PNH 91851
(Lagrimas); e: Vidal 3570; f: Koorders 18161; g—j: Whitmore c.s. TCW 3436: k: Kostermans 13772:
1: Kostermans 13866: m: S 34468]. — Scale bars for a, b, e-g, j, k, m= 2 cm; c, d, h, i, ] = 1.7 mm.

KEY TO THE SUBSPECIES

la. Fruits globose or broadly ellipsoid, 1.5—2.5(-3) cm long, pseudostalk + absent or
short and 2—4 mm broad; pericarp 2—4 mm thick. Leaves glabrescent below. — Phil-
ippines [& S Taiwan: Botel Tobago Is.] ......---+++++--- a. subsp. simiarum

b. Fruits (broadly) ellipsoid or ovoid, 3—3.5 cm long, pseudostalk stout, 4-5 mm broad;
pericarp 4-5 mm thick. Leaves glabrescent below, or with dispersed hairs. —
Sulawesi, Bacan, Sula I., Philippines (Zamboanga) .......--. c. subsp. celebica

c. Fruits ellipsoid, 3.5—4 cm long, pseudostalk 3-4 mm broad; pericarp 3 mm thick
(apparently much thicker when fresh). Leaves with persistent dense bright brown
indumentum below. — W & NE Kalimantan, W Sarawak .... b. subsp. calearea

568 Flora Malesiana, Ser. I, Vol. 14 (2000)

a. subsp. simiarum

Myristica simiarum A.DC., Ann. Sc. Nat. 4, 4 (1855) 29; Prodr. 14, 1 (1856) 192; Warb., Mon.
Myrist. (1897) 397, t. 13; Hayata, Icon. Pl. Formos. 3 (1913) 156; Gen. Index Fl. Formos. (1917)
61; Merr., Enum. Philipp. Flow. Pl. 2 (1923) 180; Kudo & Masam., Ann. Rep. Taihoku Bot.
Gard. 2 (1932) 89; Kaneh., Formos. Trees, rev. ed. (1936) 194; Hui-Lin Li, Woody Flora Taiwan
(1963) 195: W.J. de Wilde, Blumea 42 (1997) 184. — Myristica elliptica Wall. ex Hook. f. &
Thomson var. simiarum (A.DC.) J. Sinclair, Gard. Bull. Sing. 16 (1958) 356; 23 (1968) 62 (key)
190, f. 12A—-E, I-J.

Myristica discolor Merr., Philipp. J. Sci., Bot. 13 (1918) 218. — Type: Curran 13155, Philippines.

Gymnacranthera sulphurascens Elmer, nom. nud. in sched.; Merr., Enum. Philipp. Flow. Pl. 2 (1923)
180 in note sub Myristica simiarum (Elmer 7377). — Myristica sulphurascens Elmer, Leafl.
Philipp. Bot. 10 (1939) 3809, nom. nud. sub Myristica simiarum, Index Kew. Suppl. 10 (1947)
149 (Ramos 1609).

Myristica elliptilimba Merr., nom. nud. in sched. (Ramos & Edano 31173).

Myristica merrillii Warb., in sched. (Elmer 6357).

Myristica verrucosula Merr., nom. nud. in sched. (de Mesa 27584; Ramos & Edanio 29206; Ramos
& Pascasio 34469).

Tree 5-15 m. Leaves: upper surface olivaceous-brown or blackish (when blackish
usually grey or whitish below, sometimes scabrous, see note 2), lower surface glabrescent,
hairs scattered; (in)distinctly papillose or not. Male flowers: buds variable in size, 4.5—-8
mm long, thinly pubescent. Female flowers as in the species. Fruits globose or broadly
ellipsoid, (1.5—)2—2.5(—3) by 1.5-2(-2.5) cm, apex rounded or up to 1.5 mm long beaked,
base rounded or contracted into a pseudostalk up to 2 mm long and 2—4 mm broad;
pericarp brown or blackish, 2-4 mm thick. — Fig. 86a-e.

Field-notes— Tree, dbh to 30 cm, without stilt-roots, sometimes with buttresses. Bark
slightly rough, reddish brown; sap dark red. Leaves glossy, medium green above; glau-
cous with yellowish green midrib and nerves below. Flowers yellow or brown. Fruits
green, then orange. Testa mottled brown.

Distribution — Taiwan (Botel Tobago Is., no specimens seen); Malesia: Philippines
(Luzon, Samar, Mindanao, Batan).

Habitat & Ecology — Forest, also on slopes, rough country; 0—600 m altitude; fl. &
fr. throughout the year.

Uses — Bark applied for internal pains (Mindanao).

Notes — 1. Anthers possibly 6—10; the thecae are narrow and closely appressed so
that the number of anthers is difficult to count.

2. The type subspecies, occurring throughout the Philippines, is remarkably variable,
whereas the other two subspecies seem rather uniform.

The leaves of many specimens from Luzon dry conspicuously blackish above, and
grey or almost whitish below. Some specimens have relatively large male flowers, while
others have remarkably small flowers on slender inflorescences. The fruits dry black-
ish as well. The leaves of specimens from Samar and Mindanao are (olivaceous) brown
above and grey-brown beneath; possibly these specimens always have brown fruits when
dry. In some collections the leaves have minute whitish dots beneath, in others these are
lacking. In some specimens from Luzon the leaves (drying blackish or olivaceous) are
verruculose or scabrous, caused by crustations in the leaf tissue, but possibly this feature
has little taxonomic merit.

De Wilde — Myristicaceae (Myristica) 569

b. subsp. calcarea W. J. de Wilde

Myristica simiarum A. DC. subsp. calcarea W. J. de Wilde, Blumea 42 (1997) 184. — Type: Koster-
mans 13866, E Kalimantan, Berouw.

Tree 20-36 m. Leaves: upper surface olivaceous-brown, lower surface (grey-)brown,
indumentum dense, persistent, hairs flat-lying, brown; indistinctly papillose or not. Male
flowers bright rusty brown pubescent; buds 7-8 mm long. Female flowers not seen.
Fruits broad-ellipsoid, 3.5(—4) by 2.5—3 cm, apex blunt-acute, at base contracted into a
pseudostalk 2—4 mm long, 3—4 mm broad; pericarp dull brown, 3 mm thick (possibly
very much thicker when fresh). — Fig. 86k—m.

Field-notes — Bole to 15 m long, up to 80 cm diameter, buttresses low, to | m high.
Bark superficially fissured or very rough and deeply fissured, hard, dark brown, red-
brown, or black, pitted, 5 mm thick, peeling off in irregular strips 2-3 cm wide, 2-3 mm
thick; living bark 10-15 mm thick, undulate, yellowish brown, red-brown or dark red;
sap pale red, watery; cambium brown-yellow; sapwood 8 cm thick, dirty white or yel-
lowish white; heartwood dark brown. Lower leaf surface brown or golden. Flowers
(brown-)yellow. Fruits yellow or orange-red; seed coat brown with yellow streaks.

Distribution — Malesia: Borneo (W & NE Kalimantan, W Sarawak).

Habitat & Ecology — Forest on (old coral) limestone or sandstone, yellow sandy
soils, hilly country and low mountains, shallow clay soil; 25—700 m altitude; fl. Sept.;
fr. June—Sept.

Note — Almost all collections are from limestone or lime-containing soils; once from
sandy clay derived from granite. Distinguished by the large, brown-yellow flowers and
large fruits, and the leaves with persistent, dense, golden indumentum on the lower sur-
face. The fruits may recall those of small-fruited M. elliptica.

c. subsp. celebica (Miq.) W. J. de Wilde

Myristica simiarum A. DC. subsp. celebica (Miq.) W.J. de Wilde, Blumea 35 (1990) 255, f. 2 (13);
42 (1997) 185. — Myristica celebica Miq., Ann. Mus. Bot. Lugd.-Bat. 2, 1 (1865) 47; Warb.,
Mon. Myrist. (1897) 395, t. 15; Koord., Fl. N.O. Celebes (1898) 570 (not M. celebica Gand.,
1919 = M. fatua Houtt.). — Myristica elliptica Hook. f. & Thomson var. celebica (Miq.) J. Sinclair,
Gard. Bull. Sing. 16 (1958) 356; 23 (1968) 194, f. 12F. — Syntypes: Sulawesi: Forsten s.n.,
Teijsmann 5801 (U, lecto, here designated); Bacan: de Vriese s.n. (Myristica fallax Migq., in
sched.); Buru: de Vriese s.n. (no specimens seen).

Myristica fragrans auct. non Houtt. forma sylvestris Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1864) 205;
Ann. 2 (1865) 48 (sub Myristica celebica Miq.). — Type: Teijsmann 5872, Moluccas.

Tree 10-30 m. Leaves: upper surface olivaceous brown or dark brown, lower surface
grey-brown, indumentum thin, (late) glabrescent, or subpersistent, hairs + scattered, ap-
pressed, grey(-brown), of varying sizes; distinctly or indistinctly papillose. Male flow-
ers: moderately or densely (grey-)brown pubescent; buds 6—7 mm long. Female flowers
as in the species. Fruits ellipsoid or ovoid, 3-3.5 by 2.5 cm, apex + blunt or 1-2 mm
beaked, base contracted into a stout pseudostalk 2-4 mm long, 4-5 mm broad, pericarp
dull greyish brown to dark brown, 4-5 mm thick. — Fig. 65d, 86f-j.

Field-notes — Tree, dbh c. 40 cm, without buttresses. Outer bark brownish, 0.3—0.4
mm thick, much fissured, peeling off or not; inner bark 7-9 mm, cream or reddish, sap
reddish, clear; middle bark yellow, fibrous. Leaves whitish below. Flowers (female) golden

570 Flora Malesiana, Ser. I, Vol. 14 (2000)

yellow. Fruits green with brown indumentum when immature, when ripe orange-yellow,
orange-brown or golden.

Distribution — Malesia: Sulawesi (incl. MoenaI.); Moluccas (Bacan, Sula Is., Buru);
Philippines (Zamboanga).

Habitat & Ecology — Forest (disturbed) on alluvial flatland; 10-900 m altitude;
fl. Jan.—Sept.; fr. June—July.

Note — Subsp. celebica is close to the two other subspecies. It has larger fruits than
subsp. simiarum and from subsp. calcarea it differs in the less densely pubescent leaves.

124. Myristica simulans W. J. de Wilde

Myristica simulans W. J. de Wilde, Blumea 40 (1995) 325. — Type: Brass 28894, E Papua New
Guinea.

Tree 15 m. Twigs 3 mm diameter, rather late glabrescent, hairs shaggy, 1 mm long;
older twigs not flaking, with many small lenticels. Leaves chartaceous or thinly coriaceous,
(ovate-)oblong, 15-22 by 6-8.5 cm, base short-cuneate or broadly rounded, apex acute-
acuminate, upper surface olivaceous, glabrescent, indumentum minute, greyish, lower
surface pale greyish brown, persistent or late glabrescent, hairs interwoven, scale-like,
(brown-)grey, 0.1 mm; not papillose; dots absent; midrib slightly raised above, lateral
nerves 16-20 per side, at 60—70° to the midrib, lines of interarching and venation faint;
petiole 10-15 by 2.5-3.5 mm, late glabrescent, hairs shaggy, 1 mm, leaf bud 15 by 4
mm, hairs dense, woolly, rufous, 1(—1.5) mm. Inflorescences between the leaves, of the
Knema-type: in male a sessile or to 2 mm pedunculate, scar-covered brachyblast 2-5
mm, with (1 or) 2-10 flowers, buds almost equal in size, all parts late glabrescent or
hairs shaggy, rufous, 1 mm. Male flowers: pedicel 4 by | mm, bracteole rounded, | mm,
caducous, (sub)apical; buds ellipsoid-oblong, 8(—10) by 4.5(—5) mm, apex blunt, cleft
c. 1/4, lobes 2 mm long, 0.3 mm thick. Androecium 7 mm; androphore 2 by | mm, gla-
brous; synandrium 5 by 1.5 mm, thecae 14-16, sterile apex broadly rounded, 0.2—0.3
mm long. Female flowers and fruits not seen.

Field-notes — Leaves below at first brown, later grey. Flowers fragrant.

Distribution — Malesia: E Papua New Guinea (Milne Bay Prov., Modewa Bay, Gara
River); known only from the type.

Habitat & Ecology — Riverine rain forest; sealevel; fl. Dec.

Note — Myristica simulans was formerly confused with M. buchneriana that differs
in shorter hairs on twig apex and inflorescences, smaller male flowers (S—6 mm long)
with shorter hairs, and weaker hairs on the lower leaf surface. In appearance and male
flowers the species also looks like M. inopinata, but in that species the synandrium has
a distinct, acute, sterile apex, and the androphore a collar of dense hairs at the base.

125. Myristica sinclairii W. J. de Wilde

Myristica sinclairii W.J. de Wilde, Blumea 40 (1995) 327, f. 7. —Type: NGF 44076 (Kairo), Papua
New Guinea.

Tree 4-5 m. Twigs 1-1.5(—2) mm diameter, glabrescent, hairs shaggy, 1 mm long;
older twigs neither fissuring nor flaking; lenticels not apparent. Leaves (thinly) charta-
ceous, (elliptic-)oblong, 6-12 by 2—4 cm, base attenuate, apex ( acute-)acuminate; upper

De Wilde — Myristicaceae (Myristica) 571

Fig. 87. Myristica sinclairii W.J. de Wilde. a. Leafy twig with male inflorescences; b. (sub)mature
male bud shortly before anthesis; c. ditto, opened, showing androecium, note persistent bracteole;
d. androecium taken from flower at anthesis; note largely pubescent androphore which is longer
than synandrium, and conspicuous sterile apex; e. apex of female flowering twig (flowers submature);
f. branched twig with submature fruit; g. mature fruit [a-d: NGF 44076 (Kairo); e, g: NGF 29302
(Henty & Vandenberg); f: Stauffer, Whitemore & Pennington 5552). — Scale bar for a, e—g = 2 cm;
b—d = 1.65 mm.

Se. Flora Malesiana, Ser. I, Vol. 14 (2000)

surface olivaceous or brown, lower surface yellowish brown by persistent fine felty hairs
mixed with appressed stouter hairs 0.5 mm, partially late glabrescent leaving a grey
undersurface; not papillose; dots absent; midrib flat above; lateral nerves 10-15 per
side, at 45(—60)° to the midrib, flat and indistinct above, lines of interarching and vena-
tion indistinct on both surfaces; petiole 0.6—-15 by 1-2 mm; leaf bud 8-13 by (1.5—)2-2.5
mm, hairs conspicuous, | mm long. Inflorescences between the leaves, axillary or to 1.5
mm supra-axillary, + rusty pubescent, mostly to 3 mm pedunculate, almost sessile, end-
ing in a simple or forked scar-covered brachyblast to 5 mm long; in male: with clusters
of 3—5 flowers, buds of various sizes; bracts 1 mm or less, pubescent, caducous; in
female (from infructescences): peduncle short, |—3-flowered; flowers with scurfy hairs
0.1-0.4 mm. Male flowers: pedicel often + curved, 1.5—3(—3.5) mm long, bracteole ovate,
(0.5—)1-1.5 mm long, persistent or late caducous; buds (ovoid-)oblong, 7-8 by 1.5—2
(—2.5) mm, narrowed from the middle to the apex, base broadly rounded, cleft c. 1/7,
lobes membranous, 0.7—1 mm, c. 0.3 mm thick, perianth inside smooth or in the lower
half finely warty-bullate. Androecium slender, 6—6.5 mm; androphore 3—4 by 0.4—0.5
mm, hairs dense, 0.5—0.8 mm; synandrium 2-3.5 by 0.5—0.6 mm, thecae 8-10(-12),
sterile apex + slender, + acute, 0.6 mm, reaching the base of the perianth lobes. Female
flowers (from perianth remnants under immature fruit, Stauffer c. s. 5552): pedicel 3 mm
long; buds 6 mm, lobes | mm long. Fruits solitary, ellipsoid, 2.5 by 2 cm, hairs dense,
rusty, scurfy, 0.2—0.4 mm, stouter (0.5 mm) towards the base and apex; pericarp 1—1.5
mm thick, seeds not seen (probably 1.5—2 cm); fruiting pedicel slender, 5 mm long. —
Fig. 87.

Field-notes — Slender tree, branches horizontal, exudate from twigs colourless; older
twigs smooth, brown; exudate reddish; wood pinkish straw. Leaves yellowish brownish,
or + glaucous below (with thin brownish indumentum). Flowers yellowish or brownish.
Fruits with bright (yellow-)brown or orange indumentum.

Distribution — Malesia: Papua New Guinea (endemic to Morobe Prov.: Bulolo).

Habitat & Ecology — Understorey tree in Castanopsis dominated (ridge) forest; alti-
tude 1000-1600 m; fl. Jan., Aug.; fr. Jan.—Feb.

126. Myristica smythiesii J. Sinclair

Myristica smythiesii J. Sinclair, Gard. Bull. Sing. 23 (1968) 316, f. 42; W.J. de Wilde, Blumea 42
(1997) 185: Tree Fl. Sabah & Sarawak 3 (2000) 470. — Type: SAN 17440 (Ashton, Smythies &
Wood), Sabah.

Tree 8-25 m. Twigs 2—-3(—4) mm diameter, glabrescent, hairs 0.30.5 mm; older twigs
dark brown or blackish, finely longitudinally cracking, sometimes thinly flaking; lenticels
absent. Leaves (thinly) coriaceous, elliptic-oblong or oblong-lanceolate, 8-20 by 2-6
cm, base rounded or short-acute, apex acute(-acuminate) (sometimes + blunt); upper
surface olivaceous, late glabrescent on midrib towards base, lower surface with persist-
ent indumentum, hairs grey-brown or cinnamon, interwoven, scale-like, 0.1—0.2 mm,
mixed with scattered dendroid hairs 0.3—0.4 mm; not obviously papillose; dots absent;
midrib little raised above, lateral nerves 12—15(—18) per side, at 60—70° to the midrib,
faint, flat or sunken, lines of interarching and venation faint or invisible on both sur-
faces; petiole 7-15 by 1.5-2.5 mm; leaf bud 10-15 by 2.5-3 mm, hairs coarse, 0.5(-1)
mm. Inflorescences between the leaves or below, with brown-red hairs 0.5(—1) mm, of

De Wilde — Myristicaceae (Myristica) 313

the Knema-type: sessile or to 2 mm long pedunculate, single or paired brachyblasts,
bracts subpersistent, c. 1.5 mm long; in male: to 10 mm long, at apex with some flowers,
buds of various sizes; in female 2~3 mm long, with 1—4 flowers; flowers with hairs 0.3—
0.5(-1) mm. Male flowers: pedicel (2—)3 by 1 mm, bracteole subcircular, entire or faintly
3-topped, 1.5 by 2 mm, subpersistent; buds + ovoid to ellipsoid-oblong, 4-5 by 2-2.5
mm, cleft 1/4—1/2, lobes 0.3 mm thick. Androecium 3.5 mm; androphore c. | mm, to-
wards base with dense hairs 0.3—0.5 mm; synandrium 2.5 by 0.6 mm, thecae 10-12,
sterile apex blunt, 0.2 mm long, glabrous or with few minute hairs 0.1 mm. Female
flowers: pedicel c. 1 mm long, bracteole minute, buds ovoid, c. 4 mm, ovary ovoid, in-
cluding short stigma 2.5 mm long, minutely pubescent. Fruits solitary, almost sessile,
broadly ellipsoid, 2.5-3.5(-3.8) by 2—2.4 cm (remnant of the style + acute, glabrous,
blackish brown, 0.5 mm long), hairs mealy, 0.2-0.5 mm; pericarp 1-2 mm thick; seeds
(broad-)ellipsoid, 2.5 cm, red-brown, glossy; fruiting pedicel rather stout, 3-6 mm long.

Field-notes — Medium-sized tree, sometimes with small buttresses or stilt-roots. Bark
smooth, with fine longitudinal striation, narrowly fissured, or flaky, dark grey or brown-
black; outer bark thin; inner bark soft, red-brown; sapwood soft, orange-yellow; heart-
wood deep red. Flowers brown, cream inside. Fruits pale yellow-green, brown hairy.

Distribution — Malesia: Borneo (Brunei; NE Sarawak, 2nd Div.; C & E Kaliman-
tan).

Habitat & Ecology — Primary mixed dipterocarp forest, riverbank forest, hill slopes
and heath forest, peat swamp forest; on yellow sandy clay or loam soil, or shallow peat
overlying clay; 0-700 m altitude: fl. & fr. Apr.—July, Oct.

Note — Myristica smythiesii is close to M. beccarii and M. cinnamomea, all three
species with an indumentum of (pale) cinnamon scale-like hairs on the lower leaf sur-
face. Myristica beccarii differs in its usually rounded or emarginate leaf base, coriaceous
leaves with stronger and more distinct nerves, and somewhat stouter twigs: M. cinna-
momea is distinguishable by its longer petioles, a branched-type inflorescence with a
peduncle, angular perianth buds, and larger fruits with thick pericarp.

127. Myristica sogeriensis W. J. de Wilde

Myristica sogeriensis W. J. de Wilde, Blumea 40 (1995) 328, f. 3h, h’. — Type: Forbes 396, E Papua
New Guinea , Central Prov., Sogeri region.

Shrub or treelet, 2-3 m tall. Twigs 0.5—1(—1.5) mm diameter, bluntly angular, early
glabrescent, hairs scale-like, less than 0.1 mm; older twigs neither cracking nor flaking:
lenticels inconspicuous or absent. Leaves membranous or thinly chartaceous, (elliptic-)
oblong, 5—12 by 1.5—4 cm, base attenuate, apex acute-acuminate: upper surface olivaceous
or brown, lower surface brownish grey (discolorous), seemingly glabrous, but glabrescent,
hairs sparse, less than 0.1 mm; not papillose, dots absent; midrib raised above, lateral
nerves 10-15 per side, at 45-80° to the midrib, flat or sunken above, bright brown and
contrasting in colour below, lines of interarching and venation distinct: petiole 6-10 by
(0.5—)1(-1.5) mm; leaf bud 5-8 by 0.5(—1) mm, hairs less than 0.1 mm. /nflorescences
between the leaves, + of the Knema-type; in male: peduncle slender, 1-2 mm long with
small, slender, simple or forked scar-covered, glabrescent brachyblasts 3-4 mm long,
ending with 1-3 flowers, buds about equal in size; female: a sessile or to 1 mm peduncu-

574 Flora Malesiana, Ser. I, Vol. 14 (2000)

late minute wart-like brachyblast, 1—-3-flowered; flowers with + sparse hairs less than
0.1 mm, looking as if nearly glabrous. Male flowers: pedicel 6(—7) by 0.3—0.4 mm, brac-
teole | mm long, persistent, 1-2 mm below the apex; buds tubiform, 6.5 by 1(—1.5) mm,
apex subacute, the basal half somewhat narrowed, cleft c. 1/6, lobes | mm long, 0.2 mm
thick, perianth inside at base faintly warty-ridged. Androecium slender, 6 mm; androphore
3.5 by 0.3 mm, with minute hairs in the lower half; synandrium 2(—2.2) by 0.5 mm,
thecae c. 10, sterile apex subacute, 0.5—0.6 mm. Female flowers not seen. Fruits 1—3 per
infructescence, + fusiform, (3—)3.5 by 1.5 cm, apex acute, basal part tapered into a 5-8
mm long pseudostalk, hairs minute, dull-rusty, mealy, 0.1 mm or less; pericarp 1—1.5
mm thick; seeds ellipsoid, 1.4 cm; fruiting pedicel 3-4 by 2 mm, faintly fissured and
lenticellate, bracteole scar about median. — Fig. 63h.

Distribution — Malesia: E Papua New Guinea (endemic to the Sogeri region, Central
Prov.); known from two collections.

Habitat & Ecology — Foothill forest; c. 400 m altitude; fl. Mar.

Notes — 1. Resembling and related to M. pumila, a species occurring in the same
area and with larger and longer pedicelled male flowers with a caducous bracteole,
(sub)sessile male inflorescences, and fruits borne on a 20-25 mm long thread-like pedi-
cel (much shorter in the present species). Recent collecting has shown that the Sogeri
region harbours several local endemic species, sometimes with a strikingly similar
foliage.

2. Because the (male) inflorescences are short-pedunculate, M. sogeriensis has been
entered twice in the keys, in the group with sessile inflorescences and in that with stalked
inflorescences.

128. Myristica sphaerosperma A.C. Sm.

Myristica sphaerosperma A.C. Sm., J. Arnold Arbor. 22 (1941) 71; J. Sinclair, Gard. Bull. Sing. 23
(1968) 247, f. 23; Foreman, Contr. Herb. Austral. no. 9 (1974) 42, p.p.; Handb. Fl. Papua New
Guinea 1 (1978) 210, p.p.; W.J. de Wilde, Blumea 40 (1995) 330. — Type: Brass 4174, Papua
New Guinea.

Tree 10-30 m. Twigs subterete or bluntly angled, (2—)3—5 mm diameter, early or late
glabrescent, hairs powdery, 0.5 mm long; older twigs sometimes longitudinally crack-
ing, not flaking; inconspicuously lenticellate. Leaves chartaceous or coriaceous, ovate-
elliptic to elliptic(-oblong), rarely oblong, 10—25(—32) by 4—9(-10.5) cm, base (short-)
cuneate or (broadly) rounded, apex acute-acuminate, upper surface olivaceous or brown,
lower surface grey-brown, shaded by cinnamon (sub)persistent indumentum, hairs in-
terwoven or widely spaced, scale-like, grey-brown, 0.1—0.2 mm, mixed with scattered
+ dendroid brown hairs to 1 mm; irregularly papillose (appearing non-papillose); dots
absent: midrib + flat above, at (60—)70—80(—90)° to the midrib, lateral nerves (10—)15—
25 per side, flat or sunken, lines of interarching distinct, venation barely visible; petiole
15-25 by 2.5-4 mm; leaf bud 20 by 3-4 mm, hairs rusty 0.5(-1) mm long. Jnflores-
cences between the leaves, + of the Knema-type: a simple, scar-covered brachyblast
2-3 mm long, sessile or to 5 mm pedunculate, late glabrescent; in male (Clemens 4527):
with a cluster of 5 or 6 flowers, buds of various sizes; female inflorescences similar to
male, 1—3-flowered; flowers with hairs 0.5—1 mm. Male flowers: pedicel 8-10 by 11.5
mm, bracteole 2 mm, caducous; buds oblong-ellipsoid, 10-11 by 3-4 mm, cleft (1/5—)

De Wilde — Myristicaceae (Myristica) Sis

1/4. Androecium 6 mm long; androphore 2 mm, finely brown pubescent at base; synan-
drium 4 mm long, thecae 8—12(?), sterile apex minute. Female flowers: pedicel 5—10
by 2 mm; buds ovoid-oblong, 10 by 5 mm, apex subobtuse; ovary ovoid-oblong, 4-6 by
3-4 mm, with stiff hairs (0.5—)1 mm long, stigma 2 mm. Fruits single, subglobose or
broadly (ob)ovoid, sometimes faintly beaked, or somewhat narrowed towards the base,
6-8 by 5-6.5 cm, hairs dense, mealy, rusty, 0.5(—-1) mm long; pericarp 5—12 mm thick;
seeds subglobose, 3-3.5 cm diameter; fruiting pedicel + stout, 7-15 by 5—10 mm, late
glabrescent, + fissured or flaking when old.

Field-notes — Tall tree with horizontal branches. Bark grey-brown or black, not or
little fissured or peeling; slash cream; sap red, or clear turning brown on exposure; wood
creamy-white. Flowers yellow or brown, inside cream. Leaves greyish below. Fruits
8—9 by 6 cm, brown or brownish green. (Immature) fruits large, pointed at both ends, aril
white.

Distribution — Malesia: New Guinea (Papua Barat: Bird’s Head, Japen I., Wandammen
Peninsula, Idenburg R., Cycloop Mts; Papua New Guinea: Morobe Prov.).

Habitat & Ecology — Mountain forest of ridges and slopes; midmontane forest, oak
forest; clay-soil; (300—)800-1600(-1800?) m altitude; fl. & fr. throughout the year.
Locally common.

Notes — 1. Myristica sphaerosperma is related to M. womersleyi which is distin-
guishable by the longer indumentum on the lower leaf surface and fruits with a shorter
stalk. Another related species is M. mediovibex, in the key placed far apart because of its
smaller leaves with short indumentum below without emergent hairs, and its smaller
fruits.

2. Some specimens have only a thin indumentum on the lower leaf surface, composed
of scattered scale-like hairs; old leaves may become quite glabrous. Schram BW 13492
(Wandammen Peninsula) has narrower and thinner leaves, and was collected at a rather
low altitude for the species, c. 300 m.

129. Myristica subalulata Miq.

Myristica subalulata Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 47. — Type: Zippelius s.n., West
New Guinea.
For more references and synonyms see under the varieties.

