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NEW YO«K
BOTANICAL
6AIUDiM

FLORA
MALESIANA

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.F55
 THYMELAEACEAE (Ding Hou, Leyden)

Shrubs, trees, or lianas, rarely undershrubs or herbs, with a very strongly

developed and layered, fibrous, tough bast ("Seidenbast", silky fibres). Leaves
opposite or decussate, spiral or alternate, very rarely some ternate, simple,
entire, exstipulate, articulated at the base, glandular-punctate in Gonystyloideae.
Inflorescences terminal, axillary or extra-axillary, or on internodes, sometimes
on brachyblasts, simple or rarely branched, sessile or peduncled, racemose,
umbelliform, spicate, capitate, or fascicled, obviously basically racemose; flowers
rarely solitary, sometimes cauliflorous and condensed into glomerules, bracteate
(bracts sometimes forming an involucre) or ebracteate. Flowers bisexual (rarely
unisexual by abortion and polygamodioecious or dioecious in extra-Mai. spp.),
homomorphic, rarely heteromorphic, regular, tubular, campanulate or in-
fundibuliform, tube very short in Gonystyloideae, or with almost free sepals in
extra-Mai. spp., mostly caducous, some circumsciss in the lower part, or persistent
(sometimes enveloping the ripe fruit in extra-Mai. spp.), sometimes slit lengthwise
in fruit, 4-5(-6)-lobed, the lobes imbricate (rarely valvate in some extra-Mai. spp.),
equal or rarely the interior 2 slightly smaller, erect or reflexed. Corolla absent or
represented by free or united petaloid appendages, isomerous and alternating with
the calyx lobes, or double in number and arranged in pairs opposite the calyx
lobes, rarely more (Gonystylus), fleshy or membranous, filamentous or oblong,
entire or lobed, rarely united into a ring, inserted at the throat of floral tube or
slightly lower, sometimes behind the stamens, or absent. Stamens 2 only, or
4-c\D, in Malaysia (except in some Gonystyloideae) mostly diplostemonous, in
two or in one series, if in two series then at two different levels, the upper ones
opposite the calyx lobes and the lower ones alternate with them, sessile or fila-
mentous; filaments filiform or slightly flattened, entirely or partly adnate to the
floral tube; anthers 2-celled, basi- or dorsifixed, obtuse or apiculate, introrse,
hippocrepiform (Gonystyloideae), or extrorse (extra-Mai. spp.), dehiscing length-
wise, usually free, sometimes the lower Y^-Yz adnate to the tube (Aquilaria
cumingiana). Disk hypogynous, membranous or subcarnose, annular, cupular,
lobed, free and scale-like, or none. Ovary superior, 1-2-celled, 3-5(-8)-celled in
Gonystyloideae and extra-Mai. spp., sessile or shortly stalked; style filiform,
caducous, sometimes very short or obscure, terminal or excentric, in Gonystyloideae
sometimes accompanied by 'parastyles' at the base; stigma capitate, subglobose,
oblong, subclavate or pyramidal, entire and smooth, or slightly emarginate,
sometimes papillose. Ovules solitary in each cell, with axial or parietal placenta-
tion, pendulous from near the top, sometimes partly or entirely and laterally
adnate to the placenta, the micropyle towards the top and outward. Fruit a drupe
or drupaceous, a berry, or a capsule, either apically or laterally emerging from
the floral tube, 1- or 2(-3)-seeded, or 3-5(-8)-seeded in Gonystyloideae and
extra-Mai. spp. pericarp membranous, pulpy, coriaceous, or fibrous. Seeds with
;

a caruncle-like or tail-like appendage, usually with an aril in Gonystyloideae, the

seed usually hanging out by one end on a thin, string-like funicle in Aquilarioideae;
testa usually crustaceous, black, often with rather irregular ridges, glabrous or
short-hairy in some spp. o^ Aquilarioideae; albuminous or exalbuminous. Embryo
straight; cotyledons plano-convex; radicle short, superior.

(1)
 Flora Malesiana [ser. I, vol. 6^

Distribution. About 50 genera with about 500 species, chiefly developed in south and tropical
Africa and Australia; it is almost cosmopolitan.
In addition to subfam. Gonystyloideae which contains 3 genera with 21 species and has been treated in
this Flora (I, 4, 1953, 349-365) by Airy Shaw, there are 9 genera with 46 species in Malaysia.
Aqiiilaria, Gyrinops, Enkleia, and Linostoma are confined to Malaysia and the southern part of tropical
continental SE. Asia. Wikstroemia is widely distributed, from eastern Asia at about 37° N
southward
throughout Malaysia to northern and eastern Australia and the Pacific Islands (Bonin, Guam, Palau,
Hawaii, Tahiti, Marquesas, Tonga, Samoa, Fiji, Norfolk I., and New Caledonia).
Daphne is distributed in Europe, northern Africa, through central Asia, eastward to China and Japan,
and southward to Malaysia.
Phaleria is developed chiefly in Malaysia and Fiji, westward to Ceylon {P. capitata), southward to
eastern Australia, and eastward as far as Palau, Samoa, and Tonga.
Drapetes shows the typical pattern of S. Pacific subantarctic distribution: South America (Fuegia
and Falkland Is.), New Zealand, Tasmania, SE. Australia, and Malaysia (New Guinea and Borneo).
Pimelea is chiefly confined to Australia, with some outlying species in New Zealand, and two others
extending northward to Malaysia (Timor, Sumba, New Guinea, and Luzon in the Philippines).
Ecology. Most Malaysian species are of small to moderate size, while a few species oi Aquilaria,
Gyrinops, and Gonystyloideae are trees up to 45 m tall. They usually occur scattered, but Gonystylus

bancanus may occur gregarious, sometimes forming pure stands. They are chiefly constituents of primary
and secondary rain-forests, while Gonystylus bancanus occurs predominantly in freshwater swamp and
peat forest; recently J. A. R. Anderson found Linostoma longiflorum in peat swamp forest in Sarawak.
Most of the species occur at low and medium altitudes, some of them ascending into the montane zone
{e.g. Phaleria capitata 0-1200 m and Linostoma pauciflorum 0-1300 m), or even confined to the montane
zone {,e.g. Daphne composita commonly recorded from 1200-2000 m, and Aquilaria apiculata from
1800 m). A few are restricted to the upper montane and subalpine rain-forest {e.g. Daphne luzonica
2000-2500 m, and Wikstroemia brachyantha 1400-2800 m). Drapetes ericoides is commonly reported
from the subalpine to alpine zone from 3000-4450 m.
As to climate, most of the species are confined to everwet regions, some also extend to seasonal areas
{e.g. Phaleria capitata, Gyrinops versteegii, and Wikstroemia indica), while Wikstroemia androsaemifolia
and Phaleria octandra chiefly occur under seasonal climatic conditions.
Pollination. Insect-pollination is indicated by the brightly coloured, generally many-flowered
inflorescences, the sweet scent, the occurrence of floral heteromorphism, and the usual presence of the
hypogynous disk (^^e Rendle, Classif. Fl. PI. 2, 1952, 371). I have no records of observations on Malay-
sian species.
Dispersal. Though no direct evidence has been recorded from Malaysia it can be indirectly inferred
that the red or black coloured drupaceous fruits of Wikstroemia, Phaleria, and Daphne will be dispersed
endozoically by birds or other animals. See Ridley, Disp. (1930) 401, 466, 472, and Guppy, Observ.
Nat. Pac. 2 (1906) 348. Wikstroemia indica has, probably through this agent spread from the Botanic
Gardens at Bogor but its area is only slowly, though steadily, extending into a circle with Bogor in its
focus; its radius of c. 60 km was reached only after several decades.
Another dispersal class is represented by species of Linostoma and Enkleia. In Linostoma the in-
florescences consist of a few inconspicuous flowers subtended by a pair of thin, cream-coloured or
rose-pink coloured leaf-like bracts. In anthesis they possibly act as a show apparatus attractive to
pollinators. They become pale and papery when the fruit is ripe, and are detached, adhering to the fruit,
so as to be blown away separately. In Enkleia, a lofty climber, the pair of bracts below the inflorescence
is very inconspicuous during anthesis, but in fruit (one developing only) the peduncle below the small

nut lengthens considerably and the bracts grow to large, stiff, coriaceous leaves (fig. lOe). When the fruit,
on its peduncle, with the two bract leaves attached, separates from the plant, it rotates rapidly, drifting
away in the wind, across the forest to some distance (Ridley, I.e. 92-93). Though the structure is most
peculiar, effect (for longer distances) must not be overrated as winds are scarce in the tropical rain-
its

forest, the apparatus is rather heavy, and as soon as it descends in the canopy it will come down, gradu-
ally, in a vertical line.
A third, very interesting dispersal class is represented by the capsular fruits o^ Aquilaria and Gyrinops,
in which the seeds dangle from the apex of the fruit valves on filiform funicles, the glossy seeds having
typically contrasting dark colours and possessing tails or other aril-like structures, probably of a pale
colour, as is also found in Gonystyloideae (fig. 1 and 22). This structure is doubtless a curious adaptation
to zoochorous dispersal, but unfortunately no observations have as yet revealed more exact data on its
functioning.
Galls. DocTERS VAN Leeuwen (Zoocec. N.I. 1926, 397, f.735) recorded a leaf-gall caused by a gall-
midge in Phaleria laurifolia (= P. octandra). The leaves bear spherical galls, 2-3 mm in diam.
Heteromorphous flowers. The flowers are heteromorphous in Phaleria macrocarpa. Two kinds of
flowers are commonly found on different plants of that species, viz possessing exserted stamens and a
short style and short stamens and an exserted style.
There is a sheet in Leyden Herbarium identified as "'Phaleria neumanni F. v. M." collected by W. Dunn
s.n. (in Nov. 1909) at Acacia D'K, New South Wales, which has three separate branchlets with similar
Dec. 1960] Thymelaeaceae (Ding Hou) 3

vegetative parts and two forms of flowers just like the above-mentioned case. It is not clear whether they
were collected from the same plant.
Wood-anatomy, den Berger, Determinatietabel houtsoorten van Malesie, Veenman, Wageningen
(1949) 20 {Aquilaria); Desch, Mai. For. Rec. 15- (1954) 607; Leandri, Ann. Sc. Nat. Bot. X, 12 (1930)
125 (hand lens); Metcalfe & Chalk, Anat. Die. 2 (1950) 1169 & 1178; Moll & Janssonius, Mikr.
—
Holzes 5 (1934) 413. By Janssonius I.e. Gonystylus is referred ioihQ Thymelaeaceae, n\diin\y because of
the characteristics shown by the pit pairs; Metcalfe & Chalk I.e., although recognizing common
features, are treating Gonystylaceae as a separate family. — C.A.R.-G.
Morphology. In order to avoid confusion, it is advisable to give a concise explanation of some
terms which are used in the descriptions of this revision. These terms serve for convenience of descriptive
purpose.
Floral tube. —
The vascular bundles going to the ovary are clearly different from those of the tube
above the pedicel; the tube contains the vascular bundles of the outer whorls, it is 'appendicular' and
not 'axile' in origin. Therefore, the tube is not an invaginated receptacle {fide Leandri, Ann. Sc. Nat.
Bot. X, 12, 1930, 235). Miss Heinig (Am. J. Bot. 38, 1951, 125) confirmed the 'appendicular' origin of
the tube which is composed of the fused bases of the sepals and adherent filaments. In the following I
have called the tubular part of the flower the 'floral tube' and its lobes 'calyx lobes'.
Petaloid appendages. —In some genera there are petal-like structures, situated either at the throat or
on the receptacle surrounding the ovary. In this treatment, they have been designated as 'petaloid
appendages'. Miss Heinig suggested {I.e. 127) them to represent special enations of the sepals.
Disk. —In some genera and species there is a cup-shaped or free, thin structure at the base of the
ovary, which has here been designated as 'disk'. According to Miss Heinig {I.e. 128) this structure is
probably a part of the androecium.
—
Chalazal fold. A mature seed-coat is formed by the outer integument and the inner integument;
the latter is composed of a sclerenchymatous layer and a reticular layer {cf. Guerin, Ann. Jard. Bot.
Btzg 29, 1916, 29).
In all the seeds (at least in Malaysia) there is, at the basal part or chalazal end, either a caruncle-like
thickening (in most of the genera) or a tail-like appendage (in some species of Aquilarioideae). The
formation of the tail-like appendage has been interpreted in difi"erent ways. Gilg (in E. & P. Pfl. Fam.lll,
6a, 1894, 223) assumes it to be the downward elongation of the integument. Lecomte (Bull. Soc. Bot.
Fr. 61, 1914, 414-418) accepted it as the elongation of the lower part of the ovule. Domke (Bibl. Bot.l 11,
1934, 37, t.V, f.43a-h) believed it to be formed by a more or less deeply transverse fold of the testa and
designated it as "chalazal fold". However, the ontogeny of this appendage has not been well understood
and further morphological and anatomical studies are needed.
—
Albumen. Endosperm is found in most of the seeds although it is often a very thin layer, predo-
minantly found on the dorsal surface of the cotyledons; it is very abundant in the seeds oi Pimelea
{ef. Guerin, Ann. Jard. Bot. Btzg 29, 1916, 31-32, t.4). The absence of endosperm is rather rare (some
Phalerias).
Cytology. As far as is known a basic number of chromosomes in the family seems to be n = 9,
which found in Wikstroemia, Gnidia, and Daphne; Edgeworthia has n = 18. In Wikstroemia indiea
is

Fagerlind found also an apomictic triploid 2 n = 27 (Hereditas 26, 1940, 1-50).

—
Taxonomy. Subdivisions. According to Domke (Bibl. Bot. Ill, 1934, 103-104) the family is sub-
divided into 4 subfamilies, viz Gonystyloideae, Aquilarioideae, Gilgiodaphnoideae, and Thymelaeoideae.
With the exception of Gilgiodaphnoideae, the other three subfamilies all have some representatives in
Malaysia.
The 3 genera of trib. Aquilarioideae-Mierosemmatidae, all endemic in New Caledonia, seem to be
more closely related to subfam. Gonystyloideae than their arrangement in two distinct subfamilies would
suggest; they lack the pellucid dots and the petaloid appendages of the latter. But Solmsia has the typical
parallel nervation, venation, and leaf texture as in Gonystylus, and the nervation and texture of the leaves
of Mierosemma and Deltaria is resembling that of Amyxa. Furthermore, the macroscopical structure
of the bast fibers in the three genera of Microsemmatidae resembles that of Gonystyloideae and is not so
fine as in typical Aquilarioideae. Finally, the fruit in Aquilarioideae is 2-celled, against 3- or more-celled
in both Microsemmatidae and Gonystyloideae.
1. Subfam. Gonystyloideae has been treated in this Flora (I, 4, 1953, 349-365) by Airy Shaw.
Leaves mostly pellucid-punctate. Flowers with a short or inconspicuous tube. Petaloid appendages
7-40, deltoid to linear-subulate, rarely joined into a low, entire annulus, inserted at the base of the floral
tube. Disk 0. Stamens 8-80; filaments free. Ovary (2-)3-5(-8)-celled. Fruit a capsule. Seeds without
chalazal fold, usually with aril. Endosperm 0. {Gonystylus, Amyxa, and Aetoxylon.)
2. Subfam. Aquilarioideae. Leaves not pellucid-punctate. Flowers with a short to cylindric tube or
sepals free. Petaloid appendages scale-like, free or rarely united, inserted at the throat of the tube or
slightly below it or none. Stamens (in the Mai. spp.) at most 10,diplostemonous or haplostemonous;
filaments (in Mai. spp.) partly or entirely adnate to the tube. Disk 0, or ring-shaped. Ovary (in Mai.
spp.) 2-celled. Fruit a capsule. Seeds usually with a conspicuous chalazal fold, and a thin funicle, without
aril. Endosperm 0, or present. {Aquilaria and Gyrinops.)
3. Subfam. Thymelaeoideae. Leaves not pellucid-punctate. Floral tube funnel-shaped or cylindric.
 Flora Malesiana [ser. I, vol. 6^

Petaloid appendages obscure and ridge-like or represented by scales. Stamens at most 10, usually diplo-
stemonous, rarely haplostemonous or hemistemonous; filaments partly or entirely adnate to the tube.
Ovary 1-2-celled. Fruit a drupe or drupaceous. Seeds mostly without or rarely with a small chalazal fold.
Endosperm 0, or present. (Linostoma, Enkleia, Phaleria, Wikstroeniia, Daphne, Drapetes, and Pimelea.)
Generic delimitation in Thymelaeaceae proves sometimes to be very difficult on account of the fact that
though the majority of the species of one genus might be distinguished from those of another genus by
two or even more good characters, there are frequently one or two species - or even different specimens
of one species - which form an exception and are transitional in all but one character. Consequently
such genera are then sharply separated by one character only, which is an unsatisfactory situation.
For instance in Phaleria the petaloid appendages are rim-like, but they are distinct in P. pentecostalis
Leandri. In Aquilaria the opposed case occurs, viz that they are distinct in all species except in A.
urdanetensis where they are rim-like. In Aquilaria the anthers are always free from the tube except in
A. cumingiana where they are partly adnate to the tube in part of the specimens! Also in Aquilaria the
petaloid appendages are free except in part of the specimens of ^4. cumingiana; the same phenomenon is
observed in Gyrinops where they are free, but in G. moliiccana and G. decipiens they are usually united.
Dr B. Peterson, Lund, who is working on the African Thymelaeaceae, told us of similar difficulties
encountered in defining genera in that area. He wrote (May 1959): "I have devoted much time to generic
delimitation in this family. As I have examined more and more African material (c. 15.000 sheets)
I have found that the limits are in some cases so vague that it has appeared unavoidable to merge several

genera. It is often rather easy to give a specific epithet but very difficult to come to a decision of the
generic name. For example the only generic characters in Gnidia, Lasiosiphon, and Arthrosolen, and some
smaller genera, are the number of calyx lobes and the presence or absence of petaloid appendages. And
these are not at all enough to keep these genera separate. Sometimes these characters do not even hold
for the type species. Gilg and later Staner proposed that these genera should be united but other
botanists have not followed their suggestions. In my monograph of Gnidia I will merge seven genera."
Aquilaria and Gyrinops seem to be very closely allied, the first being diplostemonous, the second
haplostemonous, which is the only constant character. Hallier /. found this difference not sufficient
for generic distinction and united these genera. Dr Peterson found in Africa a similar case, viz between
Gnidia and Struthiola of which the first is diplostemonous, the second haplostemonous. He "never
found any trace of staminodes in Struthiola. In some species of Gnidia, however, usually, but not always,
the upper whorl of stamens is abortive. All species oi Struthiola have a whorl of hair round each petaloid
appendage. This arrangement is not found in Gnidia except for a single species as far as I have found.
This will be placed in a separate section."
In Dr Peterson's opinion Struthiola and Gnidia, though properly only distinguished 'absolutely' by
one character, should not be united; if that were done, the consequence would be that still more genera
had to be merged in the complex which would lead to an unsatisfactory situation. In this revision I
have not followed Hallier /. in uniting Aquilaria and Gyrinops.
The difference between Wikstroemia and Daphne seems, by being merely vegetative, still more feeble,
the chief distinction being the opposite phyllotaxis in Wikstroemia, notwithstanding the note by Staff
(Bot. Mag. 156, 1933, sub t. 93 13, p. 2). If it is realized that the phyllotaxis varies widely within the single
genus Pimelea, it is tempting to merge Wikstroemia and Daphne.
The merging of Aquilaria and Gyrinops and of Daphne and Wikstroemia might give a better reflection
of the natural affinities, as the single character separating the components of these pairs effects, in my
opinion, not a natural segregation.
Specific delimitation in Thymelaeaceae is in many cases also extremely difficult, specially because it
has appeared that characters not only vary within a single species, but also within the flowers of one
single specimen, as for example the shape of the disk in Wikstroemia aurantiaca {cf. Staff, I.e.). I have
encountered several similar cases in other species and Dr Peterson communicated to have a similar
experience with African representatives which has led him to a severe reduction of accepted species.
Specific delimitation in Malaysia proved particularly difficult in Wikstroemia and Phaleria; for
W. indica I have accepted much wider specific limits than my predecessors.
Affinities with other families.— For a detailed review and discussion of the relationship of Thyme-
laeaceae and other families, one should consult the works of Domke (Bibl. Bot. Ill, 1934, 1-3, 16) and
Heinig (Am. J. Lot. 38, 1951, 113 & 131).
According to Miss Heinig's studies on the floral morphology the polypetalous and polystemonous
condition of the primitive members of the Thymelaeaceae and the modified parietal placentation suggest
a derivation from some polymerous parietalean family such as, possibly, the Flacourtiaceae; there seems
also a possible relationship with the Tiliaceae.
Erdtman (Pollen Morph. & Pi. Tax. 1952, 43) stated that there is a more or less close relationship
between Thymelaeaceae and Euphorbiaceae, especially the crotonoid members of the latter.
Uses. The heartwood of some species of Aquilaria and Gonystylus contains aromatic substances and
is used as incense {cf. Burk. Diet. 2, 1935, 197-205). The scented portions are only found in irregular

small parts of the heartwood and are obviously caused by some abnormality (infection by fungi or
insects?) and they occur not in all trees. Schuitemaker described the occurrence of scented thymelaea-
ceous wood in West Borneo and the ceremonials connected with the collecting of it (Tectona 26, 1933,
 Dec. 1960] Thymelaeaceae (Ding Hou) 5

851-892, fig. 1-6). The strong barks of some species are used for weaving, walls of huts, paper-making,
and tying purpose. Wood of Gonystylus bancamis is used for internal building construction; it is one of
the most important timber exports of Sarawak and Brunei.
Note. Sterile material has a limited value and can sometimes hardly be identified even to the genus
with certainty, viz in Aquilaria-Gyrinops and Phaleria-Wikstroemia. Flowering or fruiting material is
essential for identification.
I am indebted to Dr J. Leandri for putting his valuable manuscript notes at my disposal.

KEY TO THE GENERA

Based on flowering and fruiting material ^

1. Leaves not pellucid-dotted. Stamens and petaloid appendages adnate to or inserted on the floral tube.
Fruits 1-2-celled.
Ligneous, perennial. Inflorescences without involucral bracts, or (in Phaleria and Daphne composita)
2.
with free ones. Stamens 4 or more.
3. Stamens twice the number of the calyx lobes.

4. Fruit a loculicidal capsule. Petaloid appendages usually distinct and always densely pubescent
or puberulous 1. Aquilaria

4. Fruit a drupe or drupaceous. Petaloid appendages if present always glabrous.

5. Ovary 2-celled (rarely one cell abortive in Phaleria perrottetiana). Fruits (l-)2-seeded. (Petaloid

appendages none, or obscure and rim-like.) 2. Phaleria

5. Ovary always 1 -celled. Fruits 1 -seeded.
6. Usually climbing shrubs. Inflorescences usually provided with 2 leafy bracts on each branch.
Petaloid appendages well developed. Ovary densely pubescent.
7. Stamens in two series. Style obscure or shorter than the ovary ....
3. Enkleia
7. Stamens in one series. Style several times as long as the ovary ....
4. Linostoma
6. Erect shrubs. Inflorescences without leafy bracts. Petaloid appendages none. Ovary glabrous
or only hairy at the top.
8. Leaves opposite. Disk lobed, scale-like, lobes free or united in pairs 5. Wikstroemia .

8. Leaves alternate. Disk ring-like or cup-shaped 6. Daphne

3. Stamens the same number as the calyx lobes.
9. Shrubs or trees. Leaves ovate-oblong to lanceolate, 1 V^-24 by (1/3-) 1-3 cm. Ovary densely hairy,
2-celled; style terminal. Fruit a loculicidal capsule, protruding either from the top or from the
split side of the floral tube 7. Gyrinops
9. Dwarf-shrub. Leaves linear, 3-5 by 2/3 mm. Ovary hairy at the upper half or only at the top,
1 -celled; style lateral. Fruit a drupe, developing inside the floral tube 8. Drapetes...
2. Annual herbs. Inflorescences with 4, partly united involucral bracts. Stamens 2 9. Pimelea .

1. Leaves pellucid-dotted. Stamens free. Petaloid appendages inserted on the receptacle. Fruits 3-5

(-8)-celled.
10. Leaves decussate, sometimes some subopposite; nervation lax and open. Flowers subumbellate.
Calyx lobes valvate. Petaloid appendages fused in a ring. See vol. 4, p. 365 11. Aetoxylon .

10. Leaves spiral or alternate. Inflorescences thyrsoid or racemose. Calyx lobes imbricate or subvalvate.
Petaloid appendages 7-40.
11. Leaves with few, spaced nerves. Petaloid appendages 10, more or less in pairs. Parastyles subulate-
corniform. Fruits long-beaked. See vol. 4, p. 363 and this vol. p. 47 10. Amyxa ...
11. Leaves with numerous parallel nerves, veins distinctly prominent. Petaloid appendages 7-40,
not in approximate pairs. Parastyles if present very small and clavate. Fruits not beaked. See vol. 4,
p. 350 12. Gonystylus

KEY TO THE GENERA

Based on sterile material
1. Leaves not pellucid-dotted.
2. Ligneous, perennial plants.
3. Leaves at least 15 mm
long, penninerved, not linear, at least 1 mm petioled. Lowland or montane
shrubs or trees.
4. Lateral nerves and intermediate veins more or less parallel.
5. Leaves alternate or spiral. Erect shrubs or trees 1. Aquilaria 7. Gyrinops &
5. Leaves opposite, rarely also with some subopposite ones. Liana, very rarely erect shrubs or
small trees 4. Linostoma
4. Lateral nerves curved, intermediate veins reticulate or cross-bar like (Enkleia).
6. Leaves strictly opposite or decussate 2. Phaleria 5. Wikstroemia &
6. Leaves alternate or spiral, or at least not all strictly opposite.

(1) In some genera the floral characters can usually easily be studied in the fruiting state as the floral
parts are generally persistent.
6 Flora Malesiana [ser. I, vol. 6^

7. Liana, often provided with hooks. Cross-bar veins subparallel 3. Enkleia

7. Erect shrub or small tree. Venation reticulate 6. Daphne
3. Leaves small (3-5 by 2/3 mm), with 7-9 more or less parallel, longitudinal nerves, sessile, linear.
Subalpine dwarf-shrub 8. Drapetes
2. Annual herbs 9. Pimelea
I. Leaves pellucid-dotted.
8. Leaves opposite or subopposite. See vol. 4, p. 365 11. Aetoxylon
8. Leaves alternate or spiral.
9. Leaves with few, spaced nerves, veins rather obscure. See vol. 4, p. 363 and this vol. p. 47
10. Amyxa
9. Leaves with numerous more or less parallel nerves, veins distinctly prominent. See vol. 4, p. 350.
12. Gonystylus

1. AQUILARIA
Lamk, Encycl. 1 (1783) 49, nom. gen. conserv.; ibid. 2 (1786) 610; Domke, Bibl.

Bot. Ill (1934) 118, map 2; Quis. J. Arn. Arb. 27 (1946) 402.— Agallochum
RuMPH. ex Lamk, Encycl. 1 (1783) 48, nom. gen. rejic.—Ophispermum Lour. F1.
Coch. 1 (1790) l%\.—Gyrinopsis Decne, Ann. Sc. Nat. Bot. II, 19 (1843) 41;
Quis. J. Am. Arb. 27 (1946) 404.— Decaisnella O.K. Rev. Gen. PI. 2 (1891)
5^4.— Aquilariella van Tiegh. Ann. Sc. Nat. Bot. VII, 17 (1893) 216; Bull. Soc.
Bot. Fr. 40 (1893) 77. Aquilaria sect. Agallochum Hallier /. Med. Rijksherb.
n. 44 (1922) 15. Aquilaria sect. Gyrinopsis Hallier/. I.e. 16. Aquilaria sect.
Amphinoman Hallier /. I.e. 18. —Fig. 1.
Shrubs, treelets or trees. Innovations always pubescent but usually glabrescent.
Leaves on the lateral twigs alternate, penninerved nerves distinct or obscure,;

simple or sometimes branched, usually shghtly curved, ascending towards the

margins and joining several intramarginal veins; veins and veinlets numerous,
parallel or subparallel; margins wavy, slightly recurved and thickened. Inflores-
cences axillary or supra-axillary, sometimes on internodes, terminal, or rarely
cauliflorous, sessile or short-peduncled, simple or rarely branched, umbelHform
or paniculiform, usually without bracts, rarely with a few small ones. Flowers
usually 5-merous, pedicelled, articulated at the base of the pedicel. Floral tube
cupular to tubular, persistent, in fruit sometimes splitting on one side, outside
puberulous or pubescent, inside puberulous with reflexed hairs arranged in
lengthwise lines towards the upper part. Calyx lobes (4-)5(-6), reflexed or erect,
usually shorter than or rarely as long as the tube. Petaloid appendages twice as
many as the lobes, free, or united in a ring {A. cumingiana), inserted at the throat
of the tube, lanceolate, ovate, semi-orbicular, or rim-like (A. urdanetensis), each
pair opposite the calyx lobe, usually densely pubescent or puberulous. Stamens
twice as many as calyx lobes, emerging from the tube at the same level as the
appendages, rarely emerging shghtly below them, sometimes behind them,
sessile or filamentous, equal in length or sepalous ones longer than the others;
filaments filiform, sometimes shghtly swollen at the upper end; anthers linear-
oblong, dorsifixed, free (but in A. cumingiana the lower I/3-V2 adnate to the tube);
connective broad over the whole length of the anther. Disk none or rarely ring-
like. Pistil included. Ovary sessile or stiped, ovoid, oblanceolate or ellipsoid,
densely short-puberulous, 2-celled (or incompletely 2-celled in extra-Mai. spp.);
style terminal, obscure or distinct, gradually dilated to the ovary, densely
puberulous towards the base; stigma distinct, globose, capitate, pyramidal, or
oblong, black. Ovule attached near the top of the septum and partly adnate to
it. Fruit a loculicidal capsule, globose, obovoid, or oblanceolate, rugose or smooth,
Dec. 1960] Thymelaeaceae (Ding Hou)

Fig. 1. Aquilaria beccariana van Tiegh. a. Habit, x 2/3, b. opened flower, two anthers removed, X 3,
c. dehiscing fruit emerging from top of floral tube with one seed dangling out, nat. size. A. brachyantha
(Merr.) Hall./, d. Opened flower, x 3. A. hirta Ridl. e. Opened flower, X 3,/. dehisced fruit, nat.
size. A. microcarpa Baill. g. Dehisced fruit, nat. size. A. malaccensis Lamk. /;. Dehisced fruit, nat.
size. A. cumingiana (Decne) Ridl. /. Dehisced fruit emerging from lateral slit of floral tube, one
seed dangling out, nat. size (a-b San A 1726, c SF 29381, d FB 19562, e Bunnemeyer 7575, /Cuming
1617, g San 16965).

sometimes slightly compressed laterally, protruding either from the top or from
the split side of the floral tube, distinctly stalked, densely puberulous to glabrous;
pericarp coriaceous or woody. Seeds 2, or 1 by abortion, ovoid or ellipsoid; testa
crustaceous, sometimes downy, bearing a caruncle-like or tail-shaped appendage
at the base; usually the whole seed and sometimes a portion of the appendage is
adnate to the septum, with an obscure or a distinct funicle; in the latter
laterally
on the end of the thin funicle in open capsules;
case the seeds dangle out of the fruit
albumen none or scant; cotyledons thick, plano-convex.
Distr. About 15 spp., India (Bengal and Assam), Burma (Tenasserim), Indo-China (Cambodia,
Annam, and Cochinchina), China (Hongkong and Hainan), and widely distributed in Malaysia.
Ecol. In forests at low and medium altitudes, some species occurring from 1000-1700 m.
8 Flora Malesiana [ser. I, vol. 6^

KEY TO THE SPECIES

1. Flowers cupular or bell-shaped, 4-6 mm
long, the lobes usually as long as the tube. Stamens distinctly
filamentous, filaments of the episepalous ones at least as long and usually longer than the anthers.
2. Calyx lobes reflexed in anthesis. Ovary densely pubescent; style absent or obscure.
3. Fruits obovoid, 3-4 by 2'/^ cm. Seed with a tail-like, slightly twisted and pubescent appendage c.
10 mm long. Episepalous stamens longer than the petaloid appendages 1. A. malaccensis
.

3. Fruits slightly obcordate, ^-IVi by 1-1 Va crn. Seed with a caruncle-like, glabrous appendage c.
2 mmlong. Episepalous stamens usually shorter or as long as the petaloid appendages.
2. A. microcarpa
2. Calyx lobes always erect. Ovary slightly pubescent; style distinct, filiform and almost as long as
the ovary 3. A. brachyantha
1. Flowers short-tubular to cylindric, (5-6-) 7-1 5 mm
long, the lobes usually V2-V5 the length of the tube.
Stamens sessile or subsessile, filaments rarely up to V2 as long as the anthers, in A. urdanetensis the
episepalous ones as long as the anthers.
4. Calyx lobes c. 1/2 the length of the tube. Seed with a short caruncle-like appendage at the base.
5. Petaloid appendages obscure, rim-like. Filaments of the episepalous stamens sometimes as long
as the anthers. Style distinct. (Fruits globose, contracted at the base into a distinct stalk.)
4. A. urdanetensis
5. Petaloid appendages distinct, semiorbicular or ovate to oblong, V1-V2 as long as the anthers.
Stamens sessile or subsessile. Style absent or very short.
6. Flowers 8-10 mm long, densely pubescent outside. Ovary slightly obovate-oblong, truncate at
the apex, densely covered with a layer of densely set, reflexed, short hairs mixed with some ap-
pressed, straight, long hairs. Stigma pyramidal, sessile. Leaves densely pubescent beneath.
5. A. citrinaecarpa
6. Flowers 5-6 mm long, sparsely puberulous or glabrous outside. Ovary slightly elliptic-oblong,
gradually narrowed towards the top, covered only with densely set, appressed, straight long hairs.
Stigma capitate or globose, on a very short style. Leaves slightly pubescent, glabrescent, or
glabrous beneath, very rarely densely pubescent.
7.Fruit with a distinct stipe as long as or longer than the floral tube. Floral tube usually not splitting
in fruit. Pedicels at least as long as the flowers 6. A. apiculata
7. Fruit sessile or on a short stipe (c. 2-3 mm), not longer than the floral tube. Floral tube in fruit
splitting on one side. Pedicels usually shorter than the flowers.
8. Fruits slightly obovoid or broadly ellipsoid, gradually narrowed to the base, sessile or some-
times with a very short stipe. Floral tube in fruit sometimes transversely curved and calyx lobes
usually reflexed. Leaves 10-20 by 3-51/2 cm, the nerves scarcely distinguishable from the
intermediate veins 7. A. filaria
8. Fruits globose, contracted at the base into a short, slender stipe. Floral tube usually flat in fruit
and lobes erect. Leaves 4y2-\5 by 1-41/2 cm, nerves 7-12 pairs, distinct from the intermediate
veins 8. A. parvifolia
4. Calyx lobes V3-V5 the length of the tube, c. ^/g-Vs in A. beccariana. Seeds with an elongated or tail-
like appendage (except in A. cumingiana).
9. Fruits oblanceolate, 2-31/2 by 1-1 24 cm, attenuate to the base and narrowed into a stipe which is
usually longer than the floral tube. Seeds ovoid or ellipsoid-oblong, brownish hairy or puberulous,
with a tail-like appendage. Petaloid appendages free and inserted at the same level as the stamens.
Anthers free from the floral tube.
10. Undersurface of the leaves and the fruits densely pubescent. Leaves acute. Petaloid appendages
deltoid, 1/3-I/2 the length of the anther, long-hairy, the hairs as long as the appendages or longer.
Seeds cuneate to the base and attached to a glabrous, elongate appendage c. 10 mm
long.
9. A. hirta
10. Lower surface of the leaves and the fruits sparsely pubescent, glabrescent, or glabrous. Leaves
acuminate.
11. Seeds narrowed to the base and elongated into a long (c. 15 mm), glabrous or subglabrous
appendage. Petaloid appendages unknown 10. A. rostrata
11. Seeds narrowed to the base and separated from the tail-like, hairy appendage c. 10 mmlong
by a short, thin stipe-like constriction. Petaloid appendages oblong, almost as long as the
stamens, shortly puberulous 11. A. beccariana
9. Fruits subglobose, globose, slightly obovoid or ellipsoid, 1% by 1 1/3 cm, contracted at the base,
sessile or with an obscure stipe. Seeds broadly ovoid, planoconvex, glabrous, c. 1 by ^4 cm, with a
small caruncle-like appendage. Petaloid appendages short, usually united in a ring. Lower 1/3-I/2 of
the anthers usually adnate to the floral tube 12. A. cumingiana
Dec. 1960] Thymelaeaceae (Ding Hou)

1. Aquilaria malaccensis Lamk, Encycl. 1 (1783) seed, separated from it by a short, thin, stipe-like
49, t. 356; DC. Prod. 2 (1825) 59; Meisn. in DC. constriction.
Prod. 14 (1857) 602, exd. citat. of Benth.; Miq. Distr. India (Bengal and Assam), Burma
Fl. Ind. Bat. 1, 1 (1858) 883; Kurz, Nat. Tijd. (Tenasserim), and Malaysia: Sumatra (Simalur,
N.I. 27 (1864) 171; For. Fl. Burm. 2 (1877) 236; Sibolangit, Palembang, and Banka), Malay
RiDL. Trans. Linn. Soc. Lond. 11, 3 Bot. (1893) Peninsula (common), N. & E. Borneo, and the
341; GiLG, Bot. Jahrb. 18 (1894) 506, f. 8, B; Philippines (Luzon).
ibid. 28 (1900) 145; Boerl. Handl. 3 (1900) 112; Ecol. Primary forests at low and medium
RiDL. J. Str. Br. R. As. Soc. /;. 35 (1901) 73; altitudes up to 270 m.
Gamble, J. As. Soc. Beng. 75, ii (1912) 264; Uses. According to Heyne, I.e. and Ridley
KooRD. Exk. Fl. Java 2 (1912) 656 (erron. record); (1901, the tree yields a celebrated incense
I.e.)
Merr. Philip. J. Sc. 10 (1915) Bot. 44; Int. Rumph. wood which is obtained from the center of an old

(1917) 381; Brown, Minor Prod. Philip. Forests 1 or dying tree. It is said to be caused by a disease
(1920) 403; Merr. En. Born. (1921) 417; Hall./. which gains entry through the old decaying
Med. Rijksherb. n. 44 (1922) 16; Merr. En. branches. Its greatest use has always been for
Philip. 3 (1923) 130; Ridl. Fl. Mai. Pen. 3 (1924) fumigating and it is highly valued in the Orient
147; BuRK. & Henders. Card. Bull. S.S. 3 (1925) for ceremonial purposes. It also furnishes a
417; Heyne, Nutt. PI. ed. 2 (1927) 1 149; Henders. beautiful, silvery bast used for making ropes and
Card. Bull. S.S. 4 (1928) 314; Metcalfe, Kew clothes. The bast is highly prized for its strength
Bull. (1933) 5; Corner, Ways. Trees (1940) and durability.
632; Quis. J. Arn. Arb. 27 (1946) 403; Merr. J. Vern. Calambac, clung karas, gaharu, galoop,
Arn. Arb. 31 (1950) 270. Agallochuin secundarium garu, karas, kayu gaharu, kekaras, kepang, laroo,
coiuamense & A. malaicense Rumph. Herb. Amb. mengkaras, tabak, taras gharu, tengkaras, M,
2 (1741) 34-35, t. \Q.—A. ovata Cav. Diss. (1789) sigsigi,Borneo; Sumatra: alim, Batak, halim.
377, t. 22A.—A. secundaria DC. Prod. 2 (1825) 59; Lamp., kareh, Minangk.; Malayan eaglewood tree,
Miq. Fl. Ind. Bat. 1, 1 (1858) 883; Boerl. Handl. E.
3 (1900) \\1.— Agallochuin malaccense O.K. Rev. Note. The vegetative parts of this species are
Gen. PI. 2 (1891) 583. Aquilariella malaccensis similar to those of ^4. microcarpa in the herbarium,
van Tiegh. Ann. Sc. Nat. Bot. VII, 17 (1893) 216; and I cannot find any good character to separate
Bull. Soc. Bot. Fr. 40 (1893) 77.— Fig. Ih. them.
Tree up to 40 m by 60 cm. Bark smooth,
whitish; branchlets slender, pale brown, pubes- 2. Aquilaria microcarpa Baill. Adansonia 1
cent, glabrescent. Leaves chartaceous, sub- (1875) 304; Gilo, Bot. Jahrb. 28 (1900) 145;
coriaceous, glabrous, sometimes pubescent be- Merr. En. Born. (1921) 417; Domke, Bibl. Bot.
neath, glabrescent, shining on both surfaces, 1 1 1 (1934) t. 4 f. 43f; Quis. J. Arn. Arb. 27 (1946)

elliptic-oblong to oblong-lanceolate, 7V2-12 by 403. Aquilariella microcarpa van Tiegh. Ann.

2'/2-5'/2 cm; base acute, attenuate, or obtuse; Sc. Nat. Bot. VII, 17 (1893) 216; Bull. Soc. Bot.
apex acuminate, acumen up to 2 cm; nerves 12-16 Fr. 40 (1893) 77. Aquilariella borneensis van
pairs, rather irregular, often branched, curving Tiegh. Ann. Sc. Nat. Bot. VII, 17 (1893) 217;
upward, elevated beneath, plane or obscure above, Bull. Soc. Bot. Fr. 40 (1893) 11.— A. borneensis
veins distinct beneath, invisible above; petiole Gilg in E. & P. Pfl. Fam. Ill, 6a (1894) 224;
4-6 mm. Inflorescences terminal, axillary or supra- Boerl. Handl. 3 (1900) 112; Merr. En. Born.
axillary, sometimes on internodes, usually branch- (1921) 417.— Fig. Ig.
ed with 2 or 3 umbels and each with about 10 Tree up to 40 m
by 80 cm. Bark grey, super-
flowers, more rarely a simple umbel; peduncle or ficially fissured;branchlets light brown, puberu-
common peduncle 5-15 mm; pedicels slender, lous, glabrescent. Leaves subcoriaceous. shining
3-6 mm. Flowers green or dirty-yellow, cam- and glabrous above, rather dull, glabrous or
panulate, 5-6 mm long, scattered puberulous sometimes scattered hairy beneath, elliptic-oblong
outside. Floral tube nearly glabrous within, to obovate-oblong or oblanceolate, 4V2-10 by
distinctly 10-ribbed. Calyx lobes ovate-oblong, 1 V2-4'/2 cm; base cuneate to attenuate; apex acute

2-3 mm long, densely puberulous within, almost to acuminate, the acumen up to 1 cm; nerves
as long as the tube, reflexed. Petaloid appendages 12-19 pairs, sometimes branched, slightly curved
oblong or slightly ovate-oblong, c. 1 mm long, and ascending to the thickened margin, pro-
slightly incurved, densely pilose. Stamens 1 Y^-l minent beneath, visible above; veins and veinlets
mm long, the episepalous ones longer than the rather irregular, subparallel, distinct beneath,
others; anthers linear, obtuse, as long as or shorter obscure above; petiole 3-5 mm, pubescent.
than the filaments. Ovary ovoid, 1-1 '/2 mm long, Inflorescences axillary or supra-axillary, terminal,
densely pubescent; style obscure; stigma capitate. or on short lateral branchlets, usually branched,
Fruits obovoid or obovoid-oblong, rounded at the rarely simple, peduncle short or up to 1 cm,
apex, cuneate to the base, 3-4 by IVi cm, usually 6-1 1-flowered. Flowers white, light-yellow or
compressed, pubescent outside, glabrescent; peri- yellow. 5 mm
long; pedicels c. 5 mm, puberulous.
carp woody, the suture face c. 6 mm wide. Seed Floral tube puberulous outside, sparsely puberu-
proper ovoid, including the beak 10 by 6 mm, lous inside. Calyx lobes ovate or oblong, obtuse,
densely covered with red hairs, the beak c. 4 mm densely puberulous on both surfaces. Petaloid
long, the appendage twisted and as long as the appendages almost as long as the stamens or
10 Flora Malesiana [ser. I, vol. 6^

sometimes slightly longer, ovate or oblong, densely elliptic-oblong to broadly lanceolate, 4-9 by
hairy. Stamens 1-1 '/a mm, alternately long and 1 1/2-3^ cm; base cuneate to attenuate; margins
short; anther c. '/a mm, usually shorter than the slightly thickened and shining on both surfaces;
filament, rarely as long or longer. Pistil ovoid, apex acuminate, the acumen up to 1 cm, pointed
IV2-2 mm long. Ovary densely pubescent; style or obtuse at the tip; nerves and veins undistin-
obscure or rarely very short; stigma capitate. guishable, numerous, divaricate, subparallel, some
Fruits subcordate, slightly compressed, 8-12(-16) of them branched, distinct beneath, obsolete
by 10-12(-15) mm, l-(2-)seeded; persistent floral above; petiole 3-4 mm. Inflorescences axillary,
tube sometimes splitting on one side. Seed ovoid, sessile or very short peduncled, usually with a few,
6 by 4 mm, densely brownish pubescent; caruncle- very small, caducous bracts, 2- or 3-flowered;
like appendage 2 mm long. pedicels c. 31/2 mm, sparsely pubescent. Flowers
Distr. Malaysia: Malay Peninsula (Singapore), short-tubular, 5-6 mm long, yellowish. Floral
Sumatra (Sidjungdjung, Palembang, and tube 31/2-4 mm long, sparsely pubescent outside,
Lampongs), Billiton, Banka, and throughout pubescent towards the base and at the mouth
Borneo. inside. Calyx lobes 5(-6), ovate and obtuse,
Ecol. Lowland forests up to 200 m. sparsely pubescent outside, densely puberulous
Vern. Sumatra: tengkaras, M, liepang, Banka; inside, 1 1/2-2 mm long. Petaloid appendages
Borneo engkaras, Dayak, karas or sigi-sigi, Bugis,
: obscure, rim-like. Stamens free from the tube
kumbil, garu, tulang, M. slightly below the petaloid appendages, 2/3-1 V2
mm, episepalous ones with a filament shorter than
3. Aquilaria brachyantha (Merr.) Hall. /. Med. the anther or as long as it, the others sessile.
Rijksherb. /;. 44 (1922) 16; Quis. J. Arn. Arb. 27 Pistil c. 2 mm long. Ovary ellipsoid, densely
(1946)403. Gyrinopsis brachyantha Merr. Philip. puberulous, narrowed into a short style c. 1 mm;
J. Sc. 7 (1912) Bot. 313; Elm. Leafl. Philip. Bot. 5 stigma nipple-like. Fruits globose or slightly
(1913) 1629; Merr. En. Philip. 3 (1923) 130; obovate, glabrous when mature, c. 8 mm diam.,
DoMKE, Bibl. Bot. 1 1 (1934) t. 2, f. 8; t. 5, f. 43e.
1 with a slender stipe 3-6 mm; persistent floral tube
—Fig. Id. splitting on one side. Seed ovoid, plano-convex,
Small tree or shrub up to 2 m. Branchlets black, c. I cm long, with a short caruncle-like
glabrous, yellowish brown to brownish when dry, appendage.
the tips usually pubescent. Leaves chartaceous to Distr. Malaysia: Philippines (Mindanao: Mt
subcoriaceous, shining on both surfaces, oblong, Urdaneta), twice collected.
elliptic-oblong, or lanceolate, 8-16 by 2-4'/2 cm; Ecol. In the mossy forest on exposed ridges,
base acute or obtuse; apex acuminate; nerves and c. 1700 m {cf. Merr. I.e.).

veins numerous, homogeneous, slightly elevated Vern. Mahgod, makolan, Mbo.

beneath, obscure or invisible above; petiole c. Note. Known only from the authentic collec-
5 mm. Flowers greenish, small, axillary, 1 to tions, Elmer 14195 (lectotype) and 13742 (para-
several in a fascicle on a very short peduncle; type).
pedicels 1-3 mm, pubescent. Floral tube cam-
panulate, 3-4 mm long, pubescent or puberulous 5. Aquilaria citrinaecarpa (Elmer) Hall. /. Med.
on both surfaces, usually glabrescent outside. Rijksherb. //. 44 (1922) 18. Gyrinopsis citrinae-
Calyx lobes 5, slightly oblong or ovate-oblong, as carpa Elmer, Leafl. Philip. Bot. 5 (1913) 1631;
long as the tube or sometimes slightly longer. Merr. En. PhiUp. 3 (1923) 130; Quis. J. Arn. Arb.
Petaloid appendages linear or oblong, c. 1 long, mm 27 (1946) 405.
densely pubescent. Stamens 1-1 1^ long, mm Small tree up to 8 m. Young branches densely
filamentous, the episepalous ones slightly longer olivaceous tomentose, glabrescent. Leaves sub-
than the others. Ovary ovoid, c. 1 1/2 long, mm coriaceous, dull, olivaceous, and densely pubes-
slightly pubescent; style distinct, filiform, c. 1 mm; cent beneath, shining, reddish-brown, and glabrous
stigma capitate. Fruits narrowly obovoid, com- above, elliptic-oblong, or slightly obovate-oblong,
pressed, 1 14-1 1/2 by 3/^-1 cm. Seed including the 6-lO(-12) by 2i/2-4(-5i/2) cm; base cuneate; apex
caruncle-like appendage c. 1 cm long, pubescent, acute; margins slightly recurved; nerves 15-20
persistent floral tube splitting on one side. pairs, on the lower surface obscure, rarely distinct,
Distr. Malaysia: Philippines (Luzon: Cagayan slightly ascending towards the margin; veins and
Prov.), twice collected. veinlets obscure or visible beneath, obscure above;
Ecol. In primary forests at low altitudes. petiole c. 3 mm. Inflorescences terminal and
axillary, sessile or with a very short peduncle,
4. Aquilaria urdanetensis (Elmer) Hall. /. Med. densely hairy, usually with a few small bracts,
Rijksherb. /;. 44 (1922) 16. Gyrinopsis urdanetense 3-6-flowered; pedicels 2-4 mm, pubescent.
Elmer, Leafl. Philip. Bot. 5 (1913) 1630; Merr. Flowers 8-10 mm long, greenish. Floral tube
En. Philip. 3 (1923) 131; Quis. J. Arn. Arb. 27 cylindric, densely pubescent outside and towards
(1946) 405. the base inside. Calyx lobes oblong or ovate,
Slender shrub, up to 7 m. Bark dull grey and 3-31/2 mm long, densely puberulous on both sur-
smooth. Young branchlets whitish pubescent, faces. Petaloid appendages oblong or ovate, about
glabrescent. Leaves chartaceous, shining on both 14 the length of the anthers, densely villose.
surfaces, young leaves pubescent beneath es- Stamens sessile, 1 1/2 mm long. Ovary slightly
pecially on the nerves and veins, glabrescent. obovate-oblong, densely puberulous, c. 3 mm
Dec. 1960] Thymelaeaceae (Ding Hou) 11

long, style none; stigma pyramidal, black. cent. Leaves subcoriaceous, glabrous or scattered
Capsules 1 %by 1 14 cm, bright yellow or citrine, hairy rarely pubescent beneath, oblong, elliptic-
obtuse to subtruncate at the apex, attenuate to the oblong to lanceolate, rarely oblanceolate-oblong,
base, sometimes slightly compressed, persistent 10-20 by 3-5 '/2 cm; base obtuse to cuneate; apex
floral tube splitting on one side. Seeds deltoid, 8 shortly acuminate; nerves and veins usually homo-
by 7 mm, plano-convex, acute to the apex, almost geneous, slightly elevated beneath, obscure above;
truncate at the base, with a very short caruncle- petiole 3-5 mm. Inflorescences axillary and extra-
like appendage at the base. axillary, umbelliform or condensed paniculiform,
Distr. Malaysia: Philippines (Mindanao), once rarely cauliflorous, very short-peduncled, (l-)3-7
collected. (-cv))-flowered pedicels 2-5
;
mm, pubescent.
Ecol. On moist, compact soil of forested ridges, Flowers yellowish-green or white, infundibular,
c. 1300 m. 5-6 '/2 mm
long. Floral tube sparsely pubescent
Vern. Agododan, Mbo. outside, glabrescent. Calyx lobes oblong to slightly
ovate, c. 2 mm
long, densely puberulous on the
6. Aquilaria apiculata Merr. Philip. J. Sc. 20 upper part and the margins outside, and the whole
(1922) 41 1 ; En. Philip. 3 (1923) 30; Quis. J. Arn.
1 surface Petaloid appendages oblong or
inside.
Arb. 27 (1946) 403. deltoid, c. 1 mm
long, densely villous. Stamens c.
Shrub or small tree up to 3 m. Branchlets 1 mm long, the episepalous ones with short, fleshy
reddish brown, pubescent, glabrescent. Leaves filaments, the others sessile or subsessile. Ovary
papery, glabrous above, sparsely or scattered obovoid, c. 3^4 mm
long, densely villous; style
pubescent beneath, elliptic-lanceolate, 8-14 by very short or obscure; stigma capitate. Fruits
2Vi-4 cm; base cuneate to attenuate; apex acu- ellipsoid to obovoid or subglobose, slightly com-
minate, the acumen up to 2 cm, tip obtuse; nerves pressed, rugose, 1 y^-l '4 by 1 14 cm, sparsely hairy,
8-16 pairs, curved and ascending to the margin, glabrescent, narrowed to the base, sometimes on a
prominent beneath, visible above; veins distinct, very short stipe, yellow. Seeds deltoid, including
sometimes visible beneath, obscure above; petiole the appendage c. Y^ by Y^ cm, plano-convex, black,
3-5 mm. Inflorescences axillary, sessile or short- with a very short caruncle-like appendage.
peduncled, usually with a few small bracts,
3-6-flowered; pedicels 6-7 mm, puberulous.
Flowers 5-6 mmlong. Floral tube short-tubular,
4-5 mm long, puberulous outside and inside,
glabrescent. Calyx lobes ovate to oblong, 1 V2-2
mm long, densely puberulous on both surfaces.
Petaloid appendages semi-orbicular to ovate,
V1-V2 the length of the stamens, hairy. Stamens
c. 1 mm long, sessile or the episepalous ones on
short filaments. Ovary slightly obovate, 3 mm
long, densely pubescent, slightly attenuate towards
the base, acute and narrowed towards the apex;
style very short; stigma capitate. Fruits yellowish
orange, ellipsoid, slightly compressed, developing
on top of a slender V2-\ cm long stipe, protruding
from the floral tube, 1 2/3 by 1 cm persistent floral
;

tube entire or sometimes splitting at one side.

Seeds ovoid, 8-9 by 6 mm, dark-brown, with a
caruncle-like appendage c. 2 mm
long. Fig. 2. Localities of Aquilaria filaria (Oken)
Distr. Malaysia: Philippines (Mindanao: Merr.
Bukidnon Prov.).
Ecol. In dry and mossy forests, 1100-1800 m. Distr. Malaysia: Philippines (Dinagat I. and
Bucas Grande I.), Moluccas (Morotai, Ceram, and
7. Aquilaria filaria (Oken) Merr. J. Arn. Arb. 31 Ambon), and New Guinea (Sorong, Babo, and
(1950) 283, excl. syn. Gyrinopsis brachyantha Kapor). Fig. 2.
—
Merr. Cortex filarius Rumph. Herb. Amb. 7 Ecol. In lowland forests, once collected in open
(1755) 13. Pittosporum ferrugineum var. (i swamp forest (Sorong: Pleyte 393), up to 130 m.
filarium DC. Prod. 1 (1824) 347, excl. Rumph. t. 7 Vern. Age, Sorong, bdkuin, Morotai, lason,
cit.\ Don, Gen. Hist. 1 (1831) ilA.— Pittosporum Ceram, kasjik, Tehid lang., malowassi, Uliassers.
filarium Oken, Allg. Naturgesch. 3^ (1841) 299.— Notes. In the description of Cortex filarius
A. tomentosa Gilg, Hot. Jahrb. 28 (1900) 145.— Rumph. (Herb. Amb. 7, 1755, 13) Rumphius re-
Gyrinopsis brachyantha {non Merr. 1912) Merr. corded the bark with strong bast and the leaves
Int. Rumph. (1917) 380, quoad specim. —
Gyrinopsis with more or less parallel veins which agree with
acuminata Merr. Philip. J. Sc. 17 (1920) 294; the characters of Aquilaria. He described the fruit
En. Philip. 3 (1923) 130.-/4. acuminata Quis. J. as 2-celled; one of the cells being empty and filled
Arn. Arb. 27 (1946) 403. with pulp, the other having two seeds. However,
Shrub or tree up to 17 m
by 50 cm. Young in Aquilaria the ovary is 2-celIed and each has
branchlets light-brown, pubescent and glabres- only one ovule. Based on the description, the
12 Flora Malesiana [ser. I, vol. 6^

common name 'Malowassi' and the usage, Heyne 8-9 by 61/4-7 mm, dark-brown, smooth, shining,
(Nutt. PI. 1927, p. 1151) identified it as a Gyrinopsis with an obscure caruncle-like appendage at the
sp. (= Aquilaria). base.
Merrill (Int. Rumph. 1917, 380) in inter- Distr. Malaysia: Philippines (Luzon: B.S.
preting RuMPHius' Cortex filarius, with the re- 26876, 41562, 76441— type (A), Wenzel 1201).
presenting specimen (Robinson's PI. Rumph. Amb. Ecol. On forested slopes, 1000 m.
n. 274), referred it to Gyrinopsis brachyantha Note. Very closely related to A. filaria, but
Merr. (= Aquilaria brachyantha). The leaves of easily separated from the latter by the persistent
these two species are very similar to each other. floral tube with erect calyx lobes and the smaller
See also Barker (F1. Mai. I, 5, 1957, 359-360). leaves with distinct lateral nerves.
J. Smith initiated the error to combine the plate
of Cortex foetidus Rumph. (t. 7) with the de- 9. AquCaria hirta Ridl. J. Str. Br. R. As. Soc. n. 35
scription of Cortex filarius Rumph. referring them (1901) 78; Gamble, J. As. Soc. Beng. 75, ii (1912)
both to Pittosporum ferrugineum (in Rees, Cycl. 265; Ridl. F1. Mai. Pen. 3 (1924) 148; Domke,
27 art. Pittosporum, 1814), although he remarked Bibl. Bot. Ill (1934) t. 5 f. Aig.—A. moszkowskii
already that the "thready bark" ascribed to it by GiLG, Notizbl. Berl.-Dahl. 5 (1908) 84; Quis. J.
Rumphius does not occur in Pittosporum. This Arn. Arb. 27 (1946) 403.— Fig. le-f.
error was continued by de Candolle, I.e. Tree up to 14 m
with whitish and rather smooth
In reviewing Oken's work in 1950, Merrill bark. Young branchlets light brown, covered with
(J. Arn. Arb. 31, 1950, 283) pointed out that the silky hairs, glabrescent. Leaves subcoriaceous,
description of Pittosporum filarium Oken was dull and pubescent beneath especially on the
wholly taken from Rumphius' Cortex filarius and midrib, nerves and veins, sometimes glabrescent,
concluded that Gyrinopsis brachyantha Merr. shining on the upper surface, elliptic-oblong,
should be added to the synonymy of Aquilaria ovate-oblong, 6I/2-I4 by IVi-^Vi cm; base
(Oken) Merr.
filaria cuneate to obtuse or rounded; apex acuminate,
On
examining Robinson's specimen indicated the acumen up to 1 Vi cm, mucronate, pointed at
above and another specimen collected by Teys- the tip; nerves 16-30 pairs, irregular, sometimes
mann {s.n.. Bo) from Soja, Ambon, however, it branched, elevated beneath, visible to obsolete
appears that A. filaria is distinct from the Phi- above, slightly curved upward and towards the
lippine A. brachyantha. A. filaria is characterized margin; veins distinct or visible beneath, obscure
by calyx lobes about half the length of the tube, or not visible above; petiole 5-7 mm, thickened,
stamens sessile or very short-filamentous, ovary curved, pubescent. Inflorescences sessile or up to
densely villous, and the style obscure, while A. 10 mm peduncled, pubescent, 5-14-flowered;
brachyantha is characterized by calyx lobes about bracts small. Flowers white {fide Kep. 71521) or
as long as the tube, stamens distinctly filamentous light yellow {fide Bunnemeijer 7575), up to 2 cm
and the filaments as long as or longer than the pedicelled, pubescent. Floral tube cylindric 6-8
anthers, ovary sparsely hairy, and the style distinct mm long, densely pubescent outside and towards
and as long as the ovary. the base inside, ribbed and sparsely villose within
at the upper part. Calyx lobes ovate and obtuse,
8. Aquilaria parvifolia (Quis.) nov. comb.— 2-3 mm
long, densely pubescent outside and
Gyrinopsis parvifolia Quis. J. Arn. Arb. 27 (1946) densely puberulous inside. Petaloid appendages
405. inserted slightly behind the stamens, ovate or
Shrub c. 1 mtall, branches light brown or semi-orbicular, densely villous, almost as long as
reddish-brown. Leaves subcoriaceous, slightly the stamens, sometimes even slightly longer.
pubescent beneath, glabrous above, elliptic- Stamens sessile, c. 1 mm
long, oblong, connective
oblong, ovate-oblong, or lanceolate, 41/2-15 by dark-brown. Pistil clavate, 5 mm long; ovary
1-41/2 cm; base acute to cuneate; apex narrowly densely puberulous; style absent; stigma capitate.
acute to acuminate; nerves 7-12 pairs, slightly Fruits protruding from the floral tube, oblan-
curving upward, distinctly elevated beneath, slightly ceolate, abruptly acute at the apex, attenuate to
elevated above; veins distinct beneath, invisible the base, including the stipe 3 1/2-5 by 1 cm, densely
above; petiole c. 5 mm, sparsely pubescent, gla- golden puberulous; pericarp coriaceous. Seeds
brescent. Infructescences axillary, terminal, some- ovoid, 10 by 6 mm, puberulous, glabrescent,
times extra-axillary, umbelliform, short-peduncied, shortly beaked at the apex, cuneate at the base,
sometimes almost sessile, each with 1 to 4 fruits; black and shining, with a long glabrous appendage
pedicel c. 3 mm, puberulous. Persistent flower c. 10 mm long.
short-tubular, 5-6 mm long. Floral tube sparsely Distr. Malaysia: Malay Peninsula (Trengganu,
puberulous on both surfaces. Calyx lobes ovate or Pahang, Johore, and Singapore; lectotype:
ovate-oblong, 1/2-2
1 mm long, densely puberulous MuRTON 2, Sing; paratypes: Ridley 3837, Sing
inside and on the margins and tips outside. Petaloid and Ridley 11020, K, Sing), E. Sumatra (Sena-
appendages orbicular or deltoid, c. 1/3- 1/2 the length maninik), Riouw, and Lingga.
of the stamens, villous, the hairs slightly longer Ecol. Hill slopes, from the lowland up to
than the appendage. Stamens sessile, c. 1 mm long. 300 m.
Fruits sUghtly obovoid or globose, 1-1 1/2 by 1-1 Va Vern. Chamdan, changang, kayu chandan,
cm, yellowish, rugose when dry, constricted at the sahare, M; Sumatra: karas.
base into c. 2 mm long stipe. Seeds broadly ovoid. Note. A. moszkowskii Gilg was described on a
 Dec. 1960] Thymelaeaceae (Ding Hou) 13

specimen collected by Moszkowski (12, B)

sterile nerves (10-) 15-25 pairs, curving and ascending
at Senamaninik, eastern Sumatra. I have not seen towards the margin, elevated and prominent
the type, but the locality and Gilo's detailed des- beneath, distinct above; veins loosely reticulate;
cription agree very well with the present species, petiole 5-7 mm. Inflorescences axillary or extra-
especially the silky hairs occurring on the under- axillary, branched and up to 1
1/2 cm peduncled,
side of the leaf which is peculiar to this species. short-paniculiform, pubescent; pedicels 3-7 mm,
pubescent. Flowers 7-12 mm long, yellowish,
10. Aquilaria rostrata Ridl. FI. Mai. Pen. 3 (1924) greenish or yellowish-white. Floral tube cylindric,
148. 10-costate, sparsely hairy outside. Calyx lobes
Tree. Branchlets pubescent, glabrescent. Leaves slightly ovate, puberulous inside, 2-3 mm
long,
subcoriaceous, glabrous, rather shining on both densely puberulous on both surfaces, sometimes
surfaces, lanceolate, rarely ovate-oblong, 6'/2-10 glabrescent on the outside. Petaloid appendages
by 2 1/2-4 '/2 cm; base obtuse, cuneate to attenuate; oblong, c. 1 mm long, densely short-hairy. Stamens
apex acuminate, the acumen up to 1 '/2 cm; nerves usually sessile, rarely with very short filaments,
16- many pairs, simple or rarely branched, spread- almost as long as the petaloid appendages. Disk
ing or slightly curved and ascending, elevated ring-like to cupular, densely puberulous. Pistil c.
beneath and visible above; veins visible beneath 5 mm long, with a distinct stipe c. 2 mmlong, the
and obscure above; petiole 31/2-7 mm. Pedicels c. stipe accrescent and elongated. Ovary ellipsoid,
3 mm, brownish hairy. Floral tube cylindric, 6 mm attenuate to the base, gradually narrowed at the
long, splitting on one side, glabrous outside, apex; stigma capitate. Fruit protruding from the
sparsely puberulous inside. Calyx lobes slightly top of the floral tube, ellipsoid or obovoid, 2-31/2
oblong, c. 1 1/2 rnm long, puberulous on both sur- by ly^ cm, slightly puberulous and glabrescent,
faces. Petaloid appendages unknown. Stamens narrowed to the base into an elongate stipe up to
sessile. Fruits (young) obovate-oblong or oblan- 1/2 cm, acuminate to the apex, usually slightly
1

ceolate, including the stipe 3 by %-l 1/2 cm, contracted in the middle; floral tube entire, very
brownish hairy outside, long-narrowed towards rarely splitting on one side (Kadir A
3601). Seeds
the base, apex beaked. Seeds slightly ellipsoid- black, ovoid, 10 by 5 mm, sparsely puberulous,
oblong, 10 by 4 mm
{excl. the appendage), acuminate to the apex, with an elongate tail c.
brownish, puberulous, acuminate, base attenuate 5mm long, attached at the center of the appendage,
and elongate into a slender appendage, glabrous. the appendage slender, c. 1 cm long, densely
Distr. Malaysia: Malay Peninsula (Pahang, reddish-brown pubescent.
Wray's Camp, Gunong Tahan, Ridley 16264, Distr. Malaysia: Sumatra (Palembang), Malay
type, K, Sing). Peninsula (Johore), and common in Borneo.
Note. As mentioned by Ridley the specimens Ecol. Primary forests, rarely in swampy forest
are poor. No material has been collected since the (Johore: S.F. 29008, K), from the lowland up to
type. I have seen two sheets of the type number 825 m.
and one other sterile unnumbered sheet. Only Vern. Merkaras puti. Sum., galiaru, gumbil,
young fruits are available, with the persistent floral njabak, M, tanduk = garu. Born.
tube. Unfortunately, the petaloid appendages and Notes. This species is characterized by the
stamens of them were eaten by insects except the cylindric floral tube, the oblong and puberulous
basal parts of two sessile stamens in one flower. petaloid appendages which are almost as long as the
From the available material, it is impossible to sessile or subsessile stamens, and the stiped pistil
verify the number and shape of the petaloid with a short, puberulous, ring-like disk at its base.
appendages and the number of stamens. The type specimen of the present species was
This species, as pointed out by Ridley, is collected by Beccari (PB 2339, Fi) from Sarawak.
characterized by the long-beaked fruits. In ad- It has rather small leaves (8I/2-I3I/2 by 1/2-4 cm)

dition, the floral tube is longer than the lobes, and young flowers. The type of A. grandifolia
and the stamens are sessile. (Grashoff 693, Bo) collected in the swamp forest,
Palembang, S. Sumatra, has larger leaves (17-27
11. Aquilaria beccariana van Tiegh. Ann. Nat. Sc. by 6-8 1/2 cm) and young flowers. Many specimens
Hot. VII, 17 (1893) 217; Bull. Soc. Bot. Fr. 40 collected in theMalay Peninsula {e.g. S.F. 29008,
(1893) 77; Gilg, Bot. Jahrb. 28 (1900) 145; Boerl. 29195, 29381, 29470) and Borneo {e.g. bb 34916,
Handl. 3 (1900) 112; Becc. Nelle Foreste (1902) Endert 3319, 4035, C.F. 34453, Purseglove
592; Merr. En. Born. (1921) 4X6.— A. grandifolia P4752, Rutten 68, Patrick Ping San A 1726,
Domke, Notizbl. Berl.-Dahl. 11 (1932) 348.— and Wood San 15218) have flowers and fruits in
A. cumingiana var. parviflora Airy Shaw, Kew different stages of development and their leaves
Bull. (1940) 261. Gyrinopsis grandifolia Quis. J. show a variable size. From this additional material
Arn. Arb. 27 (1946) 406.— Fig. la-c. we can clearly infer that only one species is
Tree up to 20 m
tall and 36 cm diam. with grey represented.
and smooth bark. Young branchlets pubescent. Aquilaria cumingiana var. parviflora was based
Leaves papery to subcoriaceous, glabrous on both on Haviland 3092 (type) and several other
surfaces, sometimes scattered pubescent beneath, collections from western Borneo. All the specimens
oblong, oblong-lanceolate, or elliptic-oblong, cited in the original description agree with the
rarely elliptic, (7-)ll-27 by (3-)6-8i/2 cm; base present species and are quite different from A.
cuneate to attenuate; apex acute to acuminate; cumingiana.
14 Flora Malesiana [ser. 1, vol. 6^

Fig. 3. Phaleria capitata Jack. X 2/3, Z>. opened flower, x ''/g, c. ovary with disk, x 7, (/. stamens,
a. Habit,
X 7, e. fruit, nat. size,/, longitudinal section of fruit, one seed removed to show meshes of
seed, x 2, g.
endocarp, x 2. P. elegans L. M. Perry, h. Opened flower, x '/g, /. punctate bract, X 2/3. P. macro-
carpa (ScHEFF.) Boerl. j-k. Opened flowers, dimorphous, x ^/g, /. longitudinal section of fruit, one seed
removed, x 2/3. P. octandra (L.) Baill. m. Opened flower, X ^/g, n. fruit, nat. size, o. longitudinal
section of fruit, one seed removed, x 2 (a Bakhuizen van den Brink 2294, b-d C.H.B. XI-B-III-8,
e-g C.H.B. XI-B-XVII^3, h Brass 24484, i Brass 24483,7 Atasrip 139, k-l C.H.B. VIII-G-93, m-o
Walsh 36).
Dec. 1960] Thymelaeaceae (Ding Hou) 15

12. Aquilaria cumingiana (Decne) Ridl. J. Str. Br. short-stiped, densely hairy. Ovary obovate, at-
R. As. Soc. n. 35 (1901) 80; Hall./. Med. Rijks- tenuate to the base; style continuous with the
herb. n. 44 (1922) 17. Gyrinopsis cumingiana ovary, obscure or distinct; stigma capitate. Fruits
Decne, Ann. Sc. Nat. Bot. II, 19 (1843) 41, t. 1 globose, slightly obovoid, or ellipsoid, rugose,
f. B, 13-21; Meisn. in DC. Prod. 14 (1857) 603; protruding laterally from the split floral tube,
MiQ. Fl. Ind. Bat. 1, 1 (1858) 883; F.-Vill. Nov. 1 %
by 1 '/'3 cm. Seeds broad-ovoid, plano-convex,
App. (1880) 183; Vidal, Phan. Cuming. (1885) 1 by Y^ cm with a short caruncle-like appendage.
140; Rev. PI. Vase. Filip. (1886) 230; Merr. Bull. Distr. Malaysia: S. Borneo (Sampit region),
Bur. For. Philip. (1903) 41; Elm. Leafl. Philip.
1 Philippines (common), and Moluccas (Morotai
Bot. 5 (1913) 1629; En. Philip. 3 (1923) 131; and Halmaheira).
Holthuis & Lam. Blumea 5 (1942) 216; Quis. J. Ecol. In primary forests at low and medium
Arn. Arb. 27 (1946) 405; Med. PI. Philip. (1951) altitudes.
636. Decaisnella cumingiana O.K. Rev. Gen. PI. Uses. According to Quisumbing, I.e., the bark
2 (1891) 584. Gvrinopsis cumingiana var. pubes- and roots are used to stop bleeding from wounds.
cens Elm. Leafl. Philip. Bot. 5 (1913) 1629; Merr. The bark, wood, and fruit are used as a substitute
En. Philip. 3 (1923) 131. Gyrinopsis decemcostata for quinine.
Hall./. Med. Rijksherb. n. 44 (1922) 17; Domke, Vern. Ala/ian, maga-an, palisan. Tag., bago,
Bibl. Bot. 1 (1934) t. 2 f. 9; t. 4 f. 26^.— Gyrinop-
1 1 Mbo., binukat, Ak. Bis., butlo, Neg., dalakit, S.L.
sis pubifolia Quis. J. Arn. Arb. 27 (1946) 406.— Bis., magwalem, Sub., pamaluian, Bag.; giba
Fig. li. kolano, Halmaheira.
Shrub or small tree up to 5 m. Bark ashy grey, Note. The type specimen of Gyrinopsis pubifolia
mottled and smooth. Young branchlets densely Quis. is B.S. 75314 (A). According to Quisumbing
pubescent, glabrescent. Leaves chartaceous to (/.c) it was collected at Mt Abucay, Catanduanes,
subcoriaceous, glabrous on both surfaces or some- at c. 1600 m, September 11, 1928 (the field data
times pubescent on the lower surface; oblong- on the label of this specimen are simply indicated
lanceolate, elliptic-oblong, or ovate-oblong, rarely as "Catanduanes, M. Ramos & G. Edano, July-
obovate-oblong, 14-18 by 5'/2-8'/2 cm; base Sept. 1928"'); it has rather young flowers with the

cuneate, rarely rounded; ape.x acute to acuminate, lower part of the anthers united with the floral
acumen up to 1 1/2 cm; nerves 12-18 pairs, slightly tube. There is another specimen (B.S. 75516,
curved and ascending to the margin, elevated and Sing) which is similar to the above one and bears
distinct beneath, slightly elevated above; veins and the same field data; it has both young and mature
veinlets numerous, irregularly forked; petiole 4-6 flowers and has been distributed as Gyrinopsis
mm. Inflorescences simple or sometimes branched, cumingiana Decne {= A. cumingiana) with which
few- to many-flowered; peduncle short, c. 5 mm I agree. In comparing these two specimens, it
long, rarely subsessile; pedicels 3 mm, setose. appears that Gyrinopsis pubifolia is conspecific
Flowers whitish, 13-16 mm
long. Floral tube with A. cumingiana.
cylindric, puberulous outside, glabrescent, densely
or sparsely pubescent inside, the retrorse hairs Excluded
appressed, sometimes distinctly 10-costate inside, Aquilaria? bancana MiQ. Sum. (1861) 355 is ac-
usually with irregular, sulphureous, wart-like cording to Airy Shaw (FI. Mai. I, 4, 1953, 361)
excretions. Calyx lobes ovate or oblong, obtuse, = Gonystylus bancanus (MlQ.) KuRZ (Thyme-
2-3 mmlong, densely puberulous on both surfaces, laeac).
sometimes glabrescent. Petaloid appendages short, Aquilaria? macrophylla MlQ. Sum. (1861) 356
usually united in a ring, rarely free or united at is according to Airy Shaw (Fl. Mai. I, 4, 1953,
the base, about half as long as the stamens, den- 354) = Gonystylus macrophylla (MiQ.) Airy Shaw
sely hairy, the hairs longer than or as long as the (Thymelaeac).
appendages. Stamens sessile, 1-2 mm
long, free Aquilaria pentandra Blanco, Fl. Filip. ed. 1
from the tube, slightly below the appendages or at (1837) is according to Barker (Fl. Mai. I, 5,
the same level with them, the lower V3-V2 of the 1957, 355) = Pittosporum pentandrum (Blco)
anther usually adnate to the tube. Pistil c. 7 long, mm Merr. (Pittosporac).

2. PHALERIA
Jack, Mai. Misc. 2 (1 822) 59 reimpr.
; Hook. Comp. Bot. Mag. ( 1835) 56 Domke,
1 1 ;

Bibl. Bot. Ill (1934) 123, t.4 f.36h, map 6; Merr. J. Arn. Arb. 33 (1952) 239.—
Drimyspermum Reinw. Syll. PI. Ratisb. 2 (1825) 15; Rchb. Norn. Bot. Hort.
2 (1841) 65, as Drymispermum.—Pseudais Decne, Ann. Sc. Nat. Bot. II, 19 (1843)
40.—Leucosmia Benth. in Hook. Lond. J. Bot. 2 (1843) 23\.— Dais {non Linne)
aiict. —Fig. 3-9.
Shrubs or trees. Leaves decussate or opposite. Inflorescences terminal or
axillary, sometimes cauliflorous, capitate, fascicled or umbelliform, peduncled.
16 Flora Malesiana [ser. I, vol. 6^

rarely sessile, peduncles usually with decussate, persistent, reddish-brown, glabrous

bracteoles towards the base and gradually increasing in size and more spaced
towards the upper parts, sometimes 4 or more involucral bracts at the uppermost
part of the peduncle surrounding the flowers. Flowers monomorphous rarely
heteromorphous, white, sessile, articulated at the base. Floral tube infundibuli-
form or cyhndric, glabrous or puberulous on both surfaces. Calyx lobes 5, rarely
4 or 6, slightly unequal. Petaloid appendages obscure and rim-like, or none, rarely
distinct {P. pentecostalis Leandri, an extra-Mai. sp.). Stamens in two series,
usually filamentous and exserted, sometimes included, rarely sessile; anthers
oblong, dorsifixed. Disk cupular, submembranous. Ovary ovoid or ellipsoid,
glabrous or hairy at the apex, 2-celled or rarely 1 -celled by abortion, once found
3-celled in P. octandra; style terminal, filiform, sometimes exserted; stigma
capitate, papillose. Fruits drupaceous, 2- or 1 -seeded, exocarp and mesocarp
fibrous and fleshy (sometimes hard in the herbarium), endocarp coriaceous and
hard. Seeds exalbuminous cotyledons thick and hemispherical.
;

Distr. About 20 spp., distributed in Ceylon (P. capitata), SB. Asia, through Malaysia to Australia,
Micronesia {P. nisidai), and the Pacific (as far as Samoa and Tonga).
Ecol. In rain-forests, rarely in seasonal forests, from the lov/land up to 1400 m.
Note. The generic name Drimyspennum has in literature frequently been mis-spelled as Drymispermum.
No attempt has been made to indicate this erroneous etymology, except where new species or com-
binations have been proposed.