Tree 3—10(—25) m. Twigs + flattened, with two raised lines or wings to 1(—2) mm high
between the petioles, usually hollow and with ant-swellings (these faint or absent in var.
hagensis), (3—)4—8(-10) mm diameter, glabrescent, hairs 0.1 mm or less; older twigs
sometimes shallowly fissured, lenticellate. Leaves membranous, chartaceous or (sub)-
coriaceous, elliptic(-oblong), 15-40 by 5—13(—16) cm, base cuneate or (narrowly) rounded,
sometimes shallowly cordate, apex acute-acuminate; upper surface olivaceous or brown,
lower surface grey-brown, glabrous or pubescent; neatly or irregularly papillose or not
papillose; dots absent; midrib flat above, lateral nerves (13—)17-25 per side, at 50—80°
to the midrib, flat or sunken, lines of interarching and venation distinct or not; petiole
(10—)15—25 by (2-)3—5 mm; leaf bud 10-15(—20) by 2-3 mm, hairs 0.1(—0.2) mm. /n-
florescences between the leaves or rarely below, of the Knema-type: a sessile or to 4mm
pedunculate (or S~20 mm in var. pedunculata), woody, wart- or worm-like, scar-cover-
ed brachyblast, 10(—20) by 5 mm, sometimes 2- or 3-armed, pubescent or glabrescent,

576 Flora Malesiana, Ser. I, Vol. 14 (2000)

Fig. 88. Myristica subalulata Miq. var. subalulata. Male twig with immature flowers (bracteoles
partly still present), ant-swellings, and ant-openings; stem partly opened to show ant-cavities with
two coccids [Polak MP 750]. — Scale bar = 2 cm.

De Wilde — Myristicaceae (Myristica) 577

ending in few- or many-flowered subumbels (in male to 20-flowered, in female 1—3-
flowered), the buds either + equal in size or of various sizes; flowers with hairs 0.1—0.2
mm, sometimes (sub)glabrous. Male flowers: pedicel 8-16 by (0.6—)1 mm, bracteole
sometimes + 3-topped, 2(-3) mm, persistent or caducous, (sub)apical; buds (mostly)
membranous, (ellipsoid-)oblong or + tubiform, 8—15 by (2—)2.5—5 mm, apex + obtuse or
subacute, not or faintly angled, cleft 1/6 or less, lobes 1.5—2 mm long, 0.2—0.5 mm thick.
Androecium slender or + stout, 7-12 mm; androphore 2—6 by 0.4—1 mm, subglabrous or
variable with scattered hairs less than 0.1 mm, either only at the base or in the lower half;
synandrium (4—)5—7 by 0.5—1.5 mm, tapering into acute sterile apex 0.5—1 mm long,
thecae 8-24. Female flowers: pedicel 1-5(—7) mm, bracteole 2—3 mm, (sub)apical; buds
ovoid-oblong, narrowed to the apex, 4—7(—8) by 2.5-3.5 mm, lobes 1—1.5 mm; ovary
narrowed to the apex, 3-3.5 mm long, minutely pubescent, stigma 1.5(—2) mm. Fruits
solitary or in clusters of up to 6, subglobose, ellipsoid or broadly ovoid, 1.5—2
(—2.5) by 1-1.5 cm, narrowed to the apex, with the style-remnant remaining as a short
apiculum, (fruits 1-3 per cluster, 2.5-3 cm long, with pedicel to 10 mm long in var.
paucifructa), hairs persistent, mealy, rusty, 0.1—0.3(—0.4) mm, rarely glabrescent, some-
times lenticels in the basal part; pericarp 2(—3) mm thick; seeds (broadly) ovoid or ellip-
soid, 1.5(—2) cm; fruiting pedicel 2-5(—10) by 1—2(—2.5) mm, generally with conspicu-
ous raised lenticels. — Fig. 88.

Distribution — Malesia: Moluccas (Aru and Kai Islands), throughout New Guinea
(incl. Misool; excl. Bismarck Archipelago). There are five varieties.

Habitat & Ecology — Altitude 0—2000 m.

Note — Widely distributed, with a large altitudinal range, characterized by the (sub)-
alulate twigs mostly with ant-swellings. Much of the variation, e.g. in the leaf texture,
presence or absence of papillation on the lower leaf surface, stoutness, and length of
flower pedicel, seems gradual, and the subdivision of this species into varieties 1s arbi-

trary.
KEY TO THE VARIETIES

la. Twigs with faint swellings or without; ridges faint or fairly distinct. Leaves and
flowers coriaceous. — Papua New Guinea (Western Highlands Prov.: Mt Hagen and
paciniby ac: 2 000.401) \erien, titsosak inte Ae eee b. var. hagensis

b. Twigs with ant-swellings, generally with an opening ..................---. 2
2a. Male buds not particularly narrow, 8-14 by (3—)4—5 mm; bracteole generally cadu-
cous; androphore about half as long as the synandrium; anthers 7-12. — Lowland to
POOO muvaltitude’.aices.celd ence Sed We Ek SEOs Bea cee ear 3

b. Male buds slender, tubiform, 12—15 by (2—)2.5—3.5 mm; bracteole persistent or cadu-
cous; androecium slender, androphore (much) longer than half the length of the synan-
drium; anthers 4—6. [Fruits not known, possibly 1.5—2 cm long.] — Lowland and

lowenmmontane areas: mila nod. Seek ORs ot Ro A Ee eee 4

3a. Fruits 2.5-3 cm long. [Flowers not known with certainty, possibly similar to type
sAnety.||—> Montane areas. |e 2s: See. 3 BE Ee d. var. paucifructa

b. Fruits 1.5—2(—2.5) cm long. — Lowland and montane areas ... a. var. subalulata
AggMaleintlorescencéssessilep wc J See ee c. var. leptantha

b. Male inflorescences 5-20 mm long pedunculate .......... e. var. pedunculata

578 Flora Malesiana, Ser. I, Vol. 14 (2000)

a. var. subalulata

Myristica subalulata Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 47; Warb., Mon. Myrist. (1897)
484, t. 19; Markgr., J. Arnold Arbor. 10 (1929) 214; Bot. Jahrb. Syst. 67 (1935) 163; J. Sinclair,
Gard. Bull. Sing. 23 (1968) 385, f. 64, 65; Foreman, Handb. Fl. Papua New Guinea | (1978)
210: W.J. de Wilde, Blumea 35 (1990) 255, f. 2: 14; 40 (1995) 330; 43 (1998) 176, f. 1.

Myristica macrophylla Zipp. ex Miq., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 47, nom. nud.

Myristica myrmecophila Becc., Malesia 2 (1884) 37, t. 1. — Syntypes: several Beccari numbers
(FI), West New Guinea.

Myristica heterophylla K. Schum. in K. Schum. & Hollr., Fl. Kaiser Wilhelmsland (1889) 45, p.p..
nom. inval. [non Myristica heterophylla Fern.-Vill. (1880) = Knema glomerata}.

Doubtful synonym: Myristica costata Warb., Bot. Jahrb. Syst. 18 (1893) 191; Mon. Myrist. (1897)
A87. t. 19: K. Schum. & Lauterb., Fl. Schutzgeb. Siidsee (1900) 327; Markgr., Bot. Jahrb. Syst.
67 (1935) 166. — Type: Hellwig 247 (B, lost), Sattelberg near Finschhafen.

Tree 3-10(—25) m. Twigs as those of the species, perforated ant-swellings always
present. Leaves membranous, chartaceous, or (sub)coriaceous, lower surface distinctly
papillose (lens!), or not. Flower buds of various sizes within one inflorescence, bracteole
sometimes persistent (see note). Male flowers: buds 8-14 by (3—)4—5 mm. Androecium
7-10 mm; androphore shorter than the synandrium, 2—2.5 mm, subglabrous or minutely
pubescent to various degrees; synandrium (3.5—)5—6 by 1—1.5 mm, thecae 14-24, sterile
apex ().5—1 mm long. Fruits solitary or in clusters of up to 6, subglobose or ovoid 1.5-
2(-2.5) by 1-1.5 cm, apex narrowed or rounded, style-remnant persistent as a 1-3 mm
long beak; fruiting pedicel 2-9 mm long. — Fig. 59d, e, 61a, 65e, 66d, 88.

Field-notes — Shrub or small erect tree of lowest or middle layer of the forest, rarely
with stilt-roots, branches sometimes tiered, horizontal, often drooping. Bark (dark) brown,
smooth or finely vertically fissured, or finely tesselated, sometimes with pustulate lenticels
or circular raised ridges; inner bark orange-brown or salmon; exudate clear, sometimes
resinous, turning (pink-)brown on exposure; wood straw or cream, turning (salmon-)
brown, often pinkish mottled by exudate (vessels standing out pinkish against a white
background). Leaves grey or glaucous below. Flowers succulent, cream or yellow(-brown),
cream inside. Fruits either brown, orange- or yellow-brown, or (pale) chestnut.

Distribution — Malesia: as the species.

Habitat & Ecology — Lowland, (sub)montane forest on flatland, slopes, and ridges;
in riverside forest, bamboo regrowth, and Araucaria- and Nothofagus—Castanopsis-domi-
nated forest: also in secondary regrowth; on clayey soils, soils over limestone, volcanic
and peaty soils; most commonly on well-drained soil, occasionally in temporarily inun-
dated or swampy locations; 0—2100 m altitude; fl. & fr. throughout the year.

Characteristically the twigs are hollow and have swollen parts with small round or
slit-like perforations, the swellings inhabited by small black ants. Occasionally olive-
coloured swollen galls are found on the top of the twigs.

Note — Myristica subalulata var. subalulata comprises most of the material of the
common species M. subalulata. The type variety is variable in itself: — Twigs with at
every 4 or 5 internodes a swollen hollow part, with a characteristic hole or a rough-lined
slit, inhabited by small black ants. If ant-swellings are lacking in a specimen, this iS pos-
sibly due to incomplete sampling of the material. The swellings are presumably initiated
and grown by the plant and will be subsequently occupied by the ants (see De Wilde,
1998). — Leaves. The texture of the leaves is variable according to the provenance,

De Wilde — Myristicaceae (Myristica) 579

(sub)coriaceous in specimens from montane areas. The lower leaf surface is either strong-
ly and regularly, or irregularly whitish papillose, or but faintly or not papillose (lens!).
Myristica subalulata is one of the few species in which this character is not constant. In
non-papillose leaves the papillae may not have been developed (or only insufficiently
so); conversely, the papillae may be present so profusely and densely that they entirely
cover the lower leaf surface so that this appears to be non-papillose. Distinct regular
papillae are most apparent in lowland specimens; irregular papillation is common in
lowland and montane material, while leaves that are non-papillose with a smooth sur-
face predominate in specimens from montane and upper montane regions. — Indumen-
tum. The indumentum is inconspicuous, and can be seen only with magnification. De-
veloping leaves have sparse or dense, scale-like, pale brown or greyish hairs, 0.1(—0.2)
mm or less. They are early shed, especially in lowland specimens. In collections from
montane areas the indumentum is densest and often longer persistent as well. — Flow-
ers. The pedicels of male flowers, but especially those of female flowers and fruits, tend
to be longer towards the east of the distributional area. The flowers of part of the male
specimens of an eastern distribution, i.e. Papua New Guinea, are more slender, with
smaller perianth and longer pedicels, but this feature seems to intergrade. Plants with
conspicuous slender male perianths belong to var. /eptantha. — Bracteole. In most speci-
mens the bracteole is early falling; in a few collections, linking up with the bulk of the
material of var. subalulata, the bracteole is persistent or late caducous (e.g. NGF 13207,
48143, Docters van Leeuwen 9181, 11297, from distant localities, in lowland as well as
montane areas). — Fruits. Shapes and sizes are variable, but the fruits are always small,
1.5-2(-2.5) cm long. Subglobose, ovoid, and (ovoid-)oblong fruits occur; ovoid fruits,
narrowed to the apex and ending in the short beak formed by the style-remnant are most
frequent. The fruiting pedicel may be short or long, to 7 mm, mostly slender, but may
become quite thick, caused by coarse lenticel-like outgrowths. Specimens with fruits
longer than 2.5 cm belong to var. paucifructa.

b. var. hagensis W. J. de Wilde

Myristica subalulata Miq. var. hagensis W.J. de Wilde, Blumea 40 (1995) 330; 43 (1998) 177, 181.
— Type: Hoogland & Pullen 5831, Papua New Guinea.

Tree 8-15 m. Twigs 5—7 mm diameter, ridged or not, always + angular; ant-swellings
faint and without opening, or absent. Leaves (thinly) coriaceous, comparatively small,
the largest 24 by 7 cm, lower surface subglabrous, not papillose. Inflorescences: brachy-
blast thick, wart-like, to 10 mm long; flower buds rather robust, resembling those of the
type variety, slightly differing in size within one cluster, bracteole 2(—4) mm, caducous,
perianth leathery. Male flowers as the species. Fruits solitary or 2 (or 3) together, broadly
ellipsoid, 2 by 1.5 cm, apex pointed; fruiting pedicel 5—7 by 2(-3) mm, smooth, not or
but little lenticellate.

Field-notes — Bark 3—5 mm thick, dark brown or dark purple-brown, with numerous
longitudinal fissures; sap red; underbark purple-brown; inner bark pale brown; blaze
straw; wood pink or straw with orange dots, or pale brown, easy to cut. Flowers hard,
cream or olivaceous. Fruits reddish brown or light yellow tinged with brown.

Distribution — Malesia: Papua New Guinea (Western Highlands Prov.: Mt Hagen
and vicinity).

580 Flora Malesiana, Ser. I, Vol. 14 (2000)

Habitat & Ecology — High mountain forest, Nothofagus- or Castanopsis-forest, 1800—
2150 m altitude; fl. & fr. May, July, Aug.

Notes — 1. The status of var. hagensis is provisional. It is close to typical M. subalulata
as shown by the male flowers. All specimens (restricted to Mt Hagen and vicinity) are
distinguishable by the absence of perforated ant-swellings in the twigs and the coriaceous,
rather small leaves. Possibly the ant species belonging to M. subalulata does not occur
at the high altitudes of the present variety.

2. The lower leaf surface which appears to be non-papillose, may be so strongly and
densely papillose that the separate papillae are difficult to discern as such.

c. var. leptantha W. J. de Wilde

Myristica subalulata Miq. var. leptantha W.J. de Wilde, Blumea 40 (1995) 331; 43 (1998) 177. —
Type: van Royen & Sleumer 7747, Irian Jaya, Bird’s Head.

Tree 5-7 m. Twigs as those of the species. Leaves membranous or chartaceous, the
lower surface distinctly papillose or not. Inflorescences sessile, without peduncle; flower
buds slender, of various sizes (according to age) within one inflorescence, bracteole per-
sistent or caducous. Male flowers: buds lanceolate-tubiform, 12—15 by 2.5—3(—3.5) mm,
apex narrowed, + blunt, lobes 1-1.5 mm long. Androecium slender, 10-12 mm, andro-
phore 4—5 by 0.4—0.6(—0.8) mm, almost completely pubescent, hairs sparse, pale, 0.1
mm or less; synandrium 4—6 by 0.5—1 mm, thecae 8-12, contiguous or rather spaced,
sterile apex slender, | mm long. Female flowers and fruits not known.

Field-notes — Slender tree with few, short, excurrent branches. Leaves slightly glau-
cous below. Flowers cream or white, brown suffused.

Distribution — Malesia: New Guinea; known from three male-flowering collections
of widely scattered localities in Papua Barat (Bird’s Head) and Papua New Guinea (West-
ern Provy., Palmer River, and Sepik Prov.).

Habitat & Ecology — Forest of mountain slope, forest undergrowth on lower ridge,
or on steep riverbank; 100—840 m altitude; fl. July, Nov.

Note — Characterized by slender male flowers. The lower leaf surface may be papil-
lose or not. In Darbyshire & Hoogland 8297 the bracteole is persistent in mature (but
not yet open) flowers.

d. var. paucifructa W. J. de Wilde

Myristica subalulata Mig. var. paucifructa W.J. de Wilde, Blumea 40 (1995) 332, 43 (1998) 177.
— Type: BW 1144] (Vink), West New Guinea.

Tree 5-15 m. Twigs as those of the species. Leaves membranous or subchartaceous,
the lower surface glabrous or somewhat pubescent, not or but indistinctly papillose.
Inflorescences: flower buds somewhat variable in size within one inflorescence, bracte-
ole caducous. Male flowers not known. Fruits 1-3 per infructescence, (ovoid-)ellipsoid,
2.5—3 by 1.7—2 cm; fruiting pedicel 3-9 mm long.

Field-notes — Slender trees, twigs hollow, with many small black ants producing an
unpleasant smell. Bark smooth or vertically cracked; exudate red; inner bark (blaze)
chestnut or reddish; wood dark cream. Leaves glaucous or coppery green below. Fruits
brown, seeds dark brown.

De Wilde — Myristicaceae (Myristica) 581

Distribution — Malesia: Moluccas (Kai Is.); widely distributed in Papua Barat and
Papua New Guinea, including the Papuan Islands.

Habitat & Ecology — Mountain forest of gullies, slopes and ridges; 800-1200 m
altitude; fl. & fr. throughout the year. Locally common.

Note — The variety paucifructa accommodates specimens with few fruits per in-
fructescence, the fruits being large in comparison to those of the type variety. Vegetatively
and in flowers the specimens of the present var. paucifructa merge with those of var.
subalulata. Possibly var. paucifructa has developed polytopically. All specimens are
from montane areas.

e. var. pedunculata W. J. de Wilde

Myristica subalulata Migq. var. pedunculata W.J. de Wilde, Blumea 40 (1995) 332, f. 2b; 43 (1998)
177. — Type: Sands et al. 1483, Papua New Guinea.

Tree 4 m. Twigs as those of the species. Leaves thinly chartaceous, lower surface not
papillose. Inflorescences worm-like, the scar-covered part unbranched, to 4 cm long,
peduncle glabrous (glabrescent), somewhat flattened, 5-20 mm long; flower buds slen-
der, of various sizes within one inflorescence, bracteole + 3-topped, 3 mm long, caducous.
Male flowers: pedicel 11 by 0.5 mm; buds tubiform, 15 by 2—2.5 mm, apex + acute,
somewhat angled, lobes 1 mm long. Androecium slender, 12 mm; androphore 6 by 0.4
mm, almost completely pubescent, hairs sparse, pale, less than 0.1 mm; synandrium 5
(—6) by 0.7 mm, thecae c. 12, sterile apex 0.3 mm long. Female flowers and fruits not
seen. — Fig. 66e.

Field-notes — Bark brown; young wood olive-green. Leaves dark green above, paler
below. Inflorescence stiff, pointing downwards from the branches, rachis swollen, cof-
fee-brown; flower buds butter-cream, on pale green pedicel.

Distribution — Malesia: Papua New Guinea (Gulf Prov., Kikori subprov., c. 200 km
upstream Purari River); known only from the type.

Habitat & Ecology — In lowland rain forest including Terminalia, Elaeocarpus,
Albizia, and Pometia; growing near small tributary stream; 140 m altitude; fl. Aug.

Note — The present var. pedunculata is noticeable for the pedunculate inflorescences,
a character mostly used on species level. In this character var. pedunculata somewhat re-
sembles M. cornutiflora, a species also with long tubiform male flowers.

130. Myristica subcordata Blume

Myristica subcordata Blume, Rumphia 1 (1837) 186; W.J. de Wilde, Blumea 40 (1995) 333. —
Type: Zippelius s.n. (4 sheets in L) (Myristica cordifolia Zipp. in herb., nom. nud., p.p.), West
New Guinea.

For more references and synonyms see the varieties.

Tree (5—)10—30 m. Twigs 3—5(—10) mm diameter, glabrescent, hairs 0.1—0.6(—1) mm,
older twigs finely or coarsely lenticellate, bark often longitudinally cracking, occasion-
ally flaking; lenticels present. Leaves membranous, chartaceous, or subcoriaceous, elliptic-
oblong to lanceolate, 12—35(—40) by 4-14 cm, base (short) cuneate, rounded, or shallowly
cordate (frequently so in var. subcordata and morindiifolia), apex acute-acuminate; up-
per surface olivaceous or brown, lower surface with persistent indumentum, hairs dense,

582 Flora Malesiana, Ser. I, Vol. 14 (2000)

somewhat powdery, (grey-)brown, 0.1—0.4(—0.6) mm; not papillose; dots absent; midrib
flat above (sometimes raised towards the apex), lateral nerves 18—30(—35) per side, at
60—70° to the midrib, in cordate leaves those towards the base frequently at nearly 90° to
the midrib, lines of interarching faint or distinct, venation indistinct; petiole 10-25 by
2.5—5 mm: leaf bud 10-30 by 2.5—5 mm, hairs 0.1—0.5(—0.6) mm; cataphyll-like leaves
often present on the twig below the normally developed leaves, especially in var. morin-
diifolia. Inflorescences between the leaves or below, of the Knema-type: a short, simple
or forked, scar-covered brachyblast (in male) up to 15 mm long, sessile or to 5 mm long
pedunculate, with short rusty indumentum, glabrescent; bracts 1.5 mm, caducous; in
male: ending in a cluster of 5-10 flowers, buds of various sizes; female similar to male,
fewer flowered; flowers with flocculose hairs 0.2-0.5(-1) mm. Male flowers: pedicel
5-12 by I-1.5 mm, bracteole + ovate (sometimes faintly 3-topped), 2—2.5 or 4—5 mm
long (see note under var. morindiifoliia), caducous, scar apical or 1-2 mm below; buds
(ovoid-)oblong or nearly tubiform, 6.5—8(—10) by 3-4 mm, cleft 1/5—1/4, lobes (1-)1.5-
2 mm long, 0.2—0.3 mm thick. Androecium slender, (4—)4.5—6.5 or 7.5-8 mm; androphore
2—4 mm long, in the lower 1/3—1/2 with hairs 0.1-0.5 mm; synandrium 2.5—4 by 0.7-1.2
mm, thecae 10-12, sterile apex acute or blunt, sometimes faintly lobed, 0.2—0.4(-1) mm
long, or absent. Female flowers: pedicel 1 mm long (longer in var. gigacarpa), bracteole
3-4 mm, caducous; buds ovoid, 4—5(—6) by 4—5 mm; ovary ovoid, 3.5—4 by 3.5—-4 mm,
hairs 0.2—0.6 mm. Fruits single or in clusters of 2 or 3(-5), (ovoid-)oblong, rarely sub-
globose, (3.5—)4—-5.5 by 2.5-3.5 cm, hairs (dark) rusty, + flocculose, 0.1-0.5(—1) mm;
pericarp leathery, sometimes + woody, 3—7(-10) mm thick; seeds ellipsoid, 2.5—3 cm;
fruiting pedicel (3—)4—10(— 20 in var. gigacarpa) by 3-5 mm, late glabrescent, bracteole
scar usually below the apex.

Field-note — Frequently with stilt-roots.

Distribution — Malesia: throughout New Guinea (incl. the Bismarck Archipelago).

Habitat & Ecology — Generally at low altitudes, but var. morindiifolia up to 1000
(—1400) m.

Notes — 1. A variable species in which tentatively four varieties are distinguished.
The type variety is upheld for those specimens which (almost) match the original type
specimens. Provisionally, most material which falls outside var. subcordata sensu stricto
is merged into the very variable var. morindiifolia. Of var. subcordata male flowers are
still unknown; those described by Sinclair (1.c.: 308) pertain to a specimen now under
M. fatua. Additional explanatory notes are given under the varieties. Myristica subcordata
varies in general habit and in details of flowers and fruits, e.g. in length of hairs of
indumentum and length of the fruiting pedicel. It has solid twigs, and no ant-inhabita-
tion. Stout specimens, especially of var. morindiifolia, may be confused with certain
myrmecophilous species. The taxonomic status of var. gigacarpa and var. rimosa is pro-
visional. Of these two varieties male flowers are lacking, and they may represent sepa-
rate species; perhaps var. rimosa may be a variety of M. verruculosa.

2. The varieties subcordata and morindiifolia were accepted by Sinclair (1.c.) as dif-
ferent varieties under his broad concept of M. fatua. From Miquel (1865) onwards, the
original two species described by Blume (1837) have been mostly regarded as particu-
larly closely related, and treated under one name, M. fatua B subcordata. In the present
treatment M. subcordata is accepted as a distinct species beside M. fatua, following
Warburg (1897: 420).

De Wilde — Myristicaceae (Myristica) 583

KEY TO THE VARIETIES

la. Pericarp 7(—10) mm thick. —Papua New Guinea; montane .... b. var. gigacarpa
b. Pericarp 3-6 mm thick. — Lowlands and lower montane areas ...........-.. 2
2a. Twigs pale brown, bark rough by numerous closely set lenticels. — Papua Barat
ME RCMEM ELC AG ese sen 25.4, are Giers 9.2 aE Syauena > kee cee eane eee d. var. rimosa

b. Twigs brown, bark with dispersed lenticels ..........--.- +--+ e ese eeeeeees 3

3a. Leaves subcoriaceous, margin revolute on drying. Indumentum of fruits with woolly
hairs 1(-1.5) mm long. Indumentum of leaf bud and innovations with hairs | mm
long. Fruiting pedicel 1-2(-3) mm. — W & S Papua Barat, SW Papua New Guinea
MPeSeieTOVs)lOWlANUS 2... ee a te ech a em cues a. var. subcordata

b. Leaves membranous or chartaceous, margin usually flat. Indumentum of fruits with
short or long hairs, 0.1—0.5(—1) mm. Indumentum of leaf bud and innovations with
short or rather long hairs, 0.1—0.5 mm. Fruiting pedicel (3—)4-10 mm long. —
Throughout New Guinea, incl. Bismarck Archipelago and Papuan Islands; lowlands,
rpm TAOO) I ee ope were eects 2 oe ee Bee c. var. morindiifolia

a. var. subcordata

Myristica subcordata Blume, Rumphia | (1837) 186; Warb., Mon. Myrist. (1897) 420, t. 19, p.p.;
W.J. de Wilde, Blumea 40 (1995) 333; 43 (1998) 181. — Myristica fatua Houtt. var. subcordata
(Blume) Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 46, p.p.; J. Sinclair, Gard. Bull. Sing. 23
(1968) 307, f. 38 A-C (excl. male fl.); Foreman, Contr. Herb. Austral. no. 9 (1974) 37; Handb.
Fl. Papua New Guinea | (1978) 196.

Myristica wallacea Warb. var. keyensis Warb., Mon. Myrist. (1897) 531. — Type: Warburg 20720
(lost).

Tree. Twigs (dark) brown; older bark with dispersed lenticels. Leaves subcoriaceous,
17-28 by 6-13 cm, base subcordate or rounded, margin revolute, the lowermost nerves
often almost rectangular to the midrib, lower surface with dense flocculose indumentum,
hairs to 1 mm long; petiole comparatively short, 10 mm long; leaf bud with hairs 1(—1.5)
mm long. Male flowers not known. Fruits + sessile, 3.5—4 cm long, (broadly) rounded at
base, hairs coarse, shaggy, |—-1.5 mm long; fruiting pedicel 1-2(—3) mm long, bracteole
scar (sub)apical.

Field-notes — Bark paper-thin, faintly peeling, fissured or not; inner bark pink or
blood-red brown; wood straw; sap red, brown on exposure. Leaves glaucous and brown
powdery below. Fruits hairy, copper brown.

Distribution — Malesia: New Guinea (SW Papua Barat: western part of Bird’s Head;
Papua New Guinea: Western Prov.).

Habitat & Ecology — Forest on flatland, level land inundated in the wet season,
forest on stony clay; 0-200(—300) m altitude; fr. Apr.—July, Nov. Locally common.

Note —BW 10768 (Ch. Koster) from Mt Krabo, S of Manokwari, at 300 m, deviates
in its chartaceous (not coriaceous) leaves, at base not cordate; the fruit, however, matches
the type. Soegeng 351 and LAE 73930 (both S New Guinea) deviate by their more spaced
lateral nerves, which are comparatively faint and not much projecting below.

b. var. gigacarpa W.J. de Wilde

Myristica subcordata Blume var gigacarpa W.J. de Wilde, Blumea 40 (1995) 333; 43 (1998) 181.
— Type: Streimann 8548, Papua New Guinea.

584 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 12-20 m. Twigs 5-10 mm diameter, brown. Leaves chartaceous or subcoriace-
ous, 28-32 by 8-12 cm, margin flat, lower surface with dense indumentum, hairs 0.1 mm
or less. Leaf bud with hairs 0.1 mm or less. Flowers not known. Fruits solitary, axillary
to lower leaves, ellipsoid, 5.5—6 by 3.5-4 cm, hairs 0.1 mm or less; pericarp + woody,
7(-10) mm thick; fruiting pedicel stout, 15-17(—20) by 6 mm, + lenticellate, bracteole
scar 6 mm below apex.

Field-notes — Straight-boled tree. Leaves brown below. Fruits rusty brown.

Distribution — Malesia: Papua New Guinea (Eastern Highland Prov.).

Habitat & Ecology — Primary and advanced regrowth forest; along creek, on slope
on limestone; 1200-1800 m altitude; fr. Aug., Sept.

Note — Possibly this variety merits the status of a separate species, but more material
(including flowers) is needed to make a final decision.

c. var. morindiifolia (Blume) W. J. de Wilde

Myristica subcordata Blume var. morindiifolia (Blume) W.J. de Wilde, Blumea 40 (1995) 334; 43
(1998) 181. — Myristica morindiifolia Blume, Rumphia 1 (1837) 186, ‘morindaefolia’. —
Myristica fatua Houtt. var. morindiifolia (Blume) J. Sinclair, Gard. Bull. Sing. 23 (1968) 286,
f. 34. —Type: Zippelius s.n. (3 sheets in L = Myristica cordifolia Zipp., in herb., nom. nud., p.p.,
remainder = var. subcordata, see the notes by Sinclair, l.c.), New Guinea.

Myristica multinervia A.C. Sm., J. Arnold Arbor. 22 (1941) 70. — Type: Brass 3914, New Guinea.