KEY TO THE SPECIES

I . Inflorescences terminal and/or in the leaf axils of the terminal node, sometimes also occurring in
the upper two nodes, rarely in several nodes in P. octandra, sometimes cauliflorous in P. capitata and
P. coccinea. Only one peduncle in each axil, bearing (6-)8-many flowers. Flowers homomorphic.
Fruits usually small, less than 3I/2 by 2 cm; pericarp thin, less than V2 cm thick.
2. Floral tube pubescent outside, very rarely glabrescent. Fruits ellipsoid and apiculate at both ends,
often spindle-shaped.
3. Involucral bracts 8 or more, large, 2'/2-3'/2 by 1-2 cm. Floral tube wide, 12-15 mm diam. at the
throat 1. P. elegans
3. Involucral bracts usually 4 or 5, smaller, 34-IV2 by I/3-I cm. Floral tube narrow, 2-6 mm diam.
at the throat.
4. Inflorescences 8-10(-15)-flowered, very rarely many-flowered. Flowers lV2-2(-2i4) cm long;
calyx lobes Vi{-Vi) the length of the tube. Ovary glabrous. Leaves elliptic-oblong, elliptic-
lanceolate, or obovate, (4-)13-26 by (li/4-)3-8 cm; nerves 9-11 pairs . 2. P. octandra
4. Inflorescences 20-many-flowered. Flowers 3-4 V^ cm long; calyx lobes V^-Va the length of the tube.
Ovary usually hairy at the top, rarely glabrescent. Leaves oblong-lanceolate, oblanceolate or rarely
ovate-oblong, 111/2-33 by 3 14- 14 cm; nerves (8-) 13-22 pairs . . 3. P. perrottetiana
2. Floral tube glabrous outside. Fruits subglobose, ovoid, or ellipsoid, usually rounded or obtuse at
both ends, sometimes apiculate at the apex (acute or acuminate towards both ends mP. sogerensis).
5. Stamens and style always included. Stamens sessile or short-filamentous. Style usually not longer
than the tube 4. P. nisidai
5. Stamens and style exserted. Stamens long filamentous.
6. Flowers IVi-^Vi cm long. Calyx lobes %-V^ the length of the tube. Ovary glabrous or hairy at
the top.
7. Inflorescences usually 8-flowered, sometimes cauliflorous and many-flowered. Ovary glabrous.
Fruits subglobose, 1-1 '/a cm in diam.; endocarp perforated (fibrous strands interlaced, leaving
distinct meshes) 5. P. capitata
7. Inflorescences 20-many-flowered, rarely cauliflorous. Ovary usually hairy or puberulous at the
top. Fruits ellipsoid, 1 '/2-2 by 1 cm, usually blunt at both ends, sometimes apiculate at the apex;
endocarp not perforated (fibres uniformly arranged, not leaving open spaces) 6. P. coccinea
6. Flowers 1 V2-2 cm long. Calyx lobes usually Vii-Vi) the length of the tube. Ovary hairy at the top.
Fruits acute or acuminate towards both ends.
8. Inflorescences with usually more than 20 flowers. Leaves 12-33 by 4-14 cm. Stamens c. 10 mm
exserted beyond the tube. Calyx lobes c. 3 times as long as broad . 3. P. perrottetiana
8. Inflorescences with 6-10 flowers. Leaves 7-16 by lYi-l cm. Stamens c. 5-6 mm exserted. Calyx
lobes c. 2-21/2 times as long as wide 7. P. sogerensis
Dec. 1960] Thymelaeaceae (Ding Hou) 17

Inflorescences axillary and occurring in the leaf axils of several nodes along the branches or branchlets,
sometimes cauliflorous; peduncles 1-3 or sometimes several or many in each axil, each peduncle
bearing 2-5(-8 flowers. Flowers usually heteromorphic (sessile stamens and an exserted style, or exserted
)

stamens and a short style, and their intermediate forms). Fruits large, 3-5 '/2 by 3-4 '/2 cm; pericarp
thick, 1-1 1/2 cm 8. P. macrocarpa

1. Phaleria elegans L. M. Perry, J. Arn. Arb. 39 1, 1 (1858) 885; Blume & de Vriese, Ann. Hort.
(1958) 422, f. c & d.— Fig. 3h-i. Bot. (Fl. Jard.) 2 (1859) 33, with p].—Drimysper-
Shrub or small tree, 2-3 m, sparsely branched. iniim bitrnianni Decne, Ann. Sc. Nat. Bot. II, 19
Branchlets reddish-brown, terete, glabrous, hollow. (1843) 40, as Drymispermiim; Bleeker, Nat.
Leaves chartaceous to subcoriaceous, glabrous, Geneesk. Arch. N.I. 2 (1845) 75; Meisn. in DC.
oblanceolate, rarely elliptic-lanceolate, 18-30 by Prod. 14 (1857) 605. Drimyspermum bhimei
51/2-9 cm; base attenuate; apex acute to short- (non Decne) Hassk. Nat. Tijd. N.I. 10 (1856) 885.
acuminate; margins recurved; nerves 8-12 pairs, — Drimyspermum ambiguum Meisn. in DC. Prod.
elevated on both surfaces, spreading, curved to- 14 (1857) 605, as Drymispermum. —
Drimysper-
wards the margin; veins slightly elevated on both mum longifolium Miq. Fl. Ind. Bat. 1, 1 (1858)
surfaces, rather widely reticulate; petiole stout, 885, as Drvmispermum. — P. laurifolia Hook. /.
about as thick as the branchlet, red when fresh Bot. Mag. (1869) t. 5787; Val. Ic. Bog. 4 (1913)
(fide Brass), reddish-brown to black when dry. 21 1, t. 368, f. 1-3.— P. ambigita Hook./. Bot. Mag.
Inflorescences terminal and/or in the axils of the (1896) t. 1A1\.—P. longifolia Boerl. Handl. 3
terminal node, subsessile to short-peduncled (1900) 111. P. laurifolia var. javanica Val. Ic.
(peduncle 5-9 mmin fruit), with small, decussate Bog. 4 (1913) 212, t. 368, f. 4-9; in K. &
V. Bijdr.
bracts towards the base. Involucral bracts cream- 13 (1914) 46; Hall. /. Med. Rijksherb. n. 44
coloured (fleshy in bud), 8 or more, arranged in (1922) 23. P. octandra var. laurifolia Ware, ex
whorls, those of the inner whorl longer, oblong or VON Malm in Fedde, Rep. 34 (1934) 282.— P.
ovate, 2 1/2-3 '/2 by 1-2 cm, obtuse or acute, parvifolia Back. Blumea 5 (1945) 494.— Fig.
slightly puberulous towards the upper part inside, 3m-o.
especially near the margins, glabrescent, some- Shrub up to 5 m by 5 cm. Leaves chartaceous to
times pellucid-dotted, 6-20-flowered. Flowers in- subcoriaceous, elliptic-oblong, narrowly elliptic
fundibular, 3%-41/i cm long. Floral tube 1 '4-1 1/2 or obovate, (4-)13-26 by (li/2-)3-8 cm; base
cm diam. at the throat, densely pubescent outside attenuate, rarely short-acute; apex acuminate;
and towards the base inside. Calyx lobes sparsely nerves 9-11 pairs, slightly ascending and curved;
pubescent outside and densely pubescent inside, veins rather widely reticulate, slightly elevated
± oblong, c. 9 by 5 cm, obtuse. Petaloid ap- below, plane and visible above; petiole 6 mm.
pendages rim-like. Stamens included, filamentous, Inflorescences usually terminal and/or in the axils
6-10 mm, free from the tube at the throat or of the terminal node, sometimes in the axils along
slightly below it; anthers oblong, about 1/2 mm the branchlets (Beumee 2405, Bo), 8-10(-15--v))-
1

long. Disk cup-shaped, crenulate, c. 2 mm long. flowered; peduncles very short to up to 1 1/4 cm,
Pistil included. Ovary glabrous, ellipsoid, c. 3 mm usually with decussate, small, lanceolate scales at
long, gradually narrowed towards the apex; style the base gradually increasing in size apically. In-
filiform, c. 3 cm; stigma globose. Fruit ellipsoid to volucral bracts 4, rarely 5, ovate or obovate,
fusiform, 2%-3i/3 by 1 1/2-1% cm, slightly com- 8-12 by 4-11 mm, puberulous on the upper part
pressed, acuminate on both ends. Seeds broadly of both surfaces, persistent, rarely caducous after
ovate or obovate, or semiglobose, plano-convex, anthesis. Flowers (1-)1 1/2-214 cm long. Floral
10 by 7-10 mm, sharply pointed at the apex. tube cylindric, slightly swollen at the base, usually
Distr. Malaysia: New Guinea (Goodenough pubescent on both surfaces, sometimes glabres-
I.). cent. Petaloid appendages sometimes rim-like.
Ecol. Undergrowth of an oak forest and oc- Calyx lobes 4 or 5, oblong or slightly elliptic or
casional in gullies in the forest, 1600-1750 m. obovate, 4-7 by 2-3 mm, densely puberulous out-
Note. This species is characterized by 8 or side and at the upper part inside. Stamens and
more, large involucral bracts which are caducous pistil long-exserted (up to 8 mm) in anthesis.
after anthesis, a wide floral tube which is densely Disk cup-shaped, membranous. Pistil 2-21/2 cm
pubescent outside, and the included stamens and long. Ovary glabrous, ellipsoid, narrowed into the
style. filiform style; stigma capitate. Fruits ellipsoid,
ovoid, sometimes slightly compressed, 11-16 by
2. Phaleria octandra (L.) Baill. Adans. 11 (1875) 9-15 mm, acute or attenuate and pointed towards
321; Merr. Philip. J. Sc. 19 (1921) 361.— Dais both ends, usually 2-celled, 2- or 1 -seeded, once
octandra Linne, Mant. PI. 1 (1767) 69; Burm. /. found 3-seeded (Valeton 123, Bo).
Fl. Ind. (1768) 104, t. 32, f.2.— Dais dubiosa Distr. Australia: North Queensland (Michael
(non Bl.) Decne, Herb. Timor. (1834) 41; 1250, Bo), and Malaysia: throughout Java,
Spanoghe, Linnaea 15 (1841) 335. Drimysper- Madura, Bawean, and Lesser Sunda Islands (Bali
mitm laiirifolium Decne, Ann. Sc. Nat. Bot. 11, to Timor and Tanimbar), Moluccas (Halmaheira),
19 (1843) 39, t. 1, f. 1-12, as Drymispermiim; and South New Guinea (Daru I.). Fig. 4.
Bleeker, Nat. Geneesk. Arch. N.I. 2 (1845) 75; Ecol. In beach-forest, common on sandy soil
ZOLL. Syst. Verz. 2 (1854) 117; Miq. Fl. Ind. Bat. of teak forest in E. Java, rarely in primary and
 li Flora Malesiana [ser. I, vol, 6^

secondary mixed forests, from the lowland up to Valeton (in K. &

V. Bijdr. 13, 1914, 40), one can
600 m, in Jamdena up to 800 m, in Timor up to occasionally find glabrous flowers among the
1000 m. hairy ones (cf. Koorders 30200 /3, 30145 ^, and
Buwalda 7291).
In the Kew Herb, there are two specimens of
cultivated origin identified as Phaleria laurifolia.
One of them has flowers with a floral tube glabrous
outside which has been used for the plate in
CuRTis's Bot. Mag. t. 5787; the other has flowers
with a floral tube pubescent outside. In other res-
pects their characters entirely agree. It is not clear
whether they were collected from the same or
from diff"erent plants.
According to Valeton (Ic. Bog. 4, 1913, 47)
P. octandra is slightly more xerophytic than the
others and obviously cannot well maintain itself in
rain-forest and the specimens which had been
introduced in the Bogor Botanic Gardens have
perished. The fibers which constitute the endo-
Fig. 4. Localities of Phaleria octandra (L.) Baii l. and mesocarps form a rather thick layer, with a
(+) and P. perrottetiana (Decne) F.-Vill. (•). smooth, compact, shining inner surface, more
loose than the peripheral parts but leaving no
Vern. Kopinan, mritja sunda, pantjal pamor, J, meshes among them.
kaju pateng, Kangean, mandalika, maupulang, In 1893 H. Hallier (320, Bo) once collected it
Bawean; Lesser Sunda Islands: daun wempe, nu along the border of the Palace Garden (Herten-
impi, Jamdena, koffifui, Timor, lolong, Sumba; kamp) adjoining the Botanic Gardens at Bogor;
pepigeow, Halmaheira, Sawai lang. it must have escaped or been derived from
Note. Dais octandra was first published by cultivation {cf. Valeton in K. &. V. Bijdr. 13,
Linnaeus in 1767 (Mant. PI. 1, p. 69). There is a 1914, 47). One specimen was collected by
specimen in the Linnean Herbarium which bears Hoogerwerf (42, Bo) in the beach forest of
LiNNAEUs's handwriting and agrees with the Udjungkulon, W. Java, which is apparently the
original description with the exception that some western limit of this species.
of the flowers are sparsely pubescent outside. P. octandra is closely related to P. perrottetiana.
In 1768, one year later. Dais octandra appeared Some specimens collected at Sumbawa (Elbert
in Burman's F1. Ind. p. 104, t. 32 f. 2. There are 3903, 3949, 3994, 4099, 4127) have 1 5-cNi-flowered
two specimens of Burman's in the Herb. Delessert inflorescences. One specimen collected in the
at Geneva; one of them bears in Burman's Tanimber Is. (S. Moluccas) (Buwalda 4369), has
handwriting "Dais octandra'' and the other has a large leaves (20 by 9 cm) which are similar to
label "Java Kleinhoff". These two specimens are those of P. perrottetiana both in size and shape.
similar to each other and may belong to one
collection. They are also similar to the specimen 3. Phaleria perrottetiana (Decne) F.-Vill. Nov.
in the Linnean Herbarium mentioned above but App. (1880) 183; Merr. Sp. Blanc. (1918) 378;
the flowers appear more pubescent on the outside Brown, Minor Prod. Philip. For. 1 (1920) 403;
of the floral tube. I assume Burman has sent his Merr. En. Phihp. 3 (1923) 131; Philip. J. Sc. 29
drawing with one of his specimens - which might (1926) 404; Merr. & Perry, J. Arn. Arb. 22
have been the one which he had used for the des- (1941)265. Drimyspermum perrottetianumDECNE,
cription - to Linnaeus, as Linnaeus cited "Burm. Ann. Sc. Nat. Bot. II, 19 (1843) 40, as Drymisper-
Ind. t. 33, f. 2" in the description. mum; Bleeker, Nat. Geneesk. Arch. N.I. 2 (1845)
In 1876 Baillon (Adansonia 11, p. 321) rightly 75; Meisn. in DC. Prod. 14 (1857) 605; MiQ. Fl.
transferred Dais octandra to Phaleria as P. Ind. Bat. 1, (1857) m6.—Dais laurifolia {non
1

octandra. He mentioned only Burman's publica- Jacq.) Blanco, Fl. Filip. (1837) 375, ed. 2 (1845)
tion and overlooked that of Linnaeus. According 263, ed. 3, 2 (1878) 125. Drimyspermum urens
to priority we should accept Linnaeus as the {non Reinw.) Scheff. Ann. Jard. Bot. Btzg 1
author of Dais octandra and the specimen in the (1876) 46. Drimyspermum coccineum {non Dais
Linnean Herbarium as the holotype. coccinia Gaudich.) Becc. in d'Albertis, New
Because of some minor discrepancies between Guinea 2 (1880) 398, as Drymispermum, quoad
Burman's {I.e.) description and drawing, and both specim.— P. blumei {non Benth.) Hemsl. Bot.
of them also not exactly agreeing with the spec- Chall. 3 (1885) 244.— P. splendida Val. Ic. Bog. 4
imens preserved in Burman's herbarium, Decais- (1913) 219, t. 370 A-B.
NE (Ann. Sc. Nat. Bot. II, 19, 1843, 40) based a Shrub, sometimes a tree, up to 8 m. Branches
new species, Drimyspermum burmanni, on the and branchlets glabrous and dark-brown. Leaves
specimens in the Burman Herbarium. However, chartaceous, oblong-lanceolate, oblanceolate, or
the discrepancies have no value for defining ovate-oblong, 111/2-33 by 31/2-I4 cm; base
species: the floral tubes are usually pubescent cuneate, rounded; apex acuminate; margins
outside but rarely glabrous and, as pointed out by slightly recurved; nerves (8-) 13-22 pairs, distinct
 Dec. 1960] Thymelaeaceae (Ding Hou) 19

and elevated on both surfaces, curving and as-

cending towards the margin, veins and veinlets
slightly reticulate; petiole 1 Vi cm, slightly winged.
Inflorescences terminal and/or axillary at the
terminal node, sometimes in the axils of the
upper two nodes of the branch, solitary, very
rarely more than one in the same axil. Peduncle
up to 31/2 cm; bracts small, lanceolate, 3 mm
long, densely decussate at the basal part, per-
sistent. Involucral bracts 4, caducous after an-
thesis, rarely persistent, oblong, obovate-oblong,
15 by 8 mm, obtuse at the apex, apiculate or
obtuse, densely puberulous towards the upper
part on both surfaces. Flowers (2-)3^i/2 cm
long. Floral tube pubescent outside, villose at
the lower half or lower 2/3 inside. Calyx lobes
5-9 mm long. Stamens and style c. 10 mm
exserted beyond the tube. Disk cup-shaped, some-
times consisting of 6 or 7 free lobes (PNH 33763).
Ovary usually hairy at the apex or on one side of
the ovary, usually 2-celled, sometimes 1 -celled by
abortion. Fruits usually 1 -seeded, ovate, gradually
narrowed towards the apex, acute at the base,
IV2-3 by \y^-\y^ cm. Seed ellipsoid, plano-
convex, 10 by 8 mm.
Distr. Louisiade Archipelago (Sudest I.),
Admiralty Is. and Ma/a^^.y/a.- New Guinea (through-
out), Moluccas (Kai Is. and Ceram), PhiUppines
(throughout), and N. Borneo (Banguey I. and
Lahat Datu). Fig. 4.
Ecol. In rain-forest at low and medium alti-
tudes, one collection (NGF 8511) at 1140 m
(Western Highlands, New Guinea).
Vern. New Guinea: kwareo, Wanigela, bearoa,
Gabobora; Philippines: aligpagi, Davao, bdgo.
Bat., tuba, Cag.
Note. Some specimens collected at Davao
Prov., Mindanao, Philippines (B.S. 48963, 49614
and Merrill 11616) seem to be a distinct local
form of P. perrottetiana; this form differs from
the typical one only by the rather small flowers
(c. 2 cm long) and the floral tube which is glabrous

outside. Because of the presence of two kinds of

flowers P. perrottetiana has been placed in the key
twice.
Some specimens {viz Carr 11353, 11672, and
HooGLAND 3788) bear large fruits (c. 3 by 1% cm)
with rather thick pericarps (c. 2 mm). Their
enormous might have been caused by the
size
attack of insects, as there are always hole(s) on
the pericarps and excrements of insects inside the
fruits.

4. Phaleria nisidai Kanehira, F1. Micron. (1933)

248, f. 116; Bot. Mag. Tokyo 47 (1933) 675.
Shrub or small tree, up to 3 m. Branchlets
smooth, glabrous, yellowish-green to reddish-
brown. Leaves chartaceous, greenish when dry
and glabrous on both surfaces, eUiptic-oblong,
rarely lanceolate, 10-18 by 21/2-61/2 cm; base
obtuse, acute or cuneate; apex acuminate; nerves
6-10 pairs, curving and ascending towards the
margin, elevated beneath, plane and distinct
Fig. 5. A tree of Phaleria sp. in bridal attire. above, veins reticulate, usually rather dense,
Kebun Raya Indonesia, Bogor. elevated beneath, distinct or obscure above.
20 Flora Malesiana [ser. I, vol. 6^

Inflorescences terminal and/or in the axils of the Micron. (1933) 248. P. urens Koord. Minah.
terminal node, umbelliform, 10-12-flowered; pe- (1898) 577; Suppl. 2 (1922) t. 98; Suppl. 3 (1922)
duncles c. 2-4 mmwith a pair of opposite, green, 48. P. cauliflora Bedd. For. Man. Bot. (= Fl.
and obovate-oblong bracts (4 by 2 mm) at the Sylv. vol. 3) (1873) 180, t. 25, f. 5; Trim. Fl. Ceyl. 3
upper part; involucral bracts 4, green, glabrous, (1895) 459; Hook./. Fl. Brit. Ind. 5 (1886) 199.—
caducous after anthesis, oblong or elliptic-oblong, Fig. 3a-g, 6.
8-10 by 4-6 mm. Flowers 2-3 V2 cm long. Floral
tube cylindric, slightly dilated towards the top,
glabrous outside, puberulous or pubescent inside.
Calyx lobes 5(-4), oblong, ovate or orbicular,
2-4'/^ by 2-3 mm, puberulous on the margins
and top outside and the whole surface inside.
Stamens included; filaments '/2-3 mm; anthers c.
1 mm long. Disk cup-shaped, crenate, mem-
branous. Pistil usually shorter than the tube,
rarely exserted. Ovary ovoid, c. 21/2 rnm long,
hairy at the apex; style filiform; stigma oblong or
slightly globose. Fruits globose or slightly obo-
void, 1 V2-2 by 1 Vi-l cm, slightly compressed,
constricted at the base into a 3 mm long stipe.
Seeds broadly ellipsoid, 51/2 by 7 mm.
Distr. Western Carolines (Palau: Kanehira
2445, K), in Malaysia: D'Entrecasteaux Is.
(Normanby I.: Brass 25827, K, L), New Britain
(Gazelle Pen.: Waterhouse 908, K), Louisiade
Archipelago (Misima I.: Brass 27505; Rossel I.:
Brass 28284, L), and New Guinea (Eastern
Highlands: NGF 9564).
Ecol. Rain-forests, lowland up to 600 m.
Note. All specimens cited above match very
well with Kanehira's description and his fine
drawing as well as his collection from the type
locality (2445, K). This species is characterized
by the leaves with rather dense venation and the
Fig. 6. Phaleria capitataJack. Botanic Gardens,
flowers with included stamens and pistils.
Singapore, Febr. 1952 (Photogr.
M. R. Henderson).
5. Phaleria capitata Jack, Mai. Misc. (1822) 59;
reimpr. in Hook./. Comp. Bot. Mag. 1 (1835) 156;
K. & V. Bijdr. 13 (1914) 41 S. Moore, J. Bot. 63
; Shrub or small tree, up to 9 m by 16 cm.
(1925) Suppl. 89; Heyne, Nutt. PI. (1927) 1152.— Branchlets reddish-brown. Leaves chartaceous,
Dais dubiosa Bl. Cat. (1823) 69; Bijdr. (1826) 651 glabrous, in dry state reddish-brown above, pale
Hassk. Cat. Hort. Bog. (1844) 94; Filet, PI. Bot. brown beneath; elliptic-oblong, (1 1-)15'/2-21(-26)
Tuin Weltevr. (1855) 50. Drimyspermum mens by (3'/2-)5I/2-7(-10) cm; base acute to attenuate,
Reinw. Syll. PI. Ratisb. 1 (1825) 15; Decne, Ann. rarely rounded; apex narrow acute to acuminate,
Sc. Nat. Bot. II, 19 (1843) 39; Bleeker, Nat. acumen l'/4-2'/2 cm; margins sometimes recurved
Geneesk. Arch. N.I. 2 (1845) 74; Meisn. in DC. in dry state; nerves 8-10 pairs, elevated beneath,
Prod. 14 (1857) 604; Holthuis & Lam, Blumea 5 slightly elevated above; veins loosely reticulate,
(1942) 216. Drimyspermum blumei Decne, Ann. distinct beneath, obscure above; petiole 5 mm.
Sc. Nat. Bot. II, 19 (1843) 39, as Drymispermum; Inflorescences usually terminal and/or in the leaf
Bleeker, Nat. Geneesk. Arch. N.I. 2 (1845) 74; axils of the terminal node, solitary, sometimes
ZoLL. Syst. Verz. 2 (1854) 117; Meisn. in DC. cauliflorous, subsessile or on very short (f. 3 mm)
Prod. 14 (1857) 604; Miq. F1. Ind. Bat. 1, 1 (1857) peduncles, with decussate, small bracts at the
885. P. dubiosa Zoll. Nat. Geneesk. Arch. N.I. base. Involucral bracts 4, oblong, ovate or
1 (1844) 616. Drimyspermum laurifolium (non obovate, 6 by 3 mm, usually caducous after
Decne) Hassk. Nat. Tijd. N.I. 10 (1856) 155.— anthesis sometimes persistent, usually 8-flowered.
Drimyspermum phaleria Meisn. in DC. Prod. 14 Flowers 2'/2^'/2 cm long. Floral tube cylindric,
(1857) 604, as Drymispermum; Miq. F1. Ind. Bat. gradually enlarged towards the top, glabrous on
1, 1 (1858) 884. Drimyspermum cauliflorum both surfaces. Calyx lobes oblong or elliptic, 6-7
Thw. En. Ceyl. PI. (1860) 251, as Drymispermum. by 2-3 '/2 mm> puberulous inside and towards the
—P. cumingii F.-Vill. Nov. App. (1880) 183; upper part and margins outside, sometimes
ViDAL, Phan Cuming. Philip. (1885) 140; Rev. glabrous outside. Stamens and style usually ex-
PI. Vase. Filip. (1886) 230; Brown, Minor Prod. serted sometimes up to 5 mm. Pistil sometimes
Philip. For. 1 (1920) 403; Merr. Bull. Bur. For. shorter than the tube or about as long as it
Philip. 1 (1903) 43; En. Philip. 3 (1923) 131; (Bakhuizen van den Brink 5063, Bakhuizen
Kanehira, Bot. Mag. Tokyo 45 (1931) 331; Fl. VAN DEN Brink/. 688, and San A 3140). Disk
 Dec. 1960] Thymelaeaceae (Ding Hou) 21

cup-shaped. Ovary ellipsoid, glabrous, apex D'AIbertis, New Guinea 2 quoad

narrowed into the filiform style; stigma capitate, —(1880)
basionym, as Drvmispermum. P. revohita Boerl.
398,

1 •/2 by 1 mm. Fruits subglobose, 1-1 Vi cm in Handl. 3 (1900)'lll; Val. Ic. Bog. 4 (1913) 215,
diam., sometimes short-acute at the apex, usually t. 369. P. amboinensis Merr. Philip. J. Sc. 1 1

2-celled, 2-seeded; endocarp inside with distinct (1916) Bot. 29A.—P. platyphvlla Merr. Philip. J.
meshes. Sc. 14 (1919) 429; En. Philip. 3 (1923) 131.—
P. subcaudata Merr. & Perry, J. Arn. Arb. 22
(1941) 265.
Small tree, up to 5 m. Branchlets red-brownish,
usually hollow. Leaves chartaceous to coriaceous,
glabrous on both surfaces; obovate, elliptic,
elliptic- to lanceolate-oblong, obovate- to oblan-
ceolate-oblong, ovate-oblong, 151/2-26 by 6-ll'/2
cm; base rounded to cuneate; margins slightly
recurved; apex short-acute, acute, up to 2 cm
acuminate; nerves 8-14 pairs, prominent and
elevated beneath, slightly elevated or plane above,
curving and ascending towards the margins;
veins loosely reticulate; petiole thick, c. 5-7 mm
long. Inflorescences terminal or/and axillary at the
terminal node, rarely cauliflorous, (15-)20-many-
flowered; peduncle up to 2'/2 cm. Bracts small,

Fig. 7. Localities of Phaleria capitata Jack.

lanceolate, ovate to obovate, 3-10 mm
long;
involucral bracts 4, elliptic- or obovate-oblong,
D i s Ceylon, Carolines (Palau.^f/e Kanehira),
t r.
obtuse, 4-12 by 2-5 mm, caducous after anthesis,
and Malaysia: Malaya (cult, and natur.), Sumatra, rarely persistent. Flowers IVi-'iVi cm long. Floral
Java, Borneo, Philippines, Celebes, Moluccas tube glabrous outside, glabrous or sparsely
(Buru), and New Guinea (Waigeo I.). Fig. 7. puberulous to pubescent inside. Calyx lobes
Ecol. In primary and secondary forests, from obovate-oblong or oblong, 5-7 by 2-3 mm,
the lowland up to 1200 m. reflexed, usually puberulous on both surfaces.

Uses. (C/. Heyne, I.e.; Burk. Diet. 2, 1935, Stamens and style up to 8-10 mmexserted. Disk
1703). Tough fibres of the bark have been used for cup-shaped. Ovary villous or puberulous at the
cordage and tying material. The fruits are sweet top, 2-celled, very rarely 1 -celled. Fruits ellipsoid,
and edible. The seeds are used for scurfy eruptions 1 V2-2 by 1 cm, blunt at both ends, sometimes
in children. apiculate at the apex. Seeds ovoid, 6'/2 by 5 mm.
Vern. Sumatra: siiwa lansat,Simalur, rimbo Distr. New Britain and Malaysia: Philippines
siiloh, Lampong; Java: godong-laweh, kakapassaii, (Panay and Mindanao), Moluccas (Sula Is.,
ki-angkrieng, ki taiigkieh, S, kojoian, lawe, lawean, Ambon, Ceram, and Key Is.), New Guinea
lawe-lawe, ulati, J; Borneo: djariim djariim; (Sorong, Waren, Hollandia, Central and South
Celebes: suka, Bug., sunsiian, susuati, Minah.; Division, Nabire, Sepik region).
Moluccas: la'awana, Sangi & Talaud Is. Ecol. In rain-forests from the lowland up to
Notes. The mesocarp of the fruit is fleshy and 300 m.
rather soft. The endocarp consists of interwoven
fibres which form a characteristic network; 7. Phaleria sogerensis S. Moore, J. Bot. 61 (1923)
through the meshes one can see the testa of the Suppl. 43.
exposed seed (cf. Valeton, Ic. Bog. 4, 1914, 21?). Shrub up to 2 m. Branchlets reddish- to dark-
Fig. 3g. brown, glabrous. Leaves papery, rarely sub-
There are two specimens collected by Cokker coriaceous, glabrous on both surfaces, sometimes
(S.F. 28481 and 32776, Sing) at Johore, Malay brownish or brownish-green when dry, elliptic-
Peninsula, which might have escaped or been oblong, oblong-lanceolate to lanceolate; 7-16 by
derived from cultivation. 2'/2-7 cm; cuneate to attenuate towards the base;
acuminate at the apex, sometimes with an acumen
6. Phaleria coccinea (Gaudich.) F. v. M. Descr. c. 2 cm long; nerves 6-10 pairs, spreading towards

Not. 2 (1885) 9, quoad basionym; K. ScH. & the margin and then curved upwards, prominent
Laut. FI. Schutzgeb. (1900) A59.—Dais coccinea and elevated beneath, slightly elevated above;
Gaudich. Voy. Uranie (1826)443, t. 44. Pseudais veins reticulate, distinct beneath, obscure above;
coccinea Decne, Ann. Sc. Nat. Bot. II, 19 petiole c. 5 mm. Inflorescences terminal, and/or
(1843) 41; Bleeker, Nat. Geneesk. Arch. N.I. in the leaf axils of the terminal one or two nodes,
2 (1845) 77; Meisn. in DC. Prod. 14 (1857) 603; 6-10-flowered; peduncle none to 12 mm; in-
MiQ. Fi. Ind. Bat. 1, (1858) ?,%2.~Drimysper-
1 volucral bracts 4, slightly obovate, c. 5 by 3 mm,
mum revohitum T. & B. Nat. Tijd. N.I. 27 (1864) usually caducous, sometimes persistent. Flowers
30. Drimyspermum cumingii Meisn. in DC. Prod. 1 1/2-2(-2'/2) cm long. Floral tube glabrous on both

14 (1857) 605, as Drymispermum. P. vriesii — surfaces, rarely puberulous inside. Calyx lobes
Baill. Adansonia 11 (1875) 329.—/'. zippelii slightly oblong, 5-8 mm long, densely puberulous
Baill. I.e. —
Drimyspermum coccineum Becc. in inside and on the margins outside. Stamens and
22 Flora Malesiana [ser. I, vol. 6^

Fig. 9. Phaleria macrocarpa (Scheff.) Boerl.

Fig. 8. Phaleria macrocarpa (Scheff.) BOERL.
with its beautiful, glossy, bright-red fruits.
Kebun Raya Indonesia, Bogor
Cultivated in Kebun Raya Indonesia, Bogor.
(C.H.B. VIII-G-93).
K. ScH. & Hollr. F1. Kais. Wilh. Land (1889)
pistil slightly exserted. Disk cup-shaped, crenate, 93; Warb. Bot. Jahrb. 13 (1891) 337.—P. papuana
c. Yi mm long. Ovary usually pubescent or Ware, ex K. Sch. & Laut. F1. Schutzgeb. (1900)
puberulous at the apex, rarely glabrescent or 460; GiLG, Nova Guinea 8 (1910) 411.—
glabrous. Fruits ellipsoid, acute or acuminate P. sp. —
GiLG, I.e. P. calantha Gilg, I.e. P. —
towards both ends, 1 V2 by 1 cm, rarely the basal wichmannii Val. Ic. Bog. 4 (1913) 222, t. 371.—
part of the endocarp inside with small meshes. Fig. 3j-l, 8-9.
Seeds ellipsoid, plano-convex, 8 by 5 mm
with a Shrub or small tree, up to 18 m by 15 cm.
caruncle-like appendage, c. 1 V2 rnm long. Young branches hollow. Leaves chartaceous to
Distr. Malaysia: NewGuinea (Sogere, Kanosia, subcoriaceous, glabrous, ovate-oblong, elliptic-
Koitaki, Boridi, and HoUandia). oblong, lanceolate, oblong-lanceolate, 10-25 by
Ecol. In forests at low and medium altitudes, 3-10 cm; base cuneate or rounded; apex shortly
sometimes up to 1400 m. acute to acuminate, acumen up to 2 cm; nerves
6-1 1 pairs, spreading towards the margins, some-
8. Phaleria macrocarpa (Scheff.) Boerl. Handl. 3 times their distal end curving upwards and united
(1900) 111; L. M. Perry, J. Arn. Arb. 39 (1958) loop-like, elevated and distinct on both surfaces;
420, fig. a-b. D rimysper mum macrocarpum veins loosely reticulate, sometimes subperpendi-
Scheff. Ann. Bot. Card. Btzg 1 (1876) 46, as cular to the midrib, slightly elevated on both
—
Drymispermum. P. octandra {non (L.) Baill.) surfaces. Inflorescences terminal and in the axils
 Dec. 1960] Thymelaeaceae (Ding Hou) 23

along the branchlets, sometimes cauliflorous, 1 to common and can also be observed in for example
5, rarely more peduncles in each axil; peduncles bb 25746, BW 5468 from Hollandia, and NGF
or up to 2'/2 cm (cf. bb 25746), each 2-5(-8)- 7298 from Morobe Distr. The vegetative and
flowered; invoiucral bracts 4, small, obovate, morphological characters of the type agree with
oblong, caducous, 7 by 2 mm. Flowers Vi-^ cm 1 those of P. calantha and P. wichmannii.
long. Floral tube glabrous on both surfaces, some- The type of P. wiclimaumi was collected by
times puberulous and hairy on the inside towards Atasrip (139, Bo, L; cult, in Hort. Bog. under
the base. Calyx lobes oblong, 4 by 2 mm, reflexed, II. VIII. G. 75) in northern Dutch New Guinea;
densely puberulous inside and on the margins its flowers have a long, exserted style and almost

outside. Stamens sessile or up to 6 mm exserted. sessile stamens.

Ovary glabrous; style shorter than or as long as The type of P. calantha was collected by
the tube or c. 5-10 mm exserted. Fruits sub- Versteeg (1939, Bo, L. K; cult, in Hort. Bog.
globose to broadly ellipsoid or rounded, some- under n. VIII. G. 93) at Merauke, southern Dutch
times slightly obovate and stipe-like narrowed New Guinea; its flowers have long filamentous
towards the base, 3-5 '/2 by 3-4 '/2 cm, exocarp stamens and a style shorter than or as long as the
woody when dry. Seed subgiobose or slightly floral tube. The fruits of these two 'species'
ovate, c. Vi by 1 '/i cm.
1 collected from the cultivated plants mentioned
Distr. Malaysia: common in western New above are very similar, large and fleshy. I assume
Guinea. that the flowers are heteromorphous and these
Ecol. Primary and secondary forests, from the two 'species' represent two forms of one species.
lowland up to 550 m, once at 1260 m. Moreover, two other forms have been found,
Uses. It is cultivated in Sabron near Hollandia viz one with both stamens and style exserted as
on clay soil at 120 m. The bark is used by the represented for example by Kloss s.n. (13/1 &
Papuans for making bags (BW 5468). 16/1, 1912 [1913], BM) and Hoogland 4520 (Bo,
Vern. Daloiti, Sentani, kotteh, Djair, matoniek, G, L), and another one with short-filamentous
Andjai. stamens and a short style, represented by NGF
Note. Phaleria macrocarpa was first described 9564 (L).
as Drimyspermiim macrocarpum by Scheffer
{I.e.) based on fruiting specimens collected by Excluded
Teysmann (7786, L, Bo) near Dore, western New Phaleria axillaris Elmer, Leafl. Philip. Bot. 8
Guinea. These young fruits are ellipsoid or (1915) 2840 is according to Merrill (En. Philip. 3,
slightly obovoid (15 by 12 mm) and narrowed 1923, 535) = Tricalysia tinagaoensis Elmer {Rii-
stipe-like towards the base. This shape is not un- biac).

3. ENKLEIA
Griff. Calc. J. Nat. Hist. 4 (1844) 234, in note; van Tiegh. Bull. Soc. Bot. Fr.
40 (1893) 69; Domke, Bibl. Bot. HI (1934) 121, t.4 f.36n & map 3.—Linostoma
subg. Linostoma KuRZ, J. As. Soc. Beng. 39, ii (1870) 83; reimpr. Flora 53 (1870)
312.—Linostoma Wall, ex Endl.: Benth. & Hook./. Gen. PI. 3 (1883) 197,
p.p.—Macgregorianthus Merr. Philip. J. Sc. 7 (1912) Bot. 312.— Fig. 10.
Lianas. Leaves alternate, sometimes opposite towards the upper part of the
branchlets, penninerved with oblique and subparallel cross-bar veins. Inflores-
cences paniculiform, terminal, lax, bearing a few flowers on the top of the ramifica-
tions, each of which always bears a few conduplicate or involute, linear, lanceolate
or oblong bracts; these bracts are subopposite, opposite or alternate, usually
perpendicular to the ramification and sometimes slightly curved upwards when
young, the basal two accrescent and leafy in fruit, horizontally spreading or
sometimes reflexed, slightly enlarged at their attachment (not enlarged in the extra-
Mai, sp.). Flowers 5-merous, articulated at the base of the pedicel. Floral tube
cyhndric, shortly puberulous outside, glabrous inside. Calyx lobes puberulous
on both surfaces. Petaloid appendages twice as many as calyx lobes - or the same
number and then each bifid -, Hnear or oblong, membranous, entire or emarginate,
inserted at the throat of the tube. Stamens included, twice as many as calyx lobes,
in two series, the upper series free from the tube and inserted just below the throat,
the lower series a little below the upper series, sessile, subsessile or short-filamen-
tous; filaments if present slightly broadening towards the base of the anthers;
 24 Flora Malesiana [ser. I, vol. 6^

Fig. 10. Enkleia malciccensis Griff, a. Habit, X %, b. part of stem with hook-lii<e
branchiets, x 2/3,
c. opened flower, x 6, d. stamens, x 13, e. infructescence with two leafy bracts, X 2/3,/. fruit, X 2,
g. cross-section showing structure of pericarp, x A.~E. paiiiculata (Merr.) Hall./.. A. Opened flower!
X 6, /. stamens, x 13 (a Keith 9234, b Maingay 1308/2, c-d Heyne sm., e Elmer 20834,/ Kostermans
7034, /;-/ ZiPPELius 148^).

anthers linear or oblong, slightly apiculate or obtuse; connectives distinct on the

dorsal side and almost as broad as the two locules. Disk none or obscure, some-
times represented by some minute scales. Ovary sessile, ellipsoid or ovoid, densely
Dec. 1960] Thymelaeaceae (Ding Hou) 25

hairy; style terminal, distinct; stigma oblong. Fruits ovoid or ellipsoid, pro-
minently ribbed and reticulate (in the herbarium), surrounded at the base by
the torn remains of the floral tube, with thin exocarp and hard endocarp. Seediht
same shape as the fruit, testa membranous.
Distr. Species 3, distributed in the Andaman Is., Burma, Siam, Indo-China, and Malaysia: Sumatra,
Malay Peninsula, Borneo, Philippines, and New Guinea.
Ecol. In lowland forests.

KEY TO THE SPECIES

1. Leaves subcoriaceous to coriaceous, rather dark often red-brown when dry, usually broad-elliptic
and bluntish. Floral tube not twisted after anthesis. Stamens sessile or shortly filamentous in the
upper series; anther longer than the filament, acute and apiculate at the top. 1. E. raalaccensis
1. Leaves chartaceous, pale (greenish or light brown) when dry, usually ovate-oblong and rather acutish.
Floral tube twisted after anthesis. Stamens distinctly filamentous in the upper series; anther shorter
than or as long as the filament, obtuse to truncate at the top 2. E. paniculata

1. Enkleia malaccensis Griff. Calc. J. Nat. Hist. 4 Uses. Said to give an inferior scented 'gaharu'
(1844) 235, in note; Gilg in E. &
P. Pfl. Fam. Ill, v/ood( fide Heyne, I.e.). The bast fibers can be used
6a (1894) 231; Gamble, J. As. Soc. Beng. 75, ii for tying purpose.
(1912) 262, excl. sviu; Hall./. Med. Rijksherb. Vern. Akar kareh hitam, akar panas, akar
n. 44 (1922) 24; Ridl. F1. Mai. Pen. 3 (1924) 147; puchong kapiir, garu hitaja, kapang akar, M,
Fischer, Kew Bull. (1932) 182; Burk. Diet. 1 tementak akar, Banka, terap akar. Sum. Borneo
;

(1935) 925; Leandri, Rev. Int. Bot. Appl. & aka dian, Kaya, akar garu, Dusun, tuba-tuba,
Agr. Trop. 29 (1949) 505; Proc. 8th Pac. Sc. Bajau.
Congr. Manila 4 (1957) 585. Lasiosiphon scan-
dens Endl. Gen. PI. Suppl. 4, 2 (1847) 67, nam. 2. Enkleia paniculata (Merr.) Hall. /. Med.
illegit.; Meisn. in DC. Prod. 14 (1857) 598; Miq. Rijksherb. 44 (1922) 26. Macgregorianthus
n.

F1. Ind. Bat. 1, 1 (1858) 881.— £. malayana Griff. panicidatusMerr. Philip. J. Sc. 7 (1912) Bot. 312;
Not. As. 4 (1854) 363. Linostoma scandens Gilg in E. & P. Pfl. Fam. Nachtr. 4 (1915) 212;
KuRZ, J. As. Soc. Beng. 39, ii (1870) 83; reimpr. Merr. En. Philip. 3 (1923) 132. E. zippeliana
Flora 53 (1870) 371; Hook. /. Fl. Brit. Ind. 5 Hall. /. /.c— Fig. lOh-i.
(1886) 198, excl. syn.; Boerl. Handl. 3 (1900) 111; Climbing shrub. Branchlets puberulous, glabres-
Heyne, Nutt. PI. (1927) 1152.— £. riouwemis cent. Leaves chartaceous to subcoriaceous, when
Hall. /. Med. Rijksherb. n. 44 (1922) 25.— dry the upper surface light-brown, glabrous and
E. coriacea Hall. /. I.e. —
Fig. lOa-g. shining, the lower surface somewhat paler, dull,
Climber up to 30 mby 10 cm. Branchlets some- sparsely puberulous, glabrescent; ovate-( rarely
times transformed into hook-like organs, reddish elliptic-)oblong, 51/2-II by 3-5 cm; base acute to
brown. Branchlets, inflorescences, and young obtuse; apex acute to ± acuminate; nerves 11-15
leaves always ferrugineous-pubescent. Leaves sub- pairs, slightly ascending towards the cartilaginous
coriaceous to coriaceous, upper surface dull, oli- margin and united with it, elevated beneath,
vaceous when dry, undersurface usually brownish, slightly elevated or plane above; veins slightly
or reddish brown, pubescent on both surfaces, elevated beneath, plane or slightly impressed
sometimes glabrescent; usually broad-elliptic and above; petiole 5-8 mm, densely puberulous.
bluntish, 51/2-14 by 3-7 cm, obtuse or rounded at Inflorescences in the upper axils, up to 28 cm long,
both ends, rarely short-acute; nerves 12-20 pairs, densely puberulous; leafy bracts oblong, 2 ',2-6 by
distinct and plane beneath, visible and impressed %-2 cm. Pedicels 3-5 mm. Flowers pale-green, c.
above, veins distinct beneath, obscure above; 1 cm long. Floral tube cylindric and distinctly
petiole 6-12 mm, pubescent. Inflorescences ter- costate inside, c. 8 mmlong, densely puberulous
minal, up to 30 cm long, flowers (4-)6-8(-14) on outside, glabrous inside, twisted after anthesis.
each ramification; leafy bracts chartaceous, Calyx lobes oblong, c. 2 mm long, densely pu-
oblong, 4—5 by 1-2 cm. Flowers c. 8 mm
long, berulous outside and on the margins or sometimes
yellowish or whitish, short-pedicelled. Calyx lobes the whole surface inside. Petaloid appendages
c. 2 mm long, 3 of them larger and ovate, 2 smaller oblong, membranous, emarginate or slightly erose
and lanceolate. Petaloid appendages 10, linear, c. at the top, '/3-I mm
long. Stamens of the upper
% mrn. Stamens c. 8 mm, sessile or shortly series 1 1/2-2 mm
long with filaments as long as or
filamentous. Pistil included, c. 3 mm
long; ovary longer than the anthers, those of the lower series
c. 2 mm; style short; stigma slightly capitate. c. 1 mm long, sessile or shortly filamentous;
Fruits ovoid, 1 1/4 by 1/2 cm. anthers c. '2 mm
long, obtuse or slightly apiculate.
Distr. Andaman Is., Burma (Tenasserim and Ovary ovoid or ovoid-oblong, c. 2 '2 mm long,
Prome, fide Kurz), Indo-China (Laos and densely pubescent; style filiform, c. 1 '/2 mm;
Cambodia), and Malaysia: Sumatra (Palembang), stigma obovoid, papillose. Fruit ovoid, 1 '/2 by
Riouw, Banka, Malay Peninsula (Singapore and 1 cm.
Malacca), and Borneo. Distr. Malaysia: Philippines (Luzon) and
Ecol. In lowland forests. western New Guinea.
26 Flora Malesiana [ser. I, vol, 6^

Ecol. In hill-side forests (Philip.) or lowland geographic distribution. However, from the two
m.
rain-forest, 50 sheets of the type of £. paniculata (B.S. 12360, Bo,
Note. According to Hallier /. (/.r.) E. zippe- L) as compared with the type of E. zippeliana
liana similar to E. panicitlata, but would differ
is (ZipPELius 148/a. L), it appears that these dif-
in leaves shortly and sparsely puberulous beneath, ferences are only quantitative. There are only
with conspicuous and slightly prominent network young flowers on the Philippine specimens and the
of veins on both surfaces, branchlets, petioles and floral characters are similar to those in the New
panicles minutely rusty (not gray-)tomentose, and Guinea specimen in the same stage.

4. LINOSTOMA
Wall. [Cat. (1831) no 4203, nomen] ex Endl. Gen. PI. (1837) 331; Suppl. 4,
2 (1847) 67; Benth. &
Hook. /. Gen. PL 3 (1883) 197, p.p.; Domke, Bibl. Bot.
111(1934)120, map 3.—Nectandra {non Berg. 1767) Roxb. [Hort. Beng. (1814)

Fig. 1 1. Linostoma longiflorum Hall./., a. Habit, X 2/3. L. pauciflorum Griff, b. Habit, X 2/3, c. upper
part of opened flower, schematic, showing positions of stamens and petaloid scales, x 3, d. the same,
in detail, x 4, e. pistil, x 4,/. disk at base of ovary, x 8, g. stamens, x 13 (a Haviland 1759, b-g
H. M. BuRKiLL 240).
Dec. 1960] Thymelaeaceae (Ding Hou) 27

(90),nomen] Fl. Ind. ed. Carey 2 (1832) 425, non Roland, ex Rottb. Luw- —
stoma sect. Eulinostoma Meisn. in Mart. Fl. Bras. 5, (1855) 71. Psilaea MiQ.
1

Sum. (1861) 355. Luiostoma siibg. Nectandra [{non Berg.) Roxb.] Kurz, J. As.
Soc. Beng. 39, ii (1870) 83; reimpr. Flora 53 (1870) 372.—Fig. 11.
Lianas, rarely erect shrubs. Leaves opposite or subopposite, glabrous, with fine
parallel nerves; margins somewhat reflexed. Inflorescences umbelliform or
paniculiform, few-flowered, usually on the terminal part of the lateral branchlets,
rarely axillary; bracts 2, rarely 3 or 4, discoloured, opposite or alternate. Flowers
cylindric, lobes 5, imbricate, then spreading; pedicels articulated at the base.
Peta/oid appendages 10, long club-shaped or fihform, inserted at the throat of the
tube. Stamens twice as many as the calyx lobes, unequal in length, free from the
tube at the throat; filaments long and slender, usually exserted, broadened into
the connective; anthers oblong, slightly separated by the connective except at the
top. Disk obscure, sometimes just a short toothed ring at the base of the ovary.
Ovary stipitate, oblong or slightly obovate-oblong, densely hairy; style long,
filiform; stigma capitate. Fruits ovoid or globose, surrounded by the cleft base of
the floral tube; pericarp red, crustaceous. Seeds of the same shape as the fruit;
testa membranous.
Distr. About 6 spp., distributed in Siam (Chiengmai, Singora, and Dulit), Burma (Tenasserim,
Silhet,and Chittagong), southern Indo-China (Annam, Laos, Cochin-China), and Malaysia: Malay
Peninsula, Sumatra, and Borneo.
Ecol. In primary and secondary forests, once found in swamp forest, from the lowland up to 1300 m.
Taxon. The genus has been subdivided into two sections by Hallier /. (Med. Rijksherb. /;. 44,
1922, 27) and the two species in our region both belong to the sect. Psilaea (Miq.) Hall. /. I.e. 28.

KEY TO THE SPECIES

1. Bracts leafy, ovate, as large as the ordinary leaves, 23-40 by 14-20 mm, usually opposite or sub-
opposite, covering at least the lower half of the flower. Flowers 2-2 '/^ cm long 1. L. pauciflorum
1. Bracts small, lanceolate, much smaller than the ordinary leaves, 10 by 3 mm, alternate or opposite,
at most covering the base of the flower. Flowers 3-3 '/2 cm long . . 2. L. longiflorum
. .

I. Linostoma pauciflorum Griff. Calc. J. Nat. translucent and with less lateral nerves than the
Hist. 4 (1844) 234, in note; Gamble, As. Soc.
J. leaves. Peduncles 5-10 mm; pedicels c. 7 mm.
Beng. 75, ii (1912) 261; Meisn. in DC. Prod. 14 Flowers green to greenish-white. Floral tube
(1857) 600; Miq. Fl. Ind. Bat. 1, 1 (1858) 882; slender, slightly narrowed towards both ends,
Kurz, J. As. Soc. Beng. 39, ii (1870) 83; For. Fl. 12-15 mm long, usually glabrous on both
Burm. 2 (1877) 334; Hook./. Fl. Br. Ind. 5 (1886) surfaces. Calyx lobes linear. Petaloid appendages
198; BoERL. Handl. 3 (1900) 107, 111; Ridl. J. club-shaped or filiform, c. 5 mm
long. Stamens
Str. Br. R. As. Soc. n. 59 (1911) 164; Fl. Mai. Pen. 7-10 mm, usually exserted. Ovary including the
3 (1924) 146. Psilaea dalbergioides Miq. Sum. stipe 4-6 mm long; style 15-18 mm, terminal or
(1861) 355. L. leucodipterum Hall. /. Med. slightly sublateral; stigma capitate. Fruits ellip-
Rijksherb. rt. 44 (1922) 28; Airy Shaw, Kew Bull. soid, c. I14 cm long, narrowed to both ends.
(1940) 262.— Fig. llb-g. Distr. S. Siam (Singora, Dulit), Burma
A climber up to 24 m, rarely a shrub or small (Tenasserim and E of Tounghoa in the Martaban
tree up to 7 m {cf. H. M. Burkill & Shah 240). Hills), and Malaysia: Sumatra (Simalur), Malay
Branches long, slender, black when dry. Leaves Peninsula (Pedis, Kedah, Dindings, Penang, and
chartaceous, glabrous, rather glaucous beneath, Singapore), and Borneo (W. Borneo),
elliptic, l'/2-4(-6) by l-2(-3i4) cm; apex obtuse Ecol. In primary and secondary forests, from
and mucronate or shortly acute; base acute, the lowland up to c. 1300 m.
cuneate, obtuse, or rounded; petiole c. 2 mm. Yern. Bebora, kakat betul, kakrat biitu or butol.
Inflorescences terminal or rarely axillary, (l-)2-4- perakat betui, tuba bara, M.
flowered, nodding, umbelliform, provided with Use. The Burmese use it medicinally (r/. Burk.
2 opposite or subopposite leafy bracts at the lower Diet. 2, 1935, 1352; field note on Curtis 3197).
half of the peduncle, besides sometimes a small
linear bract (5-7 mm long) at the top of the 2. Linostoma longiflorum Hall. /. Med. Rijks-
peduncle; leafy bracts usually smaller than but herb. /;. —
44 (1922) 29. Fig. 11a.
sometimes as large as the ordinary leaves, ovate or A slender climbing shrub. Leaves chartaceous,
oblong-ovate, 2 1/^-4 by 1 1/2-2 cm, whitish when dry, glabrous, dull on both surfaces, ovate, 3-3 V2 by
28 Flora Malesiana [ser. I, vol. 6^

l'/4-2!/2 cm; apex obtuse and mucronate; base oblanceolate, c. 10 mm long. Petaloid appendages
rounded, sometimes shortly acute. Flowers 10, club-shaped, c. 6 mm long. Stamens c. 10 mm,
solitary or sometimes2, axillary, or terminal on usually exserted. Ovary c. 1 mm long, surrounded
the short branchlets in the inferior leaf-axils of the at the base by a very short disk; style long filiform,
branch; peduncle c. 5 mm, provided with 2 small 3'/^ cm; stigma capitate. Fruit unknown,
bracts; pedicels c. 1 mm. Flowers green. Floral Distr. Malaysia: Borneo (Sarawak),
tube slightly ellipsoid, 2-21/2 cm long, glabrous Ecol. Primary peat-swamp forest, at low
outside, sparsely pubescent inside. Calyx lobes altitude {fide J. A. R. Anderson 9047).

5. WIKSTROEMIA
Endlicher, Prod. Fl. Norfolk. (1833) 47, as Wickstroemia, nee Schrader 1821
(Theac), nee Sprengel 1821 (Comp.), nom. gen. cons.; Gen. PI. (1837) 332,
Suppl. 4 (1847) 68; Domke, Bibl. Bot. Ill (1934) 124, t.4 f.36r& s,map 6, excl.
syn. Stellera L. n. 2. Capura Linne, Mant.
2 (1771) 149, nom. gen. rejic.
PI. —
Diplomorpha Meisn. Denkschr. K. Bayer. Bot. Ges. Regensb. 3 (1841) 289.
Fig. 12.
Shrubs or undershrubs, sometimes trees. Leaves opposite or decussate, very
rarely ternate, of various texture and shapes. Inflorescences terminal and/or
axillary, fascicled or solitary, spicate, racemose, umbelliform or capitate, often
ebracteate. Flowers subsessile or distinctly pedicelled, 4- or 5-merous; pedicel
articulated. Floral tube cylindric or tubular, sometimes slightly funnel-shaped,
usually caducous after anthesis, rarely persistent for some time. Petaloid
appendages O. Calyx lobes usually in two pairs, imbricate, the external ones
cucullate and usually slightly longer than the inner ones. Stamens sessile or filamen-
tous, twice as many as the lobes, included, in two distinct series, usually both free
from the upper half of the tube; anthers oblong, basifixed. Disk membranous,
cup-shaped and shghtly crenate or dentate, deeply lobed, or free and scale-like.
Pistil sessile, rarely short-stiped, included. Ovary usually ellipsoid, glabrous or
hairy at the top, 1 -celled; style terminal, short, distinct or obscure; stigma large,
capitate or disciform, rarely cyUndric to ovoid. Fruits drupaceous, sometimes
surrounded by the dried remains of the floral tube pericarp fleshy or membranous.
;

Seeds of the same shape as the fruit; embryo with thickened or flattened cotyledons
and short or slightly elongated hypocotyl.
Distr. About 70 spp., in SE. Asia, through Malaysia to Australia, Fiji, and Polynesia.
Taxon. All Malaysian species belong to subg. Wikstroemia. Sect. Euwikstroemia Meisn. in DC.
Prod. 14 (1857J 543.— 5«6g. Euwikstroemia Domke, Bibl. Bot. 1 1 1 (1934) tab. facing p. 58 (Type species:
W. australis Endl.).
Nomencl. In Linnaeus's Sp. PI. (1753) Addenda 559, there are two species described under the genus
Stellera, viz 1. S. passerina L. from Europe, and 2. S. chamaejasme L. from Siberia. As these two species
belong to different genera, the generic typification of Stellera L. and its delimitation has caused much
controversy and confusion. As far as 1 could trace, Fasano (Atti Ac. Sc. Fis. Mat. Napoli 1787, 1788,
235) has been the first to point out that these two species do not belong to one genus; he proposed a new
genus Ligia (= Thymelaea) typified by Stellera passerina L. and left S. chamaejasme L. in Stellera.
In 1844, C. A. Meyer (Bull. Ac. Imp. Sc. St. Petersb. 1, 1843, 359; reimpr. Ann. Sc. Nat. Bot. II,
19, 1843, 49) again clearly indicated Stellera chamaejasme L. as the type species of the genus Stellera.
This has been followed by Meisner (in DC. Prod. 14, 1857, 548), Lecomte (Not. Syst. 3, 1914, 212),
Staff (in Curtis's Bot. Mag. 1924, t. 9028), and Hitchcock & Green (Proposals by British Botanists
1929, 150).
Domke merged Stellera L. with Wikstroemia, in transferring its type species, 5. chamaejasme L.,
to Wikstroemia, retaining the taxon as a separate subgenus Chamaejasme [Amman] Domke (cf. Notizbl.
Berl.-Dahl. 11, 1932, 362; Bibl. Bot. Ill, 1934, tab. facing p. 58, and p. 124). Accordingly he made the
new combination, W. chamaejasme (L.) Domke. As Rehder has correctly pointed out (J. Arn. Arb. 15,
1934, 106-107) this is against the Rules of Nomenclature, because \i Stellera and Wikstroemia are united
for taxonomic reasons, Stellera has priority over Wikstroemia, unless Stellera is proposed as a nom. gen.
rejic.
Dec. 1960] Thymelaeaceae (Ding Hou) 29

-1959

Fig. 12. Wikstroemia brachyantlui Merr. —

a. Habit, X 2/3, b. opened flower, x •/,. W. androsaemifolia
Decne. c. Habit, X 2/3, d.opened flower, x 3.— W. tenuiramis MiQ. e. Habit, x 2/3,/. opened flower,
X 3, g. pistil with scale-like disk at base, X 7 {a-b Clemens 32439, c Coert A\, d Rant s.ii., e-J For.
Dep. N. Borneo 4173).

Rehder and Pobedim (F1. U.S.S.R. 15, 1949, 502) have, however, retained Wikstroemia and
{I.e.)
Stellera astwo distinct genera.
If Domke's system will be followed, it will be desirable to conserve Wikstroemia against Stellera.
Uses. The bark is used for tying purpose, rope-making, and is also used in the manufacture of bank-
notes and other strong papers {cf. Brown, Min. Prod. Philip. For. 1, 1920, 403).
30 Flora Malesiana [ser. I, vol. 6^

Notes. Fagerlind (Hereditas 26, 1940, 38 & 48) found an agamogenic clone of IV. indica which is
an intraspecific triploid (2n = 27).
The tropical African Englerodaphne Gilg, reduced by Domke (Bibl. Bot. Ill, 1934, 134) to Gnidia,
looks astonishingly like Wikstroemia and seems to differ from it only by the presence of petaloid ap-
pendages.
KEY TO THE SPECIES
1. Nerves running towards the margin and merging into an intramarginal vein.
2. Leaves membranous to papery, lanceolate to narrow-lanceolate (3-81/2 by 1/2-21/2 cm). Peduncle
slender, terete. Pedicel articulated at the middle or the upper half, after falling of the flower or fruit
leaving a short stalk on the rachis 1. W. lanceolata

2. Leaves subcoriaceous to coriaceous, rarely chartaceous, elliptic- or ovate-oblong, rarely lanceolate

(5-15 by 2'/2-5 cm). Peduncle stout, slightly angular and gradually thickened towards the apex.
Pedicel articulated at the base, after falling of the flower or fruit leaving a prominent scar on the
rachis 2. W. brachyantha

1. Nerves running towards the margin and then curving upwards, not merging into an intramarginal

vein.
3. Leaves usually ovate, elliptic to lanceolate, (P/4-I5V2 by 3^-5 cm), membranous to chartaceous,
rarely subcoriaceous; usually olivaceous to light-brown; apex always acute to acuminate; margins
not cartilaginous. All internodes of the branchlets usually distinct and more than 1 cm long.
4. Inflorescences usually axillary and occurring in several subsequent leaf axils along the branchlets,
sometimes also terminal in addition. (Leaves rather discoloured). 3. W. tenuiramis
. .

4. Inflorescences usually terminal, and/or in the axils of the terminal node.

5. Flowers articulated at the top of the pedicel, after falling of the flower or fruit leaving a short
stalk on the rachis. (Leaves ovate or ovate-oblong, 4-14 by 31/2-5 cm; base usually obtuse or
cuneate, rarely subcordate. Flowers 11/4-2 cm long) 4. W. ovata

5. Flowers articulated at the base or near the base of the pedicel, after falling of the flower or fruit
leaving a prominent scar or a short protuberance on the rachis.
6. Inflorescences racemose; rachis usually elongating, '/2-4 cm long, usually many-flowered,
sometimes also associated with few-flowered inflorescences.
7. Inflorescences usually erect, or slightly curved at the upper part, very rarely nodding from
the base. Flowers loosely arranged on the rachis. Ovary usually glabrous. Anthers usually
apiculate 5. W. polyantha

7. Inflorescences nodding from the base. Flowers densely arranged on the rachis. Ovary hairy
at the top. Anthers usually obtuse 6. W. venosa

6. Inflorescences umbelliform; rachis not elongating, very short or less than 1/2 cm long, few-
flowered.
8. Flowers (15-)18-22 mm
long. Stamens usually sessile; anthers 1 1/2-2 mm
long. Leaf base obtuse,
occasionally shallow-cordate, very rarely attenuate 7. W. meyeniana
8. Flowers 9-15 mm
long. Stamens distinctly filamentous; anthers I-II/2 mm
long. Leaf base
acute, attenuate or obtuse.
9. Flowers 9-12 mm
long, puberulous outside. Leaves acute at both ends, l%-5i/2(-8) by V^-lVi
(-4) cm 8. W.
androsaemifolia
9. Flowers c. 15 mm
long, almost glabrous outside at maturity. Leaves acuminate at the apex,
attenuate, acute or obtuse at the base, (4-)6-15i/2 by (2i/2-)3-5 cm 9. W. ridleyi
. .

3. Leaves usually obovate- or elliptic-oblong, oblanceolate, elliptic, or rarely ovate, li/4-4V2(-7) by

'/2-2(-3i/2) cm, subcoriaceous, brown to reddish-brown; apex usually rounded or obtuse, rarely
acute; margins usually cartilaginous; internodes of the branchlets usually obscure or very short,
2-5 mm long (usually transversely fissured). 10. W. indica

1. Wikstroemia lanceolata Merr. Publ. Govt Lab. an intramarginal vein; veins obscure or distinct,
PhiUp. 29 (1905) 31; Philip. J. Sc. 1 (1906) Suppl. spreading and loosely reticulate; petiole short,
101; 5 (1910) Bot. 366; Brown, Min. Prod. I-21/2 mm, appressed-hirtellous. Inflorescences
Philip. For. 1 (1920) 404; Merr. En. Philip. 3 umbelliform to shortly spicate, terminal, very
(1923) 133.— If. angustissima Merr. Philip. J. Sc. rarely axillary; peduncles very short, sometimes
7 (1912) Bot. 92; En. Philip. 3 (1923) 132. up to 1 1/2 cm, (l-)3-5(-20)-flowered; pedicels c.
An undershrub up to 4 m. Young branchlets 1-11/2 mm, appressed-hirtellous. Flowers green or
densely appressed-pubescent and glabrescent. yeUowish-green, 6-15 mm long, puberulous,
Leaves membranous to papery, glabrous, rarely glabrescent outside. Calyx lobes ovate or oblong,
sparsely pubescent on the midrib beneath, obtuse, c. 1 mm long. Stamens sessile or short-
lanceolate narrowly lanceolate, 3-8I/2 by
to mm
filamentous, c. 3/^-11/2 long. Disk 2 free scales,
1/2-21/2 cm; base obtuse; apex acuminate; nerves linear or slightly oblong. Ovary ovoid, slightly
10-16 pairs, sometimes branched and irregular, hairy at the apex; style obscure; stigma sub-
slightly elevated beneath, visible or obscure above, globose. Fruits short-ovoid, c. 8 by 5 mm, usually
obliquely spreading to the margins and united into glabrous sometimes sparsely hairy at the top,
Dec. 1960] Thymelaeaceae (Ding Hou) 31

pericarp fleshy. Seeds 6 long. mm glabrous. Leaves membranous to papery, glabrous,

Distr. Malaysia: Philippines (Palawan, Min- in the dry state the upper surface subolivaceous or
doro, and Luzon). light-brown, rather shining, lower surface dirty-
Ecol. Common on forested slopes at low and white or light-green, rather dull, sometimes light-
medium altitudes up to 1300 m. brown on both surfaces, ovate-oblong, elliptic-
Vern. Philippines: karanpinig, Neg., maragawa, oblong, broadly-elliptic, or lanceolate, rarely
saldgip, salago, Tag., siika, tuka. Ilk. ovate, 6-12 by 1 Vi-^Vi cm; base cuneate, acute or
obtuse; apex acuminate, the acumen up to c.
2. Wikstroemia brachyantha Merr. Philip. J. Sc. 1 cm; nerves 7-12 pairs, rather irregular, slightly
13 (1918) Bot. 313; En. Philip. 3 (1923) 132.— elevated, rarely indistinct on both surfaces,
W. crassifolia Merr. ex Domke, Bibl. Bot. 1 1 obliquely ascending close towards the margin and
(1934) tab. facing p. 58.— Fig. 12a-b. then curved upward; veins loosely anastomosing,
Shrub or small tree up to 3'/2 m
by 2'/2 cm. reticulations usually obscure on both surfaces;
Branchlets puberulous, light brown. Branches petiole c. 4 mm. Inflorescences usually axillary
reddish-brown and usually transversally fissured. and occurring in several leaf axils along the
Leaves subcoriaceous to coriaceous, rarely branches or branchlets, sometimes also terminal
chartaceous, in dry condition both surfaces or on the top of a reduced or very short branchlet
olivaceous-brown to brownish, glabrous and with bract-like reduced leaves, 1-5-flowered;
shining, elliptic-oblong to ovate-oblong, rarely peduncle very short to 1 1/2 cm, appressed-pu-
lanceolate, 5-15 by 2'/2-5 cm; base obtuse to berulous. Flowers 10-13 mm long, yellowish, or
cuneate; apex acuminate; nerves 12-15 pairs, cream {fide Clemens 20980), subsessile. Floral
elevated and prominent beneath, slightly elevated tube scattered-puberulous outside, glabrescent.
or plane above, obliquely spreading towards the Calyx lobes ovate-oblong, 2-3 mm long. Stamens
margin and united with the intramarginal vein; with c. V2 rnm space between the two whorls,
veins anastomosing, almost as prominent as the those of the upper series sessile or sometimes some
nerves; petiole 2-3 mm, glabrous. Inflorescences of them shortly filamentous, those of the lower
terminal or/and in the leaf axils at the terminal series always shortly filamentous; anthers linear,
node, distinctly peduncled; peduncles stout, 1-1 1/2 rnrn long, acute or slightly apiculate. Ovary
brownish-pubescent, sometimes angular or flatten- oblong or slightly obovoid-oblong, c. 2 mm long,
ed and gradually thickening towards the top, glabrous or a few hairs at the top; style very short
usually bent slightly downward, sometimes with 1 or sessile; stigma capitate, papillose. Fruits yellow,
or 2 bracts below the 6 to 8 apical flowers. Flowers green or orange, ovoid, c. 8 by 5 mm.
yellowish, yellowish-green or green, subsessile. Distr. Malaysia: Sumatra (Menggala), Banka,
Floral tube 10-12 mmlong, sparsely puberulous Borneo (N. Borneo, Brunei, Sarawak, and S.
outside, glabrous inside. Calyx lobes 4, ovate- Borneo: Sampit).
oblong, 1 Vi-i mm long, obtuse. Two series of Ecol. In forests, swampy land, and hills, from
stamens close to each other, on very short fila- the lowland up to 1600 m.
ments; anthers oblong, obtuse, IVi rnrn long. Uses. According to Boorsma {I.e.) it provides a
Disk 2 free, oblong, 2-lobed scales. Ovary ellipsoid scented wood which is used only occasionally.
or slightly obovoid-oblong, 3 mm
long, hairy at The wood is harder than that of Aquilaria and
the top; style distinct, filiform, c. 1 mm; stigma scentless, but when burned it gives forth a frag-
globose, papillose. Fruits red, broadly ellipsoid, rance similar to that of Aloe-yNOod. In Banka the
12 by 9 mm. bark is used for making ropes.
Distr. Malaysia: Borneo (Kinabalu) and Vern. Injat, Brunei, kaju lingau, Menggala,
Philippines (Luzon and Catanduanes). menameng, tementak tindat, Banka.
Ecol. In the Philippines in primary forests at Note. This species can easily be distinguished
low and medium altitudes, in Borneo (Mt Kina- from related ones by the axillary inflorescences
balu) in damp and mossy forests at 1400-2800 m. which occur in several leaf axils along the branch-
Note. This species is characterized by its lets or branches, the more or less discoloured
usually thick leaves with prominent venation es- leaves, and the usually indistinct venation.
pecially on the lower surface, a marginal vein on Mrs Clemens once noted this species to be a
each side, and the stout peduncle. vine (//. 31292) but I believe this to be due to er-
roneous information by her native collectors.
3. Wikstroemia tenuiramis MiQ. Sum. (1861) 141
& 354; Boorsma, Bull. Dep. Agric. I.N. n. 1 (1907) 4. Wikstroemia ovata C. A. Mey. [Bull. Ac. Imp.
19; Heyne, Nutt. PI. (1927) 1152; Burk. Diet. Sc. St. Pdtersb. CI.Ph.-M. 1 (1843) 357; reimpr.
(1935) 2258.— W^. acuminata Merr. J. Str. Br. R. Ann. Sc. Nat. Bot. II, 20 (1843) 50, nomen] ex
As. Soc. n. 76 (1917) 99; En. Born. (1921) 417; Meisn. in DC. Prod. 14 (1857) 544; Miq. F1. Ind.
Un. Cal. Publ. Bot. 12 (1929) 1X9,.— W. dementis Bat. 1. 1 (1858) 880; F.-Vill. Nov. App. (1880)
Merr. J. Str. Br. R. As. Soc. /;. 76 (1917) 99; En. 182; ViDAL, Phan. Cuming. (1885) 140; Rev. PI.
Born. (1921) 417; Heine, Pfl. Clemens Kinabalu Vase. Filip. (1886) 230, excl. syn. Daphne aquilaria
(1953) 69.— Fig. 12e-g. Blanco; Merr. Philip. J. Sc. 1 (1906) Suppl. 101;
Shrub or small tree, up to 10 m. Branchlets Sp. Blanc. (1918) 279; Brown, Min. Prod. Philip.
light-brown to dark-brown, sparsely pubescent, For. 1 (1920) 404; Merr. En. Philip. 3 (1923) 133,
glabrescent. Branches smooth, reddish-brown. excl. citation of Vidal, Synopsis; Quis. Med. PI.
32 Flora Malesiana [ser. I, vol. 6^

Philip. (1951) 637. Daphne indica (non Linne) 5. Wikstroemia polyantha Merr. Philip. J. Sc. 10
Blanco, 837) 309, ed. 2 (1 845) 21 5, ed. 3,
F1. Filip. (1 (1915) Bot. 332; En. Philip. 3 (1923) ]33.— W.
2 (1878) 38. Daphne foetida {non Linne) Blanco, candolleaua {non Meisn.) Ridl. Trans. Linn. Soc
I.e. 308, as phaetida, II. cc. 217, 37. Bot. n, 3 (1893) 341, as candollei, corr. p. 456
Shrub up to 5 m
by 7-8 cm. Young branchlets J. Str. Br. R. As. Soc. n. 35 (1901) 180; Fl. Mai
appressed-hirtellous, glabrescent or glabrous. Pen. 3 (1924) 145; Burk. & Hend. Gard. Bull
Leaves membranous or papery, glabrous, rarely S.S. 3 (1925) 417; Heyne, Nutt. PI. (1927) 1152
sparsely pubescent on the midrib beneath; ovate to BuRK. Diet. (1935) 2258; Symingt. J. Mai. Br
ovate-oblong, 4-14 by 3'/2-5 cm; base usually R. As. Soc. 14 (1936) 358.— PF. junghuhnii {non
obtuse or cuneate, rarely subcordate; apex acumi- MiQ.) K. & V. Bijdr. 13 (1914) 58, sphalm. jiing-
nate; nerves 8-12 pairs, curved and ascending, huhniana. — W. ridleyi {non Gamble) Gibbs, J
slightly elevated below, distinct above; veins Linn. Soc. Bot. 42 (1914) 132; Merr. En. Born
reticulate, distinct beneath, visible or obscure (1921) A\l.— W. calva Back. Blumea 5 (1945) 494
above, petiole 3 mm, sparsely appressed-hirtellous. Shrub or small tree up to 7 m by 7V2 cm
Inflorescences terminal, short-spicate or umbelli- Branchlets reddish to dark-brown, sparsely pu
form, peduncled, sparsely puberulous, sometimes berulous, glabrescent. Leaves membranous, char
with 1 or 2 caducous bracts, 7-20-flowered. taceous, rarely subcoriaceous, glabrous on both
Flowers 114-2 cm long, greenish, yellowish, at the surfaces, rarely scattered hairy beneath especially
upper end of the peduncle, short-pedicelled. on the midrib, in dry condition light-brown to
Floral tube cylindric, sparsely puberulous outside. dark-brown above, paler beneath, ovate-oblong,
Calyx lobes oblong, obtuse, 2-4 mm
long. Stamens elliptic-oblong, or lanceolate, 6-9(-12) by 1 Vi-Wi
sessile or on short filaments; anthers 1-1 V2 vnm, (—41/2) cm; base acute to cuneate, sometimes ob-
slightly apiculate. Disk 2 free, oblong, scales. tuse or rounded; apex acuminate, rarely acute;
Ovary ellipsoid, 2-3 mm
long, hairy at the apex; margins sometimes slightly recurved; nerves 8-15
style distinct, filiform, %-l 14 mm; stigma capitate. pairs, irregular, often branched, elevated beneath,
Fruits subglobose to slightly ellipsoid, 8-10 by distinct or plane above, obliquely ascending tow-
6-8 mm. ards the margin; veins obscure on both surfaces,
Distr. Malaysia: Borneo (North Borneo, sometimes as distinct as the nerves; petiole 2-4
Sebattik I., Sampit, and Pulu Lampei) and the mm, sparsely pubescent. Inflorescences terminal
Philippines (Palawan, Mindoro, Luzon, Negros, or/and in the axils at the terminal node, spicate,
and Mindanao). Fig. 13. gradually elongating, up to 4 cm, rarely to 6 cm,
Ecol. In thickets, primary and secondary erect or slightly curved, very rarely nodding
forests at low and medium altitudes up to 800 m. (S.F. 20726), with 6 to many flowers; peduncle
distinct, sparsely hairy. Flowers c. 10 mm
long,
yellow, yellowish-green, or rarely white {fide
Ridley), loosely arranged on the rachis; pedicels
very short, '/2-I mm, puberulous, articulated at
the base. Floral tube scattered puberulous outside,
glabrous inside. Calyx lobes oblong or ovate-
oblong, 1 V2-3 V2 mrri long. Anthers linear, c.
1 mm long, the two series close to each other;
filaments c. V2 the length of the anther. Disk 2
free, linear or obovate-oblong, c. 1 mm
long,
irregularly lobed or dentate scales. Ovary ellipsoid
or obovoid, l'/2-2V2 rnm long, glabrous or
sparsely hairy at the top; style distinct, as long as
or slightly longer than the stigma; globose or
slightly oblong, c. Vi mm long, papillose. Fruits
ovoid, red, c. 8 1/2 by 5V2 rnm.
Distr. Malaysia: Malay Peninsula (Pahang,
Kedah, Perak, Johore, Kelantan, Selangor, and
Fig. 13. Localities of Wikstroemia ovata C. A. Mey. Gunong Korbu), Java (western part), North
ex Meisn. Borneo, and Philippines (Luzon).
Ecol. In forests, from the lowland up to 2200 m.
Uses. This plant has been used by the Filipinos Use. The wood yields incense.
The leaves are a strong purgative
as a purgative. Vern. Chandan pelaudok, M.
when chewed and swallowed and one bowel Notes. Backer {I.e.) has pointed out that the
movement is produced for every taken. The fresh name " Wikstroemia junghiihniana" {non MiQ.)
bark or branches of this plant are tied round given by Koorders & Valeton {I.e.) was an error
about the neck of a patient to relieve bronchial because they intended to identify their specimens
catarrh (r/. Garcia, Philip. J. Sc. 51, 1933, (Koorders 26824 y3 and J. J. Smith 292, Bo) as
485-494; Quis. I.e.). W. junghuhnii MiQ. (= W. androsaemifolia
Vern. Philippines: arandon. Ilk, dapnit, suka, Decne). The sheets cited by Koorders & Valeton
Bon., salago. Tag.; Borneo: gelam hutan, Brunei, under " W. junghuhniana'' as mentioned above
pait-pait, Bajau. differ from W. junghuhnii MiQ. and belong to a
Dec. 1960] Thymelaeaceae (Ding Hou) 33

different species; therefore, Backer provided Fl. Gen. I.-C. 5 (1915) 167; Leandri, Proc. 8th
them with a new name " Wikstroemia calva" to Pac. Sc. Congr. 4 Bot. (1957) 582, inch var.—
indicate the 'completely glabrous ovary tip". W.fenicis Merr. Philip. J. Sc. 13 (1918) Bot. 312;
However, whether the ovary is glabrous or hairy En. Philip. 3 (1923) 132.
at the apex is not a constant character as some Shrub up to 3 m. Young branchlets sparsely
specimens {e.g. van Steenis 4187 & 12930) have pubescent, glabrescent, sometimes glabrous. Leav-
both kinds of ovaries. Specimens collected at high es papery, glabrous, olivaceous, shining, lanceolate
altitude have thicker and reddish-brown leaves. rarely elliptic-lanceolate, (5-)9i/2-13 by (II/2-)
The species is characterized by the long-spicate 3'/2-4'/2 cm; base obtuse occasionally shallow-
inflorescences which are gradually elongating cordate or attenuate;apex acuminate; nerves
during flowering time. It is closely related to W. 9-14 pairs, slightlycurved and ascending, elevated
nutans Champ, from Kwangtung, China, diff"ering beneath, visible or obscure above, veins reticulate,
from it by the larger leaves, distinct style, and the slightly elevated beneath, obscure above; petiole
approximate insertion of the two whorls of 3-4 mm. Inflorescences terminal and axillary, um-
stamens. belliform, sometimes occurring on short, reduced
The inflorescences are erect or slightly curved branchlets associated with reduced leaves (2-3 cm
with the exception of one specimen (S.F. 20726) long) and resembling a leafy panicle, (2-)5-6
collected in Pahang at c. 2100 m, which has both (-lO)-flowered; peduncle up to 3 cm, densely ap-
erect and nodding ones. pressed-hirtellous. Flowers greenish-yellow, (15-)
18-22 mm long, densely puberulous outside.
6. Wikstroemia venosa Merr. & Perry, J. Arn. Floral tube cylindric. Calyx lobes narrow-
Arb. 22 (1941) 266. oblong, 2-31/2 rnm long. Stamens usually sessile
Shrub, c. m. Young branchlets densely yellow-
1 rarely some of them on short filaments, 1/2-2
1 mm
ish-brown puberulous; older ones reddish- or long. Disk 2 free, oblong scales. Ovary slightly
dark-brown, puberulous, glabrescent. Leaves obovoid, 2-21/2 mm long, hairy at the apex; style
chartaceous or subcoriaceous, lower surface of the distinct, 1/2-I mmlong; stigma oblong, 1/3-1/2 mm
young leaves densely pubescent on the midrib and long. Fruit ovoid, c. 8 by 6 mm.
scattered pubescent on the lamina, glabrescent; Distr. Indo-China and Malaysia: Philippines
in dry condition light brown or brown above and (Luzon to Mindanao).
shining, glaucous beneath; ovate-oblong to Ecol. In primary humid forests at low and
lanceolate, 2-7 by 1-3 cm; base obtuse or cuneate; medium altitudes, up to c. 400 m.
apex acute to acuminate; nerves 7-10 pairs, rather Vern. Philippines: sagii. Tag., salago, Bik., Tag.
irregular, obliquely spreading towards the margin
and then slightly curved ascending, elevated 8. Wikstroemia androsaemifolia Decne, Ann. Sc.
beneath, plane or slightly depressed above; petiole Nat. Bot. II, 20 (1843) 50; in Jacq. Voy. Bot.
c. 2 mm long, puberulous. Inflorescences terminal, (1844) 146; Bleeker, Nat. Geneesk. Arch. N.I. 2
nodding, sessile or up to 1 cm peduncled; spicate, (1845) 74; Meisn. in DC. Prod. 14 (1857) 546;
rachis elongated, cm, puberulous. Flowers
1/2-2 MiQ. Fl. Ind. Bat. 1, (1858) 879; Back. Bekn. Fl.
1

crowded, c. mm,
green; pedicel c.
8 mm, ar- 1 Java (em. ed.) 4A (1942) fam. 77, p. 5.~IV.
ticulated at the base, puberulous. Floral tube spanoghii Decne, Ann. Sc. Nat. Bot. II, 20 (1843)
scattered-puberulous outside and glabrous inside, 50; in Jacq. Voy. Bot. (1844) 146; Bleeker, Nat.
sometimes glabrous outside. Calyx lobes oblong Geneesk. Arch. N.I. 2 (1845) 74; Meisn. in DC.
or ovate-oblong, 2-2 '/2 mm
long. Stamens shortly Prod. 14 (1857) 545; Miq. Fl. Ind. Bat. 1, I (1858)
filamentous; anthers linear, c. 1 mm
long, obtuse, 879. Eriosolena viridiflora Zoll. & MoR. Nat.
the space between the two series c. mm. Disk % Geneesk. Arch. N.I. (1844) 615, e.xcl. syn. Hassk.
1

2 free linear scales, bilobed at the top, sometimes Cat. 117; Zoll. Syst. Verz. 2 (1854) 116.— ^f'.
disk cup-shaped and erose at the top, '/2-I Tim candolleana Meisn. in DC. Prod. 14 (1857) 544;
long. Ovary slightly obovate-oblong, c. 1 Y2 mm K. & V. Bijdr. 13 (1914) 56; Miq. Fl. Ind. Bat. 1,
long, sparsely strigose at the top; style short, c. 1 (1858) 878; Hall. /. Med. Rijksherb. n. 12
Vs mm; stigma capitate. Fruits ellipsoid, 6-8 mm (1912) 26; K. & V. Bijdr. 13 (1914) 56; Heyne,
long, slightly narrowed at both ends. Seed similar Nutt. PI. (1927) 1152; Burk. Diet. 2 (1935) 2258.
in shape to the fruit. — W.junghuhnii Miq. Fl. Ind. Bat. 1, (1858) 879;
1

Distr. Malaysia: New Guinea (Balim Valley, Hall. ./: Med. Rijksherb. n. 44 (1922) 30.—
Manokwari, Humboldt Bay, HoUandia, and Rona Fig. 12c-d.
(Central Div.)). Shrub up to 21/^ m
by 4 cm. Young branchlets
Ecol. Deforested slopes, grassland, occasionally slightly flattened at the nodes, densely appressed-
on grassy banks of streams, lowland up to 1900 m. pubescent, glabrescent. Branches terete, reddish-
brown, glabrous; axillary buds densely covered
7. Wilistroemia meyeniana Warb. in Perk. Fragm. with golden-coloured hairs. Leaves papery,
Fl. Philip.(1905) 171; Merr. Philip. J. Sc. 1 (1904) glabrous, rarely sparsely hairy on the lower
Suppl. 101 Brown, Min. Prod. Philip. For. 1
; surface and especially on the nerves and veins of
(1920) 404, t. 23; Merr. En. Philip. 3 (1923) 133. young leaves, in dry state light-greenish, light-
— Daphne cannabina {non Lour.) Schauer, Nov. brown or greenish-brown to brownish and shining
Act. Caes. Leop.-Car. 19 (1843) Suppl. 1, 411.— on the upper surface; pale-greenish, light-yellowish
W. longifolia Lecomte, Not. Syst. 3 (1914) 128; -green or light-brown and dull on the under-
34 Flora Malesiana [ser. I, vol. 6^

surface; elliptic, elliptic- or ovate-oblong, rarely 1 800 mhave leaves with distinct, densely reticulat-
broadly elliptic, iy4-5i/2(-8) by y4-2V2(-4) cm; ed venation, and flowers with distinct peduncles
base acute; apex acute to narrow-acute, very rarely and short styles.
obtuse; nerves 8-11 pairs, elevated below and Handel- Mazzetti described a diff'erent species
slightly depressed above, obliquely spreading from Yunnan, China, as W. androsaemifolia in
towards the margin and then curved upward; 1923. If this proves to be a good species it must be
veins almost as distinct as the nerves, loosely renamed.
reticulate beneath, obscure above; petiole c. 2 mm.
9. Wikstroemia ridleyi Gamble, Kew Bull. (1912)
Inflorescences umbelliform or spicate, 5-10-flower-
200; J. As. Soc. Beng. 75, ii (1912) 260; Ridl. F1.
ed, terminal and in the axils of the terminal node,
Mai. Pen. 3 (1924) 146, f. 147; Burk. Diet. (1935)
so usually 3 inflorescences at the top of the
2258.
branchlet, of which usually the middle one (some-
Shrub up to 2 m. Branchlets reddish-brown,
times also the lateral ones) is provided with a
sparsely puberulous and glabrescent. Leaves mem-
pair of bracts or reduced leaves; peduncle obscure
branous to chartaceous, usually olive-brown when
to 31/2 cm, erect or slightly curved; pedicels c.
dry, glabrous on both surfaces, rarely sparsely
1 mm, articulated at the base. Flowers light-green
puberulous on the midrib beneath, elliptic-oblong,
or yellowish-green. Floral tube slightly pubescent
lanceolate, ovate-oblong or ovate, 4-151/2 by
outside, 9-12 mm
long. Calyx lobes oblong, or
2 1/2-4 V2 cm; base attenuate, acute or obtuse; apex
slightly ovate, fleshy, 2-3 '72 mm
long, obtuse.
acuminate; nerves 7-12 pairs, slightly curved
Disk 2, rarely 3 free, linear scales. Stamens
towards the margin and then upward, sUghtly
filamentous; anther c. 1 mm
long, obtuse, the
elevated on both surfaces; veins obscure on both
space between the anthers of upper and lower
surfaces; petiole 2-3 mm, sparsely puberulous
series 1-1 1/2 mm. Ovary ellipsoid or slightly
obovoid, \V2-2y2 mm
long, pilose at the top,
when young. Flowers c. 15 mm
long, yellow or
greenish-yellow, 6-14, umbelliform on a terminal,
sometimes glabrescent; style obscure to 1 mm very short, slightly puberulous peduncle; pedicels
long; stigma globose, c. Y^ mm
in diam. Fruits red,
c. 1 mm, articulated towards the base, puberulous.
oblong, rounded.
Floral tube sparsely puberulous outside, glabres-
Distr. Malaysia: Central and East Java,
cent, glabrous inside. Calyx lobes ovate-oblong,
Madura, Kangean Arch., Lesser Sunda Islands
(Flores and Timor), Borneo (North Borneo,
obtuse, 3'/2-4 mm
long. Stamens shortly filamen-
tous, free from the tube at the upper half, the two
Koetai and Balikpapan), Celebes (Bonthain,
Manado, and G. Pangararan), and W. New
series c. 1 mmapart; anthers linear, obtuse or

Guinea.
slightly apiculate, c. 1
1/2 mm long. Disk 2 free

Ecol. In lowland forests from near the beach

linear, c. 1 mm long, 2-lobed scales. Ovary
up to 1800 m, in Celebes at 2200-2400 m.
ellipsoid or slightly obovoid, 1 1/2-2 long, mm
hairy at the apex; style very short or obscure;
Note. There are two authentic sheets of W.
stigma globose and papillose. Fruits red, ellipsoid
androsaemifolia in the Nat. Hist. Mus., Paris; one
or ovoid, 8 by 5 mm. Seeds ovoid, the same shape
is labelled as 'Java Leschenault' and the other as
as the fruit.
'Daphne n. 341'. Apparently these specimens may
Distr. Lower Siam (Telok Udang) and
belong to one collection, as they are very similar.
Malaysia: Malay Peninsula, chiefly on the east
The leaves are papery and rather discoloured,
coast (Kelantan, Trengganu, Pahang, Burau Bay).
brownish above, light-brown beneath, and not
Ecol. Sandy open coastal country.
larger than 6 by 2'/2 cm. The flowers are about
Uses. The species contains a purgative sub-
10 mm long, sessile, and fascicled or crowded on
stance and the leaves are eaten as an aperient. The
a short peduncle (c. 10 mm). The ovary is sparsely
bark is used as entering into a compound potion
hairy or glabrous at the top.
against small-pox; it is pounded and converted
The type of W. spanoghii was collected by
into a poultice for applying to boils, or merely tied
Spanoghe {s.n., L) in Timor. Its leaves are rather
round the neck to stop vomiting. It is also used as
membranous, pale-greenish beneath, light-green-
a fish-poison. The fruits are poisonous (c/.
ish above, and the size is up to 8 by 21/2 cm. It has
BURKILL).
longer, spicate inflorescences and the longest
Vern. Depu, depu pelandok, M; dalu pelandok
peduncle is c. V/i cm. The ovary is densely hairy
is a misprint {cf. Burkill, I.e.).
at the top, and the style is obscure.
Note. According to Ridley this plant was
When one compares the type specimens of these brought from Pekan (not: Penang!) to Singapore
two species they do not seem to be conspecific.
in 1890 and "ran wild for some time in Tanglin,
However, after examining a large range of spec-
Singapore".
imens, there are too many intermediate forms in Three specimens have been cited in the original
which the diff'erential characters break down.
description, collected by Ridley at Pahang
Consequently I have interpreted them as forms of
(Kwala Brawas: Ridley 1583, lectotype and
one variable species.
Pekan: Ridley s.n., Aug. 1889, paratype) and
The type specimen of W. junghuhnii MiQ.
Tringganu (Pulo Katan: Ridley s.n., Aug. 22,
collected by Junghuhn (s.n., L) on Mt Ungaran
1899, paratype).
(Central Java) and some specimens {e.g. Koor-
DERS 43148 ^, 43875 /3, 43876 /3, and van Steenis 10. Wikstroemia indica (L.) C. A. Mey. Bull. Ac.
17974) collected on mountains between 1100- Sc. St. Petersb. 1 (1843) 357; reimpr. Ann. Sc. Nat.
Dec. ]960] Thymelaeaceae (Ding Hou) 35

Bot. II, 20 (1843) 50; Meisn. in DC. Prod. 14 florescence terminal, sometimes 1-2 additional
(1857) 543; MiQ. Fl. Ind. Bat. 1, 1 (1858) 880; ones in the axils of the terminal node, few-
Benth. Fl. Austr. 6 (1873) 37; F.-Vill. Nov. App. flowered, subsessile, sometimes on a very short
(1880) 182; Vidal, Rev. PI. Vase. Filip. (1886) peduncle; pedicels 1 '/2-2 mm, articulated at the
229; FoRB. &
Hemsl. J. Linn. Soc. Bot. 26 (1894) base. Flowers green, 10-12 mm
long, sparsely
398 BoERL. Handl. 3 ( 1 900) 111; Bailey, QueensI
; puberulous outside, glabrous inside. Calyx lobes
Fl. pt 4 (1901) 1369; Bold. Zakfl. (1916) 171 2-3 mm
long, broadly ovate or oblong, obtuse.
Merr. Sp. Blanc. (1918) 279; Brown, Min. Prod Stamens very shortly filamentous, rarely sessile;
Philip. For. 1 (1920) 404; Merr. En. Philip. 3 anthers linear, c. 1 mm
long, sometimes those of
(1923) 132; Rehder, J. Arn. Arb. 15 (1934) 103 the lower series slightly shorter, obtuse rarely
Merr. Comm. Lour. (1935) 278; Back. Bekn. Fl apiculate at the apex, the two series c. 1 apart. mm
Java (em. ed.) 4A (1942) fam. 77, p. 4; Holth. & Disk 2 free, linear, c. % mm
long scales with
Lam, Blumea 5 (1942) 216. Daphne indica Linne, narrowed or obliquely truncate top, sometimes
Sp. PI. (1753) il 5.— Daphne aquilaria Blanco, FI lobed or crenate at the apex. Ovary slightly
Filip. (1837) 310; ed. 2 (1845) 216; ed. 3, 2 (1878 obovoid or elliptic, c. 1 Yj mm
long, sparsely hairy
39. — W. Meisn. Denkschr. K. Bayer
viridiflora or glabrous at the top; style very short or obscure;
Bot. Ges. Regensb. 3 (1841) 286; Decne, in Jacq stigma globose, c. Yi mm
diam. Fruits broadly
Voy. Bot. 4 (1844) 145; Meisn. in DC. Prod. 14 ellipsoid, c. 6 by 4 mm, red.
(1857) 546; Miq. Fl. Ind. Bat. 1, 1 (1858) 879 Distr. India, SE. Asia, through Malaysia to
Vidal, Rev. PI. Vase. Filip. (1886) 229; Merr Australia (N. Australia, Queensland, and N.S.
Philip. J. Se. 3 (1908) Bot. 422; Back. Ann. Jard Wales) and Melanesia (as far E as Fiji), in
Bot. Btzg Suppl. 3 (1909) 419; K. & V. Bijdr. 13 Malaysia not found in the seasonal parts: absent
(1914) 54; RiDL. Fl. Mai. Pen. 3 (1924) 145 from the Lesser Sunda Islands, in Java only found
Guillaumin, J. Arn. Arb. 13 (1932) 88; Burk in the vicinity of Bogor and once found at 100 1 m
Diet. (1935) 2259; Leandri, Proe. 8th Pae. Se near Sindanglaja as an escape from the Botanic
Congr. Manila 4 (1957) 582.— ff. ovato (non C. A Gardens, now locally thoroughly naturalized and
Mey.) Vidal, Synopsis (1883) 229.— W. indica var slowly spreading along roadsides and in other
viridifloraHook./. Fl. Br. Ind. 5 (1886) 195.— anthropogenous terrain (Backer, 1909 I.e.).
W. linearifolia Elm. Leafl. Philip. Bot. 2 (1910) 680; Ecol. In thickets and secondary growths, ob-
Merr. En. Philip. 3(1923) 133. W. pidgarensis — viously very soil-tolerant and occurring in various
Elm. Leafl. 5 (1913) 1844; Merr. En. Philip. 3 biotopes, for example on sandy soil near the
(1923) 133.— If. pachyphrlla Merr. Philip. J. Se. beach, on limestone of a ridge top, on granite
12 (1917) Bot. 297; En. Philip. 3 (1923) 133.— peaks, along river-banks, and on open hill-sides,
W. siibcoriacea Merr. J. Str. Br. R. As. Soe. //. 76 from the lowland up to 1 300 m, a few above 2200
(1917) 100; En. Born. (1921) 417; Heine, Pfl. m even up to 2700 m (in Celebes and New Guinea).
Clemens Kinabalu (1953) 69. Daphne sp. Steen. Vern. Borneo: lajak, M; Philippines: inyam,
Bull. Jard. Bot. Btzg III, 13 (1933) 254. P. Bis., aranddn, baleo. Ilk., palupo, titpuho, Iv.,
Shrub up to 3 m. Branchlets black-brown, salago. Tag., Bis., Bik., talo, Bik.; Celebes:
scattered puberulous, glabreseent, sometimes perapata or posi-posi, Manado.
transversally fissured. Internodes usually very Note. W. indica is a widely distributed species
short or even obscure. Leaves chartaceous to sub- and is very variable in its vegetative parts. As
coriaceous, in dry condition usually brown to pointed out by Bentham in a note under W. indica
reddish-brown, sometimes glaucescent, sparsely (Fl. Austr. 6, 1873, 37), "it is, however, not always
puberulous beneath, glabreseent, or glabrous, easy to determine the limits to be assigned to it".
shining above and rather dull beneath, obovate- Fagerlind (I.e.) has found apomixis in this spe-
or elliptic-oblong, oblaneeolate, elliptic, rarely cies, which may give an explanation of its great
ovate, 1 '/4^'/2(-7) by '/2-2(-3i/4) cm; base cuneate vegetative variability and difliculties involving in
to attenuate; apex rounded, obtuse, sometimes specificdemarcation. With a large number of
slightly emarginate, or acute; margins usually specimens of this species available, no sharply
cartilaginous; nerves 5-12 pairs, irregular, and defined infraspecific taxa or forms can be dis-
often branched, obliquely ascending towards the tinguished.
margin, rarely the basal or 2 nerves on each side
1 Excluded
ascending along the margin towards near the top, W. amplifolia (Schltr) Domke(Bibl. Bot. HI,
usually distinct beneath obscure above, sometimes 1934, 60) of New Caledonia (isotype: Schlechter
obscure on both surfaces; veins obscure or in- 14749, L) has erroneously been recorded for New
visible on both surfaces. Petiole c. 2 mm. In- Guinea by Domke (/.c).

6. DAPHNE
LiNNE, Gen. PI. ed. Meisn. in DC. Prod. 14
5 (1754) 167; Sp. PI. (1753) 356;
(1857) 530; Gilg, in E. &
6a (1894) 237; Domke, Bibl. Bot. HI
P. Pfl. Fam. 3,
(1934) 130, t.4 f.i & map S.—Scopolia Linne/. Suppl. (1781) 409, non Jacq. 1764,
—
nee al. Eriosolena Bl. Bijdr. (1826) 651; van Tiegh. Bull. Soc. Bot. Fr. 40
(1893) 67; Domke, Bibl. Bot. Ill (1934) 70-83, 130, t.4 f.36 C & map 10.—
36 Flora Malesiana [ser. I, vol. 6^

Fig. 14. Daphne luzonica C. B. Rob. a. Habit, x 2/3, b. opened flower, x 4. D. composita (L. /.) Gilg.
c. Habit, X 2/3, d. opened flower, X 4, e. fruit, X 2, f. seed, X 2, g. longitudinal section of fruit, X 2

{a-b BS 40335, c-rf Rahmat si Boeea 11238, e-^ SF 51832).

Daphne sect. Eriosolena Meisn. Denkschr. K. Bayer. Bot. Ges. Regensb. 3 (1841)
283; in DC. Prod. 14 (1857) 540.— Fig. 14.
Shrubs, rarely small trees or dwarf shrubs. Leaves spirally arranged, sometimes
subopposite or crowded towards the upper part of the branchlets. Inflorescences
Dec. 1960] Thymelaeaceae (Ding Hou) 37

usually capitate, ebracteate or surrounded by caducous bracts, terminal and/or

axillary, sessile or peduncled, sometimes racemose or a few flowers in a fascicle,
rarely paniculiform, usually with some linear bracteoles in the leaf axils or at the
base of the peduncle. Flowers 4-merous, sessile. Floral tube cylindric or slightly
infundibuliform, glabrous or pubescent outside, usually caducous after anthesis,
rarely persistent and surrounding the fruit (in extra-Mai. spp.). Calyx lobes 4,
erect or spreading, alternating longer and shorter. Petaloid appendages none.
Stamens 8, in two rows, sessile or on short filaments; anthers linear, dorsi- or
basifixed. Disk annular and entire, or membranous and irregularly toothed or
split, sometimes elongated on one side, or obscure, or absent. Pistil always
included in the floral tube. Ovary ovoid, sessile or slightly stalked, usually hairy
towards the top or in the upper half; sometimes glabrous; style sessile or short-
fihform, terminal, sometimes slightly lateral (in extra-Mai. spp.); stigma globose
or capitate. Drupe ovoid or ellipsoid, with fleshy or dry pericarp, endocarp
sclerified. Seed similar in shape to the fruit; testa crustaceous.
Distr. Species c. 70, distributed in the Old World on the northern hemisphere, from Europe and
northern Africa to eastern Asia and Malaysia.
Ecol. The genus is represented in Malaysia by two species of widely different affinity. D. composita
belongs to a small section Eriosolena (Bl.) Meisn. {cf. Gilg in E. & P. Pfl. Fam. 3, 6a, 1894, 238) which
is restricted to the undergrowth of the montane rain-forest of SE. continental Asia and West Malaysia,

centering in Asia. D. liizonica belongs to a section Daphuanthoides Gilg (/.c.) which occurs chiefly in
the Himalaya, China, Japan, and Formosa, and has reached northern Luzon where it occurs at high-
montane altitude.
Note. Scopolia {non Jacq. nee al.) Linne/. and Eriosolena Bl. which are congeneric and even based
on the same species (though with different type specimens) have been separated from Daphne because
the flower of this taxon possesses a tubular hypogynous disk, it being absent in Daphne. In 1841 Meisner
{I.e.) reduced Eriosolena Bl. to a section of Daphne and in 1857 (I.e.) he reduced also Scopolia L. /. to

Daphne. The reduction o^ Eriosolena has been adopted e.g. by Baillon (Hist. PI. 6, 1877, 131), Bentham
& Hooker (Gen. PI. 3, 1880, 190), Gilg (in E. & P. Pfl. Fam. 3, 6a, 1894, 238), Boerlage (Handl.
3, 1900, 105), Backer (Bekn. Fl. Java, em. ed., 4A, 1945, fam. 77, p. 5), and others.
In 1893 VAN Tieghem (Bull. Mus. Hist. Nat. Paris VII, 17, p. 195; Bull. Soc. Bot. Fr. 40, p. 68) restored
Eriosolena to generic rank, basing himself on anatomical characters of the branches and leaves and the
presence of a tubular disk.
In 1914 H. Lecomte (Not. Syst. 3, 99) agreed with van Tieghem, adding that Eriosolena was, besides,
characterized by typical, caducous, involucral bracts. In passing it may be remarked that such bracts
also occur in Daphne s. str.. Eriosolena has been further upheld by Hallier f. (Med. Rijksherb. /;. 44,
1922, 30), DoMKE (Bibl. Bot. Ill, 1934, 130), Leandri (Rev. Intern. Bot. App. Agr. Trop. 29, 1949,
503, and Proc. 8th Pac. Sc. Congr. Manila 4, 1957, 581) and some others.
In 1915, however, H. Lecomte (Bull. Mus. Hist. Nat. Paris 21, p. 291-292) reversed his opinion on
the status of Eriosolena. After having examined all the species of Daphne contained in the Paris
Herbarium, he observed that they possessed, without exception, a very clear annular disk, sometimes
developed into a truly cupular disk surrounding the base of the ovary, for example in D. papyracea
Wall, ex Steud. (D. cannabina {non Lour.) Wall.)! He further advanced that the anatomical data
found by van Tieghem (Ann. Sc. Nat. Bot. VII, 17, 1893, 185) concerning the origin of periderm, the
presence or absence of internal phloem in the leaves, the existence or absence of crystals and their nature,
although they are interesting in themselves, cannot serve for solving the question about the rank (generic
or sectional) of Eriosolena. He did not recognize it as a separate genus to which I agree.

KEY to the species

I. Flowers in distinctly peduncled (2V2-6'/2 cm) heads enveloped by two caducous, involucral bracts.
Floral tube densely appressed-hairy outside. Disk distinct, membranous, cup-shaped. Ovary densely
hairy at the top 1. D. composita
1. Flowers in sessile or short peduncled (0-3 mm) heads without involucral bracts. Floral tube glabrous
outside. Disk obscure, ring-like. Ovary glabrous 2. D. luzonica

1. Daphne composita (L. /.) Gilg in E. & P. Pfl. Tjib. 2 (1923) 201; Burk. & Holtt. Gard. Bull.
Fam. 3, 6a (1894) 238; Koord. Exk. Fl. Jav. 2 S.S. 3 (1923) 70; Burk. & Hend. ibid. 3 (1925)
(1912) 657; Gamble, J. As. Soc. Beng. 75, ii (1912) 417; Moore, J. Bot. Suppl. 63 (1925) 89; Hochr.
258; K. & V. Bijdr. 13 (1914) 49; Koord. Fl. Candollea 2 (1925) 443, //W. va/-. wo/jrana Hochr.
38 Flora Malesiana [ser. I, vol. 6^

and montana f. macrophylla Hochr. Burk.

var. ; This specimen might have come from Java as the
Diet. (1935) 765;Corner, Ways. Trees (1940) 633, epithet indicated (c/. also Wikstr. Kongl. Vet.
f. 240; Back. Bekn. Fl. Java (em. ed.) 4A (1942) Acad. Handl. 1818, 297). It is distinctly D. com-
fam. 77, p. 5; Heyne, Nutt. PI. (1927) 1152.— posita. On the back of the sheet at the upper left
Scopolia composita L.f. Suppl. (1781) 409. corner is written in Thunberg's handwriting
D. javanica Thunb. Mus. Nat. Acad. Upsal. App. "e Ceilona. Thunberg", for this reason the species
1 1 (1806) 4, nometi; Fl. Jav. (1825) \i.—D. pendula has also been listed in Thunb. Fl. Ceilan. (1825) 5.
Sm. pi. Ic. ined. 2 (1790) 34, t. 34, nom. illegit.; However, the genus does not occur in Ceylon and
WiKSTR. Kongl. Vet. Acad. Handl. (1818) 296; the record of this specimen from Ceylon is ap-
Meisn. Denkschr. K. Bayer. Bot. Ges. Regensb. 3 parently an error.
(1841) 285; in DC. Prod. 14 (1857) 540, incl. p
montana Meisn. and y concolor Meisn.; Miq. Fl. 2. Daphne luzonica C.B. Rob. Bull. Torr. Bot. CI.
Ind. Bat. 1, 1 (1858) 877; KuRZ, For. Fl. Burm. 35 (1908) 72, 75; Merr. Philip. J. Sc. 5 (1910) Bot.
2(1877) 333; Hook./. Fl. Br. Ind. 5 (1886) 194; 366; En. Philip. 3 (1923) 132; Steen. Bull. Jard.
BoERL. Handl. 3 (1900) 111; Ridl. Fl. Mai. Pen. Bot. Btzg 13 (1934) 254.— Fig. 14a-b.
3 (1924) 144. Eriosolena montana Bl. Bijdr. Slender shrub up to 1 1/2 ni. Branchlets light
(1826) 651; Hassk. Cat. Hort. Bog. (1844) 92, brown to reddish-brown, glabrous. Leaves charta-
incl.a macrophylla Hassk. and /5 minor Hassk.; ceous to subcoriaceous, glabrous on both surfaces,
ZoLL. Nat. Geneesk. Arch. N.I. 1 (1844) 616; narrow elliptic-oblong, 8-9 by V/^-lYi cm; base
Syst. Verz. 2 (1854) 116.— Z). montana Meisn. cuneate to attenuate; apex acuminate; nerves 6-8
Denkschr. K. Bayer. Bot. Ges. Regensb. 3 (1841) pairs, distinct beneath, obscure above, curved as-
284. Eriosolena composita van Tiegh. Ann. Sc. cending; petiole almost absent, up to 3 mm.
Nat. Bot. VII, 17 (1893) 196; Bull. Soc. Bot. Fr. Inflorescences sessile or up to 3 mm
peduncled,
40 (1893) 68; Merr. Contr. Arn. Arb. 8 (1934) few- to many-flowered; pedicels 1 or 2 mm. Buds
111. Eriosolena pendula Bl. ex Lecomte, Not. very acute. Flowers c. 10 mm
long, articulated at
Syst. 3 (1914)101.— Fig. 14c-g. the base. Calyx lobes ovate, 21/2-3 mm
by 2 mm,
Shrub or small tree up to 10 m by 16 cm. very acute. Stamens subsessile, or with a very short
Leaves chartaceous to subcoriaceous, usually filament; anthers 1 '/2-2 mm
long. Disk ring-like.
brownish above and glaucous beneath when dry, Ovary ellipsoid-oblong, 3 by mm, glabrous;
1

elliptic-oblong to lanceolate, (3V2-)7-14(-20) by style obscure; stigma globose. Fruit (young) ovoid,
(l|^-)2-5 cm; base attenuate; apex acuminate; 7 by 4 mm.
nerves 9-14 pairs, distinct and elevated beneath, Distr. Malaysia: Philippines (N. Luzon:
visible or obscure above, sometimes distinct on Benguet Prov.).
both surfaces; petiole 3-5 mm. Inflorescences Ecol. In the mossy forest on the higher moun-
axillary, solitary or very rarely 2 inflorescences in tains, 2000-2500 m.
an axil (c/. King's coll. 6940) involucral bracts 2,
; Note. The leaves are similar to those of D.
ovate-oblong to oblong, 1-1 V2 cm long, minutely odora Thunb., from which it diff"ers by obviously
pubescent outside; peduncle 2'/2-6V2 cm, usually smaller flowers, absence of bracts below the umbel,
nodding, with several small linear bracts at the and absence of the typical yellow tomentum on
base, (4—)7-12-flowered. Flowers light-yellowish peduncle and pedicels. From D. kiusiana Miq. it
or white, fragrant, 10-15 mm long, sessile, densely diff"ers in the glabrous flowers, from the Formosan
covered with appressed, golden-yellowish or whit- D. arisanensis Hayata by the absence of floral
ish hairs outside. Calyx lobes convolute, 2 longer bracts, the very acute calyx lobes, shorter pedicels,
and 2 shorter, lanceolate or ovate-oblong, rarely longer anthers, and larger calyx tube.
oblong, 2-A by 1 mm. Stamens sessile or with short Though Merrill (1908, I.e.) suggested that
filaments; anthers linear, 1-1 V2 mrn long. Ovary there a closely allied form in Yunnan, I have not
is

ellipsoid, 1 '/4-2 mm, densely hairy; style c. IV2 succeeded in identifying it with a Chinese species.
mm; stigma globose. Fruits ellipsoid or ovoid, Excluded
10-15 by 5 mm, black (Backer s.n.) or red
(Backer 14479). Daphne decandra Bl. Bijdr. (1825) 650 is according
Distr. India, Burma (Southern Shan States, to Sleumer (Fl. Mai. I, 5, 1954, 91) = Casearia
Tenasserim and Tounghoo), Indo-China (Annam), velutina Bl. (Flacourt.) {cf. also Miq. Fl. Ind. Bat.
China (Yunnan), and Malaysia: Sumatra, Malay 1, 1, 1858, 709).
Peninsula, Borneo, and West Java. Eriosolena aflinis ZoLL. Syst. Verz. 2 (1854) 116.
Ecol. In rain-forests (900- 1000-) 1200-2000 m. I have seen two sheets, one from Paris, with
Use. The bark is used as binding material. Zollinger's own handwriting and the type num-
Vern. Kakapasan (also used for Phaleria), ber 3209, collected in Lombok, and one at Leyden.
kemanden, S, ki-salam, J; Sum. kulei manis rimbo.
: Miquel already had
(Fl. Ind. Bat. 1,1, 1858, 878)
Note. Daphne javanica Thunb. (1806, I.e.) is a suggested it and I have to
to represent a Rubiaceae
nomen nudum. There is a specimen under that thank Dr Bakhuizen van den Brink for the final
name bearing Thunberg's handwriting in his her- reduction to Antirrhoea hexasperma(RoxB.) Merr.
barium at Uppsala, kindly sent on loan from there. En. Philip. 3 (1923) 540 {Rubiaceae).
 Dec. 1960] Thymelaeaceae (Ding Hou) 39

7. GYRINOPS
Gaertn. Fruct. 2 (1791) 276, t.l40 f.6; Domke, Bibl. Bot. Ill (1934) 119, map 2;
Quis. Arn. Arb. 27 (1946) 404. Laclmolepis Miq. Ann. Mus. Bot. Lugd.-Bat.
J.

1 (1863) m.~Brachythalamus Gilg, Bot. Jahrb. 28 (1900) 146.— Aquilaria sect.

Brachythalamus Hall. /. Med. Rijksherb. n. 44 (1922) 19. Aquilaria sect.

Gyrinops Hall. /. I.e. ^Aquilaria sect. Lachnolepis Hall. /. I.e. Fig. 15. —
Trees or shrubs. Leaves spirally arranged, usually with distinctly parallel veins
joining the several intramarginal veins; margin thickened. Inflorescences terminal
or axillary, sessile or short-peduncled, in fascicles or a few flowers at the top of a
peduncle, with 2 or 3 small caducous bracts. Flowers 5-merous, pedicels articulated
at the base. Floral tube cupular to cylindric, puberulous outside, inside puberulous
with reflexed hairs arranged in lengthwise lines towards the upper part, some-
times glabror.s. Calyx lobes 5, spreading, puberulous on both surfaces. Petaloid
appendages 5, distinct, or united in a ring (G. moluccana and G. decipiens), inserted
at the throat of the tube, alternating with the calyx lobes, usually densely hairy.
Stamens 5, episepalous, free from the tube, inserted at the same level as the
petaloid appendages or slightly below, sessile or subsessile, Hnear, basifixed. Disk
shortly cup-shaped or ring-like, scale-like, or none. Ovary ellipsoid or obovoid,
pilose, sessile or short-stiped, 2-celled; style terminal, distinct or obscure; stigma
small. Fruits a loculicidal capsule, obovoid or ellipsoid, long-stiped and emerging
from the top or from the side of the floral tube. Seeds slightly ovoid, plano-convex,
usually with a caruncle-like appendage at the chalazal end.
Distr. Species 8, distributed in Ceylon (G. walla Gaertn.), and Malaysia (Lesser Sunda Islands,
Celebes, Moluccas, and New Guinea). Fig. 16.
The distribution pattern is very similar to that of Tiichadenia (see vol. 5, p. 39).
Ecol. In forests from the lowland up to 900 m.

KEY TO THE SPECIES

1. Floral tube tubular, 12-14 mm
long. Fruits emerging from the lateral slit of the floral tube.
2. Leaves narrow-lanceolate, 5-8 times as long as wide. Flowers in a raceme 1. G. moluccana
.

2. Leaves elliptic-oblong to ovate-lanceolate, c. 2'/2 times as long as wide. Flowers in an umbel.

3.Nerves prominent and spaced beneath, 16-20 pairs. Infructescences 12-14-flowered. Pedicels c.
mm. Petaloid appendages united in a ring. Fruits ovoid-oblong, c. 22 Vi
2 mm long; valves c.
3 mm
thick at the suture 2. G. decipiens
3. Nerves obsolete and close to each other, 25-35 pairs. Infructescences 2-3-flowered. Pedicels
3-5 mm. Petaloid appendages distinct and connected only at the base. Fruits pyriform, c. 17 1/2 mm
long. Valves ^/j-^/s mm thick at the suture 3. G. lederraannii
I. Floral tube cupular, 2-5 mm long. Fruits emerging from the top of the intact floral tube.
4. Leaves usually narrow-lanceolate, II/2-IO by V5-I cm. Petaloid appendages oblong, as long as
the stamens 4. G. salicifolia
4. Leaves elliptic-oblong or ovate-oblong, very rarely lanceolate, 6-15 by 1 '/2-5 cm. Petaloid ap-
pendages shorter than the stamens.
5. Pedicels more than twice as long as the floral tube. Petaloid appendages transverse-oblong.

5. G. caudata
5. Pedicels usually shorter than the floral tube. Petaloid appendages deltoid or slightly oblong.
6. Nerves and veins usually similar. Pistil usually shorter than the floral tube. Style none. Fruits
obovoid-oblong or ellipsoid, acuminate 6. G. versteegii
6. Nerves distinct and more prominent than the veins. Pistil usually longer than the floral tube.
Style distinct. Fruits pyriform, acute, bearing the persistent, curved style. 7. G. podocarpus

1. Gyrinops moluccana (Miq.) Baill. Adansonia Aquilaria moluccana Hall. /. Med. Rijksherb.
11 (1875) 326; Gilg in E. & P. Pfl. Fam. 3, 6a //. 44 (1922) 19.— Fig. 15a-d.
(1894) 225; Boerl. Handl. 3 (1900) 111; Quis. J. Shrub. Leaves chartaceous, glabrous, oblong-
Am. Arb. 27 (1946) 404.— Lachnolepis moluccana lanceolate, (8-) 18-24 by (Ii/3-)2-3 cm; base ob-
MiQ. Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 132.— tuse; apex acuminate; nerves 23-32 pairs, slightly
40 Flora Malesiana [ser. I, vol. 6^

Fig. 15. Gyrinops moluccciim (MiQ.) Baill. a. Habit, X 2/3, b. opened flower, one anther removed, X 3,
c. attachment of stamen, X 7, c'. stamen, X 7, d. fruit bulging out of floral tube, X 2. G. versteegii
(GiLG) DoMKE. e. Opened flower, x 5. G. podocarpiis (Gilg) Domke./. Opened flower, X 5, g. frontal
and lateral view of fruit protruding from floral tube, X 2. G. decipiens Ding Hou. h. Habit, x 2/3,
/. dehisced fruit bulging out of floral tube, x 2, /. opened floral tube of fig. i, X 3. G. salicifolia Ridl.
k. Habit, X 2/3. G. caudata (Gilg) Domke. /. Habit, X 2/3, m. opened flower with characteristic long
pedicel, X 5, n. young fruit protruding from floral tube, X 2 {a-d de Vriese & Teysmann s.n., e Ver-
STEEG 1381, /-g Pleyte 567, h-j Kjellberg 889, k Kanehira & Hatusima 12443, l-m Beccari PP 911).
Dec. 1960] Thymelaeaceae (Ding Hou) 41

curved and ascending, at c. 60° to the midrib, Ecol. In rain-forest, at 100 m.

distinct or visible beneath, indistinct above; veins Note. This species is closely related to G.
it parallel.Inflorescences axillary, sometimes on moluccana by tubular flowers with
the long
the branches, simple, rarely branched, 3-5-flower- petaloid appendages united into a ring and fruits
ed; peduncle almost none up to c. 10 mm, some- emerging through the lateral slit, but differs from
times 2 or 3 in an axil; pedicels c. 4 mm. Flowers it by the characters shown in the key.

long-tubular, c. 15 mm. Calyx lobes oblong, c. The leaves are similar to those of Aquilaria
3 mm long, erect, the apex slightly incurved. beccariana.
Petaloid appendages usually united behind the
stamens, with hairs almost as long as themselves.
3. Gyrinops ledermannii Domke, Notizbl. Berl.-
Stamens inserted slightly below the appendages,
Dahl. 11 (1932) 349.
sometimes the lower part of the anthers adnate
Shrub. Leaves subcoriaceous, glabrous except
to the tube. Pistil c. 4 mmlong. Ovary ovoid or
sparse hairs at the lower parts on both surfaces
ellipsoid, gradually narrowed into a distinct style,
and the midrib beneath, oblong- or ovate-lanceo-
pilose; stigma ovoid. Fruits emerging through the
late, sometimes obovate-lanceolate, 6V2-I2 by
lateral slit of the floral tube, ovoid, shortly stalked,
2/2-5 cm; base rather acute or shortly narrowed
sparsely pilose, ovoid, '/2 by
I cm, 1- or 2-seeded.
1
towards the petiole; apex acute or acuminate;
Seeds ovoid, with a short and thick appendage
nerves spreading, obsolete, close to each other,
at the base.
among which c. 25-35 stronger pairs, curved and
Distr. Malaysia: Moluccas (Buru: Kajeli and
ascending towards the apex. Infriictescences
Halmaheira), thrice collected. Also cultivated in
pseudo-lateral or terminal, subsessile, 2-3-flower-
Hort. Bog. (from seed of Buru).
ed; pedicels thin, 3-5 mm. Floral tube cylindric,
Note. One specimen collected from a cultivated
plant in Hort. Bog. has smaller leaves (8-141/2 by
indistinctly ribbed, 13 mm
long, /2 mrn in diam.
1

Calyx lobes ovate, '/2-2 by c. Yi mm, the outer

1
1 1/3-2
'/4 cm).
lobes acute and the inner ones obtuse, pubescent
outside, tomentose inside, and also with a tuft
of hairs at the top. Petaloid appendages ± rectan-
gular, c. ^/s by V2 mrn, obtuse, connected at the
base, villose. Stamens sessile, oblong, 1-1 '/4 by
1/5 mm. Fruits ± pyriform, c. I/4 by Vi cm (in-
cluding stipe 3 mmand acute and cuspidate apex
4 mm), short pilose, compressed, irregularly,
transversely ± rugose. Seeds 2 or by abortion,
I

c. 9 mm long (including an appendage c. 3 mm

long), woolly.
Distr. Malaysia: New Guinea (Sepik R.,
Station Mt Pfingst: Ledermann 7401).
Fig. 16. Distribution of Gyrinops Gaertn. Ecol. Slope in dense, virgin forest, at the foot
2, Gyrinops decipiens, nov. sp.
of the mountain, 0-200 m
altitude.
Note. The type specimen of this species is not
ab G. moliiccana foliis elliptico- vel sub-
DilTert
available and no additional material has been
obovato-oblonga, pedunculis brevibus validis
collected in that area. The description above is
apiceque incrassatis, floribus umbellatis, antheris
extracted from the original one. From Domke's
liberis, pericarpio late suturato. Typus G. Kjell-
detailed description, this is a distinct species
BERG 889, Bo, L.— Fig. 15h-j.
characterized by the long floral tubes, distinct
Small tree, c. 4 m. Leaves chartaceous, glabrous,
petaloid appendages and the leaves with many
rarely sparsely hairy beneath, shining on both
obscure nerves. It is closely allied to G. moluccana
surfaces when dry, elliptic- or slightly obovate-
and G. decipiens by the flower with a tubular
oblong, 14-17 by 5-7 cm; base narrowly cuneate;
floral tube and the fruit emerging from the lateral
apex shortly acuminate; nerves 16-20 pairs,
slit of it, but differs from both of them by the
slightly curved or obliquely spreading towards the
characters given in the key.
margin, elevated beneath, visible sometimes ob-
scure above. Infructesceuces terminal and axillary,
umbelliform, 12-14-flowered; peduncle very short 4. Gyrinops Ridl. Trans. Linn. Soc.
salicifolia
to 2 '4 cm, thick, accrescent, knob-like thickened Bot. II, Gvrinopsis salicifolia Quis.
9 (1916) 145.
at the top; pedicels c. 2 mm. Flowers long-tubular, J. Arn. Arb. 27 (1946) 407.— Fig. 15k.
c. 15 mm long. Floral tube almost glabrous inside. Slender shrub, c. 1 m. Branchlets light brown,
Calyx lobes oblong, 3-4 mm long. Petaloid ap- pubescent. Leaves sparsely pubescent on the mid-
pendages united behind the stamens with hairs as rib and sometimes on the nerves and veins
long as themselves. Stamens inserted slightly below beneath, lanceolate to linear-lanceolate, \y2-\O
the appendages. Fruits ovoid-oblong, c. 2'/4 by by Vs-l cm; base cuneate; apex acuminate and
1
1/4 cm, acuminate to the apex, suture surface c. pointed; nerves and veins similar and equally
3 mm wide. Seeds unknown. strong, slightly visible beneath, obscure above;
Distr. Malaysia: Central Celebes (Wavatoli, petiole c. '/2 rnm- Inflorescences terminal, sessile,
Palarahi). 3-5-flowered; pedicel c. 2 mm. Flowers pale
42 Flora Malesiana [ser. 1, vol. 6^

yellow, c. 3 mmlong. Floral tube c. 2V2 long, mm 1-3 mm. Flowers white, yellowish, light greenish,
cupular, pilose outside. Calyx lobes oblong, c. or yellowish-green. Floral tube cupular, c. 3V2 mm
1 mm long, puberulous. Petaloid appendages ob- long. Calyx lobes oblong, c. 1 mm
long. Petaloid
long, c. 1/2 mrn long, shortly hairy. Stamens sessi- appendages deltoid, about half as long as the
le, as long as the petaloid appendages. Disk anther, densely hairy. Stamens sessile, c. mm.%
obscure, ring-like. Pistil c. 2 mm
long, densely Disk scale-like. Pistil c. 2'/2 mm
long, densely
short-hairy. Ovary obovate, 1 mm
long; style puberulous, except the stigma. Ovary ovoid, c.
filiform, c. 1 mm; stigma obscure. Fruit unknown. 1 mmlong, narrowed towards the apex; style ab-
Distr. Malaysia: western New Guinea (Utakwa sent; stigma ovoid. Fruits yellow, slightly obovoid
and Nabire). or ellipsoid, 2'/^ by cm, shortly acuminate to the
1

Ecol. In fringing rain-forest, 300 m. apex, attenuate to the base. Seeds ovoid, plano-
convex, 9 by 6 mm, with a caruncle-like appendage

5. Gyrinops caudata (Gilg) Domke, Notizbl. Berl.-

at the base, c. 2 mm
thick.
Distr. Malaysia: Lesser Sunda Islands (Lom-
Dahl. 11 (1932) 349; Quis. J. Arn. Arb. 27 (1946)
bok, Sumbawa, Flores, and Sumba), NE. Celebes
404. Brachythalamus caiidatus Gilg, Bot. Jahrb.
(Minahassa), and West New Guinea (Alkmaar
28 (1900) 147; in E. & P. Pfl. Fam. Nachtr. 3
Bivouac and Somula).
(1903) 238.— Fig. 151-n.
Ecol. In forests, scattered, from the lowland up
Shrub or tree up to 17 m
by 36 cm, fide BW to 900 m.
6738. Branchlets greyish, whitish pubescent and
Vern. Ketemunan, Lombok, ruhu wama, Sumba,
glabrescent. Leaves chartaceous, glabrous, dull
seke, Flores.
beneath and shining above, elliptic-oblong, ovate-
Note. Closely related to G. podocarpus; more
oblong, rarely lanceolate, 6-13 by 1 '/2-4 cm; base
material is needed to verify whether it deserves
cuneate; apex up to 1 '/2 cm, acuminate; nerves
specific distinction.
and veins scarcely distinguishable, numerous,
parallel, visible beneath, obscure above; petiole c.
7. Gyrinops podocarpus (Gilg) Domke, Notizbl.
3 mm. Inflorescences axillary or terminal, 3-10-
Berl.-Dahl. 11 (1932) 349; Quis. J. Arn. Arb. 27
flowered, sessile and peduncled, peduncle up to
(1946) 404. Brachythalamus podocarpus Gilg,
8 mm. Flowers c. 5 mm
pedicelled. Floral tube
Bot. Jahrb. 28 (1900) 146; in E. &
P. Pfl. Fam.
cupular, 2 mmlong. Calyx lobes oblong, 1 mm Nachtr. 3 (1908) 238. Aquilaria podocarpus Hall.
long. Petaloid appendages transverse-oblong, c.
/. Med. Rijksherb. /;. 44 (1922) 19; Domke, Bibl.
V2 mm long. Stamens subsessile, slightly longer
Bot. Ill (1934) t. 2 f. 10.— G. ledermannii (non
than the appendages. Ovary ovoid, densely pilose;
Domke) Merr. & Perry, J. Arn. Arb. 22 (1941)
style very short; stigma capitate.
264.— Fig. 15f-g.
Distr. Malaysia: New Guinea (Sidai and Mt
Slender shrub, 14-2 m. Leaves chartaceous,
Arfak).
pubescent beneath especially on the nerves and
Ecol. Primary forest, 5-20 m{fide BW 6738).
veins, glabrescent or sometimes glabrous, glabrous
Vern. Niwawur, Amberbaken language.
above, elliptic-oblong, narrow-oblong, slightly
Note. This species is easily recognized by the
obovate-oblong, 10-15 by 3-5 cm, base cuneate,
pedicel usually c. 2 times as long as the floral tube.
apex up to 21/2 cm acuminate; nerves 25-40 pairs,
distinct beneath, obscure above; veins and veinlets
6. Gyrinops versteegii (Gilg) Domke, Notizbl. visible beneath, indistinct above, sometimes nerves
Berl.-Dahl. 11 (1932) 349; Quis. J. Arn. Arb. 27 and veins similar. Inflorescences terminal or
(1946) 404. G. walla (non Gaertn.) Koord. axillary, sessile or with a short peduncle up to
Minah. (1898)577;BoERL. Handl. 3 (1900) HI.— 6 mm, 2-6-flowered; pedicels 2-3 mm, pubescent.
Brachythalamus versteegii Gilg, Nova Guinea 8 Flowers white, 4-5 mmlong. Floral tube cupular.
(1910) 410. Aquilaria versteegii Hall. /. Med. Calyx lobes ovate, 1-1 '/2 mmlong. Petaloid ap-
Rijksherb. //. 44 (1922) 19.— C
sp. Hall./. I.e. 20. pendages deltoid or slightly oblong, !4-'/2 mm
—Fig. 15e. long, densely whitish hairy. Stamens sessile. Disk
Shrub up to 6 m, or tree up to 21 m by 65 cm shortly cup-shaped, crenate. Pistil shortly stipitate,
{fide bb 21394, Bo). Leaves chartaceous to sub- densely hairy except the stigma, c. 4'/2 mm
long.
coriaceous, pubescent, especially on the nerves Ovary oblanceolate; style distinct, c. 1 V2 mm;
and veins beneath, glabrescent or glabrous, dull stigma capitate. Fruit (young) green, pyriform,
and yellowish-brown beneath, shining and reddish- 15 by 6 mm, densely puberulous, acute to the apex
brown above, elliptic-oblong, ovate-oblong, or and crowned by the persistent, curved style, stipe-
obovate-oblong, 5-14 by 1 '/2-5 cm; base cuneate; like, cuneate towards the base.
apex up to 2 cm narrow-acuminate; nerves and Distr. Malaysia: West New Guinea (Ramoi,
veins similar, numerous, slightly oblique and par- Sorong, Monep, and Idenburg R.).
allel; petiole short, 3-5 mm.Inflorescences sessile, Ecol. In primary forests, from the lowland up
usually terminal, consisting of 6 to 8 flowers, to 750 m.
rarely axillary, or on the branchlets; pedicels Vern. Kokkoree, Asmat language.
Dec. 1960] Thymelaeaceae (Ding Hou) 43

8. DRAPETES
Banks ex Lamk, J. (1792) 188, t.lO f.la-d; Persoon, Syn.
Hist. Nat. Paris 1 1

(1805) 148; Gaertn. Fruct. 3 (1807) 199, t. 215; Endl. Gen. PI. (1837) 33; Suppl.
4 (1847) 61; Benth. F1. Austr. 6 (1873) 35; Benth. &
Hook./. Gen. PI. 3 (1880)
196; Domke, Bibl. Bot. Ill (1934) 138, map M.^Kelleria Endl. Gen. PL Suppl.
4 (1847) 61; Domke, Bibl. Bot. Ill (1934) 137, map \6.—Daphnobryon Meisn.
in DC. Prod. 14 (1857) 566. Drapetes sect. Daphnobryon Boerl. Handl. 3
(1900) 106.— Fig. 17.
Dwarf shrubs with creeping, radiant, or i
tufted and glabrous stems, sending
out fibrous roots from beneath, the lower part of the stems marked with prominent
scars of fallen leaves. Leaves subopposite or spiral, more or less appressed, sessile,
narrow-linear, convex on the dorsal side, plane or slightly concave on the ventral
side, with 5-9 striated longitudinal nerves; apex obtuse, with a tuft of hairs;
margins ciliate especially in the young ones. Flowers aggregated in small, sessile,
terminal heads almost entirely immersed in the leaves; pedicels short, articulated
at the apex, articulation hairy. Floral tube continuous, or circumsciss above the
ovary (S. American sp.), usually pilose outside, glabrous inside, caducous after
anthesis. Calyx lobes 4, slightly spreading. Petaloid appendages inserted at the
mouthof the tube, consisting of 1-2 episepalous scales (orO). Stamens 4, free from
the tube at the mouth, ahernate with the lobes; filaments slender, basifixed,
usually longer than the anthers; anthers oblong or sometimes subglobose. Pistil
sometimes abortive. Fertile pistil usually included, rarely exserted. Ovary ellipsoid
or slightly obovoid, 1 or hairy in the upper half or at the apex; style
-celled, pilose
linear, lateral, usually longerthan the ovary, caducous after anthesis; stigma
capitate and papillose when young. Fruit a small drupe with a thin-fleshy pericarp.
Seed similar in shape to the fruit, closely enveloped by the endocarp.
Distr. Species 4, three of them in S. America (Fuegia and Falkland Is.), New Zealand, Tasmania,
and Australia, one in Malaysia (New Guinea and North Borneo). Fig. 18.
Ecol. On dry grassy, or rocky places in the mountains, in the tropics almost confined to subalpine
and alpine heights.
Note. Drapetes was described in 1792 by Lamarck {I.e.) with only one species, D. muscosiis, known
from Fuegia and the Falkland Islands in S. America. Its perianth tube {i.e. floral tube) is circumsciss
above the ovary, whence the upper part falls away after anthesis; there are no scales at the throat. The
style is terminal. Since then some other species have been described from the Old World.
In 1847 Endlicher (I.e.) based a new genus Kelleria on D. diejfenbachii Hook, from New Zealand,
which should diff"er from Drapetes sens.str. by the continuous perianth tube, the presence of 4 appendages
alternating with the stamens at the throat of the perianth tube, and the capitate stigma.
Hooker/, (in Hook. J. Bot. Kew. Misc. 5, 1853, 300) maintained that all species of Drapetes resemble
each other very closely and form one natural genus without necessity to recognize Kelleria as a distinct
genus.
In 1857 Meisner {I.e.) proposed a new genus Daplinobryon for Drapetes ericoides Hook./, from Borneo
and D. tasmaniciis Hook. / from Tasmania. This should be characterized by 8 appendages alternating
in pairs with the stamens and by the distinctly lateral style.
Van Tieghem (Bull. Soc. Bot. Fr. 40, 1893, 72) stated that the anatomical characters of the twigs
of Drapetes, Kelleria, and Daphnobryon as a group closely agree and differ profoundly from those found
in all other Thymelaeaceae. Still, because of the diff"erent origin of the periderms, different texture of
the cells in the pith, and the quantity of the lignified peridesmic fibers in the meristele in the limbs of
their leaves, he found reason to maintain the three genera as distinct.
Domke {I.e.) maintained only Drapetes and Kelleria. Drapetes being characterized floral tube cylindric,
:

circumsciss. No appendages. Style terminal. Leaves ovate, semi-amplexicaulous; S. America. Kelleria:

floral tube short-cylindric or funnel-shaped, continuous. Appendages present. Style lateral. Leaves
needle-like, not semi-amplexicaulous, spiral or decussate; New Zealand to Borneo.
Bentham {I.e. 36) pointed out that whether the perianth is circumsciss above the ovary or not "is no
more than what is admitted as sectional only in Pimelea". As for the pairs of appendages, whether th
44 Flora Malesiana [ser. I, vol. 6^

are distinct or confluent into a single entire or notched one, is not constant, as has been pointed out by
Bentham (I.e. 36) and Gilg (Bot. Jahrb. 18, 1894, 514, f. 9A). I have also observed this variation in a
single specimen (Travers s.n., IV, 1909, New Zealand), and even in a single flov^'er! Because of the
great resemblance in habit and the number of stamens being the same as that of the perianth lobes,
Bentham (I.e. 36) and Bentham & Hooker/. (Gen. PI. 3, 1880, 196) reduced Kelleria and Daphuobryon
to Drapetes. I have followed them in this treatment and believe the first and last could be distinguished
in the rank of sections.

1. Drapetes ericoides Hook./. Ic. PI. (1852) t. 895; 484; Steen. Bull. Jard. Bot. Btzg 13 (1934) 254.—
Staff, Trans. Linn. Soc. II, 4 Bot. (1894) 221; Kelleria ericoides Domke, Bot. Jahrb. 62 (1929)
Boerl. Handl. 3 (1900) 111; Gibbs, J. Linn. Soc. 485; Steen. Bull. Jard. Bot. Btzg 13 (1934) 254.—
Bot. 42(1914) 132; Merr. En. Born. (1921) 417. Kelleria patula Merr. & Perry, J. Arn. Arb. 22
Daphtwbn'on ericoides Meisn. in DC. Prod. 14 (1941) 267.— Fig. 17.
(1857) 566; Miq. F1. Ind. Bat. 1, 1 (1858)881.— Stem reddish- or dark-brown, up to 50 cm;
Kelleria papuana Domke, Bot. Jahrb. 62 (1929) young branchlets villous, glabrescent. Leaves 3-5

Fig. 18. Localities of Drapetes ericoides Hook./.

by 2/3 mm.
Inflorescences (l-)4-9-flowered; pedi-
cels short, Vi mm. Flowers white, cylindric,
c.

slightly expanded towards the apex, 3-5 long,mm

12-costate. Calyx lobes ovate, obtuse, 1-1 1/2 mm
long. Stamens c. 1 mm
long; anthers as long as or
slightly shorter than the filaments. Petaloid ap-
pendages always 8, in 4 episepalous pairs. Pistil
2-3 mmlong, the abortive ones only c. 1 mm.
Fruits ellipsoid.
Distr. Malaysia: Borneo (Mt Kinabalu) and
New Guinea (Mt Carstensz, Lake Habbema, Mt
Wilhelmina, Mt Doorman, Hagen Range, Mt
Giluwe, Mt Wilhelm, Mt Albert Edward, Mt
Victoria, and Central Div.). Fig. 18.
Fig. 17. Drapetes ericoides Hook. /.. a. Habit, Ecol. Alpine plant, on dry grasslands, shallow
x 6, c. opened flower, pistil
nat. size, b. flower, soil over rocks on sheltered grasslands, plentiful
removed, x 6, d. pistil, x 6, e. stamens, X 13, on sandy banks of streams, and in cracks of
/ stem showing leaf-scars, x 6, g. leaf, x 6 granite, usually occurring from 3000 to over
(ROBBINS 315). 4000 m.

9. PIMELEA
Banks & Soland. ex Gaertn. Fruct. 1 (1788) 186, nom. gen. cons.; Meisn. in
DC. Prod. 14 (1857) 496; Benth. F1. Austr. 6 (1873) 1; Domke, Bibl. Bot. Ill
(1934) 138, map 18, non Banksia Forst. 1776.— Thecanthes Wikstr. Kongl. Vet.
Acad. Handl. Stockh. (1818) 269, 271.— Fig. 19-20.
Dec. 1960] Thymelaeaceae (Ding Hou) 45

Fig. 19. Pimelea cornucopiae Vahl. a. Habit, X 2/3, b. inflorescence, x 2, c. longitudinal section of
inflorescence, X flower, x 4, e. opened flower, x 4,/. stamen, x 13, g. seed, x 13.
2, d. P. concreta
F. V. M. h. Inflorescence, x 2, /. longitudinal section of inflorescence, x l,j. flower, x 4, k. pistil, X 4
(ci WoMERSLEY &
VAN RoYEN 5853, b-g WoMERSLEY 8767, h-k VAN Steenis 3304).

Herbs, shrubs or undershrubs (extra-Mai. spp.). Leaves opposite or decussate,

sometimes subopposite, or alternate. Inflorescences terminal or axillary (extra-
Mai, spp.), capitate, rarely spicate or solitary (extra-Mai. spp.); involucral bracts
partially united, or free (extra-Mai. spp.), 4, or in extra-Mai. spp. 4-6(-8) or more,
sometimes wanting. Flowers bisexual, sometimes unisexual by abortion or
dioecious (extra-Mai. spp.), at least in Mai. spp. developing centripetally. Floral
tube tubular, often silky villous outside, in Mai. spp. circumsciss above the ovary.
Calyx lobes 4, spreading or erect. Petaloid appendages 0, but the throat usually
thickened or rim-like folded. Stamens 2, inserted in the throat of the tube and
opposite the 2 outer calyx lobes. Disk consisting of 4, scale-like, fihform or
46 Flora Malesiana [ser. I, vol. 6^

club-shaped lobes or wanting (extra-Mai. spp.). Ovary ovoid; style long-filiform

and attached to one side of the ovary immediately below the apex. Fruit a small
drupe. Seed in shape similar to that of the fruit, albuminous.
Distr. Subendemic in Australia and New Zealand, comprising c. 80 spp., two of which extend
to Malaysia.
Ecol. In Malaysia in grassland and savannahs in the seasonal areas, mostly at low altitude, ascending
to 1000 m.
Note. The two Malaysian species belong to subg. Thecanthes (Wikstr.) Meisn. ex Gilg (in E. & P. Pfl.
Fam. 3, 6a, 1894, 243); in the text "Untergen." was erroneously printed as "Unterfam.", cf. also de
Dalla Torre & Harms p. 340).

KEY TO THE SPECIES

1. Flowers at anthesis long-exserted from the involucral bracts, the exserted parts usually longer than
the length of the involucral lobes. Calyx lobes usually spreading. Stamens exserted; filaments at
least twice as long as the anthers 1. P. concreta
1. Flowers at anthesis usually included or slightly exserted from the involucral bracts. Calyx lobes
usually erect. Stamens included; filaments almost as long as the anthers. 2. P. cornucopiae

1.Pimelea concreta F. v. M. Fragm. 5 (1865) 73; Annual, simple or branched, up to 50 cm,

Benth. F1. Austr. 6 (1873) 6. F. brevitiiba Fawc. glabrous throughout. Leaves membranous, nar-
in Forbes, (1885) 516; Gilg in E. & P.
Wand. rowly oblong or oblong-lanceolate, or obovate-
Pfl. Fam. 6a (1894) 243, f. 84, D, E.—P. sp.
3, oblong, IV^-liVi cm by 4-8 mm; base acute to
Dammerman, Nat. Tijd. N.I. 86 (1926) 45, f. 1.— obtuse; apex acuminate and minutely apiculate;
Fig. 19h-k, 20. nerves obscure, 4-6 pairs; petiole short, c. 1 mm.
Involucral bracts 4, united at the lower third or
lower half into an obconical cup, 7-15 long,mm
the free parts ovate or deltoid, rarely obovate,
4-10 by 6-8 mm, acute or acuminate, imbricate,
the outer pair overlapping the inner pair and
usually longer than wide rarely wider than long.
Inflorescences terminal, usually more than 50-
flowered; peduncles variable in length, very short
or up to 4V2(-7) cm; pedicels flat, usually dilated
at the base, very short to 1 1/2 mm, articulated at
the top. Flowers 10-15 mm
long, inserted on the
cup-shaped part of the involucre, centripetally
developing, white to rose. Floral tube cylindric.
Calyx lobes 4, imbricate, oblong or slightly
obovate-oblong, 2-21/2 rnm long. Disk scale-like,
small, c. 1/2 mm long. Stamens 2V2-3 V2 mm. Ovary
ovoid, 1-1 mm long; style exserted; stigma sub-
1/2

by 1 mm, short-stalked.
capitate. Fruits ovoid, 2-4
Distr. Australia (Northern Territory) and
Malaysia: Lesser Sunda Islands (Sumba and
Timor). Fig. 21.
Ecol. Grassland and sandy ground, from the
lowland up to 1000 m.
Vern. Tua leu, Timor.
Note. Some specimens collected in Sumba have
the involucral bracts deltoid and slightly wider
than long.

2. Pimelea cornucopiae Vahl, En. PI. 1 (1804) 305;

R. Br. Prod. (1810) 359; Meisn. in DC. Prod. 14
(1857) 496; F. v. M. Fragm. 7 (1869) 3; Benth.
Fl. Austr. 6 (1873) 6; F. v. M. Descr. Not. 2
(1885) 8; Bailey, Queensland Fl. pt 4 (1901)
1363. Thecanthes cornucopiae Wikstr. Kongl.
Vet. Acad. Handl. Stockh. (1818) 271.— P.
philippinensis C. B. Rob. Philip. J. Sc. 6 (1911)
Bot. 345; Merr. En. Philip. 3 (1923) 134.—
Fig. 20. Pimelea concreta F. v. M. Sumba Fig. 19a-g.
(Photogr. DE VOOGD). Annual up to 50 cm tall. Leaves membranous.
Dec. 1960] Thymelaeaceae (Ding Hou) 47

lanceolate, narrowly oblong, rarely obovate- Archipelago (Misima I. and Sudest I.), and
lanceolate, (li/4-)2-3'/2 cm by 1 '/2-6(-7i/2) mm; Malaysia: Philippines (extreme N of Luzon,
apex acuminate; base obtuse; nerves obscure, Cagayan Prov. near Sanchez Mira, once collected,
3-6 pairs; petiole very short, c. mm. Inflores- 1 B.S. 7410, US) and New Guinea (Western and
cences terminal, 5-40-flowered peduncles vari-
1 ;
Central Divisions). Fig. 21.
able in length, very short, or up to 5|/2 cm; pedi- Ecol. Common on savannah ridges and grass-
cels 1-5 mm, but not dilated at the base,
flat lands, from the lowland rarely up to 570 m.
articulated at the top. Involucral bracts 4, united
at the lower third or half into an obconical cup,
7-10(-15) mm
long, the free parts ovate and
acuminate, imbricate, the outer pair overlapping
the inner pair, longer than wide, 6-8 by 2'/2-4(-7)
mm. Flowers 7-10 mm
long, inserted on the cup-
shaped part of the involucre, white, usually in-
cluded in the involucral bracts, sometimes slightly
protruding beyond them. Calyx lobes 4, imbricate,
oblong, or slightly obovate, 1-2 long. Disk mm
small and obscure. Stamens 1-1 '/2 mm, slightly
shorter than the calyx lobes. Ovary ovoid, Yj 1 mm
long; style slightly protruding beyond the floral
tube; stigma small, globose. Fruits ellipsoid, 3 by
1
'/2 mm, slightly stiped. Fig. 21. Localities of Pimelea concreta F. v. M.
Distr. Australia (Queensland), D'Entrecas- (+) and Vahl. (•) in Malaysia;
P. coniucopiae
teaux Is. (Fergusson I.), New Britain, Louisiade both species also occur in Australia.

10. AMYXA
Cf. Airy Shaw, F1. Mai. I, 4 (1953) 363, f.5.—Fig. 22.
1. Amyxa pluricornis (Radlk.) Domke, Bibl. Bot. also take place in nature. The seed colours are
Ill (1934) 116; Airy Shaw, F1. Mai. I, 4 (1953) marked :green, softish, seed dark glossy
fruit
363.— Fig. 22. brown, funicle partly very fleshy, aril-like, white.
Shaw questioned whether the fruit of this small How these colours will be at complete maturity is
Bornean tree is dehiscent. Judging from fruiting unknown; they may change at the last moment.
material collected in Sept. 1958 by M. Jacobs Tropical fruits are often devoured by animals
(//. 5376) near Belaga, in the Third Division of before maturity. When mature the attractive seed
Sarawak, this seems indeed to be the case, though with its contrasting colours will dangle out of the
the spontaneous dehiscence of the almost mature dehisced capsule hanging on the thin basal part
fruit happened during the process of drying the of the funicle from the apex of the valve; it is
specimens. But the dehiscence of the valves ap- likely to be dispersed by animals.
pears so regular along distinct sutures that there Distr. Now also twice collected by Koster-
seems to be hardly any doubt that dehiscence will mans in West Kutai, East Borneo.

'b'~>

Fig. 22. Fruit of Amyxa pluricornis (Radlk.) Domke, nat. size. When collected it was still closed, but
dehisced during drying with 3 valves showing the dark seed and the funicle which is thread-like at the
base but thickened, pale, and fleshy in its apical part. In nature the seed is in the ripe state probably
dangling from the dehisced fruit (Jacobs 5376).
48 Flora Malesiana
Excluded and Doubtful
Gnidia oppositifolia {non L.) Blanco, F1. Filip. (1837) 299, and Gnidia? philippinica Meisn. in DC.
Prod. 14 (1857) 592 are according to Exell (F1. Mai. I, 4, 1954, 555) = Termimilia polyantha Pr.
(Combret.).

Hornera Jungh. Tijd. Nat. Gesch. Phys. 7 (1840) 314.

H. glomerata Jungh. I.e. 316. Japan. —
H. umbellata Jungh. I.e. 316. Japan. —
No type specimen of either of these species has been as yet located. Dr van Steenis and Dr Hatusima
could not clarify them from the description. They might not be natives of Japan (c/. Benth. &
{in /in.)
Hook. /. Gen. PI. 3, 1880, 188-189). Mr Airy Shaw (inlitt.) assumes them to be Lauraceous.

Passerina javanicaTnuNB. Fl. Jav. 2(1825) 19. The type specimen could not be found in Thunberg's
herbarium at Uppsala (c/. Backer e.s., Blumea 6, 1950, 358) and the description is entirely inadequate.
 STAPHYLEACEAE (B. L. van der Linden, Leyden)

This smallish family, containing five genera', is almost confined to the northern hemisphere in

both the Old and New World, overstepping the equator only in Ecuador and Peru in S. America
and in Malaysia, where it is found southward to Java and New Guinea.
Among the genera Huertea is confined to Peru and the West Indies (Cuba, Haiti). Tapiscia
and Eiiscaphis are East Asian. Staphylea is widely distributed in the subtropical and temperate
zone on the northern hemisphere. Turpinia is subtropical and tropical, it is the only genus
represented in Malaysia. It is remarkable that the distributional areas of the latter two genera
seem to exclude one another save for a slight overlapping in SE. Asia.
Ecologically the members of the family may be found from the tropical lowland up into the
mountains, in Malaysia up to the upper border of the montane zone at c. 2400 m, but in the
Sinohimalayan area they may ascend to c. 3000 m. Latitudinally the northern frontier is found
at c. 50" N (Central Germany, South Canada); in S. America the southern border is found in
Ecuador and Peru.
The taxonomic position of the family has a chequered history. In the 18th century its place
was designated in the affinity of Rhamnaceae. A. P. de Candolle (1825) and Meisner (1836)
referred the family as a tribe to the Celastraceae; Endlicher (1840) had Staphyleaceae as an
order next to the Celastraceae but placed Ochranthe as a separate family near the Hypericineae.
Reichenbach (1828) had arranged it near Sapindaceae and this position was accepted by
Bentham & Hooker, and up to the present day it is referred to Sapindales by Asa Gray,
Engler, Hutchinson, etc.
LiNDLEY (1835) kept it as a separate family in the Hypericineae together with Cunoniaceae.
Hallier /. has repeatedly stressed their close affinity to the Cunoniaceae (Uber Juliania, etc.
1908, 74, 116, 182; Arch. Neerl. Sc. Ex. Nat. Ill B, 1912, 164).
The affinity with the Cunoniaceae seems to be without much doubt and is shown by the fact
that Turpinias have twice been wrongly described as members of the Cunoniaceae, viz as
Ochranthe and Kaernbachia. The vegetative resemblance is large as both families have stipulate,
decussate, pinnate leaves and terminal inflorescences. Besides, the differential characters are
few, mainly 5 stamens in Staphyleaceae and diplostemonous flowers in Cunoniaceae, the 3-celled
ovary against a mostly 2-celled one, and more vague diff'erences in filaments and fruit. Leaves
of Staphyleaceae are herbaceous and the articulations of petiole and rachis shrink in the her-
barium; leaves of Cunoniaceae are generally coriaceous and the junctions do not shrink. In his
key Hutchinson diflferentiates Cunoniaceae and Staphyleaceae by having pendulous and
ascending ovules respectively, but the importance and constancy of this character seems doubtful.
(Krause, p. 272). The pollen structure of Staphyleaceae is it similar to that of several other
families, for example Celastraceae (Erdtman, 1952).
The wide separation of Staphyleaceae (near Sapindales) and Cunoniaceae (near Rosales) is
not only unsatisfactory from a taxonomical standpoint, but also anatomically. Dr. Metcalfe,
Kew, has made an anatomical investigation as far as available material permitted and his
conclusion is that there are marked anatomical differences between Turpinia and the Sapindaceae
and that both Staphyleaceae and Cunoniaceae show more mutual affinity than either of them
with the Sapindaceae. In his opinion "an anatomist could not disagree with a taxonomist who
wished to 'remove' the Staphyleaceae from the 'vicinity' of the Sapindaceae" (cf. Nova Guinea
n.s. 10, 1959, 212).
Plantgeographically Staphyleaceae represent a marked northern counterpart to the Cunonia-
ceae which is largely a southern hemisphere family. Such 'pairs' turn up repeatedly as our
studies of plant geography advance: Fagus and Nothofagus, Dillenia and Hibbertia, Ericaceae

(1) Gagnepain has described a sixth genus, Triscaphis, from Indo-China (Not. Syst. 13, 1948, 190;
Fl.G6n. I.-C. Suppl. 1, 1950, 999, fig. 128 3-8). According to the description this is almost certainly not
staphyleaceous through its exstipulate, spiral leaves and 3-merous flowers; it might be anacardiaceous.
According to Lemee it would be sapindaceous (Diet. Suppl. 10, 1959, 213)
50 Flora Malesiana [ser. I, vol. 6^

Fig. 1. Details of Malaysian species of Tiirpitm. —T. bonieensis (Merr. & Perry) van der Linden.
a. Habit, X V2> b. open nerves and venation, X Yi, c. flower, x 5, (^. (//7/o, in section, x 5,e. fruit with
cross-section, nat. size,/, stamen, x 10. T. montana (Bl.) Kurz. g. Leaf showing its closed nervation,
X 1/^, h-i. stamens, X 10. T. pomifera (Roxb.) T>C.j-k. Fruit and its section, nat. size. T. sphaerocarpa
Hassk. l-m. Fruit and its section, nat. size, n. embryo, x 4. T. ovalifolia Elmer, o. Fruit and its section,
nat. size. T. stipulacea van der Linden, p. Defoliated part of twig showing persistent stipules {a, c-d,
/Clemens 30070, b, e Clemens 28840, / Kerr 2521, j-k Lam 2855, o Ridley 15906, p SF 27516).

and Epacridaceae, etc. The meeting point of the areas of each pair and their overlapping margins
are almost always found in the vicinity of the tropical zone. It would seem that the birthplace of
must have been the tropics from where their ancestors have been branched off with a
these pairs
northward and southward directed distribution, respectively giving way to a subsequent devel-
opment (diversity) in antipodial centres. Other explanations for the phenomenon of these
Dec. 1960] Staphyleaceae (van der Linden) 51

antipodial pairs of affinity seem less likely, viz if the pairs had to be explained as random
parallel development which is extremely unlikely if we take the close taxonomical affinity of each
pair into consideration. And the mere suggestion that the phenomenon is due to 'coincidence'
seems not worthy to consider.
N'egetatively Turpinia show s a marked, structural resemblance with Sambiicus. both possessing
decussate, simply-pinnate, frequently herbaceous, toothed lea\es. with stipules and gland-like
(though stipules are not always well represented in the latter genus and some Turpinias
stipels
have simple leaves). Properly the sympetaly and inferior ovary of Sambiicus separates them
mainly in the reproductive section, but these two 'characters' are gradually losing their unique
value as essentials for natural affinity. Some time the relationship between Sambucus and the
Staphyleaceae should be scrutinized more carefully by modern methods. van Steenis.

1. TL RPIMA
Vent. Choix (1803) 31, t.31, noiii. gen. conserv. prop.; Merrill & Perry. J. Am.
Arb. 22 (1941) 543; J. Krause\ in E. & P. Pfl. Fam. ed. 2, 20b (1942) 306;
Bakh.. \. D. Linden & \an Steenis, Taxon 9 (1960) 57-58. Triceros Lour.
Fl. Coch. 1 (1790) 184, nom. gen. rejic. prop, non Griff. 1854; Spreng. Syst.
1 (1825) 947; Moritzi. Syst. Verz. (1846) 15; Baill. Hist. PI. 5 (1874) 342, 343,
excl. Euscaphis; Maza. Dice. Bot. Nom. Vulg. Cub. & Puerto-Riq. (1889) 15; c/.
also O.K. Rev. Gen. PI. 1 (1891) 148.— Z)a/n7?2/?e/eaRoxB. [Hon. Beng. 1814. 17,
nomen, "Dalrxmpelia'']V\. Corom. 3 (1820) 76, t.279. Ochranthe Lisdl. Bot. Res. 8
(1836) X.im.—Hasskarlia Meisn. PI. Vase. Gen. 2 (1843) 348; cf. Walp. Ann. 1
(1849) 753.— Maurocenia ^Triceros O.K. Rev. Gen. 1 (1891) 149.— Kaernhachia
Schltr. Bot. Jahrb. 52 (1914) 151, nom. illeg., non O. Kuntze, Rev. Gen. 1 (1891)
62. nom. illeg.; Engler. in E. & P. Nat. Pfl. Fam. ed. 2. 18a (1930) 241. fig. 140;
cf. HoOGL., VAN DER LiNDEN & Steen. Nova Guinea n.s. 10 (1959) 211-212.

Fig. L
E\ergreen trees or shrubs with terete, pithy twigs; pith terete. Indument. if
present, consisting of simple hairs. Stipules interpetiolar, 2 to each node, partly
inserted in the axil of the petioles, imbricate, entire, rarely 2-tipped, early caducous
(except in T. stipulacea). leaving a distinct annular scar. Leaves decussate, simple
or odd-pinnate; petiole sulcate. Articulations (base of petiole, nodes of rachis)
shrinking in the dry state. Two small glands (sometimes called stipels) on the
rachis near the insertion of the petiolules and also 2 near the base of the leaflet.
Leaflets 3-11 in compound leaves, herbaceous to subcoriaceous, mostly 2-3 times
as long as wide, penninerved. midrib prominent, apex acute to acuminate, base
obtuse to rounded, sometimes cuneately decurrent, margin glandularly serrate,
dentate or crenate. Panicles axillary, terminal or subterminal. mostly glabrous.
Bracts small. Pedicels with or without 1 or 2 minute bracteoles, not articulated
with the flower, apically widened into the short obconical receptacle. Flowers
regular, bisexual, 5-merous. Sepals persistent, free, imbricate, the outer ones
broader than the inner ones, ovate, broadly attached at the base, rounded at the
apex, fleshy, more or less ciliate at the margin. Petals free, imbricate, spathulate
or oblong-elliptic, or obovate, equal-sized, narrowly attached, membranous, more
or less ciliate at the margin, longer than the sepals, caducous. Stamens 5, epi-

(1) Kralse, I.e., has cited all subsequent different spellings oi Dairy mpelea, Ochranthe, Turpinia, etc.
and cited them erroneously as synonyms. I have omitted their mention, to avoid a complicated, un-

necessary formality.
52 Flora Malesiana [ser. I, vol. 6^

sepalous, equal; filaments linear, gradually widened to the base, (in Mai. spp.)
glabrous, inserted close to the disk, caducous; anthers rounded or ovate, with
spreading cell-bases, dorsifixed, dehiscing lengthwise, introrse, sometimes dis-
Disk annular, glabrous and crenate, fleshy. Ovary superior,
tinctly apiculate.
and styles closely appressed but not connate, the
(2-)3(-4)-celled, the three cells
combined stigmas 3-lobed. Ovules 1-cv) in each cell, anatropous, attached on the
dissepiment very close to the axis, in 2 vertical rows. Fruit up to 2i/) cm diam.,
with a more or less fleshy pericarp (in dry state rather hard), indehiscent, globular,
slightly 3-lobed, sometimes crowned by the horn-like conical style remains. Seeds
l-oo in each cell, of various shape, mostly roundish or reniform, or compressed,
yellow-brown to dark-brown when dry; hilum large; endosperm present; coty-
ledons flat, roundish.
Distr. Probably c. 30-40 spp., occurring in the continental-Asiatic and throughout the Malaysian
tropics (from Ceylon to S. Japan southward to Java and New Guinea) and in the Central and South
American tropics (West Indies, Columbia, Ecuador, Peru), in the West Pacific northward to S. China
on the continent but to Formosa, Riukius, and S. Japan (Kyushu: Yakushima, at 32" N) in the more
mild oceanic climate (Kuro Sjio current). Absent in the Pacific and the Australian continent. Though
Turpinia is not found in North America the trans-Pacific disjunction belongs obviously to a disrupted,
former North Pacific distribution, similarly to that of Stapliylea.
Some species have a wide distribution, for example T. montana (Bl.) Kurz, from Java and Sumatra
northward to Indo-China, Hongkong, and Yunnan, and T. pomifera (Roxb.) DC. in SE. Asia and
Malaysia (rare) where it has often been confused with T. sphaerocurpa Hassk., a species confined to
Malaysia. The other species in Malaysia are all of a more restricted distribution.
The greatest density of species is found in Borneo (among which 2 endemic species on Mt Kinabalu);
New Guinea, though situated at the southeastern end of the generic distribution, possesses 2
endemic species. In the Lesser Sunda Islands Turpinia is very rare which is probably due to the fact
that Turpinicis avoid countries subject to seasonal climatic conditions. Fig. 2.
Ecol. Obviously all species are constituents of tropical to subtropical rain-forest areas. They are
evergreen and frequently of small stature, belonging to the undergrowth or substage. But some species
may attain large size, up to 26 m, once noted 35 m with a free bole of 22 m, diam. 50 cm. They shun
the areas subject to periodical drought and none has been found in several of the Lesser Sunda Islands
or similar areas in South New Guinea.
As to altitude Turpinia prefers the montane zone, the highest stations in Malaysia being at c. 2400 m
in New Guinea and on Mt Kinabalu. In the Himalaya and Yunnan representatives may ascend to
3000 m (and in Yunnan even to 3300 m, cf. Krause).
Ridley does not cite any records for dispersal of Turpinia, but it is most likely that the fleshy, some-
times edible fruits will be devoured by birds and olher animals and that the very hard-shelled seeds will
be able to stand the passing of the intestinal duct and be dispersed endozoically.
As to pollination nothing is known definitely, but Fyson (Fl. Nilg. Puln. Hill-tops 1, 1915, 91) recorded
for T. nepalensis that the disk is producing an appreciable amount of honey attracting insect visitors.
The flowers lack, it is true, singly, attractive colours and size, but the inflorescences are large and some-
times very many-flowered and the honey scent may be a powerful agent for attracting insects, in a com-
parable way as is known from the equally unattractive flowers of Leea, as I have been informed by
Dr M. A. LiEFTiNCK. Lorzing repeatedly noted the flowers of T. sphaerucarpa to be sweet-scented.
M orph. Stipules. Merrill & Perry mention in their key {I.e. 545) that the stipules would be connate
in some Papuan species, but this rests on erroneous observation.
Characters which are suitable for specific distinction are not particularly showy, but they prove to
be very constant; some are unexpected. They are:
(1) Number of ovules per cell, for example T. montana has always 2, T. pentandra 4(-5), and T. brachy-
petala 8(-7). In T. sphaeroearpa the number is less constant.
(2) Size and shape of flower parts (calyx, petals, anthers).
(3) Size and shape of the fruits and the thickness of the pericarp.
(4) Stipules in T. montana are very small and hairy and with a bifid apex, in others they are glabrous
and entire. Unfortunately in many specimens and even of some species no stipules are available for
examination. T. slipulaeea possesses by exception obviously persistent stipules.
(5) The leaves provide reliable characters in only rather few species, for example in T. montana
and T. nitida, they are simple in T. simplieifolia.
(6) Hairiness occurs in a few species but the indument is very short and sparse, it is only typical in
T. grandis.
The place of insertion of the inflorescences has generally been accepted as terminal and I can confirm
Dec. 1960] Staphyleaceae (van der Linden) 53

this in a general way. However, there are specimens in which there are 2, or twin pairs of peduncles in
anthesis which are only seemingly terminal, but really separated by a small terminal bud visible between
the bases of these peduncles, if such a flowering branch starts growing after the anthesis the infructes-
cences become lateral and the central terminal bud prolongs the twig and produces a new flush with
some leaf pairs and the same mode of pseudo-apical reproduction is repeated.
Galls. A few zoocecidia have been described by Docters van Leeuwen (Zoocec. Neth. Ind. 1926,
331, fig.), viz a stem gall caused by a gall midge on T. moutana; two leaf galls caused by gall midges and
a witches' broom caused by a gall mite on T. sphaerocarpa.
Vern. The vernacular names have little use generally for the species identity; I have only taken up
those which I have found on sheets which I have identified, as in literature taxa have frequently been
combined under one name (with all the vernaculars) pertaining to diflferent species.
Uses. The timber is available only in small quantity and dimension and is of inferior quality with a
low durability; it is only exceptionally used. By its quick growth Tiirpinia has been tried out as a pioneer
for reaff"orestation on devastated mountain slopes in Central Java (Pangentjongan on Mt Galunggung
and Kl^dung on Mt Sindoro) which were promising, according to Koorders's report.
Wood Anat. den Berger, Meded. Proefstat. Boschw. 13 (1926) 95, Determinatietabel houtsoorten
van Malesie, Veenman, Wageningen (1949) 30; Desch, Mai. For. Rec. 15- (1954) 524 (hand lens);
Heimsch, Lilloa 8 (1942) 163; Metcalfe & Chalk, Anat. Die. 1 (1950) 445; Moll & Janssonius 2
(1908) 416.— The presence of the primitive features: many-barred scalariform perforation plates, fiber
tracheids, long-tailed rays, do not favour the position of the family near to the Sapindales. Janssonius
I.e. recognized the large difiTerences between the two families; Heimsch I.e. p. 182, 189 (erroneously cited

by Metcalfe & Chalk I.e. p. 446) suggested affinity to Celastrales but critical studies are necessary;
Desch I.e. p. 525 stated that too many genera of the Celastrales do not show affinity to Turpinia. —
C.A.R.-G.
Notes. Practical research with Turpinia has proved extremely difficult by the inadequacy of the
herbarium material which is often scrappy, in bud, or only bearing either flowers or fruit, bearing
witness that collectors do not sufficiently realize the manifest need of collecting complete material for
scientific purpose. It is lamentable that this has led to description of new species, and creation of types,
on an insuflScient basis: the types of T. parviflora, T. simplieifolia, T. unifoliata, and of T. laxiflora were
described without fruit. Admittedly 1 am committing myself in this revision, but the two new species
proposed here have such outstanding vegetative characters that I feel excused.
The venation of mature leaves is rather characteristic in herbarium materials. Leaflets have mostly a
rather wide open venation beneath with prominence in various degree in proportion to the order of the
veins (primary, secondary, tertiary, etc.). The upper surface of the leaflets shows mostly no prominence
of the small veinlets.
However, in a number of species the veinlets of different order are all about equally strongly prominent
on both surfaces giving the surface under the lens a reticulation approaching that of a tessellate structure.
This is characteristic for T. borneensis, T. nitida, T. grandis, T. stipulacea. and T. ovalifolia.
The measurements of the flowers given in this revision are based on those of mature ones, boiled from
herbarium material.
It is rather remarkable that several specimens of the Euphorbiacea Bischofia javaniea Bl. have in the

herbarium been identified as Turpinias though having spiral leaves and no annular, stipular scars.
As the vegetative parts offer only occasionally distinctive characters collectors should secure fertile
material, preferably with both flowers and fruit. Observations are desirable on the size and shape of the
stipules and the mode of regeneration of the flush and inflorescences.
The framing of a separate key on sterile material did not seem of much practical use and has
been omitted.
54 Flora Malesiana [ser. I, vol. 6^

KEY TO THE SPECIES

1. Leaves all and always simple, lanceolate-oblong, 10-15 by 3'/4-5'/2 cm, on both sides with prominent
venation. Sepals and petals c. 2Y2-3 mm long. Filaments up to 2 mm. Ovules 2 per cell.
1. T. simplicifolia
1. Leaves l-cvi-jugate, the upper ones under the inflorescence exceptionally 1-foliolate.