Tree. Twigs brown; older bark with dispersed lenticels. Leaves membranous or
chartaceous, margin usually flat, lower surface with persistent indumentum, hairs 0.1—
(.4(—0.6) mm; petiole 1-2.5 cm long; innovations and leaf bud with hairs 0.1-0.5 mm.
Male flowers as those of the species. Fruits single or in clusters, (ovoid-)oblong, 4—5.5
cm long, hairs to 0.1—0.5(—1) mm; pericarp 3-6 mm thick; fruiting pedicel (3—-)4-10 mm
long, bracteole scar + below the perianth.

Field-notes — Straight-boled tree with horizontal branches, crown narrow, with or
without stilt-roots or buttresses. Bark dark brown or black, smooth, or with lenticels, or
with shallow vertical fissures, or rarely with small flakes (faintly peeling); inner bark
fibrous, reddish brown; sapwood pale; sap red, sometimes sticky; no heartwood. Flow-
ers with yellowish (brown) indumentum, inside greenish; with fetid smell or sweetly
scented. Fruits rusty tomentose; seeds brown-black.

Distribution — Malesia: throughout New Guinea, including Bismarck Archipelago
and Papuan Islands.

Habitat & Ecology — (Tidal) forest of riversides, (periodically inundated) marsh for-
est (stout specimens), also dryland forest; on sandy or clayey soil, soil over limestone, or
voleanic scoria; also in strand vegetation (smaller-leaved forms); 0-500 m altitude (but
BW 13626, Wandammen Peninsula, 950 m, NGF 21540, Morobe Prov., c. 1400 m); fl. &
fr. throughout the year.

Notes — 1. Specimens intermediate with var. subcordata occur all over the area, but
especially in New Britain (specimens with rather long-haired fruits).

2. The figures in Sinclair (1968: 295, f. 37) for M. fatua var. papuana all represent
specimens now referred to M. subcordata var. morindiifolia.

3. Male plants of var. morindiifolia from western New Guinea (especially Bird’s Head)
have somewhat shorter perianths, and smaller bracteole as compared to specimens from

De Wilde — Myristicaceae (Myristica) 585

eastern localities in Papua New Guinea. Coarse, stout specimens generally originate
from wet soils, e. g. riversides, swamp forest, or forest on (periodically) inundated ground.
Specimens with small leaves with a cuneate base, from all over the area, including some
from Bird’s Head, may strongly differ in general habit. Nearly always these small-leaved
forms are from well-drained soils, and possibly they represent a separate (subspecific)
taxon.

Some specimens from New Britain, e.g. NGF 6652, 22463, LAE 74233, have com-
paratively long petioles; the indumentum of the lower leaf surface and the fruits is par-
ticularly flocculose, with hairs to 1 mm long. These specimens are clearly different in
general aspect compared to the specimens here treated under var. subcordata. Male flow-
ering specimens from the Bismarck Archipelago are unknown.

4. The feature of cataphyll-like little-developed leaves on the twigs below the normal
leaves (i.e., of each new shoot the first leaves remain undeveloped), is frequent and
characteristic for var. morindiifolia.

5. Small-leaved specimens, sterile or with immature fruits, may be difficult to sepa-
rate from M. inutilis.

d. var. rimosa W.J. de Wilde

Myristica subcordata Blume var. rimosa W.J. de Wilde, Blumea 43 (1998) 181. — Type: de Vogel
9757, Irian Jaya, Bird’s Head.

Twigs terete, 5-8 mm diameter; older twigs at first blackish brown, soon pale and
conspicuously rough, flaky, and fissured by numerous elongate lenticels. Leaves coria-
ceous, elliptic-oblong or oblong(-lanceolate), 17-30 by 5—9.5 cm; lower surface with
pale yellowish brown dense short-felty hairs 0.1 mm; petiole short, 5(—10) by 3-5 mm,
dark brown. Inflorescences (female) of the Knema-type: sessile, 2- (or 3-)flowered. Fe-
male flowers + sessile; buds ovoid-oblong, narrowed towards the apex, hairs | mm long.
Fruits solitary, ellipsoid, 2.5-3.5(—4) by 1.5—2.5 cm, apex subacute, hairs powdery, (light)
brown, 0.5 mm; pericarp 3 mm thick; fruiting pedicel thick, 1-2 mm long, c. 5 mm thick,
frequently with the perianth persistent under the fruit, bracteole scar not seen.

Distribution — Malesia: Papua Barat (western Bird’s Head).

Habitat & Ecology — Forest on sandy or clayey soil, presumably all collections from
forest on limestone bedrock; 50—300 m altitude; fl. & fr. May.

Note — Var. rimosa is clearly distinguishable by the rough, flaky and lenticellate-
fissured bark of the twigs. The persistent perianth (with hairs 1 mm long) under the fruit
is remarkable.

131. Myristica succedanea Blume

Myristica succedanea Reinw. ex Blume, Rumphia | (1837) 186; Scheff., Ann. Jard. Bot. Buitenzorg
1 (1876) 46, p.p.; Warb., Ber. Pharm. Ges. (1892) 219; Die Muskatnuss (1897) 369, t. 3 f. 8;
Mon. Myrist. (1897) 474, t. 17; K. Heyne, Nutt. Pl. Ned. Indié (1927) 647; J. Sinclair, Gard.
Bull. Sing. 23 (1968) 238, f. 21; W.J. de Wilde, Blumea 35 (1990) 255, f. 2: 15, 3f, 4; 42 (1997)
185. — Type: Reinwardt s.n., Moluccas.

Myristica radja Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1864) 206, p.p. (non Rumph.); Warb., Mon.
Myrist. (1897) 453, sub syn. Myristica speciosa Warb. [Pala radja Teijsm., Natuurk. Tijdschr.
Ned. Ind. 23 (1861) 337]. — Type: Teijsmann s.n., culta Bogor, ex Bacan, p.p.

Flora Malesiana, Ser. I, Vol. 14 (2000)

586

De Wilde — Myristicaceae (Myristica) 587

Myristica succedanea Blume var. brevifolia Scheff. & Teijsm., Ann. Jard. Bot. Buitenzorg | (1876)
61, nom. nud. (Teijsmann 7586).

Myristica speciosa Warb., [Ber. Pharm. Ges. (1892) 219, t. 11; Die Muskatnuss (1897) 365, t. 3]
Mon. Myrist. (1897) 453, t. 17; K. Heyne, Nutt. Pl. Ned. Indié (1927) 647. — Syntypes: Beccart
Fl acc. nos. 7730, 7731, Warburg 18297 (B, lost); Teijsmann s.n., Moluccas.

Myristica schefferi Warb., [Ber. Pharm. Ges. (1892) 220; Die Muskatnuss (1897) 372;] Mon. Myrist.
(1897) 477, t. 17; K. Heyne, Nutt. Pl. Ned. Indié (1927) 647. — Syntypes: Teijsmann s. n. (Pala
onin); Warburg s.n. (culta, lost); Beccari acc. nos. FI 7694-7698 (culta BO) (see Sinclair: 241).

Myristica lakilaki Murata & Nitta, Acta Phytotax. Geobot. 40 (1989) 177. — Type: de Vogel 3668,
Moluccas, Bacan I.

Tree 8—20 m. Twigs 2—3.5 mm diameter, early glabrescent, hairs scattered, 0.1 mm;
older twigs somewhat cracking, with many inconspicuous lenticels. Leaves chartaceous
or usually (sub)coriaceous, elliptic or oblong, 9—22 by 3.5—10(—12) cm, base narrowly
rounded or cuneate, apex acute-acuminate; upper surface (olivaceous or yellowish) brown,
lower surface greyish, (late) glabrescent, hairs dense, scale-like, 0.1 mm or less, mixed
with few brownish hairs 0.1—0.2 mm; papillose or usually not; dots absent; midrib raised
above, lateral nerves 8—12(—15) per side, at 45—80° to the midrib, flat or sunken above,
lines of interarching and venation indistinct; petiole 10-20 by 1.5—3 mm; leaf bud (5—)
7-15 by 2-3 mm, hairs 0.1 mm. Inflorescences with hairs 0.1 mm, between the leaves or
below, or sometimes in to 5 cm long short-shoots resembling a compound inflorescence,
ending in a vegetative leaf bud; male and female inflorescences somewhat as in M.
fragrans: short-pedunculate, either simple or at the apex 2- (or 3-)forked, or each of the
branches forking again, with age developing into slender or thickish scar-covered up
to 15 mm long brachyblasts ending with 1-3 (or 4) flowers, buds almost equal in size;
bracts caducous; in male: peduncle 5—15(—20) by 2—2.5 mm, in female (1—)3-10 mm
long; flowers with hairs less than 0.1 mm. Male flowers: pedicel 7-12(-14) by 1(-1.5)
mm, bracteole rounded or reniform, 2—3 mm, persistent or late caducous; buds (ovoid-)
oblong, 7-11 by 4-6 mm, the apex rounded, cleft 1/5—1/8, lobes 0.4—0.8 mm thick.
Androecium 5—8 mm long; androphore 2—2.5 mm long, minutely pubescent or almost
glabrous; synandrium somewhat conical, 4-6 by 1.5—2(—2.5) mm, thecae 16-20(—24),
sterile apex blunt or irregular in shape, 0.3—0.5(—1) mm. Female flowers: pedicel (2—)4—
12 by 1.5 mm, bracteole late caducous; buds 7(—10) by 5-6 mm, cleft 1/5—1/4; ovary
narrowly ovoid, (3—)4—6 by 2—3(-3.5) mm, hairs sparse or dense, less than 0.1 mm, style
slender, 1.5 mm. Fruits single or paired, rather stout, ovoid-ellipsoid, coarsely wrinkled
on drying, 4.5-8 by 3.5—4.5 cm, apex + blunt or subacute, (usually) glabrescent, hairs
scattered, scale-like, 0.1 mm long or less, usually persistent towards the insertion of the
stalk; pericarp 10(—15) mm thick; seeds ellipsoid or oblong, (2—)2.5—3(—4) cm; fruiting
pedicel stout, 4-11 mm long. — Fig. 65f, 82f, 89.

Fig. 89. Myristica succedanea Blume. a. Leafy twig with male inflorescences; b. two male buds
with small persistent bracteoles, closed and opened; c. leafy twig with male inflorescences and
immature buds; d. opened bud of the same; e. apical part of twig with female inflorescences axillary
to reduced leaves; f. two female buds, closed, and opened showing pistil; note thinly hairy ovary;
g. apical part of leafy twig with infructescence with two developing fruits; h. mature fruit (dried)
[a, b: de Vogel 3626; c, d: de Vogel 3544; e: Woerjantoro 98 (culta); f: de Vogel 3642; g, h: de Vogel
3628). — Scale bar for a, c, e, g, h = 2 cm; for b, d, f = 3.3 mm.

588 Flora Malesiana, Ser. I, Vol. 14 (2000)

Field-notes — Clear bole to 15 m, very straight, without buttresses (but according to
Sinclair, l.c.: 239, usually with stilt-roots). Bark smooth or strongly peeling off with
+ round flakes, or bark with longitudinal cracks 5 mm apart; outer bark 1—2.5 mm thick,
dark brown: inner bark 6-7 mm thick, reddish with white rays; sap little, reddish, wa-
tery; sapwood pale reddish or yellowish with red rays, gradually passing into the slightly
darker heartwood. Young parts of twigs pale green with brown indumentum. Leaves
glossy dark green above, silvery below. Flowers (female) cream or yellowish green,
with brownish tinge, fragrant; ovary minutely rusty pubescent. Fruits yellowish green.

Distribution — Malesia: northern Moluccas (Ternate, Tidore, Bacan).

Habitat & Ecology — Lowland and montane forest; forest with little undergrowth
on level places on ridge; deep clayey soil, bedrock grey schists; steep slopes or ravines;
0-1100 m altitude; fl. Oct.; fr. July—Oct.

Uses — Said to be formerly cultivated on a small scale in Halmahera; nuts aromatic.

Note — This species is homogeneous, characterized among other species of the M.
fragrans-type of inflorescence (M. argentea, M. fragrans, M. impressinervia, partly) by
the + stout twigs and large flowers. It is also somewhat similar to M. pubicarpa from
nearby Halmahera and Obi.

132. Myristica sulcata Warb.

Myristica sulcata Warb., Mon. Myrist. (1897) 538, t. 19; J. Sinclair, Gard. Bull. Sing. 23 (1968)
396, f. 66: Foreman, Contr. Herb. Austral. no. 9 (1974) 42; Handb. Fl. Papua New Guinea 1
(1978) 211 (excl. syn. Myristica undulatifolia J. Sinclair); W.J. de Wilde, Blumea 40 (1995) 334;
43 (1998) 182. — Type: Chalmers s.n., Papua New Guinea, Central Prov.

Myristica anceps Warb., Mon. Myrist. (1897) 528. — Type: Beccari 681/, Irian Jaya, Bird’s Head.

Tree 10-40 m. Twigs 2-4 mm diameter, usually 2-edged, early glabrescent, hairs less
than 0.1 mm, seemingly glabrous; older twigs + fissured and transversely cracking;
lenticels small. Leaves membranous or chartaceous, ovate-elliptic or elliptic-oblong,
12-27(-30) by 4-12 cm, base attenuate or rounded, apex acute-acuminate; upper sur-
face olivaceous or (dark) brown, lower surface greyish, early glabrescent, hairs incon-
spicuous, scattered, scale-like, 0.1 mm or less; not papillose; dots absent; midrib + flat
or raised above, lateral nerves 15-21 per side, at 45-60(—70)° to the midrib, flat or
sunken above, lines of interarching distinct or not, venation + trabeculate, usually dis-
tinct below; petiole 15-25 by 2-3 mm; leaf bud 10-15 by (1—)2 mn, hairs 0.1—0.3 mm.
Inflorescences between the leaves or below, glabrescent, indumentum brown, short; of
the Knema-type: a simple or bifurcate, scar-covered brachyblast, sessile or 1(—2) mm
pedunculate; in male: to 10 mm long, 4-5 mm wide, with flowers in clusters of 5-10
(-15), buds unequal in size; in female shorter, simple or 2-forked, few-flowered; flowers
with widely spaced hairs 0.1 mm. Male flowers: pedicel slender, (5—)6-8 by 0.6-0.7
mm, bracteole 1.5 mm, subpersistent or caducous, (sub)apical; buds ellipsoid-oblong,
4—5 by 1.5—2 mm, apex subacute or obtuse, cleft 1/4-1/3, lobes (1—)1.5 mm long, 0.1-
0.2 mm thick. Androecium slender, 3—3.5 mm long; androphore 1.5 by 0.5 mm, with
hairs 0.1 mm or less in the lower half; synandrium (1.5—)2 by 0.6(—0.7) mm, thecae 12-
18, usually contiguous, sterile apex + blunt, 0.2(—0.3) mm long. Female flowers: pedicel
3-5 mm, bracteole scar subapical; buds ovoid, 4.5 mm; ovary ovoid-oblong, hairs 0.2
mm. Fruits 1 or 2 (or 3) per infructescence, subsessile, subglobose, short pear-shaped,

De Wilde — Myristicaceae (Myristica) 589

ellipsoid, or + depressed-globose, 3—4 by (2.5—)3—4 cm, apex broadly rounded, base
rounded or narrowed, + glabrescent, hairs mealy, greyish brown, 0.1—0.2 mm; pericarp
3-5(-8) mm thick; seeds broadly ovoid-ellipsoid or subglobose, 1.5—2(—2.5) cm, with
conspicuous impressions left by the aril; fruiting pedicel rather stout or slender, 7—10 by
3-5 mm, bracteole scar subapical.

Field-notes — Bole straight, without or with buttresses up to 1.5 m high and out, 5—
10 cm thick; buttresses and many adventitious roots protruding from c. 90 cm above the
ground (once recorded). Branches horizontal. Bark (grey-)brown, or blackish, smooth
or + longitudinally fissured, or slightly flaky, with lenticels; blaze (slash) purple, pink
red, or yellow-brown; inner bark layered pinkish; exudate colourless turning red, sticky
or watery. Wood pale, white or straw, turning reddish; no heartwood or heartwood darker.
Midnerve yellow-green below. Flowers yellow, or yellow with brown dots, slightly fra-
grant. Fruits (orange-)brown or red-brown; aril slightly aromatic.

Distribution — Malesia: lowland areas of Papua Barat (Bird’s Head, Jayapura) and
Papua New Guinea (Madang, Morobe, Central, Northern, and Milne Bay Provinces).

Habitat & Ecology — Forest on flats and ridges, dry land forest; on (stony or sandy)
clayey soil, also over limestone; 0—450(—700) m altitude; fl. & fr. throughout the year. In
Bird’s Head locally common.

Notes — 1. Myristica sulcata is recognized by its sulcate or angled twigs, especially
the apical part, and the on drying conspicuously (dark) brown, seemingly glabrous leaves.

2. Jacobs 9685 and LAE 52028 (Morobe Prov.) are mutually similar; they deviate
from M. sulcata by their faintly angled twigs and ellipsoid (not subglobose) fruits with
broadly rounded apex and asymmetrically positioned stigma remnant. Possibly these
specimens represent a separate form within M. sulcata. The fruits of these specimens
have some reminiscence to those of the related M. undulatifolia.

3. Myristica sulcata may be confused with M. crassipes and M. flosculosa, two spe-
cies generally with rather angular twigs and somewhat resembling leaves.

133. Myristica sumbawana Warb.

Myristica sumbawana Warb., Mon. Myrist. (1897) 383 (key), 529, t. 15, “swmbavana’. — Type:
Warburg 16983 (B, lost); neotype: Elbert 3809 (L, here designated), Sumbawa.

Tree 10-15 m. Twigs 2.5—5 mm diameter, early to late glabrescent, hairs 0.1—0.2 mm;
older twigs rather pale and + contrasting with the dark petioles, lenticels distinct or
not. Leaves chartaceous, elliptic-oblong or (oblong-)lanceolate, 19-38 by 5—12 cm, base
rounded or attenuate, apex + acute-acuminate; upper surface dark olivaceous (greenish),
lower surface golden(-brown) or greyish brown by interwoven stellate-dendroid hairs
0.i-0.2 mm; not papillose; dots absent; midrib above slightly raised, lateral nerves 20—
25 per side, at 40—80° to the midrib, flat or sunken above, lines of interarching and
venation faint or invisible; petiole 15-30 by 2.5—4.5 mm; leaf bud 10-20 by 2.5-5 mm,
hairs 0.2-0.3 mm. Inflorescences between the leaves and below, of the Knema-type:
without peduncle, single or paired, woody, scar-covered tubercle-like brachyblasts to
5 mm long, + with hairs 0.5 mm; bracts 1-1.5 mm, caducous; in male: with flowers in
subumbels of 2-6, buds of various sizes; female similar, 1—3-flowered; flowers with
conspicuous hairs 0.4—0.5 mm. Male flowers: pedicel to 0.3 mm only, bracteole 4.5 mm
long, persistent or late caducous; buds ovoid-ellipsoid, 6 by (3.5—)4 mm, apex narrowly

590 Flora Malesiana, Ser. I, Vol. 14 (2000)

rounded, cleft c. 1/3, lobes 0.3 mm thick. Androecium + slender, 4.5 mm; androphore 1—
1.2 by 0.6 mm, with few pale hairs 0.2—0.3 mm towards the base; synandrium 3—3.2 by
1 mm, thecae 16-20, sterile apex blunt, 0.3 mm. Female flowers: sessile, bracteole broadly
rounded, 3.5 mm; buds ovoid, 5 by 4 mm; ovary ovoid, 3 by 2.5 mm, pubescent. Fruits
solitary or 2 per infructescence, subsessile, broadly ovoid or ellipsoid, 2.5-3.5 by 2.5
cm, apex narrowly or broadly rounded, often 1-2 mm beaked, hairs dense, yellow-brown
or rusty, shaggy, (1—)1.5 mm; pericarp 2-4 mm thick; seeds (Schmutz 4277) ellipsoid,
2.5 cm, aril with laciniae up to 5 mm wide; fruiting pedicel 1-2 mm long.

Field-notes — Bark rough. Fruits brown.

Distribution — Malesia: Lesser Sunda Islands (Sumbawa, W Flores).

Habitat & Ecology — (Sub)montane forest, high monsoon forest, on andesit breccia;
250-800 m altitude (the lost type recorded from 1000-1300 m altitude in montane
forest): fl: Dec:; fr. Oct Ape

Note — Myristica sumbawana is related to M. rumphii, with which it has much of the
overall habit in common including the sessile flowers and + small sessile fruits. Myr-
istica rumphii differs from M. sumbawana in the shorter indumentum of the fruits (hairs
0.5 mm long only), glabrescent lower leaf surface, and the inconspicuous, slack hairs.
The male flowers of M. rumphii are unknown, but possibly they are not very different
from those of M. sumbawana. Both M. rumphii and M. sumbawana seem related to M.
fatua, the latter differing in the persistent dense indumentum on the lower leaf surface,
and the larger fruits and pedicellate flowers.

134. Myristica tamrauensis W. J. de Wilde

Myristica tamrauensis W.J. de Wilde, Blumea 40 (1995) 334. — Type: van Royen & Sleumer 7587,
Irian Jaya, Bird’s Head.

Tree 6 m. Twigs (sub)terete, 2-3 mm diameter, early glabrescent, hairs scale-like, less
than 0.1 mm; older twigs sometimes fissured or transversely cracked; lenticels incon-
spicuous. Leaves membranous, brittle, (ellipsoid-)oblong, 15—23 by 6-9 cm, base (short)
cuneate, apex acute(-acuminate); upper surface (olivaceous) or dark brown, lower sur-
face grey-brown, glabrous (very early glabrescent); not papillose; dots just visible to the
naked eye; midrib raised above, lateral nerves 15—20 per side, at 60—80° to the midrib,
inconspicuous above, bright brown and contrasting in colour below, lines of interarch-
ing and venation faint; petiole (10—)15 by 3-3.5 mm; leaf bud 10-15 by 2-3 mm, hairs
scale-like, 0.1 mm or less. /nflorescences, male and female flowers not seen. Infructescence
(van Royen & Sleumer 7587) between the lower leaves, peduncle 3-5 mm. Fruit soli-
tary, ovoid-ellipsoid, 4.5—5 by 3.5 cm, apex + pointed, hairs dense, mealy, rusty, 0.1 mm;
seeds ellipsoid(-oblong), 2 cm; pericarp rather woody, 8(—10) mm thick; fruiting pedicel
4 mm long.

Field-notes — Low tree. Leaves greyish green, or silvery light green below. Fruits
orange or light orange-brown.

Distribution — Malesia: W Papua Barat (endemic to the mountains of northern Bird’s
Head: Tamrau Range).

Habitat & Ecology — Oak forest; 500-800 m altitude; fr. Oct—Nov.

Note — Myristica tamrauensis is a species within a group of related species with,
e.g., M. brassii, M. duplopunctata, M. millepunctata, M. neglecta, and, more remotely,

De Wilde — Myristicaceae (Myristica) 591

M. cucullata, all characterized by leaves of medium or large size classes, with blackish
dots on the lower leaf surface, and by fruits with a thick pericarp. Myristica tamrauensis
is known only in fruit. Mature male flowers are known only from M. brassit, M. duplo-
punctata, and M. neglecta. The present species has the marked, contrasting lateral nerves
on the lower leaf surface in common with M. brassii (Central New Guinea), but that
species has more coriaceous leaves and sessile inflorescences.

135. Myristica teijsmannii Miq.

Myristica teijsmannii Miq., Fl. Ind. Bat. 1, 2 (1858) 57, ‘teysmanni’; Koord. & Valeton, Bijdr.
Booms. Java 4 (1896) 180; Warb., Mon. Myrist. (1897) 516, t. 16; Koord., Exk. Fl. Java 2 (1912)
257; K. Heyne, Nutt. Pl. Ned. Indié (1927) 648; Backer & Bakh. f., Fl. Java 1 (1964) 139; J. Sin-
clair, Gard. Bull. Sing. 23 (1968) 430, f. 76; W.J. de Wilde, Blumea 42 (1997) 185. — Type:
Teijsmann s.n., Java.

Myristica hyposticta Miq., Fl. Ind. Bat. 1, 2 (1858) 55; Koord. & Valeton, Bijdr. Booms. Java 4
(1896) 178. — Type: Horsfield s.n., Java.

Tree 10-20 m. Twigs 2—4(—5) mm diameter, somewhat angular, yellowish or orange
brown, soon glabrescent, hairs 0.1 mm; older twigs longitudinally cracking, not flaking;
lenticels sometimes present, small. Leaves membranous or thinly chartaceous, elliptic-
oblong or oblong-lanceolate, 10—30(—34) by 2.5—9 cm, base narrowly rounded or cuneate,
apex acute-acuminate; upper surface olivaceous brown, lower surface grey-brown, seem-
ingly glabrous, but hairs scattered, scale-like, less than 0.1 mm; papillose; dots absent;
midrib flat above, lateral nerves 14-19 per side, at 45(-60)° to the midrib, flat above,
distinct below, lines of interarching and venation indistinct; petiole 10—20(-30) by 1.5—
3 mm: leaf bud 8-12 by 2.5 mm, hairs dense, 0.2 mm. Jnflorescences between the leaves,
sometimes on short lateral shoots, short-pedunculate, unbranched, possibly of longer
duration, glabrescent, hairs 0.1—0.2 mm; bracts 3-4 mm, caducous; in male: peduncle
(1-)5-15 mm, at apex with irregularly wart-like brachyblast (central axis usually ab-
sent) with a subumbel of (5—)10—20 flowers, buds of various sizes; female smaller, pe-
duncle to 3 mm, brachyblast to 10 mm long, few-flowered; flowers with hairs 0.2-0.3
mm. Male flowers: pedicel 4-6 mm long, bracteole broadly reniform, 4—6 mm, (sub)-
persistent; buds carnose-coriaceous, ovoid-ellipsoid, 7-8 by 4-4.5 mm, apex narrowly
rounded, cleft c. 1/3, lobes 0.4 mm thick. Androecium 5—5.5 mm; androphore 2 by |
mm, with appressed hairs, less than 0.1 mm, in lower 3/4: synandrium (2—)2.5 by 1-1.5
mm, thecae 16-24, sterile apex + blunt, 0.3(-0.5) mm long. Female flowers: pedicel
stout, 2 mm long; buds ovoid, 4-6(—9) by 4-5(—7) mm, cleft c. 1/3, lobes 0.4-0.5 mm
thick; ovary ovoid, 2.5-3.5 by 2-2.5 mm, minutely pubescent. Fruits 1 (or 2) per infruc-
tescence, subsessile, subglobose or broadly ovoid, 3-4.5 cm diameter, hairs dense, dark
rust-coloured, 0.1 mm or less; pericarp 4-8 mm thick; seeds short ellipsoid, 2.5—3 cm;
fruiting pedicel 5 mm.

Field-notes — Trunk often with aereal roots or stilt-roots. Outer bark with a few
flakes, but no ridges or furrows; living bark reddish brown, 0.7 cm thick; sap copious,
red-brown. Flowers yellow brown. Fruits brown.

Distribution — Malesia: C & E Java.

Habitat & Ecology —A rare tree of mixed lowland and submontane forest; altitude
50-1000 m; fl. Oct.-May; fr. May—June.

592 Flora Malesiana, Ser. I, Vol. 14 (2000)

Notes — 1. Close to M. andamanica Hook. f. (Andaman Islands) and M. crassa, with
similar unbranched inflorescences; M. andamanica differs in its non-papillose lower
leaf surface.

2. In the Teijsmann collections at BM, some (large-sized) female flowers are found
mixed with male flowers in one inflorescence.

136. Myristica tenuivenia J. Sinclair

Myristica tenuivenia J. Sinclair, Gard. Bull. Sing. 23 (1968) 327, f. 46; Foreman, Handb. Fl. Papua
New Guinea | (1978) 211; W.J. de Wilde, Blumea 40 (1995) 335. — Type: Brass 27658, Misi-
ma I.

Tree 10-30 m. Twigs (sub)terete or 2-lined, 2.5—3(—5) mm diameter, early glabrescent,
hairs 0.1—0.2 mm; older twigs with few scattered small lenticels. Leaves membranous,
chartaceous or subcoriaceous, obovate-elliptic or oblong, 8-17 by 3-7 cm, base short-
attenuate or rounded, apex (short-)attenuate with blunt tip; upper surface olivaceous or
(dark) brown, lower surface greyish cinnamon, indumentum persistent, hairs scale-like,
0.1 mm or less; not papillose; dots absent; midrib above flat or slightly raised, lateral
nerves 16—23 per side, at 60—80° to the midrib, thin and sunken above, (hardly) raised
below, lines of interarching and venation faint; petioles 16-22 by 2—3.5 mm; leaf bud
15-20 by 2.5 mm, hairs 0.1-0.3 mm. Inflorescences (known only from female flowers
and infructescences) between the leaves and below, of the Knema-type: a sessile, simple
or 2—3-forked, scar-covered, woody knob-like brachyblast, 1-3 mm diameter, with hairs
0.2-0.5 mm, with a fascicle of 4-10 flowers, buds of various sizes or almost equal in
size. Male flowers not seen. Female flowers: with hairs 0.2—0.4 mm; pedicel 1-4 mm
long, bracteole 1.5 mm long, caducous; buds hard-carnose, ovoid, 3—4 by 2.5-3 mm,
lobes 1.5 mm long; ovary ovoid, 2 by 1.5 mm, hairs 0.5 mm or less. Fruits 1 or 2 per
infructescence, broadly ellipsoid or (sub)globose, 3—4 cm long, hairs rusty, 0.2-0.5 mm,
partly glabrescent; pericarp 4-8 mm thick, fleshy or woody, seeds ellipsoid, 1.5—2.5 cm,
fruiting pedicel rather stout, 2-7 mm long.