2. Stipules large, ovate-triangular, c. 1 '/s ciri long, persistent, stout (fig. Ip). Leaflets (7-)9, rather thin,

elliptic, c. 9 by 5 cm. Venation on both sides fine-reticuiate (tending to be tessellate) and prominent
(in the herb.). Petals c. 2'/2-3 mm long. Stamens c. 2'/2 rnm long; anthers ^j^ mm. Ovules 4 per cell.
2. T. stipulacea
2. Stipules early caducous. Not this combination of characters.
3. Upper internodes, stipules, petioles and petiolules, and inflorescence with a very short (puberulous)
but continuous indument (also in fruit). Leaflets 3-1, large, 15-23 by 10-16 cm, ovate to ovate-
oblong, sharply dentate; venation on both sides dense and prominent (tending to be tessellate).
Leaf-articulations hardly shrinking in the herb.. Fruits globular, without horns, c. 1-1 V2 cm diam.;
pericarp c. V2-I mrn diam. Seeds pale to pale brown, rather large, 10-18 per fruit. (Flowers un-
known) 3. T. grandis
3. Such indument absent. Not this combination of leaf-size and venation characters.

4. Flowers small (petals c. IV2-2V2 mm long; anthers c. Vi mm). (Mature fruits not crowned by
3+ distinct horns).
5. Pericarp (mostly much) thinner than 1 mm (fig. le). Stamens 1 '/2-2 mm.

6. Stipules 2-3 mm long, pubescent, shortly bifid at the apex. Ovules 2 per cell (exceptionally
3-4 in a few flowers of an inflorescence). Fruit c. 8 mm diam.; testa ^U-Vi mm diam.. Nerves
patent and+ straight, connected by regular loops of equal strength and prominence (fig. lo).
Leaflets mostly thin and herbaceous (mostly pale green) in the herb., rather narrow-elliptic
oblong, c. 5-10(-16) by 1 /2-5(-7) cm. Inflorescences delicate . . . . 4. T. montana
6. Stipules c. 4'/2-9 mm long, glabrous, entire. Ovules 4-8 per cell. Fruit c. 8-15 mm diam. Testa
mm
'/2-I diam.. Nerves curved-ascending, not connected by regular loops of equal strength
(fig. la, b). Leaflets chartaceous to thin-coriaceous. Inflorescences less delicate.

7. Venation dense and prominent with small areoles and tending to be tessellate on both surfaces.
mm
Anthers distinctly apiculate, c. '/2 long. Ovules 4 in each cell. Seeds large, c. 9 mm averagely
mm
testa ^/s-l diam.. Leaflets mostly distinctly widest below the middle, often towards the
base 5. T. borneensis
7. Venation rather lax with wider areoles and veins of various degree in prominence, the finest not
prominent, not tending to be tessellate. Stamens 1 Ya-I mm. Anthers not apiculate, c. Vi-Vi mm
long. Ovules 6-7-8 in each cell. Seeds c. 5 mm, testa 1/2 mm diam.. Leaflets widest in the
middle, wide and large, brown in the herb., shallow-dentate. Inflorescences 30-45 cm long.
6. T. laxiflora
5. Pericarp thicker than 1 mm (fig. 1 1). Stamens (generally) 2-3 mm. Anthers c. 0.5 mm, little to
distinctly apiculate. Ovules 6-8 per cell. Seeds c. 5 mm, testa c. V2-I mm diam.. Leaflets elliptic,
not very wide, rather coarse-dentate, upper surface often greyish or metallic, discolored.
7. T. sphaerocarpa
4. Flowers large (petals 3-4 mm, anthers at least % mm).
8. Venation of dried leaves distinctly prominent on both sides.
9. Pericarp thinner than 1 mm. Fruit ± globose, crowned by 3 distinct horns, c. 7-12 mm diam..
Leaflets 3, shining, large (10-20 by 4-81/2 cm), with 8-10 very patent, regular nerves prominent
on both sides, more or less straight in the lower half, upwards curving and distinctly arching,
coriaceous. Anthers 1 mm, apiculate. Pericarp c. V^-Y^ mm diam.. Ovules 4(-5) per cell. Testa
mm
3/4-1 diam 8. T. nitida
9. Pericarp 1-2 mm diam.. Fruit small, oval, globular, crowned (at least when young) with ap-
proximate style horns, c. 4-10 mm diam.. Leaflets 3, 4, or 5, mostly not shining on both sides,
7-20 by 4-10 cm; nerves 5-7, prominent below, spreading, ascending from the base, not dis-
tinctly looped, chartaceous. Anthers ^4 or 1-1 V4 mm, not (or slightly) apiculate. Ovules 5 or
6(-8) per cell. Testa 0.3-0.4 mm diam 9. T. ovalifolia
8. Venation in the herbarium not distinctly prominent at both sides.
10. Pericarp thick, at least 5 mm diam. (fig. Ij). Ovules (5-)6(-7) per cell. Full-grown fruit large.
Plant glabrous 10. T. pomifera
10. Pericarp 1-2 mm diam. Ovules 4(-5) or 8(-7) per cell. Full-grown fruit 10-15 mm diam.
Plants often puberulous.
11. Ovules 4(-5) per cell. Fruit with 3 distinct horns, at least when young. Leaflets IVi-^Vi
(-6I/2) cm wide 11. T. pentandra
11. Ovules 8(-7) per cell. Fruit without style horns. Leaflets (3'/2-)5V^-8 cm wide.
12. T. brachypetala
 Dec. 1960] Staphyleaceae (van der Linden) 55

1, Turpinia simplicifolia Merr. Philip. J. Sc. 27 Tree c. 18 m

high. Leaves 4-5 jugate; leaflets
(1925) 34. T. uiiifoliaui Merr. & Chun, Sunyat- ovate, glabrous, acuminate, rounded at the base,
senia 2 (1934) 37. margin denticulate, 6-10 by IVi-^Vi cm; petiole
Tree up to 4 m. Leaves simple, lanceolate- up to 8 cm petiolules of the lateral leaflets 3-7 mm.
;

oblong to oblong-elliptic, glabrous, acuminate, Stipules large, 17 by 10 mm, persistent, woody,

decurrent at the base, slightly dentate, 10-15 by glabrous. Panicles axillary up to 25 cm long;
3 1/2-6 cm; petiole up to 4 cm long, glabrous. pedicels 1-2 mm long. Sepals ovate-oblong, green,
Stipules small, glabrous, 2-3 long. Panicles mm suffused red down
to the middle of the segments,
axillary, up to 18 cm long. Sepals elliptic, rounded, 2-21/2 by 11/2-2 '4 mm. Petals obovate-oblong,
scarcely ciliate, c. 21/2 mm
long. Petals oblong- creamy-yellow, 21/2-3 mm
long. Stamens c. 2/2
elliptic, ciliate, 2'/2-3 mm
long. Stamens c. lYj mm mm long; filaments c. 2 mm; anthers round,
long; filaments 2 mm. Ovary 3-celled, each cell slightly apiculate, 2/5 mm long. Ovary 3-celled,
with 2 ovules. Fruit globular, c. 8 cm diam., with 4 ovules per cell. Fruit unknown.
without style-horns; pericarp very thin, 0.1-0.2 Distr. Malaysia: North Borneo (Mt Kinabalu,
mm thick. near Tibabah R.), once collected, 2100 m;//. June.
Distr. Hainan; in Malaysia: Philippines Notes. A remarkable species deviating from all
(Luzon), thrice collected. others by its obviously persistent, very large
Ecol. In forests. stipules. The large number of leaflets and the 4
Notes. The type (Loher 12992) bears an in- ovules per cell remind of the Papuan T. pentandra,
florescence and slightly immature foliage. In but this species has another venation and much
comparing the characters it seems not specifically larger anthers. It is possibly more allied to T.
different from that of T. unifoliata Merr. & Chun borneensis with which it shares the prominent
(S. P. Ko 52249, NY) from Hainan, which has venation and 4 ovules, but this species has nar-
more mature leaves but an inflorescence which is rower, more pointed leaf-shape, smaller flowers,
in bud; the sizes of the floral parts (stamens) are and finer, shorter inflorescences, besides lacking
obviously for this reason somewhat smaller than the unique stipules.
those of T. simplicifolia.
Another(fruiting) specimen from Hainan (F. C. 3. Turpinia grandis v. d. Linden, nov. sp.
How 73403, Sing) was preliminarily referred by Arbor 10-18 m alta. Internodia ultima, petioli et
Merrill to the same species, but from its venation infnictescentiae indumenta denso brevi muniti.
and leaf-shape I conclude this to represent T. Folia 1 3 foliolata, foliolis magnis grosse-serratis,
indochinensis Merr. This seems to be extremely 15-23 X 10-16 cm, venulis utrinque prominentibus.
close to T. formosana Nakai which has only 4 Infructescentiae 30 cm longae. Fructus globosus
ovules per cell, T. indochinensis having 8. It might ecorniculatus, I-IV2 cm diam., pericarpio c. V2-I
probably be better to reduce the latter as a variety mm crasso. Semina flavescenti-brunnea, 10-18 in
to T. formosana. fructo singula. T. Endert 4669, L.
:

Masamune (Fl. Kainant. 1943, 178) reduced Tree 10-18 m. Upper internodes, stipules,
T. unifoliata to T. formosana Nakai, J. Arn. Arb. petioles, and infructescences with a very short
5 (1924) 80. In my opinion this is a different (puberulous) but continuous indument. Leaves
species by possessing 4 ovules per cell, larger 1-3-folioiate; leaflets ovate to ovate-oblong,
flowers (petals 4 mm), lesser nerves (7-9 pairs), acuminate, rounded at the base, sharply dentate,
and hairy stamens. I have not seen the type 15-23 by 10-16 cm; venation on both sides dense
(Wilson 10130), but Tanaka & Shimada 13554 and prominent; petiole up to 7 cm long; petiolules
and 11180, which agree with the description of of the lateral leaflets V2-1 1/2 cm. Stipules small, c.
T. formosana. 5 mm long. Flowers unknown. Infructescences
Theleaves are astonishingly resembling those axillary, sub-terminal or terminal, up to 30 cm
of certain specimens of T. borneensis, but this long. Fruit globular, without horns, c. 1-1 1/2 cm
species has 5 leaflets, petals less than 2 long, mm diam.; pericarp 1/2-I mm
thick. Seeds pale to pale-
and 4 ovules per cell. brown, rather large, 10-18 in each fruit.
Its closest alliance is, as Merrill & Chun Distr. Malaysia: Central E. Borneo (W. Kutai:
already remarked,with other 1-foliolate
not Kiau; Mt Kemul; Long Petah), thrice collected.
species (such as T. arguta), but with T. nepalensis Ecol. On forested river-banks and in deep
from which it diff'ers by a more regular nervation, ravines, 450-1000 m.
finer inflorescence, 2 ovules per cell, and simple Note. Superficially somewhat resembling T.
leaves. laxiflora but with 1-3 leaflets. Besides, T. la.xiflora
has a loose, only slightly prominent venation,
2. Turpinia stipulacea v. d. Linden, now sp. — smaller fruits, and sepia-coloured seeds. It is
Fig. Ip. probably closest allied to T. borneensis. but it
Aspeciebus omnibus differt stipulis maximis from that species by
differs its indument, leaf-size,
persistentibus lignescentibus. Folia 4-5-jugata, and number of ovules.
6-10 X 21/2-.5 cm. Inflorescentiae robustae,
foliolis
usque ad 25 cm longae. Sepala 2-2V2 X 1Vi-2Va 4. Turpinia montana (Bl.) Kurz, J. As. Soc. Beng.
mm. Petala 2\2~3 ''"" longa. Stamina c. 2V2 mm 44, ii (1875) 182, a genuina; Koord. Exk. Fl. Java

longa, aniheris suborbicularibus subapiculatis. 2 (1912) 528; Atlas Baumarten 1 (1913) t. 92;
Ovula 4 in utroque loculo. T: SF 27516 Carr, Sing. Fl. Tjib. 2 (1923) 149; Hochr. Candollea 2 (1925)
56 Flora Malesiana [ser. I, vol. 6^

412, incl. f. arborescens et f. scandens; Merr. which, however, has never only 2 ovules per cell.

Contr. Arn. Arb. 8 (1934) 93; Backer, Brittonia 3 T. montana var. borneensis Merr. & Perry
(1938)81; Merr. J. Arn. Arb. 19(1938)42; Merr. I have found to represent a distinct species under
& Perry, J. Am. Arb. 22 (1941) 552; Back. Bekn. that epithet.
Fl. Java (em. ed.) 7 (1948) fam. 152, p. 1.— Whether the Chinese T. glaberrima Merr.
Zanthoxvlum {Xanthoxvlum) montatuim Bl. Bijdr. (Lingn. Sc. 1931, 312; ibid. 14, 1935, 27; J.
J. 7,

(1825) 248; MiQ. Fl. Ind. Bat. 1, 2 (1859) 670, cf. Arn. Arb. 22, 1941, 552) diff"ers from T. montana
Radlk. Sitz. Ber. K. Bay. Ak. Wiss. 16 (1886) is not certain as several numbers Merrill &
305-306 (1887). Zanthoxvlum seirulatum Bl. Perry mentioned to belong to it (Tsiang 2715,
Bijdr. (1825) 249; MiQ. I.e. —
Triceros cochin- How 71654, 73218, Chun & Tso 43918) represent
chinensis (non Lour.) Moritzi, Syst. Verz. (1846) in my opinion T. montana.
15. Maurocenia zollingeri O.K. Rev. Gen. PI. 1
(1891) 147, 150, Backer, Brittonia 3 (1938) 81.
cf.
—T. parva K. &
V. Bijdr. Booms. 9 (1903) 249;
Back. Schoolfl. (1911) 272; Craib, Fl. Siam. En.
1 (1926) 338.—Evodiaparviflora Craib, Kew Bull.

(1915) 425.—.? T. gracilis Nakai, J. Arn. Arb. 5

(1924) 79.— r. parviflora Craib, Fl. Siam. En. 1
(1931) 339.— Fig. Ig-i.
Shrub or tree up to 15 m. Leaves 3-7-foliolate,
below the inflorescence exceptionally with only 1
leaflet; leaflets elliptic to oblong, glabrous,
acuminate, decurrent at the base, dentate, 3-10
(-15) by \V2-IV2 cm; nerves straight, very
regularly connected by a looped intramarginal
vein; petiolules of the lateral leaflets '/2-3 cm.
Stipules c. 3 mm, puberulous, with a short but
sharply bifid apex. Panicles axillary, open, up to
c. 18 cm long. Sepals ovate, glabrous, ciliate,
1-114 by IV4-2 mm. Petals obovate, glabrous,
cihate, 1-1 V^ by 1 Vi-2 mm. Stamens c. 1 V2 mm Fig. 2. Demarcation of the Indo-Malaysian part
long; filaments 1-1 V4 mm; anthers V2-Vi mm, not of the distributional area of Turpinia, showing
or only very slightly apiculate. Ovary (2-)3(^)- also the number of species in each island or island
celled, each cell with 2 ovules. Fruit globular, group; above the hyphen the number of endemic
sometimes with 3 radial lines from the top, 8-10 species, below it other species.
mm diam.; pericarp thin, to V2 thick; moremm
than one seed developed; testa 0.2-0.3 thick. mm
Distr. Deccan Peninsula (Pulney Hills), Burma, 5. Turpinia borneensis (Merr. & Perry) v. d.
Siam, Indo-China, China (Yunnan, Kwantung, Linden, nov. comb. —
T. montana var. borneensis
Hainan), Hong Kong, in Malaysia: Sumatra to Merr. & Perry, J. Arn. Arb. 22 (1941) 553. T:
Central and W. Java (most frequent in W. Java). Clemens 29391 bis.— Fig. la-f.
Ecol. In primary montane rain-forest, 750-2300 Tree up to 15 m. Leaves 3-5(-7)-foliolate;
m. Fl. Oct.-Jan., fr. March-Aug. leaflets ovate-oblong to almost lanceolate, glab-
Vern. Puhun putag, ki bantjet leutik, S. rous, distinctly acuminate, rounded at the base
Notes. By its thin, very regularly looped nerves (in the Philippine specimens sometimes cuneately
and delicate inflorescence easily recognized. On decurrent), dentate, 7-17 by 2'/2-6 cm; primary
the mainland a few specimens have been found nerves and the reticulating veins distinctly pro-
with somewhat thicker, shorter leaves and more minent on both sides; petiolules of the lateral
contracted inflorescences (in drier cUmate?); leaflets c. I/4-2 cm long. Stipules c. 5 mm, glabrous.
venation and ovules are however exactly matching Panicles axillary, open, up to c. 20 cm. Sepals
those of T. montana. This form has been described ovate, ciliate, 2 by 1 1/2 mm. Petals obovate, ciliate,
as a distinct species T. parviflora Craib from Siam 1% by 1'4 mm. Stamens \Ya mm
long; filaments c.
(Kerr 2527, K) and is also found in Indo-China 1
1/2 mm; anthers 1/2 mm
long, apiculate. Ovary
(Pierre 907, L). The size of the flower parts is (2-)3-celled, each cell with 4 ovules. Fruit globular,
slightly smaller than that in T. montana, but this is sometimes with 3 radial lines from the top, 8-15
due to the fact that both specimens cited above mm diam.; pericarp thin, 0.1-0.2(-0.5) diam. mm
are in bud. There is no question that this species Seeds large brown; testa 3/^-1 mm
thick.
is ever scandent. The number of leaflets may Distr. Malaysia: Borneo and the Philippines.
occasionally be up to 1 1 (Garrett 792, Thailand). Ecol. In rain-forests, up to 1600 m.
A collection from the Pulney Hills (A. Sauliere Note. Merrill & Perry already remarked that
115, Bo, K, L) certainly represents T. montana; their T. montana var. borneensis might deserve
it has the same venation, glabrous stamens, specific rank. Its venation, prominent on both
constantly 2 ovules per cell, and 3-4 mm
long bifid sides, ovate-oblong leaf, its curved nerves, and
stipules. 4 ovules per cell (in T. montana always 2) remove it
There is a distinct affinity with T. nepalensis distinctly from T. montana. The latter is an ex-
Dec. 1960] Staphyleaceae (van der Linden) 57

clusively montane species distributed from SE. 1834): Merr. Contr. Arn. Arb. 8 (1934) 94.—
Asia along the Sumatran tracii to Java. Fig. ll-n.
Large shrub or tree, up to 20 m high, 55 cm
6. Turpinia laxiflora Ridl. J. Str. Br. R. As. Soc. diam. Leaves 3-5-7(-l l)-foliolate, brownish-red
n. 82 (1920) 179; Fl. Mai. Pen. (1922) 512;
1 when young; leaflets ovate, oblong or elliptic,
Merr. & Perry, J. Am. Arb. 22 (1941) 553.— glabrous, more or less acuminate, base obtuse or
T. pomifero var. spliaerocarpa (non Hassk.) obtusely rounded or decurrent, more or less
King; King, J. As. Soc. Beng. 65, ii (1896) 453, (sometimes not) dentate, 3-18 by 2-10 cm; petio-
pro parte. lules of the lateral leaflets 3-15 mm, glabrous or
Tree, up to 24(-35) m
by 50 cm. Leaves 3-7- puberulous. Stipules large, 8-9 long, glabrous. mm
foliolate; leaflets ovate-elliptic, glabrous, dentate, Panicles axillary, open, up to 30 cm long. Sepals
acuminate, rounded at the base, 8-24 by 31/2-12 ovate, ciliate, 1 1/2-2 by 1 1/2-21/2 mm.
oblong Petals
cm; petiolules of the lateral leaflets 4-20 mm, to elliptic, thin, more or by V2 mm.
less ciliate, 21/2 1

glabrous. Stipules 5-9 mm long, glabrous. Panicles Stamens 21/4-3 mm long; filaments I-IV2 mm;
axillary, lax, long, up to 45 cm, the ultimate anthers I/2 by V^ mm, generally more or less
branches minutely puberulous. Sepals ovate, apiculate. Ovary (2-)3(-4)-celled, each cell with
sparsely ciliate, n/2-2 by 1-2 mm. Petals oblong, 5-6(-8) ovules. Fruit globular, mostly with three
distinctly ciliate, 21/4-2 1/2 by 1-1 1/4 mm. Stamens grooves from the top, c. 1-1 1/2 cm diam.; pericarp
l%-2 mm long; filaments 1/2-1% mm; anthers
1 1-3 mmthick. Seeds brown, mostly several per
1/3- 1/2mm long, roundish, not apiculate. Ovary fruit; testa 1/2- 1 mm thick.
3-celled, each cell with (6-)7-8(-9) ovules. Fruit Distr. Malaysia: Sumatra, Malay Peninsula,
globular, in dry state mostly wrinkled, V^-l cm Borneo, Java, Lesser Sunda Islands (Bali, Flores),
diam.; pericarp (very) thin, 0.2(-0.9) mm diam. Celebes, Philippines, Moluccas (Ceram, Ambon).
Seeds several in each fruit; testa c. I/2 mm thick. Ecol. Frequent in rain-forests, on various soil
Distr. Malaysia: N. Sumatra (also Simalur I.) types,50-2000 m.
and Malay Peninsula (Perak: Larut). Vern. Ki bangkong, ki bantjet, ki keujeup, hi
Ecol. In rain-forests, at low altitudes, up to pongpasang, ki tjehai, tjawane sore, S; godong
150 m, once at 900 m. T. laxiflora has been noted bantjet, J; langkiang etem, rebung, tutuh sirawi,
by Ridley («. 6214, coll. a. 1894) to change its Simalur, ae, kua, Endeh.
foliage with new flush appearing simultaneously Notes. This species has often been confused
with young inflorescences. Young leaflets, when with T. pomifera but the diff"erences in size of the
they first appear, are narrow elliptic-lanceolate! flowers and thickness of the pericarp make it
As no later similar data have been reported it desirable to distinguish these taxa as two species;
seems premature to conclude that it is deciduous. their areas overlap. No attempt has been made to
Vern. Arilan-buluh, arilaii pajo item, arilan unravel all confusions in the references.
sitobiidlung, (awd)arilan uding, awd mdtdn nanas, In the Malay Peninsula and Sarawak T.
lahulung, matan-nanas pajo, Simalur, kaju long- sphaerocarpa is mostly represented by a slightly
gakan, k. songgak, Sumatra, k. rebiuig, Pasemah distinguishable form described as T. latifolia but
(Palembang). I cannot find any fitting characters to delimit it

Note. Closely allied to T. sphaerocarpa but against T. sphaerocarpa.

obviously distinct by a diff"erent pericarp, larger A specimen from Ceram (Eyma 2150) shows a
inflorescences, and generally wider leaflets which remarkable resemblance to the Papuan species but
are brown in dry state. it has small flowers (2/2 mm) and constantly 6
ovules per cell. In fruit these species can hardly be
7. Turpinia sphaerocarpa Hassk. Flora 25, ii distinguished.
(1842) Beibl. 1, p. 42; Miq. Fl. Ind. Bat. 1, 2 In Sumatra some specimens have been found
(1859) 593; Ridl. J. Mai. Br. R. As. Soc. 1 (1923) which are shortly but rather stiffly hairy on both
58; Baker/. J. Bot. 62 (1924) Suppl. 24; Merr. & leaves and inflorescences; they represent a minor
Perry, J. Arn. Arb. 22 (1941) 548.— Dairy mplea variety, var. pubescens, nov. var. No great impor- Vja.>f

javanica Hassk. PI. Jav. Rar. (1848) 439.-7". tance can be attached to this variety, although it is \^^ei
pomifera [non (Roxb.) DC. 1825]: Miq. Sum. occupying a geographically coherent area in
(1861) 201, 513; K. &
V. Bijdr. Booms. 9 (1903) Central Sumatra. A specimen from Mt Kinabalu *'^''^'

245; Back. Schoolfl. Java (1911) 272; Koord. (Clemens 34462) and one from Celebes (Palu: ^^^^
Exk. Fl. Java 2 (1912) 528; Atlas Baumart. 1 bb 28283) deviate in having only the inflorescence ^
(1913) t. 93; Fl. Tjib. 2 (1923) 149; Hochr. puberulous, besides having stamens 11/2-1% mm,
CandoUea 2 (1925) 412; Back. Bekn. Fl. Java anthers% mm.
(em. ed.) 7 (1948) fam. 152, p. 1; Heine, Pfl. Of Merrill's record of T. nepalensis in N.
Kinabalu (1953) 57. Maurocenia sphaerocarpa Sumatra I saw only one cited number (Bangham
O.K. Rev. Gen. PI. 1 (1891) 147, \49.—T. pomifera 981) which I refer to T. sphaerocarpa. This is a
var. sphaerocarpa King, J. As. Soc. Beng. 65, ii fruiting specimen; the leaf is typically discoloured
(1896) 453, pro nomen, partim. —
T. latifolia Wall. as in r. sphaerocarpa. The ripe fruits are distinctly
[Cat. 4939] ex Ridl. J. Str. Br. R. As. Soc. n. 82 too large for T. nepalensis and may well fit the
(1920) 178; Fl. Mai. Pen. 1 (1922) 512.-7. size and structure of T. sphaerocarpa, but they
sambucifolia Elmer, Leafl. 9 (1934) 3217.-7. have a distinctly 3-horned tip which is unusual
nepalensis (non [Wall. Cat. 4277] ex W. A. & for the latter species. The horns are not widely
58 Flora Malesiana [ser. I, vol. 6^

spaced as in T. cochinchinensis which species has similar toMalay Peninsula specimens of 'T.
also no discoloured leaves. trifoliata"but its flowers are smaller and there are
only 4 ovules per cell; it is unfortunately only
8.Turpinia nitida Merr. & Perry, J. Arn. Arb. 22 described after flowering material which in all its
(1941) 549. characters is extremely close to T. cochinchinensis
A recumbent treelet, m high. Leaves
41/2-6 (Lour.) Merr., all species having 4 ovules per
oblong to elliptic,
1-5-foliolate; leaflets coriaceous, cell. The latter species has much larger fruits
glabrous, denticulate, acuminate, rounded at the (1-2 cm diam.) with spaced horns; T. nepalensis
base, upper surface shining, 11-25 by 5'/2-10!/2 has small fruits c. 1/2 cm diam., also provided with
cm; nerves prominent, arcuately ascending, then small remains of the style bases.
parallel with the margin; petiole up to 6 cm long,
glabrous; petiolules of the lateral leaflets 12-20 10. Turpinia pomifera (RoxB.) DC. Prod. 2 (1825)
mm. Stipules small, glabrous, c. 5 long. mm 3; Wall. Cat. 4267; Kurz, J. As. Soc. Beng. 44,
Panicles axillary, subterminal (or terminal), up to ii(1875) 182, excl. svn.; Hayata, Ic. PI. Form.
1

22 cm long. Pedicels 2-21/2 mm. Sepals oblong- (1911) 160; ?KiNG, J. As. Soc. Beng. 65, ii (1896)
elliptic, ciliate, 2-4 by 1 1/2-21/2 mm. Petals oblong, 453, pro nomen, excl. var. sphaerocarpa; Kanjilal
ciliate, 4 by 2 mm. Stamens 21/2-31/2 mm long; & Das, Flora of Assam 1, 2 (1937) 309; Merr. &
filaments c. 3 mm; anthers by 1 mm, dis-
c. 1 Perry, J. Arn. Arb. 22 (1941) 546; Holthuis &
tinctly apiculate. Ovary (2-)3-celled, each cell with Lam, Blumea 5 (1942) 205; Gagn. Fl. Gen. I.-C.
4(-5) ovules. Fruit globular, with 3 distinct, some- Suppl. 1 (1950) 993. Dalrympelea pomifera
times closely appressed horns; pericarp i/4-% mm RoxB. Beng. (1814) 17] PI. Corom. 3
[Hort.
thick. Seeds large, two or more in each fruit; (1819) 76, 279 {Dalrvmpelia).—Maurocema
t.

testa 3^-1 mm thick. pomifera O.K. Rev. Gen. PI. 1 (1891) 147, 149.—
Distr. Malaysia: North Borneo (Mt Kinabalu). Turpinia sp. Merr. For. Bur. Bull. (Philip.) 1
Ecol. Fairly frequent on wet forest ridges, (1903) 34.— Fig. Ij-k.
900-1800 m. Tree, c. 10-20 m. Leaves 3-5-7(-9)-folioIate;
Note. In size of the leaflets resembling T. leaflets elliptic-oblong, glabrous, distinctly acu-
grandis (also occurring on Mt Kinabalu), but minate, decurrent (sometimes a little rounded) at
difl'ering in the shape of the fruit, the venation of the base, dentate, 12-25 by 6-10 cm; petiolules of
the leaflets, and the absence of an indument. the lateral leaflets 3-10 mm. Stipules triangular,
4-5 mm. Panicles terminal, subterminal or axillary,
9. Turpinia ovalifolia Elmer, Leafl. Philip. Bot. 2 10-30 cm long, sometimes slightly puberulous.
(1908) 490; Merr. &
Perry, J. Am. Arb. 22 Sepals ovate, unequal, fleshy, scarcely ciliate,
(1941) 544.-7. trifoliata Ridley, J. Str. Br. R. 2-31/2 by 11/4-2 mm. Petals oblong, thin, scarcely
As. Soc. n. 82 (1920) 178; Fl. Mai. Pen. 1 (1922) ciliate, 3-31/2 by 1-1 1/2 mm. Stamens 3 long; mm
511.— r. hicida Nakai, J. Arn. Arb. 5 (1924) 80.— filaments c. 21/2 mm; anthers ^/j-l long, mm
? T. pachyphylla Merr. Philip. J. Sc. 27 (1925) 33, mostly subapiculate. Ovary 3-celled, each cell with
ex descr. —Fig. lo. (5-)6 ovules. Fruit globular, mostly with 3 grooves
Tree c. 8 m
or sometimes higher. Leaves 3-5 from the top, up to 25 cm (perhaps to 37 mm);
(-7)-foliolate; leaflets rounded to elliptic, glabrous, pericarp very thick, to c. 5 mm
diam. Seeds small
acuminate, sometimes with an abrupt acute point, and brown, glossy.
rounded at the base, margin crenate with fine, Distr. Continental Asia from the East
whitish, callous points, (2i/2-)4-20 by (l-)4-10 Himalaya eastward; in Malaysia rare: Sumatra,
cm; petiolules of the lateral leaflets 4-15 mm, Java, Central Celebes (Nuha Distr.: Kjellberg
glabrous. Stipules small, 21/2-3 mm
long, glabrous. 2303) and Minahassa, Talaud Is, and Philippines
Panicles mostly short, <:. 10-15 cm long (in fruit (Mindanao: Ahern 354; Luzon: Camarines Sur:
to 30 cm), rather dense. Sepals ovate, ciliate, BS 76375).
2-41/2 by 1 1/2-3 mm. Petals obovate-oblong, thin, Ecol. In forests, 0-2100 m; fl. March,//-. Sept.-
ciliate, 31/2 by 2 mm. Stamens 2/2 long; mm Oct.
filaments I-IV2 mm; anthers -^ or 1-1 14 long, mm Vern. Ki bangkong, ki renggang, S, lampasia,
not (or slightly) apiculate. Ovary (2-)3-celled, Minahasa.
each cell with 5 or 6(-8) ovules. Fruit with 3 short Notes. T. pomifera which is a common species
horns on top, 4-12 mm
diam.; pericarp thick on the Asiatic continent is rare in Malaysia. It is
1-4 mm; testa 0.3-0.4 mm
thick. characterized by a very thick pericarp closely
Distr. Malaysia: Malay Peninsula (Nyalas, enveloping the seeds, and besides it differs from
Selangor), Philippines (Luzon, Palawan). T. sphaerocarpa, with which it has frequently been
Notes. I cannot well separate T. ovalifolia and confused in Malaysia, by larger flowers and much
T. trifoliata; the type of the former is extremely larger anthers. In sterile state it is impossible to
poor; the flowers Elmer described are not present tell them apart.
in any of the isotypes I have had on loan for study. The only specimen of the Malay Peninsula
Though there is a slight difference in the size of the which might represent true T. pomifera is King's
anthers I cannot discriminate the Malayan spec- coll. 4243 (Sing), but the material is too inade-
imens from those of the Philippines. The species quate to be conclusive.
can be expected to occur in Borneo. In the W. Deccan (Ghats and Nilgiris) and
T. robusta Craib from Siam is habitually very Ceylon the records of T. pomifera have been
Dec. 1960] Staphyleaceae (van der Linden) 59

straightened out by Gamble (Kew Bull. 1916, 135; constantly differing in the number of the ovules,
FI. Madras pt 2, 1918, 241); it appears that two the shape of the fruit (horns), and width of the
different species are concerned, viz T. nepaleusis leaflets.
and a new species, T. molabarica Gamble, which
is also the single one occurring in Ceylon, charac- 12. Turpinia brachypetala (Schltr) v. d. Linden,
terized i.a. by hairy filaments. Nova Guinea 10 (1959) 212.
n.s. Kaernbachia
brachypetala Schltr, Bot. Jahrb. 52 (1914) 153,

11. Turpinia pentandra (Schltr) v. d. Linden,

f. 5 A-G; Engler, in E. & P. Pfl. Fam. ed. 2,

Nova Guinea n.s. 10 (1959) 212. Kaenihachia

18a (1930) 241, f. 140 A-G.— T. versteeghii Merr.
pentandra Schltr, Bot. Jahrb. 52 (1914) 151, f. 5
& Perry, J. Arn. Arb. 22 (1941) 554.
Tree, 4-26 m; bark brownish grey, flaky.
H-N; Engler, in E. & P. Pfl. Fam. ed. 2, 18a
Leaves (l-)2-3-jugate (rarely paripinnate by ab-
(1930) 241. T. papuana Merr. & Perry. J. Arn.
sence of the terminal leaflet); leaflets ovate-oblong,
Arb. 22 (1941) 554.—.^ T. papuana Harms in E. &
serrate, glabrous, obtuse, more or less acuminate,
P. Pfl. Fam. ed. 2, 20b (1942) 312, descr. germ.,
obtuse to rounded at the base, (6i/2-)8-17 by
sine typ., homon. illeg.
41/2-81/2 cm; petiole 3/2-9 cm long, very short-
Tree, 15-20(-25) m. Leaves (l-)2-3-jugate
hairy or glabrous; petiolules of the lateral leaflets
(rarely paripinnate by absence of the terminal
leaflet); leaflets oblong or ovate-oblong, glabrous,
4-9 mmlong; young foliage glossy brownish
green. Stipules triangular, glabrous or sparsely
shallowly serrate, distinctly obtusely acuminate,
hairy outside, 4-7 by 3-6 mm. Panicles axillary,
obtuse to rounded at the base, 6-12(-15) by
2 1/2-4 14 (-6 '/2) cm; petiole 2^i/2(-7) cm, very
open, up to 30 cm long. Pedicels 1/2-3 long. mm
Sepals ovate, ciliate, white, 3/2-4 by 2-2^4 mm.
short-hairy to glabrous; petiolules of the lateral
Petals obovate-oblong, spathulate, thin, sparsely
leaflets 3-5(-8) mm. Stipules triangular, glabrous
ciliate, white, 31/2-4/2 by I/2-IH mm. Stamens
or sparsely hairy outside, 5-7 by 3-4 mm. Panicles
axillary, open, up to 25 cm long. Pedicels Yj-'i mm 31/2-4/2 mm
long; filaments 3-4 mm; anthers %-

long. Sepals broad-obovate, ciliate, 3-3 '/2 by

0.9 mm long, distinctly apiculate. Ovary 3-celled,
each with (7-)8 ovules. Fruit globular, some-
cell
1 1/2-2 mm. Petals obovate, rounded or obtuse at
times with 3 radial lines from the top, 1-1 1/2 cm
the apex, thin, ciliate, 4-41/2 by l%-2 mm.
Stamens ^Vz-^Va mm long; filaments 21/2-3 mm;
diam.; pericarp c. 2 mm
thick. Seeds mostly 3
or more.
anthers -)4-l mm long, apiculate. Ovary 3-celled,
Distr. Malaysia: New Guinea.
each cell with 4(-5) ovules. Fruit globular, mostly
Ecol. In primary rain-forests, locally frequent,
with 3 distinct horns (remains of the styles), c.
300-2000 m.
11/2 cm diam.; pericarp c. 2 mm thick. Seeds
Note. We have at present rather numerous
mostly l(-2).
collections from various places all over New
Distr. Malaysia: New Guinea.
Guinea from both T. pentandra and T. brachypetala.
Ecol. In primary rain-forests, locally frequent, They are doubtless closely allied, but there appear
1200-2500 m. to be no intermediates. If only fruiting material is
Vern. Naun, Waria (Mt Hagen). available T. brachypetala can hardly be dis-
Note. Very close to T. brachypetala, but tinguished from T. sphaerocarpa Hassk.
 CAPPARIDACEAE (M. Jacobs, Leyden)

Herbs or shrubs, often climbing, rarely trees. Indument, if present, consisting of

simple (unicellular or multicellular) hairs (sometimes capitate-glandular), stellate
hairs, or appendages (Cleome). Leaves spirally arranged, petioled, simple,
palmately dissected, or compound, entire, penninerved, in Sfixis pellucid-
dotted. Stipules thorny, or minute, or wanting. Inflorescences racemose, terminal
or lateral, rarely the flowers axillary, or sometimes serial. Bracts, if present, small
and caducous, rarely with stipular bracteoles. Flowers bisexual but sometimes
the gynoecium reduced (in extra-Mai. spp. staminodes may occur), actino-
morphic with a tendency towards zygomorphism, especially in the receptacle
and in the position of the petals, mostly in bud until anthesis, but in Crateva
opening at a very early stage. Sepals 4, either equal or in 2 whorls of 2 and then
the outer pair enveloping the bud and slightly different from the inner pair, or
(in Stixis) in 2 equal whorls of 3, free, rarely the outer pair connate in bud. Petals
4 or (in Stixis) absent, free, often unguiculate, equal, or sometimes 2 of the
petals slightly asymmetrical and adjoining at the base. Receptacle more or less
conical, often with peculiar protrusions, such as (in Malaysia) a small anterior
disk in Capparis, or a long anterior tubular gland in Cadaba, or a ring in Crateva.
Stamens (4^)6 to cv), in Malaysian genera all fertile, either free or their base

connate with the gynophore in a very short to very long androgynophore;

anthers dorsifixed, often near the base, introrse, 2-locular, dehiscent lengthwise,
connective inconspicuous. Ovary generally on a long gynophore, to sessile, ovoid
to cylindrical, with a small, simple, sessile stigma, 2-6-carpellate, in Malaysia
1-3-locular. Ovules mostly oo, on parietal, rarely axillary placentas, campy-
lotropous, with 2 integuments, a third, thin, innermost seed-coat of tracheal tissue
being present at least in certain examined cases. Fruit a capsule, or a berry with
tough exocarp. Seeds cv), rarely 1 (Stixis), mostly coiled-reniform, poor in endo-
sperm; embryo curved, horseshoe-shaped or coiled, the cotyledons mostly
involute or plicate, or coiled, or one partly enveloping the other; testa in seeds of
dry fruits mostly sculptured and sometimes with an elaiosome, otherwise smooth.
Distribution. About 45 genera and approximately 700 spp. in tropica! and subtropical regions.
The largest genera are Capparis (over 250 spp.), Cleome (over 150 spp.), Maenia (over 50 spp.), and
Boscia (over 35 spp.). Capparis and Cleome are both best-represented in the neotropics; another large
centre is Africa. Monotypic genera are extremely numerous in this family of the 45 genera acknowledged
:

by Pax & Hoffmann 20 (44 %) are monotypic. Even if some might be reduced in future studies the
figure will remain remarkably high. SE. Asia and Australia are the poorest in monotypic genera, either
area having 3; it is not certain that all these six monotypic genera can be upheld. These six monotypic
genera are also endemic and only SE. Asia has one more endemic genus. On account of these facts,
I presume that the family occupied these parts of the world later than Africa and the neotropics.
Of the 5 Malaysian genera, Crateva, Capparis, and Cleome are pantropic. Cadaba extends from Africa
to Australia, and only Stixis is limited to the SE. Asian rain-forest area.
Ecology. Most of the Capparidaceae are xerophiloiis, and it is the world's drier regions that are
richest in representatives. In Malaysia Cadaba and partly Capparis show such a xerophilous distribution.
It is notable how many species of Capparis avoid the everwet Sunda shelf land and are distributed around

it. This tendency also holds for Cadaba, but not for Crateva and Stixis.
Most Capparidaceae are heliophilous. Next to the above-mentioned tendency to xerophily. adaptations
to other conditions occur. A few Capparis spp. prefer more or less light forest, as do the species of 5n'.v/5.
Crateva seeks the proximity of rivers, both in the everwet and monsoon areas. Cleome seems in
Malaysia anthropochorous. Most Capparidaceae are lowland plants; a few ascend as high as 1500 m
or slightly higher.
Pollination. The type of flower with numerous radiating stamens, as it occurs in Capparis, is,
flower-biologically, likely to represent the basic condition in this family. The production of nectar seems
62 Flora Malesiana [ser. I, vol. 6^

to represent a later development. Radlkofer, who gave an extensive description of the flower of Capparis
micracantha, found that the sweet nectar excreted by the disk can only be reached through a very small
slit nearly halfway up between the upper petals (Sitz. Ber. Bay. Ak. Wiss. 14, 1884, 1 1 1-1 16). Fig. 17a, d.

This, the great distance between the anthers and the nectar, and the difficulty for insects to land on the
flower, especially at nocturnal anthesis, renders this species apt to be pollinated by Sphingids, as has been
repeatedly stated to occur in Capparis. In C micracantha and C pubiflora the basal median part of the
upper petals has a red, pink, or yellow honey-guide. The nocturnal flowers of C. spinosa and C. lucida
are sweet-scented and produce nectar. Cadaba capparoides, with its long, tubular, nectar-hiding gland
and dark, versatile anthers, possesses indeed the most perfect adaptations in this field. Fig. 26, 27.
Since the Rhoeadales apparently lack the potentiality of producing a style, a gynophore became
necessary to elevate the ovary into the sexual zone. The peculiar position of the petals, which are all
pointing upwards (in Cadaba, in Cleome p.p., in Crateva there being a tendency to it) seems useful to
form a showier beacon to the nocturnal butterflies.
Next to the above described sphingophilous type (as here found in Cleome gyiiaudra, C. speciosa, and
C spinosa, all noctiflorous), another type has developed in the Cleomoideae where the stamens are not
exceeding the petals; this type is better adapted to pollination by bees (the other Mai. spp.).
It is noteworthy that in extra- Malaysian species of Cleome and of Cadaba ornithophily was observed,
and in the South African Cleome natalensis psychophily, i.e. pollination by diurnal butterflies.
Thanks are due to Dr L. van der Pijl, who supplied most of the above information.
Dispersal. Ridley cites very few facts about dispersal of Capparidaceae. It seems certain that
Crateva fruits are dispersed by water. The berries of Capparis, which can attain considerable size in some
species, have sometimes a lively colour (yellow, orange, or bluish-black) and dispersal by birds and/or
bats seems likely. Several species of Cleome have seeds provided with an elaiosome and are supposed to
be dispersed by ants. Their seeds are sometimes provided with small bristles on the sculptured testa and
easily adhere to cloth etc. and seem therefore fit for epizoic dispersal.
Morphology. M. Y. Orr (Notes Bot. Gard. Edinb. 12, 1921, 249-257) found that in several genera
of Capparidaceae, i.a. in Cleome, Polanisia (notably in our spp. Cleome viscosa and Cchelidonii), Capparis
and Crateva, the embryo is completely enveloped by a thin sheath of parenchymatic cells with such
thickenings in the wall as there are found in the water-conducting tracheal tissue. It is assumed that this
layer plays a part in the water supply of the embryo. In the cells of this third innermost seed-coat the
tracheal thickenings are found in the periclinal walls in the Cleomoideae, and in the anticlinal walls in the
Capparidoideae. The same phenomenon was found (I.e. 259) in seeds of the Resedaceae. It may be of
value with regard to classification and judgment of relationships.
The phenomenon of partial sterility is worthy of attention and of closer study. We can observe here
the beginning of an evolutionary development which may finally lead to a complete segregation of the
sexes as in monoecious or dioecious plants. In nearly all Malaysian Capparidaceae only a small part of
the flowers set fruit, except in Cleome spp. where the fertility is complete or almost so. Apparently some
flowers, though not obviously diff'erent in structure, are more fit to produce a ripe fruit than others. In
the Bogor Botanic Gardens I could observe three ligneous species, all represented by a single plant;
of these Capparis lucida regularly produced fruits with viable seeds, whereas in Capparis pubiflora and in
Crateva nurvala not one fruit ever developed. Since all three are native in Java, I am not inclined to
believe that eff"ective pollinators would be absent. Rather I think that the two non-fruiting plants are
self-sterile. Experimental work is needed to reach a conclusion.
In several extra- Malaysian Capparidaceae reductions in the androecium occur. In several Malaysian
species we see reductions in the gynoecium. In part of the flowers of Crateva the whole gynophore with
ovary is shed in an early stage of development. Capparis micracantha and C. scortechinii produce some
flowers with a very short gynophore and a deaf ovary. In Cleome gynandra it is mostly the apical flowers
of a raceme that possess a much smaller, sessile and sterile ovary (see also p. 105).
Stoudt published a valuable study on alternation of sexes and intermittent production of fruits in
Cleome spinosa, dealing with the intricate pattern of reductions both in gynoecium and androecium
(Am. J. Bot. 10, 1923, 57-66).
Phytochemistry. Characteristic features for the Capparidaceae are the following: presence of
myrosin cells, isothiocyanates (mustard-oils) and their parent glucosides (glucocapparin), the frequent
occurrence of deposits of calcium salts (carbonate, sulfate, oxalate) and the absence of leucoanthocyanins
and tannin-like compounds in leaves and stems. These features point to a distinct affinity with the
Cruciferae. This is strengthened by the fact that in both families there is found accumulation of quater-
nary ammonium compounds {e.g. tetramin in Capparidaceae, sinapin in the Cruciferae). If sinapin and
erucic acid (in seed oil) could be demonstrated in Capparidaceae this would suggest still closer affinity
between the two families. Hutchinson's evaluation in ascribing the agreement to parallelism is very
—
improbable from a phytochemical point of view. R. Hegnauer.
Taxonomical affinity. The hitherto generally acknowledged place of the Capparidaceae is in the
order Rhoeadales, with the Cruciferae, Resedaceae, Tovariaceae, Moringaceae, Papaveraceae, and
Fumariaceae; the hierarchic pattern of these relationships varies only slightly with diff'erent authors.
Hutchinson in his concept of two main trends of affinity in the angiosperms, the divisions Herbaceae
and Lignosae, has recently proposed to break up the Rhoeadales (Fam. Fl. PI. ed. 2, 1, 1959, 224). He
Dec. 1960] Capparidaceae (Jacobs) 63

separated the herbaceous Cruciferae, Resedaceae, Papaveraceae, and Fumariaceae from the largely
ligneous Capparidaceae, admitting only a superficial resemblance with the Cruciferae, due to parallel
evolution. In his circumscription the order Capparidales comprises the Capparidaceae, Tovariaceae,
and Moringaceae; it is related to Pittusporales, Tamaricales, Violates, and Polygalales.
The anatomical evidence relating to the affinity of Capparidaceae and Cruciferae is meagre but signifi-
cant, as Metcalfe & Chalk note that the "presence of myrosin cells in certain genera suggests that the
Capparidaceae and Cruciferae may have affinities with one another, and also with the Resedaceae where
similar cells occur" (Anat. Die. 1, 1950, 94).
As pointed out above by Dr Hegnauer the phytochemical data run parallel with the anatomical and
taxonomical evidence. This agreement can in our opinion not be interpreted by parallel development.
It seems likely that the Capparidaceae are a tropical, probably old stratum from which the Cruciferae

represent a specialized branch. This could also well agree with the geographical distribution
pattern of the families, the Capparidaceae being mostly tropical, the Cruciferae mostly temperate.
Wood anatomy, den Berger, Determinatietabel houtsoorten van Malesie, Veenman, Wageningen
(1949) 48 {Crateva) & 51 (Capparis); Desch, Mai. For. Rec. 15' (1941) 70 (hand lens); Metcalfe &
Chalk, Anat. Die. (1950) 91; Moll & Janssonius, 1 (1906) 175.— C.A.R.-G.
1

Uses. No plant of appreciable economic significance is found among the Capparidaceae. Some
minor uses will be dealt with under the species.
Notes. Collectors are requested to pay attention to flower biology. In Crateva too little is known
about the mature fruit and its properties, and of the variability in leaves on one and the same tree.
As to Capparis, a search in the Malay Peninsula and North Sumatra will yield much to complete our
knowledge of certain species. It is highly desirable to make field observations on the structure of flowers
in most species, and to mention the state of maturity whenever fruits are collected. Some Capparis
species produce juvenile or sterile twigs different from the flowering ones; these are badly known and
of much importance.

key to the genera

1. Leaves simple. Plants ligneous.
2. Stamens free, i.e. not connate with the gynophore and consequently after anthesis leaving no scars
on its base. Fruit a berry with leathery pericarp 2. Capparis
2. Stamens at the base connate with the gynophore, leaving scars which remain still distinctly visible
in fruit.
3. Androgynophore 10 mm or longer. Stamens 5-7. Sepals not reflexed. Petals with a long claw.
Fruit cylindrical, ^:-seeded 3. Cadaba
3. Androgynophore 1 to a few mm long. Stamens ^. Sepals reflexed. Petals absent. Fruit ellipsoid,
1-seeded 4. Stixis
I. Leaves palmately compound or dissected.
4. Ligneous plants. Leaves compound. Flowers opening in a very early stage. Fruit globular to ellipsoid,
indehiscent, fleshy 1. Crateva
4. Herbaceous plants. Leaves dissected. Flowers mostly in bud until anthesis. Fruit cylindrical,
dehiscent, dry 5. Cleome

1. CRATEVA
LiNNE, Gen. PI. ed. 5 (1754) 203; Sp. PI. 1 (1753) 444; Hamilton, Trans. Linn.
Soc. 15 (1827) 116 {Crataeva); KuRZ, J. Bot. 12 (1874) 193; Corner, Card. Bull.
S. S. 10 (1939) 15.— Fig. 1-4.
Small to medium-sized trees, facultatively shortly deciduous and then flowering
when bare; glabrous. Branchlets terete with distinct leaf-scars. Stipules small,
caducous. Leaves 3-foliolate, the top of the long petiole sometimes bearing gland-
like appendages on the upper surface. Leaflets sessile to shortly stalked, the lateral
ones basiscopically asymmetrical, sometimes with more or less distinct pellucid
dots. Raceme terminal, corymbiform, either with arrested growth or growing
through and developing into a leafy twig with lateral flowers. Flowers sustained
by bracts, rarely by leaves, pedicelled, opening at a very early stage of development,
floral parts not persistent. Bracts stipulate. Receptacle wide; disk dish-shaped and
incurved. Sepals equal, ovate-spathulate, green. Petals equal, unguiculate, more
or less ovate to rhomboid with narrowed base, first white, later cream-coloured,
the lower (anterior) pair tending to take a transversal (horizontal) position.
64 Flora Malesiana [ser, I, vol. 6^

Fig. 1. Crateva nurvala Ham. var. nurvala (Cult. Hort. Bog. IV-F-76; Jacobs, 1955).

Stamens (8-) 12-30, filaments at the very base connate with the gynophore, long,
filiform, spreading. Gynophore approximately as long as the stamens. Ovary
1-locular, the 2 placentas sometimes intruding to about halfway the lumen but
not coalescent. Stigma conspicuous, flat, soon after anthesis obsolete. In fruit
pedicel, torus and gynophore woody and more or less thickened, the last with
a whorl of filament-scars near the base; the gynophore mostly not stretching.
Berry large, 1 -celled, with tough, sometimes papillate skin. Seeds densely packed;
embedded in pulp, horseshoe-shaped, smooth or crested, one cotyledon larger,
curved round the other.
Distr. A genus of c. 6 spp., pantropical but neither in Australia nor in New Caledonia, the area of
the 3 Indo-Malaysian spp. extending from Ceylon, Western India, South China, South Japan, the
Ryukyus, Formosa, and Hainan, through Malaysia to Tahiti.
Ecol. Mostly in periodically inundated lowland forest near rivers, below 700 m. In dry regions
shortly deciduous, the flowers then appearing simultaneously with the flush. Also cultivated for or-
namental purposes and presumably occasionally introduced.
Dec. 1960] Capparidaceae (Jacobs) 65

Notes. In part of the flowers the gynophore is shed shortly before anthesis, leaving a scar. In some
specimens only the apical flowers remain bisexual. Few flowers set fruit.
It appears to me that there are, in Indo-Malaysia, only few species, widely distributed, which show in

many respects a considerable variability, for example in the size of the floral parts and, to a less degree,
in the sculpture of the seed.
Several authors stated that in some cases the ovary is 2-celled, and they attributed specific value to this
'character'. The matter is, however, that the two parietal placentas intrude into the lumen to a varying
degree, sometimes so deeply as to divide it seemingly into 2 locules, but as far as known, the placentas
do never actually fuse.

KEY TO THE SPECIES^

1. Leaflets with fewer than 1 1 up to 10 cm long, with up to 20, rarely 40, flowers.
pairs of nerves. Rachis
2. Leaflets generally sessile or subsessile, occasionally on petiolules up to 5 mm, very rarely 13 mm.
Blade thin-herbaceous, green when dry, (5'/2-)8'/2-16(-27) cm. Fruit 6-12(-15) cm long, papillate,
greyish when dry 1. C. religiosa'
2. Petiolules 4-5(-6) mm. Blade herbaceous to subcoriaceous, mostly red-brownish when dry, 51/2-IO
(-14) cm. Fruit 31/2-4 cm long, smooth, red-brownish when dry 2. C. odora /. axillaris
.

1. Leaflets with (7-)10-15(-22) pairs of nerves. Rachis 10-16 cm, 20-90-flowered. Leaflets below paler
and duller than above, firmly herbaceous to subcoriaceous. Fruit c. 5 cm long, papillate, finally more
or less smooth, greyish when dry 3. C. nurvala var. nurvala

1/^ cm, nerves 4-6 pairs. Stamens

1. Crateva religiosa Forst. /. PI. Escul. Ins. Oc. pair 1 1/2-2 by 1-1

Austral. (1786) 45, {Crataeva); FI. Ins. Austral. (10-)13-18(-30); filaments 41/2-III/2 cm, pink or
Prod. (1786) 35; DC. Prod. 1 (1824) 243, p.p.; purple towards the top; anthers 2/2-6 by 1 V2 mm,
non Hook./. F1. Br. Ind. 1 (1872) 172; Laut. Bot. sometimes recurved. Gynophore 4-7 cm; ovary
Jahrb. 52 (1914) 110; Merr. Philip. J. Sc. 11 4-6 by 11/2-2/2 mm, subcylindrical, sometimes
(1916) 272; Walker, Imp. Trees Riukiu (1954) 95, ovoid, contracted below the stigma 1 1/2 in mm
f. 45.— C. mewbraiiifolia MiQ. Sum. (1861) 387, diam. Pedicel in fruit (4-)5-7/2(-8) cm, 3-4(-5)
158; Illustr. (1870) 21 Koord. Minah. (1898) 343;
; mm thick; torus 7-11 mm
wide; gynophore
Laut. Bot. Jahrb. 52 (1914) 111; Merr. En. Born. 5/2-8/2(-14) cm long, sometimes cylindrical, c.
(1921) 280; Corner, Ways. Trees (1940) 181.— 3-5 mm diam., or gradually thickened and up to
C. mcicrocarpa KuRZ, J. Bot. 12 (1874) 195, t. 148 1 cm thick at the top. Fruit subglobular to (ob-)
f. 8-10; King, J. As. Soc. Beng. 58, ii (1889) 397; ovoid, 6-12(-15) by 5/2-9/2 cm, wall in the unripe
RiDL. Fl. Mai. Pen. 1 (1922) 125; Merr. Philip. stage up to 7 mm thick, at maturity probably not
J. Sc. 29 (1926) 371; Corner, Card. Bull. S.S. 10 thicker than 1-1 1/2 mm, smooth when very young
(1939) 16 {Crat. B).—C. hausemaimii K. Sch. Bot. but soon covered with flat, pale, dry papillae,
Jahrb. 9 (1888) 201; K. Sch. &
Hollr. Fl. Kais.
Wilh. Land (1889) 50; Warb. Bot. Jahrb. 13
(1891) 318; K. Sch. &
Laut. Fl. Schutzgeb.
(1901) 335, />./7.; Valeton, Bull. Dep. Agric. Ind.
N^erl. 10 (1907) 15; Laut. Bot. Jahrb. 52 (1914)
110.— C. speciosa Volkens, Bot. Jahrb. 31 (1901)
463; Kaneh. J. Dep. Agr. Kyushu Imp. Univ. 4
(1935) 321.— Fig. 2a.
Tree (l-)5-15(-30) m. Stipules 1/2-I mm,
subulate. Leaflets thin-herbaceous, when dry on
both sides of the same greenish colour, much
varying in size on one and the same tree, (51/2-)
8</2-16(-27) by (3-)4-10i/2 cm, central leaflet
oblong, obovate, the base narrowly decurrent, the
apex shortly (incidentally up to 2'/2 cm) acuminate,
often mucronulate; nerves 7-11 pairs; petiole
(3'/2-)6'/2-10 cm, on sterile twigs often longer,
up to 22 cm; petiolules 0-5(-13) mm. Flowers with
a few to over a dozen; rachis 3-5(-14) cm; lower
flowers inserted above the axil of normal leaves,
the others subtended by an early caducous bract.
Pedicels 2-9 cm. Bracts 10 by 1-1 1/2 mm, 3-5 mm
petioled. Sepals ovate, obtuse to acute, 4-7 by
1 1/2-3 mm. Petals once recorded orange, 5-20 mm Fig. 2. Lateral leaflets of Crateva, a. C. religiosa
stalked, blade broadly ovate to elliptic, acute to Forst. f., b. C. odora Ham./, axillaris (Presl)
obtuse, upper pair 2-3(-4) by l-2(-2.3) cm, lower Jacobs, c. C. nurvala Ham. var. nurvala, all X ^/j.

(1) See also 4. C. hygrophila, under imperfectly known species.

66 Flora Malesiana [ser. I, vol. 6^

sometimes giving the impression of a thin, dull, that a variety exists with fruits as small as a
yellowish grey crust. Seeds more or less asymme- thumb's length. Another record about such small
trically cordate, 10-19 by 5-17 mm
diam. and fruits could not be found. The few field notes give
4—8 mmthick, dorsally with a keel rather narrow the colour of the young fruit as pale green, once
and sparsely tuberculate to very broad and dark mauve, and of the ripe fruit white. According
densely tuberculate (sometimes rather smooth), to VAN Royen in New Guinea the fruits are light
the sides smooth to shallowly grooved. green with yellowish scales, hard pericarp, spongy
Distr. India (E. Himalaya), Burma, Lower endocarp, and possess floating capacity. In
Siam, Indo-China, Ryukyus, through Micronesia Bornean specimens seeds were measured 1 by
(Marianes & Carolines) and Melanesia (Solomons) %-l cm in diam. and V2 cm thick; in one specimen
to Polynesia (Fiji, Samoa, Society Is., Gambler from Java the seeds were V2 by I-l Ya cm in diam.
1

Is.); inMalaysia: Sumatra, Riouw Arch., Malay and 1/2 cm thick.

Peninsula, W. Java (twice collected), Borneo (also About the identity of the Palawan specimens I
Banguey I.), S. Philippines (Palawan, Sulu Arch.), am not certain. The material available, Elmer
Celebes (also Kabaena and Muna Is), Moluccas 12650 and Cenabre c.s. FB 27861, is not in a very
(Talaud, Sula, Buru, E. Ceram, Ambon, Kai, Aru, good state and seems to be intermediate in charac-
and Tanimbar), New Guinea (also Normanby I. ters between C. religiosa and C
odora. Pro-
and Salawati), New Ireland. visionally, on account of the large seeds, the com-
Ecol. The species seems to be frequent in paratively large leaflets with a base somewhat
Borneo, New Guinea, and the Solomon Is where decurrent on the petiolule and a short-acuminate
it is often found in periodically inundated forest top, I assume them to belong to C. religiosa,
along rivers, rarely in secondary or primary dry- notwithstanding the brownish colour of the leaves
land forest; one record from the beach in Sarawak, in the Elmer specimen and the almost smooth
where it seems to attain smaller size. Mostly below surface of the seeds.
100 m, but up to 700 m. Fl. fr. in all months. Mr N. G. BissET observed in Tanimbar that
Incidentally cultivated. the fruit pulp had a burning taste, and that the
Vern. Kepayan Malaya, iijesta, S,
(ayer), seeds contained a very high amount of alkaloids,
tigarun, and W. Borneo, kenohan
tjigaron, S. while starch was practically absent.
seguntu, Kutei, niakendem alus, malasiit, sang-
kiauw, Minahasa, tjandaule, SE. Celebes: Toko- 2. Crateva odora Ham. Trans. Linn. Soc. 15 (1827)
laki, kamfooiju,Mangoli, bala-lehe, Muna, 118.
papangi-nasii, Talaud, ombo-ombo, SW. New
Guinea, ai-yumba, bam-baimovi, Solomons, pua /. axillaris (Presl) Jacobs, stat. nov. —
C. tapia
veoveo, Tahiti. (non L.) Bl. Bijdr. 2 (1825) 54; Miq. Fl. Ind. Bat.
Uses. In the Solomon
Is the leaves are heated 1, 2 (1858) 102.— C. axillaris Presl, Rel. Haenk. 2

and applied of ear-ache, and the fruits are

in case (1835) 85; F.-Vill. Nov. App. (1880) 10.— C.
used against constipation. In Yap the fruits are religiosa {non Forst. /.) Blanco, Fl. Filip. (1837)
eaten. The raw fruit is used as fish-bait in W. 399, ed. 2 (1845) 279, ed. 3, 2 (1878) 154, t. 176;
Borneo. F.-ViLL. Nov. App. (1880) 10; Vidal, Sinopsis
BuRKiLL briefly refers (under C. macrocarpa) to Atlas (1883) 13, t. 6 f. C; Phan. Cuming. (1885) 94;
occultpower ascribed to Crateva species in India Rev. PI. Vase. Filip. (1886) 48; Merr. Sp. Blanc.
and Polynesia where it is planted round temples, (1918) 158; En. Philip. 2 (1923) 210; Quis. Med.
to which also Forster's epithet refers (Diet. p. PI. Philip. (1951) 341.— C. tumulorum Miq.
676). Illustr. (1870) 21, t. 11; K. &V. Bijdr. 4 (1896)
Notes. The species very variable in floral
is 269; Back. Schoolfl. (1911) 64; Bekn. Fl. Java
parts, in the shape of the fruit, and the size of the (em. ed.) 4A (1942) fam. 45, p. 6.— Fig. 2b, 3.
seeds. Small tree, 3-10(-30?) m. Stipules falcate, small.
The leaves are mostly (in Malaya to a less Leaflets on slender stalks 4— 5(-6) mm long, her-
degree) so thin in texture that a herbarium spec- baceous to subcoriaceous, in herb, dull, red-
imen with undamaged leaves is extremely rare brownish; 5i/2-10(-14) by 2'/2-5(-7) cm, elliptic
and there is no difference in colour between the to oblong, rarely lanceolate; base cuneate, top
upper and the lower surface. Besides, the seeds, abruptly acutely c. 1-1 Vi cm acuminate; central
however variable, have always a more or less leaflet mostly the largest, broadest at or somewhat
developed crest of warts on the dorsal side. above, rarely below the middle; lateral leaflets
The fruit is once recorded to be compressed. In strongly asymmetrical; nerves generally 5, some-
West Malaysia the subglobular to ellipsoid shape times up to 7, rarelyup to 10 pairs; petiole slender,
seems to prevail, whereas in eastward regions the (3i/2-)6-8(-10) cm, on top occasionally with a
fruits are more elongate. In Malaya the seeds also triangular gland. Inflorescences on small twigs,
seem to be smaller than in the eastern specimens. afterwards whether or not growing through,
From the Solomon Is the fruit is reported to be flower-bearing part c. 3(-10) cm long, with a few
sausage-shaped 14 by 4 cm, with unpleasant odour to about forty flowers partly above the axils of
when cut; in W. New Guinea van Royen noted young leaves or bracteate. Pedicels 4-5(-7) cm.
that "the smell of the fruit fills the forest with a Sepals ovate, acute, c. 4-6 by lV2-2'/2 cm. Petals
soury scent not unlike durian". From Yap suborbicular to broadly ovate-elliptic, (1 V2-)
VoLKENS mentioned a fruit 18 by 10 cm and added 21/2-3 by 1-2 cm, the top blunt to rounded or
Dec. 1960] Capparidaceae (Jacobs) 67

Fig. 3. Crateva odoia Ham./, axillaris (Presl) Jacobs. Sumbawa, 1934 (de Voogd 1911).

sometimes notched, base rounded and (rather) ab- doubt that it has been introduced long ago;
little
ruptly narrowed into the stalk 6-10 mm. Stamens it sometimes found planted on graves.
is

21-25, 3-4 cm long, anthers c. 3 by 1 mm, re- Ecol. Prefers dry, shrubby places. Mostly deci-
curved. Gyiiophore (2-)3^(-6'/2) cm, ovary sub- duous, the flowers then appear just before the
globular to oblong, c. 2-5 by 2 mm, constricted young leaves break out, but the blossoming seems
below the stigma c. V2 mm broad. In fruit the
1 not to show periodicity. In the Malay Peninsula it

gynophore slightly thicker than the pedicel, c. is only rarely found in fruit.

2-3 mm all over, torus c. 1

thick mm
wide. Fruit Uses. QuisuMBiNG, Med. PI. Philip. (1951) 341,
globular, (up to?) 3V2-4 cm diam. (in Indo- mentioned quite a few minor medical applications.
Chinese specimens up to 6 cm), pericarp leathery, Vern. Sempal wadak, J (for the genus, reliable);
c. 1 mmthick, smooth during the whole develop- kenialo-kemalowan, sekar bulan, Kangean, saling-
ment, at maturity probably red when fresh, bogog. Tag. (Philip.).
brownred when dry. Seeds rather irregularly Notes. From C. religiosa distinguished vege-
horseshoe to deeply kidney-shaped, about 6 by 2 tatively by its leaves being firmer in texture, often
mm, smooth. red-brownish in the herbarium, especially the
Distr. The species occurs throughout India, specimens from Java and the Lesser Sunda Is.,
Ceylon, Burma, Indo-China, S. China, Formosa, and giving a more graceful impression, due to the
and Hainan, the Malaysian form in S. India smaller size, the slender distinct petiolule, and the
(Coimbatore), Ceylon, Malay Peninsula (Malacca: acute longer tip of the leaflets. In fruit it is diflTerent
cultivated), Java (W. and E. part, cultivated), by its shorter gynophore and by the surface of the
Madura, Kangean Arch., Lesser Sunda Is. fruit which is smooth and not papillate; this
(Sumbawa), Philippines (N. Palawan, Mindoro, character being more reliable than those provided
Luzon, most provinces, Guimaras, Mindanao). by the (often immature) seeds.
There are far more collections from Luzon than In one specimen, cultivated in Hort. Bog. under
from any other locality. It looks as if the species is IV-F-81, the largest leaflet was 17 by 9 cm. Its
native in the Philippines. It cannot be verified adult leaves were only slightly reddish tinged, the
whether it is introduced or native in Kangean and tip being more or less abortive or absent. This
Sumbawa; in Madura, Java, and Malaya there is seems to prove once more that species may show
68 Flora Malesiana [ser. I, vol. 6^

deviations unknown from the wild state merely by hardly lengthened (rarely up to 10 cm), 3-4(-5)
cultivation in a botanic garden. mm thick, only slightly thicker towards the top;
The closest affinity is with the African C. pedicel mostly thinner, torus 7-8 mm
wide. Fruit
adamonii DC. which could even be looked upon unknown in fully mature state, ellipsoid, rarely
as a subspecies. In C. adaiisonii the inflorescence ovoid, (up to?) 5-51/2 by 4-41/2 cm, pericarp 4—5
rachis is + 4-5 cm, and finally set with the thick mm thick, covered with a thin, dull, yellow-
callous scars of the shed pedicels; this top of the
rachis often starts to grow again vegetatively
afterwards. The petals are about 2-3 by 1-2 cm
in all, the stamens 15-20, c. 3 cm long, the fruit
greenish and not reddish in the dry state. On ac-
it seems appropriate to
count of these diff'erences
keep them apart.

3. Crateva nurvala Ham. Trans. Linn. Soc. 15

(1827) 121, (Crataevci Niirvala); W. & A. Prod. 1
(1834) 23; KuRZ, J. Bot. 12 (1874) 195; Gagn.
Not. Syst. 8 (1939) 213; Fl. Gen. I.-C. Suppl. 1
(1939) 157, t. 14 f. 8.

Fig. 4. Crateva nurvala Ham. var. nurvala. Flower,

var. nurvala. C. nurvala Ham. I.e.; MiQ. Illustr.
front view (Cult. Hort. Bog. IV-F-76; Jacobs.
(1870) 20; K. & V. Bijdr. 4 (1896) 266; Back.
1955).
Schoolfl. (1911) 64; Hall./, in Winkl. Bot. Jahrb.
49 (1913) 369; Merr. En. Born. PI. (1921) 280;
Back. Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, greyish crust breaking into minute particles which
p. 6.— Niirvala Rheede, Hort. Malab. 3 (1682) seem to peel oflT sooner or later, leaving the surface
—
49, t. 42. C. religiosa (iwn Forst./.) Bl. Bijdr. 2 smoothish. Seeds deeply horseshoe-shaped, 6-9
(1825) 54; Ridl. Fl. Mai. Pen. 1 (1922) 125; mm long and wide, 2-3 mm
thick, dorsally with a
Rendle, J. Bot. 63 Suppl. (1924) 5. —C
nmgyw crest of sharp irregular protrusions.
[a// (Lour.) DC.?] Hassk. PI. Jav. Ran (1848) 179; Distr. India (Deccan, Sikkim, Assam), Burma,
MiQ. Fl. Ind. Bat. 1, 2 (1858) 102; Sum. (1862) S. China (Yunnan, Kweichow?), Hainan, Indo-
158, 387. C. religiosa non Forst./. var. nurvala China, Siam; in Malaysia: Sumatra (P. Weh,
(Ham.) Hook./ & Thoms. Fl. Br. Ind. 1 (1872) East and West Coast Res., Palembang), Malay
172, p.p.~C. loplwsperma KuRZ, J. Bot. 12 (1874) Peninsula, W. to E. Java, and Borneo. Cultivated
195, t. 147 f. 4-6, an var. propria'^.; Corner, Gard. throughout the area, for ornamental or magic
Bull. S.S. 10 (1939) 16 {Crat. A); Ways. Trees purposes.
(1940) 181, f. 48.— Fig. 1, 2c, 4. Another variety occurs in India (mainly in the
Tree, (8-)10-15(-20) m, to 40 cm diam. Branch- Northeast), Burma, Siam, Indo-China, S. China
lets slightly zig-zag. Stipules minute, late caducous, (Yunnan, Kwangtung, Fukien), Hainan, and S.
acute with broad base. Leaflets firmly herbaceous Japan (probably this species).
to subcoriaceous, (0-)3-6(-10) mm stalked, mostly Ecol. Mostly along streams in shady places,
lanceolate, sometimes oblong, rarely linear, sometimes behind the sea-shore, at low altitudes
(4i/2-)9-15(-28) by (1 i/2-)3-5(-6V2) cm; base up to 600 m. Seems to be rare everywhere. Flower-
acute, top acuminate, tip acute; central leaflet ing and fruiting time irregular.
broadest about or below, rarely above the middle, Vern. Si baluak, Sumatra, badat, dalaili), dalur,
lateral ones more or less symmetrical; nerves Mai. Pen., barun(d)aj, ki /lowe, S, djaranan, sempal
(7-) 10- 15 (-22) pairs; surface below duller and wada, J, pingos, sasagali, sebelii, tigarun, Borneo.
paler than above, sometimes on both sides with Uses. The wood seems to find local application
minute grey-brown scattered papillae. Petiole as timber. In Malaya the root, leaf, or bark are
(4-)5 1/2-9 '/2(- 14) cm, vigorous, broadly sulcate, boiled with oil and applied externally in case of
on top bearing numerous pale to light brown 'sakit angin". In Siam the fruit is used as fish-bait
gland-like appendages up to 1 mm long. In- and the young leaves are pickled in salt. See
florescence ultimately c. 10-16 cm long, bearing BuRKiLL (Diet. 1, p. 676) for other minor uses.
20-100 flowers. Bracts early caducous, 5-9 by According to Filet, cf. Heyne, Nutt. PI. p. 682,
%-l V2 mm, acute, their stipules minute, longer the bark contains an acrid, bitter substance and is
persistent. Pedicels 4-7 cm. Sepals 2-31/2 by pounded in water, used as a skin-irritant against
II/4-II/2 rnrn, ovate, acute, somewhat narrowed high fever, etc.
at the base. Petals 5-12 mmstalked, blade (8-) Notes. This species is vegetatively characterized

15-30 by (5-) 15-22 mm, suborbicular or sub- by nerf-pattern the leaves have underneath a
its :

rhomboid to elliptic, ovate, base rounded, ab- typical aspect by the thin but vigorous and prom-
ruptly narrowed into the stalk, top obtuse. inent nerves, more numerous than in other
Stamens 15-25, 3i/2-4i/2(-6) cm; filaments purple; species, while the insertions of the intermediate
anthers c. 2-3 by I mm. Ovary ellipsoid to cylin- veins on the midrib are also far more distinct.
drical, c. 5-6 by 2/2 mm; stigma dark purple. Besides, the lower surface is more pale, dull, and
Gynophore 3V^-5V^ cm, in fruit probably not or greyish or glaucous than the upper, the difference
Dec. 1960] Capparidaceae (Jacobs) 69

being far greater than in the other Cratevas. top; nerves ± 6-7 pairs, as the midrib narrow,
In none of the herbaria consuUed any type prominent above. Inflorescence rachis practically
material of C. loplwsperma was found. Kurz none. F/ovt'^r.y unknown. Pedicels in fruit 5 '/2-6'/4
named as type specimen "Gustav Mann, from cm, gynophore A^/^-SY^ cm, both blackish, woody,
banks of the Koolsee River, Kamroop, Assam, ± 2-2 1/2 mm in diam. Fa7//7 (immature) cylindrical
Fr. July." From the description it is clear that one umbonate, ± 9 by 2'/2 cm, dull brown-purplish
of the two varieties of C/;///r«/fl was meant; pro- with numerous white lenticel-like specks, c. 1
'/2

bably it was var. nurvala. Corner's interpretation mm thick. Seeds irregularly horseshoe-shaped,
of KuRz's species as C. nurvala s. lat. was correct; ±11 by 8 by 3 mm, the outer side with some
his cited material no doubt belongs to var. nurvala, coarse sculpture.
but perhaps Kurz himself had Hamilton's Distr. Burma (Pegu); in Malaysia: Penang
'C. H/»7c>r///flm' at hand, which is more frequent in (King's Coll. 1412).
Assam than var. nurvala. Note. There are only two collections; both are
inadequate but match very well. They come nearest
Imperfectly known to C. religiosa, but I feel reluctant to refer them to
this species. They may represent an abnormal,
4. Crateva hygrophila Kurz, J. As. Soc. Beng. 41, deviating form
ii(1872) 292; J. Bot. 12 (1874) 196, t. 148 f. 6, 7;
J. As. Soc. Beng. 48, ii (1874) 33; Fl. Burm. 1 Excluded
(1877) 67.
Leaves herbaceous, dull reddish brownish, Crateva marmelos L. Sp. PI. (1753) 444 =
by 31/2 cm, leaflets sessile, not
largest leaflet I21/2 Aegle marmelos (L.) Correa in Trans. Linn. Soc.
much asymmetrical, narrowed towards base and 5 (1800) 223 {Rutaceae).

2. CAPPARIS
TouRN. ex LiNNE, Gen. PI. ed. 5 (1754) 222; Sp. PI. 1 (1753) 503; Lamk, Encycl.
1 (1785) 604; DC. Prod. 1 (1824) 245; MiQ. Illustr. (1870) 22-36, t. 12-19; Gagn.
in Morot, J. Bot. 21 (1908) 53; Pax Hoffm. in E. &
P. Pfl. Fam. ed. 2, 17b &
(1936) \12.—Busbeckea Endl. Fl. Norfolk. (1833) 64.— Fig. 5-25.
Shrubs, often sprawling or climbing, rarely small trees, mostly hairy but
glabrescent. Leaves simple, with a pair of stipular thorns which are occasionally
wanting, sometimes these persistent on knobs on the main stems; nerves arcuating
and interlooping near the margin. Flowers pedicelled, arranged in
fairly regularly
serial, supra-axillary rows (flowering basiscopically!), or in racemes with the
pedicels subumbellately conferted towards the top, the subumbels sometimes
paniculate, or more rarely flowers solitary, axillary. Bracts mostly present but early
caducous, rarely 2 basal bracteoles. Sepals biseriate, mostly imbricate, the outer
pair mostly strongly concave, the inner pair flattish, rarely (in sect. Busbeckea)
the outer pair connate in bud. Petals 4, variously imbricate, rather delicate, not
unguiculate, the two adaxial ones (upper pair) with asymmetrical base coherent
and surrounding a small disk, the two abaxial ones (lower pair) quite free. Recept-
acle (torus) slightly thickened, ±
conical, with a more or less developed adaxial
disk. Stamens co, rarely (7-)8(-12) free, radiating, longer than the petals,
glabrous; anthers small. Gynophore about as long as the stamens, sometimes
longer, not or very little stretching in fruit, exceptionally abortive, irregularly
coiled in bud. Ovary 1-locular, placentas 2-6, mostly 4, with cnd ovules; stigma
sessile, small. Berry in Mai. globular to ellipsoid, rarely elongate, with leathery or
corky pericarp, -celled. Seeds (1-)^^, embedded in pulp, obliquely reniform,
1

rather large, with circinnate embryo.

70 Flora Malesiana [ser. I, vol. 6^

Distr. Presumably about 250 spp. in the tropics and subtropics of both hemispheres, especially in
America and Africa. Another centre of development, with c. 40 spp., is found in Burma and Indo-China.
In Malaysia 23 spp. are recognized, among which only a few are confined to one island or province,
the development being richest in the Philippines where also the infraspecific variability is greater than in
other islands. A few species are shared by SE. Malaysia and northern Australia. There are only few
species which are chiefly distributed in the large rain-forest area of western Malaysia (Malaya, Sumatra,
W. Java, and Borneo) and most species avoid that area because of their preference for dry and seasonal
climatic conditions. Fig. 6.
Ecol. Only few spp. are adapted to primary rain-forest conditions; they are most frequent in helio-
philous, warm and dry habitats under seasonal climatic conditions, for example in coastal vegetation,
in savannahs, hedges, light forest, secondary forest, thickets, and forest borders, in the lowlands and
hills, the highest record being 1700 m.
Disp. The often large pulpy fruits may have various colours at maturity, for instance bluish-black in
C. lanceolaris DC, deep-yellow in C. trinervia Hook./., orange in C.ftoiibunda Wight, and are likely
to be eaten and (?)dispersed by animals. In the last-named species Elmer noted "fruits opened and seed
eaten by birds".
Pollination. Though the flowers are often scented and the inflorescences are in several species very
showy, no observations on flower visitors have come to my knowledge. The honey is very much concealed
under the coherent asymmetrical bases of the adaxial petals which surround a small disk.
Two species, viz C. liicida (Banks ex DC.) Benth. and C. spinosa var. mariana (Jacq.) K. Sch., are
known to be noctiflorous. C. ervcibe Hall./., C. micracantha DC, and C. pubiflora DC. were observed
to flower in the daytime; they may be open and scenting during the night as well.
Indument. The characters of the hairs covering at least the young parts of many species are often of
specific value. In Asiatic species the indument consists only of hairs, but for instance in C. breynia Jacq.
of tropical America the young parts are densely scaly like a Durio twig. Short-stalked, stellate hairs,
with 3-4 arms arising from a 1 -cellular base are found in 10. C. sepiaria and 20. C. quiniflora (also having
many hairs with 2 arms, like the malpighiaceous balance-hairs); 14. C pubiflora (hairs very long and
silky); 15. C. pyrifolia (hairs slender, glassy, with unequal arms); 19. C. zeylanica (hairs thicker, less
regularly shaped); 22. C. spinosa var. mariana (hairs small, white, soft, irregularly stellate, the arms
somewhat twisted; hook-like hairs as depicted in E. & P. Pfl. Fam. 17b, f. 78A, were not seen in my
limited material). Simple hairs, patent unless stated otherwise, occur in: 1. C. scortechinii (hairs soft,
mostly straight, more or less erect); 2. C. trinervia (hairs rather long and straight, but near the base bent
in an arbitrary direction); 8. C. floribunda f. indiita (hairs soft and somewhat twisted); 7. C. lanceolaris
(like the last, but straighter and generally shorter); 9. C. lobbiana (hairs stiff, straight, and very unequal);
6. C. longestipitata (hairs appressed, singularly short and thin); 5. C. cantoniensis (hairs appressed,
mostly unbranched, small, soft, twisted); 23. C. lucida (hairs small, somewhat twisted). Species not
mentioned above are glabrous or almost so.
Taxon. A complete subdivision cannot yet be given, because the Old World species need further
study. If we follow de Candolle's subdivision, then spp. 1-22 come into the sect. Capparis {Eucapparis
Plum, ex DC), with imbricate sepals.
Of these, spp. 4-15 fall into subsect. Corymbosae DC. with the flowers in subumbels, spp. 16-20 into
subsect. Seriates DC. (incl.^ecr. Monostichocalyx Radlk.) with the flowers in supra-axillary rows, sp. 22
into subsect. Pedicellares DC. with axillary flowers.
Sp. 23 is the only Malaysian representative of the Australian sect. Busbeckea (Endl.) B. H. with &
the outer pair of sepals connate in bud.
Spp. 1, 2, 3, and 21 are as yet difficult to place in this classification.
Note. There are a few field observations that the completely mature fruit would dehisce by valves in
some species. I have not seen any material confirming this.

KEY to the species

1. Flowers solitary in the leaf axils.

2. Leaves oblong. Sepals 5-7 mm
long 21. C. larutensis
2. Leaves suborbicular. Sepals longer than 10 mm 22. C. spinosa
1. Flowers in serial rows, in axillary subumbels or panicles, or in terminal racemes or panicles (rarely

depauperated to 1 flower in 23. C. lucida).

3. Inflorescences exclusively terminal on normal twigs.
4. Sepals longer than 7 mm. Gynophore longer than 2'/2 cm.
5. Pedicels '/2-I cm. Fruit c. 10 cm diam. Gynophore c. 5 cm 1. C. scortechinii

5. Pedicels 1-6 cm.

6. Sepals in bud connate, 10-15 mm
long, glabrous. Leaves shorter than 9 cm, base acute.
23. C. lucida
6. Sepals in bud imbricate. Leaves longer than (6-) 10 cm.
7. Plant glabrous. Leaves oblong to lanceolate, coriaceous.
Dec. 1960] Capparidaceae (Jacobs) 71

8. Gynophore 3-4'/2 cm. Sepals 8-12 mm long. Leaves with a hardened tip.
12. C. callophyUa
8. Gynophore 2-31/2 cm. Sepals 5-6(-10) mm long. Leaf-tip not particularly thickened.
13. C. zippeliana
Innovations hairy, glabrescent. Leaves obovate, subcoriaceous, subtriplinerved, (6-) 10-14 cm
7.
long 2. C. trinervia
4. Sepals shorter than 7 mm. Gynophore shorter than 21/2 cm.
9. Flowers in a panicle. Leaves longer than (8-)10 cm.
10. Gynophore 4-12 mm. Fruit IY2 cm diam 4. C. erycibe
10. Gynophore 2-3 V2 cm. Fruit c. 31/2 cm in diam 13. C. zippeliana
9. Flowers in simple subumbels. Leaves shorter than 8 cm. Gynophore shorter than 2 cm.
11. Stamens 30-45. Gynophore (5-)6-10(-13) mm. Twigs vigorous, hairy, zig-zag, mostly with
strong thorns 10. C. sepiaria
11. Stamens 12-15. Gynophore c. 15 mm. Twigs slender, glabrescent, straight, with weaker thorns.
11. C. diifusa
3. Inflorescences axillary or supra-axillary, sometimes also terminal.
12. Pedicels (or their scars) serial, in supra-axillary rows.
13. Sepals glabrous or ciliate. Young leaves glabrous.
14. Sepals shorter than 5 mm, glabrous. Fruit tuberculate. Thorns, if present, recurved.
16. C. buwaldae
14. Sepals longer than 5 mm, ciliate. Fruit smooth. Thorns patent 18. C. micracantha
. .

13. Sepals puberulous outside.

15. Young leaves hairy, at least underneath. Fruit ±
globular.
16. Fruit smaller than 12 mm. Thorns directed upwards, straight or slightly curved. Sepals 4-5 mm
long 15. C. pyrifolia
16. Fruit larger than 15 mm. Thorns, if present, recurved.
17. Stamens more than 30. Sepals longer than 6 mm 19. C. zeylanica
17. Stamens 7-8. Sepals shorter than 5 mm 20. C. quiniflora
15. Young leaves glabrous. Fruit elongate 17. C. cucurbitina
12. Flowers in sessile racemes or in subumbels or panicles.
18. Flowers in short, sessile, axillary racemes.
19. Gynophore at anthesis hairy all over. Rachis of the inflorescence inconspicuous.
14. C. pubiflora
19. Gynophore at anthesis glabrous. Rachis of the inflorescence thick, 1-2 '/2 cm long.
3. C. brachybotrya
18. Flowers in subumbels or in panicles, rarely in a small leafy raceme.
20. Leaves (8-)13-20(-26) cm long, often reddish tinged when dry.
21. Gynophore 3-4'/2 cm. Sepals 8-12 mm
long. Leaves with a hardened tip. 12. C. callophyUa
21. Gynophore 2-31/2 cm. Sepals 5-6(-10) mm
long. Leaf-top not particularly thickened.
13. C. zippeliana
20. Leaves (2-)4-12(-16) cm long, green to yellowish when dry.
22. Gynophore at anthesis hairy all over. Fruit up to 2 cm long . 14. C. pubiflora
. .

22. Gynophore at anthesis glabrous, at least in the upper half.

23. Gynophore shorter than 1/2 cm.
1 Fruit 1/2-2 cm diam.
24. Stamens c. 8. Sepals 3-4 mm
long. Fruit 1/2-2 cm diam.
1 .8. C. floribunda
. .

24. Stamens cn:>. Sepals 4-5 mm

long. Fruit 1 1/2 cm or smaller.
25. Twigs stout, markedly zig-zag, greyish-hairy, with vigorous thorns. Leaf-top rounded,
notched. Midrib flattish above 10. C. sepiaria
25. Twigs slack, approximately straight, glabrescent, with weaker thorns. Leaf-top acuminate.
Midrib narrowly sulcate above 5. C. cantoniensis
23. Gynophore longer than 1 1/2 cm. Fruit (lV2-)2-3i/2 cm diam.
26. Twigs glabrous or with appressed pubescence. Thorns, if present, recurved.
27. Leaves 6i/2-12(-16) cm long, the margin somewhat recurved. Sepals at least 6 mm long.
Fruit 2-31/2 cm diam 7. C. lanceolaris
27. Leaves 5-6 cm long, flat.
Sepals c. 3 mm
long. Fruit unknown 6. C. longestipitata
.

26. Twigs generally with short, patent pubescence. Leaves 4-8(-13) cm long, ovate, subcordate.
Fruit c. 1 1/2 cm diam. Thorns, if present, straight 9. C. lobbiana

1.Capparis scortechinii King, J. As. Soc. Beng. 58, 2-4 mm. Leaves (sub)coriaceous, pubescent when
ii(1889) 394; Ann. R. Bot. Gard. Calc. 5 (1896) young, soon glabrescent, ovate to obovate, 2-5
118, t. 135; RiDL. Fl. Mai. Pen. 1 (1922) 122. times as long as broad, 6i/2-12(-21) by n/2-5
Climbing shrub, 2-10 m. Twigs straight, an- (-7/2) cm; base narrowed, acute; top rounded to
gular and pubescent, terete and glabrescent when narrowed, more or less acuminate, dark-mucro-
older; internodes 1-3 cm. r/jor/w strong, recurved, nate; midrib flat above; nerves c. 5-6 pairs;
72 Flora Malesiana [ser. I, vol. 6^

Fig. 5. Capparis zeylanica L.a. Flowering twig, the leaves still young, X 2/3, b. flower, nat. size, c. fruiting
branch with adult leaves, the fruit opened, X 2/3.
Dec. 1960] Capparidaceae (Jacobs) 73

petiole c.1 cm, brown pubescent, late glabrescent. basal part; nerves (6-)7-8(-10) pairs, glabrous to
Raceme terminal, 2V2-IO cm long, brown pubes- early glabrescent; petiole (7-)10-14(-18) mm,
cent all over. Bracts 10 by 1 '/2-2 mm, sometimes hairy as the twig. Flowers (3-)5-10(-l5) in a
larger and resembling small leaves. Pedicels terminal raceme. Pedicels rather vigorous, in the
5-10 mm, leaving a prominent scar, in fruit woody. axils of the upper leaves or in the axils of very
Sepals densely pubescent outside, with mem-
branous, ciliate margin 1 mm
broad, outer pair
coriaceous, orbicular, 8-11 mm
diam., inner pair
subcoriaceous, ovate, 2-6 by 4 1/2 mm, densely
pubescent outside. Petals pink, c. 8-9 by 4'/2-6
mm, obovate, notched, with cuneate base, sparsely
pubescent inside, or glabrous. Stamens 35-50, c.
15 mm. Gynoplwie c. 5-61/2 cm (occasionally
abortive), in fruit transversely wrinkled, woody,
thickened up to 16 mm; ovary ovoid, 1% by ^/^
mm. Fruit globular, c. 10(-12i/2) cm diam., peri-
carp woody, 2-21/2 cm thick, smooth, yellow.
Seeds cv;, c. 2Vi by P,{}-2'/2 by c. 1 cm.
Distr. Malaysia: ? Banka, Malay Peninsula
(Perak, Trengganu, Pahang, Selangor, and
Penang).
Ecol. In rain-forests, up to 1400 m.
Vern. Menawul, Banka, suso/i Pahang.
beriiga,
Fig. 7. Approximate area of distribution of
Capparis trinervia Hook. /. & Th. (broken line)
Note. An entirely glabrous specimen
sterile
collected by Teysmann s.n. (Bo) in Banka, pro-
and localities of C. quiniflora DC. (dots within
bably belongs here. In King's type collection both continuous line), the latter also in Fiji.
flowers with an abortive and a developed gyno-
phore are represented. soon caducous c. 4 mm long narrow bracts, c.
2-4 cm long, widened at the top, hairy. Buds
approximately globular, pointed. Sepals c. 9-12
by mm, outside densely orange-yellow puberu-
1 1

lous, the outer pair coriaceous, the inner pair

thinner, flattish, suborbicular, with membranous
margin. Petals light red, obovate, c. 12-15 by
8-9 mm, puberulous towards the base on both
sides, the margins crisp, except at the base. Torus
c. 5(-6) mm wide, flattish. Stamens c. 60-70, c.
27 mm long, whitish. Gynophore ly^-AVi cm long,
light red, glabrous, ovary ellipsoid to ovoid, 2V2
mm long, with umbonate stigma, green, glabrous.
Fruit globular, deep yellow, 3i/^ cm in diam.,
pericarp soft-woody, 4 mm thick. Seeds 15-17 by
14 by 6 mm.
Distr. Indo-China (Tonkin, Laos, Annam),
Fig. 6. Species density of the genus Capparis in
Burma (Tenasserim); in Malaysia: NE. Sumatra
Malaysia. The number above the hyphen reters (Tinggiradja, N of lake Toba), Malay Peninsula
to the number of endemic species, that below the (Perak: G. Booboo). Fig. 7.
hyphen to the number of other species in each Ecol. A creeper reported clinging to trees in
province. C. longipes Merr. (incompletely known; dense jungle, and from dry, almost bare lime-
from the P.I.) has not been incorporated. cinder ledges, at low altitude.
Note. The Malaysian specimens differ slightly
from those of Indo-China where the leaves have
2. Capparis trinervia Hook./. & Th. F1. Br. Ind. 1 3-5 pairs of nerves and the buds are not
(1872) 175; Kurz, F1. Burm. 1 (1874) 64; Gagn. pointed.
Fl. Gen. I.-C. 1 (1908) 193; Suppl. 1 (1939) 168. —
C. kiinstleri King, J. As. Soc. Beng. 58, ii (1889) 3. Capparis brachybotrya Hall. /. in Fedde, Rep.
396; RiDL. Fl. Mai. Pen. 1 (1922) 122. 2 (1906) 59; Laut. Bot. Jahrb. 52 (1914) 112.
Climber or shrub, up to 4 m. Young parts Branchlets stout, terete, slightly zig-zag, gla-
ferruginous-tomentose, soon glabrescent. Twigs brous, internodes 2i2-5 cm. Thorns slightly re-
almost straight, often angular, internodes c. 3-5 curved, minute to 2 mm long. Leaves coriaceous,
cm; thorns l-2(-3) mm, patent or slightly re- elliptic, oblong or sublanceolate, light green when
curved. Leaves subcoriaceous, elliptic to oblong, dry, glabrous, 13-211,2 by 5-10i2 cm; base ob-
slightly obovate, (6-)10-14 by (254-)3 1/2-5 1/2 cm; tusely acutish to rounded; top rounded and 2-10
base cuneate, often subtriplinerved, top acuminate mm acuminate, with a dark and stiff" tip; nerves
with acute tip c. %
cm; midrib above sulcate in the (6-)7-9(-10) pairs, depressed above. Petiole stout,
74 Flora Malesiana [ser. I, vol. 6^

Fig. 8. Capparis erycibe Hall./, a. Habit, X 2/3, b. flower, X 6, c. fruits, X 2/3 (after Jacobs 4838).
Dec. 1960] Capparidaceae (Jacobs) 75

\-lY2 cm. Racemes axillary, sometimes serially in Climber, about l'/^(-14?) m. Twigs slack,
twos; rachis stout, (l-)2'/2 cm, up to 20-flowered, brownish, brown-puberulous, especially when
pale-puberulous, glabrescent, at the very base young, slightly zig-zag; internodes about 3-5 cm.
surrounded by conferted bracts. Bracts subulate,
small, early caducous; bracteoles stiff, subulate,
minute, later caducous. Pedicel l-2'/2 cm, slightly
thickened towards the top, puberulous, glabres-
cent. Flowers (pinkish) white, scented. Buds sub-
globular, occasionally apiculate, 1 cm diam.
Sepals c. 11-14 by 6-10 mm, more or less mu-
cronulate, outer ones subglabrous, inner ones out-
side puberulous along the margin. Petals 12-20 by
6-10 mm, outside puberulous, especially in the
upper part; upper petals the smallest, in the basal
median part densely puberulous inside; lower
petals glabrous inside. Disk fleshy, c. 1 mm
diam.,
glabrous. Stamens (80-) 100-1 60, 25-35 mm
long,
filaments white. Gyuophore 2-3 1/2 cm; ovary ovoid,
2-3 by 1 mm, stigma I/2 mm, all glabrous.

/. brachybotrya.
Leaves elliptic, 1.7-2.1 times as long as broad,
14-22 by IVi-X^Vi cm, with rounded base. Fruit
on a gynophore '/2-2'4 cm, ellipsoid with narrow-
1

ed base and short, abruptly acuminate top, 4—51/2

by 3'/4 cm, pericarp leathery. Seeds cv, c. 10 by 7
by 5 mm, light brown.
Distr. Malaysia: Moluccas (Kai Is), New
Guinea (Vogelkop and Batanta Is).
Ecol. Primary and secondary forest, up to
250 m.
Note. The fruits belonging to the type specimen
were separated from it and got lost; the only
collection of ripe fruits now available was made by
VAN RoYEN. He informed me that the short
peduncle with several fruits on their long stalks
was quite showy. The fresh fruits are somewhat
larger than the dry ones, red, smooth, and they
possess the abruptly pointed base and apex.

/. angustifolia (Hall. /.) Jacobs, stat. now C. —

brachybotrya var. angustifolia Hall. /. in Fedde,
Rep. 2 (1906) 60; Laut. Bot. Jahrb. 52 (1914) 12. 1

—Fig. 9.
Leaves oblong, 2.4-3.3 times as long as broad,
I3-2IV2 by 5-7 cm, with obtusely acutish to
rounded base. Gynophore 41/2 cm; fruit (almost
mature?) elongate, 4 by 1 1/2 cm, pericarp thin.
Seeds cv), c. 8 by 7 by 4 mm.
Distr. Malaysia: Celebes (Pangkadjene, Mina-
hasa), New Guinea (Vogelkop).
Ecol. Coastal forest. fV^is^-

Note. In sterile state very similar to C

micra-
cantha, but easily distinguished by the axillary Fig. 9. Capparis brachybotrya Hall. /. /. angusti-
inflorescence with a thick rachis, C. micracantha folia (Hall. /.) Jacobs, node with leaf and
having a supra-axillary row of pedicels or their inflorescence, x 2/3 (Kostermans 2819).
scars.
Thorns up to 2 mm
long, recurved, mostly wanting.
4. Capparis erycibe Hall. /. Bull. Herb. Boiss. 6 Leaves herbaceous to subcoriaceous, often reddish
(1898) 216; Back. Schoolfl. (1911) 62; Koord. brown when dry, glabrous above, glabrescent
Exk. Fl. Java 2 (1912) 294; Back. Bekn. Fl. Java beneath, broadest above, sometimes at the middle,
(em. ed.) 4A (1942) fam. 45, p. 10.— C. paniculata 1.8-2.5(-2.8) times as long as broad, (9"/2)-12-16
RiDL. J. Fed. Mai. St. Mus. 10 (1920) 129; Ridl. (-20) by AYi-^y'i cm. base mostly narrowed,
Fl. Mai. Pen. 1 (1922) 124; Baker, J. Bot. 63 sometimes rounded, acute to obtuse; top rounded
Suppl. (1924) 5.—Fig. 8. to acuminate, mucronate; midrib above mostly
76 Flora Malesiana [ser. I, vol. 6^

narrowly sulcate; nerves c. 6-8 pairs, glabrous and acutish, mucronate; midrib above sulcate all

above, glabrescent beneath; petiole 4-10 mm, over; nerves inconspicuous, 6-9(-ll) pairs;
brown-pubescent, late glabrescent. Panicle mostly petiole 5-6(-10) mm, hairy as the twig. Flowers
brown-puberulous on a c. 5-10 cm long, slender sometimes fragrant, in axillary subumbels which
peduncle terminal on a twig of which the upper are often arranged in a terminal panicle c. 15-20
part has mostly lost its leaves, c. 10-20 by 5-15 cm. cm long, each a few cm peduncled, sparsely hairy.
Pedicels slender, 4-18 mm. Bracts minute, some- Pedicels slender, 1/2-2 cm. Bracts subulate, 1-2
times wanting. Flowers white to greenish, some- mm long, caducous; bracteoles basal, minute,
times tinged reddish. Buds globular, c. 4 mm diam. sometimes wanting. Buds globular, 4-5 mm diam.
Sepals 4-6 by 2'/2-3 mm, the outer pair sometimes Outer sepals 4'/2-6(-7) mm diam., sometimes
sparsely puberulous outside; the inner pair with sparsely puberulous at the base outside, inner
broad membranous margin, glabrous. Petals 4 sepals elliptic to obovate, 5-7(-8) by 4-6 mm, with
(occasionally single flowers with 5 or 6), 4'/2-6 by membranous, ciliate margin. Petals white (some-
1-4 mm, suborbicular to subspathulate, sometimes times greenish or pinkish?), (3'/2-)5-6'/2 by
puberulous, especially at the base. Stamens 20^0, 2-3(-4) mm. mostly obovate, pubescent. Stamens
5-6(-8) mm. Ovary ovoid to spindle-shaped, 2 by 20^5, 15-25(-32) mm long, filaments white.
1 mm, glabrous; gyiwplwre 2-5 mm, glabrous, in Gynophore A-\l mm, ovary approximately ellip-
fruit 4-12 mm
long, with the pedicel and torus a soid, 1/2 by
1 mm, both glabrous. Fruit globular
1

little incrassate.Mature //7//7 not known, globular, to ellipsoid, 1-1 '/2 cm diam., pericarp thin,
c. Vi1 cm diam., pericarp thin, leathery, smooth or leathery, smooth. Seeds one to few, globular and
Seeds \-A, subglobose to angular,
finely papillate. 5-6 mm
to elliptic and 10 by 7 by 5 mm.
9-10 by 7-9 by 4-5 mm. Distr. India (Sikkim, Khasia, and Assam),
Distr. Indo-China (Annam), in Malaysia: Burma (Dawna Range), S. China, Hainan, Indo-
S. Sumatra (Lampongs), Malay Peninsula China, and the Andamans; in Malaysia: Central
(Pahang, Kelantan), Borneo (Sarawak), Java Sumatra, Java, Lesser Sunda Islands (Bali,
(scattered). Lombok), Celebes, Philippines (Mindanao),
Ecol. Forests, often on limestone, up to 600 m. Moluccas (Sula Is: Mangoli, Burn). Fig. 10.
Fl.Dec-June,//-. July-Nov. Apparently a rare plant. Ecol. Forests and forest edges, frequently in the
Vern. Endog-dogan, J, lorteloran, Md. shade, seems to prefer moist places. Fl. fr. Jan.-
Note. The material of C. paniciilata Ridl. is Dec.
somewhat different from the Javanese C. erycibe In the Himalaya, Sumatra, and Java it is found
by its being less hairy, by the longer pedicels, the between (700-)l200 and 1750(-2000) m. In Hong
glabrous sepals, the more orbicular petals, and the Kong and vicinity it occurs near sea-level, and the
smaller number of stamens; its fruit is unknown. same is probably the case in the Philippines and
The only Bornean specimen, collected by Havi- the Sula Is.
LAND (Sar), and the only Sumatran specimens,
Forbes 1696 and 1719A, agree quite well with the
type of C
paniculata.

5. Capparis cantoniensis Lour. Fl. Coch. (1790)

331; ed. Willd. (1793) 404; DC. Prod. 1 (1824)
253; Merr. Comm. Lour. (1935) 173.— C. salac-
censis Bl. Bijdr. (1825) 54; MiQ. Fl. Ind. Bat. 1,
2 (1858) 101; Illustr. (1870) 23, t. 12A (excl. var.
celebica Miq., quae est C. lanceolaris); Back.
Schoolfl. (1911) 61; Koord. Exk. Fl. Java 2
(1912) 294; Ridl. J. Fed. Mai. St. Mus. 84 (1917)
15; Back. Bekn. Fl. Java (em. ed.) 4A (1942) fam.
45, p. 9. —
C. piimila Champ, in Hook. J. Bot. Kew
Card. Misc. 3 (1851) 260; Hook./. & Th. Fl. Br.
Ind. 1 (1872) 177; Gagn. Fl. Gen. L-C. 1 (1908)
188.— C. hasseltiana Miq. Illustr. (1870) 24, t. 13.
— C. celebica MiQ. I.e. 26.
Slack climber, 3-20 m. Twigs almost straight,
angular and puberulous when young, terete and Fig. 10. Distribution of Capparis cantoniensis
glabrescent when older; internodes c. '/2-4 cm.
1 Lour. The three areas north of Malaysia form
Thorns patent to recurved, 2-5 mmlong, es- one whole.
pecially on flowering branches minute or wanting.
Leaves subcoriaceous, oblong to lanceolate, Vern. Endog endogan, kidjeruk, segore tjalut
(2.3-)2|/2-4(-5) times as long as wide, sometimes (?), sanik lakik, Md.
sigar djalak, J,

ovate, rarely obovate, above glabrous, beneath Notes. In the two Lombok specimens, collected
sparsely puberulous but soon glabrescent, young by Elbert, the leaf-top is rounded to acutish.
leaves salmon-reddish, (4-)5i/2-10'/2 by (11/2-) A few specimens from Mindanao were originally
2-3'/2(-4) cm; base obtuse to acutish, top narrow- identified as C. sepiaria, and indeed show resem-
ed, acuminate, tip often blunt, sometimes slender blance to that species, as the leaves are compara-
Dec. 1960] Capparidaceae (Jacobs) 77

lively small with a slightly notched top. On ac- Scandent shrub, few (up to 20?) m
high, rarely
count of the midrib, however, which is narrowly self-supporting and c. '/2 rn high. Twigs mostly
1

sulcate, I reckon them to belong here. They are overhanging, straight, fulvous to red-brownish
from altitudes below 300 m. puberulous when young, sooner or later (rarely
not) glabrescent but nearly always vestiges of the
6. Capparis longestipitata Heine, Mitt. Bot. pubescence persistent near the leaf axils; inter-
Staatssamml.Munch. Heft 6 (1953) 210; Pfl. nodes c. 2-5 cm. Thorns mostly present, recurved,
Samml. Clemens (1953) 41. up to 7 mm long. Leaves subcoriaceous, above
Young
parts with a greyish, very short in- glabrous, beneath mostly glabrous, sometimes
dument, glabrescent; twigs terete, straight, inter- fulvous-puberulous, whether or not glabrescent,
nodes c. '/2 cm; thorns hardly or not developed.
1 (1.5-)2.0-4.0(-4.7) times as long as broad, broad-
Mature leaves firmly herbaceous, glabrous, ob- est in the middle, (3>/2-)6'/2-12(-16) cm long,
long to slightly obovate, c. 5-6 by 2'/2(-3) cm, (l%-)2-4(-7) cm broad; base acutish to rounded
base rounded, top acuminate, tip 4-7 long, mm or subcordate, top acuminate, rarely rounded to
acute; midrib above sulcate in the basal half, subemarginate, tip mostly acute-mucronate, marg-
otherwise flat; nerves 5-7 pairs, thinly prominent in often markedly recurved, especially towards the
on both sides, reticulation distinct; petiole c. top; midrib above sulcate mostly all over, rarely
1 cm long, hairy as the twig. Subumbels c. 15- flat; nerves 6-10 pairs, hardly visible; petiole
flowered, in the axils of the higher leaves and ('/4-)%-l(-l Vi) cm, hairy as the twigs. Subumbels
some terminal, c. IV2-2V2 cm peduncled. Pedicels mostly axillary, sometimes terminal, mostly
c. 8-15 mm, on small distinct cushions; bracts simple, sometimes branched; peduncle 2-7 cm,
subulate, few mm
long, very soon caducous. Buds occasionally leafy and up to 15 cm, more or less
globular, c. 3 mm
diam. Outer pair of sepals c. glabrescent. Pedicels (1 V4-)2-2'/2(^-3) cm, gla-
3 mm diam., outside densely greyish puberulous, brous, sometimes each with a pair of distinct
the inner ones smaller and flattish and hairy only thorns at the base. Bracts small, caducous, narrow,
in the centre. Petals c. 4 by 2 mm, glabrous, white. hairy. Buds globular, 5-6(-8) mm
diam. Flowers
Stamens c. 20-30, c. 1 mm
long. Gynophore white, yellow-white, pink or red, whether or not
20-25 mm, glabrous; ovary spindle-shaped, sub-
ovoid, 1 '/2 mm
long, glabrous with a knob-shaped
stigma. Fruit unknown.
Distr. Indo-China (Nhatrang in S. Annam,
once coll.); in Malaysia: North Borneo (Mt Kina-
balu, once coll.).
Ecol. In forest, at 1000 and 1500 respectively. m
Note. The Indo-China specimen, Chevalier
38671, differs slightly in having thorns 3-4 mm,
the leaves ^U-\^l2 cm acuminate and brown-pu-
berulous on the midrib underneath, the sepals
5-5',., by 3'/2-4 mm, the petals 6 by 2 and pu- mm
berulous inside, -^ 18 stamens 1/2-2 cm, and the 1

gynophore 3 cm long.
I have compared C. longestipitata with C.
cantoniensis but its gynophore is longer, its
midrib is only grooved in the basal half, and it has
hardly any thorns. From C. lanceolaris it differs by
the far smaller flowers. Fig. 11. Distribution of Capparis lanceolaris DC.

7. Capparis lanceolaris DC. Prod. I (1824) 248; fragrant. Sepals c. 6-7(-10) by 5 mm, with mem-
Mio. Fl. Ind. Bat. 1, 2 (1858) 101 Back. Schoolfl. ; branous margin, glabrous; outer pair herbaceous,
(1911) 62; KooRD. Exk. Fl. Java 2 (1912) 294; inner pair flatter and thinner, rarely minutely
Back. Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, ciliate.Petals obovate, somewhat oblique, thin,
p. 9. — C. subspiuosa RoxB. Fl. Ind. ed. Carey 2 8-11 by 4-6 mm, puberulous towards the base,
(1832) 568.— C. roxbitrghii (non DC.) Span. especially inside. Stamens c. 20 (-40), 2-3 cm.
Linnaea 15 (1841) 166. —
C. elliptica Span. I.e. — Cv/76i/?/7o/e(2-)3-4(-5) cm, ovary ellipsoid, 1-2 mm
C. platyacanlha TuRCZ. Bull. Soc. Nat. Moscou long, both glabrous. Fruits few; pedicel, torus, and
27, 2 (1854) 323.— C. calloplnlla (non Bl.) Miq. gynophore but little incrassate. Fruit (sub)globu-
PI. Jungh. (1855) 397; Fl. Ind. Bat. 1, 2 (1858) 101, lar, 2y4-3y2 cm, bluish black. Seeds 3 or more, c.
p.p. —C salaccensis var. celehiea Miq. Illustr. 8 by 6 by 5 mm.
(1870) 23, t. 12b.— C. oblongata Merr. Gov. Lab. Distr. Malaysia: M.-E. Java (also Nusa Ba-
Publ. Philip. //. 35 (1906) 15; Philip. J. Sc. (1906) 1 rung), Madura, Lesser Sunda Islands (Timor,
Suppl. 58; En. Philip. 2 (1923) 212.— C. copelandii Damar, Jamdena), Celebes (also Salajar and
Elmer, Leafl. Philip. Bot. 2 (1910) 680; Merr. En. Buton), Philippines (not known from Palawan),
Philip. 2 (1923) 210.— C. torricellensis Laut. Bot. Moluccas (Ceram, Nusa Laut), New Guinea (N.
Jahrb. 52 (1914) 112.— C. viridis Elmer, Leafl. part from Manokwari to the Sepik and Papua,
Philip. Bot. 8 (1919) 3076. also Schouten I.), New Ireland. Fig. II.
n Flora Malesiana [ser. I, vol. 6^

Ecol. Secondary (or primary) forests, thickets, arranged in a terminal panicle up to 15 by 10 cm,
or hedges, mostly on dry, calcareous, rocky soil, with some additional smaller axillary panicles.
also in coastal vegetation, up to 700 m, once at Bracts linear, 2-6 mm, sooner or later caducous.
1650 m. Buds globular, 3-4 mm
diam. Sepals 2-4 by
Vern. Gagaan, tjantelan, J, kengkeng, Md, 1 1/2-2
'/2 mm, ovate, patent and persistent for
nonoh mukeh, Timor; Philippines: sulu-siilu. Bag., some time after anthesis, outer ones patent, not or
kamit-kabag, Dum. narrowly membranous-margined, inner ones
Notes. In sterile state very similar to C. sometimes broader, with a broad, membranous
floribitnda, but this species has commonly some- margin. Petals very thin, oblong or ovate, 3-5 by
1 1/2-2 mm. Stamens 7-9(-12), 6-8 mm. Gynophore
what longer petioled, mostly ovate leaves narrow-
ing gradually towards the apex, whereas a sharply 4-6 mm (in Ceylon up to 10 mm); ovary ovoid,

acuminate leaf tip is typical for C. lanceolaris, acutish, 1-1 1/2 by 1/2-I mm, both glabrous. In
especially in the Philippines. Some sterile New fruit the pedicel, torus and gynophore only
Guinean specimens resemble C. zippeliana (see slightly thickened. Fruit globular, ± 1 i/2-2(-2i/2)
there). cm diam., soft, fleshy, pericarp coriaceous, smooth
In the Philippines some deviating specimens orange. Seeds 1-3, ± 13 by 10 by 6 mm.
have been described as distinct species, C. oblon-
gata, C. viridis, and C. copelandii. In the first two
the innovations are glabrous to early glabrescent,
the leaf base is obtuse to subcordate, the apex is
narrow and sharply acuminate, the sepals are
9-10 mm long, the inner pair being sometimes
minutely ciliate; sometimes also small thorns are
found in the inflorescence! Besides, in some
specimens identified as C. oblongata, the sub-
umbels are conferted towards the end of the twigs
and are merely subtended by minute puberulous
bracts, resulting in impressive inflorescences
10-15 cmlong. This latter feature, however,
varies even in different duplicates of the type
collection (R. Meyer FB 2632). Two other
Philippine specimens have densely pubescent
twigs and leaf underside; this material (in fruit)
was described as C. copelandii.
The specimens from Java are characterized by
generally well developed thorns which are also
Fig. 12. Localities of Capparis floribunda Wight
found in the inflorescence and by comparatively
(•) and of its/, induta Jacobs ( + ). The locality
small leaves (31/2-8 cm).
near the Malay Peninsula (Koh Ha) is approx-
imate. Also in the Deccan and Formosa.
8. Capparis floribunda Wight, 111. Ind. Bot. 1
(1840) 35, t. 14; Hook. /. &
Th. F1. Br. Ind. 1
(1872) 177; F.-Vill. Nov. App. (1880) 11.— Distr. Ceylon, Deccan, Indo-China (Annam),
Crataeva octandra Blanco, F1. Filip. (1837) 400, Formosa, Peninsular Siam, Andamans; in Malay-
ed. 2 (1845) 280. ed. 3, 2 (1878) 155, nan sia: Java (only Kangean Arch.), Philippines
Capparis octandra Jacq. —
C. luzonensis TuRCZ. (Debangan I. near Palawan, Babuyan I., Luzon),
Bull. Soc. Imp. Nat. Mosc. 27,2(1854) 324; Merr. Moluccas (Halmaheira, Tidore, Sula Is, W.
Sp. Blanc. (1918) 159; En. Philip. 2 (1923) 212, Ceram, Kai Is). Fig. 12.
incl. var. ampla Merr. I.e. —
C. andamanica King, Ecol. This showy plant seems to prefer dry
Ann. R. Bot. Gard. Calc. 5 (1896) 119, t. 137. country and rocky soil, sometimes in coastal
— C. oligostema Hay. Ic. PI. Form. 3 (1913) 22. vegetation, in the lowlands; once at 1600 m. Fl.fr.
Jan. -Dec.
/. floribunda. Elmer found "many fruits opened and seeds
Young parts glabrous, rarely very soon glabres- eaten by birds".
cent.— Shrub or climber (?), few m high. Twigs
straight or slightly zig-zag, terete, smooth, inter- /. induta Jacobs, /. nova.
nodes 2-5 cm; thorns small, recurved, mostly Af. floribunda difl'ert indumenta denso fulvo
wanting. Leaves firmly herbaceous, broadest at puberulo in partibus juvenilibus.
the middle or sometimes below, mostly narrowed A dense, fulvous-puberulous indument on
in the upper half, mostly 2'/2-3 times as long as twigs, young leaves, and inflorescences. Outer
wide, (4-)6-10(-13) by (1 1/2-)2!/2^(-6) cm; base sepals at the base somewhat hairy and petals
mostly rounded and more or less acute, top slightly ciliate.
variable; midrib above often narrowly sulcate; Distr. Indo-China (Annam); in Malaysia:
nerves 7-9 pairs; margin often slightly recurved; E. Java (Surabaja Residency, Muning, fl. ix-1927,
petiole 1/2-1 '/2(-2) cm. Flowers numerous, white, DE VooGD 673 (L, type; Bo)).
fragrant, in small subumbels, c. 1-2 cm stalked, Vern. Wangon lanang, J. Fig. 12.
Dec. 1960] Capparidaceae (Jacobs) 70

Notes. The species is characterized by the primary forests, bamboo thickets, etc., up to
glabrous appearance, with small or without thorns, 600 m.
the large number of small flowers with 8 stamens, Vern. Manungal-lalaki, Mindoro.
subpersistent sepals, and a short gynophore which Notes. A variable species which can always be
is more vigorous than in C. sepiaria; the latter distinguished from C. lanceolaris by the straight,
species has also a lighter coloured fruit. though sometimes little developed thorns.
The only specimen known from the mainland Capparis palawanensis Merr. represents a form
of Java is the specimen described as /. induta with unusually large leaves, but transitions with
Jacobs. In sterile state it can look like C. lanceo- the average leaf-size occur. C. littoralis Merr.
laris (see there). represents obviously a glabrous form; in some
The leaves are variable in width. Turczaninow specimens of typical lobbiana the leaves are also
described C. luzonemis on the specimen with the soon glabrescent. There is but poor material of
narrowest leaves that I have seen, the leaf index C. ilocana Merr.; the type and another specimen
being 3.7-4.1. Merrill's var. ampla actually agree mutually very well and seem to represent
represents the normal leaf-shape; intergrading another glabrous paramorph with almost abortive
specimens are by no means rare. thorns and subcoriaceous leaves.

9. Capparis lobbiana Turcz. Bull. Soc. Nat. 10. Capparis sepiaria Linne, Syst. Nat. ed. 10, 2
Moscou 27, 2 (1854) 323; Rolfe, J. Bot. 23 (1885) (1759) 1071; Sp. PI. ed. 2, 1 (1762) 720; Burm. /.
210; ViDAL, Rev. PI. Vase. Filip. (1886) 47.— Fl. Ind. (1768) 118; DC. Prod. I (1824) 247,
C. sepiaria var. acuta Vidal, I.e. —C
littoralis inch ^ glabrata; Roxb. Fl. Ind. 2 (1832) 568; W.
Merr. Philip. J. Sc. 7 (1912) Bot. 270; En. Philip. & A. Prod. 1 (1834) 26; Decne, Nouv. Ann. Mus.
2 (1923) 211.— C. loheri Merr. Philip. J. Sc. 7 Hist. Nat. Paris 3 (1834) 427; Herb. Timor.
(1912) Bot. 270; En. Philip. 2 (1923) 21 1.— C. pala- Descr. (1835) 99; Span. Linnaea 15 (1841) 166;
wanensis Merr. Philip. J. Sc. 10 (1915) Bot. 304; Jacquem. Voy. Ind. Bot. 4 (1844) 21, t. 24; Gray,
En. Philip. 2 (1923) 212.— C. ilocana Merr. U.S. Expl. Exp. Bot. 1 (1854) 70; Miq. Fl. Ind.
Philip. J. Sc. 13 (1918) Bot. 13; En. Philip. 2 Bat. 1, 2 (1858) 101; Sum. (1860) 159; Illustr.
(1923) 211.— Fig. 13. (1870) 27; Hook./. &Th. Fl. Br. Ind. 1 (1872) 177,
Climber l'/2-3(-4) m high. Twigs slender, incl. var. vulgaris; KuRZ, Fl. Burm. 1 (1877) 66;
straight or slightly zig-zag, with leaves not seldom Fern.-Vill. Nov. App. (1880) 11, cur?i var.;
in two rows, densely clothed with up to '/2 mm Naves in Blanco, Fl. Filip. ed. 3 (1880) t. 209;
long, straw-coloured hairs, rarely almost glabrous ViDAL, Phan. Cuming. (1881) 94; Sinopsis Atlas
or soon glabrescent; internodes l-2(-3) cm. (1883) 13, t. 6 f. A; Hemsl. Bot. Chall. 3 (1884)
Thorns almost straight, up to 2, rarely 5 mm, thin, 120; Vidal, Rev. PI. Vase. Filip. (1886) 47; King,
dark, with lighter base, pointing upwards, ex- J. As. Soc. Beng. 58, ii (1889) 393; KuRZ ex
ceptionally downwards. Leaves herbaceous, broad- Prain, J. As. Soc. Beng. 59, ii (1890) 277, incl. var.
est below, rarely at the middle, (1. 5-) 1.8-2. grandifolia; ibid. 60, ii (1891) 302; ibid. 62, ii
(^.3) times as long as wide, glabrous or sparsely (1893) 65; Back. Fl. Bat. (1907) 59; Merr. Philip.
hairy above, hairy beneath, especially on the J. Sc. 3 (1908) Bot. 77; Gagn. Fl. G6n. I.-C. 1
nerves, 4-8(-15) by 2-3'/2(-5) cm; base cuneate, (1908) 191 Back. Schoolfl. (191 1) 61 Ridl. J. Str.
; ;

sometimes rounded, rarely acute, top narrowed, Br. R. As. Soc. «. 59 (1911) 68; Koord. Exk. Fl.
acute to (rarely long-) acuminate, often with a Java 2 (1912) 294; Merr. Fl. Manila (1912) 215;
nerf-tip; midrib above shallowly sulcate in the Ridl. Fl. Mai. Pen. 1 (1922) 122; Merr. En.
lower part, yellowish, sparsely hairy underneath; Philip. 2 (1923) 212, excl. var. acuta Vidal in
nerves 6-8 pairs; petiole 2-4(-12) mm, hairy as are synon., quae est C. lobbiana; Craib, Fl. Siam. En.
the twigs. Subumbels axillary; peduncle '/2-6 cm, 1 (1925) 83; C. T. White, J. Arn. Arb. 10 (1929)
thin, bearing sometimes one or more small leaves, 217; Gagn. Fl. G^n. I.-C. Suppl. 1 (1939) 165;
hairy. Pedicels slender, '/2-3 cm, mostly with
1 Back. Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45,
distant hairs, rarely glabrous. Bracts inconspicu- p. 9.— C. umbellata R. Br. ex DC. Prod. 1 (1824)
ous, narrow. Buds globular to depressed-globular, 247; C. T. White, Proc. Roy. Soc. Queensl. 34
5-6 mm diam. Flowers white or pale pink. Sepals (1922) 31.— C. emarginata Presl, Rel. Haenk.
persistent for a short time, outer pair 5-6 mm 2 (1835) 85, non A. Rich.; Fern.-Vill. Nov. App.
diam., herbaceous, hairy (or rarely glabrous), (1880) II; Merr. En. Philip. 2 (1923) 211.—
inner sepals 6-7 mm
diam., thinner, ciliate. Petals — C. retusella Thw. En. PI. Zeyl. (1864) 16
very thin, ovate, 2-4 by 21/2-3 mm, ciliate. Stamens = C sepiaria var. retusella Thw. I.e. 400; Fern.
15 to 60, 8-12 mmlong, anthers 1-1 1/2 by V2-I -ViLL. Nov. App. (1880) 11. — C. subacuta Miq.
mm. Torus obscure. Gynophore 1 '/2-3(-43/4) cm, Illustr. (1870) 35, p.p., quoad specim. halmah.
glabrous; ovary ovoid, c. 2 by 1 mm. In fruit — C. trichopetala Val. Bull. Dep. Agr. Ind.
the pedicel, torus, and gynophore only slightly N^erl. 10 (1907) 14 = C. sepiaria var. trichope-
incrassate. Fruit globular, c. 1/2 cm diam., peri- tala Val. I.e. 72; Laut. Bot. Jahrb. 52 (1914)
1

carp leathery, thin, smooth, glossy, blackish. 112.— C. affinis Merr. Philip. J. Sc. 10 (1915)
Seeds few, subellipsoid, c. 1/2 cm long. Bot. 303; En. Phihp. 2 (1923) 210.
Distr. Malaysia: Endemic in the Philippines Wide, much-branched shrub, few m high, some-
(not known from the Sulu Arch.). times climbing. Young shoots densely fulvous or
Ecol. Prefers dry, rocky conditions, also in greyish puberulous, sooner or later glabrescent;
80 Flora Malesiana [ser. I, vol. 6^

Fig. 13. Capparis lobbiana Turcz. a. Branch with fruits and flowers, X 2/3, b. bud, X 3 (Williams 382
and BS 85241).
Dec. 1960] Capparidaceae (Jacobs) 81

twigs stout, zig-zag, terete; internodes \Y2-iV2 Ecol. Drier places in thickets, hedges, teak-
cm. Thorns generally vigorous, recurved, 3-5 mm forests, etc., in the lowlands, often near the seaside,
long. Leaves firmly herbaceous to subcoriaceous, solitary or in groups, obviously bound to seasonal
when dry mostly greyish green (in some Philippine climatic conditions. When in fruit, often a good
specimens brownish), elliptic, sometimes obovate, deal of the leaves is shed.
or ovate, exceptionally linear, hairy as the twigs, Vern. Poka(n), Md.
upper surface glabrescent first, often with scattered Uses. The plant is said to be have some medi-
minute warts, (l'/2-)1.8-2.3(-4) times as long as cinal use {cf. QuisuMBiNG, Med. PI. Philip. 340).
wide, (1 1/2-)3 1/2-5 1/2(-10) by (1-)1 i/2-2i/2(-4) cm; Notes. It seems not unlikely that this species
top mostly rounded, nearly always notched, occurs also in Africa under the names corym- C
rarely blunt; midrib flattish above, sometimes bosa Lamk and C. tomentosa Lamk, but this
slightly sulcate at the base; nerves 4-6(-8) pairs, question deserves further research.
not very distinct; petiole 2-4(-7) mm. Subumbels In the eastern part of Malaysia specimens are
few- to 20-flowered, at the end of small lateral found which are intergrading between typical C.
twigs, rarely terminal. Bracts small, hairy, early sepiaria and the Australian C. umbellata; they
caducous. Pedicels Y^-2(-2y2) cm, glabrous. Buds have straighter stems with less developed or often
globular, 4(-5) mm
diam. Sepals ovate, 4-6 by wanting thorns, larger, ovate, soon glabrescent
3-5 mm, occasionally finely ciliate, outer pair leaves, fewer flowers in a shorter stalked and more
herbaceous with narrow membranous margin; often terminal and (sub)sessile subumbel. In a few
inner pair somewhat smaller, very thin, mem- specimens the midrib is sulcate all over.
branous towards the margin. Petals AVj-lVi by C. sepiaria and C. cantoniensis are easily dis-
1 "4-3 mm, very thin, white, more or less pubes- tinguished by the midrib, flattened in the former
cent, especially outside at the base of the upper and narrowly sulcate in the latter.
pair. Torus inconspicuous. Stamens 30-45, 7-12 In one sterile specimen collected byGAUDicHAUD
mm, filaments white, anthers 1 '/^ by 1 mm. near Manila the leaves are extremely narrow, viz c.
Gynophore (4-)6-10(-13) mm, glabrous; ovary 10 times as long as wide (6-7 mm). A similar
ovoid, V2-2 by
1 mm, glabrous. In fruit the torus
1 phenotypic variation was found in C. quiniflora.
and gynophore somewhat incrassate, the pedicel
hardly so. Fruit rather fleshy, (sub-)globular,
11. Capparis diffusa Ridl. J. Str. Br. R. As. Soc.
1-2-seeded, 1-1 V2 cm diam., pericarp subcoriace-
n. 59 (1911)68; Fl. Mai. Pen. 1 (1922) 122;
ous, smooth, whitish-yellowish to almost blackish.
Henders. J. Mai. Br. R. As. Soc. 18 (1939) 35.
Seeds ± 8 by 5 by 4 mm.
Shrub or climber; twigs slender, sparsely
brown-puberulous when young; internodes 2-4
cm. Thorns recurved, 1-3 mm
long. Leaves elliptic,
ovate to obovate, c. 5-8 by 21/2-4 cm; base
rounded to blunt, top blunt, minutely retuse, some-
times subacuminate; midrib above subprominent,
sulcate at the base, brown-puberulous when
young on both surfaces, or glabrous; nerves 5-6
pairs, thin; petiole 3-4 mm, sulcate above, brown-
puberulous when young. Umbel terminal, sessile,
3-5-flowered. Bracts minute, caducous. Pedicels
filiform, 2-5 cm, glabrous. Buds globular, 3-4 mm
diam. Sepals c. 4 mm
long, outer pair glabrous;
inner pair elliptic, ciliate. Petals white, oblong, c.
4-5 mm, hairy inside. Stamens 12-15, c. 1 1/4 cm,
anthers small, white. Gynophore c. 1 1/2 cm,
glabrous; ovary subglobular, acute, c. I long.mm
Fruit unknown.
Fig. 14. Distribution of Capparis sepiaria L. in
Distr. Malaysia: N. Sumatra (P. Weh), Malay
Malaysia.
Peninsula (Perils), twice collected.
Ecol. On (limestone) rocks. Fl. Dec.
Distr. Ceylon, India (Deccan, Sind, Punjab)
to SW. Burma (Diamond I.), Andamans, SE.
China (Kwangtung), Indo-China, Siam, and 12. Capparis callophylla Bl. Bijdr. 2 (1825) 53;
Hainan to Australia (Arnhem Land, Queensland, KooRD. Minah. (1898) 342; Back. Schoolfl.
and New South Wales to c. 32° SL), in Malaysia: (191 62; Bekn. Fl. Java (em. ed.) 4A (1942) fam.
1)
Malay Peninsula (Kedah, Kelantan), North 45, p. 10.— C. tylophylla Spreng. Syst. Nat. 4
Borneo, Java (in W
only twice found at the N. (1827) 204; Miq. Choix (1864) t. 2; Fl. Ind. Bat.
coast), Madura, Kangean Arch., Lesser Sunda 1, 2 (1858) 101; Ulustr. (1870) 22; Koord. Exk.
Islands (Bali, Nusa Penida, Sumbawa, Alor, Fl. Java 2 (1912) 294; Merr. En. Philip. 2 (1923)
Timor, Leti, Babar, Wetar), Celebes (also Salajar 213. C. cumingii Merr. &
Rolfe, Philip. J. Sc. 3
I. and Buton), throughout the Philippines, (1908) Bot. 101.— C. turczaninowii Elmer, Leafl.
Moluccas (Halmaheira, Buru, Ambon, Kai Is), Philip. Bot. 5 (1913) 1755.— C. mucronata Elmer,
New Guinea (near Merauke, S. Papua). Fig. 14. I.e. 1757.— C. robusta Heine, Mitt. Bot. Staats-
82 Flora Malesiana [ser. I, vol. 6^

samml. Munch. Heft 6 (1953) 211; Pfl. Samml. 13. Capparis zippeliana Miq. lUustr. (1870) 25
Clemens (1953) 41. t. Ann. Jard. Hot. Btzg 1 (1876) 5
14; ScHEFF.
Climbing, glabrous shrub, few m high. Twigs K. SCH. & HoLLR. Fl. Kais. Wilh. Land (1889) 49
stout, (reddish-)brown when dry; internodes 2-7 K. ScH. & Laut. Fl. Schutzgeb. (1900) 336
cm. Thorns often wanting, if present recurved, up Laut. Bot. Jahrb. 52 (1914) 114, f. la-d, inch var.
to 5 mm
long. Leaves coriaceous, often reddish novohibernica, f. le, et var. novobritannica, f. If
brown when dry, broadest in the middle, (l'/2-) Pax & HoFFM. in E. & P. Pfl. Fam. ed. 2, 17b
2-3(-3i/2) times as long as wide, (8-)14i/2-20 (1936) 178, f. 90.— C. dahlii Gilg & K. Sch.
(-26) by (3i/2-)5-9(-14) cm; margin slightly re- Notizbl. Berl.-Dahl. 1 (1896) 208; K. Sch. &
curved; base more or less acute to rounded, some- Laut. Fl. Schutzgeb. (1900) 335.— C. carolinensis
times subcordate, top rounded (or emarginate) Kaneh. Bot. Mag. Tokyo 48 (1934) 919, f. 6.
and mostly with a hardened tip or shortly and Small climber. Twigs slender, fairly straight,
bluntly acuminate; midrib above fiat to sulcate; glabrous, rarely glabrescent; internodes 2-5 cm.
nerves c. 6-7 pairs, subprominent above; petiole Thorns mostly wanting, if present slightly recurved,
l-2(-2i^) cm, stout, dark-coloured, rough. In- up to 3 mm. Leaves firmly herbaceous, often dull
florescence a more or less leafy, terminal panicle red-brownish when dry, broadest in the middle,
of subumbels. Pedicels 2-4'/2 cm. Bracts early (1.7-)2.2-2.8(-3.5) times as long as wide, (8-)
caducous. Buds globular, 8-12 mm
diam. Flowers 13-20(-26) by (4i/2-)5-8(-10) cm, glabrous or
white, pink or red-brown. Outer sepals coriaceous, rarely thinly puberulous below; margins sub-
subpersistent, 10-13 by 8 mm, rough; inner sepals revolute; base rounded or acutish or subcordate,
subcoriaceous, 12-14 by 6-10 mm, thinner towards top (rarely sharply) acuminate, or rarely emar-
the margin. Petals oblong to obovate-subspathu- ginate; midrib above mostly sulcate in the basal
late, 22-35 by 6-11 mm. Torus 4-5 mm
broad. part, to flat; nerves 5-9 pairs; petiole {Y^-)
Stamens c. 50-80, filaments light red, 3 1/2-4 1/2 cm, l-l'/2(-l%) cm, sulcate above, glabrous, rarely
anthers 2-3 by mm. Gynoplwre 3-4'/^ cm, ovary
1 glabrescent. Subumbels up to 10-flowered, mostly
ellipsoid, c. 4 by 2-3 mm, both glabrous. In fruit sparsely puberulous, sometimes axillary but
the pedicel (especially towards both ends), the mostly arranged in a terminal panicle, slender,
torus, and the gynophore towards the top, con- 2-7(-12) cm peduncled. Pedicels l-4i/2(-6) cm.
siderably incrassate. Fruit globular to ellipsoid, Bracts small, subulate, hairy, caducous. Buds
5-61/2 cm long, pericarp 2-5 mm
thick, smooth, globular, 5-6(-10) mm
diam. Sepals subpersistent,
yellowish orange. Seeds cv), c. 1 by 1/2 cm. outside sparsely hairy or glabrous, with mem-
D i s t r. Formosa (a variety), in Malaysia: Sumatra branous margin, 5-6(-12) mm
diam.; outer pair
(Palembang), W. Java, Madura, North Borneo, subcoriaceous, one often initially enveloping the
Celebes, and throughout the Philippines. Fig. 1 5. bud for 2/3; inner pair suborbicular, herbaceous.
Except in the Philippines, the species seems to Petals thin, obovate, 6-8 by 21/2-4 mm, white,
be rare. minutely puberulous on both sides, base narrowed,
Ecol. Mostly on dry, often calcareous soil, but top rounded (sometimes crisped?). Stamens c.
also in moister habitats, in secondary forest, etc., 25^5, (li/2-)2i^-3 cm long, white or pale pink;
from the lowland up to 700, in N. Borneo up to anthers 1 1/2-2 1/2 by 1/2 mm. Gynophore 2-3 V^ (-4)
1700 m. Fl. fr. Jan.-Dec. cm, often thinly puberulous at the base, soon
Vern. Manunggal, Mindoro, sani, Minahasa. glabrescent; ovary ellipsoid, 1 1/2 by 1 mm. In
fruit the pedicel, torus, and gynophore are rather
incrassate. Fruit ovoid or ellipsoid, c. 41/2 by 3I/2

cm, top sometimes umbonate, pericarp thin,

leathery, smooth, red. Seeds cvi, c. 1 2 by 9 by 6 mm.
Distr. Micronesia (Palau), in Malaysia: S.
Moluccas (Tanimbar and Kai), New Guinea,
New Britain, New Ireland, and Solomons (Ulawa).
Fig. 20.
Ecol. Primary and secondary rain-forests,
sometimes on rocky soil, below c. 1200 m alt.
Fl. fr. Jan.-Dec.
Vern. Dschiriguh, Constantinhafen, wendos,
Manokwari, walakenaru, Solomons.
Note. In the sterile state much resembling C.
lanceolaris, though the different average leaf-size
is generally a good character; besides, C. lan-
ceolaris is pubescent on the mature twig near the
leaf-insertion which is glabrous in C. zippeliana.

14. Capparis pubiflora DC. Prod. 1 (1824) 246;

Decne, Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834)
246; Herb. Timor. Descr. (1835) 98; Deless. Ic. PI.
Fig. 15. Localities of Capparis callophylla Bl. ;
3 (1837) t. 12; Span. Linnaea 15 (1841) 165; MiQ.
a variety in Formosa. Fl.Ind.Bat. 1,2(1858) 100; Illustr. (1870) 27, t. 15,
Dec. 1960] Capparidaceae (Jacobs) 83

inch var. sumatrana et var. moluccana MiQ. I.e. long. Sepals herbaceous, concave, 41/2-7 by
28; King, J. As. Soc. Beng. 58, ii (1889) 394, incl. 2'/2^ rnm, outside puberulous, outer pair acute
var. perakensis ScoRT.; Hall./, in Koord. Minah. or blunt, sometimes slightly cucullate, inner pair
(1898) 343; Ridl. J. Fed. Mai. St. Mus. 10 (1920) with blunt or rounded top. Petals thin, mostly
129; Back. Bekn. Fl. Java (em. ed.) 4A (1942) obovate, 7 — 10 by 3-4 mm, outside soft-hairy at
fam. 45, p. 13. —
C. nigricans Span. Linnaea 15 top and margins. Torus conical, disk c. 1 mm.
(1841) 165; MiQ. Fl. Ind. Bat. 1, 2 (1858) 100.— Stamens 20-30(-50), filaments 15-20(-25) mm,
C. cerasifolia A. Gray, U.S. Expl. Exp. Bot. 1 anthers elliptic, by '/2 mm. Gynophore 15-25 mm,
1

densely tomentose, in fruit glabrescent; ovary

ellipsoid, IV2 by V2 mm, hairy as the gynophore,
1

the small, knob-shaped stigma glabrous. Pedicel

and gynophore not much incrassate in fruit.
Fruit subellipsoid, 12-21 by 10-19 mm, shortly
umbonate, pericarp leathery, verruculose, glabrous,
black, sometimes with a reddish or bluish tinge.
Seeds 5-15(-25), c. 6 by 5 by 4 mm, smooth.
Distr. Siam, Indo-China, Hainan, in Malaysia:
Central & S. Sumatra (Gajo Lands, once collect-
ed), Malay Peninsula (Perak, Kelantan, Pahang,
Selangor), North Borneo, Java (E of Lembang,
also Nusa Barung), Madura, Lesser Sunda Islands
(Bali, Lombok, E. Sumbawa, Timor), Celebes,
Philippines (Palawan, Luzon, Mindanao with
Pujada I.), Moluccas (Ceram, Ambon, Halma-
heira), New Guinea (Vogelkop). Fig. 18.

Fig. 16. Cappari.s piibifiura DC. (Cult. Hort. Bog.

XV-J-B-II-6; Jacobs, 1956).

(1854) 71 C. Mueller in Walp. Ann. 7 (1868) 189

;

{crassifolia,sphalma); Merr. Philip. J. Sc. 3 (1908)

Bot. 77; En. Philip. 2 (1923) 210.— C. brachyscias
TuRCz. Bull. Soc. Nat. Moscou 27, 2 (1854) 323.
— C. lasiopoda Turcz. I.e. 'ill; Fern.-Vill. Nov.
App. (1880) 11; ViDAL, Phan. Cuming. (1885) 94;
Rev. Pi. Vase. Filip. (1886) 48.— C. myrioneura
Hall. /. in Fedde, Rep. 2 (1906) 60, p.p.—C.
dealbata {non DC.) Back. Schoolfl. (1911) 62.—
C. perakensis (Scort. ex King) Ridl. Fl. Mai.
Pen. 1 (1922) 124.— C. borneensis Merr. PI. Elm.
Born. (1929) 91.—Fig. 16-17.
Shrub 2-5 m
high. Young shoots densely
fulvous-tomentose, soon (rarely late) glabrescent;
twigs often zig-zag, terete; internodes 1-3 cm.
Thorns straight or slightly curved upwards,
patent, 3-6 mm
(wanting or very small in the
Philippine specimens). Leaves herbaceous to sub-
coriaceous, glabrous with yellowish nerves,
(1.7-)2. 7-3(^.4) times as long as wide, often
obovate, rarely ovate, (5-)8-16(-27'/2) by 2 1/2-6
(-9) cm; base acute to blunt, sometimes rounded,
top acuminate, tip up to 2 cm, acute to blunt;
midrib sometimes sulcate above; nerves (6-)7-9
(-13) pairs, reticulations distinct; petiole 5-8
(-11) mm. Flowers either l-5(-10) in short,
axillary racemes, with a persistent, densely tomen-
tose, subulate bract and 2 smaller basal bracteoles, Fig. 17. Capparis pubiflora DC,
flower, a. Corolla,
or sometimes this inflorescence replaced by a front view, stamens and gynophore removed, X '/j,
short young twig with axillary flowers; racemes b. calyx (one sepalremoved) and disk, x 2,
and young twigs with a number of minute subulate c. calyx and corolla from the back, x 21/2,
bract-like leaves at the base. Pedicels V^-3(-5) d. flower, near-longitudinal section, X lYi (after
cm, glabrescent. Buds globular to ovoid, c. 5 mm living plant. Cult. Hort. Bog. XV-J-B-II-6).
84 Flora Malesiana [ser. I, vol. 6^

Ecol. More or less dry places, such as hedges, Shrub, sometimes climbing, IVi-^-Vii-'^Vi) m;
roadsides, teak forests, brushwood, jungle, up to twigs straight, terete, with minute, stellate,
400(-600) m. Fl. June-July, fr. March. ferruginous hairs, glabrescent; internodes 2-5 cm.
Vern. (Daun) poka, saneg-sanegan, Md, Thorns patent, straight or slightly curved upwards,
djenggotan, waan-waanan, J, bangol bangol, Bali; l-3(-4) mm. Leaves on lateral branches mostly
Minahasa: maha-limu, Ratahan, malemo, Ton- distichous, more or less firmly herbaceous,
sawang, tiitiuiean woring, Tontemboan, sahamiiu- elliptic, when young with minute, stellate, fulvous-
tei; wama pusu, Halmaheira, holibui, Mbo. ferruginous indument giving it a farinaceous as-
Notes. Some specimens from Sumatra and the pect, glabrescent, (1.2-)1.7-2.2(-3) times as long
Malay Peninsula, and generally those from as wide, often ovate, sometimes obovate, 5-9 Vi
Celebes, the Moluccas, and the Philippines, have (-15) by IVi-AVii-^Vi) cm; base rounded to blunt,
markedly larger, obovate leaves; they cannot be top acuminate with a mostly blunt tip Vz-^Vi cm
long; nerves yellow to light-brown, midrib some-
times sulcate above; nerves c. 5 pairs, veins reti-
culate; petiole c. 1/2 cm, densely greyish brown
tomentellous. Flowers 2-4, serial. Pedicels (1-)
XVi-li-lVi) cm, thin, densely hairy, glabrescent
except for the somewhat broadened top. Buds
globular, 4—5 mm diam. Sepals elliptic-ovate, 4-5
by 2'/2-4 mm, minutely hairy outside, 3-nerved,
outer pair (especially the posterior one surround-
ing the disk) slightly larger and more obtuse than
the inner. Petals elliptic to oblong, 6-8 by 2-4 mm,
very thin, on both sides floccose-hairy, white
tinged pale yellow, green, or violet, upper pair
mostly slightly smaller than the lower pair, the
base thickened with a mostly yellow-coloured,
later red honey-guide. Disk bilobed, fleshy,
roundish, up to 2 mm diam. Stamens c. 20,
15-23 mm long, filaments pale, anthers 1 mm,
sordidly blue. Gynophore 18-20(-25) mm; ovary
1 by 34 mm, stigma obtusely conical, Vi rnm high,

Fig. 18. Distribution of Capparis pubiflora DC. both glabrous. In fruit neither the pedicel, nor the
gynophore much incrassate. Fruit about globular,
distinguished taxonomically from the population 8-12 mm diam.; pericarp minutely rugose when
in Java and the Lesser Sunda Islands. dry, glossy, black when ripe (once reported red),
C. Mueller, I.e., copied A. Gray's name glabrous. Seeds 2-6, 6 by 3-4 by 2 mm, smooth,
C. cerasifolia as C. crassifolia. Pax Hoffmann & glossy brown.
continued this mistake in a remarkable way when
stating: "Die Ueberleitung zu den Philippinen
bilden die 2 Arten der Sulu-Insein: C. cerasifolia
A. Gray und C
crassifolia A. Gray" (in E. & P.
Nat. Pfl. Fam. ed. 2, 17b, 1936, 178).

15. Capparis pyrifolia Lamk, Encycl.

Bot. 1 (1785)
606, quoad r^; DC. Prod. (1824) 246; Deless.
1

Ic. PI. 3 (1837) t. 1 nou W.

1 ; &
A. Prod. 1 (1834)
25, quae est C. grandiflora Wall, ex Hook. /. &
Th. — C. acuminata Willd. Sp. PI. 2 (1799)
1131; DC. Prod. 1 (1824) 247; Hook./. Th.&
Fl. Br. Ind. 1(1872) 178; Back. Fl. Bat. (1907) 56;
Voorl. (1908) 13; Schoolfl. (1911) 63; Koord.
Exk. Fl. Java 2 (1912) 293; Back. Bekn. Fl. Java
(em. ed.) 4A (1942) fam. 45, p. 7. —
C. zeylanica
{nan L.) DC. Prod. 1 (1824) 247, quoad specim.
javan.; W. & A. Prod. 1 (1834) 25.— C. foe tida
Bl. Bijdr. 2 (1825) 52; Miq. Fl. Ind. Bat. 1, 2
(1858) 99; Gagn. Fl. Gen. I.-C. I (1908) 184;
Craib, Fl. Siam. En. 1 (1925) 80; Gagn. Fl. Gen.
I.-C. Suppl. 1 (1939) 161.— C. dasypetala Turcz. Fig. 19. Distribution of Capparis pyrifolia Lamk.
Bull. Soc. Nat. Moscou 27, 2 (1854) 322.— C.
oxyphylla Miq. PI. Jungh. 4 (1855) 397; Fl. Ind. Distr. Siam, Indo-China (Annam, Cochin-
Bat. 1, 2 (1858) 100.— C. horrida {non L. /.) Miq. china); in Malaysia: N. Sumatra (Laubalang?),
Illustr. (1870) 34, pro var. cc pro parte et synon. C. Java, Kangean Arch., Madura, Lesser Sunda
foetida et C. oxyphylla. Islands (Bali, E. Sumbawa). Fig. 19.
Dec. 1960] Capparidaceae (Jacobs) 85

Ecol. In the lowlands and hills in dry places, Ecol. Forests, jungle, along rivers, from the
in teak forests, brushwood, hedges, on limestone lowland up to c. 1600 m. Fl. fr. Jan. -Dec.
hills, fairly common in Java, up to c. 850 m. Fl. Vern. Dunggol manok, Sarawak.
especially Sept., no flowers collected Febr.-May; Uses. Two collectors mention the fruit(pulpj
//•. July-Febr. to be edible.
Vern. Kaju tudjiih, S, gagahan, gedangan, Note. In vegetative characters much resembling
kedeling, lorowan, risini, tjantelan, waan-waaiwn, C micracantha ssp. korthalsiana but distinguished
wcin-uwanan, J, sanek, sanik-lakek, Md, kaluang- by the recurved thorns, the narrower leaves with
kaloangan, pokak-pokadn, Kangean. long and slender tip, and the nerves being more
Uses. According to Heyne (Nutt. PI. 1927, prominent towards the margin.
682) the white wood is sold in Djakarta as 'kaju
tudjuh' against bile and stomach ache, and an 17. Capparis cucurbitina King, J. As. Soc. Beng.
extract of the rasped wood is taken against 58, (1889) 395; Ann. R. Bot. Card. Calc. 5 (1896)
ii

dizziness. Fruit once reported to be sweet and 119, t. 136; RiDL. Fl. Mai. Pen. 1 (1922) 124.
edible. Scandent, 6-10 m
high; twigs slightly zig-zag,
Notes. cannot always easily be observed
It (nearly) glabrous; internodes c. 2 cm. Thorns
that the flowers are serial because the row is recurved, 2-3 mm
long, sharp. Leaves herbaceous,
sometimes very short. This could lead to con- obovate, c. 21/2 times as long as broad, glabrous,
fusion with C
pubifiura, which is characterized, c. 9- 18 1/2 by 31/2-71/2 cm; base generally rather
however, by the hairy gynophore and a fruit over abruptly rounded and acute, top rounded and
12 mm
diam. rather abruptly acuminate, tip narrow and acute,
1-1 1/2 cm; midrib above sulcate in the basal part;
16. Capparis buwaldae Jacobs, sp. nov. nerves 6-8 pairs, thinly subprominent above,
Glabra. Spinae stipukires breves, recurvatae. reticulations distinct on both sides; petiole
Folia 6± mm oblongata, apice acute
petiulata, Vi-Va cm. Flowers in mature state unknown, pale
acuminata; costa nervique majores supra sulcati. green, yellow inside, 2-3, serial. Pedicels 2-3 cm.
Flores 2-4 in serie supraaxillare dispositi, minores; Buds ovoid, acute, 4 mm
long. Sepals ovate-
alabastra globosa acuta 3 mm
diametro; pedicellus lanceolate, acuminate. Petals broadly elliptic,
gynophoriumque fructiferum vix incrassatum. Fruc- obtuse. Stamens c. 20. In fruit neither the pedicel
tus globosus vet ellipsoideus, umbonatus, 2-4 cm nor the torus incrassate, the gynophore 11-17 mm,
longus, irregulariter tuberculatus; semina majora. slightly thickened, obviously articulated with the
Typus: Hallier /. 2573 (Bo, holotype; L, iso- fruit. Fruit irregularly spindle-shaped, (31/2-)
type). 51/2-71/2 by cm, the base slightly acu-
11/2-21/2
Climbing shrub or liana, 2-15 m
high, glabrous; minate-tapering, the top umbonate; pericarp 1

twigs terete, slightly zig-zag; internodes 1-8 cm. mm thick, leathery, yellow to glossy orange.
Thorns recurved, 1-3 mm
long, often wanting. Seeds ^j, ovoid, 9 by 7 by 7 mm.
Leaves firmly herbaceous, oblong, sometimes Distr. Malaysia: Malay Peninsula (Perak),
ovate or obovate, (2.2-)2.4-4(-5) times as long as few collections.
wide, 6-13(-23i/2) by (1 1/2-)2'/2-4'/2(-8) cm; base Ecol. Dense mixed lowland jungle, up to
rounded to acute, top narrowed, acuminate, tip 200 m.
acute, often mucronulate, 3-20 long; midrib mm Notes. The only flowering material which we
sulcate above; nerves (4-)6-9 pairs, conspicuously have is King's Coll. 8824 (Cal). Contrary to
looped-arcuating; petiole 4-7(-10) mm. Flowers King I found the sepals not puberulous but
white, as a rule on slender, short, lateral twigs, glabrous.
serial, 2-4, supra-axillary. Pedicels slender, Diff'ers vegetatively from C. micracantha ssp.
l-3(-4) cm long, glabrous. Buds globular, acute, c. korthalsiana in that the leaves of the latter are
3 mm diam. Sepals ovate, 3-5 by 2-3 mm, with a much coarser, and thicker, while in C. cucurbitina
slightly thickened and more or less acute top, the leaf tip is longer and more slender.
outer pair almost glabrous at the margins, inner
pair slightly ciliate. Petals very thin, 4-6 by 2-3 18. Capparis micracantha DC. Prod. 1 (1824) 247.
mm, white, outside more pubescent than inside,
often minutely ciliate. Disk bilobed, fleshy, by 1 KEY TO THE SUBSPECIES
1 Vi mm. Stamens 20-30, c. 2 cm long. Gynophore
13-20 mm, ovary ellipsoid to globular, c. XVi by 1 . Ripe fruit (sub)globular. Stamens c. 20-35(-60).
3/i-l mm, smooth, stigma Vi mm
long, all gla- Sepals mostly obtuse ssp. micracantha
brous. In fruit the pedicel hardly incrassate and I. Ripe fruit elongate, acute. Stamens 60-100.
the gynophore slightly so. Fruit orange or red, Sepals acute to acuminate. West Malaysia.
globular to ellipsoid, 3-51/2 by 2 1/2-3 '/2 cm, ssp. korthalsiana
shortly umbonate at the top and sometimes also
at the base; pericarp woody to coriaceous, 2-3 ssp. micracantha. C. micracantha DC. Prod. 1

mm thick, more or less tuberculate. Seeds ^j, (1824) 247; Bl. Bijdr. 2 (1825) 52; Spreng. Syst.
embedded in whitish pulp, 10 by 6-8 by 5-6 mm, Veg. 4, 2 (1827) 204 (micrantha. sphalma, non
smooth, brown. A. Rich.): Miq. Fl. Ind. Bat. 1, 2 (1858) 99;
Distr. Malaysia: throughout Borneo (also P. Hook./. & Th. Fl. Br. Ind. 1 (1872) 179; KuRZ,
Laut). J. As. Soc. Beng. 43, ii (1874) 31; Radlk. Sitz.
86 Flora Malesiana [ser. I, vol. 6^

Ber. Bayer. Ak. Wiss. 14 (1884) 118, 128; Vidal or dark violet. Disk bilobed, c. 1 mm. Stamens c.
Rev. PI. Vase. Filip. (1886) 47; King, J. As. Soc 20-35(-60); filaments 18-30 mm, white; anthers
Beng. 58, ii (1889) 394; Pax in E. & P. Pfl. Earn \V2-2y2 by 1/2 rnm, grey or bluish. Gynopliore
3, 2 (1891) 231; Brandis, Ind. Trees (1906) 36 15-30(-35) mm, ovary ovoid to ellipsoid, c. 3 by
Merr. Philip. J. Sc. 1 (1906) Suppl. 58; Back. F1 2 mm, both glabrous, sometimes vestigial. In
Bat. (1907) 57, incl. (Bl.) Hall./.
var. callosa fruit the gynophore 4-6 mm diam., the pedicel
Gagn. Gen. I.-C. 1 (1908) 186; Back. Schoolfl.
El. thinner. Fruit globular to ellipsoid, with 4 longitu-
(1911)63;RiDL. J.Str. Br. R. As. Soc. n. 59(1911) dinal sutures, 3-6 '/2 by 3-41/2 cm, yellow, orange,
68; Hall./. Med. Rijksherb. 12 (1912) 18; Koord. or red; pericarp smooth, 2 mm thick, when dry
Exk. El. Java 2 (1912) 293; Merr. El. Manila woody-coriaceous, pulp juicy. Seeds c. 6-8 by
(1912) 215; Sp. Blanc. (1918) 160; Ridl. El. Mai. 41/2-7 by 3-5 mm, red to shiny black, smooth.
Pen. 1 (1922) 123, f. 1 1 W. H. Brown, Min. Prod.
;

Philip. Eor. 2 (1921) 282; Merr. En. Philip. 2

(1923) 212; Parkins, For. El. Andam. (1923) 82;
Geerts-Ronner, Trop. Natuur 13 (1924) 167,
f. 8; Craib, El. Siam. En. 1 (1925) 82; Merr.
Philip. J. Sc. 29 (1926) 371 ; Gagn. El. Gen. I.-C.
Suppl. (1939) 162; Corner, Ways. Trees (1940)
1

180, f. 47; Back. Bekn. El. Java (em. ed.) 4A

(1942) fam. 45, p. 7; Quis. Med. PI. Philip. (1951)
339.— C. billardierii DC. Prod. 1 (1824) 247;
MiQ. El. Ind. Bat. 1, 2 (1858) 99.— C. callosa Bl.
Bijdr. 2 (1825) 53; MiQ. El. Ind. Bat. 1, 2 (1858)
99; Illustr. (1870) 29, t. 16; Naves in Blanco, El.
Eilip. ed. 3 (1877-83) t. 180; Radlk. Sitz. Ber.
Eig. 20. Distribution of Capparis micracantha DC.
Bayer. Ak. Wiss. 14, 1 (1884) 131; K. V. Bijdr. & ssp. micracantha (continuous line, add Formosa)
4 (1896) 262; Koord. Gedenkb. Jungh. (1910)
and of ssp. korthalsiana (MiQ.) Jacobs (broken
] 67. ^C. flexuosa Bl. Bijdr. 2 (1825) 53; non L.,
nee Vellozo; Hassk. PI. Jav. Rar. (1848) 178;
line). Localities of Czippeliana Miq. (dots).

MiQ. Analecta 3 (1852) 1 El. Ind. Bat. 1, 2 (1858)

;

98; Illustr. (1870) 30; Radlk. Sitz. Ber. Bayer. Distr. Burma, Siam, Indo-China, S. China,
Akad. Wiss. 14, 1 (1884) 101, 129.— C. odorata Formosa (a variety), Hainan, Andamans, in
Blanco, El. Eilip. (1837) 439, ed. 2 (1845) 305, Malaysia: S. Sumatra, Malay Peninsula (except
ed. 3, 2 (1878) 201; Eern.-Vill. Nov. App. Java (in
in the S. corner), W
only a few coastal
(1880) 11; Merr. Philip. J. Sc. 3 (1908) Bot. 77.- localities, Prinsen I., Nusa Kembangan),
also
C. forsteniam MiQ. Illustr. (1870) 32, t. 18, excl. Madura, Kangean Arch., Lesser Sunda Islands
syn. C. ovalifolia Zipp., quae est C. zeylanica L. Lombok, E. and W. Sumbawa, W. Elores,
(Bali,
Hall. /. in Koord. Minah. (1898) 343.— C. Komodo, Alor, Timor, Wetar), North Borneo
roydsiaefolia KuRZ, As. Soc. Beng. 39, ii (1870)
J. (also Banguey), throughout the Philippines,
62. C. myrioneura Hall.
/. in Eedde, Rep. 2 Celebes, Moluccas (Halmaheira). Fig. 20.
(1906) 60, p.p., excl. Koord. 16341, excl. var. Ecol. Especially in light shade (monsoon, teak,
latifolia.—C. venosa Merr. Philip. J. Sc. 10 (1915) or evergreen forest) on dry, often calcareous soil,
Bot. 305, ex descr.; En. Philip. 2 (1923) 213. also in thickets, savannahs, hedges, etc., not
Stout shrub or small tree, rarely climbing, seldom coastal, mostly below 500 m, highest
2-6(-10) m; trunk greyish, finely fissured and set record 1400 m. Fl. fr. Jan. -Dec.
with small knobs each surmounted by a thorn; An at. Raghavan investigated the vascular
branchlets terete, mostly zig-zag, when young supply of the floral parts in a 'C flexuosa' from
sparsely pubescent; internodes c. 21/2-4 cm. Java, cultivated at Kew (the plant was no longer
Thorns 1-A{-1) mm
long, patent or directed up- living in 1959) (J. Linn. Soc. Bot. 52, 1939, 247).
wards, straight or slightly curved. Leaves (sub-) MoLiscH Stated that nearly all the parenchyma
coriaceous, glabrous, 1.7-2.9(-4.1) times as long cells of the petiole, the leaves, and the stem
as wide, mostly broadest about halfway, sometimes contain a strongly refractive crystal of lime. Also
below, or rarely above the middle, 8-18(-24) by bodies of silicon are sometimes found (Ber. Deut.
4-8(-12y2) cm; base mostly rounded, sometimes Bot. Ges. 34, 1916, 154-160).
blunt to subcordate or acute, top broader or Note. C. billardierii, C. flexuosa, and C.
narrower rounded, sometimes slightly emarginate, roydsiaefolia have been described on specimens
or acute, rarely acuminate, dark-tipped; midrib with a vestigial gynophore and ovary. Radl-
subprominent above, nerves 5-7(-]0) pairs, light kofer's sect. Monostichocalyx comprised C.
green when dry; petiole 6-10(-15) mm. Flowers flexuosa Bl., C. callosa Bl., and C. micracantha
up to 6, serial. Pedicels l(-2) cm. Buds ellipsoid, DC. which are in my opinion all synonyms.
acute, 5-12 mm
long. Sepals firmly herbaceous,
51/2-I3 by 2V2-5V2 nim, ±
boat-shaped, ovate to ssp. korthalsiana (Miq.) Jacobs, stat. nov. C. —
oblong, the margins mostly hairy. Petals very thin, korthalsiana MiQ. Illustr. (1870) 31, t. 17; Merr.
oblong to lanceolate, 10-25 by 3-7 mm, white, En. Born. PI. (1921) 280.— C. finlaysoniana Wall.
honey-guide yellow, turning dark red or brownish, [Cat. (1832) 6992 B, nomen] ex Hook. /. &
Th.
Dec. 1960] Capparidaceae (Jacobs) 87

Fl. Br. Ind. 1 (1872) 179; King, J. As. Soc. Beng. a recurved, stiff, darker mucro up to 3 mm; midrib

58, (1889) 395; Ridl. Fl. Mai. Pen.

ii (1922) 124; 1 subdepressed above; nerves 3-8 pairs; petiole
Kew Bull. (1925) 77. 1/2-I '/2(-2) cm, glabrescent. Flowers developing

Fruit oblong, c. 6-17 by 2'/2-3-% cm, tapering before the leaves on young twigs, conspicuous,
to the top and sometimes to the base. Leaves 2-6, serial. Pedicels 4-20(-28) mm, hairy. Buds
acuminate, not mucronate, mostly recurved, never globular, c. 8 mmdiam. Sepals subcoriaceous,
cordate at the base. Sepals very acute and slightly outside more or less densely tomentellous, outer
cucullate at the top, the buds therefore (sub-) pair orbicular to elliptic, mostly acute, 6-11 by
acuminate. Flowers comparatively large. Stamens 5-9 mm, the posterior one (surrounding the disk)
very numerous (up to 100). the largest; inner pair elliptic to oblong with more
Distr. Malaysia: Malay Peninsula (Perak, rounded top, 6-10 by 3-7 mm. Petals very thin,
Pahang, Johore, Singapore), Central and S. oblong with rounded top, 9-1 2(-l 6) by 3 '/2-5 mm,
Sumatra, Borneo. Nowhere common. Fig. 20. white, turning pink, largely glabrous; upper pair
with a pinkish to reddish basal-median spot,
19. Capparis zeylanica Linne, Sp. PI. ed. 2(1762) hairy at the base. Disk c. 1 mm
diam. Stamens
720;DC. Prod. {\^2A)2A1 quoad specim.
1 ,
Ceyl.; 30-45(-70), 20-30(-35) mm, white, turning red;
Dunn, Kew Bull. (1916) 62; non Hook. f. & Th. anthers oblong, slightly broader at the base, c.
Fl. Br. Ind. (1872) 174, quae est C. brevispimi
1 2 by 1 mm, bluish grey. Gynophore slightly ex-
DC. —
C. horrida L. /. Suppl. (1781) 264; DC. ceeding the stamens, up to 4'/2(-5'/2) cm, basal
Prod. 1 (1824) 246; Wight, Ic. PI. Ind. Or. 1 part pale pubescent, otherwise glabrous; ovary
(1839) t. 173; MiQ. Illustr. (1870) 34, quoad var. ft ellipsoid, l'/2-2'/2 by 1-1 '/2 nim, stigma V2 mrn
ervthrodasys; Hook./. & Th. Fl. Br. Ind. 1 (1872) long. In fruit the pedicel sometimes still hairy,
178; Fern.-Vill. Nov. App. (1880) 11; Vidal, gynophore glabrous, up cm by 3-6 mm, as
to 5'/2
Rev. PI. Vase. Fiiip. ( 1886) 48; Merr. Philip. J. Sc. thick as the pedicel. Fruit globular to ellipsoid,
1 (1906) Suppl. 58; Gagn. Fl. Gen. I.-C. 1 (1908) up to 5 by 4 cm, pericarp c. 2 mm
thick, woody-
185; Back. Schoolfl. (191 1) 63; Ridl. J. Str. Br. R. coriaceous, smooth, reddish, orange or purple.
As. Soc. «. 59 (1911) 69; Koord. Exk. Fl. Java 2 Seeds ckj, 5-7 by 5-4 1/4 rnm, brown.
(1912) 294; Merr. Fl. Manila (1912) 215; Sp.
Blanc. (1918) 159; W. H. Brown, Min. Prod.
Philip. For. 2 (1921) 282; Merr. En Philip. 2
(1923) 211; Craib, Siam. En.
(1925) 81;
Fl.
1

Gagn. Fl. (1939) 161; Back.