Distribution — Malesia: Papua New Guinea (confined to the Papuan Islands). Provi-
sionally with two subspecies.

KEY TO THE SUBSPECIES

la. Twigs neither ridged nor lined, pericarp not woody ....... a. subsp. tenuivenia
b. Twigs ridged or 2-lined, pericarp granular-woody ........... b. subsp. lignosa

a. subsp. tenuivenia

Canopy tree. Twigs neither ridged nor lined. Leaves thinly chartaceous or membra-
nous, lateral nerves 15—20 pairs, + raised below. Fruits ellipsoid or subglobose, 3-4 cm
diameter, hairs mealy or scurfy, 0.2—0.4 mm, partly glabrescent; pericarp not woody,
4—8 mm thick, apparently juicy and much shrunken on drying; fruiting pedicel 2-4 by
4 mm.

Field-notes — Leaves grey-brown or brownish below. Flowers yellow. Fruits brown
pubescent, on the older wood.

De Wilde — Myristicaceae (Myristica) 598

Distribution — Malesia: Papua New Guinea: Louisiade Archipelago (Misima I., type,
and Rossel I., Brass 28528).

Habitat & Ecology — Streambank forest; the type from quartz mountain; 10-150 m
altitude; fl. Aug.; fr. Aug., Oct.

Note — In the description of M. tenuivenia Sinclair (1.c.: 329) suggested a relation-
ship to M. archboldiana, M. pedicellata, and M. smythiesii (from Borneo) on account of
the cinnamon or rusty scale-like hairs on the lower leaf surface, but possibly there only
is a remote affinity with M. pedicellata.

b. subsp. lignosa W. J. de Wilde

Myristica tenuivenia J. Sinclair subsp. lignosa W.J. de Wilde, Blumea 40 (1995) 335. — Type:
LAE 68655 (Croft et al.), Papua New Guinea, Fergusson I.

Tree up to 30 m. Twigs ridged or 2-lined. Leaves subcoriaceous, lateral nerves 20-28
pairs, faint on both surfaces. Fruits subglobose, (2.5—)3 cm diameter, hairs (0.2—)0.5
mm long, persistent; pericarp granular-woody, 5—7 mm thick, not shrunken on drying;
fruiting pedicel 5—7 by 5 mm.

Field-notes — Bole 20 m, dbh 35 cm. Bark red-brown or black, fissured and pustular;
inner bark red; exudate little, red; wood straw, pink when exposed. Leaves glossy dark
green above, dull mid-green below with rust-brown bloom, or rusty-grey. Flowers and
fruits rust-brown.

Distribution — Malesia: Papua New Guinea (S Fergusson I., type, and Rossel L.,
LAE 74561, with immature fr.).

Habitat & Ecology — Hill forest; c. 700 m altitude; fl. (female) & fr. Mar.

Note — The two specimens known of subsp. lignosa considerably deviate from those
of the type subspecies. When more material becomes available it may be found that two
separate species are concerned here.

137. Myristica trianthera W.J. de Wilde

Myristica trianthera W.J. de Wilde, Blumea 40 (1995) 335. — Type: BW 39 (Versteegh 58), Irian
Jaya, Jayapura area.

Treelet 10-13 m. Twigs 1 mm diameter, early glabrescent, hairs setose, 0.5—1 mm
long; older twigs + smooth; lenticels not apparent. Leaves membranous, ovate-elliptic
or elliptic-oblong, 6—12.5 by 2-5 cm, base + rounded or attenuate, apex acute-acumi-
nate; upper surface grey-green or brown, lower surface greyish brown, early glabrescent
or few rather long widely scattered hairs left; not papillose; dots absent; midrib flat or
somewhat raised above, lateral nerves 7-10 per side, at 50—80° to the midrib, flat and
faint above, rather faint below, lines of interarching and venation indistinct; petiole 6-10
by 1 mm; leaf bud 6—8 by | mm, hairs (0.5—)1 mm long. Inflorescences between the
leaves or below, paniculate; in male: peduncle 5—20 by 0.5—0.8 mm, ending in a simple
or forked, densely scar-covered, + pubescent brachyblast to 9 mm long, with cluster(s)
of 2—5 flowers, buds of various sizes; in female (from infructescence): peduncle 9 mm,
one- or few-flowered, unbranched; flowers with shaggy hairs (0.3—)0.5 mm. Male flow-
ers: pedicel slender, 3 mm long, bracteole + ovate, 1(—1.5) mm, persistent, (sub)apical;
perianth membranous, inside not warty; buds ellipsoid-oblong, 4—4.2 by 1.5-1.8 mm,

594 Flora Malesiana, Ser. I, Vol. 14 (2000)

apex and base subacute to obtuse, cleft 1/5—1/4, lobes 0.7—1 mm. Androecium slender,
3.8 mm long; androphore 4 by 0.4 mm, with appressed hairs 0.4 mm long; synandrium
1.8 by 0.5 mm, thecae 6, not closely appressed, sterile apex 0.1 mm. Female flowers not
seen. Infructescence simple, peduncle slender, 8-10 mm long, proceeding into a 12-14
mm long pedicel of a single fruit. Fruits + ellipsoid, 2—2.5 by 1—1.3 cm, apex short-apic-
ulate, base contracted into pseudostalk up to 5 mm; hairs dull brown, scurfy, less than
0.1 mm, at apex and base longer and shaggy; pericarp thin, | mm thick; seeds ellipsoid,
1.5 cm; fruiting pedicel slender, 14 mm long, bracteole scar 3—4 mm below the apex.

Field-notes — Flower buds green.

Distribution — Malesia: NE Papua Barat (endemic to the Jayapura area).

Habitat & Ecology — Lowland riverside or temporarily inundated forest, clayey soil;
scattered; 2—50 m altitude; fl. July, Oct.; fr. July.

Note — Resembles M. fugax but M. trianthera is distinguishable by its pilose leaf
bud, smaller flowers and fruits; the synandrium has only 3 anthers, 4—6 in M. fugax.

138. Myristica tristis Warb.

Myristica tristis Warb., Mon. Myrist. (1897) 444, t. 19. — Syntypes: Beccari 96 (FI acc. nos. 7657,
lecto, here designated; 7657a), Bird’s Head, Sorong.
For more references see under the subspecies.

Tree (3—)12—-30 m. Twigs somewhat angular (sometimes with faint ridges between
insertions of petioles), 1-2 mm diameter, reddish or yellowish brown, early glabrescent,
hairs fine-woolly, (0.1-)0.2 mm; older twigs (faintly) cracking, sometimes flaking;
lenticels inconspicuous. Leaves membranous or thinly chartaceous, elliptic-oblong or
oblong-lanceolate, 8-16 by 2-6 cm, base cuneate, apex acute-acuminate, the tip some-
times + blunt; upper surface olivaceous or (blackish) brown, lower surface (grey-)brown,
early glabrescent, hairs scattered, scale-like, 0.1 mm; not papillose; dots absent; midrib
above raised or + flat, lateral nerves 10—15(—17) per side, at 45—60° to the midrib, flat or
sunken (and faint) above, little raised below, lines of interarching and venation indis-
tinct; petiole comparatively long, 12—20 by 1—-1.5 mm; leaf bud 6-12 by 1-2 mm, hairs
0.1 mm or less. Inflorescences between the leaves and below, of the Knema-type: sessile
simple or forked brachyblasts, 2-4 mm long, with hairs 0.2 mm; bracts minute; in male:
with one or two subumbels of 3-6(—12) flowers, buds of various sizes or almost equal in
size: female similar to male, (1-)2—4(—10)-flowered; flowers with hairs 0.1—0.2 mm or
less. Male flowers: pedicel (4-)6-7 by 0.5 mm, bracteole persistent, ovate, 1-1.5 mm;
buds ovoid(-ellipsoid) or ovoid-oblong, (4—)5—6 by 2.5—3 mm, apex + acute, cleft
(1/5—)1/4, lobes 0.2 mm thick. Androecium 3-5.5 mm long; androphore 1-2 by (0.3-)
(.4-0.6 mm, with hairs 0.1—0.3 mm in the lower half; synandrium 2—3.5 by 0.6—0.8 mm,
apex blunt or acutish, thecae (8—)10-—14, sterile apex 0.1 mm. Female flowers: pedicel
3—4.5 by 0.7—1 mm, bracteole rounded, | mm, caducous, (sub)apical; buds ovoid(-ob-
long), 4(—4.5) by 2.5-3 mm, apex + blunt, lobes 1.5—1.7 mm long; ovary ovoid, includ-
ing short stigma 3—4 by 2.2-2.5 mm, hairs 0.1—0.2 mm. Fruits solitary or 2 (or 3) per
infructescence, (broadly) ellipsoid to globose, 2—3.5 by 1.5—3 cm, apex broadly rounded
or subacute, hairs brown, scurfy, 0:1—0.2 mm or less, partially (late) glabrescent; pericarp
3-8 mm thick; seeds broad-ellipsoid, 2 cm, blackish, deeply broadly grooved by impres-
sion of the aril; fruiting pedicel usually slender, 2-5 mm long.

De Wilde — Myristicaceae (Myristica) 595

Field-notes — Trunk incidentally with buttresses to | by 1 m, or with buttresses as
well as prop-roots. Bark strongly fissured or strongly flaking, flakes 0.5 by 1 cm; outer
bark brown, | mm thick; inner bark 6-7 mm thick, pale red; exudate little or much, pale
red, watery; sapwood cream or light brown, tinged red, the transition to the slightly
darker heartwood gradual. Flowers cream. Fruits yellowish, seeds greyish black.

Distribution — Malesia: Moluccas, throughout New Guinea; rare in eastern Papua
New Guinea, where it is largely replaced by the resembling M. globosa. There are five
subspecies.

Habitat & Ecology — Locally common as a solitary tree in lowland and submontane
forest: forest on limestone, (sandy-)clayey soil over limestone; 0—600 m altitude; fl.
May; fr. May—Nov.

Notes — 1. Related to M. globosa, but M. tristis is distinguishable and chiefly charac-
terized by slender twigs, membranous leaves drying frequently dark brown or blackish
above, faint venation on the pale brownish lower leaf surface (generally in M. globosa
the lower leaf surface much paler, with contrasting venation), persistent bracteole in the
male flowers (seen in the few male-flowering collections available), generally almost
globose fruits with minute scurfy hairs 0.1 mm or less, and thick pericarp, 3—8 mm thick.
Myristica tristis also resembles the variable M. lancifolia (incl. M. montana as a sub-
species), a species with thicker and differently shaped leaves, a different venation and
papillose lower surface, more cylindric (not ovoid-oblong) male buds, and rather small
ellipsoid fruits. For differences with the resembling M. lepidota subsp. montanoides, see
there. Also similar is M. cumingii (Philippines) which differs in its male buds, cleft to
nearly halfway, and its larger fruits.

2. Myistica tristis is highly polymorphic with a vast distributional area. Variation
abounds in distinctness of the lateral nerves on the lower leaf surface, the midrib above
either raised or flat, and especially in size and shape of the fruits, and thickness of the
pericarp. Most variation, especially in the fruits, is correlated with the geography. Male
flowering specimens still have been insufficiently collected.

KEY TO THE SUBSPECIES

la. Fruits globose, 3 cm diameter; pericarp S—8 mm thick. —N Moluccas ........--.
Sete oa. cie bint Hamena hg Mite naire cml d. subsp. moluccana

b. Fruist globose, less than 2.5 cm diameter, or (broadly) ellipsoid; pericarp 5 mm thick
5p ee ane een tee ricer ara acy ace tts Shee cae Crcaceae ica 2

2a. Fruits ellipsoid, 3.5 cm long; fruiting pedicel 2 by 2 mm. — N Moluccas, Bacan...
MN Re 2A a Ws me hice le CSS Msltete, Me amen ae tates e. subsp. sessilifructa

b. Fruits subglobose or broadly ellipsoid, 1.8—2.3(—2.5) cm long; fruiting pedicel 3-5
“HTT pes eye gel SNS Oe RO MaMa alee En Mae MR Tee rl Rb Arca alla eA Sacer A Ne lec fas oc 3

3a. Fruits 2 cm long; pericarp (2—)3 mm thick. Lateral nerves on lower leaf surface not
much raised but usually distinct; [midrib flat or raised above.] — W and E New
Guinea main island (incl. Bird’s Head and islands in the Geelvink Baai).........

= ig LO ET PR a Pb eRe SUN a. subsp. tristis

b. Fruits slightly larger (longer), 2.3(—2.5) cm long; pericarp (2—)3-5 mm thick. Lat-
eral nerves on lower leaf surface faint .... 0.0.0.0. 22. ee ee oe eee -

596 Flora Malesiana, Ser. I, Vol. 14 (2000)

da. Leaves glossy on drying at both surfaces; midrib raised above. Fruits brightly rusty
pubescent. Older twigs blackish, strongly finely flaking. — Central southern New
Guiiea eat ok 0 at eR Sat a b. subsp. ingambitense

b. Leaves rather dull on drying; midrib + flat or but slightly raised above. Fruits choco-
late or dull brown pubescent. Older twigs grey-brown, striate or slightly cracked, but

not flaky. — Louisiade Archipelago. ......0....00.250. c. subsp. louisiadensis

a. subsp. tristis

Myristica tristis Warb., Mon. Myrist. (1897) 444, t. 19. — Myristica globosa auct. non. Warb.:
J. Sinclair, Gard. Bull. Sing. 23 (1968) 378, f. 63, p.p., for most of the material from the Moluccas,
Papua Barat (Bird’s Head), and Mios Noem I., incl. f. 63A—C; W. J. de Wilde, Blumea 35 (1990)
258, f. 2 (16); 40 (1995) 336; 42 (1997) 186.

Older twigs grey-brown, striate or longitudinally cracked or but faintly flaky. Midrib
flat or raised above. Lateral nerves on lower leaf surface usually clearly visible; vena-
tion usually faint or absent. Male and female flowers as the species. Fruits globose or
broadly ellipsoid, 1.8—2.2 cm long, hairs dull grey-brown; pericarp (2—)3(—4) mm thick.
— Fig. 6lf, 65g.

Distribution — Malesia: throughout New Guinea, including Bird’s Head, and islands
in the Geelvink Baai; 0—400 m altitude.

b. subsp. ingambitense W. J. de Wilde

Myristica tristis Warb. subsp. ingambitense W.J. de Wilde, Blumea 40 (1995) 336; 42 (1997) 186.
— Type: NGF 31816 (Henty, Ridsdale & Galore), Papua New Guinea.

Older twigs dark brown, strongly finely flaking. Leaves membranous, brown, glossy
above: midrib above narrow, raised; lateral nerves flat, narrow, faint on both surfaces,
venation almost invisible. Male and female flowers not seen. Fruits solitary or paired,
subglobose or broadly ellipsoid, 2.5 cm long, hairs bright rust-coloured; pericarp (2—)3
mm thick.

Field-notes — Small tree about 7 m tall. Bark brown, smooth, aromatic; wood white;
exudate red. Fruit with brown indumentum.

Distribution — Malesia: New Guinea (SE Papua Barat; SW Papua New Guinea:
Western Prov., in the area of Ingambit village).

Habitat & Ecology — Regrowth forest; c. 150 m altitude; fr. June.

c. subsp. louisiadensis W. J. de Wilde

Myristica tristis Warb. subsp. louisiadensis W.J. de Wilde, Blumea 40 (1995) 337; 42 (1997) 186.
— Type: LAE 74537 (Damas & Katik), Tagula I.

Older twigs dark brown, longitudinally pliate (striate) or fissured, but not flaky. Leaves
thickly membranous, olivaceous, rather many-nerved but the lateral nerves faint on both
surfaces, venation faint; midrib flattish above. Male and female flowers not seen. Fruits
solitary, subglobose, 2.5 cm diameter, hairs dull brown, minute; pericarp 4-5 mm thick.

Field-notes —Tree 8—25 m, of subcanopy layer. Bark black, rough outside; inner bark
red; wood white with red tinge. Leaves pale green below. Fruit yellow, seed dark brown
with strong nutmeg flavour.

De Wilde — Myristicaceae (Myristica) 597

Distribution — Malesia: Papua New Guinea (Louisiade Archipelago: Tagula I.; pos-
sibly Rossel I.).

Habitat & Ecology — Lowland hilly forest subcanopy layer; (50—)100—450 m alti-
tude; fr. Mar., Aug., Oct.

Note — Brass 28302 (Rossel I.) vegetatively agrees with the type, but the fruits devi-
ate by a more ellipsoid shape and thinner pericarp.

d. subsp. moluccana W. J. de Wilde

Myristica tristis Warb. subsp. moluccana W.J. de Wilde, Blumea 42 (1997) 186. — Type: de Vogel
4058, Moluccas.

Older twigs grey-brown, striate or slightly cracked, but not flaky. Midrib raised above.
Lateral nerves on lower leaf surface weak, but clearly visible; venation faint. Male and
female flowers not seen. Fruits globose, 3 cm diameter, hairs sparse, greyish; pericarp
5-8 mm thick.

Distribution — Malesia: N Moluccas (Obi, Morotai, E Seram); up to 600 m altitude.

e. subsp. sessilifructa W. J. de Wilde

Mpyristica tristis Warb. subsp. sessilifructa W.J. de Wilde, Blumea 42 ( 1997) 186. — Type: de Vogel
3858, Moluccas.

Older twigs finely striate, not flaking, light brown. Leaves membranous, dark (black-
ish) brown, lateral nerves 10-14 per side, thin and faint, venation faint; midrib + raised
above. Male and female flowers not seen. Fruits solitary, subsessile, axillary to the (lower)
leaves, ellipsoid, 3.5 by 1.7 cm, hairs dull brown, 0.1 mm or less; pericarp 2(-3) mm
thick.

Field-notes — Solitary tree 12 m, with few small buttresses 50 cm out, clear bole 6m,
dbh 18 cm. Outer bark 0.3 mm thick, dark grey, rather fissured, not peeling off; inner
bark 4 mm thick, on section reddish; exudate little, red-brown, watery; sapwood pale
yellowish, tinged reddish, gradually passing into the slightly darker heartwood.

Distribution — Malesia: N Moluccas (Bacan).

Habitat & Ecology — Tall forest with little undergrowth on sloping hill ridge; soil
loamy with stones, with little humus; 100 m altitude; fr. Nov.

Note — The true status of this subspecies is unclear yet (see De Wilde, I.c.).

139. Myristica tubiflora Blume

Myristica tubiflora Blume, Rumphia 1 (1837) 182, t. 56; Warb., Mon. Myrist. (1897) 436; Pulle,
Nova Guinea 8 (1912) 636; Markgr., Bot. Jahrb. Syst. 67 (1935) 162, p.p.; J. Sinclair, Gard.
Bull. Sing. 23 (1968) 339, f. 51, p.p. (mixed with Hoogland 3971 = Myristica pumila); Fore-
man, Handb. FI. Papua New Guinea | (1978) 212, p.p.; W.J. de Wilde, Blumea 40 (1995) 337,
f. 8. — Type: Zippelius s.n., Irian Jaya, Bird’s Head.

Tree 3-24 m. Twigs subterete or + angular, yellowish brown, (0.5—)1—1.5 mm diam-
eter, early glabrescent, hairs less than 0.1 mm; older twigs cracking and flaking; lenticels
small, dense on younger twigs, indistinct or absent on the older twigs. Leaves membra-
nous or thinly coriaceous, (elliptic-)oblong, 5—14(-16) by 1.5-5.5(—7.5) cm, base at-

Flora Malesiana, Ser. I, Vol. 14 (2000)

598

De Wilde — Myristicaceae (Myristica) 599

tenuate or rarely + rounded, apex acute-acuminate, with + blunt or acute tip; upper sur-
face olivaceous, lower surface pale brown, glabrous; not papillose; dots absent; midrib
raised above, lateral nerves 8-13 per side, at 60—70° to the midrib, flat or sunken above,
far apart, lines of interarching and venation indistinct; petiole 10-18 by 0.8—-1.5 mm;
leaf bud conspicuously slender, 5—12 by 0.5—1 mm, hairs less than 0.1 mm. /nflores-
cences between the older leaves or below, of the Knema-type: a simple sessile scar-
covered brachyblast up to 4 mm long, glabrescent; bracts minute; in male: 1—3(—4)-
flowered, buds of slightly variable sizes; in female minute, sessile, 1- (or 2-)flowered;
flowers glabrous (or with few scattered appressed hairs less than 0.1 mm). Male flowers:
pedicel 8-11.5 mm, bracteole ovate-oblong, 1-2 mm, persistent or late caducous, 1—3.5
mm below the apex; buds membranous, oblong-lanceolate, 8.5-11 by 2—3 mm, apex
subacute or narrowly rounded, + triangular, base short-attenuate, cleft 1/6—1/8, lobes
(1-)1.5(—2) mm long, c. 0.2 mm thick. Androecium slender, 6—8 mm; androphore 2.5—4
by 0.8-1(-1.5) mm, glabrous; synandrium 4-6 by 0.8-1.5 mm, thecae c. 15, sterile apex
+ acute or lobulate, 0.2—0.3(—0.4) mm long. Female flowers (BW 1503): pedicel 2-4
mm long, bracteole + ovate, | mm, persistent, | mm below the apex; buds ovoid-oblong,
apex narrowed, faintly triangular, 6 by 3-3.5 mm, lobes 1.5-2 mm long; ovary ovoid-
oblong, 4 mm long, hairs less than 0.1 mm. Fruits solitary, oblong, 4.5—6 by 2—2.5 cm,
apex + acute, often somewhat oblique, base narrowed into a 1—1.5 cm long pseudostalk,
glabrous or glabrescent, hairs (grey-)brown, powdery, 0.1 mm; pericarp bright brown,
2-3 mm thick; seeds ellipsoid-oblong, 3 cm; fruiting pedicel slender, 2-7 mm long,
bracteole scar 1-3 mm below the apex. — Fig. 90.

Field-notes — Understorey tree, dbh to c. 40 cm; branches horizontal, twigs pendu-
lous, spreading; crown dense; without or with small buttresses, up to 30 cm out. Bark
dark brown or dark grey-black, strongly flaking; exudate red gum; wood salmon-pink or
straw. Flower buds yellow. Fruits green, yellow or pale brown.

Distribution — Malesia: mainly western half of New Guinea (Papua Barat and SW
Papua New Guinea: Western Prov.).

Habitat & Ecology — Coastal plain forest, riverine (seasonally inundated) swamp
forest, forest on coral limestone; mostly on (sandy) clay soil over limestone; 0—100
(250) m altitude; fl. throughout the year; fr. mostly July, Aug. Locally common, scat-
tered or a few trees together.

Note — Myristica tubiflora is a lowland species, characterized by leaves of a small
size class, sessile inflorescences, elongate male flowers with long pedicels, with the
(sub)persistent bracteole usually at some distance below the perianth (in most Myristica
species the bracteole is apical) and elongate fruits. Sterile specimens and those with
immature fruits may resemble M. lepidota subsp. montanoides, and some, now treated
under M. tubiflora, may belong to that taxon or vice versa. This may partly account for
the remarkable variation in leaf texture admitted for M. tubiflora: the majority of the

Fig. 90. Myristica tubiflora Blume. a. Leafy twig with male inflorescences; b. male bud (just before
anthesis) opened, showing androecium; c. leafy twig with one-flowered female inflorescences and
one developing fruit; d. female bud just before anthesis; note persistent bracteole; e. leafy twig with
infructescence with a single fruit [a, b: Atasrip 709; c, d: BW 1503 (Schram); e: LAE 51919 (Strei-
mann et al.)|. — Scale bar a, c, e = 2 cm; for b, d= 1.65 mm.

600 Flora Malesiana, Ser. I, Vol. 14 (2000)

specimens have subcoriaceous leaves, but some have brittle membranous leaves, with a
more distinct venation, reminiscent of the leaves of M. lepidota subsp. montanoides.
The latter subspecies, however, is distinguishable by its small flowers and fruits.

140. Myristica ultrabasica W. J. de Wilde
Myristica ultrabasica W.J. de Wilde, Blumea 42 (1997) 187. — Type: van Balgooy 4064, C Sulawesi.

Tree 20-30 m. Twigs 1-2 mm diameter, early glabrescent, hairs 0.1 mm or less; older
twigs coarsely striate; lenticels inconspicuous. Leaves thinly coriaceous, elliptic-oblong
or oblong(-lanceolate), (2.5—)4-7.5 by (0.8—)1—2 cm, base attenuate, apex subacute or
blunt, sometimes narrowly rounded; upper surface olivaceous or blackish brown, lower
surface greyish or reddish brown, glabrous; not papillose; dots absent; midrib above
+ flat, lateral nerves 10-15 per side, at c. 45° to the midrib, rather closely set, thin, hard-
ly visible on both sides, lines of interarching and venation invisible; petiole 4-9 by 1
mm; leaf bud 3-6 by | mm, hairs 0.1 mm or less. Female inflorescences (from infructes-
cences): between the leaves and below, of the Knema-type: a sessile short wart-like
1 mm long short-shoot, few-flowered, short-pubescent. Male inflorescences, male and
female flowers not known. Fruits solitary, (ovoid-)ellipsoid, 2—2.5 by 1.5—1.9 cm, apex
sometimes | mm beaked, base narrowly or broadly rounded, hairs persistent, dense, dull
or bright rusty, scurfy, 0.1 mm or less; pericarp 1-2 mm thick; seeds 1.7 cm, ellipsoid;
fruiting pedicel 1.5—3 mm long.

Field-notes — Crown small, restricted to apical 5 m. Bark purple-grey with fine lon-
gitudinal cracks; sap red. Fruits orange or brown.

Distribution — Malesia: C Sulawesi.

Habitat & Ecology — Forest on ultrabasic (nickel), shales; 400-750 m altitude; fr.
June—Aug.

Note — The small, thinly coriaceous leaves with faint nerves are quite typical. Myr-
istica ultrabasica (known only in fruit) is similar to M. lancifolia which differs in its
papillose lower leaf surface; it is also similar to M. bifurcata subsp. sulaica (with larger
leaves and fruits) and M. tristis (with larger, thinner leaves and much larger subglobose
fruits).

141. Myristica umbellata Elmer

Myristica umbellata Elmer, Leafl. Philipp. Bot. 5 (1913) 1816; Merr., Enum. Philipp. Flow. PI. 2
(1923) 180; J. Sinclair, Gard. Bull. Sing. 23 (1968) 174, f. 10; W.J. de Wilde, Blumea 42 (1997)
188. — Syntypes: Elmer 12820 (male), 13166 (fr.), Philippines, Palawan.

Tree 10-15 m. Twigs slightly angular, 1-2 mm diameter, early glabrescent, hairs less
than 0.1 mm; older twigs coarsely striate and longitudinally cracking; lenticels absent.
Leaves (membranous or) chartaceous, (elliptic-)oblong, 6-19 by 2—6 cm, base + nar-
rowly rounded or (acute-)attenuate, apex (sub)obtuse or (sub)acute; the upper surface
olivaceous; lower surface dull (purplish) grey-green, glabrous; obviously papillose; dots
absent; midrib flat or above slightly raised, lateral nerves 10-14 per side, at 45—60° to
the midrib, flat and indistinct above, indistinct below, lines of interarching and venation
indistinct or absent; petiole 9-20 by 1-2 mm; leaf bud 7 by 1—1.5 mm, hairs 0.1 mm or
less. Inflorescences between the leaves or below, pedunculate, unbranched or nearly so,

De Wilde — Myristicaceae (Myristica) 601

early glabrescent, indumentum minute; bracts 0.2 mm, caducous; in male: peduncle slen-
der, 4—10(—20) by 1 mm, with a subumbel of 6-15 flowers, buds nearly equal in size; in
female: peduncle (0—)2—4 mm long, 2—5-flowered; flowers with (scattered) pale grey-
brown appressed hairs less than 0.1 mm long. Male flowers: pedicel slender, 7-10 mm,
bracteole 1 mm long, early caducous; buds narrowly ovoid, 5—5.5 by 3-3.5 mm, apex
narrowly rounded, cleft 1/4(—1/3), lobes 0.2 mm thick. Androecium slender, 4.5—5 mm;
androphore slightly broadened towards the base, 2-2.5 by 0.5—-0.7 mm, hairs sparse,
pale brown, less than 0.1 mm; synandrium 2-3 by 0.5—0.6 mm, subacute, thecae 14—16
(10 anthers according to Sinclair, I.c.), + contiguous, sterile apex 0.1-0.3 mm. Female
flowers (Ridsdale SMHI 89): pedicel (6—)10—-13 mm long, bracteole caducous, 3-10 mm
below the apex; buds ovoid-oblong, narrowed towards the apex, 5-7 by 4-5 mm, cleft
c. 1/4, lobes 0.2—0.3 mm thick; ovary ovoid, 3 by 2.5 mm, hairs 0.1 mm or less, pistil
5 mm long, including the 2 mm long stigma. Fruits 1(—3) per infructescence, nearly
sessile, + ellipsoid, 3.5—4(—5) by 2(-3) cm, minutely pubescent, glabrescent; pericarp
3(—5) mm thick, seeds not seen; fruiting pedicel slender, 8-20 mm long, bracteole scar
well below the apex. — Fig. 60c.

Field-notes — Medium-sized tree with spreading branches, the ultimate numerous,
lax, and slender, or tree with high spherical crown. Bark smooth and scaling in thin
flakes, or fissured, greyish or blackish brown; inner bark reddish brown, with exudate.
Fruits (somewhat immature) ovoid, 5 cm long, yellowish green.

Distribution — Malesia: Philippines (endemic to Palawan: Mt Pulgar and vicinity).

Habitat & Ecology — Low canopy lowland and montane rain forest (with Agathis
dammara), also on limestone and ultrabasic alluvial deposits; 50-800 m altitude;
fl. Mar.; fr. Apr., May.

Note — Myristica umbellata is similar to certain forms of M. iners, a polymorphous
species. It differs from M. iners in small umbellate inflorescences (the umbel consists of
two opposite closely set clusters of flowers; only occasionally the middle branch is slightly
developed, 1-2 mm long), in slender pedicels, subglabrous perianth, early fallen bracteole,
and subglabrous androphore, and vegetatively in the slender branches with small leaves
papillose below (not papillose in M. iners), with indistinct nerves, and almost invisible
venation. The female flowers are much larger than the male, with the bractole scar far
below the perianth.

142. Myristica umbrosa J. Sinclair

Myristica umbrosa J. Sinclair, Gard. Bull. Sing. 23 (1968) 147, f. 4; Foreman, Contr. Herb. Austral.
no. 9 (1974) 42; Handb. Fl. Papua New Guinea | (1978) 212; W.J. de Wilde, Blumea 40 (1995)
337. — Type: Carr 16410, Papua New Guinea.

Tree 5—26 m. Twigs 3—4(—5) mm diameter, (early) glabrescent, hairs 0.5(—1) mm;
older twigs brown or blackish, often longitudinally cracking; lenticels frequently small
but distinct. Leaves (thinly) coriaceous, elliptic-oblong or oblong(-lanceolate), 20—50
by 7-15 cm, base short-cuneate or rounded, apex acute-acuminate; upper surface oliva-
ceous or brown, lower surface whitish or greyish, glabrescent, hairs thin, scale-like, less
than 0.1 mm; not papillose; dots absent; midrib slightly raised above, lateral nerves 16-
20(—22) per side, at 60-80° to the midrib, flat above, brown and contrasting in colour
below, lines of interarching and venation indistinct; petiole 15—25(—30) by 3-6 mm; leaf

602 Flora Malesiana, Ser. I, Vol. 14 (2000)

bud 15—20(—30) by 3-4(—5) mm, hairs 0.3—0.5(—0.6) mm. /nflorescences pedunculate,
late glabrescent, between the lower leaves or just below, sometimes axillary to cataphylls
on 1-3 cm long short-shoots (with apical sterile buds) on older wood; in male: peduncle
10-15 by 33.5 mm, striate, ending in a subumbel (composed of two adjacent sessile
partial subumbels) of 4-6 flowers, buds nearly equal in size; bracts caducous; in female:
peduncle 5—6 mm long, 1—3-flowered; flowers thinly coriaceous, with hairs 0.5 mm
long. Male flowers: pedicel 8-10 by 1.5(—2) mm, bracteole caducous; buds (immature)
ovoid-ellipsoid, 7(-8) by 5 mm, rounded at both ends, cleft nearly 1/2, lobes 3(—4) mm
long, 0.5 mm thick. Androecium 4.5 mm; androphore 0.5(—1) by | mm, glabrous, with a
ring of densely set pale hairs 0.2 mm at base (see also note 1); synandrium 4 by 2 mm,
thecae c. 30, sterile apex broadly convex, 0.5 mm. Female flowers: see note 1. Fruits:
peduncle short, fruits solitary or 2 or 3 together, ellipsoid, 5—7 by (3-)4—4.5 cm, apex
rounded or subacute, hairs rusty, mealy, (0.3—)0.6—1 mm; pericarp S—10 mm thick; seeds
ellipsoid, 3—3.5 cm; fruiting pedicel 5 mm long, + pubescent.

Field-notes— Low or medium-sized tree, branches horizontal, crown open, stilt-roots
(once recorded). Bark grey-brown or black, smooth or with longitudinal fissures; inner
bark (blaze) brown-orange; exudate red, watery; wood reddish straw. Leaves (dull or)
glossy dark green above, (silver-)grey-green or whitish below. Flowers yellowish brown.
Fruits reddish brown or brown pubescent, with flesh-coloured mesocarp, seeds black.

Distribution — Malesia: Papua New Guinea (Morobe and Northern Prov.), a local
endemic.

Habitat & Ecology — Dryland tall lowland forest, swamp forest, hillside slope with
microphyllous vegetation, lowland ridge forest, mangrove-rain forest transition; alti-
tude 0—400 m; fl. Jan., Oct.; fr. throughout the year.

Notes — 1. Foreman (1974) described the male flowers as follows: “pedicels 9-10
mm long; bracteole large, surrounding one side of the perianth, 8 mm long, caducous,
mid-brown tomentulose; buds mid-brown tomentulose, + subglobose, 10-11 mm long;
3 acute perianth lobes about half as long as perianth; staminal column 6 mm long; stalk
2.5 by 3 mm long, brown tomentose”; and the female flowers as “pedicels 4 mm long;
perianth 8 mm long; ovary mid-brown tomentulose, c. 6 mm long.” I have not seen such
mature well-developed male flowers. The measure of the stalk of the staminal column
seems unexpectedly stout. Foreman (1978: 212) described the peduncle of the male
inflorescences as 3.5 cm long, which possibly rests on a mistake; the peduncle actually
reaches 10-15 mm, as also stated by him in his 1974 publication.

2. LAE 59184 (Manus L., low altitude) distributed under the name M. cf. umbrosa or
perhaps sp. nov., comes close to M. umbrosa but it differs in general aspect: more slen-
der twigs, thinner and smaller leaves; fruits about identical with those of M. umbrosa.
Because the collection from Manus L. is so far from the other collections of M. umbrosa,
all from a restricted area in SE Papua New Guinea, it may represent an undescribed
species, but more collections should corroborate this.

143. Myristica uncinata J. Sinclair

Myristica uncinata J. Sinclair, Gard. Bull. Sing. 23 (1968) 150, f. 5; Foreman, Handb. Fl. Papua
New Guinea 1 (1978) 212, f. 92; W.J. de Wilde, Blumea 40 (1995) 337. — Type: Carr 14907,
Papua New Guinea.

De Wilde — Myristicaceae (Myristica) 603

Tree 18-30 m. Twigs 2.5—4(—5) mm diameter, late glabrescent, hairs conspicuous,
chocolate, flossy, 1-1.5 mm long; older twigs blackish, coarsely striate; lenticels small,
scattered. Leaves thinly chartaceous, oblong(-lanceolate), 17—27(-30) by 5—9(—10.5) cm,
base cuneate, apex acute-acuminate; upper surface olivaceous-brown, lower surface pale
brownish grey, + glabrescent, hairs scattered, appressed, usually some light brown flossy
hairs, 0.5—1(—2) mm, persistent on and near the midrib towards the base; not papillose;
dots absent (but see note 2); midrib flat or raised above, lateral nerves 14—18 per side, at
45-70° to the midrib, flat, lines of interarching and venation inconspicuous; petiole 15—
20 by 2.5—4 mm; leaf bud 15—20 by 3—4 mm, curved (uncinate), hairs I(—1 .5) mm long.
Inflorescences between the leaves or below; male unknown; in female: peduncle 10-12
mm long, with at the end 1-3 flowers, with bright red-brown flossy hairs 1 mm. Male
flowers not known. Female flowers: see note |. Infructescences below the leaves: pedun-
cle 10-20 by 4(—5) mm, with at the end | or 2 scars of fallen flowers, and mostly with
a single fruit. Fruits broadly ovoid-subglobose, 6-7 by (4—)5.5 cm, the very apex
+ narrowed, often + hooked, hairs dark brown, 1-1.5 mm; pericarp 10 mm thick; seeds
ovoid, 3.5 cm long; fruiting pedicel 10 by 4 mm, bracteole scar invisible or subapical.

Field-notes — Medium-sized tree without buttresses. Bark dark brown, finely fissured,
+ rough; inner bark red-brown; sap reddish; wood whitish but staining reddish brown.
Leaves dark glossy green above, + white below. Flowers solitary, brown or cream out-
side, cream inside. Fruits solitary, + globose, tip uncinate, covered with red-brown hairs.

Distribution — Malesia: Papua New Guinea (a local endemic of the region of Boridi,
Central Prov.).

Habitat & Ecology — Lower montane forest, ridge forest; Terminalia—Sloanea for-
est; 1300—1400 m altitude; fl. & fr. Sept.—Nov.

Notes — 1. Female flowers are known from the descriptions of previous authors only,
but it is suspected that these pertain to anomalous forms. Foreman (1.c.) described the
female flowers as “pedicel c. 1 cm long, perianth ellipsoid in buds, c. 20 by 5—7 mm,
split c. half-way down into 3 lobes which become highly reflexed.” Sinclair (1.c.) de-
scribed the female inflorescence as “1- or 2-flowered, with a flattened axis c. 1 cm long,
pilose or glabrescent, with coriaceous flowers, 2 cm long (being the largest flowers in
the genus), the ovary ovoid, 5 mm diam., with bi-lobed stigma (style) 2-3 mm long; the
bracteole rigid, early deciduous, leaving a conspicuous scar.” The recent collection
LAE 60175 is annotated with “flowers solitary.” The duplicate in L has a single female
flower in the axil of an undeveloped leaf, there is no peduncle and no bracteole scar on
the pedicel; however, the single-fruited (immature) infructescences in the same collec-
tion do have a peduncle, 1 cm long, but no bracteole scar on the fruiting pedicel. The
flower has a woody consistency and the ovary is 5 by 3.5—4 mm, with densely pilose
bright (red-)brown hairs 1-2 mm, partly agreeing with the description as given by Sinclair
(see above). Possibly these extraordinary large flowers, without a bracteole, may be
(partly) diseased or gallous, and normal flowers may be smaller, with a bracteole. The
fruiting pedicel of Carr 14908 (at K) has a distinct bracteole scar 2 mm below the fruit.
As stated, male inflorescences and male flowers are unknown.

2. Spot characters are the usually uncinate apical leaf bud, and the uncinate apex of
the fruits. The leaves are characteristically whitish below. Dots on the lower leaf surface
are absent, but blackish dots are present on the upper surface; the exact nature of these
dots (traumatic or not) remains to be asserted.

604 Flora Malesiana, Ser. I, Vol. 14 (2000)

144. Myristica undulatifolia J. Sinclair

Myristica undulatifolia J. Sinclair, Gard. Bull. Sing. 23 (1968) 400, f. 67; W.J. de Wilde, Blumea 40
(1995) 337. — Type: NGF 7173 (Gray & Floyd), Papua New Guinea, Gulf Prov.

Tree 6—30 m. Twigs not or but faintly angled at the very apex, 2-3(—4) mm diameter,
early glabrescent, hairs 0.1 mm; older twigs dark brown, longitudinally fissured, cracked,
or thinly flaking; lenticels small, pale, contrasting. Leaves membranous or thinly charta-
ceous, oblong-lanceolate, 12—26 by 4-9.5 cm, base cuneate or (narrowly) rounded, apex
acute-acuminate; upper surface olivaceous or brown, lower surface grey-brown or sil-
very, seemingly glabrous, hairs felty, dense, pale, scale-like, less than 0.1 mm, some-
times late glabrescent; not papillose; dots absent; midrib slightly raised above, lateral
nerves 15-25 per side, at 45—80° to the midrib, flat or sunken above, reddish brown and
contrasting in colour below, lines of interarching and venation inconspicuous; petiole
15-20 by 2(-3) mm; leaf bud 10-15 by 2 mm, hairs scale-like, 0.1 mm or less. /nflores-
cences between the lower leaves or below, of the Knema-type: a (sub)sessile (to 2 mm
pedunculate) simple or forked, scar-covered, pubescent brachyblast 3(—5) mm long; in
male: with a subumbel of (5—)10—15 flowers, buds of various sizes; bracts small, caducous;
in female: (3—)4—6-flowered; flowers with hairs 0.1—0.2 mm. Male flowers: pedicel 4—5
by 0.6—-0.8 mm, bracteole + boat-shaped, 2.5 by 1.5(—2) mm, persistent; buds obovoid-
oblong, 4(—5) by 2 mm, cleft c. 1/4, lobes 1 mm long, 0.1—0.2 mm thick. Androecium
0.5—0.6 mm; androphore (1—)1.2 by 0.4 mm, with hairs less than 0.1 mm in the lower
half; synandrium narrowed at apex, 2 by 0.5—0.6 mm, thecae 14—16(—20), sterile apex
acute, (0.2—)0.3 mm. Female flowers: pedicel 1.5 mm, bracteole 3.5 mm long; buds ovoid,
5 by 4 mm, the lobes 1.5 mm; ovary ovoid, c. 3 mm long, hairs minute, dense, appres-
sed, pale brown. Fruits solitary or 2 per infructescence, (sub)sessile, ovoid-oblong, 2.5—
3.5 by 1.7—2.5 cm, base rounded or subtruncate; hairs dense, very fine, grey-brown or
yellowish brown, 0.1 mm or less, old fruits + glabrescent, pericarp 5 mm thick; seeds
ellipsoid, (1.5—)2 cm; fruiting pedicel stout, 3-5 by 4—6 mm, rough by cracks and few
lenticels.

Field-notes — Locally tall, canopy-forming trees. Outer bark rough or fissured, not
or somewhat peeling; inner bark red; exudate pinkish brown; sapwood cream or brown-
ish pink. Leaves glossy light brown or silvery-rusty below. Flowers (buds) brown. Fruits
pale brown or greenish brown.

Distribution — Malesia: southern part of West and East New Guinea (also Bird’s
Head).

Habitat & Ecology — Primary and degraded lowland and lower submontane forest;
flatland and ridges, mixed Castanopsis forest; moreover in forest inundated in the wet
season, forest on clay soil by streams; 100—600 m altitude; fl. Mar., Apr.; fr. Apr., July—
Sept.

Notes — 1. In some specimens the thinly chartaceous leaves dry with a more or less
undulate margin, giving the leaves sometimes a denticulate appearance; in reality the
leaves never are denticulate.

2. The specimen von Rémer 96 (southern West New Guinea) has the bark of the older
twigs as in M. undulatifolia, but its very immature fruits have longer and more slender
pedicels; possibly it could be referred to M. fatua or represents a separate, yet undescribed,
new taxon.

De Wilde — Myristicaceae (Myristica) 605

145. Myristica velutina Markgr.

Myristica velutina Markgr., Bot. Jahrb. Syst. 67 (1935) 165; W. J. de Wilde, Blumea 40 (1995) 339.
— Type: Ledermann 11175 (B, lost); paratypes: Ledermann 11333a, 11395a, 12802a (B, lost)
(Ledermann 12802a, L, lecto, here designated), Papua New Guinea.

Tree 4-10 m. Twigs subterete or bluntly (tri)angular, or mostly with two sharp lines or
ridges between the petioles, (2—)3(—4) mm diameter, ant-swellings absent, early gla-
brescent, hairs 0.1 mm or less; older twigs with non-contrasting lenticels. Leaves charta-
ceous or thinly coriaceous, (obovate-elliptic-)oblong or oblong-lanceolate, 10-23 by
2.5-8 cm, base attenuate or narrowly rounded, apex acute-acuminate; upper surface oliva-
ceous or brown, lower surface grey(-brown), late glabrescent or indumentum (sub)per-
sistent, hairs flocculose, + scattered, 0.1(—0.3) mm; not distinctly papillose; dots absent;
midrib slightly raised above, lateral nerves (15—)18—22(—25) per side, at 60—80° to the
midrib, flat or sunken above, lines of interarching rather distinct, venation indistinct;
petiole 8-15 by 2-4 mm; leaf bud 10-15 by 2 mm, hairs grey-brown, 0.1 mm. /nflores-
cences between the leaves, + of the Knema-type: a (sub)sessile or to 4 mm pedunculate
scar-covered knob-like brachyblast up to 5 mm long, + glabrescent; bracts | mm, cadu-
cous: in male: with a subumbel of 3—6 flowers, with buds of + variable sizes; female as
male, buds up to 8 in a cluster, subequal in size; flowers with hairs 0.1 mm. Male flow-
ers: pedicel 4-5 mm long, bracteole persistent; buds ellipsoid-oblong or oblong-lanceo-
late, 6 or 10 mm long; see further under the subspecies. Female flowers (Kalkman a2):
pedicel 2-3 mm, bracteole minute, + caducous; buds ovoid(-oblong), 4.5 by 3 mm, lobes
1.5-2 mm; ovary ovoid, including a 3.5 mm long stigma, short-pubescent. Fruits soli-
tary or in clusters of up to 5, ovoid(-oblong), 2.5 by 1.5(-2) mm, apex + acute, often
narrowed into 2 mm long style-remnant, base rounded or sometimes shortly narrowed,
hairs persistent, mealy, rusty, 0.1(—0.2) mm; pericarp 2-3 mm thick; seeds ellipsoid,
1.5(-2) cm; fruiting pedicel 3—5(—10) by 1.52.5 mm, often thickened by numerous ir-
regular wart-like lenticels, bracteole scar apical.

Distribution — Malesia: mountains of W Central Papua New Guinea; there are two
subspecies.

Note — Resembling M. subalulata but differs in the absence of ant-swellings in the
twigs, in the short male flower pedicels, and persistent bracteole. Myristica velutina is
fairly homogeneous in habit, leaf-texture and colour, and general features of the twigs.
However, an analysis of the male flowers of Ledermann 1 2802a (in L) and those of the
later collections, show rather pregnant differences, a reason to accept two subspecies.
Possibly the description by Markgraf (1.c.) was based on specimens belonging to the
present subspecies velutina, testified e. g. by the mentioned long range in length of the
perianth, 6-9 mm. The fruits were described (Ledermann 11395a) as globose, diameter
2.5 cm, and possibly fresh fruits are proportionally broader than those found in dried
collections.

KEY TO THE SUBSPECIES

la. Twigs faintly lined or + angular. Male buds (8—)10 mm long. — E Sepik, Western &
Southeniehlands, Brov.. 6 2. one eeiiws cst) aie firs Seg a. subsp. velutina

b. Twigs sharply 2-lined or low-ridged. Male buds (5—)6 mm long. —W Sepik & Western
ECO VAR iiss ee Ee Py, Sel rs ON Oo en cs 'n re b. subsp. breviflora

606 Flora Malesiana, Ser. I, Vol. 14 (2000)

a. subsp. velutina

Twigs (sub)angular or with faint lines. Male flowers: pedicel slender, 4-5 by 0.5(-1)
mm, bracteole ovate(-oblong), 3 mm long, faintly 3-topped, persistent; buds oblong-
lanceolate, (8—)10 by 2.5(—3) mm, apex subacute, not angular, base narrowly rounded,
cleft c. 1/5, lobes 2 mm long, 0.2(—0.3) mm thick. Androecium slender, 8-9 mm;
androphore much shorter than the synandrium, 3 by 0.5 mm, almost completely set with
sparse minute appressed red-brown hairs less than 0.1 mm; synandrium fusiform, 5 by
0.8—1 mm, thecae c. 10, sterile apex slender, acute, | mm long. Fruits as the species.

Distribution — Malesia: Central Papua New Guinea (East Sepik, Western Highlands
& Southern Highlands Prov.).

Habitat & Ecology — Mid-mountain forest; 1300-2000 m altitude; fl. & fr. through-
out the year.

Note — The pale, grey(-white) lower leaf surface (young leaves may be more brown-
ish below because of indumentum) seems to be caused by a very dense whitish papillation,
so crowded that the leaf surface appears to be non-papillose, also as seen with magni-
fication. In Jacobs 8808 the lower surface is distinctly (densely) papillose.

b. subsp. breviflora W. J. de Wilde

Myristica velutina Markgr. subsp. breviflora W.J. de Wilde, Blumea 40 (1995) 339. — Type: NGF
41677 (Henty, Isgar & Galore), Papua New Guinea.

Twigs distinctly angular, with two marked lines or low ridges running in between the
petioles or the scars of petioles. Male flowers: pedicel 4 by | mm, bracteole ovate, rounded,
1.5 mm long, persistent; buds ellipsoid-oblong, (5—)6 by 2.5 mm, apex and base + rounded,
not angled, cleft c. 1/4, lobes 1.5 mm long, 0.3 mm thick. Androecium slender, 5—5.5
mm; androphore nearly as long as the synandrium, 2—2.2 by 0.6—0.7 mm, almost com-
pletely pubescent, hairs pale, scattered, less than 0.1 mm; synandrium 3 by 0.7—0.8 mm,
thecae 10-12, sterile apex + blunt, 0.3 mm. Fruits as the species.

Field-notes — Small tree, 4-10 m. Branches short, horizontal. Outer bark dark brown
or reddish brown; inner bark pink or yellow; latex red; wood straw, with (little) red latex,
or staining reddish after exposure. Leaves whitish below. Flowers brown-cream. Fruits
orange-brown or deep brown, seeds blackish.

Distribution — Malesia: W Central Papua New Guinea (West Sepik and Western
Prov.).

Habitat & Ecology — Mountain forest; forest on ridges, on latosol, fagaceous moss
forest, Nothofagus forest on swampy site; also on limestone or on well-drained old vol-
canic soil; 1500-2100 m altitude; fl. & fr. throughout the year.

Fig. 91. Myristica verruculosa W.J. de Wilde. a. Leafy twig with male inflorescences; b. leaf;
c. male bud, opened, note the irregularly warted lower inside, (sub)persistent bracteole drawn sepa-
rately; d. male inflorescence with one flower blooming; e. female bud, opened, showing pistil;
f. twig with infructescence [a—d: Polak 1283; e, f: Polak 697). — Scale bar for a, d, f = 2 cm; for
b =4 cm; for c, e = 1.65 mm.

De Wilde — Myristicaceae (Myristica)

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608 Flora Malesiana, Ser. I, Vol. 14 (2000)

146. Myristica verruculosa W. J. de Wilde

Myristica verruculosa W.J. de Wilde, Blumea 43 (1998) 177, f. 4. — Type: Polak MP 1283, Irian
Jaya, Bird’s Head.

Tree 15 m. Twigs 4—5 mm diameter, subterete with faint lines from petiole to petiole,
light brown, partially hollow with incidental ant-holes, without swellings, early glabres-
cent, hairs minute, bright brown, less than 0.1 mm; older twigs striate with scattered
lenticels. Leaves chartaceous, (elliptic-)oblong, 20-32 by 8-13 cm, base broadly rounded
or shallowly cordate, apex acute(-acuminate); upper surface olivaceous, lower surface
with persistent indumentum, hairs dense, pale, 0.1 mm or less; not papillose; dots ab-
sent; midrib slightly raised above, lateral nerves 20—23 per side, at 60—80° to the midrib,
flat or sunken above, lines of interarching and venation indistinct; petiole 20-40 by
5 mm: leaf bud slender, 15—20 mm, hairs dense, brown, stellate, 0.1 mm or less. /nflores-
cences between the leaves or below, of the Knema-type: a simple or forked brachyblast,
to 10 mm long, with stout peduncle to 5 mm, glabrescent; in male: with clusters of 10
(—20) flowers, buds of various sizes; bracts minute, caducous; in female: with sessile
clusters, 5—10-flowered:; flowers with yellow brown mealy hairs 0.1(—0.2) mm. Male
flowers: stout, pedicel 10-11 by 1.5 mm, bracteole ovate-broad-triangular or rhomboid,
acute, 4—4.5 mm, (sub)persistent; buds oblong, (10—)12 by 4-5 mm, apex + blunt to
acute, base somewhat tapered, cleft 1/3, lobes + fleshy, 4 mm long, 0.5(—0.8) mm thick,
perianth inside for the lower 1/3 finely iregularly warted. Androecium (8—)10 mm long;
androphore slender, 5—6 by 0.8 mm, in lower half with pale hairs less than 0.1 mm;
synandrium oblong, subacute, 4—5 by 1.5—2 mm, thecae c. 17, sterile apex glabrous,
+ acute, 0.2—-0.3 mm. Female flowers: pedicel 2(-2.5) mm long, bracteole ovate, + hooded,
2.5 mm long, (sub)persistent; buds ovoid(-oblong), narrowed in the apical part, 8 by
4—5 mm, lobes 1.5 mm; ovary ovoid, tapering, (including 1.5 mm long stigma lobes)
5 by 3 mm, hairs appressed 0.1(—0.2) mm. Fruits in sessile clusters of 1-3, (ovoid-)
oblong, 3.5—5 by 2.5 cm, apex acute, base broadly rounded, hairs rusty, 0.5 mm; pericarp
5 mm thick; seeds ellipsoid-oblong, 2.5—3 cm; fruiting pedicel 1-3 mm. — Fig. 91.

Field-notes — Exudate clear. Leaves pale brown below. Flowers yellow brown, | cm
long.

Distribution — Malesia: Papua Barat (Bird’s Head: E of Sorong, and Bombarai Pe-
ninsula).

Habitat & Ecology — Limestone area; in degraded forest on clay soil; 450 m altitude.

Note — Related and similar to M. bialata, M. fissiflora, M. hollrungii, M. ingrata,
and M. subcordata, and especially close to M. bialata, a species of the Bismarck Archi-
pelago; M. verruculosa is distinguishable by the fine warty structure within the perianth.
A warty inner perianth, however, is shared with M. fissiflora, but that species differs in
its deeper cleft male perianth. Myristica ingrata usually has ant-holes in the twigs, a gla-
brescent lower leaf surface, and much smaller flowers (except var. velata); M. hollrungit
grows in swampy areas, and equally has smaller flowers and (sub)glabrous leaves; M.
subcordata, with similar persistent leaf indumentum, has smaller and more narrow male
perianths, with a caducous bracteole; M. bialata has similarly stout male flowers, but
differs in the lower number (10 or 12) of thecae (16 or 18 in M. verruculosa), shorter
perianth lobes, the (almost) absence of a warty inner surface, and it has a glabrescent
lower leaf surface (densely short-felty in the present species).

De Wilde — Myristicaceae (Myristica) 609

147. Myristica villosa Warb.

Myristica villosa Warb., Mon. Myrist. ( 1897) 419, t. 14; J. Sinclair, Gard. Bull. Sing. 16 (1958) 356,
in obs.: 23 (1968) 311, f. 41; W.J. de Wilde, Blumea 42 (1995) 188; Tree Fl. Sabah & Sarawak
3 (2000) 471. — Lectotype: Beccari 1526, Sarawak.

Myristica fatua auct. non Houtt.: Mig., Ann. Mus. Bot. Lugd.-Bat. 2 (1865) 46 (quoad spec. born.
de Vriese).

Myristica borneensis Gand.. Bull. Soc. Bot. France 66 (1919) 225, in clavi (non Warb., 1897). —
Type: Korthals s.n., Borneo.

Tree (5—)10—40 m. Twigs 5-10 mm diameter, glabrescent, hairs villose, (1-)2—3 mm
long; older twigs brown or blackish, coarsely longitudinally cracking and flaking; lenticels
absent. Leaves chartaceous or thinly coriaceous, (elliptic-)oblong, (15—)20—40 by 6-13
cm, base acute or narrowly rounded, apex acute(-acuminate); upper surface olivaceous
or brown, lower surface late glabrescent or hairs persistent, dense, grey or cinnamon,
scale-like (dendroid), 0.1—0.2 mm, mixed with widely spaced unbranched uniseriate
emergent hairs 1-3 mm; not obviously papillose; dots absent; midrib + raised above,
lateral nerves 20-25 per side, at 45—60° to the midrib, sunken, lines of interarching and
venation distinct; petiole (12-)20-25 by 3.5—5 mm, late glabrescent; leaf bud 7-12 by
4-6 mm, hairs conspicuous, (1—)2-3 mm. Inflorescences between the leaves or below,
of the Knema-type: a sessile scar-covered brachyblast, pubescent; bracts ovoid or rounded,
3-4 mm, caducous; in male: simple or 2—4-furcate, 5-13 by 5-10 mm, terminally with
1-3 subsessile flowers, buds subequal in size; in female: similar, shorter, with | or 2
flowers + hidden in coarse indumentum; flowers coriaceous, with villose hairs 1-2 mm
long. Male flowers: pedicel rather thick, 3—4(—5) mm long, bracteole broadly rounded,
3-4 mm, late caducous; buds ovoid(-ellipsoid), 6-8 by 4-5(-6) mm, apex narrowly
rounded, cleft c. 2/3, lobes 1 mm thick. Androecium slender, 4-5 mm long; androphore
(0.5—)1 by 0.4—0.5 mm, at base with a collar of pale hairs | mm; synandrium 3.5—4.5 by
0.6 mm, thecae c. 12, sterile apex + blunt (0.2—)0.3—0.5 mm long. Female flowers: pedi-
cel 1 mm long, bracteole broadly rounded, 4-5 mm, + caducous; buds ovoid, 8(—10) by
(5—)6-8 mm, cleft c. 2/3; ovary (broadly) ovoid, 5—6 by 4 mm, hairs 1-1.5 mm; stigma
1(-1.5) mm. Fruits 1 or 2 per infructescence, subsessile, broadly (ovoid-)ellipsoid, apex
rounded or narrowed with somewhat uncinate apex, 4-6(—8) by 2.5—4(—4.5) cm, hairs
dense, pale brown or rust-coloured, somewhat hispid, 2—3(—4) mm; pericarp 3—6(—8)
mm thick: seeds broad-ellipsoid, 3—5 cm, aril coarsely laciniate, the laciniae finely di-
vided at the apex; fruiting pedicel up to 5 mm long. — Fig. 61c, 92.