Gen. I.-C. Suppl. 1

Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, p. 8;

Quis. Med. PI. Philip. (1951) 338.— C. dealbata
DC. Prod. (1824) 246; Decne. Nouv. Ann. Mus.
1

Hist. Nat. Paris 3 (1834) 426; Herb. Timor. Descr.

(1835) 98; Span. Linnaea 15 (1841) 165; MiQ. Fl.
Ind. Bat. 1, 2 (1858) 100; Illustr. (1870) 27.—
C. zevlanica (non L.) Roxb. Fl. Ind. ed. Carey 2
(1832) 567; Gamble, Fl. Pres. Madras 1 (1915)
18. C. aurantioides Presl, Reliq. Haenk. 2
(1835) 86; Gray, U.S. Expl. Exp. Bot. 1 (1854) 70;
Fern.-Vill. Nov. App. (1880) 11. C. linearis —
(non Jacq.) Blanco, Fl. Filip. (1837) 438, ed. 2
(1845) 305, ed. 3, 2 (1878) 200.— C. nemorosa (non
Jacq.) Blanco, Fl. Filip. (1837) 438.— C. micra-
cantha (non DC.) Blanco, Fl. Filip. ed. 2 (1845) Fig. 21. Distribution of Capparis zeylanica L. in
305, ed. 3, 2 (1878) 200, t. 188.— C. rufescens Malaysia. Add Hainan.
Turcz. Bull. Soc. Nat. Moscou 27, 2 (1854)
321.— C. erythrodasys MiQ. PI. Jungh. (1855) 397; Distr. Ceylon, India (largely E of the line
Fl. Ind. Bat. 1, 2 (1858) 99.— C. ovalifolia Zipp. Bombay-Delhi-Dehra Dun, S of the Himalaya),
ex MiQ. Illustr. (1870) 33. C. viminea (non — Burma, Siam, Indo-China, Hainan, Andaman
Hook./. &
Th.) Fern.-Vill. Nov. App. (1880) Islands, in Malaysia: Java (east of Djokjakarta),
1 1. — C mvrioneura var. latifolia Hall./, in Fedde, Lesser Sunda Islands (Lombok. Sumbawa, Semau.
Rep. 2 (1906) 61.—Fig. 5. Timor), Celebes (also Salajar I.), Philippines (Min-
Climbing shrub 2-5(-10) m. Innovations brown- doro, Luzon, Mindanao, Sulu Arch.). Uncommon
red to greyish-tomentose; branchlets mostly zig- in Indonesia, common in the Philippines. Fig. 21.
zag, glabrescent; internodes 2-7 cm. Thorns re- Ecol. Forest edges, bushes, savannahs, hedges,
curved, 3-6 mm
long. Leaves subcoriaceous, limestone hills, obviously bound to seasonal
(1.2-)1.7-2.3(-2.9) times as long as broad, ovate climatic conditions, mainly in the lowlands, up
or elliptic, rarely obovate, above soon glabrescent to 700 m. Fl. fr. Jan. -Dec.
then glossy, beneath later or not glabrescent, and Uses. Of less importance; see Quis. I.e.
dull, 4-10(-18) by 3-5'/2(-9) cm; base rounded, Vern. Philippines: Manunggal laldki. Mindoro,
sometimes subcordate, rarely acute, top acute to haralduik, Ibn., dauag, halubdgat-bdging. Tag.,
rounded, rarely slightly acuminate, generally with laginau, Bis., talaktdk, tarabtdb. Ilk.
88 Flora Malesiana [ser, I, vol. 6'

Fig. 22. Capparis qiiiniflora DC. a. Flowering branch, x 2^, b. flower from the back, X 2, c. fruit, x 2/^,
d-e. narrow leaves, x 2/3 [a-b Specht 697 from N. Austr., c Jaheri 238, d Elbert 3827, e Soehanda
exp. DE Jong 272).
Dec. 1960] Capparidaceae (Jacobs) 89

Note. From C. pyrifolia and C. micracantha In the material described by A. Gray from Fiji
easily distinguished by the recurved thorns. as C. richii the leaf-shape is remarkably variable,
one specimen having the average leaf-index, the
20. Capparis quiniHora DC. Prod. 1 (1824) 247; second having leaves 9I/2-II by l'/2-l% cm, the
Benth. F1. Austr. (1863) 94; F. v. M. Descr.
1 third having linear leaves c. 4-10 cm by 1-4 mm.
Not. Pap. PI. 1^ (1875) 5; Bailey, Queensl. Fl. 1 In West Flores a similar, also sterile linear-leaved
(1899) 57; Laut. Bot. Jahrb. 52 (1914) 112; specimen was collected (Soehanda, exp. de Jong
DoMiN, Bibl. Bot. Heft 89 (1925) 685.— C. 272, Bo, K, L); its thorns are strongly recurved,
trapeziflora Span. Linnaea 15 (1841) 165; MiQ. 1-3 mm, the leaves measure 4-10 by 0,1-0,4 cm.
Fl. Ind. Bat. 1, 2 (1858) 99.— C. subcordata Span. As I observed in several other species of Capparis
Linnaea 15 (1841) 166; MiQ. Fl. Ind. Bat. 1, 2 such sterile shoots have larger thorns than the
(1858) 99; Illustr. (1870) 34.— C. richii A. Gray, fertile twigs. Might it be that such shoots have
U.S. Expl. Exp. Bot. (1854) 69; Seem. Fl. Vit.
1 been developed under exceptionally dry weather
(1865) 6.— Fig. 22. conditions?
Climbing shrub; twigs slightly zig-zag, densely The only collection from Bali is a duplicate
minutely grey or ferruginous-tomentose, some- specimen from Bogor in Paris with a Zollinger
times glabrescent; internodes up to 9 cm. Thorns label "Leg. Teysmann, Balie, Buleling". As no
recurved, c. l-2(-3) mm, sharp, sometimes want- other sheets were seen from Bali, the location
ing, especially on flowering twigs. Leaves sub- seems to be erroneous.
coriaceous to coriaceous, often brownish green
when dry, ovate to (rarely) obovate, 1.6-4.3 times King, J. As. Soc. Beng. 58,
21. Capparis larutensis
as long as wide, initially densely minutely ferru- ii(1889) 393; Ann. R. Bot. Gard. Calc. 5 (1896)
ginous-tomentose, soon glabrescent and glossy 118, t. 134; RiDL. Fl. Mai. Pen. 1 (1922) 122.
above, (5i/2-)7-9(-12) by \^m~A(-1V2) cm; base Climber, 10-13 m. Twigs straight, brown-
rounded, subcordate, blunt, or acute, top mostly puberulous, more or less glabrescent; internodes
attenuate and acuminate, rarely blunt, acumen 1-2 cm. Thorns vigorous, recurved, 3-4 mm
long,
up to 1/2 cm, acutish with a minute, thickened glabrous, with darker, glossy, very sharp top.
mucro; midrib flat to shallowly sulcate above; Leaves firmly herbaceous to subcoriaceous,
nerves 6-8(-ll) pairs under an angle of c. 45 ; oblong, sometimes slightly obovate, glabrous,
intermediate veins often wanting, reticulations 2-3 '/2 by %-l% cm, base acute to subcordate,
distinct; margin sometimes crinkled; petiole top obtuse to acuminate, tip finely retuse; midrib
5-17 mm, hairy as the twigs. Flowers white, serial, narrowly sulcate above; nerves 4-5 pairs, hardly
sometimes developing before the leaves at the visible; margin somewhat recurved when dry;
end of young twigs, 2-10 in a row 2-12 mm long, petiole 3-6 mm, brown-pubescent. Flowers
pedicels 6-17 mm, hairy as the twigs, finally axillary. Pedicel 1 '/2-2 cm, glabrous. Sepals
glabrescent; floral leaves sometimes abortive. fleshy, 7 by 5 mm, ciliate towards the top, the
Buds c. 4 mmdiam. Sepals herbaceous, acute, outer pair ovate, the inner pair suborbicular.
ferruginous-tomentellous outside, outer pair c. Petals obovate, c. 8 by 5 mm, white then pink,
4-5 by 2-3 cm, the inner pair narrower and puberulous inside. Stamens (10?-)30, 2 1/2 cm,
flattish. Petals ovate to elliptic, 5-7 by 2-4 mm, anthers 2 mm long. Torus 2 mm
broad. Gynophore
slightly unequal, puberulous on both sides. Disk 31/2-4 cm; ovary ovoid, c. \V2 mm
long. Mature
up to 1 mm long. Stamens 7-8(-12), filaments fruit unknown, at least 13 by 10 mm, globose,
20-27 mm. Gynoplwre I-IV2 cm but sometimes umbonate. Seeds several.
abortive and only a few mm, glabrous or glabres- Distr. Malaysia: Malay Peninsula (Perak,
cent at the base; ovary ovoid, c. 1 '/2 by 1 mm, Selangor).
stigma knob-shaped. Pedicel, torus, and gyno- Ecol. Dense jungle, clinging to trees, below
phore slightly incrassate. Fruit (sub)globular, c. 200 m. Fl. Sept., Nov.
2-2% by 2-2i/2< pericarp corky-leathery, c.

11^-2'/^ mm thick, with small scattered warts. 22. Capparis spinosa Linne, Sp. PI. 1 (1753) 503;
Seeds '^, up to 7-8 by 5-6 by 4 mm. Hemsl. Bot. Chall. 3 (1885) 120 {var.).
Distr. Pacific (Fiji and New Caledonia), North
Australia to Cape York Peninsula, in Malaysia: var. mariana (Jacq.) K. Sch. Bot. Jahrb. 9
SE. Celebes, Lesser Sunda Islands (Lombok, (1888) 201; K. Sch. &Laut. Fl. Schutzgeb.
Sumbawa, Flores, Semau, Timor), Moluccas (1901) 335; Laut. Bot. Jahrb. 52 (1915) 111.—
(Kai and Tanimbar Is), New Guinea (NW. part; C. corciifolia Lamk, Encycl. 1 (1785) 609; Merr.
Papua; islands in Torres Strait). Fig. 7. Philip. J. Sc. 7 (1912) Bot. 235; Fl. Manila (1912)
Ecol. Prefers obviously coastal habitats in 216; Sp. Blanc. (1918) 159; En. Philip. 2 (1923)
drier areas, rambling on shore trees, bound to a 210; de Voogd, Trop. Natuur 25a (Jub. Uitg.)
seasonal climate, in the lowland. (1936) 72, f. 9; Merr. Philip. J. Sc. 9 (1914)
Notes. Although in fertile material the leaves Bot. 84. C. mariana Jacq. Hort. Schoenbr. 1
are variable in shape, the venation (see fig. 22a, (1797) 57, t. 109; DC. Prod. 1 (1824) 245: Decne,
d-e) is constant and very characteristic. Nouv. Ann. Mus. Hist. Nat. Paris 3 (1834) 426;
The Australian specimens are much more Herb. Timor. Descr. (1835) 98; Span. Linnaea
uniform in leaf index than those of Malaysia, 15 (1841); Blanco, Fl. Filip. ed. 2 (1845) 305;
where it does not exceed 1.8. Miq. Fl. Ind. Bat. 1, 2 (1858) 100; Teysm. Nat.
90 Flora Malesiana [ser. I, vol. 6^

Tijd. N.I. 34 (1874) 359; Naves in Blanco, Fl. the posterior one the larger and strongly saccate,
Filip. ed. 2 (1878) 201, t. 179; Fern.-Vill.
3, sometimes slightly keeled, inner pair equal,
Nov. App. (1880) 11; Safford, Contr. U.S. Nat. slightly larger than the outer pair, 1 1-20 mm
wide,
Herb. 9 (1905) 212; Burk. Diet. 1 (1935) 443.— with broad-membranous margin. Petals pure
C. sandwichiana DC. Prod. (1824) 245; A. Gray,
1 white, sometimes slightly emarginate, upper pair
U.S. Expl. Exp. Bot. 1 (1854) 69; Degener, more or less rhomboid, (3-)3'/2-4V2 by 2-A cm,
Fl. Hawaii. 1 (1937) fam. 142, with plate.— C. with thick fleshy base, lower pair 3-3 Vi by
baducca (non L.) Blanco, Fl. Filip. (1837) 438. 1 V2-l%(-3'/4) cm. Torus conical, 4-5 mm
broad,
—Fig. 23-24. the disk c. 4 by 5 mm. Stamens more than 100,
Tree? or shrub, mostly prostrate. Twigs terete, white, 41/2-5 cm, anthers 4 by %
mm, brownish.
mostly zig-zag; internodes 1-3 cm; innovations Gynophore 6-7 cm, subpubescent towards the
whitish-tomentose, glabrescent. Thorns wanting base; ovary oblong, 5-8 by 1-2 mm, glabrous.
in Mai.. Leaves suborbicular, rarely elliptic, often In fruit the gynophore incrassate until it is as thick
ovate, (1 '/2-)2'/2-6(-7'/2) cm diam., subcoria- as the pedicel, and sometimes coiled as a pig's
ceous (fleshy when fresh); base truncate to round- tail. Fruit ellipsoid to spindle-shaped, c. 5 by
ed, top rounded, seldom minutely emarginate to 1 1/2 cm, narrowed towards the apex, with 5 nerves.

acute; midrib above shallowly depressed in the Seeds cv), subglobular, c. 4 mm through.
basal part, otherwise flat; nerves c. 5-7 pairs, Distr. Scattered over the Pacific within the
thin, subprominent on both surfaces; petiole quadrangle formed by the Sandwich Is, Marianas
%-l'/4 cm. Flowers axillary. Pedicel 4'/2-7V2 cm, (Guam), Solomon Is, and Henderson I. (ENE of
glabrescent. Buds conical when young, later Pitcairn, 24°20'S, 128''20'W), NW. Australia
bulging at the posterior base, sometimes acumi- (once coll.); a different form of the species in
nate, finally 2-2 '/2 cm diam. Sepals c. 25-28 mm Australia (see notes), in Malaysia: Lesser Sunda
long, fulvously pubescent towards the base, Islands (Semau, Timor, Leti, n.v.), Philippines
glabrescent, ovate, outer pair 8-18 mm
wide. (Luzon, cultivated in a few localities; Bohol,
spontaneous along the sea-shore). New Ireland,
and New Britain (/;.v.).
C. spinosa, the well-known caper bush, has a
wide distribution from the Mediterranean through
the Near East to India (Sind and the Punjab).
Several varieties have been distinguished, cf.
BoissiER (Fl. Orient. 1, 1867, 420).
Ecol. Prefers semi-arid or seasonal conditions,
dry lavas, limestone, coastal stations, etc. A
noctiflorous species, in the lowland up to c.
350 m.
In Timor and locally in the Philippines ob-
viously quite capable of maintaining itself in
suitable places and thoroughly naturalized {vide
infra). It can be propagated by suckers or seed.
Vern. Alcaparras, Philippines, acapares, Guam,
both corruptions of the Spanish alcaparro.
Uses. In Malaysia none. According to Safford,
I.e., it appeared from the archives at Agafia in

Guam that some of the early governors of Guam

exported the pickled unripe capsules in con-
siderable quantities, employing the natives to
gather them. This is at variance with the common
habit of pickling the flower buds, and probably
the latter were meant. From Guam the plant
would have been introduced into the Philippines.
Notes. The Malaysian and Pacific population
is markedly uniform inappearance and has a
coherent area, taxonomically and geographically
well separated from the population in India,
where it occurs eastwards to about 90° E long. It
is absent between India and Timor.
Fig. 23. Capparis spinosa L. var. nmriana (Jacq.) Many paramorphs have been described in C.
K. ScH. a. Bud, just before anthesis, from the left, spinosa. The var. mariana is distinct by a com-
X 2/3, b. left petals, of the upper (adaxial) and the bination of the following characters: twigs
lower pair, as seen from within, nat. size, c. stamen unarmed, not too densely tomentose and glabres-
as it is in bud, x y^, d. bud opened, the outer cent, leaves comparatively large and orbicular,
sepals cut medianly, one inner sepal, the left petals lacking an apical spine, flowers very large with one
and the stamens removed, nat. size (Degener saccate sepal. Capparis galeata Fresen. of Arabia
26425 from Hawaii). and E. Africa has similar flowers, the odd sepal
Dec. 1960] Capparidaceae (Jacobs) 91

being even more irregularly shaped, but this plant ranean people, did not appeal to the native taste
has strong stipular thorns and an apical leaf-spine. and were therefore not further dispersed.
The species is taxonomically widely different In conclusion, there remains hardly any doubt
from all native Capparis in Malaysia and Austral- of its introduction at an early date from presum-
asia and it had and has a distinctly economic ably seeds from the Mediterranean into the
significance in the Mediterranean. From this it Marianas and dispersal by man from thence into
appears probable that the species was introduced E. Malaysia and Polynesia, as suggested by
in the Indo-Pacific as a cultigen in post-Columbian Hemsley, I.e.
time. The first records from the Pacific-E. Malay- It seems likely that introduction took place once

sian area are: only, because of the variability pattern of the

Marianas, via Sonnerat' between 1774 and species: repeated introduction would presumably
1781, have caused a greater variation of the population
Sandwich Is, Menzies, Vancouver Exp. 1792,
Timor, Cunningham, 1818,
Society Is, Beechey's Voyage, 1826,
Philippines, Blanco, before
1837.
The dates of this are significant, because it
list

appears clearly that it had not been collected

during the earliest Pacific expeditions such as
'Cook's Voyages' and the 'Boudeuse et TEtoile',
1768-1770, on which such trained botanists col-
lected as Banks, Solander, the Forsters,
CoMMERSON, etc., who did not discriminate be-
tween cultivated, introduced, and native plants,
as Merrill rightly observed in his account of
Cook's Voyages (C'hron. Bot. 14, 1954, 173). They
travelled widely in Central Polynesia, but did not
visit Guam and the northern islands. Obviously
the species was then not in Central Polynesia.
Its spreading in the Pacific, notwithstanding its
use, and its early abundance in Guam, must have
taken more than a century, because already on-
wards of 1520 there had been a connection by
Spanish ships between Acapuico in West Mexico
(from where, however, the species is not known)
and Manila via Guam in the Marianas; the regular mariana
Fig. 24. Capparis spinosa L. var. (J ACQ.)

galleon route was initiated, for return voyages, K. ScH. Flower (van der Pijl).
in 1565, with l-2(-3) ships yearly in either direc-
tion, to be abandoned in 1815 (Merrill, I.e. In addition it should be stated that the species
193). It is remarkable that the species has never occurs also on the west coast of Australia and in
been reported from Mexico. Queensland, from where it has been described as
In this respect it appears significant that the C niimmiilaria DC, a name also taken up by
earliest record from the Pacific, 1774-1784, is Bentham. It has, probably rightly, been reduced
from the Marianas, from where is also the very to C. spinosa by F. v. Mueller (Syst. Cens.
early record of its being cultivated and exported. Austr. PI. 1882, 5; 2nd ed. 1889, 8) and considered
In the Philippines there is the indication that to represent a variety of C. spinosa by F. M.
it has been imported there from Guam. Bailey (var. numnuilaria (DC.) F. M. Bailey,
RuMPHius makes no mention of it, though he Syn. Queensl. PI. 1883, 15). It has even been found
frequently referred to plants from islands outside in arid places in Central Australia. The Australian
the Moluccas proper. plants look hardly different from those of the
In Timor there is no indication of such import, Mediterranean and they are distinctly thorny.
but it is not far-fetched to assume its introduction But there is one specimen from NW. Australia
by the Portuguese. One may question why it has (Pritzel 284) which matches the Polynesian
not further spread in arid places in East Malaysia material. For this reason it seems likely that the
and the Lesser Sunda Islands. It may well be that Australian form is an introduction which took
the capers, this well-known delicacy to Mediter- place independently from that in the Pacific. As a

(1) Lamarckstated to have received his material from Sonnerat, but the latter never visited the
Marianas 1, p. 494). He attached himself, however, to Commerson and worked with him for
(see vol.
three years in Mauritius, Bourbon, and Madagascar. Jacquin also described the plant from the Marianas
but did not mention the collector's name. Therefore it remains obscure who brought plants or seeds from
the Marianas; it was obviously not brought by some early botanical explorer. But the seeds may have
been derived from the Spanish export from Guam and have been cultivated in Mauritius where Sonnerat
obtained his specimens.
92 Flora Malesiana [ser. I, vol. 6^

matter of fact the case of the rather early record somewhat incrassate. Fruit globular, IVi-^Vi cm
of this European or West Asian plant in Australia diam., pericarp c. 3 mm
thick, soft, leathery,
is matched by some other examples. In all these smooth, dull purplish brown, pulp spongy. Seeds
cases the Australian population is (mostly 7-30, sometimes subangular, c. 9 by 6 by 5 mm.
slightly) deviating and Dr van Steenis has suggest-
ed this convivial or racial differentiation to be due
to the effect of selection through isolation (this
Flora 1, 4, 949, Hi). Different imports of a cultigen
1

will likely lead to slightly different racial develop-

ment.
How the introduction in Australia has taken
place is dubious; it could be surmised that oc-
casionally ships or life-boats thrown out of their
way touched these coasts or that they resulted
from ship-wrecks. There may have been more than
one introduction.
It seems to me that C. spinosa offers an in-
teresting opportunity for introducing experimental
taxonomical investigation into ethnobotany.

23. Capparis lucida (Banks ex DC.) Benth. F1. Fig. 25. Localities of Capparis lucida (Banks ex
Austr. 1 (1863) 96; Bailey, Queensl. Fl. 1 (1899) DC.) Benth. From the New Hebrides one un-
60; Britten, 111. Austr. PL 1 (1900) 6, t. 6; confirmed record. In Australia several other
Domin, Bibl. Bot. Heft 89 (1925) 688; C. T. localities to 30" S, and one unspecified locality
White, J. Arn. Arb. 10 (1929) 2\1 .^Thylachiiim "Gulf of Carpentaria".
hicidiim Banks ex DC. Prod. 1(1824) 254.— C.
subacute! MiQ. Fl. Ind. Bat. 1, 2 (1858) 101; Distr. Australia: NW. coast (Bentham), and
Illustr. (1870) 35, t. 19, quoad specim. javan. (vide NE. coast of Queensland to 20" S. lat.; in Malay-
sub C. sepiaria); K. & V. Bijdr. 4 (1896) 260; sia a rare plant: E. Java (one locality), Lesser
Back. Schoolfl. (1911) 63; Koord. Exk. Fl. Java Sunda Islands (Komodo, Pada, Timor), SE.
2 (1912) 293 (acuta, sphalma); Back. Bekn. Fl. Celebes, New Guinea (Papua), Booby I. in Torres
Java (em. ed.) 4A (1942) fam. 45, p. 10.— C. Strait, ?Bismarck Arch. Fig. 25.
nobilis (non Benth.) F. v. M. Descr. Not. Pap. PI. Ecol. Chiefly coastal, but also in savannahs.
28 (1886) 41. Fl. fr. Jan.-Dec.
Tree or shrub, c. 2-12 m. Twigs terete, straight, The sweet-scented, white or pale lemon-yellow
densely fulvous pubescent, sooner or later glabres- flowers are typically nocturnal. Koorders noted
cent; internodes less than 2 cm. Thorns in flowering that the fruit pulp has a sourish stench, like that
specimens nearly always wanting, if present up to of rotten fish. In a living plant in Kebun Raya
2 mm. Leaves coriaceous, ovate to obovate, Indonesia (II-Q-17) I found the spongy, yellow
glabrous above, mostly very soon glabrescent pulp tasting and smelling as insipid mango. They
beneath I.7-2.5(-3.5) times as long as wide, might be eaten by bats which could add to their
(3-)5'/2-7(-9) by (2-)2 1/2-3 cm; base acute to dispersal.
cuneate, top rounded to obtuse, sometimes Notes.In the Bogor Gardens another specimen
mucronate; midrib above subprominent, nerves iscultivated (Xl-B-V-1 17); it is a tree, 5 by m
5-9 pairs, subprominent on both sides; petiole 15 cm; it stands in deep shade and never flowers.
'/4-1 Vi cm, hairy as the twigs. Raceme pubescent, The leaves of its long, overhanging twigs tend to
terminal, with up to a dozen of sweet-scented have a distichous position and are subtended by
flowers, the lower in the axils of small leaves. straight thorns 2-2'/2 mm long.
Bracts proper, if present, small, acute. Pedicels Seeds sown from n. Il-Q-17 produced seedlings
1 1/2-6 cm. Buds ovoid, c.
1/2 cm long, often with of which the shoot very similar to those of the
is
1

umbonate top. Outer pair of sepals coriaceous, just mentioned And though I have not
sterile tree.
initially connate and completely enveloping the observed the transitional stage wherein the
bud, then splitting more or less regularly, patent seedling gets unarmed twigs with normal leaves I
at anthesis, c. 10-15 by 10 mm, acuminate, consider the unarmed sterile tree as a retarded
glabrous; inner pair rather thin, flat, ovate, c. juvenile stage through the effect of shade.
11-13 by 5-6 mm, shortly acuminate. Petals The occurrence of a sterile juvenile stage with
subobovate, 18-30 by 10 mm, pubescent inside thorns seems to be typical for the mainly Austra-
and outside towards the base, white to yellowish. lian sect. Busbeckea: Bentham reported it in
Torus comparatively broad, short-conical, with a C. canescens Banks ex DC. and observed it in
I

small posterior disk. Stamens c. 50-70, 21/2-5 cm, several specimens of C. nobilis F. v. M. and C.
filaments white, anthers c. 21/2 mm
long. Gyno- mitchelliiLindl.
phore 21/2-7 cm, mostly thinly woolly in the basal de Candolle referred this species to the genus
half, glabrescent; ovary ellipsoid, acuminate, c. Thylachium on account of lacking petals; this
3-4 mm long, glabrous, stigma dark-purple. In error is perhaps due to their being very easily
fruit the pedicel, the torus, and the gynophore caducous.
Dec. 1960] Capparidaceae (Jacobs) 93

Incompletely known species which are typical for C. pubiflora I am inclined to

compare the material with that species, with

Capparis longipes Merr.^ Philip. J. Sc. 13 (1918)
which it agrees in the greyish pubescence of the
Bot. 12; En. Philip. 2 (1923) 211.
innovations, the straight thorns, and the venation
Scandent, glabrous shrub, branches slender,
pattern. It may represent a deviating form
terete, brownish or olivaceous, the ultimate
(teratological or developed in shade?)though I
branchlets c. 1 mm
diam. Thorns straight, usually
have as yet not seen any comparable paramorph
l-AYi mm. Leaves lanceolate, membranaceous to
in C. pubiflura.
chartaceous, green or greenish-olivaceous when
dry, somewhat shining, 7-11 by 2-3 cm, narrowed Excluded
upwards to the very slender apex, sharply acute- Capparis caraiidas Burm. /. Fl. Ind. (1768) 118,
acuminate, base acute; nerves c. 15 pairs, slender, 119. Merrill reduced this to Carissa carandas L.,
distinct on both surfaces, anastomosing, primary cf. Rumph. Herb. Amb. (1917) 425. Dr Baehni
reticulations lax, ultimate ones close, both kindly sent me Burman's authentic material in
distinct; petiole 2-3 mm. Infnictescetices axillary, the Geneva Herbarium on loan. The entry on
very slender, sparingly branched, up to 20 cm p. 118 may refer to one sheet which is most
long, each branch bearing a single fruit, its pedicel probably a Carissa; the entry on p. 119 refers to
c. 3 cm. Fruit globose, brown when dry, glabrous, four other sheets belonging to Carissa carandas L.
c. 12 mm
in diam. Mant. 1 (1767) 12 (Apocynaceae). According to
Distr. Malaysia: Philippines (Luzon: Abra), the labels the material was derived from cultivated
only known from the type BS 26980 Ramos (US, plants in Java.
K; L, photogr.). Capparis versicolor Griff. Notul. 4 (1854) 577;
Notes. The identity of this specimen could not Hook./. &Th. Fl. Br. Ind. 1 (1872) 175. Recorded
be established with certainty, as the holotype in from Tenasserim and from Java by Hooker, who
Manila ha« been lost, and the isotypes have no suggested its possible identity with C. salaccensis
fruit. From the description by Merrill (vide supra) Bl.
and the completely sterile isotypes it is not pos- I have seen the type in the Calcutta Herbarium
sible to form an adequate idea about the structure and agree that it is vegetatively similar to C
of the inflorescence. cantoniensis; the buds have disappeared but are
Though Merrill did not mention the bract-like assumed to have measured c. 12 diam. on mm
minute scales towards the base of the shoots pedicels c. 3 cm.

3.CADABA
FORSK. Fl. Aegypt.-Arab. (1775) 67; Pax & Hoffm. in E. & P. Pfl. Fam. ed. 2,
17b (1936) 185.— Fig. 26-27.
Shrubs, often glandular-pubescent (or with scales), occasionally with stipular
thorns. Leaves simple (or 3-merous, or wanting). Racemes terminal (or flowers
sohtary, axillary). Sepals 4, in 2 whorls, unequal, caducous, outer pair enveloping
the bud, valvate, inner pair apert. Petals mostly 4 and equal, unguiculate. Recept-
acle with an adaxial, cylindrical gland. Stamens 4-8, their base connate with the
gynophore (androgynophore); anthers comparatively large. Ovary on a long
gynophore, 1 -celled; placentas 2-4; ovules cv), 2-seriate; stigma sessile, indistinct.
Fruit mostly cylindrical, dehiscent with 2 coriaceous valves (or an indehiscent
berry). Seeds cv), subglobose, with a cartilaginous, sculptured testa; cotyledons
incumbent-convolute; radicle conical.
Distr. About 30 spp. in the drier regions of Africa, Madagascar, the Middle East, India, Ceylon,
one sp. in South Malaysia and in N. Australia.
Taxon. There are 3 sections, distinguished by the leaves (simple, 3-foliolate, or wanting) and the
number of petals and stamens. The Malaysian sp. belongs to sect. Cadaba {Eu-cadaba Endl.). The other
two sections are monotypic and occur in South Africa and the Deccan Peninsula respectively.

1. Cadaba capparoides DC. Prod. (1824) 244;

1 21; Koord. Exk. Fl. Java 2 (1912) 295; Back.
Decne. Nouv. Ann. Mus. Hist. Nat. Paris 3 Onkr. Suiker. (1931) 256, Atlas t. 266; Pax &
(1834) 427; Herb. Timor. Descr. (1835) 99; Hoffm. in E. & P. Pfl. Fam. ed. 2, 17b (1936) 186,
Deless. Ic. Sel. PI. 3 (1837) 5, t. 9; Span. Linnaea f. 806, 85 Fl. Java (em. ed.) 4A
F-G; Back. Bekn.
15 (1841) 164; Miq. Fl. Ind. Bat. 1, 2 (1858) 97; (1942) fam. 45, p. .—Fig. 26 27.
1 1

Benth. Fl. Austr. 1 (1863) 92; Miq. Illustr. (1870) Shrub, V3-3 m, procumbent to erect, very dense

(1) C. longipes Standl. Contr. U.S. Nat. Herb. 23 (1922) 303, nan Merr. 1918 = C. discolor Standl.
op. cit. 20 (1919) 182, uon Donn. Smith, 1897 = C. renominata Jacobs, nom. nov. It is a Mexican species.
94 Flora Malesiana [ser. I, vol. 6^

Fig. 26. Cadabci capparoides DC. a. Habit, X 2/3, b. flower from the left, nat. size, c. pedicel with andro-
gynophore, gynophore, fruit, x 2/3, d. seed, x 3 {a-b type specimen, Leschenault s.n., c-^ Teysmann
s.n.).

glandular-pubescent to almost glabrous. Twigs to acutish, rarely rounded, mostly mucronate;

terete,straight; internodes ^/^-lYi cm. Thorns nerves 7-9 pairs; petiole (V4-) 1/2-4 cm. Racemes
c.

straight to slightly recurved, '/2-4 mm, yellowish,

1 terminal, corymbiform; rachis 2-3 cm. Pedicels
sometimes small or wanting. Leaves simple, (1 /2-)2-3(-3i/2) cm. Bracts caducous, 2-5 '/2 mm
firmly herbaceous, ovate (to rarely obovate), stalked, blade 6-10 by '/2-2V2 rnm, lanceolate to
(1.6-)2-3(-5.2) times as long as wide, (3'/2-)6-10 ovate, acute, subtended by 2 minute thorns. Buds
(-131/2) by (iy4-)2i/2-4'/2(-8'/2) cm; base rounded ovoid, acute. Sepals glandular-puberulous outside,
to acute, sometimes obtuse, top narrowed, blunt boat-shaped, (8-)10-15(-20) by (3-)6-9(-12) mm,
Dec. 1960] Capparidaceae (Jacobs) 95

ovate, acute to acuminate, inner pair somewhat obovate, acute. Petals pure white on a green claw,
smaller, flattish, sometimes up to 1 mm clawed, all pointing upwards at the adaxial side, (41/2-)
8-15(-22) mmclawed, blade orbicular to broad-
elliptic, 5-12 by 4'/2-7 mm. Gland green, 5V2-15
by c. 1mm; top yellow, flat, transverse, one-sided,
lanceolate, 41/2-6 mm long. Stamens (5-)6(-7),
androgynophore 1-2 V2 cm; filaments 1-2 cm,
anthers 3V2-5 by 3/4 mm, all glabrous. Gynoplwre
11/2-21/2 cm; ovary cylindrical, 4'/2-6 by Yn-\ mm,
both sparsely glandular-puberulous; stigma knob-
shaped, small; placentas 2. In fruit the pedicels
hardly stretched or thickened, androgynophore
(l-3-)2-3(-3i/2) cm, gynophore (1 i/2-)2-3(-3i/2)
cm, more or densely glandular-puberulous.
less
Fruit cylindrical, dull grey-brownish, (4i/2-)7-ll
cm by 5-13 mm, ±
glabrescent, dehiscing basis-
copically, the filiform placentas persisting as a
replum. Seeds up to c. 20, 3-4 by 21/2-3 by 1/2 mm, 1

more or less deeply reniform, with shallow con-

centric ribs, dull dark brown.
Distr. Australia (N. coast and Vansittart Bay),
in Malaysia: E. Java (along the N. coast E of
Surabaja), Lesser Sunda Islands (Bali, Sumbawa,
Flores, Komodo, Timor, Leti). Fig. 29.
Ecol. Not rare in dry shrubby or grassy
habitats, sometimes coastal, also reported from
limestone, and once from periodically inundated
coastal forest, at low altitude, a typical constituent
of the flora of the monsoon area with a long dry
season.
Fig. 27. Cadciba capparoides DC. Open flower. Vern. Bangol bangoL Bali (also in use for other
The gland is in the centre (van der Pijl). Capparidaceae), paumahatas, Timor.

4. STIXIS

Lour. F1. Coch. (1790) 295, ed. Willd. (1793) 361 ; Pierre, Bull. Soc. Linn. Paris
1 (1887) 652.— Roydsia Roxb. PI. Corom. 3 (1819) 87; Hook./. F1. Br. Ind.
1 (1874) 180, 409.— Covilhamia KoRTH. Ned. Kruidk. Arch. 1 (1848) 307.— Fig. 28.
Rather small, unarmed, woody climbers, rarely shrubs. Branches lenticellate.
Leaves simple, acuminate, rather large, often with minute, whitish pustules along
the midrib, finely pellucid-dotted; petiole incrassate at the apex. Racemes or
panicles cx^-flowered, axillary or terminal, with caducous bracts. Pedicels short.
Sepals mostly 6, in two whorls of 3, valvate, at the top of the bud the outer sepals
covering the margins of the inner ones, more or less strap-shaped, densely fulvous-
tomentose on both sides, inserted on the margin of a widened, dish-shaped,
persistent torus. Corolla absent. Stamens on a short, ± cylindrical androgyno-
phore, 20-50(-c. 100), unequal, the outer ones shortest, about as long as the sepals.
Gynophore about equalling the filaments. Ovary subglobular, 3-celled, with 3
axillary placentas each bearing 5-8(-10) ovules; style simple or split into three
arms. Fruits few, on a thick woody stalk, eUipsoid, 2^/2-5 cm long, sometimes with
small persistent style, finally more or less lenticellate. Seed 1, large, embedded in
pulp, with a thin testa; cotyledons unequal, one enclosing the other.
Distr. Seven spp., in India (Khasia, Sikkim, Assam, Chittagong), Burma, Indo-China, Hainan, in
West Malaysia: 3 spp. Fig. 29.
Taxon. Hooker's division into Roydsia (style short or none, stigmas 3, free) and Alytostylis Hook./.
(style long, with 3 minute terminal stigmas), brought by Pierre to sectional level, is here abandoned.
The division is, in my opinion, artificial, as it separates S. siiaveolens (Roxb.) Pierre and S. Philippine nsis
96 Flora Malesiana [ser. I, vol. 6^

Fig. 28. Srixis ovata (Korth.) Hall./,ssp. ovata. a. Habit, X 2/3, b. flower, one inner sepal removed, X 4,
c. pedicel, torus, gynoecium,x 4, d. cross-section through ovary, x 3, e. fruit, x 2/3 (a type specimen,
KoRTHALS s.n., b-d Clemens 26647, e Clemens 26000).
Dec. 1960] Capparidaceae (Jacobs) 97

which seem to be mutually closer related than and S. obtiisifoUa (Hook. /. & Th.) Pierre,
S. suaveolens
which two species would belong to § Ruydsia. An
examination of the division of the stigmas shows that
this character is less qualitative than quantitative. A grouping according to the length of the androgyno-
phore seems to be more satisfactory, it being considerably longer in S. suaveolens and S. philippinensis
than in the rest of the species.
Anat. In leaf sections of 5. ovciia, made by Mr P. D. Burggraaf, we found that the whitish pustules
are groups of sclerenchymatic cells in the mesophyll tissue; when the leaves are dried and the mesophyll
shrinks these places become prominent.

KEY TO THE SPECIES

1. Gynophore longer than 6 mm, hairy. Leaves and ovary glabrous. Sepals 11-12 mm long.
1. S. philippinensis
1. Gynophore shorter than 5 mm, glabrous. Sepals c. 4 mm long.
2. Adult leaves hairy underneath. Ovary hairy 2. S. ovata ssp. ovata
2. Adult leaves glabrous, occasionally with a few hairs underneath on the nerves only. Ovary glabrous.
3. S. scortechinii

1. Stixls philippinensis (Turcz.) Merr. Govt Lab.

Publ. (Philip.) n. 35 (1906) 72; Philip. J. Sc. 1

(1906) Suppl. 58; En. Philip. 2 (1923) 213;

Erdtman, Pollen Morph. (1952) 97. Roydsici
philippinensis Turcz. Bull. Soc. Nat. Moscou 27,
2 (1854) 329; Fern.-Vill. Nov. App. (1880) 11;
ViDAL, Sinopsis Atlas (1883) 13, t. 6 f. B; Phan.
Cuming. Philip. (1885) 94; Rev. PI. Vase. Filip.
(1886) 48. Roydsia fluribunda Planch, ex Hook.
/. Fl. Br. Ind. 1 (1874) 409.^S.floribi/nda (Planch.
ex Hook. /.) Pierre, Bull. Soc. Linn. Paris 1

(1887) 655.
Climber 6 m by 2 cm, glabrous except the
(?),
inflorescence. Leaves subcoriaceous, elliptic to
oblong, sometimes slightly obovate, 1.3-2.2 times
as long as broad, (9-) 13-21 (-25) by (6-)8-10
(-11 '/2) cm, smooth except for a few pustules
above near the base of the midrib, top c. '/2 cm
acuminate; midrib above broadly sulcate; nerves
9-11 pairs; petiole 2-3 cm. Panicle :v -flowered, Fig. 29. Distribution of the Malaysian species of
terminal, c. 25 cm long; branches up to 15 cm, Stixis —
and of Cadaba. 1. S. philippinensis
all axes angular and fulvous-puberulous. Bracts (Turcz.) Merr., 2. S. ovata (Korth.) Hall. /.
orbicular '/t rnm to linear 3 mm. Pedicels '/2 cm. (triangles representing localities), a. ssp.fascicutata
Buds 10-11 mm
long, with cylindrical base and (King) Jacobs, b. ssp. ovata, 3. S. scortechinii
somewhat swollen, acute top. Torus 1 V2-2 mm (King) Jacobs. — Cadaba capparoides DC. (dots
broad. Sepals during anthesis reflexed halfway, representing localities).
narrow-spathulate, blunt to acute, obscurely
3-5-nerved, the base not narrowed but slightly Fl. Ind. Bat. 1, 2 (1858) 180; Merr. En. Born.
thickened. Androgynophore 2-3 mm, sometimes (1921) 381. S. fasciculata var. borneensis Heine,
hairy at the top. Stamens c. 35-40(-48), glabrous, Mitt. Bot. Staatssamml. Munch. Heft 6 (1953) 212.
filaments 10-15(-16) mm. Gynophore 8-11 mm,
densely puberulous; ovary subglobular, glabrous, ssp. ovata. — Fig. 28.
c. 2 by I '/2 mm; style lYj-li-iy^) mm, glabrous, Branchlets fulvous-tomentose, finally glabres-
with 3 small but distinct stigmatic lobes. Pedicel in cent. Leaves firmly herbaceous, oblong to lan-
fruit c. 1 cm, torus 4 mm
broad, gynophore 3 mm ceolate, mostly somewhat obovate, rather narrow,
thick, still hairy. Fruit (only some unripe and a %-P/2 cm acuminate, fulvous-pubescent, glabres-
few fragments of a ripe one are known) globular cent, (8-)12-17(-22) by 4-8(-l 1) cm; midrib above
with persistent style, up to 3'/2 cm diam.; pericarp sulcate; nerves 7-9 pairs, mostly densely set with
thin, leathery. Seedby '/2 cm.
c. 2 1 minute whitish pustules. Petiole l-2(-2'/2) cm,
Distr. Philippines (Luzon: Bataan and Laguna hairy as the twigs. Racemes or panicles axillary
Prov.; Basilan; Mindanao: Surigao, Cotabato, (rarely a terminal panicle), 6-17 cm long, axes
and Davao Prov.). Seems to be very local. Fig. 29. angular, fulvous-tomentose. Bracts linear, 2-3
Ecol. Forests at low and medium altitudes. (-4)mm, tomentose. Pedicels thin, c. 1/2 cm. Buds
Fl. Jan. -Dec, fr. March-April. obovate, 4-5 mm
long. Flowers greyish green.
Torus 1 mm
wide. Sepals reflexed, lanceolate, c. 4
2. Stixis ovata(Korth.) Hall. /. Beih. Bot. by mm, acutish, obscurely 3-nerved. Andro-
1

Centralbl. 39, (1921) 35.—Covilluimia ovata

ii gynophore c. '/2 iTim long and wide, glabrous.
Korth. Ned. Kruidk. Arch. 1 (1848) 307; Miq. Stamens 26-34, filaments IVi-'iVi mm, glabrous.
98 Flora Malesiana [ser. I, vol. 6^

Fig. 30. Cleome nitidosperma DC. a. Flowering branch, X 2/3, b. flower, lower sepal and petals removed,
right 3 stamens omitted, x flower from the adaxial side, upper sepal, petals, two stamens removed,
6, c.
lower petals partly, other sepal and stamens omitted, x 6, ci. front view of flower, x 2, e. flower, ' 4,
/. flower from the abaxial side, lower petal and 4 stamens removed, upper sepal, petals, and stamens
omitted, x 6, g-h. seed with elaiosome, x 6 (living material, Bogor).
Dec. 1960] Capparidaceae (Jacobs) 99

Gynophore 1-2 mm, glabrous; ovary ellipsoid, c. below mostly densely set with pustules; nerves
1
'/4 by mm, densely appressed-pubescent; style
1 6-7(-8) pairs, reticulations prominent; petiole
1 1/2-2 mm, slender, glabrous, stigma obscurely (1-)1 '/2-2(-2i/2) cm. Inflorescence a terminal,
3-lobed. Fruit on a woody stalk %
by 1/2 cm, slender, leafy, many-flowered panicle 15-20 cm
ellipsoid, 3-3 1/2 by 2 '/i -2 1/2 cm; pericarp %-l mm long, densely downy-puberulous. Bracts linear, c.
thick, leathery-corky, irregularly covered with 3 mm. Pedicels 2-3(-5) mm. Buds obovoid, 2V2-3
whitish crust-like fragments, glabrous, crowned mm long. Flowers fragrant. Torus c. % I mmwide.
by the style base. Seed oblong, c. 2^4 by 1 1/2 cm. Sepals reflexed, strap-shaped, somewhat obovate,
Distr. Malaysia: Borneo. Fig. 29. acute, inside pink, 3-4(-4'/2) by %-! 14 (-1 V2) mm.
Ecol. In forests, up to 1200 m. Androgynophore hardly visible, glabrous. Sta-
Vern. Sansaiig, Dusun (N. Borneo). mens (20-)25-40; filaments XVi-^Vi mm; anthers
Note. In continental Asia (Burma, Tenasserim, elliptic, c. % mm long. Gynophore %-l'/^ mm
and Indo-China) another subspecies occurs, long; ovary ellipsoid to ovoid, c. \-\Va by 1mm,
formerly distinguished as a separate species, both glabrous; style 1-1 Vi mm, slender, glabrous,
which differs from the type subspecies in having after anthesis somewhat recurved, stigmas ob-
the gynophore hirsute except at the base, flowers scure. Fruit on a woody stalk, 5-7 by 4-5 mm,
in terminal panicles 12-35 cm long or racemes over ellipsoid c. 3-4 by 2-3 cm; pericarp leathery,
12 cm long. Bract mostly 4 mm. thin, mostly smooth but sometimes spotted with
whitish, crustlike fragments. Seeds c. lYi by 1 14
3. Stixis scortechinii(King) Jacobs, nov. comb. — cm, embedded in pulp c. 3 mmthick.
Rovdsia scortechinii King, J. As. Soc. Beng. 58, Distr. Malaysia: Northern Sumatra (East and
ii (1889) 397; Ann. R. Bot. Card. Calc. 5 (1896) West Coast Res.), Malay Peninsula (Wellesley,
120, t. 139; RiDL. Fl. Mai. Pen. (1922) 121.—
1 Perak, Selangor, Negri Sembilan, Penang). Fig. 29.
Roydsia parviflora (non Griff.) King, J. As. Soc. Ecol. Dry sunny places, young secondary
Beng. 58, ii (1889) 396. forest, landslides, jungles, 100-1100 m. Fl. fr.
Shrub, often twining (to the left), (l-)3-4(-8) m. Jan. -Dec.
Branchlets downy puberulous, glabrescent. Leaves Uses. Sore eyes are treated with the juice from
firmly herbaceous to subcoriaceous, elliptic to the roots (NE. Sumatra).
oblong, glabrous, c. ( .4-)2 times as long as broad,
1 Vern. Simar silaun, Toba, andor si bumbun.
mostly obovate, (8-)IO-16(-20) by 4-7(-9) cm, Sum. E. C.
base more or less narrowed and acute, top rounded Note. KuNSTLER recorded it on his labels once
and abruptly acuminate with triangular, short, as a "tree 40-50 ft" and once as a "splendid
blunt tip; midrib above flat to finely sulcate, climber 80-100 ft"; this needs verification.

5. CLEOME
LiNNE, Gen. PI. ed. 5 (1754) 302; Sp. PI. 2 (1753) 671 R. Br. in Oudney, Denh. ;

& Clapp. Narr. Trav. Disc. Afr. (1826) App. 220; DC. Prod. 1 (1824) 238;
ScHULTES, Syst. 71 (1829) 23; Pax & Hoffm. in E. & P. Pfl. Fam. ed. 2, 17b (1936)
2\0.— Pedicel/aria Schrank in Roem. & Ust. Mag. Bot. 3, pt 7 (1790) 10.—
Polanisia {non Raf.) sensu DC. Prod. 1 (1824) 242, excl. spec. typ. P. graveolens,
cf. iLTis, Brittonia 10 (1958) 33; Pax in E. & P. Pfl. Fam. 3, 2 (1891) 224.—
Gynandropsis DC. Prod. 1 (1824) 237; Pax & Hoffm. in E. & P. Pfl. Fam. ed. 2,
17b (1936) 217.—Fig. 30-33.
Annual (or perennial) herbs, often hairy, sometimes glandular-hairy. Stipules
none or obsolete; seldom with stiff", recurved, thorn-like, stipular (?) enations at
the base of the Leaves petioled, herbaceous, in the Mai. spp. palmately
leaf.
dissected into 3-7
Flowers pedicelled, in leafy, terminal racemes or
leaflets.
panicles, the leaves apically gradually reduced, mostly slightly zygomorphic in
the position of the petals. Sepals 4, free. Petals (normally) 4, the base often
clawed. Stamens 6 to c\3, in Mai. spp. all fertile, sometimes at the base connate
with the gynophore to an androgynophore. Disk small. Ovary 1 -celled, sometimes
sessile but mostly on a gynophore; stigma knob-shaped or flattish, subsessile.
Capsule linear, terete, 2-vaIved, beaked, dehiscing from the base or from the apex,
the 2 placentae forming a persistent replum. Seeds 00, orbicular to horseshoe-
shaped (with a more or less open cleft), sometimes with a funicular elaiosome,
on the dorsal side sculptured to scaly.
100 Flora Malesiana [ser. I, vol. 6^

Distr. A pantropical and subtropical genus with over 150 spp., many of them in America, in the
Old World c. 65 spp. mostly in Africa and the Middle East; in Malaysia 8 spp., of which 2 cultivated,
and the others native or introduced. Several spp. have in recent time been introduced into other continents
as aliens and are now widely spread weeds.
Ecol. In Malaysia most species are weeds along roadsides and in fields at low altitudes.
Uses. Two spp. are cultivated as ornamentals, especially C. speciosa. Some are used as vegetables
and in primitive medicine.
Hairs. In all Malaysian Cleomes the hairs are simple and multicellular. They are densely set, patent,
and glandular in C gynandra, C. spinosa, C. viscosa, C. aculeata; in the last species the hairs are very
small and non-glandular hairs are mixed with the glandular ones. C. speciosa is almost glabrous; its
scarce vestiture resembles that of C. aculeata. In C. chelidunii the plants are covered, mostly densely,
with appressed, stiff, glassy hairs with a more or less bulbous base. In C. aspera and C. riitidosperma
there are sparse, subpatent, epidermal appendages, too large to be regarded as hairs in the strict sense.
Taxon. de Candolle (1824) distinguished Cleome, Gynandiopsis, and Polanisia. R. Brown (1826)
suggested the reduction of Gynandiupsis as a section to Cleome, except for C. gynandra on which he
based a new section Gymnogonia. The Schultescs (1829) sunk all three genera into a single genus
Cleome in which they distinguished four sections; their classification has still its merits and is preferred
here to that of Pax & Hoffmann which seems to be somewhat unbalanced. Their sections were: sect.
A. Gynandropses (DC.) Schult., with an androgynophore, to which would belong C. gynandra and
C. speciosa; sect. B. Cleomes (DC.) Schult., with 6 free stamens, which is subdivided into siibsect. 1.
Pedicellaria (DC.) Schult., with a long fleshy torus and a long gynophore, to which would belong
C. spinosa, and siibsect. 2. Silignaria (Yorsk.) Schult. w ith a small torus and a short or wanting gynophore,
to which would belong C. aspera, C. riitidosperma, and C. aculeata; sect. C. Polanisiae (DC.) Schult.,
with more than 8 non-clavate stamens, to which would belong C. viscosa; sect. D. Corynandrae (Schrad.)
Schult., with numerous, clavate stamens, to which would belong C. chelidonii.
According to Iltis the Malaysian species are to be grouped as follows: sect. Gymnogonia R.Br.:
C. gynandra; sect. Tarenaya (Raf.) Iltis: C. spinosa, C. speciosa, C. aculeata; sect. Ranmanissa (Endl.)
Griseb. C. \iscosa; sect. Corynandra (Schrad.) Schult.: C. chelidonii; sect. Rutidosperma Iltis:
:

C aspera, C. rutidosperma.
Dr Iltis (Brittonia 10, 1958, 33 and 12, 1960, //; the press) is also of opinion that there are no good
reasons to maintain the genus Gynandropsis, because the connation of the staminal base with the gyno-
phore to form an androgynophore is merely a character of degree, of quantitative value. In this last
conclusion Iltis was preceded by Woodson, who suggested to refer the bisexual species of Gynandropsis
to Cleome, and proposed an emendation of the S. American genus Podandrogyne to incorporate the
monoecious species of Gynandropsis (Ann. Mo. Bot. Gard. 35, 1948, 139-141).
As to Polanisia, Iltis pointed out that it is satisfactory to restrict Polanisia to the original concept of
Rafinesque, the type species being the N. American P. graveolens Raf. (= P. dodecandra (L.) DC).
de Candolle, followed by many later authors, had extended the concept with the cleomoid Old World
species possessing more than 6 stamens, but these possess no large adaxial gland as in the type species.
These should be referred to Cleome.
Thanks are due to Dr H. H. Iltis, who kindly put his MS at my disposal and gave additional crhical
remarks, enabling me to take advantage of his extensive knowledge of the American and other species.

key to the species •

1. Androgynophore longer than 5 mm.

2. Gynophore in flower 1-2 mm, in fruit 4-10 mm. Petals 7-15 mm long, open in bud, white.
1. C. gynandra
2. Gynophore c. 6 cm. Petals 25-35 mm long, imbricate, pink 2. C. speciosa
1. Androgynophore none or at most 3 mm long.
3. Gynophore c. 4 cm. Androgynophore 1-3 mm high. Plant with scattered prickles. 3. C. spinosa
3. Gynophore shorter than cm, or none. Stamens free.
1

4. Stamens more than 8. Ovary sessile.

5. Stamens 30 or more, filaments club-shaped. Fruit with a narrowed base; valves parallel-nerved.

Plant with scaly hairs. Flowers pink

stiff 4. C. chelidonii
5. Stamens lessthan 30, filaments almost filiform. Fruit hardly narrowed at the base, valves centri-
petal-nerved. Flowers yellow 5. C. viscosa
4. Stamens 6. Ovary on a short gynophore.
6. Plant with recurved stipular thorns, glabrous or glandular-puberulous. Flowers white to cream.
6. C. aculeata
6. Plant unarmed, with scattered, retrorse hair-like appendages, not glandular.
7. Petals violet-blue, 9-12 mm mm
long. Fruit 4 diam. Seeds with an open cleft.
7. C. rutidosperma
7. Petals white, 4-5(-7)mm long. Fruit 2mm diam. Seeds orbicular, with a closed cleft.
8. C. aspera
 Dec. 1960] Capparidaceae (Jacobs) 101

1. Cleome gynandra Linne, Sp. PI. 2 (1753) 671; many superficial concentric ribs and numerous
C. B. Robins. Philip. J. Sc. 3 (1908) Bot. 182.- irregular distinct cross-ribs. No elaiosome.
Logunsa nibiu Rumph. Herb. Amb. 5 (1747) 280, Morph. Raghavan found in this species (in
t. 96 f. 2.— C. tripliylla Linne, Sp. PI. ed. 2 (1763) S. India) that about 50 %
of the ovaries are
938.— C. peiitapltyila Linne, I.e. 938; R. Br. in sessile and sterile with only vestiges of ovules
Oudney, Denh. & Clapp. Narr. Trav. Disc. Afr. being present (J. Linn. Soc. Bot. 52, 1939, 239).
(1826) 222. Pedicellaria peiUciphylla Schrank in Mauritzon studied the embryology (Ark. Bot.
Roem. & Ust. Mag. Bot. 3, pt 7 (1790) 1 1 ; Merr. 26, /;. 15, 1935, 1).

Publ. Govt Lab. Philip, n. 11 (1905) \^.—Gynan- Distr. From Ceylon and the Punjab through-
dropsis pentaphylla DC. Prod. 1 (1824) 238; Miq. out SE. and E. Asia as far as Peiping; throughout
PI. Jungh. (1854) 397; Fl. Ind. Bat. 1, 2 (1858) 96; Malaysia. Widely introduced in the New World.
Sum. (1860) 19; Eichl. Fl. Bras. 13, 1 (1865) 261, Ecol. A weed, in dry rice-fields, along road-
t. 58 f. 3; Hassk. Neu. Schluss. Rumph. (1866) sides, near houses, from the lowlands up to c.
263; Miq. Illustr. (1870) 19; Fern.-Vill. Nov. 500 m. Fl. fr. Jan.-Dec.
App. (1880) 10; King, J. As. Soc. Beng. 58, ii Vern. Maman(g), ma mam, M, S, bobowan,
(1889) 392; Merr. Philip. J. Sc. 1 Suppl. (1906) entjeng-entjeng, J, bhubliuwan, Md; Philippines:
58; Back. Fl. Bat. (1907) 53; Schoolfl. (191 1) 60; apoi-apoian, balabalanoian. Tag. lialaya, hiilaya, ;

Merr. FI. Manila (1912) 216; Laut. Bot. Jahrb. P. tantandok, t.-a-dadakel. Ilk.
Bis.,
52 (1915) 110; Merr. Int. Rumph. (1917) 241; Uses. Some minor medicinal applications are
Sp. Blanc. (1918) 158; Ridl. Fl. Mai. Pen. 1 mentioned by Burkill, I.e. and Heyne, I.e.
(1922) 120; v. d. Pul, Trop. Natuur 19 (1930) 162. According to Ochse, I.e., the bitter leaves are
— Gyiwndropsis affinis Bl. Bijdr. 2 (1825) 51. prepared by boiling and salting as a vegetable,
C. ajfinis Spreng. Syst. Veg. ed. 6, 4, 2 ( 827) 138,
1 1 especially in Java.
—
uon DC. 1824. C. blumeaim Schult. Syst. Veg.
7' (1829) 23.— C. alliacea Blanco, Fl. Filip. 2.Cleome speciosa Raf. Fl. Ludovic. (1817) 86
(1837) 522.— C. hhimecwa D. Dietr. Syn. PI. 2 H.B.K. Nov. Gen. & Sp. PI. 5 (1821) 84, t. 436
(1840) 1065.— C. alliodura Blanco, Fl. Filip. ed. 2 — Gynandropsis speciosa DC. Prod. I (1824) 238
(1845) 363, ed. 3, 2 (1879) 307, t. m.—Gynandrop- Hassk. Nat. Tijd. N.I. 10 (1856) 118; Hort. Bog
sis gynandra Briq. Ann. Cons. Jard. Bot. Geneve Descr. (1858) 11; Miq. Fl. Ind. Bat. 1, 2(1858)96
17 (1914) 382; Merr. En. Philip. 2 (1923) 209; Fern.-Vill. Nov. App. (1880) 10; Merr. Philip
DocTERS VAN Leeuwen, Zoocecidia (1926) 211; Govt. Lab. Publ. /;. 6 (1904) 24, /;. 27 (1905) 18
Heyne, Nutt. PI. (1927) 681 Ochse & Bakh. Ind.
; Back. Fl. Bat. (1907) 54; Gagn. FI. Gen. I.-C. 1

Groenten (1931) 96, f. 57; Back. Onkr. Suiker. (1908) 174; Back. Ann. Jard. Bot. Buit. Suppl. 3
(1931) 253, Atlas t. 264; Trop. Natuur 22 (1933) ( 9 0) 403
1 1 Schoolfl. ( 9 ;) 60 Briq. Ann. Cons.
1 1 1 ;

112; BuRK. Diet. (1935) 1119; Pax & Hoffm.

1 Bot. Geneve 17 (1914) 384; Schroo, Trop.
in E. & P. Pfl. Earn. ed. 2, I7b (1936) 218; Back. Natuur 4 (1915) 108, f. 1-7; Merr. Sp. Blanc.
Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, p. 4; (1918) 158; En. Philip. 2 (1923) 209; Docters van
Merr. PI. Life Pac. World (1946) 135, f. 119. Leeuwen, Zoocecidia (1926) 211; Heyne, Nutt.
Erect, mostly widely branched, annual herb, PI. (1927) 681; Burk. Diet. 1 (1935) 1119; Back.
15-80 cm. Stem glandular-pubescent to glabrous. Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, p. 4.
Normal leaves with 5 leaflets, lowest and upper — C. speciosissima Deppe ex Lindl. Bot. Reg. 14
with 3, towards and in the inflorescence diminish- (1831) t. 1312; Burk. Diet. (1935) 5^0.— Gynan-
ing in size; leaflets thinly herbaceous, about twice dropsis pentaphylla (non DC.) Blanco, Fl. Filip.
as long as wide, obovate, e. 1-1 Vi by 1-3% cm, (1837) 523. —
C. gigantea (non L.) Blanco, ibid.
base cuneate, top rounded and ± distinctly ed. 2 (1845) 364, ed. 3, 2 (1879) 307, t. 234.
acuminate, ciliate to denticulate; nerves 5-8 pairs; Little-branched herb, c. I-I V2 m, erect, glabrous
petiole 2-10 cm, petiolules 1-3 mm, webbing at or glabrescent. Leaflets 5-7, herbaceous, sub-
the base, densely glandular puberulous. Flowers sessile, slightly webbing at the insertion, narrowed
in long, corymbose racemes, nocturnal. Pedicels towards base and top, lanceolate, c. 10-12(-I8)
thin, \V2-IV2 cm, glandular-puberulous. Sepals by 2 1/2-4 V2 cm; nerves 14-16(-25) pairs; petiole c.
2'/2-5 by Vz-^Vz mm, acute, puberulous, ciliate. 8-12(-17) cm. Raceme up to 40 cm long. Flowers
Petals with open aestivation, all pointed upwards subtended by subsessile leaves upwards gradually
towards the adaxial side, elliptic with narrowed simplified and diminishing in size. Pedicels thin,
base and rounded top, 1 Vi-S mm
stalked, 7-15 e. lYz-iVi cm. Buds cylindric, c. 1-lVi cm by
mm long in all, 1 '/2-4 mm
broad. Androgynophore 3 mm; petals imbricate, androgynophore with the
9-16 mm. Stamens 6; filaments 1 Vz-l cm; anthers base of the filaments bulging out at the anterior
linear, 2-3 mm long. Gynophore 1-2 mm; ovary side shortly before anthesis; flowers opening at
cylindric, c. 3 by 1/2 mm, in some flowers (es- dusk. Sepals patent, subulate, 5 mm, ciliate. Torus
pecially the apical ones) sessile and reduced. In c. 1
1/2 mm
wide. Petals during anthesis pointing
fruit pedicels 1-3 cm, androgynophore 13-20 mm, upwards adaxially, c. (25-)30(-35) by 5-8 mm,
gynophore 4-10 mm. Fruit cylindrical, tapering lanceolate, rounded, narrowed into a claw c.
to both ends, 2-11 cm by 3-4(-6) mm; beak 1-4 5 mm, glabrous, pink or white (sometimes referred
mm; valves with longitudinal, centripetal veins. to as /. alba). Androgynophore 5-7(-9) mm,
Seeds depressed-globular, c. '/j mm
diam., with
1 slightly incrassate towards base and top, glabrous.
a shallow and narrow cleft, black-brown, with Stamens 6, glabrous, filaments filiform, c. 5-6 cm.
102 Flora Malesiana [ser. I, vol. 6^

anthers c. 1 mm, linear. Gynophore c. 6 cm, related C

houtteaua Schlecht. (Linnaea 8, 1851,
glabrous, ovary cylindrical, few mm long, gla- 669) which would be recognizable by glabrous
brous. Fruit cylindric, c. 8-9 cm by 3 mm, valves buds and ovaries, and pink flowers.
indistinctly parallel-nerved. Seeds c. 2% mm
diam., 2 mm thick, with a fairly shallow cleft, 4.Cleome chelidonii Linne
Suppl. (1781) 300;
/.
light brown, the surface with small, pale, scattered Hook. /. & Th. Fl. Br. (1872) 170.—
Ind. 1

scales; no elaiosome. Polanisia chelidonii DC. Prod. 1 (1824) 242; MiQ.

Distr. South America from Mexico to Peru Fl. Ind. Bat. 1, 2(1858)97; Back. Schoolfl. (1911)
and Guyana, cultivated as an ornamental in 60; Heyne, Nutt. Pi. (1927) 682; Back. Onkr.
SE. Asia, up to 1500 m, in Malaysia collected in Suiker. (1931) 254, Atlas t. 264; Bekn. Fl. Java
Sumatra (East Coast), Java, Borneo (Sarawak, (em. ed.) 4A (1942) fam. 45, p. 5. Polanisia
W. Borneo), Philippines (Luzon), Moluccas (Aru angulata DC. Prod. (1824) 242; Miq. Fl. Ind.
1

Is). According to Hasskarl (1856, I.e. 119) intro- Bat. 1, 2 (1858) 97; Illustr. (1870) 20.— Fig. 31,
duced in Malaysia some years before 1855. 32a-b.
Ecol. ScHROO, I.e., observed that in Java the Widely branched herb, 15-80 cm; tap-root stout,
flowers open between 4 and 5 p.m.; small stingless white. Stems angular, with sparse, appressed, pale,
bees (Apis indiea) collect pollen but do not touch stiff hairs with a bulbous base. Leaflets firmly

the nectar; pollination would be effected by herbaceous, 6-7 below to apically only 3 or 1,
nocturnal butterflies. obovate, mostly densely appressed-hairy, central
leaflet largest, up to 4(-6) by li4(-l%) cm; base
3. Cleome spinosa Jacq. En. PI. Carib. (1760) 26; cuneate, top rounded and subacuminate to acute;
LiNNE, Sp. PI. ed. 2 (1762) 939; DC. Prod. 1 nerves 4-5 pairs; petioles upwards gradually
(1824) 239; Eichl. F1. Bras. 13, 1 (1865) 252; decreasing in length from c. 8-10 cm to almost
Merr. En. Philip. 2 (1923) 208; Back. Bekn. Fl. zero, hairy as the stem, apex and petiolules white.
Java (em. ed.) 4A (1942) fam. 45, p. 2.— C. Raceme corymbiform, flowers subtended by
sandwieensis A. Gray, U.S. Expl. Exp. Bot. 1 reduced leaves, actinomorphic. Pedicels (l-)l'/4-3
(1854) 65. cm, hairy as the stem. Buds ellipsoid, ±. obovate,
—
Herb 1 1|4 m. Stems vigorous, glandular- acute, 6-10 mm
long. Sepals narrowly imbricate,
pubescent. Stipular thorns minute (some petioles appressed, elliptic (to obovate), acuminate, 2-4
without) to 4 mmlong, sharp, recurved. Leaves mm long, sparsely scaly-hairy outside, margin
much reduced towards the inflorescence; leaflets membranous. Petals 4(-8), mostly obovate with
5-7, herbaceous, lanceolate, slightly webbed at the narrowed base and rounded top, 7-12(-15, in
base, sparsely glandular-hairy, central leaflet India -21) by 3-5 mm, glabrous, light red-purple.
largest, 6-8(-10) by l%-2|4(-3) cm; base cuneate, Stamens 30-40(-55), somewhat shorter than the
decurrent, top attenuate, acute, mucronate; petals, glabrous; filaments with a thickened top;
nerves 10-15 pairs; petiole c. 5-10 cm, sometimes anthers c. 1 mm, yellow. Ovary linear, about as
with scattered prickles as the midrib beneath. long as the stamens, glabrous. Fruit linear,
Racemes up to 40 cm. Flowers subtended by sub- parallel-nerved, narrowed at the very base, gla-
sessile, ovate-oblong, sparsely glandular-hairy brous, c. 1-3 mm
beaked. Seeds asymmetrical
bracts, gradually diminishing in size from 2 to nearly 2 mm, cleft open, dull blackish, not ribbed
V2 cm. Pedicels 2-3 cm, short glandular-hairy. but warty by scattered scales mainly on the dorsal
Buds cylindrical, c. 2% cm by 4 mm, glabrous; side. Elaiosome wanting.
petals imbricate, androgynophore with the base of Distr. India, Burma and Siam (locally), in
the stamens bulging out at the anterior side Malaysia: Central and East Java.
shortly before anthesis. Sepals patent, narrow- An at. T. S. Raghavan investigated the
triangular, (4— )6-7 mm, glandular-hairy outside. development of the female gametophyte, embryo,
Petals glabrous, on a filiform claw (5-) 10-1 2 mm, and seed, and the vascular supply of the floral
blade oblong, ±asymmetrical, 10-17 by 4-6 mm, parts (J. Linn. Soc. Bot. 51, 1937, 43-72; ibid. 52,
with cuneate base and rounded top. Stamens 6; 1939, 249).
androgynophore 1-2 mm, glabrous; filaments Ecol. Fallow sawahs, sugarcane-fields on
filiform, 3'/2-4 cm; anthers linear, 7-8 mm. heavy clays, and marls periodically drying out
Gynophore glabrous, in flower c. 4 cm, in fruit to during the pronounced dry season, in Java below
51/2 cm; ovary linear, e. 4 mm, glabrous. Fruits 100 m, locally sometimes so abundant that the
patent, cylindrical, blunt at both ends, 5V^-6V^ fields are coloured red-purple by the flowers
cm by 4 mm, valves finely and densely nerved. (Backer, 1931). Fl.fr. Jan.-Dec.
Seeds asymmetrical, 2 mm diam., almost smooth, KooPER defined a weed community characteriz-
concentric and cross ribs subprominent; elaiosome ed by Polanisia chelidonii on constantly moist,
not present. fairly to very heavy clay in sugarcane-fields (Rec.
Distr. Native in tropical America, also oc- Trav. Bot. N6erl. 24, 1927, 84 seq.).
casionally cultivated in the tropical to the tem- MiRASHi found it characteristic in the vegetation
perate zones, in Malaysia: occasionally cultivated, of freshwater swamps in India; he discussed also
known from Luzon and from Java, in the latter some anatomical details (Proc. Ind. Ac. Sc. 43B,
island once found run wild. 1956, 233-236).
Note. According to Iltis (in litt.) the Malaysian Note. The type material o{ Polanisia angulata
material would not belong to C. spinosa, but to the DC, hailing from Java, could not be located in
Dec. 1960] Capparidaceae (Jacobs) 103

the Paris Herbarium; his description is vague, ing. Leaflets tinhly herbaceous, 5-3, diminishing
but he records a field note by Leschenault upwards in size, oblong, the central
subsessile,
"flowers violet". This character occurs only in leaflet c. l-3(-5'/2) by ('4-)'/2-l(-2) cm; base
cuneate, top acute to obtuse; nerves 3-6 pairs;
petiole cm. Racemes corymbose;
('/2-)l-3(-6)
flowers in the of reduced leaves, largely
axils
actinomorphic, opening in the morning, closing
in the afternoon, ephemeral. Sepals oblong,
blunt to acute, (2"/2-)6-7(-8) by ('/2-)l-3 mm.
Petals yellow (once reported white), thin, glabrous,
oblong, (4-)7-12 by (l'4-)3-5 mm, base cuneate
or i clawed, top rounded. Stamens (8-) 10-20
(-30), glabrous; filaments (3-)5-7 mm
towards the
abaxial side, gradually increasing in length by

Fig. 31. Cleome chelidonii L. /. (India, Raipur,

Hewetson, 1950).

C. chelidonii; therefore, I agree with Backer in

referring it here.

5. Cleome viscosa Linne, Sp. PI. 2 (1753) 672;

Hook./. & Th. F1. Br. Ind. (1872) 170; F. v. M.
1

Descr. Not. Pap. PI. P (1876) 52; Ridl. F1. Mai.

Pen. 1 (1922) \\9.-'Lagciiisa alba Rumph. Herb.
Amb. 5 (1747) 280, t. 96 f. 3.— C. icosandra
Linne, Sp. PI. 2 (1753) 672; in Stickman, Herb.
Amb. (1754) 21; Burk. Diet. 1 (1935) 581.—
Polaiiisia viscosa DC. Prod. 1 (1824) 242; Bl.
Bijdr. (1825) 52; Blanco, F1. Filip. ed. 2 (1845)
364, ed. 3, 2 (1878) 308; Miq. F1. Ind. Bat. 1, 2
(1858) 97; lllustr. (1870) 20; Fern.-Vill. Nov.
App. (1880) 10; ViDAL, Phan. Cuming. (1885) 94;
Back. F1. Bat. (1907) 52, c/uoad var. a; Merr.
Philip. J. Sc. 3 (1908) Bot. 407; Back. Schoolfl.
(1911) 60; Merr. F1. Manila (1912) 216; Int.
Rumph. (1917) 240; Sp. Blanc. (1918) 158;
Heyne, Nutt. PI. (1927) 682; Back. Onkr. Suiker.
(1931) 255, Atlas t. 266, quoad/. typica.—Polanisia
icosandra W. &
A. Prod. (1834) 22; Merr. En.
Philip. 2 (1923) 209; Back. Bekn. Fl. Java (em.
ed.) 4A (1942) fani. 45, p. 5, quoad f. tvpica; Merr.
Plant Life Pac. World (1946) 135, f. 121; Quis.
Med. PI. Philip. (1951) 344.— C. acutifolia Elmer,
Leafl. Philip. Bot. 7 (1914) 2574.— C. chelidonii
(nan L. /.) Burk. Card. Bull. S.S. 3 (1924) 280;
Diet. 1 (1935) 580.— Fig. 32c-d. Fig. 32. Fruit and seed of Cleome. a-b. C. cheli-
Annual, mostly widely branched herb up to donii L. /., c-d. C. viscosa L. (a, c nat. size, b,
1 m, glandular, yellowish hairy, viscid and stink- d X 10).
104 Flora Malesiana [ser. I, vol. 6^

1-2 mm, not or only very slightly swollen at the /. deglabrata (Back.) Jacobs, stat. nov. —
top; anthers linear, c. 1 '/2-2 rnm. Ovary sessile, c. Polanisia viscosa var. deglabrata Back. Fl. Bat.
3-10 mm beaked, minutely glandular-hairy. (1907) 53; Domin, Bibl. Bot. Heft 89 (1925) 683.—
Fruit erect, (i/3-j2-3(-4) cm pedicelled, (lV2-)6-8 Polanisia icosandra f. deglabrata Back. Bekn.
(-10) cm by (2-)3-4'/2 mm, beak IVi-M-D mm; Fl. Java (em. ed.) 4A (1942) fam. 45, p. 5.
valves with distinct centripetal nerves. Seeds c. Differs from/, viscosa in being entirely glabrous;
11/4 mm
diam., 1 mm
thick, red-brown, cleft not with a distinct smell.
narrow, with strong cross-ribs and faint con- Distr. Malaysia: Malay Peninsula, Java.
centric ribs. No elaiosome.
6. Cleome aculeata Linne, Syst. Nat. ed. 12, 3
Distr. Native in the Old World, from tropical
(1768) 232; DC. Prod. 1 (1824) 241; Hassk. Nat.
Africa and S. Arabia to tropical Australia;
Tijd. N.I. 10 (1856) 1 19; Hort. Bog. Descr. (1858)
common throughout Malaysia, commonly ad- 12; MiQ. Fl. Ind. Bat. 1, 2 (1858) 96; Eichl. Fl.
ventive in the New World.
Bras. 13, 1 (1865) 259, t. 58 f. 2; Back. Ann. Jard.
Ecol. A common, very tolerant, ruderal plant, Bot. Buit. Suppl. 3 (1910) 398; Schoolfl. (191 1) 59;
on fallow along roadsides, on rubbish-
land, RiDL. Fl. Mai. Pen. 1 (1922) 120; Back. Bekn.
heaps, in fields, etc., often on sandy, sometimes Fl. Java (em. ed.) 4A (1942) fam. 45, p. 2.— C.
on calcareous soil, both under seasonally dry and hulletii KiNG, J. As. Soc. Beng. 58, ii (1889) 392;
everwet climatic conditions, sometimes near the Ann. Bot. Card. Calc. 5 (1896) 117, t. 133A.
coast or in savannahs, up to 500 m, mostly lower. — Fig. 33a.
Fl. fr. Jan. -Dec. Erect, widely branched, annual herb, Va-Vi m,
According to Ridley in Malaya dispersed rather densely covered with minute, glandular
endozoically by water-buffalos (Disp. p. 368). hairs, neither viscid nor smelling. Stem with
FosBERG observed in the Marianas that the seeds fulvous, recurved, stipular thorns e. 1 Yj mm.
were eaten by birds. Leaves upwards gradually reduced in size, ulti-
KooPER found on the sugarcane-fields in East mately simple and subsessile. Leaflets 3, sub-
Java a weed community, characterized by Polanisia sessile, thinly herbaceous, oblong, sometimes
viscosa, typical for recent, very pervious, volcanic, obovate, central leaflet c. 2i/2-3(-3!/2) by IVa-Wa
light soils valuable for sugar culture. He recorded (-2) cm; base acute to cuneate, slightly decurrent,
wilting experiments with this species under dry top acute; nerves 5-8 pairs; petiole 2-3(-4'/2) cm.
conditions (Rec. Trav. Bot. Neerl. 24, 1927, 56 Racemes few-flowered, flowers subtended by
seq., 218 seq.). simple, subsessile leaves. Pedicels 1 V2-2 cm.
Mr N. G. BissET found the seeds containing an Sepals linear, 1 1/2-214 by Va-Vi cm, minutely
appreciable amount of alkaloid. glandular-hairy outside. Petals P4-4 mm (incl.

Uses. There is a record from Java and one claw Ya-\ mm), obovate, pale yellowish. Stamens
from Hainan stating that the sap of the leaves 6, slightly exceeding the petals, anthers 1 Vi-l mm.
with water or milk is applied on the eyes. In Perak Ovary subsessile, cylindric, 2-3 mm long, gla-
the herb is rubbed on the body against rheuma- brous. Fruit on a gynophore V2-2V2 mm, cylindric
tism. In Central Sumatra the leaves and seeds are but abruptly tapering to both ends, 31/2-4 cm by
added to tobacco to stress its narcotic properties. 3 mm; valves finely parallel-nerved. Seeds asym-
Besides, the plant finds a number of other minor metrical, with closed cleft, 2-21/4 mm
diam.;
medical applications; see Heyne {I.e. p. 682) and concentric ribs faint, obtuse, cross-ribs irregular,
QUISUMBING {I.e. p. 346). scattered, incomplete, distinctly prominent; elaio-
some distinct.
Vern. Daun maman pantai, machmud panta,
Distr. Native from Mexico to N. Argentina,
maman panta, maman patih, Perak, poko kutepeng,
introduced in Malaysia: W. Java (Bogor and
Malacca, maman hiitan, M, nai ve/ai, Tamil
vicinity. According to Hasskarl (1856, I.e.)
(Mai. Pen.), dek tail eluin, Penang, daun gliengang
incidentally introduced with soil from Suriname
ajam, Palembang, maman, mamman, S, antjang
into the Hortus Bogoriensis, whence it escaped
antjang, bobowan, entjeng entjeng, etjeng, etjeug
before 1889), Malay Peninsula (Singapore), New
tembeking, J, bhubhuan, Kangean, pupuan loke,
Guinea (Papua: Kanosia, one record of 1935).
tjongblentjongan, Md, aluiru, Nenusa Is, susawi
utan, Ambon, poompito, N. Celebes; Philippines:
Cleome rutidosperma DC. Prod. (1824) 241;
7. 1
apoi-apoian, halabalanoian, silisilihan. Tag., huldya,
tuldyag, P. Bis., kabdu, Iv., lampotaki, Tagb.,
Iltis, Brittonia 12 (1960) in the press. — C. ciliata
ScHUM. & Thonn. Dansk Vid. Selsk. Afli. 4
tandandok. Ilk.
(1828) 67; Jochems, Trop. Natuur 17 (1928) 80,
Notes. BuRKiLL recorded C. ehelidonii for the f. 1, 2; Beumee, Trop. Natuur 18 (1929) 99;
Malay Peninsula (Card. Bull. S.S. 3, 1924, 280; Pax & HoFFM. in E. & P. Pfl. Fam. ed. 2, 17b
see also Diet. p. 580). The material he cited (1936) 213; Back. Bekn. Fl. Java (em. ed.) 4A
belongs, however, to C. viscosa, partly to its /. (1942) fam. 45, p. 3; Johnson & Tan, Gard.
deglabrata. Bull. Sing. 17 (1959) 325-330, fig. 1-2.— Fig. 30,
As far as could be established de Candolle 33b.
was the first to combine C. icosandra and C. Erect to spreading, widely branched, annual,
viscosa and, under Polanisia, he chose the epithet odourless herb, 14-I m; stem, petioles, and the
viscosa (1824). nerves underneath with sparse prickle-like, softish
Dec. 1960] Capparidaceae (Jacobs) 105

appendages up to 2 mm long. Leaflets 3, upwards growing in trees, where the seeds were obviously
gradually reduced and subsessile, thinly herbace- brought by ants attracted by the fatty elaiosome.
ous, ovate to subrhomboid or oblong-lanceolate, About one third of the ovaries is 2-3 mm
ciliate,central leaflet 3-3i/2(-5) by l!/2-2i/2 cm; long and sterile.
base cuneate, decurrent, top narrowed, acute to Vern. Seni walai, Tamil (Malaya).
blunt; nerves 6-9 pairs; petiole generally 3-4(-5) Note. The exact synonymy I owe to Dr Iltis
cm, slightly webbing between the short petiolules. (unpublished) who found that the type specimen
Racemes with reduced leaves. Pedicels filiform, of C. rutidosperma must have actually come from
2-3 cm, in fruit c. 3 cm, rather densely set with Sierra Leone, Africa, but was erroneously labelled
minute, gland-like hairs. Sepals 31/2-4 by Va-Va by DE Candolle "Antilles" and described by him
mm, ovate, acute to acuminate, hairy like the as from "Tabago".