Field-notes — Usually without buttresses, or with stilt-roots at base. Bark blackish,
hard, rough, flaking or longitudinally furrowed (as in Knema hookeriana and related
species), strips (1.5—-)2—3 mm wide and long, | mm thick, sometimes smooth (finely
scaly); underbark reddish brown; outer bark pinkish, laminated, soft, 5-10 mm thick;
sapwood cream or pale brown, with odour of coconut; sap red. Leaves glossy above,
silvery (brown), in old leaves glaucous below. Flowers yellow-brown, inside pale yel-
low, fragrant. Fruits yellow- or apricot-brown pubescent.

Distribution — Malesia: throughout Borneo.

Habitat & Ecology — Primary dry land (mixed dipterocarp) forest, sometimes marsh
forest; also in kerangas; on flatland, hillsides, and ridges; mostly on sand or loam soils,
sandstone derived soil, also on heavy yellow clay (Setap Shale); 20-1200 m altitude;
fl. Feb., June, Aug.—Sept., fr. May—Sept.

610 Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Myristica) 611

Note — A homogeneous species, distinguishable by the subpersistent conspicuous
indumentum of lower leaf surface, midrib (also above), twig apex, and flowers, by the
flaking dark brown or blackish bark of older twigs, by the sessile fatua-type of inflores-
cences, and by the + asymmetrical coarsely hairy fruits.

148. Myristica vinkeana W. J. de Wilde
Myristica vinkeana W.J. de Wilde, Blumea 40 (1995) 339. — Type: Eyma 4283, Irian Jaya.

Tree 17 m tall. Twigs 1.5—2(—2.5) mm diameter, glabrescent, hairs 0.1 mm; older twigs
(coarsely) striate, not flaking; lenticels small, inconspicuous. Leaves thinly coriaceous,
elliptic-oblong, 8-15 by 2—4.5 cm, base (short-)attenuate, apex acute or (long-)acumi-
nate; upper surface dark (blackish) brown, lower surface grey-brown, glabrescent, hairs
inconspicuous, widely spaced, grey or cinnamon brown, scale-like, 0.1 mm or less; not
papillose; dots absent; midrib flat or in a groove (and slightly raised) above, lateral
nerves 16-18 per side, at 60-80° to the midrib, flat or sunken, lines of interarching and
venation faint or distinct and contrasting below; petiole (6-)10—15 by 1.5 mm; leaf bud
10 by 1 mm, hairs grey-brown, 0.1 mm. /nflorescences between the lower leaves, with
dense hairs 0.1 mm; of the Knema-type: a sessile scar-covered brachyblast, less than
1 mm long; in male: with clusters of 1-3(—5) flowers, buds of various sizes; bracts ca-
ducous: in female (from infructescences): 1(—3)-flowered; flowers with hairs 0.1 mm.
Male flowers (Eyma 4283): pedicel slender, 3 mm long, bracteole comparatively large,
broadly ovate, 2-2.5 mm long, apex acute, (sub)persistent; buds ovoid(-ellipsoid) to
oblong, 5(—5.5) by 2-2.5 mm, apex + blunt, cleft c. 1/3, lobes 1.5—1.7 mm long, 0.2—0.3
mm thick. Androecium slender, 4(—4.5) mm long; androphore 2 by 0.5 mm, longitudi-
nally + ribbed, almost entirely pubescent, hairs pale brown, 0.2—0.3 mm; synandrium
2 by 0.8—0.9 mm, thecae c. 12, the sterile apex irregularly + blunt, 0.3 mm long. Female
flowers not seen. Fruits (immature) sessile, solitary, just below the leaves, + (ovoid-)
ellipsoid, 3 by 1.5 cm, apex somewhat acute, base shortly narrowed; hairs dense, grey-
brown, scale-like, 0.1 mm; pericarp 3 mm thick; seeds not seen; fruiting pedicel 10 by
2.5 mm, lenticellate, bracteole scar about median.

Field-notes — Rather common. Inner bark red. Fruits (immature) light brown.

Distribution — Malesia: Papua Barat (Wissel Lake region).

Habitat & Ecology — Mountain forest on stony sandy-clayey soil; 1600-1800 m
altitude; fl. Jan.; fr. May.

Note — Close to M. crassipes, which differs in its larger male flowers with (sub)cadu-
cous bracteole, and less hairy androphore.

Fig. 92. Myristica villosa Warb. a. Leafy twig with fruit; b. older twig showing blackish bark, con-
spicuously cracking; c. apex of leafy twig with few-flowered male inflorescences arranged dis-
tichously; d. opened male bud showing androecium beside the detached caducous bracteole;
e. opened female bud showing pistil beside the detached caducous bracteole; f. twig with
infructescence with a single ripe fruit; g. ditto, opened, showing arillate seed [a: S 39444; b, f, g:
Mogea & de Wilde 3502; c: Anderson & Hou 488, d: Paijmans 33; e: § 29211). — Scale bar for
a—c, f, g =2 cm; ford,e =2 mm.

Flora Malesiana, Ser. I, Vol. 14 (2000)

De Wilde — Myristicaceae (Myristica) 613

149. Myristica warburgii K. Schum.

Myristica warburgii K. Schum. in K. Schum. & Lauterb., Nachtr. Fl. Schutzgeb. Siidsee (1905) 267;
WJ. de Wilde, Blumea 40 (1995) 341, f. 9. — Type: Biré 21, Papua New Guinea, Sattelberg.

Tree 3-20(—25) m. Twigs usually markedly (2- or) 3-angular, |—2.5(—3) mm diameter,
early glabrescent, hairs 0.1(—0.2) mm or less; older twigs with inconspicuous lenticels.
Leaves membranous or thinly coriaceous, elliptic(-oblong) or oblong-lanceolate, base
attenuate, apex acute(-acuminate), 6-16 by (1.5—)2.5—5.5 cm; upper surface olivaceous
or (bright) brown, lower surface slightly paler, early glabrescent, hairs weak, scattered,
0.1 mm or less; not papillose; dots absent (minute dots sometimes present in young
leaves, see note | under subsp. warburgii); midrib raised above, lateral nerves 10—20 per
side, at (45—)60—-70(—80)° to the midrib, flat or sunken and inconspicuous above, little
raised below, lines of interarching and venation distinct or faint; petiole 7-15 by 1-2
mm: leaf bud 7-10 by 1.5-2 mm, hairs (0.2—)0.3—0.4 mm. Inflorescences between the
leaves or below, of the Knema-type: a sessile or to 1(—2) mm pedunculate simple or
forked scar-covered brachyblast to 4 mm long, late glabrescent (hairs 0.3 mm), in male:
subumbel of 3—6 flowers, buds of various sizes; bracts minute; in female: with 1-4 flow-
ers, almost equal in size; flowers often partially glabrescent or subglabrous, hairs 0.1
mm or less. Male flowers: pedicel slender, S—10 by 0.5—1 mm, bracteole + ovate, 1—-1.5
mm, (persistent or) caducous, apical or to 2.5 mm below apex; buds oblong-lanceolate,
+ fusiform (with base attenuate) or tubiform, the upper 1/3 narrowed, acute, + angled,
8-14 by 1.5-3 mm, cleft c. 1/8, lobes 1-2 mm long, 0.2-0.3 mm thick. Androecium
slender, 5.5—9 mm long; androphore (2—)2.5—4.5 by 0.5—-0.7 mm, hairs weak, glossy,
0.2-0.3 mm long; synandrium 3.5—5S by 0.7—0.8(-1) mm, thecae 12-16, sterile apex
+ acute or blunt, 0.1—0.5 mm long, or absent. Female flowers: pedicel 9-15 by 0.8-1.5
mm, bracteole scar 3-6 mm below apex; buds ovoid-oblong, narrowed in the upper part,
7.59 by 3.5—4 mm, lobes | mm; ovary ovoid-oblong, 6 by 2-3 mm, hairs 0.2—0.3 mm.
Fruits solitary or 2 or 3 together, long-stiped, often axillary to leaves, broadly ellipsoid,
sometimes fusiform, (1.5—)2—4 by 1.5-2.5 cm, apex rounded or narrowed, often some-
what acute with up to 2 mm long beak, base tapered or + rounded and contracted into
2-5(-7) mm long pseudostalk, hairs dense, mealy, brown, 0.1—0.2 mm or less; pericarp
1.5-3 mm thick; seeds ellipsoid, (1.5—)2 cm, attached (up to c. 5 mm) above the base of
the fruit rendering seeds and aril lobes generally in a transverse, not erect, position (see
note 2 under subsp. warburgii); fruiting pedicel slender, 10—20(—25) by 1.5—3 mm, some-
times with few lenticels, bracteole scar at about 1/4 to halfway from apex. — Fig. 93.

Distribution — Malesia: Central and East New Guinea; there are 3 subspecies.

Note — Most specimens of the present species were included by Sinclair (1968) in
M. longipes. However, M. warburgii is more closely related to M. flosculosa and M.
laevifolia, as also appears from the keys. Myristica laevifolia differs in its more coriaceous

Fig. 93. Myristica warburgii K. Schum. subsp. warburgii. a. Male flowering twig; b. immature
male bud, opened (perianth considerably longer at anthesis); c. leafy twig with one-fruited in-
fructescence; d. opened fruit, showing seed with obliquely or transversely situated aril lobes. —
M. warburgii subsp. siphonantha W.J. de Wilde. e. Part of twig with male inflorescences; f. opened
male bud: note scar of caducous bracteole [a, b: BW 28/5 (Schram); c, d: Kostermans & Soegeng
336: e, f: Sands 1048]. — Scale bar for a, c, e = 2 cm; for b, f = 1.65 mm; for d= 6.5 mm.

614 Flora Malesiana, Ser. I, Vol. 14 (2000)

leaves, smaller hairs on the leaf bud, smaller male flowers, more elongate, glabrescent
fruits, and occurs generally at higher altitudes, 1000-1800 m. Myristica flosculosa dif-
fers in e.g. leaf bud with hairs 0.1 mm or less and smaller male flowers.

KEY TO THE SUBSPECIES

la. Nerves and venation on lower leaf surface usually faint, not much contrasting in

Glo Meee. ee os oo ee ite e Phe ee ele es aes Soe 2
b. Nerves and venation on lower leaf surface distinct, much contrasting in colour. Fruits
2-3 cm long. — E Papua New Guinea, atc. 1000m ......... b. subsp. hybrida

2a. Male buds fusiform or long-urceolate, (8—)9—10.5 by 2.5—3 mm; androecium 5.5—
7.5 mm long, sterile apex 0.2 mm or less, or absent. Fruits 3-4 cm long. — E Papua
New Guinea arS0= | S00 ee oie sie trae clan no area a. subsp. warburgii

b. Male buds narrowly + tubiform, (10—)13—14 by 1.5—2 mm, narrowed in the upper
and basal 1/3: androecium 8—9 mm long, sterile apex 0.5 mm long. Fruits 2.5—3 cm
long. — Northern New Guinea, at 15-400 m ........... c. subsp. siphonantha

a. subsp. warburgii

Nerves and venation on lower leaf surface faint, not much contrasting in colour. Male
flowers: buds fusiform-urceolate, (8—)9-10.5 by 2.5-3 mm; androecium 5.5-7.5 mm
long, sterile apex to 0.2 mm long. Female flowers: pedicel 10-15 by | mm, bracteole
scar about median; buds 7.5—9 by 4 mm; ovary 6 by 2.5(—3) mm. Fruits ellipsoid-fusi-
form, 3-4 cm long, hairs 0.1—0.2 mm, pericarp 2-3 mm thick; fruiting pedicel 10-20
(-25) by 1.5-3 mm. — Fig. 93a-d.

Field-notes — Medium-sized or small tree. Bark smooth or fissured and flaky, dark
brown: inner bark red-brown; exudate little, (yellow) reddish; wood straw, sometimes
rust-red on exposure, with + thick reddish exudate; heartwood dark pink. Leaves dull or
glossy dark green above, dull mid-green below. Flowers cream(-green) or (pale) yellow;
male flowers pendent, female flowers cone-shaped, 10 mm long. Fruits golden brown or
orange-brown. Fruiting in the upper crown, flowering in the lower crown.

Distribution — Malesia: E Papua New Guinea (Morobe, Central, and Milne Bay
Prov., incl. Fergusson I.).

Habitat & Ecology — Hill and lower montane forest, Castanopsis (oak) forest; on
steep slopes; 350-1300 m altitude; fl. June, July, Dec.; fr. throughout the year.

Notes — 1. Immature or just mature leaves may have a fine dark punctation below
(lens!); however, these tiny dots are markedly smaller than, but probably essentially
similar to those found in species with a dotted lower leaf surface.

2. The seed is positioned somewhat obliquely within the pericarp, that is, attached not
basally but higher up; hence the slips of the aril are largely orientated transversely rather
than lengthwise. This has been seen also in the fully mature fruits of a specimen of
M. scripta (Jacobs 9059) and probably it occurs in other species as well.

b. subsp. hybrida W.J. de Wilde

Myristica warburgii K. Schum. subsp. hybrida W.J. de Wilde, Blumea 40 (1995) 341.—Type: Jacobs
9396, Papua New Guinea.

De Wilde — Myristicaceae (Myristica) 615

Nerves and venation on lower leaf surface contrasting in colour. Male flowers not
known. Female flowers (Jacobs 9396): pedicel 9-10 by 1-1.5 mm, bracteole scar (3—)
3.5-4 mm below the apex; buds 9 by 3.5 mm; ovary 6 by 2 mm. Fruits (broadly) ellip-
soid, (1.5—)2-3 cm long, apex 2 mm beaked, base with 2 mm long pseudostalk, hairs
mealy, light rust-coloured, 0.1 mm; pericarp 1.5—-2 mm thick; fruiting pedicel slender,
11-15 by 1.5—2.5 mm.

Field-notes — Leaves pale green below. Flowers white. Fruits (dull) orange-brown.

Distribution — Malesia: Papua New Guinea (Eastern Highlands Prov., possibly also
Sepik Prov.).

Habitat & Ecology — Mixed primary montane forest; mostly on ridges; well-drained
volcanic soil; c. 1000 m altitude; fl. & fr. Sept., Oct.

Note — Subsp. hybrida belongs with some doubt to M. warburgii, whereas it also
may be related to M. gracilipes. Male flowers of subsp. hybrida are unknown, and the
seed of the type specimen is not yet fully developed. The subspecies differs mainly in
the distinct, contrasting nervation on the lower leaf surface.

c. subsp. siphonantha W. J. de Wilde

Myristica warburgii K.Schum. subsp. siphonantha W.J. de Wilde, Blumea 40 (1995) 341, f. 9e, f.
— Type: Sands 1048, Papua New Guinea.

Nerves and venation on lower leaf surface faint. Male flowers: pedicel 10 mm, bracteole
minute, 1 mm, caducous, scar 1-3 mm below the apex; buds + tubiform, ( 10—)13—14 by
1.52 mm, narrowed in the apical and basal 1/3; androecium 8—9 mm long, sterile apex
0.5 mm long. Female flowers not known. Fruits ellipsoid(-fusiform), 2.5—3 cm long,
hairs 0.1—0.2 mm; pericarp 2-3 mm thick; fruiting pedicel 10-20 by 2 mm, bracteole
scar 1-3 mm below apex. — Fig. 93e, f.

Field-notes — Tree to 10 m tal. Bark crisp, rough, rusty, smooth or fissured, or peel-
ing off 1-2 mm thick; living bark 8 mm, dark brown, sap red, or orange; underbark
crimson, spongy; inner bark exuding red sap darkening on exposure; wood smooth, pale
(pinkish) straw, darkening on exposure. Leaves pale below. Flowers apricot cream, or
creamy green with brown or yellow indumentum. Fruits (pale) brown, seeds dark brown.

Distribution — Malesia: northern New Guinea (Papua Barat: in the area of Jayapura;
Papua New Guinea: W Sepik Prov.).

Habitat & Ecology — Locally rather common in sloping terrain, steep slopes; under-
storey treelet on sandy clay or heavy clay-loam, also on limestone hills; 15—390 m alti-
tude; fl. & fr. Mar., Aug.

Note — This subspecies may resemble M. /ancifolia (with a similar faint venation),
but differs in smaller fruits, with the seeds not transversely positioned, and the leaves
papillose below. The male flowers of subsp. siphonantha resemble those of M. cornuti-
flora, a species with a stalked inflorescence and dotted lower leaf surface.

150. Myristica wenzelii Merr.

Myristica wenzelii Merr., Philipp. J. Sci., Bot. 10 (1915) 270; Enum. Philipp. Flow. Pl. 2 (1923)
180: W.J. de Wilde, Blumea 42 (1997) 188. — Myristica fatua Houtt. var. wenzelii (Merr.)
J. Sinclair, Gard. Bull. Sing. 23 (1968) 309, f. 40. — Type: Wenzel 1152, Philippines, Leyte.

616 Flora Malesiana, Ser. I, Vol. 14 (2000)

Tree 10—15 m. Twigs 4-7 mm diameter, early glabrescent, hairs 0.1 mm; older twigs
pale grey-brown, longitudinally coarsely and irregularly warty-striate; remotely lenti-
cellate or not. Leaves + membranous or chartaceous, oblong-oblanceolate, + gradually
narrowed to the narrowly rounded or cuneate base, apex acute, (20—)25—40 by 7-12 cm;
upper surface dark olivaceous or blackish brown, lower surface grey-brown, indumentum
felty, cobweb-like, hairs scale-like, 0.1 mm or less (lens!); not papillose; dots absent;
midrib little raised above, 4—5 mm wide towards the base, lateral nerves 30-35 per side,
at 45—60° to the margin lines of interarching + distinct, venation faint or invisible at both
surfaces; petiole 17—20(—30) by 4-5 mm; leaf bud + conical, 15 by (4—)6—8 mm, hairs
0.1 mm. Inflorescences between the leaves, of the Knema-type: in male a sessile woody
brachyblast 5(—10) mm long, + glabrous, few-flowered; female not seen, flowers minute-
ly pubescent. Male flowers (immature, Wenzel 1152): pedicel 1-2 mm long, bracteole
caducous: buds ovoid, 3 mm long, not angled. Androecium 2.5 mm long; androphore
with dense appressed hairs in the lower part. Female flowers (from young fruit, FB
23019): ovary densely pubescent. Infructescence subsessile, knob-like, up to 1.5 cm
long, + below the leaves. Fruits solitary, ellipsoid, base and apex rounded, (5—)6—6.5 by
34.5 cm, hairs conspicuous, dense, bright rusty, |—1.5 mm; pericarp 10 mm thick; seeds
ellipsoid, 4 cm; fruiting pedicel 3 mm long.

Distribution — Malesia: S Philippines (Samar, Leyte, Mindanao).

Habitat & Ecology — Forests at low altitudes; fl. Oct.; fr. Oct.—Mar.

Notes — 1. Flowers and fruits approach those of M. fatua, but M. wenzelii is distin-
guishable by its stout habit, thick twigs with pale bark, large leaves with broad midrib,
with 30-35 pairs of lateral nerves. The lower leaf surface has a persistent felt-like
indumentum, resembling that of M. fatua s.s., but it is paler and shorter, with hairs only
(0.1 mm long. Mature male flowering specimens are not known.

2. Myristica wenzelii may be confused with M. philippensis; both species have a stout
habit: the latter with a pedunculate paniculate inflorescence, and the leaves glabrescent
below.

151. Myristica womersleyi J. Sinclair

Myristica womersleyi J. Sinclair, Gard. Bull. Sing. 23 (1968) 249, f. 24; Foreman, Handb. Fl. Papua
New Guinea 1 (1978) 215, f. 84, 93; W.J. de Wilde, Blumea 40 (1995) 343. — Type: NGF 11374
(Womersley), Papua New Guinea.

Tree 20-27 m. Twigs subterete or bluntly angled, 3-5 mm diameter, late glabrescent,
hairs (0.5—)1 mm; older twigs somewhat longitudinally cracking, not flaking, (finely
and) inconspicuously lenticellate. Leaves coriaceous, ovate-oblong or oblong(-lanceo-
late), 12-21 by (3-)4—9 cm, base short-attenuate or rounded, apex acute(-acuminate),
upper surface brown, lower surface grey- (or reddish) brown by dense (sub)persistent
indumentum, hairs interwoven, scale-like, 0.1—0.2 mm, mixed with scattered variously
sized emergent hairs, longly branched from the base, to 1(—2) mm; not papillose; dots
absent: midrib slender, + raised above (sometimes in a groove), lateral nerves 23-27 per
side, at 50—80° to the midrib, flat or sunken above, lines of interarching + distinct, vena-
tion somewhat scalariform, + sunken above, rendering the surface slightly bullate; peti-
ole 8-15 by 3-4 mm, leaf bud 20 by 4—5 mm, hairs | mm long. Inflorescences (from
infructescences) between the leaves or below, of the Knema-type: sessile, wart-like

De Wilde — Myristicaceae (Myristica) 617

brachyblasts, 1- or few-flowered. Male and female flowers not known. Fruits subsessile,
single, below the leaves, globose, 6—9 cm diameter, hairs dense, + rough, dark brown,
0.5—1 mm; pericarp 6 mm thick; seeds subglobose, 5.5 by 5 cm, aril with narrow seg-
ments; fruiting pedicel 3-5 by 5 mm. — Fig. 62a, b.

Field-notes — Tree with deep narrow crown, without buttresses. Bark grey, rather
hard and brittle: underbark red; inner bark dark straw-brown; wood pale straw; sap runs
clear and slowly, turning red-brown. Young foliage rusty tomentose below, old leaves
glaucous below, nerves deeply impressed. Fruits apparently not dehiscing, seeds brown,
intensely aromatic.

Distribution — Malesia: Papua New Guinea (a local endemic of Eastern Highlands
Prov.: Mt Michael, and Mt Piora).

Habitat & Ecology — Forest of slopes and ridges; on steep slope in Cunoniaceae and
Nothofagus dominated forest; 2000-2330 m altitude; fr. May, Sept., Oct.

Notes — 1. Resembles M. fusca and M. sphaerosperma in general habit and inflores-
cences, taxonomically closest related to the latter. Myristica fusca 1s distinguishable by
the papillose lower leaf surface; M. sphaerosperma differs e.g. in the irregularly papil-
lose lower leaf surface and a longer fruiting pedicel (7-15 mm). Myristica womersleyi
also resembles M. chrysophylla with similar leaves, but that species has different fruits,
24.5 cm long, with 2—5 mm long yellow brown hairs.

2. According to Foreman (1.c.) this species is, although restricted in its distribution,
locally quite common. In the Mt Piora area the seeds of M. womersleyi are used as a bait
in kapul (possum) traps. The seeds also are a possible source of spice and oil, as already
noted by Sinclair, |.c.

152. Myristica wyatt-smithii Airy Shaw

Myristica wyatt-smithii Airy Shaw, Kew Bull. No. 2 (1948) 251; W.J. de Wilde, Blumea 42 (1997)
188. — Type: KEP 52149, Peninsular Malaysia.

Tree 12-25 cm. Twigs sometimes yellowish, 2—3(—3.5) mm diameter, glabrous or ear-
ly glabrescent, hairs 0.1 mm or less; older twigs smooth with irregular cracks, + flaking;
lenticels absent or few and faint. Leaves membranous or chartaceous, elliptic-oblong or
oblong-(ob)lanceolate, (15—)20-35 by (5—)6—12 cm, base cuneate, rarely + rounded,
apex acute-acuminate; upper surface dull, greyish-olivaceous or brown, lower surface
dull, paler, glabrous; not obviously papillose; dots absent; midrib above flat or slightly
raised to the apex, lateral nerves 15-20 per side, at 45—80° to the midrib, sunken, indis-
tinct above and below (faint intersecondary nerves often present), lines of interarching
and venation indistinct; petiole 15-30 by 2-4 mm; leaf bud 10-15(—20) by 2-3 mm,
hairs 0.1 mm. Inflorescences between the leaves or (KEP 52149, 63176; FRI 20025)
axillary to fallen reduced leaves along up to 5 cm long lateral short-shoots with apical
leaf bud; paniculate, pedunculate, glabrescent, hairs 0.1 mm or less; in male: 2—4.5 cm
long, peduncle 15-20 mm, lateral branches 4-6 mm long, central axis 10-20 mm long,
with 1 or 2 spaced subumbels, the latter with 4-8 flowers, buds of variable sizes; bracts
caducous, 1-3 mm long; in female: peduncle 2-15 mm, not or hardly branched, 1-3
(—5)-flowered; flowers with hairs 0.1 mm or less. Male flowers: pedicel 4-8 mm, bracteole
rounded, 2-3 by 4 mm, with minutely ciliate margin, (sub)persistent, apical or up to
1 mm adnate to the perianth; buds ovoid, 5 by 3.5—4 mm, cleft 1/3—nearly 1/2, lobes

618 Flora Malesiana, Ser. I, Vol. 14 (2000)

(0.3-)0.4 mm thick. Androecium 3—3.5 mm; androphore 1.5 by 0.6—0.7 mm, with hairs
less than 0.1 mm towards the base; synandrium 2(—2.3) by 0.6—0.7 mm, thecae 14-16,
sterile apex + blunt, 0.2-0.3 mm. Female flowers: pedicel 3-4 mn, bracteole 1.5 by 4
mm: buds ovoid, 4.5 by 3 mm, cleft c. 1/3, lobes hard-fleshy, 0.5 mm thick; ovary ovoid,
3 by 2 mm, hairs scattered, less than 0.1 mm. Fruits | or 2 per infructescence, ellipsoid
(-oblong), 5.5-8.5 by 3.5—4.5 cm, early or late glabrescent, hairs scurfy, pale brown,
0.1(—0.2) mm or less (remnants of indumentum persistent in hidden places close to the
fruit stalk); pericarp (S—)10—15 mm thick, blackish brown; seeds ellipsoid, 4-5 cm; fruiting
pedicel stoutish, 4-10 mm long.

Field-notes — Erect tree of tall forest, no buttresses nor stilt-roots. Bark blackish
brown or greyish black, scaly or flaky (flakes flat, adherent, rather widely spaced or
gritty). Leaves glossy green, lighter below, quite glabrous. Flowers yellow. Fruits apri-
cot(-brown) or orange-red; finely scurfy.

Distribution — Malesia: Peninsular Malesia (possibly Sumatra, Borneo, see notes 3
and 4).

Habitat & Ecology — Hillsides and ridges with tall forest; also on crests, conglomeratic
ridge; 0—600 m altitude; fl. Apr., May; fr. throughout the year.

Notes — 1. This species may be confused with M. iners and M. malaccensis. Myristica
wyatt-smithii is distinguishable by its large, dull-drying leaves; the male flowers are
larger than those of M. malaccensis, smaller than those of M. iners. The latter species
furthermore differs in its smaller leaves and details of habit, twigs, leaves, flowers (incl.
androecium), and fruits. In M. malaccensis the lines of interarching of the nerves on the
lower leaf surface are more distinct.

2. Mature fruits of FRI 12553 have a rather persistent indumentum of more densely
set longer hairs, 0.1—0.2 mm.

3. Some fruiting specimens from Borneo (S 16416, 42356; SAN 16808), probably
close to M. extensa from which they differ mainly by glabrous (early glabrescent) fruits,
key out besides M. wyatt-smithii. Possibly these specimens represent a separate, as yet
undescribed species.

4. Some sterile collections from outside Peninsular Malaysia (Enggano I., off Sumatra,
and Sepilok, Sabah) may belong to M. wyatt-smithii, and are discussed by De Wilde, l.c.

DOUBTFUL AND EXCLUDED NAMES

There are a number of names in Myristica which do not belong to the family (but alter-
natively to, e.g., Annonaceae, Lauraceae, Myrsinaceae), or to other myristicaceous gen-
era, including those from Africa and South America, remembering that in the foregoing
century all Myristicaceae were described in the sole genus Myristica, chiefly by A. De
Candolle (in his Prodromus), until King published his revision of the family for British
India [Ann. Roy. Bot. Gard. Calcutta 3 (1891)]. Only since Warburg (1897) the modern
genera were defined clearly.

Excluded and doubtful names were enumerated and discussed by Sinclair [Gard. Bull.
Sing. 3 (1968) 480-488], and for New Guinea also by De Wilde [Blumea 40 (1995)
237-344]. Below follow some obscure names of true Malesian Myristicas (all New
Guinean), doubtful in synonymy and to be excluded:

De Wilde — Myristicaceae (Myristica / Paramyristica) 619

Myristica avis paradisiacae Warb. — Papua New Guinea; based on seeds only.
Myristica costata Warb. — Papua New Guinea; type lost, a doubtful species.

Myristica finschii Warb. — Papua New Guinea; a mixtum compositum now destroyed,
of loose fruits of M. inutilis subsp. papuana and a leafy twig of M. argentea.

Myristica macrocarya Warb. — Papua New Guinea, based on seeds only.
Myristica pseudo-argentea Warb. — Papua New Guinea; seeds (or fruits?) only.

Myristica resinosa Warb. — NW Papua Barat; type destroyed, a doubtful species.

PARAMYRISTICA

Paramyristica W.J. de Wilde, Blumea 39 (1994) 341-350, f. 1, 2. — Type species: Paramyristica
sepicana (Foreman) W.J. de Wilde, based on Myristica sepicana Foreman.

Trees, dioecious. Twigs lenticellate, flaky, crowded scars of cataphylls at base of in-
novations usually present. Leaves brittle when dry, lower surface pale, papillose, dots
absent: reticulation faint. Inflorescences (synflorescences) below the leaves, a compound
panicle-like short-shoot 1-5 cm long, ending ina vegetative bud; partial inflorescences
pedunculate, with flowers in subumbels of 5—10 in male, fewer in female; basal cataphylls
absent: bracts caducous. Flowers long pedicellate, bracteole absent. Male flowers: peri-
anth + urceolate, leathery, inside glabrous except at base of androphore, creamy ( ?); buds
ovoid, 5-7 mm long, cleft for 1/3 to nearly halfway, lobes 3, not reflexed in anthesis;
androecium 4(—4.2) mm long, androphore slender, 1.5—2 mm long, glabrous, except for
a dense collar of fine hairs at base, synandrium of c. 20 completely fused linear anthers,
subglobose, 22.4 mm diameter, the central column deeply and broadly excavated nearly
to the base. Female flowers not known. Infructescences 2-5 cm long, with 2—6 fruits.
Fruits ovoid-ellipsoid, 4—5.5 cm long, woolly tomentose; pericarp + thick, + woody; aril
laciniate to the base; seeds ellipsoid-oblong, 3 cm long, not variegated; cotyledons not
known. — Fig. 94.

Distribution — Malesia: Papua New Guinea; one species. — Map 6 (see p. 5).

Notes — 1. Because the compound inflorescence is a short-shoot, with a vegetative
terminal bud, the inflorescence can be named a synflorescence.

2. The deeply hollowed-out subglobose synandrium, in which the anthers curve in at
the apex, is reminiscent of many species of Horsfieldia.

Paramyristica sepicana (Foreman) W.J. de Wilde

Paramyristica sepicana (Foreman) W.J. de Wilde, Blumea 39 (1994) 348, f. 1, 2. — Myristica sepi-
cana Foreman, Contr. Herb. Austral. no. 9 (1974) 40, f. 2; Handb. Fl. Papua New Guinea |
(1978) 209. — Type: NGF 19516 (Sayers), Papua New Guinea.

Tree 5-25 m. Twigs (sub)terete, 3-6(—7) mm diameter, with minute + appressed pale
or bright rusty hairs 0.2—0.5 mm long, glabrescent, bark greyish, coarsely striate, when
older ashy or blackish, conspicuously cracked and flaky, with + concolorous lenticels;
crowded scars of bud scales at base of innovations. Leaves chartaceous, elliptic-oblong
or oblong-oblanceolate, 21-45 by (5—)8—17 cm, base rather gradually cuneate, apex

620 Flora Malesiana, Ser. I, Vol. 14 (2000)

broadly acute with acute(-acuminate) or + blunt tip; upper surface (greenish) olivaceous,
lower surface glabrous, grey(-brown) with the yellowish nerves contrasting; strongly
papillose; dots absent; midrib in large leaves to 5 mm wide, flat or slightly raised above;

Fig. 94. Paramyristica sepicana (Foreman) W.J. de Wilde. a. Leafy twig apex; note sterile apical
bud with scale and scale scars, and congested scars of scales lower down at transition with previous
innovation; b. male synflorescence; note apical bud; c. male bud, opened, and schematic longitudi-
nal section of androecium showing hollowed-out central column; d. infructescence on older wood
of twig, fruits somewhat immature [a: LAE 52983 (and NGF 48265); b, c: NGF 19516; d: LAE
73769]. — Scale bar for a, b, d = 2 cm; for c = 2 mm.

De Wilde — Myristicaceae (Paramyristica) 621

nerves 15-22 per side, at 45—60(—70)° to the midrib, slender, flat above, much raised
below, lines of interarching indistinct, venation faint on both surfaces; petiole (6—)10—
20 by 4-6 mm, brown or blackish, contrasting with light brown midrib and grey twigs;
apical leaf bud (sub)acute, 10-15 by 4—6 mm, with appressed dull brown hairs 0.5 mm,
at base with few or several bud scales, mostly with the apical part broken off. /nflores-
cences apically or below the leaves, compound; synflorescences consisting of short-
shoots (1-)2—4 cm long, always ending in a sterile vegetative bud 0.5(—1) cm long, and
with scars of cataphylls at base, the partial inflorescences 3—5, distichous, in the axils of
fallen cataphylls, with shaggy bright rusty hairs 0.5(—1) mm, peduncle flattened; flowers
with shaggy yellow-brown hairs 0.5(-1) mm long, bracts irregularly broadly rounded,
+ convex, 5-10 mm diameter, early shed, leaving distinct scars. Male synflorescences
3-6 cm long; partial inflorescences with peduncle 1—3 cm long, rachis up to 15 mm,
with 1-3 subumbels of 5—10 flowers, buds in equal stages of development. Female syn-
florescences (from infructescences) possibly always on the older wood, 1-4 cm long,
with partial inflorescences | cm long, each few-flowered; female flowers not seen. Male
flowers: pedicel slender, (5—)8—10 by 0.8(-1) mm, bracteole absent; buds ovoid, (S5—)6—
7 by (4-)5 mm, not angular, apex rounded, base broadly rounded, inside glabrous, cleft
c. 1/3 to nearly halfway, into 3 (or 4) lobes, 2.5-3 mm long, at sutures 0.2—0.3 mm thick;
androecium 4—4.2 mm long, androphore slender, 1.5(—2) by (0.3—)0.4 mm, finely stri-
ate, glabrous, except for a basal collar of dense curly brown hairs 0.3—0.4 mm long,
synandrium subglobose or broadly ellipsoid, truncate or depressed at both ends, 2-2.4
by 2 mm, at apex deeply hollowed out nearly to the base; anthers probably c. 20 (1.e.,
c. 40 linear thecae), contiguous, partially curving into the apical hollow, sterile apex ab-
sent. Infructescences 2-6-fruited, on the older wood below the leaves, 2-4(—5) cm long,
bark of branches fissured and flaking. Fruits ovoid-ellipsoid, (4—)5—5.5 by 2.5(—3) cm,
base rounded or truncate, apex narrowed, subacute, or somewhat uncinate, hairs woolly,
rusty, 1-1.5 mm long; pericarp rather woody, 5—7 mm thick, seeds ellipsoid-oblong,
3 by 1.5 cm, with impressions of the aril lobes; fruiting pedicel 3-6 by 3-5 mm, without
bracteole scar, pubescent. — Fig. 94.

Field-notes — Bole to 8 m, branches horizontal or slightly pendulous. Bark brown or
black, finely longitudinally fissured, smooth, or peeling in large sheets; inner bark light
(reddish) brown, with red sap; wood soft, straw, or pale reddish brown. Leaves glossy
and light green above, whitish, or almost glaucous below. Inflorescence covered with
brown indumentum; inflorescences and infructescences ramiflorous; flowers cream. Fruits
with dark brown or reddish brown indumentum, aril red.

Distribution — Malesia: Papua New Guinea (a restricted area in West and East Sepik
Prov.).

Habitat & Ecology — Lowland forest on ridges, on flats near creeks, and swamp for-
est dominated by Calamus species; 30-300 m altitude; fl. Mar.; fr. throughout the year.

Note — Paramyristica sepicana much resembles some species of Myristica from
New Guinea. Because of its papillose leaf undersurface it would key out in the genus
Myristica beside M. ornata. However, it does not at all resemble the latter in habit or in
fruit. The present species seems rather more related to M. markgraviana, M. carrii, and
M. hooglandii, all with a similar androecium and, indeed, in habit approaching the latter
two species. For differences between Myristica and Paramyristica, see De Wilde (l.c.:
344-347).

622 Flora Malesiana, Ser. I, Vol. 14 (2000)

LEGENDS OF THE COLOUR PHOTOGRAPHS

Photographs 1—9 (between pages 8 and 9):

1. Gymnacranthera bancana (Miq.) J. Sinclair
De Wilde & Duyfjes 20635 — Sumatra

2. Gymnacranthera farquhariana (Hook. f. & Thomson) Warb. var. farquhariana
De Wilde & Duyfjes 20542 — Sumatra

3. Gymnacranthera forbesii (King) Warb. var. forbesii
De Wilde & Duyfjes 20648 — Sumatra

4. Myristica fatua Houtt. subsp. fatua
De Wilde & Duyfjes 21921 — Hortus Bogoriensis (origin Ambon)

5. Knema glauca (Blume) Petermann var. glauca
De Wilde & Duyfjes 21923 — Hortus Bogoriensis (origin Ambon)

6. Knema kinabaluensis J. Sinclair
SAN 141918 (De Wilde & Duyfjes) — Sabah

7. Knema sumatrana (Blume) W.J. de Wilde
De Wilde & Duyfjes 19860 — Sumatra

8. Horsfieldia wallichii (Hook. f. & Thomson) Warb. — Bark slash
De Wilde & Duyfjes 20151 — Sumatra

9. Myristica iners Blume — Stilt-roots
Geesink & Santisuk 5353 — Peninsular Thailand

Photographs nos. 1-8 by W.J.J.O. de Wilde; no. 9 by R. Geesink

623

INDEX TO SCIENTIFIC PLANT NAMES

Numbers refer to page numbers. Infrageneric epithets have been entered directly under the generic
name to which they belong, preceded by the indication of their rank (sect., subsect., etc.). Infraspecific
epithets have been entered under the specific name to which they belong, preceded by the indication of
their rank (subsp., var., forma, etc. ).

Some frequently occurring author names are abbreviated here, viz.: JS (= J. Sinclair), Hk.f. & Th.
(= Hook.f. & Thomson), and WJdW (= W.J. de Wilde).

Synonyms have been printed in italics. Page numbers in bold type denote main treatment, page num-
bers with an asterisk refer to a drawing.

Cnema glomerata 135
Comacum Adans. 359
Embelia ridleyi King & Gamble
30
Endocomia WJdW 3 (map), 28,
29
canarioides (King) WJdW
30, 36*
macrocoma (Miq.) WJdW
30, 31
subsp. longipes WJdW 32,
33, 34*
subsp. macrocoma 32, 34%,
36*
subsp. prainii (King)
WJdW 32, 35, 36*
rufirachis (JS) WJdW 30,
34*.36*,38
virella WJdW 30, 39
Gymnacranthera (A. DC.)
Warb. 3 (map), 28, 40
acuminata Merr. 47
apiculata Warb. 46
bancana (Miq.) JS 40, 41, 42*
var. borneensis (Warb.) JS
41
contracta Warb. 41, 43
crassinervis Warb. 50
cryptocaryoides Elmer 344
eugeniifolia (A. DC.) JS 46
var. eugeniif 46
var. griffithii (Hk.f.) JS 45
farquhariana (Hk. f. & Th.)
Warb. 41, 44, 45
var. eugeniifolia (A. DC.)
Schouten 45, 46
var. farquhariana 45
var. griffithii (Hk. f.) Warb.
45
var. griffithii auct. 165

(Gymnacranthera farquhariana)
var. major (King) Gamble
46
var. paniculata (A. DC.)
Schouten 45, 47
var. zippeliana (Miq.)
Schouten 45, 47
farquhariana auct. 46
forbesii (King) Warb. 41, 48
var. crassinervis (Warb.) JS
49, 50
var. forbesii 49
ibutii Holthuis 33
lanceolata Merr. 405
laxa Elmer 47
macrobotrys Merr. 47
maliliensis Schouten 41, 50,
Silica
murtonii (Hk. f.) Warb. 41
var. borneensis Warb. 41
negrosensis Elmer 446
ocellata Schouten 41, 42*, 51
paniculata (A.DC.) Warb. 47
var. paniculata 47
var. zippeliana (Miq.) JS
47
stenophylla Warb. 346, 347
sulphurascens Elmer 569
urdanetensis Elmer 446
zippeliana (Miq.) Warb. 47
Horsfieldia Willd. 4 (map), 28,
29, 54
sect. Bivalves JS 56
sect. Orthanthera Warb. 55

sect. Irya (Hk.f. & Th.) Warb.

56
subsect. Euirya Warb. 57
subsect. Eyirya Warb. 56
subsect. Trivalves Warb. 57
sect. Orthanthera Warb. 57

(Horsfieldia)
sect. Pyrrhosa (Bl.) Warb. 57
subsect. Bivalves Warb. 56
subsect. Eupyrrhosa Warb.
Sy
subsect. PapillosaeWarb. 29
sect. Trivalves JS
subsect. Orthanthera
(Warb.) JS 55
subsect. Trivalves 57
acuminata Merr. 132
affinis WJdW 177
amklaal Kaneh. 132
ampla Markgr. 64, 84
ampliformis WJdW 64, 84,
85, 86*
amplomontana WJdW 71, 79,
87
amy gdalina auct. 93
androphora WJdW 71, 79,
88*, 89
angularis WJdW 64, 69, 82,89
ardisiifolia (A.DC.) Warb. 64,
81, 90, 92*
aruana (Blume) WJdW 64,
81, 82, 91
aruensis Warb. 199
atjehensis WJdW 67, 76, 93
australiana auct. 187
bartlettii Merr. 147
basifissa WJdW 65, 82, 94
batjanica Warb. 190
bivalvis (Hk. f.) Merr. 161
borneensis WJdW 71, 78, 95
brachiata (King) Warb. 69,74,
76, 77, 96
var. brachiata 96
var. laticostata JS 143
var. sumatrana (Migq.) JS
170

624

Flora Malesiana, Ser. I, Vol. 14 (2000)

(Horsfieldia)
bracteosa Henderson 203
var. bracteosa 202
var. microcarya JS 161
canariformis (Blume) Merr.
190
canarioides (King) Warb. 30
carnosa Warb. 69, 78, 98*, 99
clavata WJdW 65, 83, 100,
HOE
confertiflora Merr. 105
congestiflora A.C.Sm. 132
coriacea WJdW 72, 80. 101
corrugata Foreman 65, 84,
102*, 103
coryandra WJdW 65, 83, 104
costulata (Miq.) Warb. 71, 80,
105
crassifolia (Hk.f. & Th.)
Warb. 62, 72, 74, 76,
106*, 107
crassithyrsa 105
crux-melitensis Markgr. 65,
83, 108, 109*
decalvata WJdW 65, 81, 110
discolor WJdW 69, 78, 111
disticha WJdW 70, 79, 112
elongata WJdW 69, 73, 112,
eS
endertii WJdW 71, 79, 114
erubescens JS 187
flocculosa (King) Warb. 68,
73, 115
fragillima Airy Shaw 67, 71,
Ts US) AUIS WAI
fulva (King) Warb. 67, 73,
Wes Uy
var. paludicola (King)
Warb. 107
gigantifolia Elmer 90
glabra (Blume) Warb. 66, 72,
76, 119, 121
var. glabra 120, 121
var. javanica WJdW 120,
121
var. oviflora WJdW 120,
122
glabrescens Warb. 126
globularia (Blume) Warb.
161
var. minahassae Warb. 161
globularia auct. 200

(Horsfieldia)

gracilis WJdW 68, 77, 122,
163*
grandis (Hk. f.) Warb. 68, 73,
Tie 123
hellwigii (Warb.) Warb. 66,
84, 125
var. brachycarpa WJdW
126
var. hellwigii 124*, 126,
173
var. lignosa WJdW 126,127
var. novobritannica JS 140
var. pulverulenta (Warb.) JS
173
hirtiflora WJdW 67, 69, 76,
1P27/, 2s
inflexa WJdW 63, 82, 129,
130*
iriana WJdW 64, 82, 131
irya (Gaertn.) Warb. 62, 72,
74, 76, 80, 81, 132, 133*
iryaghedhi (Gaertn.) Warb.
62,72) 7 AS5SF136"
kerangasicola JS 178
labillardieri Warb. 132
laevigata (Blume) Warb. 66,
80, 84, 137
var. laevigata 138*, 139,140
var. novobritannica (JS)
WJdW 138*, 139, 140
lancifolia WJdW 65, 80, 141,
142*
laticostata (JS) WJdW 70, 79,
143
lauterbachii Warb. 200
lemanniana (A.DC.) Warb.
132
lemanniana auct. 169
leptantha WJdW 66, 84, 144
leptocarpa Warb. 33
leptosperma 33
macilenta WJdW 70, 74, 76,
79, 145
macrobotrys Merr. 151
macrocoma (Miq.) Warb. 31
var. canarioides (King) JS
30
var. macrocoma 32
var. rufirachis JS 38
macrothyrsa (Miq.) Warb. 72,
76, 146

(Horsfieldia)

majuscula (King) Warb. 72,
73, 76, 147
megacarpa Merr. 105
merrillii Warb. 35
minahassae (Warb.) Koord.
161
minahassae auct. 105
moluccana WJdW 63, 83,
148
var. moluccana 81, 148, 149
var. petiolaris WJdW 148,
149
var. pubescens WJdW 148,
149
var. robusta WJdW 148, 150
montana Airy Shaw 71, 79,
150
motleyi Warb. 68, 77, 151
nervosa WJdW 70, 79, 152
nesophila (Miq.) Warb. 140
nesophila auct. 200
novae-lauenburgiae Watb.
141
novo-guineensis Warb. 91,
131, 166
var. moseleyana Warb. 216
nunu Kaneh. 132
oblongata Merr. 35
obscura WJdW 72, 78, 153
obscurinervia Merr. 63, 81,
154
obtusa WJdW 70, 79, 155
odorata Willd. 135
olens WJdW 69, 82, 155
oligocarpa Warb. 68, 78, 156
olivaeformis Warb. 33
pachycarpa A.C. Sm. 65, 84,
102*, 157
pachyrachis WJdW 69, 79,
158
pachythyrsa Warb. 105
pallidicaula WJdW 67, 69, 77,
78, 159
var. macrocarya WJdW
160, 161
var. microcarya (JS) WJdW
160, 161
var. pallidicaula 160
papillosa Warb. 35
parviflora (Roxb.) JS 63, 72,
81, 161

Index to scientific names

625

(Horsfieldia)
parviflora auct. 190

paucinervis Warb. 68,77, 162,

163*
penangiana JS 63, 72, 74, 76,
78, 164
subsp. obtusifolia WJdW
165
subsp. penangiana 74, 165
perangusta WJdW 67, 73, 166
pilifera Markgr. 66, 83, 138%,
166
polyantha Warb. 140
polyantha auct. 94
polyspherula (Hk. f. em. King)
JS 70,74, 76, 80, 167, 169,
208
var. oligocarpa (Warb.) JS
156
var. maxima WJdW 168,
169
var. polyspherula 80, 168,
169
var. sumatrana (Miq.)
WJdW 168, 170
praetermissa JS 157
prainii (King) Warb. 35
psilantha WJdW 66, 83, 171
pulcherrima WJdW 68, 74,
75, 172
pulverulenta Warb. 66, 84,172
punctata WJdW 71, 72, 74,
174
punctatifolia JS 72, 74, 76, 78,
175
racemosa (King) Warb. 30
ralunensis Warb. 66, 84, 176
ralunensis auct. 141
ramosii Merr. 154
ramuensis Warb. 200
reticulata Warb. 68, 77, 163*,
77
ridleyana (King) Warb. 70, 73,
79, 178
rostrata Markgr. 201
roxburghit Warb. 190
rufo-lanata Airy Shaw 68, 77,
179
sabulosa JS 66, 77, 180, 181*
samarensis WJdW 64, 81,182
schlechteri Warb. 65, 82, 182,
183*

(Horsfieldia)

sepikensis Markgr. 71, 82,
184, 185*
sessilifolia WJdW 67, 77, 186
sinclairii WJdW 66, 83, 187
smithii Warb. 63, 81, 188
solomonensis A.C. Sm. 216
sp. Merr. 161
sparsa WJdW 67, 73, 75, 189
spicata (Roxb.) JS 63, 81,190,
Oe
var. sepikensis (Markgr.) JS
184
splendida WJdW 68, 77, 192,
194*
squamulosa WJdW 65, 83,
109*, 193
squamulosa auct. 104
sterilis WJdW 63, 77, 196
subalpina JS 72, 197
subsp. kinabaluensis WJdW
78, 197, 198
subsp. subalpina 73, 197,
198
subglobosa (Miq.) Warb. 132
var. brachiata (King) JS 96
subglobosa auct. 170
subtilis (Miq.) Warb. 65, 82,
198
var. aucta WJdW 199, 200
var. calcarea WJdW 199,
200
var. rostrata (Markgr.) JS
199, 201
var. schlechteri (Warb.) JS
182
var. subtilis 81, 82, 183%,
199
sucosa (King) Warb. 201, 202
subsp. bifissa WJdW 63,77,
202, 203
subsp. sucosa 67, 69, 73,
75, 202
sucosa auct. 189
superba (Hk.f. & Th.) Warb.
67, 73, 75, 203
sylvestris (Houtt.) Warb. 64,
80, 82, 204, 206*
var. villosa Warb. 204
talaudensis WJdW 64, 81, 207
tenuifolia (JS) WJdW 70, 79,
208, 209*

(Horsfieldia)

tomentosa Warb. 68, 73, 75,
210
triandra WJdW 67, 75, 211,
NOs
trifida A.C.Sm. 35
tristis WJdW 67, 75, 78, 213
tuberculata (K.Sch.) Warb.
65, 83, 215, 216
var. crassivalva WJdW 215,
216
var. tuberculata 214*, 215,
216
urceolata WJdW 65, 83, 217
valida (Miq.) Warb. 71, 75,
79, 218
villamilii Merr. 105
vulcanica Merr. 105
wallichii (Hk.f. & Th.) Warb.
68, 69, 74, 76, 77, 78, 219
warburgiana Elmer 90
xanthina Airy Shaw 72, 78,
220
subsp. macrophylla WJdW
220, 221
subsp. xanthina 220, 221

Knema Lour. 4 (map), 27, 29,

222
acuminata Merr. 267
alvarezii Merr. 229, 242, 243
andamanica (Warb.) WJdW
224, 244
subsp. nicobarica (Warb.)
WJdW 235, 237, 244
ashtonii JS 245
var. ashtonii 228, 240, 245
var. cinnamomea 224, 238,
245, 246*, 247
bicolor Rafin. 266
cantlyi (Hk. f.) Warb. 294
celebica WJdW 232, 243,
247, 248*
cenabrei Merr. & Quis. 274
cinerea (Poir.) Warb. 243, 249
var. alpina JS 279
var. andamanica (Warb.) JS
244
var. cordata (JS) JS 332
var. patentinervia JS
forma longipedicellata JS
347
forma patentinervia JS
319

626

Flora Malesiana, Ser. I, Vol. 14 (2000)

(Knema cinerea)
var. rubens (JS) JS 337
var. sumatrana (Blume) JS
350
var. sumatrana auct. 262,
264
communis JS 232, 235, 250
conferta (King) Warb. 233,
236, 239, 251
var. borneensis Warb. 289
var. scortechinii (King)
Warb. 340
coriacea Warb. 279
corticosa Lour. 266
curtisii (King) Warb. 227,
233, 239, 252, 253*, 254
var. amoena JS 222*, 254,
255
var. arenosa JS 254, 255
var. curtisii 236, 253*, 254
var. linguiformis JS 295
var. paludosa JS 253*, 254,
256
elmeri Merr. 222*, 224, 237,
256
elongata Warb. 289
emmae WJdW 227, 240, 257
furfuracea (Hk.f. & Th.) Warb.
225, 233, 236, 258
furfurascens Gand. 295

galeata JS 227, 238, 239, 259,

260*
geminata (Miq.) Warb. 264
gitingensis Elmer 267
glauca (Blume) Petermann
231, 235, 237, 241, 242,
261, 262

var. andamanica Warb. 244

var. bancana Warb. 351
var. glauca 262
var. nicobarica Warb. 244

var. riparia WJdW 262, 263

var. sumatrana (Blume)
Warb. 350
glauca auct. 267
glaucescens Jack 232, 235,
237, 241, 264
var. cordata JS 332
var. glaucescens
forma rubens JS 337
var. glaucescens auct. 262
var. patentinervia JS 319

(Knema)
glaucescens auct. 350

globularia (Lam.) Warb. 227,

229, 230, 234, 236, 265
glomerata (Blanco) Mert.
229, 240, 242, 267
heterophylla (Fern.- Vill.)
Warb. 267
var. pubescens Warb. 267
hirtella WJdW 232, 242, 268
var. hirtella 269
var. pilocarpa WJdW 269
var. stylosa WJdW 348
hookeriana (Hk. f. & Th.)
Warb: 225,233, 236;
270
insularis Merr. 274
intermedia (Blume) Warb.
228, 234, 236, 240, 271
var. dubia Warb. 333
kinabaluensis JS 230, 241,
272
korthalsii Warb. 228, 240,
242, 273, 274
subsp. korthalsii 273, 274
subsp. rimosa WJdW 226,
273, 274
kostermansiana WJdW 230,
241, 275
krusemaniana WJdW 233,
239, 276
kunstleri (King) Warb. 233,
238, 277, 278
subsp. alpina (JS) WJdW
28; 209)
subsp. coriacea (Warb.)
WJdW 278, 279
subsp. kunstleri 233, 278
subsp. leptophylla WJdW
278, 280*, 281
subsp. macrophylla WJdW
236, 278, 281
subsp. parvifolia (Merr.)
WJdW 243, 278, 282
subsp. pseudostellata
WJdW 278, 282
var. surigaoensis JS 344
lamellaria WJdW 225, 233,
283
lampongensis WJdW 225,
236, 238, 283

(Knema)

latericia Elmer 226, 233, 238,
284
subsp. albifolia (JS) WJdW
285, 287, 288*
subsp. latericia 230, 242,
285
var. latericia 286*, 287
var. subtilis WJdW 242,
286*, 287
subsp. ridleyi (Gand.)
WJdW 236, 285, 289
forma nana WJdW 288%,
290
forma olivacea WJdW
290
forma ridleyi 290
var. albifolia JS 287
var. latericia 285
var. latericia auct. 289
var. lunduensis JS 301
latifolia Warb. 222*, 227, 236,
240, 291, 292*
laurina (Blume) Warb. 223,
234, 236, 239, 293, 294
var. heteropilis WJdW 224,
294, 295
var. laurina 294
var. malayana Warb. 329
var. minahassae Warb. 345
linguiformis (JS) WJdW 227,
239, 295, 296*
linguiformis auct. 357
longepilosa (WJdW) WJdW
225, 238, 297, 298*
losirensis WJdW 232, 237,
299, 300*
lunduensis (JS) WJdW 226,
238, 301
luteola WJdW 230, 241, 301
malayana Warb. 231, 235,
302
mamillata WJdW 228, 239,
303, 304*
mandaharan (Miq.) Warb.
226, 234, 236, 305
matanensis WJdW 230, 242,
306, 307*
membranifolia H. J.P. Winkler
222*, 226, 228)2387239,
308*, 309
meridionalis JS 289

Index to scientific names 627

(Knema) (Knema) (Knema)

mindanaensis auct. 274
minima WJdW 229, 240,
288*, 310
missionis (King) Warb. 266
mogeana WJdW 232, 242,
310, 312*
muscosa JS 232, 238, 311
nitida Merr. 291
oblongata Merr. 226, 240,
313
subsp. oblongata 314, 315*
subsp. parviflora WJdW
314, 316
subsp. pedunculata WJdW
314, 316
oblongifolia (King) Warb.
223, 234, 316
var. monticola (King) Warb.
316
obovoidea Mert. 314
palembanica (Miq.) Warb.
262
pallens WJdW 225, 238, 317,
318*
parvifolia Merr. 282
patentinervia (JS) WJdW
23102352237, 319
pectinata Warb. 229, 240, 320
subsp. pectinata 321
subsp. vestita WJdW 321
pedicellata WJdW 233, 239,
B22") 323
peltata (Roxb.) Warb. 249
percoriacea JS 225, 238, 324
forma fusca WJdW 325
forma longepilosa WJdW
297
forma percoriacea 325
forma sarawakensis WJdW
325
piriformis WJdW 226, 241,
326
plumulosa JS 228, 234, 327
pseudolaurina WJdW 224,
234, 236, 328*, 329
psilantha WJdW 225, 238,
329
pubiflora WJdW 232, 239,
330:)3314
pulchra (Miq.) Warb. 226,
234, 241, 332

retusa (King) Warb. 224, 233,
333
riangensis WJdW 232, 241,
334
ridsdaleana WJdW 227, 242,
335
rigidifolia JS 235, 336
subsp. camerona WJdW
226, 336,337
subsp. rigidifolia 231, 336
rubens (JS) WJdW 230, 234,
237337
rufa Warb. 227, 240, 338*, 339
scortechinii (King) JS 229,
232, 235, 237, 340
sericea WJdW 224, 238, 340
sphaerula (Hk. f. & Th.) Airy
Shaw 266
steenisii WJdW 226, 243,
341, 342*
stellata Merr. 233, 243, 343
subsp. cryptocaryoides
(Elmer) WJdW 344
subsp. minahassae (Warb.)
WJdW 344, 345
subsp. stellata 344
stenocarpa Warb. 229, 242,
345
stenophylla (Warb.) JS 231,
235, 237, 346, 347
subsp. longipedicellata (JS)
WJdW 241, 347
subsp. stenophylla 347
stylosa (WJdW) WJdW 241,
348
subhirtella WJdW 226, 232,
242, 349
sumatrana (Blume) WJdW
23), 235;.237, 350
tomentella (Mig.) Warb. 230,
242, 351
tridactyla Airy Shaw 229,
240, 352
subsp. pachydactyla WJdW
353
subsp. salicifolia WJdW
3539355
subsp. sublaevis WJdW
23)0,)240).288*%.353;355
subsp. tridactyla 353, 353,
354*

uliginosa JS 228, 240, 355

umbellata Warb. 291

vidalii Warb. 267

viridis WJdW 227, 239, 356*,
357

winkleri Merr. 291

woodii JS 222*, 229, 240,
358

wrayi (King) Warb. 350

Litsea rumphii (Blume) Fern.-

Vill. 555, 556

Myristica Gronov. 5 (map), 27,

28, 359
sect. Caloneura A.DC. 57
sect. Eumyristica Hk.f. & Th.
35) I!
subsect. Horsfieldia
(A.DC.) King 55
sect. Gymnacranthera A.DC.
AO
sect. Horsfieldia A. DC. 55
sect. Irya Hk.f. & Th. 56
sect. Irya auct. 55, 57
sect. Knema Blume 222
sect. Myristica Blume 359
sect. Pyrrhosa Blume 54—57
affinis Warb. 463
agusanensis Elmer 362, 364,
375, 398, 405
subsp. agusanensis 400,
405
subsp. squamulosa WJdW
400, 405
alba WJdW 366, 377, 379,
403*, 406
albertisii Warb. 485
amboinensis Gand. 470
amplifolia Warb. 41
anceps Warb. 588
andamanica Hook. f. 367
apiculata (Warb.) Boerl. 46
archboldiana A.C. Sm. 384,
390, 407
ardisiifolia A.DC. 90
arfakensis WJdW 381, 382,
407
argentea Warb. 384, 390,
408
aromatica Lam. 470
aruana Blume 91
aruensis (Warb.) Boerl. 199