8. Cleorae aspera Koen. ex DC. Prod. 1 (1824)

''''i^*''''^^.
241; Hook./. &Th. F1. Br. Ind. 1 (1872) 169;
Back. Onkr. Suiker. (1931) 252, Atlas t. 262;
Bekn. Fl. Java (em. ed.) 4A (1942) fam. 45, p. 3.
"9
— Fig. 33c.
Erect, widely branched, annual, stinking herb,
< C
!4-% rn high. Stem, petioles, and nerves under-
neath sparsely set with small prickle-like, softish
appendages. Leaflets 3(-5), upwards reduced and
Fig. 33. Seeds of Cleoine. a. C. aciileata L.,
becoming subsessile, on short petiolules webbing
b. C. rutidosperma DC, c. C. aspera Koen. ex DC,
at the insertion, thinly herbaceous, ovate-lan-
all X 8.
ceolate, ciliate, central leaflet generally 21/2-3
(-51/2) by ^4-l(-li4) cm; base cuneate, top blunt;
nerves below. Petals violet-blue, all pointing up- nerves 4-9 pairs; petiole generally [Vj-lVii-^)
wards at the adaxial side, 9-12 (including a mm cm. Racemes with reduced leaves. Pedicels fili-
claw 2-3 mm) by IVi-^Vi mrn. Stamens 6, 7-10 form, Vi-^Vi cm, 11/2-2 cm in fruit, rather
mm; anthers linear, 2 mm, strongly curved after sparsely set with minute glandular hairs. Sepals
anthesis. Gynopliore c. 1 'A mm, ovary linear, c. 2-3 by /4-'/2 mm, acuminate to obtuse, glandular-
8-10 mm, slightly S-curved, minutely glandular, hairy. Petals white, 4— 5(-7) (including claw 1-2
beak c. mm. Fritit on a gynophore 4-8(-10) mm,
1 mm) by V^-lVz mm, oblong-lanceolate, top
cylindrical, tapering at both ends, 5-7'/2 cm by rounded. Stamens 6(-7), 4-6 mm; anthers c.
4 mm, beak 1-4 mm, valves glabrous, parallel- 1 mm, linear, curved after anthesis. Gynophore c.
nerved. Seeds dull black, asymmetrical, with a 1 mm; ovary 3-4 mm long, linear. Fruit on a
pale centre, with faint, obtuse concentric ribs and gynophore 3-6(-10) mm, 3-6 cm by 2 mm,
stronger cross-ribs, cleft open, P/i by 1 '/2 nim; cylindrical, tapering to both ends, the beak 2-8
ribs sometimes with microscopical bristles; base mm; valves parallel-nerved, glabrous. Seeds sub-
of the seed with a white elaiosome. orbicular, c. 11/2 by P/4 mm, dark-brown, with
Distr. West tropical Africa (from Guinea to small paler centre, with very narrow-elevated,
Angola and the Congo), introduced in the strongly prominent cross-ribs, and distinct obtuse
Caribbean region and occasionally found in concentric ribs, cleft closed; no elaiosome.
Malaysia: first records 1920: Belawan, Sumatra Distr. Native in India and Ceylon, in Malaysia:
East Coast; 1924: Singapore; 1946: Java (Tand- Central and East Java, Madura, and Bali.
jung Prick); 1958: wide-spread from Sarawak to The oldest Malaysian collection is by Hors-
N. Borneo. In 1946 found in Siam (100 km NW. field from Java between 1802 and 1819; the
of Kanburi), in 1948 in Burma (Insein Distr.) and second one is from Pekalongan, in 1912.
in Manila. Ecol. Obviously bound to a distinctly seasonal
Ecol. A lowland ruderal in the course of ex- climate.
tending its area with a tendency to become a Vern. ilntjeng entjeng-kebo, J (in use for all
common plant. Beumee, I.e., observed the plant Cleomoideae).
 CAMPANULACEAE (B. Moeliono, Amsterdam & P. Tuyn, Leyden)

Annuals, perennials, more rarely shrubs, small trees, or vines, mostly laticiferous,
sometimes with subterranean tubers. Leaves exstipular, simple, entire or toothed
to incised (rarely pinnatifid), spirally arranged or alternate, rarely opposite. Flowers
often blue, violet, red, or white, frequently protrandrous (rarely dioecious), axillary
or terminal, solitary or in mostly bracteate, racemose inflorescences (rarely cymes),
bisexual (rarely unisexual or dioecious), isomerous, mostly 5-merous, regular or
symmetric. Pedicels mostly with 2 bracteoles. Calyx segments mostly free, often
persistent, valvate. Petals connate to various degree, sometimes almost free (ex-
ceptionally free), valvate in bud; in strongly zygomorphous flowers the corolla
bilabiate dorsally slit and the lobes often very unequal, the lower lip often with 2
convexities near the base. Stamens adnate to the corolla or free from it, mutually
mostly partly connate (either the filaments or part of them and the anthers or
only the latter); filaments often widened at the base; anthers introrse, in zygo-
morphic flowers often unequal, often 2 or more with apical setae, further glabrous
or haired. Disk epigynous, mostly free. Ovary inferior or partly so (rarely superior),
2-5-celled. Style 1, often with hairs below the 2-5 stigmatic lobes. Ovules oo, mostly
on axile placentas (exceptionally parietal in incompletely celled ovaries). Fruit
capsular or a berry, or berry-like, mostly dehiscing at the apex with valves, or circum-
sciss. Seeds co; embryo straight; albuminous.
Distribution. Rather large family, with a worldwide distribution, with approximately 60-70
genera and roughly between 1000 and 1500 spp., the largest ones. Lobelia and Campcinitlci, counting
several hundreds of species. In Malaysia the family is sparsely represented although with one endemic
genus, Phyllochaiis, in New Guinea, and a subendemic one, Pentaphragma.
Two genera show a remarkable trans-Pacific distribution, viz Laureutia sect. Isotuma with the majority
of species Australian and one species in tropical America, and the Piatia aflfinity of Lobelia which is a
distinctly South Pacific-Subantarctic group.
Codoiwpsis on the other hand is typically East Asian, mainly Chinese, and Pcracarpa is Sino-Japanese.
Wahlenbergia is worldwide distributed. Splienoclea is a monotypic Old World swamp plant.
Ecology. The family is but little represented in the tropical lowland of Malaysia; most species
occur in the hills or in the montane zone. The highest localities are reached by Wahlenbergia confiisa
Merr. & Perry in New Guinea between 3500 and 4000 m, while some other Lobelia species are well
up in the subalpine zone.
As to the climate almost all representatives belong to the undergrowth of the rain-forest. Exceptions
are Splienoclea which grows on damp soils and is indifferent to climate. Lobelia alsinoides which is bound
to a seasonal lowland climate, and Lobelia nicotianaefolia, Wahlenbergia marginata, and W. hookeri
which are obviously bound to a seasonal montane climate.
No Malaysian Campaniilacea occupies a very special ecological niche and none occurs gregarious
to an extent worthy of note.
Flower biology. Nothing is known about this subject for Malaysian representatives. It is certain that
protrandry prevails in the family; the pollen is often shed in bud and the growing style, which is mostly
provided with hairs, pushes the pollen up above the anthers. It has been said that apids collect it and
would effect cross-pollination. In Wahlenbergia the stamens shrivel very soon. The stigma opens at a
much later stage. Autogamy is, however, not entirely excluded in this way. Docters van Leeuwen
(1933) emphasized that in Wahlenbergia marginata autogamy is the rule.
The long-tubed Lobelias are said to be adapted to be visited by birds and L. nicotianaefolia might fail
under this category, but no data are available on this species. It is remarkable that also L. montana has
scentless flowers; neither Docters van Leeuwen (1933) nor I myself have ever observed insects visiting
the showy flowers. As fruit production is abundant, Docters van Leeuwen concluded to successful
autogamy.
Seed dispersal. Part of the genera have capsular fruits, part of them possess berries; in Peracarpa
the fruit is indehiscent. In dl cases the seeds are numerous and small. As all but one are of delicate
or creeping habit and live mostly in shaded habitats the chance that wind may have an appreciable
effect can be neglected, save for the few subalpine Lobelias and Wahlenbergia which are exposed to
mountain winds.
Also for the species with berries the way of dispersal is rather obscure. I have often found the very

(107)
 108 Flora Malesiana [ser. I, vol. 6^

large showy berries of Codonopsis javanica and Lobelia montana entirely untouched and shrivelling
on the plant. There is, therefore, no indication that seed dispersal is likely to cover large distances.
Only for Lobelia aiigulata Ru:)Ley (Disp. 1930, p. 508) recorded that its small berries are, according
to the ornithologist H. C. Robinson, a favourite food of the tree partridge, Arborophila campbellii.
The distribution of this mountain bird coincides indeed remarkably well with the localities of the plant:
from the Himalaya to Formosa and the Sunda Islands. And Dr Junge tells me that there are allied
Pliasianinae in East Malaysia and Australia; in South America this group is replaced by the subfam.
OdontoplwriiKie which may take over dispersal. However, we have no proof that the seeds pass the
intestinal duct undamaged and that the eating of the berries is indeed effective for the intensive and
wide dispersal of the seeds.
Lobelia chinensis is often seen along stream banks and this would point either to dispersal of seeds
by water, or to aquatic transport of rhizomes.
Laurentia longiflora which is introduced and very common on ruderal places is also often found in
damp places near streams and ditches; its seeds may be locally transported by water, as they are said
to be water-repellent. It is sometimes also found on old walls which would point to dispersal by ants,
but as far as I know the seeds of Campanulaceae have no aril or elaiosome.
Anatomy. Metcalfe & Chalk (Anat. Die. 2, 1950, 821) confirmed the close relationship between
the Lobeliaceae and Campanulaceae which they find reflected in many similarities between the two
families. Within the Campanulaceae they admitted that Sphenoclea and Pentapliragma, which are both
not laticiferous, show some deviating anatomical characters. They expect that laticiferous canals probably
occur in all genera of the Campanulaceae. The question arises why the /w/i-laticiferous Goodeniaceae
are kept separate from this complex; by the development of the indusium below the style they obviously
represent a specialized branch of the same stock.
Chromosomes. The number of the chromosomes of the Campanulaceae and Lobeliaceae, as found
in Darlington & Wylie's 'Chromosome Atlas' (1955) provide a rather confusing picture, in contrast
with for example the allied Goodeniaceae which show in 7 publications on 4 genera constantly n = 8.
In Campanulaceae the basic numbers vary considerably, sometimes even in taxonomically coherent
genera, e.g. Phyteuma in which is found n = (6), 12, 13, 14, and in Jasione n = 6 and 7. In Campanula
the situation is even n = 8, 10, 12, 13, 14, and 17. Also within the species the numbers may vary, although
not as aneuploids, but as polyploids, with some irregularities. Polyploidy seems to be very common
indeed.
Phytochemistry. This family has not been studied intensively by phytochemists. The facts known
point on the one hand to many metabolic similarities between the two great subfamilies, Campanuloideae
and Lobelioideae. and on the other hand to marked differences. Campanulaceae sensu lato are characterized
by local accumulation of silicic acid and/or carbonate of lime in epidermal cells of the leaves, especially
in the hairs and neighbouring cells. Furthermore all Campanulaceae accumulate inulin and/or inulin-
like oligo- and polyfructosans in their vegetative organs. The tendency to accumulate fructosans is
even more accentuated in this family than in Compositae. In Campanulaceae fructosans are synthesized
not only by perennial species but also by the annual ones as Collin and Cholllet have demonstrated.
In the seeds fatty oils are stored. Flavonols and leucoanthocyanins are usually absent from the leaves
of Campanulaceae but ferulic and/or sinapis acid are of common occurrence. All Campanulaceae contain
articulated, ramified latex vessels in the phloem. The constituents of latices seem to be different in the
two subfamilies:
Campanuloideae: The chemistry of latices of this taxon is practically unknown. Phytosterols and a
phenolic glycoside, called campanulin, have been demonstrated to occur in the latex. Alkaloids or
other toxic principles are not known to be present.
A saponin, platycodin, was isolated from the roots of Platycodon grandiflorum DC. but saponins
are by no means widely distributed in the subfamily. Caflfeic acid on the other hand seems to be a phenolic
constituent frequently present in the leaves.
Lobelioideae: Many species of this subfamily are known to be toxic. The toxic principles are al-
kaloids accumulated in the latex. Many different alkaloids were demonstrated to be present in Lobelia
and Isotoma species. The medicinally used North American Lobelia inflata L. has been thoroughly
investigated for alkaloids. This plant produces about 30 alkaloids which seem all to be related structurally
to lobeline, an a, a^-disubstituted-N-methylpiperidine base. The alkaloids of all other species o^ Lobelioi-
deae are very imperfectly known. There is, however, no doubt that other types of alkaloids occur in
the subfamily. Lobelioideae are also characterized by the occurrence of chelidonic acid in the majority
of species. On the other hand caffeic acid seems to be absent from this subfamily. Ellagic acid was
found inCentropogon lucyanus Schonl. which also contains a little leucocanthocyanin in the leaves.
Some species of Lobelioideae are reported to accumulate a fair amount of rubber in their latices.
Finally it may be stated that by the accumulation of inulin-like fructosans the family is related to
Goodeniaceae, Stylidiaceae, and Compositae, and by the articulated latex bearing vessels to the Cichorieae
subfamily of Compositae. These indices of relationship are, however, rather weak. A thorough chemo-
taxonomical discussion of Campanulaceae s.l. must be postponed until the chemistry of the family has
been more thoroughly explored. — R. Hegnauer.
Taxonomy. Airy Shaw has recently suggested to 'purify' the Campanulaceae by ramngihs geneva
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 109

Pentaphragma and Sphenoclea to the status of family, suggesting that the latter genus would only be
remotely allied to Campanulaceae but more to Phytolaccaceae. Though it should be admitted that both
genera deserve a special status in the family agree with Wimmer and Hutchinson (Fam. Fl. PI. ed.
1

2, 1, 1959, 477), who keep them together in the Campanulaceae.

There seems little in favour of separating Campanulaceae and Lobeliaceae as they remain side by side.
And in the latter family there are many genera and species which can hardly be termed zygomorphous;
also there is degree in the connation of the stamens.
The Malaysian genera can be arranged as follows:
1. Tribus Campanuleae Bth.: Peiacarpa, Wahlenbergia, Codonopsis.
2. Tribus Sphenocleae Schonl. Sphenoclea.
:

3. Tribus Pentaphragmeae Schonl. Pentaphragma.

:

4. Tribus Lobelieae Bth.: Lobelia, Phyllocharis, LaureiUia.

The affinity of the Campanulaceae seems distinctly fixed with the Stylidiaceae, Goodeniaceae, and
Compositae, on morphological, phytochemical, cytogenetical, and anatomical grounds.
The classification of the major groups within the family seems also rather well established.
Generic distinction has been liable, however, to difference of opinion and in this revision has amply
been discussed under Codonopsis, Wahlenbergia, Laurentia, and Lobelia. A century ago, when few
species were known, genera could often be more or less easily defined, but as more species were found
these accepted generic distinctions have often broken down in a number of species ('links"), necessitating
a revision of generic limits. This revision of the hierarchy within the family can logically not be per-
formed without either merging genera or segregating others to bring the mutual relations into balance,
i.e. bringing taxa of the same rank on a ^
common footing. In either case we have to face name changes.
The latter procedure (segregation) has several manifest disadvantages against the former (large genera,
subdivided into sections and subsections), viz that the number of generic names becomes confusingly
large, and what is more regrettable, generic names become not any longer bound to some clear structural
plan, but to small details, which seem rather technical than structural. Segregation will also necessitate
the breaking up of genera which nobody wants to break up, because if we split up Wahlenbergia sens,
kit. into 10-20 genera, it would be absurd to leave Campanula intact, as this would greatly upset the
standard of the generic level within the family. A further disadvantage will be the increase of monotypic
genera, as for those in favour of a larger number of genera, the only means to 'remove' links is to bring
them to the generic level in their own right. But most undesirable consequences hover over the splitting
procedure, because this is not the end: many novelties wait for exploration and description, and it is
almost certain that among them there will be still more 'links' threatening the limits of the so-called
small, uniform genera. Finally I find a great disadvantage with the small genera because in a family
where the characters are so 'variable', detailed technical characters are no guarantee for common an-
cestry and are likely to be at least in part artificial. This will namely always be the case where species
are reticulately allied. In reticulate affinities, which are obviously mostly a consequence of proportionally
recent speciation\ one will always find 'links' in attempts to subdivide such an affinity. The cytogenetical
data emphasize that such a situation is found in the family.
The merging of genera is often objected with the practical argument that genera become too large
and so-called 'unwieldy'. We have, however, to accept that there are large genera on which all agree
that they should not be segregated, for example Carex, Ficus, Acacia, Rhododendron, Fimbristylis,
Astragalus, Bulbuphyllum, Dendrobium, Malaxis, Calamus, Eugenia, Eucalyptus, etc.
The classical, proper way to deal with these genera is to subdivide them into a hierarchy of in-
frageneric levels as good as we can. Segregation in such genera will never be useful, the more we learn
about them.
Most recent specialists on the Campanulaceae agree with this view (Wimmer, McVaugh, Nann-
FELDT, and others), but some are reluctant because of name changes; as said before this cannot be
remedied by segregation which will result into even more name changes. I fully agree with Dr Meikle,
—
who wrote me: "I am not happy about generic distinctions in Campanulaceae, and wonder if a resurvey
of the family could not now be considered". As far as was compatible with the scope of the present
revision we have found fit to make a small contribution towards improving generic delimitation; the
necessity of name changes should, in our opinion, never be an impediment towards a real improvement
of a taxonomic system.
In this revision, for which my two keen collaborators have examined much more extra-Malaysian
material and have studied more literature than strictly necessary, it has been shown impossible to
keep Campanumoea separate from Codonopsis, further to keep Cephalostigma and Lightfootia apart
from Wahlenbergia which itself is very close to Campanula. A more serious deviation from current usage
is the merging of Pratia with Lobelia, as the first was, by its berried fruit, mostly arranged widely apart

from the capsular genera of Lobelioideae. The same discrepancy is, however, found in Codonopsis where
berried and capsular fruits are found in one distinctly natural genus. It would appear that a subdivision
of Lobelieae based only on the structure of the fruit (capsular or berried) is artificial.

^ The remarkable development of a large number of species of Lobelia cq. genera with arboreous

habit on the high African volcanoes and the Hawaiian Islands point in the same direction.
no Flora Malesiana [ser. I, vol. 6^

Specific variability. As is usual some species have proved more variable than others and after due
examination of much extra- Malaysian material some have been accepted in a wider sense than mostly
employed, notably Wahlenbergia marginata, Lobelia angiilata, L. nicotianaefolia, Cudonopsis javanica,
and C. lancifolia, all of which show a wide area of distribution. Variable species are not unknown in
the family, as even Stearn, who is certainly not particularly in favour of extreme lumping, felt baffled
with the astonishing variability of Campanula rotiindifolia. Part of this variation is according to the
cytogenetical data due to the occurrence of chromosome races, part of it can be ascribed to raciation
whether or not effected by isolation on islands. In most cases we have refrained from giving names to
these races, as a proper naming should only be achieved after much more detailed field study combined
with cytological and experimental-taxonomical research.
Also random paramorphs are common; as has already been remarked by Schonland (in E. & P.
Pfl. Fam. 4, 5, 1894, 43) there is a great variability in the numbers of the floral parts; see also under
Wahlenbergia marginata.
Uses. Heyne I.e. has enumerated the few, minor uses recorded in Malaysia. Some exotic species are
cultivated as ornamentals; they have been enumerated at the end of the family and under Lobelia a
key has been given to five cultivated species of that genus. It must be emphasized, though, that most
Campamilaceae can in the tropics only successfully be grown in the montane zone.
Notes. Mr Moeliono is responsible for revising Lobelia, Codonopsis, and Peracarpa, Mr Tuyn for

Laiirentia, Phyllocharis, and Wahlenbergia. The drawing of the family description, the introductory
notes, and key to the genera has been my privilege. Van Steenis.

KEY TO THE GENERA

1. Anthers during anthesis free. Flowers regular.
2. Flowers in dense spikes.
3. Leaves very large and wide, asymmetrical, alternate. Spikes one-sided (scorpioid), axillary. Petals
or corolla segments valvate. Fruit an indehiscent berry. See vol. 4, p. 517. 5. Pentaphragma
3. Leaves symmetrical, lanceolate, spirally arranged. Spikes regular, not scorpioid, terminal. Corolla
lobes imbricate. Capsule circumsciss-dehiscent, opening with a lid. See vol. 4, p. 27.
4. Sphenoclea
2. Flowers solitary or in cymes, racemes or panicles.
4. Leaves at least partly (lower ones) decussate. Base of the corolla tube adnate to the ovary. Fruit
(in Mai. spp.) a berry 3. Codonopsis
4. Leaves spirally arranged or distichous. Corolla not adnate to the ovary. Fruit capsular.
5. Erect or slightly ascending herbs. Flowers blue or white, terminal. Capsule loculicid, with 2-3

apical valves 2. Wahlenbergia

5. Prostrate, creeping, delicate herb, rooting at the nodes. Flowers solitary, axillary, white. Capsule
indehiscent 1. Peracarpa

1. Anthers and mostly part of the filaments connate during anthesis. Corolla dorsally often slit to near

the base, often zygomorphic.

6. Corolla white, tube 7V2-11 cm long, narrow-cylindric, not dorsally slit, almost actinomorphic.
Stamens inserted high in the tube (in Mai. sp.) 8. Laurentia
6. Corolla much smaller, at most c. 3 cm long, generally distinctly zygomorphic, tube dorsally mostly

deeply slit. Filaments free at the base or inserted on the base of the corolla tube.

7. Flowers solitary, through connation with the leaf epiphyllous .... 7. Phyllocharis
7. Flowers arranged in various ways, not epiphyllous 6. Lobelia

1. PERACARPA
Hook. f. & Th. J. Linn. Soc. Bot. 2 (1858) 26; Peer, Bot. Jahrb. 12(1890)620,
t. VII B.
Prostrate, creeping herb; stem terete, rooting at the nodes. Leaves spiral. Flowers
5-merous, actinomorphous, solitary, axillary, sometimes terminal. Sepals free, erect.
Corolla campanulate, 5-lobed. Disk fleshy, semiglobose, 3-sulcate. Stamens 5, free
from the corolla. Ovary 3-locular; ovules cv); style simple; stigma 3(-4)fid. Capsule
pyriform, pendulous, with a thin pellucid wall, lengthwise nerved, not dehiscent,
pericarp swollen above the oblong seeds.
Distr. Monotypic, from the Himalaya to Kweichow, Japan, and Formosa, in Malaysia: Philippines
(N. Luzon), only in the mountains.

1. Peracarpa carnosa (Wall, in Roxb.) Hook. /. Jahrb. 12 (1890) 620, t. 7B; Craib, F1. Siam. En.
& Th. J. Proc. Linn. Soc. Bot. 2 (1858) 26; 2 (1936) 308; Yamamoto, Obs. Fl. Form. 13
Clarke, Fl. Br. Ind. 3 (1881) 437; Feer, Bot. (1936) 149; Hara. J. Jap. Bot. 21 (1947) 20;
Dec. 1960] CampanulaCeae (Moeliono & Tuyn) 111

Making, 111. Fl. Japan (1954) 82, incl. var. cir- campanulate, varying from blue to white; lobes
caeoides.— Campanula caniosa Wall, Roxb. in Fl. 5, for up Vi connate, nearly equal, elongate
to
Ind. 2 (1824) 102; Cat. (1828)/;. 1282; DC. Prod. to elliptic 2-y4-10 by 1-2 mm, entire, acute or
7(1839)474. — Campaiiala cinaeuides Fr. Schmidt obtuse, glabrous. Stamens: anthers c. 1 mm,
ex MiQ. Ann. Mus. Lugd. Bat. 3 (1867) 195, glabrous; filaments linear, broadened to the base,
204; FoRB. & Hemsl. J. Linn. Soc. Bot. 26 (1889) 2-4 mm, finely haired, sometimes glabrous. Ovary
9. — P. Feer, Bot. Jahrb. 12 (1890)
circaeoides obconical to campanulate, '/2-3 by 1-2 mm,
1

621; Fedorov, U.R.S.S. 24 (1957) 380.—/*.

Fl. glabrous; style simple, glabrous, rarely hairy,
luzonica RoLFE, Kew Bull. (1906) 201; Merr. with narrow stigmas. Fruit ovoid to obovoid,
& Merritt, Philip. J. Sc. 5 (1910) Bot. 392; Merr. 3-5 by 3-5 mm, with a very thin wall, pendent
En. Philip. 3 (1923) 586. on the pedicels. Seeds fusiform-ellipsoid, 1-1 '/2
A weak, branching succulent herb up to 16 cm mm, brown and smooth,
long. Leaves broad ovate to ovate, 4-30 by 4-20 Distr. SE. Asia: N. India (Himalaya), Siam,
mm, cordate or truncate, entire or crenate, acute S. China (Yunnan, Kweichow), Japan, and
or obtuse, both sides puberulous, the underside Formosa; in Malaysia: Philippines: Luzon
sometimes glabrous; petiole as long as to a little (Benguet and Lepanto: Mt Osdung; Mt Pulog)
shorter than the leaves, 4-25 mm, glabrous. and Panay.
Flowers 5-12 mm. Pedicels 6-30 mm. Sepals 5, Ecol. In moist mountain (oak) forest, at the
free, subulate, sometimes triangular, '/2 K — ^Vi) base of trees. 1450-3000 m. Fl. fr. Febr.-June.
by 1/2mm, entire, glabrous. Corolla 3-12 mm,

2. WAHLENBERGIA
SCHRAD. [PI. Sem. Hort. Ac. Gott. 1814, p. 3, nom. nud.] ex Roth, Nov. PI. Sp.
(1821) 399, nom. gen. cons., cf. Steen. Taxon 9 (1960) 125; ?D. Don, Prod. Fl. Nep.
(1825) 156; Schrad. Blumenbachia, etc. (1827) 123; Rchb. Ic. PL Rar. cent. 5«
(1827) 47, t. 480; Schrad. Comm. Gott. 6 (1828) 123; DC. Mon. Camp. (1830)
1 29 ;Brehmer, Bot. Jahrb. 53 (1915) 9; Lothian, Proc. Linn. Soc. N.S.W. 71 (1947)
20\. —Lightfootia 1'Herit. Sert. Angl. (1789) 4, t. 4, non Sw. 1788 (Flac), nee
Schreb. 1789 (Rub.); R. S. Adamson, J. S. Afr. Bot. 21 (1955) \55.— Campanula
sect. Campanopsis R. Br. Prod. (1810) 560. Cervicina Delile, Fl. d'Egypte (1813)
7, Atlas t. 5, f. 2, nom. gen. rejic. —
Cephalostigma DC. Mon. Camp. (1830) 117.
Campanulopsis Zoll. & MoR. Nat. & Geneesk. Arch. Neerl. Ind. 1 (1844)484,
nomen; cf. Steen. Bull. Jard. Bot. Btzg III, 17 (1948) 463.— Campanopsis O.K.
Rev. Gen. PI. 2 (1891) 378, nom. illeg.
Annual or perennial, erect or ascending herbs, sometimes woody at the base.
Stems simple or branched, glabrous or hairy. Leaves (in Mai.) mostly sessile, spirally
arranged, hairy or glabrous, elliptic to linear, seldom obovate or spathulate, margins
mostly thickened, shallowly dentate or almost entire. Inflorescence (or flowers)
terminal or axillary, usually sparsely flowered. Bracts narrow, small. Pedicels
glabrous or hairy. Flowers distinctly protrandrous, regular, mostly blue or white.
Calyx inferior to (in Mai.) superior, lobes 3-6 (in Mai. 3-5), acute, blunt or slightly
acuminate, triangular to linear, persistent. Corolla 5- or 4-partite, or a 3-6- (in
Mai. 3-5-) lobed tube, sometimes inside bearing slender hairs. Stamens 5, alternate
with corolla lobes, free; anthers narrow; filaments membranous, broad or broadened
at the base, ciliate, the apical part often recurved. Style about as long as corolla
tube, or longer, lengthening during anthesis, the basal part sometimes bearing long,
slender hairs, the upper portion sometimes with short bristle-like hairs, sometimes
bearing warts at or near the base of the stigmatic lobes; stigmas 2-3 (in extra-Mai.
ssp. up to 5), spreading late in anthesis. Ovary inferior (in Mai.) to superior, 2-3-
celled (in extra-Mai. spp. up to 5-celled). Capsule loculicid, opening by as many
apical valves as there are cells in the ovary. Seeds cv), elhpsoid or triangular, shining,
up to 1 mm.
112 Flora Malesiana [ser. I, vol, 6^

Distr. Large genus, possibly comprising more than 150 spp., almost cosmopolitan, with the bulk
of the species on the southern hemisphere, specially in Africa. In continental Asia and Malaysia only
a few species. Roughly it lorms a geographical and taxonomical complement to Campanula, which is
mainly a northern hemisphere genus.
Ecol. Extremely variable, preferring open localities in steppe, savannahs, and at high altitudes,
the four Malaysian species only on the mountains.
Flower biology. The flowers are distinctly protrandrous. The sticky pollen is shed already in bud,
and covers in great masses the upper portion of the style and the stigmatic lobes, conceahng their shape
and structure. The anthers are withering in early anthesis and are often not found in fully opened
flowers. The margins and/or lobes of the filaments are incurved. Their filiform apex is mostly recurved
and closely appressed to the widened central portion of the filaments. The stretching of older filaments,
which are so frequent in the herbarium, is a very delicate work. All other floral parts are growing during
flowering, which makes it very difiicult to extract exact measurements from herbarium material. The
stigmatic lobes are spreading in a later stage. The hairs that are sometimes found on the inside of the
corolla and on the style seem to fall off later.
Tax on. After ample consideration we have in this revision combined the genera Lightfootia and
Cephalostigma with Wahlenbergia and feel compelled to give our reasons for this conclusion. A century
ago, when few species were known, it was rather easy to separate and define these three genera, though
all of them are only separated from Campanula by the apical dehiscence of the fruit. Lightfootia would
be characterized against Wahlenbergia by the deeper incised corolla, the narrower corolla lobes, and
the style, which would be proportionally longer if compared with the corolla. Cephalostigma would
be closest to Lightfootia in having a narrow-lobed, very deeply incised corolla and a simple knob-like
stigma. The latter observation of De Candolle was erroneous: there are three short lobes which form
in outline a more or less globular whole.
In proportion to the tremendous increase in described species the already not particularly strong
differences have gradually become vague and Von Brehmer (1915, I.e.) in a thorough study of the
African species of Wahlenbergia came to the conclusion that a sharp separation of the genera Wahlen-
bergia and Lightfootia cannot be upheld because in both genera so many different combinations are
represented (by different species), that the differential characters imperceptibly merge into one another.
His scheme of^ style and stigma structure in the various groups of species (I.e. p. 12) showed that no
demarcation can be drawn. He maintained both names because of his 'personal aversion' to use the
nomenclaturaliy older name Lightfootia I'Herit. 1789 for the Wahlenbergias which he assumed to be
older in the phylogeny, excusing this curious conclusion by stating that in the Campanulaeeae genera
are often distinguished by "sekundare" characters. Dr R. S. Adamson, I.e., could not find more con-
vincing arguments and merely referred to Von Brehmer. In a recent letter he kindly elucidated his stand-
point, admitting that there is not any key character that would separate the two genera; for the South
African species the corolla and style characters work fairly well, but "these features break down for some
of the Lightfootia species in tropical Africa which I feel do not really belong to the genus". He con-
tinues: 'T do not feel in favour of placing the whole lot in one comprehensive genus. This would possibly
be the most logical but would be very unwieldy, would entail a lot of name changes, and would tend to
obscure some geographical lines of development. My own feeling is rather to increasing the number
of genera. The two in question could with advantage both be split up. The smaller units would admit
of key diagnosis. I am not sufficiently familiar with the extra-S. African species to attempt even an
outline".
Mr Cannon of the British Museum (Nat. Hist.), Botany Department, though making the provision
that he has no specialized knowledge of the group, kindly commented to us that in his opinion "it
does not seem that the grounds for keeping the genera separate are anything more than mere con-
venience. Presumably merging the genera would bring about a large number of new combinations,
but this can scarcely be regarded as a valid reason for not adopting what appears to be the scientificafly
correct course of action".
Finally, Dr R. D. Meikle, Kew, granted us the privilege of his opinion: "Quite frankly, I think that
all the supposed distinctions are untenable, and I would certainly unite the two genera. I suspect the
nomenclatural consequences have alone dissuaded others from doing so. I can see no good reason
for conservation of either, and even if the change calls for a lot of new combinations, that simply cannot
be helped".
To a serious matter like this, where we deal in this Flora with only few species of a large, worldwide
distributed aggregate, we think ample consideration should be given to the opinion of specialists.
What should be strived at, specially in generic distinction, is a balanced system of recognizable units
at about one level throughout the family which are structurally more or less comparable, though these
units may differ considerably in the number of species. We can expect that with increase in the number
of species described through intensified exploration, generic limits which were once clear may entirely
disappear. From the scientific standpoint we should not leave such cases unattended, but should try to
improve our insight in the family by applying a sound generic distinction.
In this case there is unanimity about the evaluation of the facts, but there is difference of opinion
about the course we should take in future. Scientifically we cannot share the aversion of Von Brehmer,
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 113

neither to required name ciianges nor to his argument of phylogeny, as for the latter the nomenclature
is entirely irrelevant.
Dr Adamson's perspective we feel we cannot share either, because we see no advantage
in segregation
in this case. Firstly it is not certain that it would really be possible, but if we
discard this doubt for a
moment, secondly it would lead to a large number of small genera ranking taxonomically below the
generic level proper and disturb the hierarchy within the family. Naturally, subgenera are generally
more "natural" than genera and sections more natural (coherent) than subgenera, and so forth descend-
ing in the ranks. But the advantage to have a large genus Wahlenbergia is that of generic balance in the
family whilst Dr Adamson's desire for coherent units can well be fulfilled by distinguishing within the
genus th2 natural subgeneric taxa in various ranks. A similar procedure has been followed within other
natural, but very large genera, as in Astragalus, Biilbopliyllum, Carex, Dendrobiiim, Ficiis, Eucalyptus,
Acacia, etc. I assume this infrageneric distinction will be similarly satisfactory in Wahlenbergia sens. lat.
and will meet the needs for distinction of natural units felt by Dr Adamson.
As to the third genus, Cephalostigma, De Candolle erroneously stated (1830, I.e. 56) that it would
deviate from Lightfootia by a "stigma en tete" (capitatum, simplex. I.e. 117); he assumed it to be inter-
mediate between Wahlenbergia and Lightfootia. Really, the stigma has afterwards always been described
as consisting of 3 lobes, as in the other genera. Consequently Cephalostigma cannot be upheld, as its
stigmatic form is also found in Wahlenbergia.
Nomencl. Though Lightfootia I'Herit. is the oldest name, it is a later homonym oi Lightfootia Sw.
(see for publication dates Fl. Mai. I, 4, 1954, cxcvii, ccxiv). Fortunately more species have been described
under Wahlenbergia than under Lightfootia, so that the smallest possible amount of transfers will be
necessary.

KEY TO THE SPECIES

1. Petals narrow, almost free. Capsules with as many lengthwise nerves as there are calyx lobes.
2. Ovary hispid. Seeds ellipsoid, compressed. Filaments 3-lobed 1. W. erecta
2. Ovary glabrous. Seeds trigonous. Filaments narrow triangular 2. W. hookeri
1. Corolla with a distinct tube, much longer than the calyx lobes. Capsules with twice as many nerves

as calyx lobes.
3. Filaments very narrow triangular. Fruit wider than long, 3-celled. Calyx lobes longer than the ovary.
Corolla tube about twice as long as the calyx lobes (sometimes longer). Corolla (stretched) 12-18 mm.
3. W. confusa
3. Filaments abruptly narrowed about halfway, the basal part about pentagonal, deltoid, or pseudo-
2-lobed. Fruit longer than wide, 2-3-celled. Calyx lobes shorter or longer than the ovary. Corolla
tube shorter or longer than the calyx lobes (never more than twice as long). Corolla (stretched)
214-12 mm (in var. grandiflora 15-25 mm) 4. W. marginata

1. Wahlenbergia erecta (Roth ex R. & S.) Tuyn, the calyx lobes. Stamens 5, anthers 0.5 by 0.2
nov.comb. —Dentella erecta Roth ex R. & S. mm; filaments 0.8 mm, membranous, broad at
Syst. Veg. 5 (1819) 25, ex descr.; Nov. PI. Sp. the base, 3-lobed, the side lobes small, the middle
(1821) 140; Cham. & Schlechtend. Linnaea 4 lobe bearing the anther. Ovary l-lVi mm, bell-
(1829) 151; DC. Prod. 4 (1830) 4\9.— Dentella shaped to obconical, patently hispid-hairy (hairs
perotifolia Willd. ex R. &Veg. 5(1819)
S. Syst. up to Vi rnm), top conical, apparently lengthening
25, descr. in syn., nom. illeg.; DC. Prod.
4 (1830) during anthesis to about 1 Vi mm; style 1 Vi mm,
419.^^F. perotifolia W. & A. Prod. (1834) 405, narrow-cylindric, thickened below the apex, with
nom. illeg.; DC. Prod. 7 (1839) 434; Wight, 3 short stigmatic lobes. Fruit 2-3 mm, obconic
Ic. PI. Ind. 3 (1844-45) 4, t. 842. Cephalostigma to bell-shaped, 5-nerved, valves 3, before opening
schimperi Hochst. ex Rich. Tent. Fl. Abyss. 2 forming a glabrous cone 1-1.7 mm
high. 5fe^.y ^J,
(1851) 2; Clarke, Fl. Br. Ind. 3 (1881) 428; compressed-ellipsoid, shining, brown, Vi mm.
Trim. Fl. Ceyl. 3 (1895) 58. Cephalostigma Distr. East Africa, ?Ceylon, India (the Deccan
erectum (Roth) Vatke, Linnaea 38 (1874) 699, and Khasya), in Malaysia: North Sumatra (Karo
quoad specim. pro parte. Fig. Ih-i. — and Toba-Batak Flighlands). According to
Annual, erect herb. Stems 5-10 (-35) cm, Trimen, I.e., probably not in Ceylon,
flexuose, branched, blunt-angular, the angles pale Ecol. Open, heath-like land and waste places,
elevated lines, patently ± hispid-hairy. Leaves 750-1400 m.
5-20 by 1 V2-6 mm, spirally arranged (sometimes Vern. Dukut tawar, Karo-Batak.
almost distichous), sessile, oblong, sometimes Note. This species has sometimes been con-
elliptic, acute, mostly obtuse at the base, sparsely fused with W. hirsuta (Edgew.) Tuyn, nov.comb.^
hairy (hairs especially along the midrib) or almost (Cephalostigma hirsuta Edgew. Trans. Linn. Soc.
glabrous, margin thickened, undulate shallowly 20, 1846, 81); the latter has also a hispid ovary
and remotely dentate. Flowers axillary and ter- but trigonous seeds,
minal. Pedicels 5-10 mm, filiform, hairy. Calyx
lobes 5, 1 '/2-3 mm, linear to narrow-triangular, ^ The name Wahlenbergia hirsuta Steud.
acute, outside sparsely patently hairy. Corolla Nomencl. ed. 2, 2, p. 782, is a nomen nudum and
5-parted, segments linear acute about as long as has no nomenclatural status.
114 Flora Malesiana [ser. I, vol. 6^

2.Wahlenbergia hookeri (Clarke) Tuyn, nov.comb.

— Cephalostigma hookeri Clarke, F1. Br. Ind. 3
(1881) 429; Danguy, F1. Gen. I.-C. 3 (1930) 690;
Craib, F1. Siam. En. 2 (1936) IQl.— Cepha-
lostigma paniculatum {uon DC.) HossEUS, Beih.
Bot. Centr. Bl. 28, ii (1911) 446.— Fig. la-g.
Slender annual herb, erect. Stems 10-20 cm,
hairy. Leaves 10-30 by 4-9 mm, (sub) petiolate,
oblong, elliptic, obtuse or acute, hairy, margin
thickened, crenate, shallowly dentate. Inflorescence
slenderly branched, glabrous or nearly so, bracts
4 by 1 mm, or (mostly) smaller, pedicels 4-30 mm,
filiform. Calyx lobes 5, 1-1.2 mm, elongate-
triangular, acute, sometimes with a few hairs at
the apex, further glabrous. Corolla 5-parted, 2
by 0.4 mm, about twice the length of the calyx
lobes. Stamens 5 anthers 0.4 by 0. 1
; mm
filaments ;

1 mm, narrow triangular, sometimes almost

filiform. Ovary 1 — 1 Vi mm,
bell-shaped to ob-
conic, glabrous; top conic, up to 1 high; mm
style 1.3-1.6 mm, narrow-cylindric, 3 short
narrow stigmatic lobes. Fruit (only one seen, in
poor condition) 2 by 2 mm, semispherical, 2-
valved. Seeds numerous, triangular, brown,
shining, 0.6 mm.
Distr. India (Behar, Khasya), N. Siam (Doi
Sutep), in Malaysia: East Java (Mt Idjen: L. van
DER PiJL 144, BO, L, K), only once collected,
June 1929.
Ecol. In reed stands i^.Themeda), probably c.
1000—1200 m alt.
Note. Diff"ers from Wahlenbergia candollei
Tuyn, nom.nov. [Cephalostigma paniculatum DC.
Mon. Camp. (1830) 117, non Wahlenbergia pani-
culata (Thunb.) DC. I.e. p. 153] in its smaller
flowers and the bell-shaped, not obconic ovary.
The shape of the seed in the Indian and the
Javanese material is elliptic, in the only specimen
from Siam (Hosseus 225) the seeds are more
obovate.

3. Wahlenbergia confusa Merr. & Perry, J. Arn.

Arb. 22 (1941) 383.— Fig. 2.
Probably perennial, usually glabrous, sometimes
stem and leaves (midrib, margin) with sparse
bristle-like hairs. Stems ascending, 10-25 cm long,
sulcate. Leaves ~
uniform in shape, thickish,
2-1 1 by '/2-3 mm. Flowers solitary. Pedicels 2-7
cm. Calyx lobes 5, i
spreading, linear-lanceolate,
acute to acuminate, usually entire, always longer
than the ovary, 2'/2-4 mm
long. Corolla tube
about twice as long as the calyx lobes, sometimes
longer, 6'/2-8 mm; lobes 5, ovate, acute, 5-10 mm.
Filaments linear-triangular, IVi-'^Vi mm. Ovary
obconical, 3-celIed, 1.2-3 mm; style slightly

Fig. 1. Wahlenbergia hookeri (Clarke) Tuyn.

a. Habit, X %, b. flower, stamens removed, x 7,
c. fruit, X 7, d. seed, X 13, e. flower, stamens
removed, x 7, /. fruit, X 7, g. seed, x 13.
Wahlenbergia erecta (Roth ex R. & S.) Tuyn.
h-i. Flowers, stamens removed, x 7, /. fruit,
X 7, k. seed, x 13 {a-d van der Pul 144 (E.
Java), e-g. HossEUS 224 (Siam), h-k Lorzing
7190 (N.Sumatra).
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 115

32; Alston, Fl. Ceyl. Suppl. 6 (1931) 176;

Docters van Leeuwen, Verb. Kon. Ak. Wet.
A'dam sect. 2, 31 (1933) 267; Hochr. Candollea
5 (1934) 290, cum var.; Steen. Bull. Jard. Bot.
Btzg III, 13 (1934) 179; Craib, Fl. Siam. En. 2
(1936) 307; Lothian, Proc. Linn. Soc. N.S.W.
71 (1947) 212, cum var.; Back. Bekn. Fl. Java
(em. ed.) 8 (1949) fam. 183, p. 5; Hatch, Trans.
Proc. R. Soc. New Zeal. 79 (1952) 368, cum var.;
YuNCKER, Bull. Bern. P. Bish. Mus. 220 (1959)
261. Campanula marginata Thunb. Fl. Jap.
(1784) 89. Campanula gracilis Forst. Prod.
(1786) 84; R.Br. Prod. (1810) 561, incl. var.—
Campanula quadrifida R.Br. Prod. (1810) 561.
Campanula dehiscens RoxB. [Hort. Beng. (1814)
85] e.\ Wall. As. Res. 12 (1816) 571, cum ic;
Fl. Ind. 2 (1824) 96. Campanula agrestis Wall.
in Roxb. Fl. Ind. 2 (1824) 97; Cat. (1829) 1292.—
Campanula lavandulaefolia Reinw. ex Bl. Bijdr.
(1825) 116.— W. gracilis [Schrad. Blumen-
bachia (1827) 38, /// obs.] DC. Mon. Camp.
(1830) 142, incl. var.; Prod. 7 (1839) 433; Jungh.
Nat. Geneesk. Arch. Need. Ind. 2 (1845) 48, cum
var. hirsuta Jungh.; Hook./. Fl. Tasm. (1860) 1

239; Bth. Fl. Austr. 4 (1869) 137; Clarke, Fl.

Br. Ind. 3 (1882) 429; Ware. Bot. Jahrb. 18
(1893) 212; Hemsl. J. Linn. Soc. 30 (1894) 183;
Bailey, Queensl. Fl. 3 (1900) 922; Cheeseman,
New Zeal. Fl. (1906) 402; den Berger, Trop.
Natuur 6 (1917) 104 f. 4; Stearn, Diet. Card.
R.H.S. 4 (1951) 1151.— W. quadrifida (R. Br.)
DC. Mon. Camp. (1830) 144; Prod. 7 (1839) 433;
N. E. Brown, Gard. Chron. 54 (1913) 316;
Domin, Bibl. Bot. Heft 89 (1929) 1192; Lothian,
Proc. Linn. Soc. N.S.W. 71 (1947) 1\Q.— W.
dehiscens (Roxb.) DC. Mon. Camp. (1830) 145
Prod. 7 (1839) 434; Lothian, Proc. Linn. Soc
Fig. 2. yVahleubergia confiisa Merr. & Perry.
N.S.W. 71 (1947) 2\6.— W. agrestis DC. Mon
a. Habit, x 2/,, ^. longitudinal section of flower,
Camp. (1830) 145; Prod. 7 (1839) 434; Wight
X 2, c. "filament, 6 (Brass 9399).
Ic. PI. Ind. Or. 4 (1850) 1175; Hook. /. & Th
Proc. Linn. Soc. Lond. 2 (1858) 21; Lothian
shorter than the corolla tube, 5'^-7 mm; stigma Proc. Linn. Soc. N.S.W. 71 (1947) l\5.— iV.
3-lobed, (l-)l'/2-3 mm long. Fruit slightly wider lavandulaefolia DC. Mon. Camp. (1830) 144
than long, 3-valved, 2-3 mm diam.. crowned by Prod. 7 (1839) 433; Zoll. Nat. Geneesk. Arch
the recurved calyx lobes. Neerl. Ind. 2 (1845) 561.— W. indica DC. Mon
Distr. Malaysia: West New Guinea (vicinity Camp. (1830) 146; Prod. 7 (1839) 434; Wight
of Mt Wilhelmina). Ic. PI. Ind. Or. 4 (1850) 1176; Lothian, Proc

Ecol. Wet, grassy, open places along streams, Linn. Soc. N.S.W. 71 (1947) 209.— H^. sieheri
once noted on sandstone, 3400-4000 m. DC. Mon. Camp. (1830) 144; Prod. 7 (1939)
Note. Merrill compared his new species with 433; Lothian, Proc. Linn. Soc. N.S.W. 71 (1947)
W. eurycarpa Domin (Bibl. Hot. Heft 89, 1929, 219. —W. multicaulis Bth. in Hiigel. En. PI.
638) but already remarked that the fruit in the latter Nov. Holl. (1837) 75; DC. Prod. 7 (1839) 433;
would appear too long. Domin himself compared N. E. Brown, Gard. Chron. 54 (1913) 337;
his species with W. sieheri and gave the differences Lothian, Proc. Linn. Soc. N.S.W. 71 (1947) 229;
with W. multicaiilis which species I have reduced Stearn, Diet. Gard. R.H.S. 4 (1951) 2258.— C<:/m/7fl-
to W. nuirgiiiata. As Domin neither mentions the nula sieheri Dietr. Syn. PI. 1 (1839) 152.— Campa-
shape of the filaments nor the relative length of nula indica Dietr. I.e. 753. Campanula littoralis
the corolla tube, and as his material is not avail- Labile. PI. Nov. Holl. 1 (1844) 49, t. lO.—Cam-
able to me, Irefrain from giving a definite panulopsis cyanea Zoll. &
Mor. Nat. Geneesk.
opinion. Arch. Neerl. Ind. 1 (1844) 484, nomen.— W.
simplicicaulis de Vriese in Lehm. PI. Preiss. 2
4. Wahlenbergia marginata (Thunb.) DC. Mon. (1846) 241 Lothian, Proc. Linn. Soc. N.S.W. 71
;

Camp. (1830) 143; Prod. 7 (1839) 433; Sieb. F1. (1947) 209. Lohelia duhia de Vriese in Lehm.
Jap. 2 (1845) 179; Koord. Exk. Fl. Java 3 (1912) PI. Preiss. 2 (1846) 242.— Lightfootia gracilis
300; Nannfeldt. Act. Hort. Gothob. 5 (1929) Miq. Fl. Ind. Bat. 2 (1856) 567, cum var. lavan-
116 Flora Malesiana [ser. I, vol. 6^

Fig. 3. Wahlenbergia marginata (Thunb.) DC. Grassy place near lava rocks, Mt Lawu, East Java
(Arens).

diilaefulia MiQ. Ccimpanopsis marginata O.K. basal part of the filaments about pentagonal or
Rev. Gen. 2 (1891) 378, cum var. rigida, nom.
PI. obtrapezoid, abruptly narrowed, the upper por-
illeg. —Cervicina gracilis Britt. 111. Bot. Cook tion filiform, 0.8-2 mm. Ovary obconical bell-
2 (1901) 56, t. m.—W. bivalvis Merr. Philip. shaped, 2-3-celled; 1-5 mm; style 1.6-5 mm;
J.Sc. 1 (\906)SuY>p\.2A2.— W.gracilenta Lothian, stigma 2-3-lobed. Fruit obconical to bell-shaped,
Proc. Linn. Soc. N.S.W. 71 (1947) 217.— Fig. 2-3-valved, II/2-IO by 1-5 mm.
3-5. Distr. Widely distributed from China and
Perennial. Stem 3-60 cm, glabrous or hairy, or Japan through tropical SE. continental Asia and
only pilose in the lower part. Leaves linear to Malaysia to New Caledonia, Tonga, Australia,
elliptic, the lower ones usually tending to be the Kermadecs, and New Zealand; in Malaysia:
broadest, decreasing in size upwards, 1/5-51/2 cm Java (from Mt Patuha eastwards). Lesser Sunda
by 1/2-5 mm. Inflorescence 1- to few-flowered. Is (Bali, Lombok, Sumbawa, Flores, Alor, Timor),
Pedicels 1-15 cm. Calyx lobes 3-5, linear-lanceo- S. Moluccas (Aru Is), SW. Celebes (Mt Bonthain),
late, acute or ±^ obtuse (China), usually entire, Philippines (Luzon), and New Guinea.
sometimes sparsely dentate, usually about as MiQUEL (Sum. 1860, 234) recorded it from
long as the corolla tube, sometimes shorter or up Sumatra, but this seems erroneous.
to twice as long, 0.8-4 mm. Corolla tube 1-5 mm, Clarke & Hemsley recorded W. gracilis even
lobes 3-5, ovate or elliptic, acute or acuminate, from S. Africa, but I have seen no records from
1.2-7(?) mm. Anthers narrow, V^-2V^ mm long; Africa; this is probably in confusion with W.
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 117

Such shoots may appear frequently in burnt

savannahs and grasslands.
Besides this ontogenous variability there is also
an astonishing individual variability, for instance
in the degree of the hairiness of stem, leaves, and
floral parts, which has sometimes been used as the
only characteristic to separate species (Lothian,
1947; Stearn, 1951). The development of the
indument is not reliable, as it may show
large variation in flowers of a single plant. Hairi-
ness of stems and leaves seems to be very much
dependent on the habitat.
There is further much variation to be observed
F-ig. 4. Distribution of Walileubergia marginata in both the absolute and relative dimensions of
(Thunb.) DC. with localities dotted in the area nearly all the floral parts, without an appreciable
covered by the map. geographical correlation (raciation). This variation
has been drawn for the shape of the filaments
gracilis E. Mey., a nomen nudum and a later in fig. 5. It must be taken into consideration that
homonym. the filaments and their margins are bent inwards
Ecol. In Malaysia only in the mountains (100, already in a very early stage (cf. Decaisne, Rev.
800-900-) 1000-3500 m, in light Eucalyptus and Hort. 3, 1849, 41 fig.); in herbarium material it
Casuarina forest, along forest edges, among rocks, is very difficult to flatten them out because of

on lava-streams, in grassfields, and along trails as their delicacy and entangled fringe-like marginal
an apophyte. In India curiously recorded from hairs.
sea-level upwards (Clarke, I.e.), in Malaysia only length of the calyx lobes may vary as much
The
found at low altitude in the Aru Is. Fl. fr. Jan.- as 1
1/2 mm
in different flowers of a single plant
Dec, specially May-July. during anthesis (Hlb 958-307-469).
It seems that this plant prefers regions with a From the Philippines Merrill has described
feeble to strong dry season which may explain W. biva/vis Merr. which would be characterized
its absence in the wet rain-forest core of West by capsules with 2 apical valves. In the syntypes
Malaysia (Sumatra, Malaya, Borneo). According of Merrill 4361 (Bo, L, Ny) which 1 could
to DocTERS VAN Leeuwen self-pollination is the examine I have only found capsules with 3 valves.
rule in Java. In a number of other specimens ranging through
Vern. Angkeb, kerckan laiuing, patikan, telugi, Celebes, the Lesser Sunda Is, and S. Moluccas
J (local), djiikut riut, S; Papua:
ik, Dunantina, to New Guinea and Thursday Island I have found
iki, Asaro, Kefamo, goiekul, Chimbu, Masul, indeed 2-valved capsules and all these specimens
kulkal, Hagen, Togoba. have very slenderly branched inflorescences
Uses. According to Backer used against skin (except Elbert 1083 from Lombok): Celebes:
diseases on Mt Dieng. BOnnemeijer 11528, partly; Lombok: Elbert
Notes. In the past many authors have already 1332; Aru Is: Buwalda 5354; SE. New Guinea:
pointed to the high degree of variability of this Forbes 847 Thursday Island Jaheri s.n. Warrior
; : ;

species, e.g. De Candolle (1830), Junghuhn Island: Le Guillou Warrior 16.

(1845), Hooker /. (1860), Bentham (1869), But there are other less delicate-branched
Bailey (1900), Cheeseman (1906), and Stearn specimens which have in a single plant 2- and 3-
(1951).
This variability concerns both habit and mor-
phological characters. The distinction between
annual and perennial species, as advanced by
N. E. Brown (1913) and followed by Lothian
(1947) is not reliable and is impracticable. Spe-
cimens flowering in their first year are very thin
and meagre with very small flowers; they have
often been found together in exactly the same
spot with vigorous old plants (Brass 32353, v.
Steenis 18425, 18426). Annual dwarfs with tiny
stems and small flowers are often found as krem-
nophytes on steep talus. Another source of such
dwarfs may be that seeds have germinated in the
wrong season and were forced into flower in a Fig. 5. Wahlenbergia marginata (Thunb.) DC.
juvenile stage, before having attained a good, Geographical survey of stamen types (boiled
full-grown vegetative state. material), a. From Hunan (China), b. Hongkong.
Further the woody shoots of old obviously c. Hongkong, d. Central Java (Mt Sumbing),
perennial plants may produce such thin shoots ('. Lombok, /. Timor, all x 10 (a Fan & Li 57,
(r/. Bth. Fl. Austr. 4, p. 137; Clarke, Fl. Br. b Lamont 405, c Wright s.n., d Lorzing 3. e
Ind. 3, p. 430; and Bailey. Queensl. Fl. 3, p. 922). Elbert 1332, f Forbes 4048).
118 Flora Malesiana [ser. I, vol. 6^

valved capsules: Lombok: Elbert 1557; China: mainly which are somewhat
in the calyx lobes,
Herb. Bog. sine coll. 133584; Celebes: sine coll. shorter compared with the corolla tube {cf.
136. Further there are also slenderly branched Lothian, I.e. and Melville, I.e.). This character,
specimens with 3-valved capsules: Celebes: however, proved to be very variable in the
BuNNEMEiJER 11528, partly, van Steenis 10400. Australian material which I could examine. The
For these reasons W. bivalvis Merr. cannot be relative length of calyx lobes and corolla tube
maintained. differs sometimes considerably in different
Chromosomes. According to the Chromo- flowers of the same specimen (e.g. Hoogland
some Atlas by Darlington & Wylie (1955) IV. 3074, 3075), whereas in the typical trichogyna
gracilenta, W. coiisimilis, W. quadrifida, and W. forms the corolla tube is sometimes much longer
gracilis would have 2 n = 18, 36, 54, and 64 than the calyx lobes.
chromosomes respectively, with the basic numbers It seems that W. trichogyna Stearn, Card.
9 and (?) 8 which might point to the occurrence Chron. Ill, 130 (1951) 169.— Campanula gracilis
of chromosome races. (non FoRST.) Sims, Bot. Mag. 18 (1803) t. 691.—
Campanula vincaeflora Vent. Jard. Malm. (1803)
var. grandiflora Tuyn, nov.var. —
W. consimilis t. 12, nom. illeg. — W. gracilis var. vincaeflora DC.
Lothian, Proc. Linn. Soc. N.S.W. 71 (1947) Mon. Camp. (1830) 142; Prod. 7 (1839) 433.— If.
223; Melville, Hot. Mag. 172 (1959) t. 343.— vincaeflora (Vent.) Decne, Rev. Hort. 3 (1849) 4,
W. gloriosa Lothian, I.e. 224. cum ic. col.; Lothian, Proc. Linn. Soc. N.S.W.
Corolla longior, c. lV2-2'/2 cm longa. 71 (1947) 220, differs only in having a hairy ovary
Similar to the species, but the corolla (in Mai.) and is better subordinated as subvar. trichogyna
l%-2 cm long, the tube c. 6-10 mm, usually c. (Stearn) Tuyn, nov. subvar. It is native in East
lV^-2 times as long as the calyx lobes (3-5 mm). Australia and has not yet been found in New
Calyx lobes always longer than the ovary (2-2 V2 Guinea.
mm). Fruit broad-obconical to bell-shaped, Cruttwell 776 and Brass 22258 have the calyx
41/2-6 by 4—5 mm. lobes clearly dentate and the broadened part of
Distr. Australia (S. New South Wales to S. the filaments seems longer than usual.
Australia and Tasmania), in Malaysia: New As there are found sometimes specimens with
Guinea. a few teeth at the margin of the calyx lobes (e.g.
Notes. Also in this variety the hairiness is Clemens 4873, 5870) and the filaments are very
variable. variable throughout the species, there seems no
Var. grandiflora seemed to differ from the reason to create a new variety.
descriptions of W. consimilis and W. trichogyna

3. CODONOPSIS
Wall, in Fl. Ind. 2 (1824) 103; Komarov, Act. Hort. Petrop. 29 (1908)
Roxb.
98; Chipp, Linn. Soc. Bot. 38 (1908) 374; Anthony, Not. R. Bot. Card. Edinb.
J.

15 (1926) 173; Nannfeldt, ibid. 16 (1931) 149; Bot. Tidsskr. 34 (1940) 381.— G/o-
socomia D. Don, Prod. Fl. Nepal. (1825) 158. Campanumoea Bl. Bijdr. (1826)
726.— Cyclocodon Griff. Not. PI. As. 4 (1854) 279; Kurz, J. As. Soc. Beng. 46, ii
(1877) 209.— Fig. 6-7.
Perennial, erect or climbing herbs with tuberous roots. Leaves opposite, at least
in part. Flowers usually large, peduncled, solitary, axillary, or in cymes on short,
leafless branches. Calyx adnate to the ovary (or below it), persistent, 4-6 spreading
lobes. Corolla wide-campanulate, the base adherent to the ovary, 4-6 lobes, white,
greenish, or tinged purple. Stamens (4-)6, free, inserted near the base of the corolla
tube on the ovary; filaments dilated at the base; anthers free. Ovary 4-6-celled;
placentas axile, thick, with many ovules; style cyHndric; stigmas 4-6 short and
broad lobes. Fruit a berry, subglobose or cylindrical, with truncate top, indehiscent
or an acutish, apically dehiscent capsule. Seeds many, small, ellipsoid.
Distr. About 30 spp. from Turkestan to India, SE. & NE. Asia, Hainan, Formosa, and Japan; 2 spp.
in Malaysia.
Taxon. To establish the proper name and circumscription for this genus I was faced with two aspects,
viz the view taken by several early authors who combined Codonopsis and Campanumoea and those
who kept them separate; among the latter are Chipp, I.e., and Komarov, I.e., among the former for
instance Hooker/. (111. Him. PI. 1855, 116). A third view was taken by Griffith, followed by Kurz,
who also distinguished two genera but of other circumscription.
The only distinction between Codonopsis and Campanumoea is the structure of the fruit, opening
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 119

by 5 small, apical valves (slits) in the first and indehiscent in the latter. When capsular, the valves are
formed protruding apical part of the fruit which is crowned by the style.
in the conically
In order to find out whether this is a really 'natural' character 1 have gone through practically all
species of both genera. If it was really a natural character the species of each genus should by necessity
be more closely allied inter se than with any species of the other genus. This appeared not to be the
case, as the distinction on the fruit characters cuts through another character, viz whether the plants
are really twiners (as in C. javanica) or semi-erect or sprawling (as in C. celebica). It thus happens that
Campcuninwea hmcifolia shows in foliage and habit more resemblance with for instance Codonopsis
purpurea Wall, in RoxB. than with other Cainpanumoeas; reversely Campanumoea javanica shows a
close resemblance to Codonopsis viridis.
A second point in favour of merging the genera is the fact that within Codonopsis in the restricted
sense, Chipp could illustrate not less than 4 very distinct floral types, which, each separately, could
serve for the same aim as a generic one-character difference. Fig. 6. This would be useless, considering
the unmistakably close relationship of all the species together and their distinction against other cam-
panulaceous genera.

Fig. 6. The five floral types in Codonopsis, with diff~erent place of insertion of calyx and corolla, schematic,
largely after Chipp. a-d. Types used for subdivision in Chipp's key. I.e. 375, e. structure in C. parviflora,
after Griffith, Ic. t. 481 (Cyclocodon distans Griff.).

The third point is the fact, as I have found in the relationship between Pratia and Lobelia, that ap-
parently the character of the "berry' versus the 'capsule' does not carry much important generic weight
in certain groups of this family.
two genera I feel strengthened by a succinct note by Diels, who remarked under
In merging the
Campanumoea "von folgender Gattung(= Codonopsis) wohl nicht zu trennen".(Bot. Jahrb. 29, 1901,606).
A similar conclusion was reached by Nannfeldt, who made much study of Codonopsis. He wrote:
"In my opinion the most logical treatment should be to unite these genera (i.e. Codonopsis and Cam-
panumoea), but in order to avoid disagreeable nomenclatural changes it is perhaps more practical to
maintain Campanumoea as a distinct genus on account of its baccate fruits".
As far as I can see all combinations under Codonopsis have been made save one, so that the nomen-
clatural consequences are very slight indeed; the taxonomical gain is, however, considerable.
Chipp, Anthony, and Nannfeldt have subdivided the genus into sections and series. have refrained I

from fitting in this scheme the Malaysian Campanumoeas. I do not believe that they can be main-
tained under one subgenus or section, as they diff"er greatly in habit, one being sprawling and with small,
white flowers (''Cyclocodon ), the other having the typical twining habit and large, greenish flowers of
the "pilosula" group of Nannfeldt.
In such genera in which the species show reticulate affinity, it appears diflficult to make natural sub-
divisions reflecting common ancestry ('Sippen').
Morph. The species of this genus gave occasion to make an observation on the remarkable nature
of the inferior ovary, often called 'calyx tube' by Anglo-Saxon botanists. As Chipp has illustrated four
floral types can be distinguished in Codonopsis, depending on the insertion of the calyx and corolla,
v/2 (1) both calyx and corolla adnate to the ovary halfway, (2) ditto so up to the ovarial apex, (3) corolla
adnate to the apex, but calyx only half-way so, (4) corolla adnate to the ovarial apex, but calyx inferior
and properly free. Fig. 6.
In no instance a free calyx 'tube' is formed.

KEY TO the species

1. Twining. Leaves with cordate base. Flowers 1.5-3.8 mm, wide-campanulate, greenish.
1. C. javanica
I. Sprawling or erect. Leaves decurrent at the base. Flowers ^/4-l'/2 cm, white. 2. C. lancifolia

1. Codonopsis javanica (Bl.) Hook. /. III. Him. panumoea javanica Bl. Bijdr. (1826) 727; DC.
PI. (1855) 116, t. 168; Miq. F1. Ind. Bat. 2 (1857) Prod. 7 (1839) 423; Hassk. Cat. Hort. Bog.
566; Merr. Lingn. Sc. J. 6 (1930) 289.— Cam- (1844) 106; van Houtte, FI. Serres 12 (1857)
120 Flora Malesiana [ser. I, vol. 6^

157, t. 1264; Hook./. &

Th. J. Proc. Linn. Soc sometimes acuminate, serrulate to serrate, some-
Bot. 2 (1858) 18; KuRZ, J. As. Soc. Beng. 46, ii times crenate, on both sides puberulous to hairy,
(1877) 209; Clarke, FI. Br. Ind. 3 (1881) 435 sometimes glabrous, underside often pruinose;
FoRB. &Hemsl. J. Linn. Soc. Bot. 26 (1889) 8 petiole IV2-6V2 cm. Flowers axillary, l'/2-3.8
DiELS, Bot. Jahrb. 29 (1901) 606; Making, Bot mm. Pedicels 1-5 V^ cm. Calyx lobes oblong to
Mag. Tokyo 22 (1908) 155, incl. var. japonica lanceolate, 1-23 by 3^ mm, entire or denticulate
1

KooRD. Exk. Fl. Java 3 (1912) 300; Fl. Tjib. 3 to serrate, glabrous or puberulous to hairy.
(1923) 54; Heyne, Nutt. PI. 2 1427
(1927) Corolla 12-35 mm, more than half-way lobed,
Danguy, Fl. Gen. I.-C. 3 (1930) 694 &
78,10 f. greenish-white or yellowish-white, outside some-
HocHR. Candollea 5 (1932) 290; Merr. Lingn. times hairy, tube inside with purplish streaks or
Sc. J. 11 (1932) 60; Craib, Fl. Siam. En. 2 (1936) veins, lobes ovate, acuminate. Anthers c. 2-4 mm;
307; Masamune, Fl. Kain. (1943) 330; Back. filaments 3-8 mm, linear, broadened towards the
Bekn. Fl. Java (em. ed.) 8 (1949) fam. 183, p. 3; base, glabrous. Style 5-10 mm, glabrous or hairy
Makino, 111. Fl. Japan (1954) 81, cum var. C. — with 3-6 ovate-acute to elliptic lobes. Ovary
cordata Hassk. Nat. Tijd. N.I. 10 (1856) 9; Ned. (3-)4-5(-6)-celIed, the outside partly concealed
Kruidk. Arch. 4 (1856) 1; MiQ. Fl. Ind. Bat. 2 by the corolla, campanulate, visible part from the
(1857) 566; Walp. Ann. 5 (1858) 393; Bot. Mag. outside 3-4 by 3-4 mm, glabrous. Fruits sub-
89 (1863) t. 5372. Campaiuimoea cordata MiQ. globular, Y2-IV2 by I-2V2 cm, red to dark-purple
Sum. (1862) 599; Maxim. Bull. Ac. Imp. Sc. to black-bluish, at the base sustained by the
Patersb. 12 (1868) 68; Craib, Kew Bull. (191 1) 404. persistent and patent or sometimes reflexed wine-
— Campaiuimoea Japonica Maxim. Bull. Ac. Imp. red calyx lobes and crowned by the withered
Sc. Petersb. 12 (1868) 67, non Siebold et Morren, corolla; walls membranous to coriaceous, some-
1863.— C.co/-i///o//flKoMAROV, Act. Hort. Petrop.29 times fleshy; placentas fleshy. Seeds ovoid to
(1908) 108. Campaiuimoea maximowiczii Honda, ovoid-cylindrical, c. 1 mm, reticulate, light brown.

Bot. Mag. Tokyo 50 (1936) 389.— Fig. 7. Distr. In SE. Asia (from the Deccan through
A sinistrorse-twining plant, up to 2 (ac- m the Himalayan region, Burma, and Siam to South
cording to KooRDERS 6-14 m
long); stem 1 '/2-4 and Central China), Japan, Hainan; in Malaysia:
mm diam., glabrous. Leaves opposite, higher up Central to South Sumatra (Mts Kerintji and
often spirally arranged, ovate to oblong-ovate, Dempo), Java (from Mt Pangrango eastwards).
2.6-8 by 2-5 cm, base cordate, apex obtuse, acute. Ecol. In open forest, mostly on forest edges,
in secondary forest and thickets, sometimes in
grassy fields, 900-2200 m. Fl. fr. Jan.-Dec.
Vern. Kitjepot, susu munding,
srintil-srintil,
S, gutji, indil-indil, sigerpolo, urek-urek polo, J.
Uses. The fruits are eaten and the tuberous
roots are used for drugs.

2. Codonopsis lancifolia (RoxB.) Moeliono, nov.

—
comb. Campanula lancifolia RoxB. FI. Ind. 2
( 824) 96
1 DC. Prod. 7(1839) ASS. —Campaiuimoea
;

celebica Bl. Bijdr. (1826) 727; DC. Prod. 7 (1839)

423; Clarke, Fl. Br. Ind. 3 (1881) 436; Boerl.
Handl. 2 (1891) 259; Staff, Trans. Linn. Soc. II,
4 (1894) 188; Gamble, J. As. Soc. Beng. 74, ii
(1905) 53; Robinson, Philip. J. Sc. 3 (1908)
Bot. 216; Craib, Kew Bull. (1911) 404; Diels,
Bot. Jahrb. 55 (1917) 121; Merr. En. Born.
(1921) 585; Ridl. Fl. Mai. Pen. 2 (1923) 202;
Merr. En. Philip. 3 (1923) 587; Danguy, Fl.
Gen. I.-C. 3 (1930) 693; Craib, Fl. Siam. En. 2
(1936) 307; Masamune, En. Pi. Born. (1942) 123;
Back. Bekn. Fl. Java (em. ed.) 8 (1949) fam. 183,
p. 4.— C. truncata Wall. [Cat. (1829) //. 1301,
nomen] ex DC. Mon. Camp. (1830) 122; Prod.
7 (1839) 423; Miq. Fl. Ind. Bat. 2 (1857) 566.—
C. albiflora Griff. Not. PI. As. 4 (1854) 279.—
Cyclocodon adnatus Griff. I.e. 278. C. celebica —
Miq. Fl. Ind. Bat. 2 (1857) 566.— C. leucocarpa
Miq. I.e. 565. Cyclocodon truncatum Hook. /. &
Th. J. Proc. Linn. Soc. Bot. 2 (1858) 18.—
Cyclocodon lancifolium Kurz, Flora 55 (1872)
303; J. As. Soc. Beng. 46, ii (1877) 2\0.—Cam-
Fig. Codonopsis javanica (Bl.) Hook. /. a.
7. panumoea axillaris Oliv. in Hook. Ic. Ill, 8
Flower, sustained by the star-shaped calyx
b. fruit (1888) t. 1775; Hemsl. J. Linn. Soc. Bot. 26
and topped by the marcescent corolla, all X 2/3. (1889) 7. Campanumoea truncata Diels, Bot.
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 121

Jahrb. 29 (1901) 606; Making, Bot. Mag. Tokyo seen), Sumatra, E. Java (Pantjur Idjen, once
8 (1904) 21; Merr. Philip. J. Sc. 7 (1912) 104.— found), N. Borneo (Kinabalu), Celebes, Philip-
Campciinimoea laricifulia Merr. En. Philip. 3 pines (N. Luzon; Panay; Mindanao), Moluccas
(1923) 587; Lingn. Sc. J. 5 (1927) 181; Pap. (Buru and Ambon), and New Guinea.
Mich. Ac. Sc. 20 (1935) 111; Yamamoto, Obs. Ecol. In open terrain or along forest-borders
Fl. Form. 13 (1936) 147; Masamune, FI. Kain. and trails, on tali and near streamlets, rarely
(1943) 33; Hara, En. Sperm. Jap. 2 (1952) 96. in forest regrowth and young secondary forest,
Sprawling or erect herb, with a hollow stem, often in wet places, 280-1500 m. Fl.fr. Jan. -Dec.
c. 3 m high, nearly glabrous or hairy, mostly Vern. Gorc/ang-gordang, M
(Toba), emiapagar,
branched. Leaves opposite, petioled, lower ones Palu (W. Cel.); Philippines: lakoronbolan. Buk.
ovate to ovate-elliptic, higher ones elliptic, often Note. There are two subspecies which largely
bract-like, 3-14 by 1-4 cm; acute to acuminate, replace one another geographically.
bluntly edged at the base, coarsely serrate to
serrulate, upperside glabrous, underside glabrous ssp. lancifolia. — All synonyms except C. celebica
or puberulous to hairy; petiole 3-15 mm, glabrous. and C. leiicocarpa.
Flowers 7-15 mm, axillary, solitary or in cymes Leaves generally large. 8'/2-16'/2 by 3-7 '/t cm,
of 3, through reduction of the higher leaves re- towards the apex of the stems not very much
sembling a terminal panicle. Pedicels or peduncles reduced, never glaucous beneath, entirely glabrous.
(I-)3-6 cm, glabrous, with two bracteoles, V2-2 Calyx lobes coarsely dentate to subpectinate.
cm long, linear to elliptic, glabrous or hairy. Filaments distinctly widened downwards, either
Buds winged, ovate. Calyx lobes 4-7, lanceolate, triangular or roundish, long-hairy inside.
4—10 by mm, entire to dentate-serrate to pinnati-
1 Distr. Continental Asia, in Malaysia: N. Su-
fid, carnosulate, glabrous to puberulous. Corolla matra (Atjeh, Toba), Philippines, and Moluccas
white, pale pink, lilac, outside glabrous, corolla (Buru, Ambon).
tube as long as the lobes, lobes ovate to trian-
gulate, acute, entire. Stamens 4-6 mm. Anthers ssp. celebica (Bl.) Moeliono, comb. nov. Cam- —
as long the filaments. Filaments linear to
as —
paniimoea celebica Bl. I.e. C. celebica (Bl.)
broadened at the base in various degree (deltoid Miq. — C. leiicocarpa MiQ.
to ovate), the broad forms inside with long hairs Leaves averagely smaller, 5'/2-8(-l Vi) by 1

at the base. Style glabrous or scabrid. Ovary 11/^_23^(_33^) cm, towards the apex distinctly
cupular to campanulate, 2-4 mm diam., glabrous shorter giving the apical part of the stem the ap-
to puberulous. Berry ± globular with a flattened pearance of a panicle, glaucous and hairy beneath
apex, the persistent calyx adnate about half-way, or glabrous. Calyx lobes entire to toothed. Fila-
greenish, turning to white, c. 1/2-1 cm diam. ments linear to triangular-broadened towards the
Seeds many, testa fine-reticulate. base, glabrous (once found short hairy inside.
Distr. SE. Asia (India: Himalaya and Sikkim; OUWEHAND 215).
Burma) to S. China, Formosa, and Hainan; in Distr. Siam (Kerr 1217). in Malaysia: Central
Malaysia: Malay Peninsula (Selangor: Taiping and South Sumatra, SE. Java, Borneo. Celebes,
Hills; Perak: Semangkok Pass, ex Ridley, not Philippines, Moluccas, and New Guinea.

6. LOBELIA
LiNNE, Gen. PI. (1754) 401; Sp. PI. 2 (1753) 929, uon sensii Adans. Fam. PI. 2

(1763) 157 et Miller, Gard. Diet. ed. 8 (1768), cf. nom. gen. cons. n. 8716; DC.
Prod. 7 (1839) 357; Wimmer, Ann. Naturh. Mus. Wien 56 (1948) 317; Pfl. R. Heft
107 (1953) 40S.—Pratia Gaudich. Ann. Sc. Nat. 5 (1825) 103; Presl, Prod. Mon.
Lob. (1836) 46; Wimmer, Pfl. R. Heft 106 (1943) \04.~Dortmaiina Adans. Fam.
PI. 2 (1763) 134; Steud. Nomencl. (1840) 526; O. K. Rev. Gen. 2 (1891) 379 (Dort-
—
manuid). Rapuutium Miller, Gard. Diet. ed. 8 (1768); Presl, Prod. Mon. Lob.
(1836) U.—Piddmgtouia DC. Prod. 7 (1839) 341 Miq. Fl. Ind. Bat. 2 (1857) 572.—
;

Isolobus DC. Prod. 7 (1839) 352.— Speirenia Hook/. & Th. J. Proc. Linn. Soc.
Bot. 2 (1858) 27.
Herbaceous, annual or perennial, sometimes woody below, rarely arborescent (Afri-
ca, Hawaii), often laticiferous. Leaves spirally arranged, alternate (distichous), or in
rosettes. Flowers axillary or in racemes or panicles, with or without bracts, 5-merous,
epigynous, ^, rarely unisexual (some Australian spp. dioecious), protrandrous.
Calyx lobes well-developed. Corolla gamopetalous, zygomorphic, with a dorsal
slit mostly to or near the base; limb with 2 dorsal lobes, mostly diverging from
122 Flora Malesiana [ser. I, vol. 6^

the 3 others which form a trifid whole consisting of a ventral lobe and 2 laterals;
lobes valvate in bud, connate to various degree. Stamens 5, akernate with the corolla
lobes, free or adnate to the corolla tube; filaments linear, sometimes broadened
at the base; anthers basifixed, introrse, 2-celled, cells opening lengthwise. Disk
absent. Ovary 2-celled; style 1, at the apex below the 2 stigmas provided with 'col-
lecting hairs', during anthesis lengthening with appressed stigmatic lobes through
the anther tube. Ovules oo, axile, anatropous. Fruit fleshy to dry in various degree,
i.e. a berry or an apically 2-valved capsule, crowned by the persistent calyx lobes.

Seeds oo, provided with endosperm.

Distr. Probably more than 200 spp., mostly in the tropics and subtropics, especially in America.
Ecol. The Malaysian species are mostly hill and mountain plants in everwet country, except L.
alsinoides, which is typical for lowland seasonal climatic conditions, and L. zeylanica, which grows from
the lowland up into the montane zone.
The flowers are resupinate, even before they are open. There are 'collecting hairs' under or at the
base of the stigmas of the bifid style. By the lengthening of the style, the unopened stigmas enter into
the anther tube, pushing the pollen out of the tube. This occurs before the stigmas are receptive, for
receptiveness is acquired long after the pollen is shed.
Nomencl. Unfortunately former typification of some common species has been unsatisfactory which
has necessitated to accept several name changes. Besides, the taxonomic changes proposed here, by
merging Pratia with Lobelia, and severe reduction of the number of species of Pratia, have resulted in a
rather complicated synonymy.
Taxon. Hitherto the genus Pratia has almost unanimously been distinguished from Lobelia by the
fruit, it being baccate and non-dehiscent against being capsular and apically 2-valved in Lobelia.
By the subdivision of the Lobelioideae into berried and capsular genera, advanced first by De Can-
DOLLE (1839) and followed almost unanimously up to the monograph by Wimmer (1943, 1953) Lobelia
and Pratia have become distant allies. The lively coloured berries of the Pratias which are native in
the Pacific area of the southern hemisphere represent a showy character, but it should be added that
flowering specimens cannot be distinguished from certain Lobelias which are also prostrate and rooting,
humble plants in the same area.
In my opinion it seems unquestionable that they are very closely allied. This opinion is sustained by
the curious fact that in the abundant material of Pratia angiilata, which I could study, the Asiatic and
West Malaysian specimens have definitely berry-like, indehiscent fruits, but those of Celebes and New
Guinea show in otherwise 'inseparable' specimens various transitions in the mature state (as shown
by ripe seed inside). In East Malaysia the fleshy pericarp becomes 'drier' and thinner, almost mem-
branous, and the lengthwise nerves become more pronounced; the apex of the fruit which is in typical
Pratia berries flat, becomes convex and tends to become tardily 2-valved. In one case of profuse material
(Eyma 1161) fleshy berries and true capsules could be found in one collection. In this area also the
variability of other characters of the species (hairiness, size of the flowers, leaves, pedicels, etc.) is wider
than in West Malaysia and continental Asia, which have, obviously, a more homogeneous 'marginal'
population. The tendency to possess a conical ovary is continued towards the South Pacific.
These observations have induced me to merge Pratia with Lobelia. It would be of profound interest
if additional field observations could be made to verify my conclusions.