628

Flora Malesiana, Ser. I, Vol. 14 (2000)

(Myristica)
atrescens WJdW 385, 390,
409

atrocorticata WJdW 388, 394,

410
avis paradisiacae Warb. 619
bancana Miq. 41
basilanica WJdW 367, 375,
411
batjanica (Warb.) Boerl. 190
bduerlenii Warb. 484
beccarii Warb. 368, 372, 413
bialata Warb. 382, 398, 414
var. bialata 412*, 415*,
416
var. brevipila WJdW 412%,
416, 417
bifurcata (JS) WJdW 365,
376, 389, 417
subsp. bifurcata 367, 377,
379, 384, 403*, 418
subsp. sulaica WJdW 367,
377, 379, 418, 419
bivalvis Hk. f. 161
borneensis Warb. 362, 371,
419, 420*
borneensis Gand. 609
brachiata King 96
brachypoda WJdW 386, 391,
402*, 422
bracteata A. DC. 543
bracteata auct. 523
brassii A.C. Sm. 380, 422
brevistipes WJdW 386, 392,
402*, 423
buchneriana Warb. 388, 394,
397, 398, 424
byssacea WJdW 388, 392,
425
caesia Span. 249
cagayanensis Merr. 367, 375,
426, 427*
calocarpa Migq. 459
canariformis Blume 190
canarioides King 30
cantleyi Hk. f. 294
cantleyi auct. 333
carnosa (Warb.) Boer]. 99
carrii JS 395, 428
celebica Gand. 463
celebica Miq. 569

(Myristica)

ceylanica A. DC.
var. cagayanensis (Merr.)
JS 426
var. ceylanica auct. 447
chalmersii Warb. 482
chrysophylla JS 386, 389,
394, 396, 429
subsp. chrysophylla 401*,
430
subsp. entrecasteauxensis
(JS) WJdW 398, 430, 431
var. entrecasteauxensi 431
cimicifera R. Br. 499
var. acutifolia Warb. 499
var. insipida (R. Br.) Warb.
499
cinerea Poir. 249
cinnamomea King 364, 369,
370, 374, 431
clemensii A.C. Sm. 384, 389,
433
coacta WJdW 389, 392, 434
colinridsdalei WJdW 365,
374, 434
collettiana King 170
commersonii Blume 541
concinna JS 386, 391, 435
conferta King 251
var. borneensis (Warb.)
Boerl. 289
conspersa WJdW 381, 436
contracta (Warb.) Boerl. 43
cookii Warb. 484
coriacea (Warb.) Boerl. 279
cornutiflora JS 381, 382, 437
subsp. cornutiflora 399%,
438
subsp. elegans WJdW 438,
439
corticata WJdW 363, 372,
439
corticosa (Lour.) Hk. f. & Th.
PI I3) 77/1!
var. amplifolia Miq. 351
var. ceramensis Miq. 351
var. lanceolata Miq. 262
var. sumatrana (Blume)
Mig. 350
var. tomentella Miq. 351
corticosa auct. 262, 267, 302
costata Warb. 578, 619

(Myristica)

costulata Miq. 105
crassa King 365, 368, 371,
440, 441*
crassifolia Hk.f. & Th. 107
crassinervis (Warb.) Boerl. 50
crassipes Warb. 383, 387,
391, 393, 442, 444
subsp. altemontana WJdW
444, 445
subsp. crassipes 401*, 444,
548*
subsp. marronia WJdW
444, 445
cucullata Markgr. 380, 445
cucullata auct. 454
cumingii Warb. 364, 367,
375, 446
var. floribunda Airy Shaw
494
curtisii King 252, 254
cylindrocarpa JS 385, 390,
448
dasycarpa WJdW 382, 449,
450*
depressa WJdW 364, 369,
370, 451
devogelii WJdW 366, 373,
452
discolor Merr. 568
diversifolia Miq. 512
duplopunctata WJdW 380,
452
subsp. duplopunctata 453
subsp. versteeghi1 WJdW
453, 454
edulis F. Muell. 204
elliptica Hk. f. & Th. 361,
369, 370, 454, 455*
var. celebica (Miq.) JS 569
var. elliptica 459
var. simiarum (A.DC.) JS
565, 568
elliptilimba Merr. 569
ensifolia JS 384, 389, 457
eugeniifolia A. DC. 46
euryocarpa Warb. 485
extensa WJdW 362, 371, 457
fallax Warb. 363, 371, 459
faroensis Hemsl. 561
farquhariana Hk.f. & Th. 44,
45, 46

Index to scientific names

(Myristica)
farquhariana auct. 46, 47
fasciculata WJdW 383, 460
fatua auct. 463, 609
fatua Houtt. 366, 368, 397,
460
subsp. affinis (Warb.)
WJdW 373, 462, 463
subsp. fatua 372, 374, 376,
377, 378, 401*, 403*,
461, 462
var. affinis (Warb.) JS 463
var. fatua 462
var. ingens Foreman 495
var. macrocarpa Miq. 462
var. morindiifolia (Blume)
JS 584
var. morobensis JS 434
var. morotaiensis JS 529
var. papuana Markgr. 503
var. quercicarpa JS 550
var. sangowoensis JS 557
var. spanogheana (Miq.) JS
556
var. subcordata (Blume)
Mig. 583
var. wenzelii (Merr.) JS 615
ferruginea King 41
filipes WJdW 386, 393, 463
finschii Warb. 408, 503, 619
firmipes JS 388, 392, 464
fissiflora WJdW 382, 465
subsp. fissiflora 466
subsp. kostermansii WJdW
466, 467
fissurata WJdW 365, 376,
378, 403*, 467
flavovirens WJdW 381, 382,
468
flocculosa King 115
flosculosa JS 387, 393, 402*,
469, 548*
flosculosa auct. 425
forbesii King 49
fragrans Houtt. 365, 376, 379,
399*, 403*, 470, 471*
fragrans auct. 569
frugifera WJdW 362, 374,472
fugax WJdW 385, 391, 473
subsp. fugax 474
subsp. septentrionalis
WJdW 474, 475

(Myristica)
fulva King 119
furfuracea Hk. f. & Th. 258
furfuracea auct. 333
fusca Markgr. 396, 475
fusiformis WJdW 385, 391,
476
subsp. fusiformis 402*,
476, 477
subsp. pseudostipitata
WJdW 401*, 402*, 476,
477
garciniifolia Warb. 395, 477,
478*
geminata Miq. 264
geminata auct. 347
gigantea King 363, 369, 370,
371, 372, 479
glabra Blume 119, 121
var. grandifolia Miq. 121
var. sumatrana Mig. 170
glabra De Vriese 271
glauca Blume 261, 262
var. bancana (Warb.)
Boerl. 351
var. sumatrana (Blume)
Mig. 350
glauca auct. 271, 556
glaucescens (Jack) Hk.f. &
Th. 264
glaucescens auct. 266, 267,
302; 321
globosa Warb. 388, 394, 480
subsp. chalmersii (Warb.)
WJdW 482
subsp. globosa 482
globosa auct. 514, 596
globularia Blume 161
var. subglobosa (Miq.)
Miq. 132
globularia Lam. 265
globularia auct. 169
glomerata Mig. 135
glomerata Thunb. 135
glomerata auct. 426
gracilipes JS 387, 393, 483
grandiflora Wall. 543
grandis Hk.f. 123
griffithii Hk. f. 45
griffithii auct. 46

guatteriifolia A. DC. 364, 369,

372, 374, 483

(Myristica)

hackenbergii Diels 517
hellwigii Warb. 125, 126
heritieriifolia Lecomte 494
heterophylla Fern.-Vill. 267
heterophylla K.Sch. 485, 578
heterophylla auct. 426
hollrungii Warb. 398, 399%,
485, 486*
hooglandii JS 395, 486*, 487
hookeriana Hk.f. & Th. 270
horsfieldii Blume 135
horsfieldia auct. 107, 219
hyposticta Mig. 591
impressa Warb. 367, 373,
488, 489*
impressinervia JS 365, 373,
489*, 490
inaequalis WJdW 396, 491,
492*
incredibilis WJdW 396, 493
iners Blume 363, 369, 371,
372, 375, 400*, 493
iners auct. 267
ingens (Foreman) WJdW
397, 495
ingrata WJdW 383, 496
subsp. ingrata 412*, 497
subsp. velata WJdW 497
inopinata JS 396, 498
insipida R. Br. 367, 377, 380,
386, 391, 403*, 499, 500*
integra Wall. 170
intermedia Blume 271
var. minor Miq. 262
inundata WJdW 388, 389,
394, 502
inutilis A.Gray 368, 389, 392,
397, 502
subsp. papuana (Markgr.)
WJdW 377, 378, 397,
503
var. foremaniana WJdW
504
var. papuana 504
irya Gaertn. 132
var. crassifolia Hk.f 107
var. longifolia King 132
var. wallichii King 132
iryaghedhi Gaertn. 135
iteophylla Miq. 271
javanica Blume 132

630

Flora Malesiana, Ser. I, Vol. 14 (2000)

(Myristica)
kalkmanii WJdW 398, 505
kjellbergii WJdW 367, 373,
505
koordersii Warb. 367, 373,
506
koordersii auct. 488
korthalsii (Warb.) Boerl. 273,
274
kunstleri King 277, 278
labillardieri (Warb.) Boerl.
132
laevifolia WJdW 385, 386,
391, 394, 401*, 402*, 507
laevigata Blume 137
laevis WJdW 362, 375, 508
subsp. badia WJdW 509
subsp. laevis 509
lakilaki Murata & Nitta 587
lanceolata Wall. 266
lancifolia Poir. 366, 377, 379,
391, 510, 511
subsp. kutubuensis WJdW
SOR tt
subsp. lancifolia 386, 403*,
510, 511
subsp. montana (Roxb.)
WJdW 403*, 510, 512
var. bifurcata JS 417
var. clemensii (A.C. Sm.)
JS 433
var. montana JS 512
lancifolia Merr. 405
lasiocarpa WJdW 389, 512
laurella Gand. 471
laurifolia auct. 426
laurina Blume 293, 294
var. amboinensis (Warb.)
Boerl. 294
var. bancana (Watrb.) Boerl.
294
var. borneensis Mig. 294
var. longifolia Mig. 294
laurina auct. 267
laxiflora Merr. 405
lemanniana Wall. 132
lepidota Blume 513, 514
subsp. lepidota 368, 377,
378, 388, 392, 403*, 514
subsp. montanoides (Warb.)
WJdW 366, 377, 379,388,
393, 403*, 514

(Myristica)

leptophylla WJdW 381, 515
littoralis Miq. 484
longepetiolata WJdW 366,
374, 515
longifolia auct. 258
longipes Warb. 385, 390,
402*, 516
lowiana King 363, 370, 372,
517
lowiana auct. 440
luzonica Blanco 543
macgregorii Warb. 499
macrocarpa Blume 543
macrocarpa Wall. 459
macrocarya Warb. 619
macrocoma Mig. 31
macrophylla Miq. 578
macrophylla Roxb. 462
macrothyrsa Miq. 146
madagascariensis auct. 543
maingayi Hk. f. 363, 370, 518
majuscula King 147
malaccensis Hk. f. 362, 369,
371°520;'521
subsp. malaccensis 520,
521
subsp. papillosa WJdW
520, 521
malayana (Warb.) Boerl. 302
mandaharan Migq. 305
markgraviana A.C. Sm. 395,
§21;,522*
mas (non Rumph.) Labill. 561
mascula De Vriese 462
maxima Warb. 362, 364, 369,
370, 371, 400*, 401*, 523
mediovibex WJdW 389, 525
var. kosteriana WJdW 384,
390, 525, 526
var. mediovibex 384, 525
mediterranea WJdW 386,
392, 526
merrillii Warb. 569
microcarpa Zipp. 514
millepunctata WJdW 380,
527, 528*
mindanaensis Warb. 367,
375, 377, 37193 40375529
mindanaensis auct. 274
mindorensis Merr. 447
missionis King 266

(Myristica)

montana Roxb. 512
var. papuana (Scheff.)
Warb. 515
montanoides Warb. 514
morindiifolia Blume 584
moschata Thunb. 470
motleyi (Warb.) Boerl. 151
motleyi Warb. 479
muelleri Warb. 482
multinervia A.C. Sm. 584
murtonii Hk.f. 41
var. borneensis (Warb.)
Boerl. 41
myrmecophila Becc. 578
nana WJdW 387, 393, 530
neglecta Warb. 380, 531
nesophila Migq. 33, 140
nesophila auct. 134
nitida Merr. 447
nivea Merr. 367, 375, 377,
379, 532
notha auct. 135
oblongifolia King 316
var. monticola King 316
odorata de Vriese 135
officinalis L.f. 470
oligocarpa (Warb.) Boerl.
156
olivacea WJdW 385, 390,
532
olivaeformis (Warb.) Boerl.
149
ornata WJdW 395, 533
ovicarpa WJdW 386, 391,
534
pachycarpidia WJdW 383,
535
pachyphylla A.C. Sm. 381,
382, 536
pachythyrsa (Warb.) Boerl.
105
palawanensis Merr. 484
palembanica Migq. 262
paludicola King 107
pandurifolia H.J.P. Winkler
521
paniculata A.DC. 47
papillatifolia WJdW 385,
391, 537
papillosa (Warb.) Boerl. 35
papuana Scheff. 511

Index to scientific names

(Myristica)

papyracea JS 361, 371, 537,
5307

parviflora Roxb. 161

paucinervis (Warb.) Boerl.162

pectinata (Warb.) Boerl. 320

pedicellata JS 388, 392, 539

peltata Roxb. 249

pendulina Hk. f. 204

perlaevis WJdW 361, 376,
378, 540

philippensis Lam. 363, 374,
541

philippensis auct. 426, 521

philippinensis Gand. 471

pilosella WJdW 384, 389,
542

pilosigemma WJdW 367,
375, 543

pinnaeformis Zipp. 204

plumeriifolia Elmer 462

polyantha WJdW 387, 394,
544

polyantha (Warb.) Boerl. 140

polyspherula Hk.f. 167, 169

pseudo-argentea Warb. 619

psilocarpa WJdW 395, 544

pubicarpa WJdW 365, 376,
378, 404*, 545, 546*

pulchra Miq. 332

pulverulenta (Warb.) Boerl.
172

pumila WJdW 387, 393,
401*, 548*, 549

pygmaea WJdW 384, 389,
550

quercicarpa (JS) WJdW 396,
401*, 550

racemosa King 30

radja Mig. 585

resinosa Watb. 619

reticulata (Warb.) Boerl. 177

retusa King 333

ridleyana King 178

ridleyi Gand. 289

riedelii Warb. 484

robusta WJdW 365, 376, 378,
404*, 551

rosselensis JS 385, 390, 552

subsp. minutiflora WJdW
553
subsp. rosselensis 553

(Myristica)

roxburgii (Warb.) Boerl. 190
rubiginosa King 123
rubrinervis WJdW 362, 364,
374, 375, 554
var. duplex WJdW 554, 555
var. rubrinervis 554, 555
rufa (Warb.) Boerl. 339
rumphii (Blume) Kosterm.
867, 372,555,996
var. florentis WJdW 556,
557
var. rumphii 556
salicifolia Willd. 204
sangowoensis (JS) WJdW
366, 376, 378, 404*, 557
sarcantha WJdW 383, 558*,
559
schefferi Warb. 587
schlechterii WJdW 386, 393,
560
schleinitzii Engl. 385, 390,
395, 561, 562*
schumanniana Warb. 484
scortechinii King 340
scripta WJdW 377, 378, 381,
382, 563
var. incrassata WJdW 564
var. scripta 564
sepicana Foreman 619
sericea Warb. 503
simiarum A. DC. 364, 374,
565, 568
subsp. calcarea WJdW 372,
567*, 569
subsp. celebica (Miq.)
WJdW 373, 374, 376,
Byen404 5667, 507 =,
569
subsp. simiarum 374, 566*,
567, 568
simulans WJdW 397, 570
sinclairii WJdW 84, 389, 570,
oh Ge
smithii (Warb.) Boerl. 188
smythiesii JS 368, 372, 572
sogeriensis WJdW 387, 391,
393, 402*, 573
sorsogonensis Merr. 405
sp. Hk.f. 254
spadicea Blume 462
spanogheana Migq. 556

(Myristica)

spanogheana auct. 561
speciosa Warb. 585, 587
sphaerosperma A.C. Sm. 397,
574
sphaerula Hk.f. & Th. 266
spherocarpa Wall. 132
spicata Roxb. 190
spuria Blume 446
stenocarpa (Warb.) Boerl.
345
stenophylla (Warb.) Boerl.
347
suavis King 440
subalulata Miq. 365, 376,
378, 382, 575; 56
var. hagensis WJdW 383,
Ss SI.
var. leptantha WJdW 577,
580
var. paucifructa WJdW
577, 580
var. pedunculata WJdW
412*, 577, 581
var. subalulata 399*, 401%,
AQ4*, 412*) 57672577,
578
subalulata auct. 467
subcordata Blume 392, 397,
581, 583
var. gigacarpa WJdW 583
var. morindiifolia (Blume)
WJdW 583, 584
var. rimosa WJdW 583, 585
var. subcordata 583
subglobosa Migq. 107, 132
sublanceolata Mig. 494
subtilis Mig. 198, 199
succedanea Blume 375, 376,
379, 404*, 546*, 585,
586*
var. brevifolia Scheff. &
Teijsm. 587
sucosa King 201, 202
sulcata Warb. 387, 393, 398,
588
sulphurascens Elmer 569
sumatrana Blume 350
sumbawana Warb. 366, 368,
372, 589
superba Hk.f. & Th. 203
sycocarpa Miq. 459

Flora Malesiana, Ser. I, Vol. 14 (2000)

632

(Myristica) (Myristica)
sylvestris Houtt. 204 ultrabasica WJdW 366, 373,
sylvestris auct. 543 600
tamrauensis WJdW 380, 590 umbellata Elmer 362, 375,
teijsmannii Miq. 362, 368,591 400*, 600

tenuivenia JS 388, 392, 592
subsp. lignosa WJdW 592,
593
subsp. tenuivenia 592
tingens Blume 161
tomentella (Miq.) Boerl. 351
tomentosa Hk. f. & Th. 210
tomentosa Thunb. 462
tomentosa auct. 294
trianthera WJdW 384, 389,
593
tristis Warb. 367, 377, 379,
388, 394, 594, 596
subsp. ingambitense WJdW
596
subsp. louisiadensis WJdW
596
subsp. moluccana WJdW
595997
subsp. sessilifructa WJdW
5958597
subsp. tristis 401*, 404%,
595, 596
tuberculata K.Sch. 215, 216
tubiflora Blume 386, 393,
S97, 98h

umbellata (Warb.) Boerl. 291
umbrosa JS 395, 601
uncinata JS 395, 602
undulatifolia JS 397, 604
valida Mig. 218
velutina Markgr. 383, 605
subsp. breviflora WJdW
605, 606
subsp. velutina 605, 606
verruculosa WJdW 398, 607*,
608
verrucosula Merr. 569
villosa Warb. 365, 371, 401*,
609, 610*
vinkeana WJdW 387, 394,611
vordermanii Warb. 494
vrieseana Mig. 132
wallacea Warb. 503
var. keyensis Warb. 583
wallichii Hk. f. & Th. 219
warburgii K.Schum. 387,
393, 613
subsp. hybrida WJdW 387,
393, 614
subsp. siphonantha WJdW
612*, 614, 615

(Myristica warburgil)
subsp. warburgii 612*, 614
wenzelii Merr. 366, 374, 615
womersleyi JS 396, 401*,
616
wrayi King 350
wyatt-smithii Airy Shaw 362,
369, 617
zippeliana Miq. 47
Myristicaceae R.Br. 1-621
Myristiceae |
Pala radja Teijsm. 585
Palala Rumph.
aruana Rumph. 91
globularia Rumph. 161
quinta 161
sexta Rumph. 351
Paramyristica WJdW 5 (map),
28, 619
sepicana (Foreman) WJdW
619, 620*
Pyrrhosa Endl. 54
glabra (Blume) Hassk. 121
globularia (Blume) Hassk.
161
horsfieldii (Blume) Hassk.
135
Sterculia decandra Blanco 267
glomerata Blanco 267
Tetranthera rumphii Blume
Dos 50

List of revised families in Flora Malesiana Series I (Seed Plants)

Aceraceae 4392
Actinidiaceae s.s. 4: 37
Aizoaceae 4; 267
Alismataceae 5: 317; 6: 915
Alliaceae WIS STS)
Alseuosmiaceae 10: 335
Amaranthaceae
4: 69,593: 6:915; 8: 549
Amaryllidaceae 11: 353
Anacardiaceae 8: 395
Ancistrocladaceae 4; 8
Aponogetonaceae
ACTS 7/5 213)
Araliaceae—I Deal. 9553
Araucariaceae 10: 419
Aristolochiaceae 10: 53
Balanophoraceae
7: 783; 9: 554
Basellaceae 5: 300
Bat(id)aceae 5: 414
Betulaceae 59 AOvs (8 D7
Bignoniaceae 8: 114; 9: 554
Bixaceae s.s. 4: 239
Boraginaceae 13: 43
Burmanniaceae
4: 13,592; 5: 553; 9: 554
Burseraceae 5: 209, 567
Oe OTs Ws BANE Ge Sa>
Butomaceae 5p il ie
Byblidaceae 739.
Caesalpiniaceae 12: 409
Callitrichaceae 4; 251
Campanulaceae
6: 107, 928; 9: 556
Cannab(in)aceae Ale yp)
Cappar(id)aceae 6: 61; 7: 822
Caprifoliaceae
Ais: 6: 928: 9: 556
Cardiopteridaceae 1 93
Celastraceae 6: 227, 389, 930
Centrolepidaceae 5: 421
Ceratophyllaceae 4: 41
Chenopodiaceae
4: 99,594: 6: 932; 9: 557
Chloranthaceae 10: 123
Chrysobalanaceae 10: 635
Clethraceae TaSs9
Cochlospermaceae 4: 61
Combretaceae
4: 533; 5: 564; 6: 932

Coniferales 10:
Connaraceae
5: 495; 6: 933; 9:
Convolvulaceae 4: 388
5: 558; 6:
1S Sse Ge
Coriariaceae 11:
Cornaceae 8:
Corynocarpaceae
4: 262; 5:
Crassulaceae 4: 197; 9
Cruciferae 10:
Crypteroniaceae 8:
Ctenolophonaceae 10:
Cupressaceae 10:
Cyperaceae 7:435; 9
Daphniphyllaceae 13:
Datiscaceae 4:
Dichapetalaceae 5
Dilleniaceae 4: 141; 7
Dioscoreaceae 4:
Dipsacaceae 4
Dipterocarpaceae 9
Droseraceae
Ae Vis S83 557s Ve
Elaeagnaceae 10:
Elatinaceae 4:
Epacridaceae 6:
Ericaceae 6: 469,
8: 549; 9: 562; 10:
Erythroxylaceae
5: 543; 8:
Fagaceae 12 265%, 9:
Flacourtiaceae Se I
6: 943; 7: 827; 9
Flagellariaceae
4: 245; 9:
Geitonoplesiaceae 12:
Geraniaceae 6: 445; 9:
Gnetaceae 4: 336; 6:
Gonystylaceae 4:
Goodeniaceae 58 335s
6: 949; 7: 827; 9
Haemodoraceae
5: 111; 10:
Haloragaceae 18 239).
Hamamelidaceae Se
Hernandiaceae 12:
Hydrocaryaceae 4:
Hippocrateaceae 6:

3187)

S)5)1/

i599

936
558
385

85

Hydrocharitaceae ai

6: 952; 7:

9: 566; 10:
Hydrophyllaceae 4:
Hypericaceae 8: 1; 10:
Icacinaceae He ie Be
Illiciaceae 13:
Iridaceae 8: 77; 10:
Ixonanthaceae 10:
Juglandaceae 6:
Juncaceae 4: 210; 9:
Juncaginaceae 4:
Labiatae 8: 301; 9:
Leeaceae We

Legum.-Mimosoideae 11:

Legum.-Caesalpinioideae

12:
Lemnaceae We
Lentibulariaceae 8:
Liliaceae s.s. 9:
Linaceae 10:
Loganiaceae
6: 293,953; 9:
Lophopyxidaceae ae
Loranthaceae 13:
Lowiaceae 12e
Magnoliaceae 10:
Malpighiaceae SP
Martyniaceae 4:
Meliaceae 12:
Menispermaceae 10:
Mimosaceae 11:
Monimiaceae 10:
Moringaceae 4:
Myoporaceae 4
Myricaceae 4
Myristicaceae 14:
Najadaceae 6
Nyctaginaceae 6
Nyssaceae 4
Ochnaceae 18
Olacaceae 10: 1,
Onagraceae 8:
Opiliaceae 10:
Oxalidaceae 78 iS
Papaveraceae 5
Passifloraceae 7
Pedaliaceae 4:216; 7
Pentaphragmataceae 4
Pentaphylacaceae 5

634

Pentastemonaceae 11:
Philydraceae 4:
Phytolaccaceae 4:
Pinaceae 10:
Pittosporaceae 5: 345; 6:
Plumbaginaceae 4:
Podocarpaceae 10:
Podostemaceae 4: 65; 6:
Polemoniaceae 4:
Polygalaceae 10:
Pontederiaceae 4:
Portulacaceae aie
Primulaceae 6:
Proteaceae Se
Punicaceae 4:
Rafflesiaceae 13:
Restionaceae 5:
Rhizophoraceae

5: 429; 6: 965; 8:
Rosaceae 11:
Sabiaceae 10:

Salicaceae se IO7/
Salvadoraceae 4: 224
Sapindaceae 11: 419
Sarcosperma(ta)ceae 4: 32
Saururaceae 4: 47
Schisandraceae 13: 185
Scyphostegiaceae
S297; 6: 967
Simaroubaceae 6: 193, 968
Sonneratiaceae
A: 280551324637973
Sparganiaceae 4: 233; 10: 718
Sphenocleaceae 430627
Sphenostemonaceae 10: 145
Stackhousiaceae 4:9/35
Staphyleaceae 6: 49
Stemonaceae 11: 399
Stylidiaceae 4:529; 6: 976
Styracaceae 4: 49; 9: 568
Symplocaceae
8: 205; 9: 569; 10: 718

Taccaceae Ws
Taxaceae 10:
Thymelaeaceae
4: 349; 6: 1,976; 7:
Trapaceae 4:
Trigoniaceae 4:
Trimeniaceae 10:
Triuridaceae 10:
Turneraceae 4:
Typhaceae 4: 242: 6:
Ulmaceae 8:
Umbelliferae 4: 113,
5: 555; 6:
7: 830; 9:
Valerianaceae 4:
Violaceae
7: 179, 831; 10:
Viscaceae 13:
Xyridaceae
4: 366, 598; 9:
Zy gophyllaceae 4:

806
347

830
43
59

B27

109

255

982
Sil

B95

983

569

255

720
403

571
64

List of revised families in Flora Malesiana Series II (Ferns and Fern allies)

Azollaceae 3:
Cheiropleuriaceae 3:
Cyatheaceae iA:
Davalliaceae Sk
Equisetaceae a:

Gleicheniaceae
Isoetaceae
Lindsaea group
Lomariopsis group
Matoniaceae

Bo
Se Ueleleicie ‘esl (sie

Plagiogyriaceae 3
Polypodiaceae 3
Schizaeaceae 1:
Tectaria group 2
Thelypteridaceae 1:

331

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0224 7862

ISBN 90-71236-47-1

Flora Malesiana

Compiled and published under the auspices of
Foundation Flora Malesiana

MALESIANA