In specific delimitation I have allowed far more variation than Wimmer, Skottsberg, and some
other authors.

KEY TO THE SPECIES

1. Stems developed.
2. Stems erect or rarely ascending, not rooting at the nodes.
3. Stems terete or ribbed, not 2-3-angled. Rather coarse plants.
4. Leaves mainly in a dense, persistent rosette, i sparsely long-white-pilose along the margin
and on the upper surface. Stems angular, unbranched. Cauline leaves oblanceolate, upwards
gradually smaller, 2-5 by '/2-I cm. Perennial plant, with a strong rhizome.
2. L. sumatrana
4. Flowering stems without such a rosette. Leaves glabrous or hairy but not long-white-pilose,
elliptic-lanceolate, mostly much larger.
5. Flowers solitary axillary, 2-4|/2 cm long. Berry globose, 7-12 mm diam., on a reflexed pedicel,
violet, finally black-purple 9. L. montana
5. Flowers in terminal panicles or terminal or leaf-opposed racemes.
6. Racemes terminal, later leaf-opposed. Fruit a berry 10. L. borneensis
6. Racemes or panicles terminal. Fruit capsular 1. L. nicotianaefolia
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 123

3. Stems 2-3-angled, sometimes winged along the edges. Rather delicate, annual herbs.
7. All anthers bearded at their apex. Seeds trigonous.
8. Free basal part of the 2 anterior filaments twice as broad as that of the others and densely
haired. Stem sharply triangular-winged. Stem, leaves, pedicels, and calyx glabrous or sparsely
hairy. Leaves ovate-oblong or even narrower, sessile, narrowed or contracted towards the base.
3. L. alsinoides
Free basal part of all filaments of equal width. Stem terete, under each leaf with 2 narrow ridges.
8.
Stem, leaves, pedicels, and calyx crisped-patent-hairy. Leaves ovate to roundish with very
blunt to subcordate base set off against a distinct petiole (l-)3-20 long mm
4. L. zeylanica
7. Only the two anterior anthers bearded at their apex. Seeds ellipsoid 5. L. heyniana
. .

2. Stems creeping or ascending, rooting at the nodes.

9. Leaves spirally arranged. All anthers bearded at their apex 4. L. zeylanica
9. Leaves distichous. Only 2 anthers bearded at their apex.
10. Leaves sessile, densely set (almost equitant), semi-amplexicaulous, fleshy. Plant glabrous.
Filaments glabrous 12. L. conferta
10. Leaf-base not semi-amplexicaulous.
11. Leaves glabrous, sessile, with acute base, subentire, oblong to elliptic, rather narrow. Testa
smooth 6. L. chinensis
Leaves exceptionally glabrous, at least the lower almost always petioled, mostly ovate to rounded,
11.
with a rounded (very rarely acutish) base.
12. Corolla hardly twice as long as the calyx lobes, campanulate, with ovate-deltoid lobes. Hairs
more-celled. Testa smooth 8. L. brachyantha
12. Corolla at least c. 3 times as long as the calyx lobes, not campanulate, with narrow-lanceolate
or ovate-lanceolate lobes. Hairs 1-celled. Testa finely reticulate ...
11. L. angulata
1. Stemless. Flowers solitary axillary, c. V2 cm long. Leaves very thin, roundish, petioled.
7. L. archboldiana

1. Lobelia nicotianaefolia Roth e.\ R. & S. Syst. Soc. Bot. 25 (1890) 41; Danguy, Fl. Gen. I.-C.
Veg. 5 (1819/720) 47; Roth, Nov. PL Sp. (1821) 3 (1930) 675; Wimmer, Pfl. R. Heft 107 (1953)
143; Wall, in Roxb. Fl. Ind. 2 (1824) 110; PI. 653. L. excelsa Lesch. ex Roxb. Fl. Ind. 2
As. Rar. 2 (1831) 43; G. Don, Card. Diet. 3 (1824) 114, non Bonpl. (1813); Wall. PI. As.
(1834) 709; DC. Prod. 7 (1839) 381; Wight, Rar. 2 (1832) 42; G. Don, Gard. Diet. 3 (1834)
lllustr. 2 (1850) 111, t. 135 f. 1-10; Hook./. & 709; DC. Prod. 7 (1839) 381 Wight, Ic. 4 (1850)
;

Th. J. Proc. Linn. Soc. Bot. 2 (1858) 29; Hook. t. 1173-4; Thwaites, En. PI. Zeyl. (1860) 170;
/. Bot. Mag. 92 (1866) t. 5587; Clarke, Fl. Br. Clarke, Fl. Br. Ind. 3 (1881) 427.— L. stimulans
Ind. 3 (1881) 427, incl. vcir. tric/umdra; Trim. Ham. ex D. Don, Prod. Fl. Nepal. (1825) 157.—
Handb. Fl. Ceyl. 3 (1895) 57; Pearson, J. Linn. L. piirpiirescens Wall. Cat. (1828) 1307, non R.Br.
Soc. Bot. 34 (1898) 348; Merr. & Merritt, (1810), nomen nudum ab auctore ipso ibidem
Philip. J. Sc. 5 (1910) Bot. 392; Merr. En. Philip. relictum p. 157. L. colorata Wall. PI. As. Rar.
3 (1923) 588; Skottsb. Medd. Goteb. Bot. 2 (1831) 42; DC. Prod. 7 (1839) 380; Hook. /.
Triidg. 4 (1928) 9-13, f. 1, 8-11, 12b, incl. var. & Th. J. Proc. Linn. Soc. Bot. 2 (1858) 28;
macrostemon. I.e. 13, f. 13-14; Alston, Fl. Ceyl. Drury, Handb. Ind. Fl. 2 (1866) 108; Clarke,
Suppl. (1931) 175; Elmer, Leafl. 9 (1934) 3180, Fl. Br. Ind. 3 (1881) 426; Wimmer, Pfl. R. Heft
incl. var. mollis; Kausik, J. Ind. Bot. Soc. 17 107 (1953) 655, incl. var.— Rapuntium pyramidale
(1938) 161-168; Wimmer, Pfl. R. Heft 107 (1953) Presl, Prod. Mon. Lob. (1836) 23. Rapuntium
643, incl. var. nicotianaefolia, bibarbata, & tri- nicotianaefolium Presl, I.e. 24. Rapuntium colo-
chandra; Santapau, Rec. Bot. Surv. Ind. 16 —
ratum Presl, I.e. lA. Rapuntium wallicliianum
(1953) 157.—L. pvramidalis Wall. As. Res. 13 Presl, I.e. 24. Rapuntium roseum Presl, I.e.
(1820) 376; D. Don, Bot. Mag. 50 (1823) t. 2387; 24. Rapuntium lesehenaultianum Presl, I.e. 24.
Wall, in Roxb. Fl. Ind. 2 (1824) 113; D. Don, L. wallichii Steud. Nomencl. 2 (1841) 62, nomen
Prod. Fl. Nepal. (1825) 57; Wall. PI. As. Rar. superfl. ad L. purpurescens Wall. —
L. robusta
2 (1831) 42; G. Don, Card. Diet. 3 (1834) 709; Wall, ^.v Voigt, Hort. Suburb. Calc. (1845) 367,
DC. Prod. 7 (1839) 381, incl. var. /3; Hook. /".
non Graham ( 831). L. eurostos Voigt, I.e. 367.
1

& Th. J. Proc. Linn. Soc. Bot. 2 (1858) 29; L. aromatica Moon [Cat. PI. Ceyl. (1824) 14,
Clarke, Fl. Br. Ind. 3 (1881) 426; Forbes & nomen] ex Wight, Ic. 4 (1850) 2, t. 1172; Hook.
Hemsley, .T. Linn. Soc. Bot. 26 (1889) 3; Skottsb. /. & Th. J. Proc. Linn. Soc. Bot. 2 (1858) 29;
Medd. Goteb. Bot. Tradg. 4 (1928) 17, 21, f. 12e, Alston, Fl. Ceyl. Suppl. (1931) 175.-1. tri-
25-31; Danguy, Fl. G6n. I.-C. 3 (1930) 676, chandra Wight, Ic. PI. Ind. Or. 4 (1853) t. 1171;
f. 76 1-4; Craib, Fl. Siam. En. 2 (1936) 304; Skottsb. Medd. Goteb. Bot. Triidg. 4 (1928) 16,
Wimmer, Pfl. R. Heft 107 (1953) 646.— L. rosea f. 12d, 18.— £. wallichiana Hook. /. & Th. J.
Wall, in Roxb. Fl. Ind. 2 (1824) 115; PI. As. Proc. Linn. Soc. Bot. 2 (1858) 29; Kurz, J. As.
Rar. 2 (1831) 42, t. 152; G. Don, Gard. Diet. 3 Soc. Beng. 46, ii (1877) 211.— L. ereeta Hook./.
(1858) 29; Drury, Handb. Ind. Fl. 2 (1866) 108, & Th. J. Proc. Linn. Soc. Bot. 2 (1858) 28, non
excl. syn.: Kurz, J. As. Soc. Beng. 46, ii (1877) de Vriese (1845); Clarke, Fl. Br. Ind. 3 (1881)
212; Clarke, Fl. Br. Ind. 3 (1881) 427; J. Linn. 426. Dortmannia lesehenaultiana O. K. Rev.
124 Flora Malesiana [ser. 1, vol. 6^

Fig. 8. Lobelia lucoiianaefulia Roth ex R. & S. Bandarawele, Ceylon, c. l-lVi m tall, 1956.

Gen. (1891) 972.— Dortmannia erecta O. K. I.e.

1 79; Ann. Naturh. Mus. Wien 56 (1948) 367, incl.
972. Dortmannia lucoticimiefoHa O. K. I.e. 973. var. sarasinoriim; Pfl.R. Heft 107 (1953) 652.—
Dortmannia rosea O. K. I.e. 973. Dortmannia L. leueanthera Kerr, Kew Bull. (1936) 34; Craib,
eolorata O. K. I.e. 973. Dortmannia pyramidalis Fl. Siam. En. 2 (1936) 304; Wimmer, Pfl. R. Heft
O. K. I.e. 380. L. seguinii Leveille & Van, in 107 (1953) 641.— L. palustris Kerr, Kew Bull.
Fedde, Rep. 12 (1913) 186; Wimmer, Pfl. R. Heft (1936) 35; Craib, Fl. Siam. En. 2 (1936) 304.—
107 (1953) 648, incl. var. —
L. fossarum Wimmer, L. eamptodon Wimmer, Ann. Naturh. Mus. Wien
Akad. Anz. Wien n. 14 (1924) 3. L. eryliae 56 (1948) 366; Pfl. R. Heft 107 (1953) 631.—L.
Fischer, Kew Bull. (1928) 141; Wimmer, Pfl. R. beddomeana Wimmer, Pfl. R. Heft 107 (1953)
Heft 107 (1953) 652.—L. lesehenaiiltiana Skottsb. 645.— Fig. 8-9.
Medd. Goteb. Bot. Tradg. 4 (1928) 4, f. 3-7; A
coarse herb, up to 2(-4i/2) m. Stem terete at
Wimmer, Pfl. R. Heft 107 (1953) 659.—£. doniana the base, angular towards the apex, hollow, simple
Skottsb. Medd. Goteb. Bot. Tradg. 4 (1928) 19, or apically branched. Leaves spirally arranged,
f. 12, 19-24, nomen illegit. —L. philippinensis densely set in juvenile plants, oblong to narrow-
Skottsb. I.e. 13, f. 12c, 15-17; Steen. Bull. Jard. lanceolate, the lower ones sometimes obovate-
Bot. Btzg III, 13 (1934) 178; Steup, Trop. Natuur oblong, gradually decurrent towards the petiole-
27 (1938) 142, f. 60; Toxopeus, Ac. 109, f. 21.— like base, 50-10 by 8-4 cm to 9-3 by 3-3^ cm;
L. epilobioides Wimmer, in Fedde, Rep. 38 (1935) acuminate towards the apex (tip acute or blunt),
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 125

cohering.) Filaments free and hairy at the base,

upwards puberulous or hairy, 8-10 mm; anthers
glabrous or hairy on the connectives, the two
anterior ones with a hair tuft on top, from dorsal
to ventral 3-4 to 2-3 mm. Ovary roundish cupular
or narrower, 7-12-nerved, glabrous to densely
hairy, 4-10 by 2-4 mm; style glabrous. Capsule
cylindric-cupular, %-l cm by 4-6 mm, glabrous
or hairy. Seeds flattened ellipsoid, '/2-5/8 mm,
brown and smooth.
Distr. SE. Asia (from the Deccan to S. China),
Formosa, in Malaysia: Philippines (Luzon and
Biliran), Celebes (Central part and SW. peninsula
as far as Mt Bonthain). Fig. 10.
Ecol. Open places on ridges in mossy forest,
often on grassy mountain slopes and hill sides,
600-2300 m. Fl. fr. Dec.-Aug.
Vern. Philippines: Adlabong, katlahung, kany-
uong, Ig., baliiiyiiiigyung, balyongyoiig, lungog-
Bon.
litngog, siibasob,
Uses. The milky juice is said to be poisonous
(Eyma in sched.).
Notes. After having studied many specimens
both from Malaysia and continental Asia I have
come to the conclusion that this complex represents
one widely distributed and very polymorphous
species, both in Asia and in Malaysia. Characters
used by former authors to distinguish microspecies
often do not hold in one single specimen bracteoles:

may be present or absent, may be inserted at the

base or in the middle of the pedicels; sepals may
be toothed or not, plants may be branched or
unbranched, hairy or glabrous. The differentiating
characters of 'species' in this complex have gone
into hair-splitting detail.
In general there are in SE. Asia two groups
according to the size of the flowers, but they do
not show a geographical replacement. There are
Fig. 9. Lobelia nicotianaefulia Roth ex R. & S. many local forms but in my opinion they should
SW. Celebes. 1937. not be named.
Kausik (J. Ind. Bot. Soc. 17, 1938, 161-167),
serrate-toothed along the margin, hairy on both who has studied the gametogenesis and embryo-
sides, especially on the nerves, rarely glabrous, logy of this species, stated that the chromosome
decreasing in size towards the apex. Racemes
up to 45 cm long, together often forming a large,
leafy panicle; rachis angular, hairy. Bracts under
the lower flowers exceeding the pedicels and connate
with them at the base, the higher ones shorter and
higher connate, finally linear-subulate. Pedicels
1-2 1/2 cm, obliquely patent, ascending, terete,
hairy; bracteoles 2, minute, 3-12 mmlong (in
Indian material often caducous or absent).
Flowers V^-3% cm long, variable in colour, pale,
I

whitish, blue, dark purple, or rose. Sepals linear

to lanceolate, 4-12 by 1 mm, entire to distinctly
toothed, acute, glabrous or hairy. Corolla -^4-3
cm, outside glabrous or hairy, inside hairy to
densely hairy, often with two gibbosities; dorsal
lobes half as high connate with the lateral ones as
compared with their junction with the ventral
lobe, three times as long as the other lobes or
even longer, linear; ventral and lateral lobes about
equal in length, the lateral ones slightly falcate, Fig. 10. Distribution of Lobelia nicotianaefolia
all three ovate-acute, with a slightly crenate Roth ex R. & S., localities in Malaysia indicated
margin. (In immature corollas the lobes are by dots.
126 Flora Malesiana [ser. I, vol. 6^

number of a plant collected at Koppa (Mysore) 2. Lobelia sumatrana Merr. Not. Nat. Ac. Sc.
is n = 14. Philad. 47 (1940) 9; Wimmer, Pfl. R. Heft 107
(1953) 654.— L. sp. Steen. Tijd. Kon. Ned. Aardr.
Gen. 55 (1938) 800. Fig. 11. —
Erect, perennial herb, 15^0 cm, with a firm
rootstock. Stem angular, glabrous to sparsely pi-
lose. Rosette leaves narrowly spathulate to obovate-
lanceolate, tapering towards the base, shallowly
crenate-dentate, blunt, 2-5 by 1/2-I cm; cauHne
ones erect, narrower, more distinctly sessile and
smaller, all pilose. Raceme unbranched, up to c.
10-flowered, rachis 5-10 cm, glabrous. Bracts
herbaceous, ovate to elliptic-oblong, resembling
the leaves but smaller, 5-15 mm
long. Pedicels
terete, pilose, 5-8 mm, with two small, linear,
hairy bracteoles. Flowers IV2-21/2 cm long. Calyx
lobes oblong, c. 5 by 1 V2-2 mm, blunt to broadly
triangular at the apex, shallowly crenate, glabrous
or pilose. Corolla 3/^-2 cm, slit to the base, lilac
or pale purple, purple-veined, inside and on the
nerves and margins outside hairy, dorsal lobes
connate with the lateral ones for 4-5 mm, lateral
lobes connate with the ventral one for 7-9 mm;
dorsal lobes narrowly lanceolate, boat-shaped,
9-11 by 2 mm, acute, lateral and ventral lobes
about equal, oblong-lanceolate, 6-8 by 1 Vi-l mm.
Filaments 6-8 mm, up to 1/3 free and hairy,
anthers from dorsal to ventral 4-2 mm, dorsally
hairy, the ventral ones with an apical hair tuft.
Ovary l-AVz by 2-4 mm, trumpet-shaped to cam-
panulate, distinctly ribbed, glabrous to densely
hairy; style glabrous. Capsule c. 5 by 4 mm,
cupular, campanulate to ovoid, hirsute to pilose.
Seeds V2 by Vi mm, flattened ellipsoid, light brown
and smooth.
Distr. Malaysia: N. Sumatra (Gajo Lands:
Mts Losir, Kemiri, Goh Lembuh).
Ecol. Mountain heaths and meadows (blangs),
common, 2400-3300 m. Fl. fr. Febr.-May.
Note. Obviously related to certain SE. Asiatic
forms of L. nicotianaefolia described as L. colorata
Wall, (specially Clarke 42477), differing by the
angular stem, large rhizome, smaller narrow
spathulate leaves, and a persistent rosette of leaves.

3. Lobelia alsinoides Lamk, Diet. Bot. 3 (1791)

588; R. & S. Syst. Veg. 5 (1819) 60; DC. Prod.
7 (1839) 378; Sond. Fl. Cap. 3 (1865) 539,
excl. svn. L. trialata; Wimmer, Ann. Naturh.
Mus. Wien 56 (1948) 360; Pfl. R. Heft 107 (1953)
571, f 93g = icon, specim. Lamarck.; Santapau,
Rec. Bot. Surv. Ind. 16 (1953) 158.—L. filiformis
(noil Lamk) Cav. Ic. 6 (1801) 7, t. 51 1, f. \.—L.
filiformis var. luzoniensis Pers. Syn. 2 (1807) 214;
R. & S. Syst. Veg. 5 (1819) 61; G. Don, Gard.
Diet. 3 (1834) 713; DC. Prod.7 (1839) 368;
MiQ. Fl. Ind. Bat. 2 (1856) 577; F.-Vill. Nov.
App. 4 (1880) 121.— L. trigona Roxb. [Hort.
Beng. (1814) 85, nomen] Fl. Ind. 2 (1824) 111;
Fl. Ind. ed. Carey (1832) 506; G. Don, Gard.
Diet. 3 709; DC. Prod. 7 (1839) 359;
(1834)
Hook. /. 4 (1841) t. 358, excl. syn. L.
Ic. PI.
trialata et L. heyneana; Wight, Ic. 4 (1848) t.
Fig. 1 1. Lobelia stiinatraim Merr. Mt Losir, Gajo 1170; MiQ. Fl. Ind. Bat. 2 (1857) 574; Hook./.
Lands (N.Sumatra), 1937. & Th. J. Proc. Linn. Soc. Bot. 2 (1858) 27, p.p.;
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 127

MiQ. Sum. (1861) 234; KuRZ, J. As. Soc. Beng. glabrous or sparsely hairy. Corolla 4-12 mm,
46, ii (1877) 211; Clarke, F1. Br. Ind. 3 (1881) varying from bright blue to violet (sometimes, in
423; Trim. F1. Ceyl. 3 (1895) 56; Koord. Exk. Celebes, white), inside hairy (except for the
Fl. Java 3 (1912) 302; Yamamoto, Obs. Fl. specimens described as L. hosseusii which are
Formosa 13 (1936) 148, excl. syn. L. trialata et glabrous, its var. villosa excluded), with 2 (white)
L. Iieyneana. —
L. triangulata RoxB. Hort. Beng. gibbosities, dorsally split to the base, dorsal
(1814) 16, nomen nudum. —
L. stipiilciris Roth, in lobes connate with the lateral ones for 3-4 1/2 mm,
R. &
S. Syst. Veg. 5 (1819) 67; Nov. PI. Sp. lateral lobes connate with the ventral one for
(1821) 144; Wall. Pi. As. Rar. 2 (1831) 43.— 3'/2-6 mm; dorsal lobes 1-3 mm, falcate-oblong
Rapuiitium alsinoides Presl, Prod. Mon. Lob. or falcate-ovate, acute or acuminate, margin entire
(1836) 22.— L. sp. Griff. Not. Pi. As. 4 (1854) or subentire, lateral and ventral lobes oblong or
281.— L. griffithii Hook./. & Th. J. Proc. Linn. ovate, l-2'/2 mm, acute or acuminate, entire or
Soc. Bot. 2 (1858) 28; Kurz, J. As. Soc. Beng. subentire. Filaments 3-5 mm, free to half-way up
46, ii (1877) 211, incl. var. genuina et var. dopa- or higher, connate, the two anterior ones ^^ twice
trioides; Clarke, Fl. Br. Ind. 3 (1881) 424; Ridl. as broad as the anthers, the outside patent hairy;
J. Str. Br. R. As. Soc. /;. 33 (1911) 124; Craib, dorsal anthers l'/4, ventral ones 1 mm, each
Kew Bull. (1911) 404; Ridl. Fl. Mai. Pen. 2 anther at the apex with a hair tuft, otherwise
(1923) 201; Danguy, Fl. Gen. L-C. 3 (1930) glabrous or hairy. Ovary l-lVi by l'/2-2'/2 mm,
682, f. 75, 7-9; Craib, Fl. Siam. En. 2 (1936) oblong, trumpet-shaped to cupular, glabrous or
303; WiMMER, Pfl. R. Heft 107 (1953) 569.— L. sparsely hairy; style glabrous. Capsule hemi- —
dopatriuides Kurz, J. As. Soc. Beng. 39, ii (1870) spherical,2-3 by n/2-3 mm. Seeds V2 by 1/3
77; Flora 55 (1872) 302; Craib, Fl. Siam. En. 2 mm, trigonous, brown.
(1936) 303. L. microcarpa Clarke, Fl. Br. Ind. Distr. SE. Asia (Ceylon and the Deccan to
3 (1881) 424; Koord. Exk. Fl. Java 3 (1912) 302; S. China: Kwangtung), Hainan, and Formosa, in
Danguy, Fl. Gen. I.-C. 3 (1930) 681; Merr. & Malaysia: Malaya (Perils: Chupeng; Singapore,
Perry, J. Arn. Arb. 22 (1941) 385; Wimmer, Wimmer, I.e.), W. Java (Indramaju), Celebes
Pfl. R. Het^t 107(1953) 574.— L. terminalis Clarke,
Fl. Br. Ind. 3 (1881) 424; Craib, Kew Bull.
(1904) 404; Danguy, Fl. Gen. I.-C. 3 (1930)
680; Craib, Fl. Siam. En. 2 (1936) 306; Wimmer,
Pfl. R. Heft 107 (1953) 513.—Dortmannia grif-
fithii O.K. Rev. Gen. PI. 2 (1891) 380.— Z)o/7-
mannici trigone/ O.K. I.e., incl. var. microcarpa et
terminalis. — Dortmannia alsinoides O.K. I.e. 972.
L. luzoniensis (Pers.) Merr. En. Philip. 3 (1923)
588; Wimmer, Pfl. R. Heft 107 (1953) 543.—/..
chevalieri Danguy, Bull. Mus. Paris (1929) 263;
Fl. Gen. L-C. 3 (1930) 683, f. 76, 9; Wimmer,
Pfl. R. Heft 107 (1953) 569.— L. thorelii Wimmer,
in Fedde, Rep. 26 (1929) 3,t. 71 f. 3.— L. hosseusii

Wimmer, I.e. 2, t. 71 f. 5; Danguy, Fl. Gen. L-C.

3 (1930) 681; Craib, FI. Siam. En. 2 (1936) 304,
incl. var.;Wimmer, Pfl. R. Heft 107 (1953) 574.—
L. hainanensis Wimmer, Ann. Naturh. Mus. Wien Fig. 12. Distribution of Lobelia alsinoides Lamk
56 (1948) 348; Pfl. R. Heft 107 (1953) 506.— L. with localities in Malaysia dotted.
chinensis (non Lour.) Hance, J. Linn. Soc. Bot.
13 (1873) 110; Forbes &
Hemslfy, J. Linn. Soc. (SW. & SE. peninsula), Philippines (Mindanao,
Bot. 26 (1889) 3; Danguy, Fl. G6n. L-C. 3 (1930) Davao), S. New Guinea (N of Merauke, Wuroi,
680, f. 75, 10-12. L. radicans {non Thunb.) Oriomo R., Lake Daviumbu, Borovia). Fig. 12.
Hosseus, Beih. Bot. Centralbl. 28, ii (191 1 ) 446.— Ecol. In West Java and in SW. Celebes during
Fig. 16c. the wet season in marshy grassland often do-
Erect to ascending, unbranched to strongly minated by 'siir (Sorghum nitidum) and/or
branched herb, 3-35(-40) cm; stem 3-angIed and Cyperaeeae; in S. Celebes also along margins
winged. Leaves 1/2-I V2 by '/2-2 cm, sessile or short- of dry rice-fields. In New Guinea this species is
petioled, contracted or narrowed to the base, found on swampy grounds, together with Erio-
acute or blunt, entire or toothed, glabrous to eaulon, Utricularia, and several Cyperaeeae; on
sparsely hairy; basal leaves ovate-oblong, wet places in savannahs with Melaleuca, Acacia,
cordate or upwards sometimes sublan-
elliptic, Eucalyptus stands and scattered Antidesma
ceolate, up to by 0.3 cm. Flowers 8-12 mm,
1 ghaesemhilla trees. From sea-level up to 1000 m,
axillary, often in the higher axils, and then restricted to regions subject to a dry monsoon.
forming a lax terminal raceme. Pedicels l-3'/2 cm, Nov.-Aug.
Fl. fr.
3-angied, glabrous or slightly patent-hairy. Note. A polymorphic species with intergrading
Bracteoles basal, minute to 2 mm
long and linear, forms, so we can a range o\' subsequent
find
often caducous. Calyx lobes triangular, subulate, 'species', beginning from the typical L. griffithii,
214-314 by V^-l mm, entire, sometimes ciliate. with its scale-like leaves to the normal broad-
128 Flora Malesiana [ser. I, vol. 6^

ovate leaves of L. dopatrioides and the petiolate Gen. PI.

1 (1891) 973. Dortmannia zeylanica
leaves of L. terminalis. I cannot trace any geo- O.K. 380, pro nomen, excl. sclied.
I.e. Dortman- —
graphical or ecological replacement of these nia subcuneata O.K. I.e. 973. Dortmannia
forms. The shape and size of the bracteoles have trigona O.K., pro var. affinis O.K. I.e. 380. L.
also lead to superfluous names: the Indian speci- barbata Warb. Bot. Jahrb. 13 (1891) 444; K.
mens have relatively long bracteoles, which in- ScH. & Lalt. Fl. Schutzgeb. (1900) 59?,.— Pratia
duced Roth to name this form L. stipularis. torricellensis K. Sch. & Laut. Nachtr. (1905)
L. luzoniensis (Pers.) Merr. was based upon a 402. Pratia ovata Elmer, Leafl. Philip. Bot. 2
specimen in herb. Nee from Luzon near Santa (1909)593; Merr. Philip. J. Sc. 1 1916) Bot. 317;
1 (

Cruz de la Laguiia. Merrill mentioned two En. Philip. 3 (1923) 589.— Pratia begonifolia
recent collections from Luzon (Lagufia) and {non Lindl.) Hosseus. Beih. Bot. Centralbl. 28,
Mindanao (Davao), citing Copeland 368 and ii (1911) 477.
Weber 1472 respectively. Wimmer quoted the last Creeping to ascending herb, 20-90 cm, stems
mentioned number under L. alsinoides, Pfl. R. often rooting at the lower nodes, terete, higher up
Heft 107 (1953) 573. I have seen Copeland 368, sometimes angular, glabrous to sparsely hairy.
which was distributed as a Wahlenbergici (NY); Leaves spirally arranged, ovate to broad ovate,
it has all anthers bearded and trigonous se3ds and (%-)l-6 by (V2-)l-3'/2 cm; base cordate to
undoubtedly represents L. alsinoides. This con- truncate or sometimes decurrent, apex acute,
firms my opinion, made from the plate and sometimes blunt, margin subentire to subdentate
description, that L. luzoniensis is conspecific with to repand-dentate; upper surface puberulous or
L. alsinoides. Cavanilles's figure of the plant is glabrous, underneath puberulous, especially the
twice enlarged; he mentions the short seta on nerves, or glabrous; petiole terete, 1-20 mm,
top of the corolla lobes and hairy anthers. sometimes puberulous to appressed-hairy. Flowers
axillary, 7-12 mm. Pedicels terete, 1-2 cm,
4. Lobelia zeylanica Linne, Sp. PI. 2 (1753) 932; ebracteolate. Calyx lobes lanceolate to oblong
OsBECK, Dagbok Ostindisk Resa (1757) 241: triangular, patent-hairy, sometimes gla-
acute,
BURM./. Fl. Ind. (1768) 186; R. «& S. Syst. Veg. 5 brous, 3-5 by V^ mm, entire to subdentate, with
(1819) 64; RoxB. Fl. Ind. 2 (1824) 113; Wall. curved hairs to dentate-ciliate. Corolla 5-9 mm,
PI. As. Rar. 2 (1831) 43; G. Don, Card. Diet. purplish to pale or creamy, inside glabrous to
3 (1834) 709; Kurz. Flora 55 (1872) 302; J. As. subglabrous with two gibbosities, dorsally slit
Soc. Beng. 46, ii (1877) 211, incl. var. affinis; to the base: dorsal lobes connate with the lateral
Yamamoto, Obs. Fl. Form. 13 (1936) 149; Merr. ones for 2-31/2 mm, lateral lobes connate with the
& Perry, J. Arn. Arb. 22 (1941) 386; Green- ventral one for 4-7 mm; dorsal lobes oblong or
wood, ibid. 30 (1949) 78.— L. succulenta Bl. elliptic, 2'/2-3 by 1-1 Vi rnm, falcate, entire to
Bijdr. (1826) 728; DC. Prod. 7 (1839) 373; MiQ. wavy, acute, outside on the nerves with long
Fl. Ind. Bat. 2 (1856) 577; Koord. Exk. Fl. Java hairs, lateral and ventral lobes spathulate to ovate,
3 (1912) 303; Ochse, Trop. Groent. (1925) 22, 1-2 by 1-2 mm. entire, at the centre of the ventral
fig.; Ochse & Bakh. Ind. Groent. (1931) 92, lobe a white spot surrounding a purple stripe.
f. 54; Hochr. Candollea 5 (1932) 292; Merr. Filaments 3-5 mm, for more than % free; free
Pap. Mich. Ac. Sc. 20 (1935) 111; Craib, Fl. parts narrowed towards the base, all of equal
Siam. En. 2 (1936) 305; Wimmer, Ann. Naturh. width, haired; anthers from dorsal to ventral
Mus. Wien 56 (1948) 361, incl. var.; Back. 1-1 1/2 mm to -^4-1 mm, hairy; all anthers at the
Bekn. Fl. Java (em. ed.) 8 (1949) fam. 184, p. 3; apex with a hair tuft. Ovary cupular to obconical,
Wimmer, Pfl. R. Heft 107 (1953) 576, incl. var. 1 V4-3 by 1 mm, scattered hairy to hairy (glabrous
lobbiana et f. glabra. —
L. affinis Wall. [Cat. in specimens from Hainan); style glabrous.
(1828) 35 et 158 no 1131] exG. Don, Gard. Diet. Capsule obconical to oblong ovate, 3-7 by 2-4
3 (1834) 709; non Mirbach, 1805; DC. Prod. 7 mm, with distinct nerves and a membranous
(1839) 360; MiQ. Fl. Ind. Bat. 2 (1856) 574; pericarp. Seeds trigonous, 0.4 by 0.3 mm, light-
Hook./. & Th. J. Proc. Linn. Soc. Bot. 2 (1858) brown; after withering the nerves and remains
27;Drury, Handb. Ind. Fl. 2(1866) 108; Clarke, of the placenta persistent.
Fl. Br. Ind.(1881) 424, incl. var. lobbiana;
3 Distr. SE. Asia (from Ceylon and the Deccan
Trim. Fl. Ceyl. 3 (1895) 57; Gamble, J. As. Soc. through the Himalaya to SE. China: Kwangtung,
Beng. 74, ii (1905) 52; Craib, Kew Bull. (1911) Kwangsi, Chekiang), Hainan, and Formosa, al-
404; Koord. Exk. Fl. Java 3 (1912) 121 ;S. Moore, most throughout Malaysia, but not yet found in
Trans. Linn. Soc. Bot. II, 9 (1916) 88; Diels, the Lesser Sunda Islands and the Moluccas; also
Bot. Jahrb. 55 (1917) 121; Ridl. Fl. Mai. Pen. 2 in the Fiji Is.
(1923) 200, f. 89; Danguy, Fl. Gen. I.-C. 3 Ecol. On shaded grassy grounds and moist
(1930) 679; Merr. Lingn. Sc. J. 6 (1930) 332.— places, under everwet climatic conditions. In
Rapuntium affine Presl, Prod. Mon. Lob. (1836) coffee, tea, and rubber plantations, in open places
13. Rapuntium succulentum Presl, I.e. 13. in primary forest along streams, in shaded ravines,
Rapuntium zeylanicum Presl. I.e. 13. L. sub- etc., from the lowland up to 1500, rarely 1650-
cuneata Miq. Fl. Ind. Bat. 2 (1857) 51A.—L. 2000 m. Fl. fr. Jan.-Dec.
lobbiana Hook. /. &
Th. J. Proc. Linn. Soc. Vern. Belimbing tanah, M, rantji djadjar, J,
Bot. 2 (1858) 28; Drury, Handb. Ind. Fl. 2 djukut bulu mata kerbo, ramo kujali, S.
(1866) 110. —
Dortmannia succulenta O.K. Rev. Use. According to Ochse the young leaves are
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 129

eaten as 'lalab' (steamed vegetable) with rice. two other Osbeck specimens were located in the
Notes. There is in Hterature no unanimous Riksmuseet Stockholm, both marked "ze/7a/7/ca"
opinion about the typification of this Linnean and "7S zeylanica" respectively. The latter speci-
species and this even induced Craib to suggest men has possibly actually been in the hands of
to reject it as a numen cunfusiim (Fl. Siam. En. 2, Linnaeus.
1936, 305-306). De Candolle accepted it as
conspecific with L. ciffinis Wall, which is in turn 5. Lobelia heyniana R. & S. Syst. Veg. 5 (1819)
conspecific with L. succideiitci Bl.; he excluded 50; non Spreng. 1825; Bl. Bijdr. (1826) 728;
from it LiNNAEUs's reference to Seba which he G. Don, Card. Diet. 3 (1834) 709 (lieyneana);
found fit to describe as L. sebae DC, now ac- WiMMER, Ann. Naturh. Mus. Wien 56 (1948) 344;
cepted as a synonym of Muiiopsis simplex (L.) Pfl. R. Heft 107 (1953) 474, incl. var. div.; San-
WiMMER. In this interpretation he was followed TAPAU, Rec. Bot. Surv. Ind. 16 (1953) 158.—L.
later by several others, for instance Kurz, and decurrens Roth, Nov. Sp. (1821) 145; non Cav.
recently Merrill & Perry. 1801.— L. micrantlia Hook. Exot. Fl. 1 (1823)
The other interpretation started with Clarke, t. 44; non H.B.K. 1818.— L. trialata Ham. ex
who, for his revision of the Indian Lobelias, D. Don, Prod. Fl. Nep. (1825) 157; G. Don,
examined the material of L. zeylanica in the Lin- Card. Diet. 3 (1834) 709; DC. Prod. 7 (1839)
nean Herbarium, and stated (Fl. Br. Ind. 3, 1881, 360; Clarke, Fl. Br. Ind. 3 (1881) 425, incl. var.
425):
—
"that Linnaeus's excellent specimen of the lamiifolia; Craib, Kew Bull. (1911) 404; Koord.
species is named L. zeylanica in his own hand, but Exk. Fl. Java 3 (1912) 302; Bold. Zakfl. (1916)
the name has been altered (erroneously) by Sir 41; Gamble, Fl. Pres. Madras 4 (1921) 736;
J. E. Smith to L. anceps, an Australian species." Craib, Fl. Siam. En. 2 (1936) 306. L. subincisa
Craib, the Linnean specimens, Wall. [Cat. (1828) n. 1320, nomen] ex DC.
two sheets:
I.e.,
—re-examined
"on one sheet marked L. zeylanica Prod. 7 (1839) 367; MiQ. Fl. Ind. Bat. 2 (1856)
are 2 specimens but different plants, viz one what 575. Rapiintium reinwardtianum Presl, Prod.
we call now "£. siicculenta" Bl. and a second Mon. Lob. (1836) 14. Rapuntium trialatum
non-campanulaceous plant." ."Pinned to this . . Presl, I.e. 13. Rapuntium arenarioides Presl,
"L. zeylanica'' sheet is another, on which is I.e. n.—L. arenarioides DC. Prod. 7 (1839) 367.—

written, in Linnaeus's hand, "18", i.e. the number L. reinwardtiana DC. I.e. }i()l MiQ. Fi. Ind. Bat.
;

of L. zeylanica in Sp. PI. The plant on this sheet 2 (1856) 565; Koord. Exk. Fl. Java 3 (1912) 303.—
belongs to L. zeylanica as usually understood L. inconspicua Rich. Tent. Fl. Abyss. 2 (1851) 8.
to-day." L. zeylanica (non L.) Moon, Cat. (1824) 14, nomen;
This latter is obviously the specimen which Clarke, Fl. Br. Ind. 3 (1881) 425, incl. var.
Clarke designated as the lectotype. It is referred walkeri; Trim. Fl. Ceyl. 3 (1895) 56; Koord.
to by Savage (Cat. Linn. Herb. 1945, 165) under Exk. Fl. Java 3 (1912) 302; Bold. Zakfl. (1916)
1015 as no 42. According to Savage it is not 41; Haines, Bot. Bihar & Orissa 4 (1922) 501,
indicated on any of the Linnean sheets which incl. var. aligera; Back. & Sloot. Theeonkr.
came from China and was collected by Osbeck. (1924) 213, fig.; Danguy, Fl. Gen. I.-C. 3 (1930)
Merrill & Perry rightly considered that, 678, incl. var. parviflora; Hochr. Candollea 5
though Linnaeus derived his specific name from (1932) 292; Back. Bekn. Fl. Java (em. ed.) 8
Campanula ceylanica, senecionis folio, flore (1949) fam. 184, p. 2. L. subracemosa Miq. Fi.
piirpiireo Sera,' Thes. I: 37, t. 22, f. 12. 1734, Ind. Bat. 2 (1857) 576, incl. var. rigidior.—L.
yet, at the same added a question mark
time, he dichotoma Miq. I.e. 576; Wimmer, in Fedde,
to this reference; his description was based wholly Rep. 38 (1935) 78; Ann. Naturh. Mus. Wien 56
on a plant collected by Osbeck, near Canton, (1948) 345, incl. var. aligera et var. pilosella;
in China, which represents a species totally dif- Pfl. R. Heft 107 (1953) 476, incl. var.—L. trigona
ferent from the form Seba illustrated (J. Arn. {non RoxB.) Hook. /. &
Th. J. Proc. Linn. Soc.
Arb. 22, 1941, 386). They concluded that the Bot. 2 (1858) 27, p.p.; Thwaites, En. PI. Zeyl.
Osbeck specimen(s) are unquestionably the type (I860) 169, p.p.; Benth. Fl. Hongk. (1860) 197,
of L. zeylanica. p.p.; HossEUS, Beih. Bot. Centralbl. 28, ii (1911)
In the rather detailed description Osbeck gave 466, p.p. —Dortmannia zeylanica O.K. Rev. Gen.
the following characters: "Perianthiiim subtus . . . PI. 2 (1891) iSO.— Dortmannia trialata O.K. I.e.
villosum filamenta
. . . duo basi villosa
. . . Cau- . . . 973. Dortmannia inconspicua O.K. I.e. 972.
lis teres Folia
. .cordata
. petiolata."' This
. . . Dortmannia reinwardtiana O.K. I.e. 973. L.
definition leaves no doubt that Osbeck's plant aligera Haines, J. As. Soc. Beng. n.s. 15 (1920)
must agree with L. succulenta Bl. 316.— L. bialata Merr. Philip. J. Sc. 7 (1912)
This is corroborated by an Osbeck collection Bot. 105, ex descr.; Wimmer, Pfl. R. Heft 107
in the Bergius Herbarium, which we could examine (1953) 474.— Fig. 16d.
through the courtesy of Prof. Florin, Stockholm. An ascending or erect herb, 5-50(-60) cm; stem
On the back is written: ^''ceylanica. Lobelia 3-angled and winged. Leaves spirally arranged,
foliis ovatis obtusis peiiolatis crenatis caule dif- the lower ones rhomboid to broad-ovate, upwards
fuse: In China agris aquosii oryzae legi 1751 elliptic to linear-lanceolate, decurrent at the base,
Osbeck." The specimen doubtless represents L. acute at the apex. Vj-^ by '4-3 cm, serrate to
succulenta Bl. serrate-dentate, glabrous or sparsely haiiy,
Through the diligence of Dr Norlindh, Lund, especially the higher ones; petiole up to 3 mm.
130 Flora Malesiana [ser. I, vol. 6^

Prod. 7 (1839) 360; Forbes &

Hemsley, J. Linn.
Soc. Bot. 26 (1889) 2; Diels, Bot. Jahrb. 21
(1901) 607; Merr. Lingn. Sc. J. 5 (1927) 181;
Craib, Fl. Siam. En. 2 (1936) 302; Yamamoto,
Obs. Fl. Form. 13 (1936) 148; Wimmer, Pfl. R.
Heft 107 (1953) 609.—L. erimis (non L.) Thunb.
Fl. Jap. (1784) 325. L. erinoides {non L.) Thunb.
I.e. 325. L. radicans Thunb. Trans. Linn. Soc.
2 (1794) 330; R. & S. Syst. Veg. 5 (1819) 60;
RoxB. Fl. Ind. 2 (1824) 110; Clarke, Fl. Br. Ind.
3 (1881) 425; Forbes & Hemsley, J. Linn. Soc.
Bot. 26 (1889) 3; Koord. Exk. Fl. Java 3 (1912)
302; Bold. Zakfl. (1916) 41; Koord. Fl. Tjib. 3
(1918) 55; Back. & Sloot. Theeonkr. (1924)
212, fig.; Alston, Fl. Ceyl. Suppl. (1931) 175;
Merr. Lingn. Sc. J. 7 (1931) 325; Back. Bekn.
& Fl. Java (em. ed.) 8 (1949) fam. 184, p. 2; T.
Fig. 13. Distribution of Lobelia heyniana R. S.
Makino, 111. Fl. Japan (1954) 80. L. campanu-
Flowers 4'/2-12 mm, axillary, often forming a loides Thunb. Trans. Linn. Soc. 2 (1794) 331;
terminal raceme. Pedicels V2-2 cm, 3-angled, Ker, Bot. Reg. 9 (1823) t. 733; G. Don, Gard.
patent, glabrous or hairy with 2(-l), small, linear Diet. 3 (1834) 709.—/.. caespitosa Bl. Bijdr.
to minutely reduced bracteoles. Calyx lobes (1826) 729; DC. Prod. 7 (1839)366; MiQ. Fl. Ind.
lanceolate-elliptic, 2-4 by 14-1 mm, acute, entire Bat. 2 (1856) 575. Rapuntium caespitosiim
to subdentate, glabrous to sparsely hairy. Corolla Presl, Prod. Mon. Lob. (1836) 13. Rapuntium
31/^-10 mm, pale-purple, lilac, sometimes white, campanuloides Presl, I.e. 13. Rapuntium chinense
inside with 2 gibbosities and a dark spot, nearly Presl, I.e. 13. Rapuntium radicans Presl, I.e.
glabrous to hairy, slit to the base; dorsal lobes \A.—Isolobus radicans DC. Prod. 7 (1839) 353.—
connate with the lateral ones for n/2-5 mm, Isolobuskerii DC. I.e. 353. Isolobus roxburglnanus
lateral lobes connate with the ventral one for 2-7 DC. I.e. 353; Hook./. & Th. J. Proc. Linn. Soc.
mm; dorsal lobes linear to lanceolate, 1-4 by Bot. 2 (1858) 27. Isolobus eampanuloides DC.
1/^-1 mm, often reflexed, sometimes longitudinally Prod. 7 (1839) 353. Isolobus caespitosus Hassk.
folded, small compared with the others, lateral
and ventral lobes spathulate to ovate, 1-7 by
'/4-2V2 mil' acute or blunt, entire to subcrenate.
Filaments 3-7 mm, up to half-way free, of equal
width, glabrous or hairy. Anthers from dorsal
to ventral 1-1 '/i to '/2-3/4 mm, glabrous or hairy,
only the 2 anterior ones with an apical hair tuft.
Ovary trumpet- to cup-shaped, 1 '/2-4 by l-2'/2
mm, glabrous, sometimes hairy; style glabrous.
Capsule obconical to obconical-campanulate,
3-8 by 2-2'/2 mm. Seeds semi-ellipsoid, '/2 by
'4 mm, smooth, brown.
Distr. E. Africa (Erytrea, Abyssinia, Kenya
to Tanganyika), SE. Asia (Ceylon and the Deccan
Peninsula to S. China), Malaysia North to Central
:

Sumatra, Java (from Mt Patuha eastward), Lesser

Sunda Islands (Bali: G. Agung; Lombok: G.
Pusuk, G. Sembalung; Sumbawa: Batu Sulang;
Timor: Noiltoko, G. Mutis), Philippines (Luzon:
Bontoc, Vanoverbergh 902, type of L. bialala
Merr., non vidi), and W. New Guinea (Merauke).
Fig. 13.
Ecol. In moist and swampy places, in marshes,
on steep slopes and in light forest, 500-2700 m.
Fl. fr. Jan. -Dec.
Vern. Djukut mata kebo, kitombe, S, kremo,
krinjingan, kiikiinnang, J.

Note. Small specimens possess often lax

racemes and leaves as in some specimens of L.
alsinoides.

6. Lobelia chinensis Lour. Fl. Coch. (1790) 514, Fig. 14. Lobelia chinensis Lour. Bogor (West
ed. Willd. (1793) 628; R.& S. Syst. Veg. 5 (1819) Java), in dense sods in the Tji Liwung valley
41; G. Don, Gard. Diet. 3 (1834) 709; DC. (1928).
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 131

Bonpl. 7 (1859) ]SO.—Praria radicans G. Don,

Card. Diet. 3 (1834) 100.— Pratici thimbergii G.
Don, I.e. 700. Dortmannia chinensis O.K. Rev.
Gen. PI. 2 (1891) 380. Dortmannia campanuluides
O.K. I.e. 380. Durtmannia radicans O.K. I.e. —
Fig. 14.
Aglabrous, branched, caespitose or prostrate
rooting herb; stem 5-15 cm, terete, with two
longitudinal ridges. Leaves alternate (distichous)
with decurrent sessile or subsessile base, elliptic-
ovate or lanceolate (especially apically), 13-33
by 2-6 mm; acute or blunt, subentire to shallowiy
toothed towards the apex. Flowers axillary, 7-15
mm, on one stem mostly 1-2, only one opened
at a time. Pedicels terete, 6-37 mm, erect, ebrac-
teolate. Calyx lobes narrow-triangular, 1 '/2-3 by
'/2-% mm, dentate at the base. Corolla 5-12 mm,
white to pale-purple (red in China), outside
glabrous, inside hairy, sometimes with 2 green
gibbosities, slit to the base; dorsal lobes connate
with the lateral ones for 4-4 '/2 mm, lateral lobes
connate with the ventral one for 4'/2-6 mm;
dorsal lobes 4 '/^-8 mm, linear, sometimes, reflexed,
blunt, lateral and ventral lobes 3-6 mm, equal,
linear to lanceolate, Filaments 5-6 mm,
acute. Fig. 15. Lobelia archboldiana (Merr. & Perry)
for more than half-way free, all of equal width,
Moeliono. a. Habit, x 2V2> b. young flower,
the 2 anterior ones patent hairy, with a seta or "'
3, c. open flower, x 3 (Clemens 12442).
hair tuft at the top; anthers from dorsal to ventral
2-1 mm, glabrous or hairy. Ovary 2-5 by mm, 1

trumpet-shaped, glabrous; style hairy at the base. obtuse at the apex, very thin, glabrous or sparse-
ly hairy above, glabrous underneath; petiole
Capsule on a recurved pedicel, 4-6 mm, obconical,
glabrous. Seeds 3/8-1/2 mm, ellipsoid, somewhat 3-7(-23) mm, glabrous or sparsely pilose. Flowers
5-6I/2 iTim, axillary or terminal. Pedicels 2-12 mm,
compressed, dark-brown, smooth.
Distr. SE. to E. Asia (from the Deccan to glabrous or hairy. Calyx lobes linear-lanceolate,
China and Japan), in Malaysia: Malay Peninsula with 1-2 pairs of teeth, hairy, 2-3 mm. Corolla
(Singapore) and Java (eastwards to Mt Dieng), red to dark wine-red, the tube 2-3 mmlong,
according to Backer & van Slooten since long outside glabrous to sparsely hairy, inside glabrous,
introduced from continental Asia. dorsally split to1 mm from the base; dorsal lobes
Ecol. On moist, grassy places, along water- half-way connate with the lateral ones, their free
courses and on cultivated land, on rice-fields part lanceolate, 2-3 mm, reflexed, lateral and
and and cinchona plantations, 500-1600 m,
in tea ventral lobes also half-way connate, their free
occasionally at lower altitudes. According to part lanceolate, 3-3.2 mm, acute. Stamens c.
Backer & van Slooten no capsules with ripe 2 mm, filaments for Y^ of their length free, gla-

seeds are found in Java. Van Steenis has found brous; anthers from dorsal to ventral l'/4-% mm;
a plant at Bogor (/;. 2151) with ripe capsules and 2 anterior ones with a short bristle and some hairs.

seeds. I also saw a sheet in the Leyden Herb. Ovary campanulate to trumpet-shaped, l-2'/2 by
(Schiffner 2728, from Tjibodas) with ripe capsules 1-2 mm, sparsely hairy; style glabrous. Capsule
and seeds. It remains questionable whether these roundish to ovoid, 3 by 3 mm, sparsely hairy,
seeds are viable. with a thin wall. Seeds roundish to ovoid, c. V2
Propagation normally vegetative. According
is
by V2 mm,
smooth, dark-brown to black.
to Backer & van
Slooten it descends along Malaysia: E. New Guinea (Murray
Distr.
water-courses to lower altitudes, even to the Pass, Wharton Range; Rawlinson Range, Morobe
lowlands, by stolons or stems dispersed by water. Prov.), rare, twice collected.
However, such lowland habitats are only tem- Ecol. Under a rock wall on grassy bank of a
porary. creek and in deep holes above deep water-courses,
Vern. Djukiit mata keiijeiip, hi tomhe, S. 2840-3600 m. Fl. fr. July-Aug.

7. Lobelia archboldiana (Merr. & Perry) Moe- 8. Lobelia brachyantha Merr. & Perry, J. Arn.
liono, nov. comb. —
Pratia archboldiana Merr. Arb. 22 (1941) 385; Wimmer, Pfl.R. Heft 107
& Perry, J. Arn. Arb. 30 (1949) 59; Wimmer, (1953) 487.— Fig. 16a-b.
Pfl. R. Heft 107 (1953) 765.— Fig. 15. Tiny, creeping, branched herb; stem terete,
Stemless, delicate plant, sometimes with stolons, rooting, sparsely pilose. Leaves a\\.err\aXQ, orbicular-
2-4 cm. Leaves in rosette, ovate to roundish, reniform, 2V2-5 by 2 ''2-6 mm. base cordate to
'/2-I V2 cm diam., subcordate or truncate at the truncate, margin subundulate to dentate, upper
base, wavy to denticulate along the margin. surface sparsely pilose, beneath glabrous; petiole
132 Flora Malesiana [ser. I, vol. 6^

(1881) 423; Koord. Exk. Fl. Java 3 (1912) 304;

Fl. Tjib. 3, 2 (1918) 56; Danguy, Fl. Gen. I.-C.
3 (1930) 677, f. 75, 1-5; Hochr. Candollea 5
(1932) 293, incl. f. variegata; Hend. Gard. Bull.
S.S. 7 (1933) 109; docters van Leeuwen, Verb.
Kon. Ak. Wet. A'dam sect. 2, 31 (1933) 240;
Steen. Bull. Jard. Bot. Btzg 111, 13 (1934) 179;
Merr. Contr. Arn. Arb. 8 (1934) 165; Wimmer,
Pfl. R. Heft 106 (1943) 116, incl. f. variegata et
var. cyanocarpa; Back. Bekn. Fl. Java (em. ed.)
8 (1949) fam. 184, p. 5. Piddingtonia montana
MiQ. Fl. Ind. Bat. 2 (1857) 573; Buijsman, Flora
—
106 (1913) 127, excl. syn. Piddingtonia patens
MiQ. Fl. Ind. Bat. 2 (1857) 573.—Speirema
montaniim Hook. /. &Th. J. Proc. Linn. Soc.
Bot. 2 (1858) 27; Clarke, J. Linn. Soc. Bot. 15
(1876) 147. Piddingtonia cyanocarpa Hassk.
Bonpl. 7 (1859) 179.— Fig. 17.
Erect, mostly branched, medium-sized to
coarse, nearly glabrous herb, c. 1-2 m; stem terete.
Leaves spirally arranged, ovate-oblong to lanceo-
late, 3'/2-12 by 1 V2-4V4 cm, decurrent at the base,
dentate to subdentate, acute to acuminate; petiole
c. 1/2 cm, glabrous to appressed-hairy. Flowers
axillary, solitary, secund, scentless. Pedicel 2-4V2
cm. during anthesis erect. Calyx lobes lanceolate,
Fig. Lobelia brachyantha Merr. & Perry.
16.
5-1 1 by 1-1 1/2 mm, obliquely patent, entire, acute.
a. Habit,x 2V2» b. flower, / 5. L. alsinoides
Corolla V2-3 cm, purplish to lilac-blue with
Lamk. c. Stamens, x 5. L. heyniana R. & S. 1

d. Stamens, x 5 (a-b Brass 11570, c van Steenis

6705, d DE VooGD 2615).

1-3 mm. Flowers c. 5 mm, axillary. Pedicels

terete, 4-5 mm, hairy, ebracteolate. Calyx lobes
oblong-triangular, 2-2 '/2 by 1/2—/4 mm, with a
distinct tooth at the base, sparsely pilose. Corolla
subcampanulate, 4 mmlong, dark purplish-red,
outside pilose, inside the 3 anterior lobes papillose,
dorsally not entirely split to the base; dorsal
lobes connate with the lateral ones for 1 mm,
lateral lobes connate with the ventral one for 2 mm
all lobes ovate-deltoid, 1 1/2 mm, acute to blunt.
Filaments 1 Vj mm, up to 1/3 free, equal, glabrous
or hairy. Anthers from dorsal to ventral 1-% mm,
glandular hairy; the 2 anteriors at the apex finely
hairy (according to Merrill & Perry 'setigeris').
Ovary cupular, by 1 V2 mm, long-hairy; style
I

glabrous. Capsule cupular-ovoid, 3 by 2 mm.

Seeds ellipsoid to roundish, 0.6-0.8 mm, smooth,
light-brown.
Distr. Malaysia: New Guinea (Bele River, near
Habbema Lake), once found.
Ecol. Creeping on bare rock on a sparsely
vegetated limestone precipice, 2350 m.
Note. The pilose indument of this plant is
very typical, because the hairs are more-celled.
The structure of the corolla is also very unlike
that of the other Malaysian species by its very
short tube which gives the corolla a campanulate
shape.

9. Lobelia montana Reinwardt ex Bl. Bijdr.

(1826) 728; DC. Prod. 7 (1839) 3S6.—Praria Fig. 17. Lobelia montana Reinw. ex Bl. a. Apex
montana Hassk. Flora 25, 2 (1842) Beibl. 1, p. 23; of flowering stem, X 2/3, b. ripe fruit, natural
Cat. Hort. Bog. (1844) 106; Clarke, F1. Br. Ind. 3 poise, X 2/3.
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 133

white-shaded margins, outside glabrous, inside J.Linn. Soc. Bot. 42 (1914) 100; Merr. En. Born.
hairy and with two pilose gibbosities, dorsally (1921) 585; Wimmer, Pfl. R. Heft 106 (1943)
slit to 1-2 mm
from the base; dorsal lobes connate 117; ibid. Heft 107 (1953) 765; Lam, Blumea 5
for 1-3 mm, connate with the lateral lobes for (1945) 568.
7-10 mm, lateral lobes connate with the ventral Coarse, erect half-shrub, c. 1-1 V2 m, branching,
one for 9-12 mm; dorsal lobes linear-lanceolate, sometimes reclining; stem glabrous, slightly to
9-10 mm, fiat or crisped, margins glabrous, lateral strongly angular and furrowed. Leaves spirally
and ventral lobes oblong-lanceolate, the ventral arranged, elliptic-lanceolate, 9-1 by 2^4-3 1/2 cm,
1

one broadest, 7-12 mm, crisped and ciliate, slightly dentate, decurrent at the base, acute to
acuminate. Filaments 4-7 mm, for 1/3 of their acuminate at the top, glabrous to puberulous,
length free, glabrous; anthers from dorsal to darker green at the upper side, paler beneath,
ventral 5-3 mm
long, hairy; the two anteriors glossy on both surfaces; petiole 3-7 mm. Peduncle
besides at their apex with a hair tuft. Ovary angular and furrowed, inflorescence a terminal
campanulate to hemispherical, 3-5 by 3-61/2 mm, raceme or pseudo-axillary by sympodial growth,
glabrous; style glabrous. Berry globose, V^-lVz erect, faintly puberulous. Bracts lanceolate, 6-7
cm diam. (living c. l-lVi cm), on long (over 6 cm), by 1-2 mm, slightly dentate. Pedicels 5-10 mm,
patent, afterwards recurved pedicels, violet, later thin, whether or not with 1 or 2 tiny lanceolate
dark purple. Seeds broad-ellipsoid to ovoid, c. bracteoles in the basal half. Flowers c. 2V2 cm.
'/2mm long, brown, smooth. Crt/r.v lobes lanceolate, 6 '/2-7 by 1 V2 mm, longitudi-
Distr. India (from the Deccan to the Himalaya) nally nerved, subentire, obtuse to acute, faintly
to Indo-China (Tonkin), S. China (Yunnan), and puberulous. Corolla 1^4-2 '4 cm, blue purple or
Malaysia: Malay Peninsula (Cameroon High- white with purple at the base of the margin of
lands), Sumatra, and Java (eastwards to Mt the segments or lavender tinged, inside with 2
Dieng.) Fig. 18. gibbosities whether or not papillose; dorsally
split to 0-2 mm
from the base; dorsal lobes
connate with the lateral ones for 2-4 mm, lateral
ones connate with the ventral one for 7-11 mm;
dorsal lobes oblong-lanceolate, 6-10 mm, lon-
gitudinally folded, recurved, the margins crisp to
flat, obtuse to subacute, glabrous or slightly
ciliate, lateral lobes oblong-lanceolate, 6-10 by
2 mm, recurved, ventral lobe oblong-lanceolate,
8-12 by 2-2 '/2 mm
whether or not incurved.
Filaments up to half their length free at the base,
c. 5 mm, glabrous; anthers from dorsal to ventral
4-2 mm, at the apex a tuft of hairs. Ovary 5 by
5 mm, campanulate to hemispherical, faintly
puberulous; style glabrous. Berry 3-5 (ma- mm
Seeds Y^-l mm, globose to
ture?), globose, green.
broad ellipsoid, slightly angular, reticulate.
Distr. Malaysia: Borneo (N. Borneo: Mt
Kinabalu; E. Kutei), Celebes (Central part and
SW. peninsula). Fig. 18.
Fig. 18. Distribution of Lobelia montana Reinw. Eco In secondary jungle, on
I. forest edges, along
ex Bl. (continuous line, localities in Malaysia forest paths, and in open grassfields, (5(H)-)900-
dotted), and Lobelia borneensis (Hemsl.) Moe- 1800 m. Fl. fr. Jan.-Dec.
liono (dotted line, localities in Malaysia indicated Vern. Akar maga pawang, N. Borneo.
by crosses). Note. The nearest alliesofthisand the foregoing
species are obviously found in remote places,
Ecol. Open or half-shaded places in mountain China, New Zealand, and Guatemala!
forest, light spots in mossy forests, 1400-2600 m.
According to Docters van Leeuwen (1933) the 11. Lobelia angulata Forst. Fl. Ins. Austr. Prod.
species is autogamous in Java. (1786) 58, //. 309; R. & S. Syst. Veg. 5 (1819) 65;
Vern. Ki bewo, ki leuntja, tetera, S, kemalun, A. Rich. Fl. Nouv.-Zel. (1832) 227; G. Don,
tjelengan, J. Gard. Diet. 3 (1834) 713; DC. Prod. 7 (1839)
Note. KuRZ recorded to have found this 366. L. nummularia Lamk, Diet. Bot. 3 (1789)
specieson the summit of Mt Menumbing in 589; R. & S. Syst. Veg. 5 (1819) 64; Bl. Bijdr.
Banka I. (Nat. Tijd. Ned. Ind. 27, 1864, 206); (1826) 727; G. Don, Gard. Diet. 3 (1834) 709.—
though have not seen
I his collection this record L. begonifolia Wall. Asiat. Res. 13 (1820) 377;
must rest on an error. in Roxb. Fl. Ind. 2 (1824) 115; D. Don, Prod.
Fl. Nep. (1825) 158; Wall. PI. As. Rar. 2 (1831)
10. Lobelia borneensis (Hemsl.) Moeliono, nov. 43. L. javanica Thunb. Fl. Jav. (1825) 9, cf.
comb. — Pratia borneensis Hemsl. in Hook. Ic. Blumea 6 (1950) 360.— L. obliqiia Ham. ex D.
PI. 16 (1886) t. 1532; Staff, Trans. Linn. Soc. Don, Prod. Fl. Nep. (1825) 158.— L. pratiam
Bot. II, 4 (1894) 188, incl. var. grandiflora; Gibbs, Gaudich. Ann. Sc. Nat. Bot. 5 (1825) 103.—
134 Flora Malesiana [ser. I, vol. 6^

Pratia repens Gaudich. I.e. 103; in Freyc. Voy.

Bot. (1826) 456, t. 79; G. Don, Card. Diet. 3
(1834) 340, incl. var.; Hook. /. Fl. Antarct. 1
(1844) 42; Reiche, Fl. Chil. 5 (1910) 63.—L.
rugulosa Graham, Edinb. N. Phil. J. (1829)
\S6.—Fratia begonifolia Lindl. Bot. Reg. (1830)
t. 1373 G. Don, Gard. Diet. 3 (1 834) 699 (begoniae-
;

folia); Presl, Prod. Men. Lob. (1836) 46; Clarke,

Fl. Br. Ind. 3 (1881) 442; Ridl. J. Str. Br. R. As.
Soc. n. 33 (1900) 102; Merr. Philip. J. Sc. 1
(1906) Suppl. 241; Burkill &
Holttum, Gard.
Bull. S. S. 3 (1923) 56 {begouiifolia); Ridl. Fl.
Mai. Pen. 2 (1923) 201; Danguy, Fl. Gen. I.-C.
3 (1930) 674; Craib, Fl. Siam. En. 3 (1936) 302.—
L. reniformis Cham. Linnaea 8 (1833) 210; DC.
Prod. 7 (1839) 365. L. hederacea Cham. Linnaea
8 (1833) 2\l.~Pratia hederacea G. Don, Gard.
Diet. 3 (1834) 699; Hook. /. & Arn. J. Bot. 1
(1834) 277; Presl, Prod. Mon. Lob. (1836) 46;
DC. Prod. 7 (1839) 340, incl. var. elliptica; Hook.
/. Fl. Antarct. 1 (1884) 43; Kanitz, in Mart. Fl.
Bras. 6, 4 (1878) 135, t. 40 f. 1; Wimmer, Rev.
Sudamer. Bot. 2 (1935) 96; Pfl. R. Heft 106
(1943) 114. Pratia serpyllacea Presl, Prod.
Mon. Lob. (1836) 46. Rapuiuium angiilatum
Presl, I.e. 30. Rapuntium reniforme Presl, I.e.
15. Rapuntium nummularium Presl, I.e. 30.
Piddingtonia nummularia DC. Prod. 7 (1839) 341;
MiQ. Fl. Ind. Bat. 2 (1857) 572; Hook./. & Th.
J. Proc. Linn. Soc. Bot. 2 (1858) 26; Hassk.
Bonpl. 7 (1859) 180; Benth. FI. Hong. (1861)
196. L. littoralis R. Cunn. ex A. Cunn. Ann.
Nat. Hist. I, 2 (1839) 50; Regel, Gartenfl. 38
(1888) 662, f. \A%.— Pratia zevlanica Hassk.
Flora 25, 2 (1842) Beibl. 1, p. 23; Cat. Hort.
Bog. (1844) 106. Pratia arenaria HooK. /. Fl.
Antarct. 1 (1844) 106, t. 29 //; icon, errore Pratia
arenosa; Fl. Nov. Zel. 1 (1853) 157.— Pratia
angulata Hook./. Fl. Antarct. 1 (1844) 43; Fl.
Nov. 1 (1853) 157, incl. var. arenaria: Handb.
Zel.
Fl. Zeal. (1867) 172; Cheeseman, Man. New
New
Zeal. Fl. (1906) 397; 111. New Zeal. Fl. (1908) t.
16; Wimmer, Pfl. R. Heft 106 (1943) 110, incl. Fig. 19. Lobelia angulata Forst. Mt Patuha, West
var. —Piddingtonia palliardii Lehm. Hamb. Gar- Java (de Voogd). •
tenz. 8 (1851) 337; Linnaea 25 (1852) 310; Walp.
Ann. 5 391.— L. rotundifolia Banks &
(1858) (1910) 99, 50; Wimmer, Pfl.R. Heft 107 (1953)
Sol. ex Hook./ Fl. Nov. Zel. 1 (1853) 158.— 483, inel.
f.

var. brevipilis. —Pratia papuana S.

L. horsfieldiana MiQ. Fl. Ind. Bat. 2 (1857) 577.— Moore, Trans. Linn. Soc. 9 (1916) 88; Diels,
II,

Pratia nummularia A. Br. & AscHERS. Index Bot. Jahrb. 55 (1917) 125; ibid. 62 (1929) 493;
Sem. Hort. Berol. (1861) Append. 6; Kurz, J. Wimmer, Pfl. R. Heft 106 (1943) Pratia UA.—
As. Soc. Beng. 46, ii (1877) 210; Koord. Exk. wollastonii S. Moore, Trans. Linn. Soc. II, 9
Fl. Java 3 (1912) 303; Bold. Zakfl. (1916) 41; (1916)89; Wimmer, Pfl. R. Heft 106(1943) 111.—
KooRD. Fl. Tjib. 3, 2 (1918) 55; Heyne, Nutt. IPratia podenzanae S. Moore, J. Bot. 55 (1917)
PI. 2 (1927) 1428; Merr. Lingn. Sc. J.
1 (1927) 306; Wimmer, Pfl. R. Heft 106 (1943) 111.—L.
Yamamoto, Obs. Fl. Form. 13
181; (1936) 150; arfakensis Gibbs, Arfak (1917) 183; Kanehira &
Wimmer, Pfl. R. Heft 106 (1943) 112; Lam, Hatusima, Bot. Mag. Tokyo 57 (1943) 128;
Blumea 5 (1945) 569; Back. Bekn. Fl. Java (em. Wimmer, Pfl. R. Heft \Q)1 {\957,) Ai7<.—L. paradoxa
ed.) 8 (1949) fam. 184, p. 3; Blumea 6 (1950) 360. Wimmer, in Fedde, Rep. 26 (1929) 2; Pfl. R. Heft
— Pratia Unnaeoides Hook. / Handb. New Zeal. 107 (1953) 483.— Fig. 19-20.
Fl. (1867) 172. Pratia reniformis Kanitz, in Polymorphous, creeping and branching herb;
Mart. Fl. Bras. 6, 4 (1878) 136, t. 40 f. 2; Wimmer, stem terete, up to 1/2 m long, rooting at the
Rev. Sudamer. Bot. 2 (1935) 96; Pfl. R. Heft 106 nodes, glabrous or hairy. Leaves alternate, round
—
(1943) 115. Z,.///;«fleo/We5PETRiE,Trans. New Zeal. to ovate, broad-ovate or reniform, 2-25 by 2-35
Inst. 23 (1890) 405; Cheeseman, Man. Fl. New mm, cordate, truncate, or even decurrent at the
Zeal. (1906) 400; Petrie, Gard. Chron. Ill, 47 base, crenate, dentate or subdentate to subentire
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 135

at the margin, acute or rounded at the top, ubrigens so geringfiigig, dass nach genauem
glabrous to hairy; petiole 0-25 mm, glabrous or Studium der Formenkreis, wahrscheinlich um-
densely hairy. Flowers 6-18, axillary, solitary. fassender Arten angenommen werden miissen."
Pedicels i/2-5(-7) cm, ebracteolate, glabrous or (Bot. Jahrb. 55, 1917, 125; ibid. 62, 1929, 493).
hairy. Calyx lobes linear-lanceolate to triangular,
1-1 1 mm, entire to distinctly dentate with up to 3
teeth at either side, acute or blunt, glabrous to
puberulous. Corolla 4 1/2-15 mm, outside and
inside glabrous or hairy, sometimes with two
gibbosities inside, dorsally split to the base or
nearly so; dorsal lobes connate with the lateral
ones for 1 Vi'^ mm, laterals connate with the
ventral one for 2-12 mm; dorsal lobes lanceolate,
2-14 mm long, falcate and reflexed, lateral and
ventral lobes spathulate, obovate, or lanceolate,
2-12 mm, the ventral one generally broadest.
Various fruits of Lobelia angulata Forst.
Fig. 20.
Filaments 3-11 mm, very variable, free for Ya
a-c. Fleshy berry to capsule, from Celebes,
of their length, equal in width, sometimes narrowed
d~e. capsular New Guinea specimens, all X 2V2
to the base, or the two anteriors broader, some-
times the two anteriors adnate to the corolla,
{a-c Eyma 1161, d Brass 10622, e 4789).NGF
glabrous or hairy or only the two anterior ones
Also WiMMER himself, in identifying a Philip-
hairy; anthers from dorsal to ventral 1-2 to l-'/2
pine specimen which was astonishingly resembling
mm, glabrous or hairy, only those of the two
Pratia nummularia but possessed fruits with valves
anteriors with a hair tuft or with a seta and some
on top, came obviously reluctantly to the conclu-
hairs. Ovary trumpet-shaped to obconical or
sion that this character settled it as a Lobelia;
ovoid or campanulate, 1-4 by 1-3 mm; glabrous
he named it L. paradoxal
to densely hairy; style glabrous or sparsely pilose.
Beuzenberg & Hair recently found the fol-
Fruit an indehiscent or dehiscent capsule or a
lowing chromosome numbers: for L. linnaeoides
subcarnose to baccate purple fruit, (l-)6'/^-16
(Hook. /.) Petrie n = 7 and for Pratia angulata
by (l-)5-13 mm, glabrous to hairy, ellipsoid to
(FoRST.) Hook./, n = 35 (N. Zeal. J. Sc. 2, 1959,
globose, sometimes flattened at the apex. Seeds
532, 537).
flattened-ellipsoid to ellipsoid, nearly mm, 1

brown, reticulate to fine-reticulate.

12. Lobelia conferta Merr. & Perry, J. Arn.
Distr. Very widely distributed, through SE.
Arb. 59 (1949) 59.— Pratia conferta Wimmer,
Asia (Nepal, Sikkim, Khasia, Burma, Siam, Indo-
Pfl. R. Heft 107 (1953) 764.— Fig. 21.
China) to China (Yunnan, Kwangsi, Hainan,
Tiny, prostrate, fleshy, glabrous herb, with
Formosa), throughout Malaysia, Borneo and the
short stems. Leaves distichous, conferted, semi-
Moluccas excepted, Australia, Tasmania, New
amplexicaulous, sessile, oblong-lanceolate, entire,
Zealand, and adjacent islands, to S. America.
Ecol. Moist open or shaded places, in mountain
4-5 mmby 1-2 mm, blunt to obtuse, smooth and
forest, mossy forest, along river-banks and forest
shining. Flowers 5 mm
long, in the upper leaf
axils. Pedicels 2-5 mm, ebracteolate. Calyx lobes
trails, 600-3300 m.
Vern. Adntingan, ki tomhe, kitonie, kiiweiing,
linear to lanceolate, l'/^-2'/2 mrn by 1 mm, entire,
blunt to slightly Corolla 4-4V2 mm,
acute.
ramo keujeup, djiikut mata keiijeiip giinung, tangkal
glabrous, very light purple, inside with two
siitji, S, ketrus aliis, manikan, serintil, urek urek
gibbosities, dorsally split to the base; dorsal lobes
polo, J; Philippines: gubagitbai, kaiiapa, pisa, Ig.,
tutugi,Bon.; tikiritoka, Kopanko, New Guinea.
Notes. An extremely polymorphous species.
As reported in the note to the description of the
genus, it is in this species that the fruits are
varying not only in shape, but even in structure
in the New Guinean and some Celebes specimens,
which also vary in floral characters. In the abun-
dant material I have studied, I cannot make any
distinct demarcation and I am convinced that it
will be wise to consider the specimens as belonging
to one species. It may be possible that more
numerous, detailed field data will enable to segre-
gate infraspecific taxa.
The polymorphism had already been indicated
by DiELS, who, in his revision of the Papuan
species, remarked:
—
"Diese drei Arten (Pratia
papuana, P. nitmmiilaria, und P. angulata) sowie Fig. 21. Lobelia conferta Merr. & Perry, a.
die P. wollastoiiiivon derNassaukette, undmehrere Habit, X 21/2. b.flower, x 5, anther tube opened
andere bei Pratia diagnostisierte Spezies, sind in front (Brass 4417).
136 Flora Malesiana [ser. I, vol. 6^

connate with the lateral ones for 1 1/2-2 mm; lateral 1-2 14 cm. Native in America, mostly cited
lobes connate with the ventral one for 2-2 '/2 mm; as L. fulgens Willd. L. cordigera Cav.
dorsal lobes lanceolate, Vi-'^Vi mm, reflexed,
sometimes of unequal length; lateral and ventral Excluded
lobes equal, lanceolate to ovate, 1 '/2-2'/2(-3) mm
by 3/4-1 mm, reflexed. Filaments \V2-i mm, half- Lobelia anceps Thunb. Prod. (1794) 40, non
way free, linear, glabrous; anthers from dorsal to Linn./. 1781; Miq. FI. Ind. Bat. 2(1856)578 =
ventral l'/2-l mm, glabrous, the two anterior Lobelia alata Labill. Nov. HoU. PI. 1 (1804) 51,
anthers finely haired at the apex. Ovary trumpet- t. 72; WiMMER, Pfl. R. Heft 107 (1953) 469.
shaped to cupular, '/2-%rnm long. Fruit not seen. MiQUEL, I.e., quoted this species as possibly
Distr. Malaysia: New Guinea (Owen Stanley occurring in the Sunda Islands. According to
Range, SW. slope of Mt Albert Edward), once WiMMER it is an extra-Malaysian species with a
found. distribution in S. Africa, Australia, and Chili.
Ecol. Wet grasslands, 3680 m. Lobelia piimila BuRM. /. Ind. (1768) 186,
Fl.
t. 60, f. 3 (type in G) = Mazus piimilus (Burm.
/.) Steen. Nova Guinea n.s. 9 (1958) 31 {Scro-
KEY TO SOME phulariaceae).
CULTIVATED SPECIES Lobelia tetragoiia Bl. Bijdr. (1826) 729; Hassk.
(in ail spp. both anterior anthers with bearded tip) Cat. Hort. Bog. (1844) 105 =
Lobelia cliffortiana
Linne, Sp. PI. 2 (1753) 931; Miq. Fl. Ind. Bat. 2
1. Corolla tube less than 7 mm
long. Pedicels (1857) 577; Wimmer, Pfl. R. Heft 107 (1953) 526.
glabrous. Leaves petiole-like decurrent. Blume described this species as an endemic
2. Corolla tube 5-7 mm
long. Flowers axillary. from Mt Gedeh, in West Java, but his specimens
Pedicels 1 '/2-5 cm. Calyx tube 2-4 mm, lobes have doubtless been erroneously localized and
3'/2-ll mm. Corolla blue or violet, rarely were derived from specimens cultivated in the
white. Stem triangular. Leaves 1 VS-S by 1/3-1 Botanic Gardens at Bogor or possibly naturalized
cm, oblong lanceolate, entire to serrate. in their vicinity. It is a native of Central America
Annual, native in S. Africa. L. erinus L. and has been introduced as an ornamental plant
2. Corolla tube 3-31/2 mm, inside hairy. Pedicels into several other countries at an early date and
less than 1 V2 cm. Calyx tube 2-3 by 2 mm, seems to have frequently naturalized.
lobes 2-2 '/2 by Y2 mm, acute, entire. Corolla Rapuntium longifolium Presl, Prod. Mon. Lob.
blue or blue-violet. Stem quadrangular, (1836) 26.— L. longifolia DC. Prod. 7 (1839) 382;
fistulose. Leaves broad-ovate, rather coarsely Miq. Fl. Ind. Bat. 2 (1856) 578; F.-Vill. Nov.
sinuate-dentate, IVi-'iVi by 1-2 cm. Native App. 4 (1880) 121; Merr. En. Philip. 3 (1923)
of Central America, early introduced as an 588 = Lobelia graminea Lamk, Diet. Bot. 3 (I79I)
ornamental . . L. cliffortiana L.
. . 583; Wimmer, Pfl. R. Heft 107 (1953) 413.
1. Corolla tube longer than 10 mm. Pedicels Presl based this species on a specimen from
hairy. Leaves sessile, the upper ones longer the Malaspina Expedition collected by Haenke.
than 5 cm. Perennials. According to Merrill it was erroneously localized
3. Flowers all axillary. Pedicels 21/2-8 cm, in Luzon and came really from Central America.
shorter than the sustaining leaf. Sepals erect, Wimmer has examined the type specimen (in
21/2-31/2 mm. Corolla tube bright red, 18-21 Mus. Prague) and has referred it to L. graminea
mm; lobes yellow to orange, 12-14 mm, lower Lamk, which is a native of Panama and Mexico.
lip short-3-lobed, posterior lobes linear. Rapuntium haenkeanum Presl, Prod. Mon.
Leaves ovate-lanceolate, very shallowly den- Lob. (1836) 26.— L. haenkeana DC. Prod. 7
tate, 6-12 by 13/4-3 cm. Erect shrub 1 1/2-21/2 (1839) 382; Miq. Fl. Ind. Bat. 2 (1856) 578;
m. Native in Mexico. L. laxiflora H.B.K. F.-Vill. Nov. App. 4 (1880) 121; Merr. En.
3. Upper flowers in bracteate, terminal racemes. Philip. 3 (1923) 588; Wimmer, Pfl. R. Heft 107
Pedicels shorter than 3 cm. Sepals longer than (1953) 685. Dortmannia haenkeana O.K. Rev.
5 mm. Lower lip over halfway 3-split. Herbs. Gen. PI. 2 (1891 ) 972 = Lobelia nelsonii Fernald,
4. Flowers purple, rarely white. Pedicels 6-8 Proc. Am. Ac. Arts Sc. 36 (1901) 503 = Lobelia
mm. Racemes 5-20 cm, lower bracts folia- laxiflora var. nelsonii McVaugh, North Am. Fl.
ceous. Sepals erect, 8-10 mm, with recurved 32A (1943) 97.
ciliate margins. Corolla tube 12-15 mm, to Presl this species was based upon
According
lower lip 10-12 mm, at the base with 2 white a specimen from Luzon. Merrill, I.e., already
convexities. Leaves oblong, 6-15 by 2-5 cm. suspected that this record was based on a Malaspina
Native in N. America L. syphilitica L. Expedition plant from tropical America er-
4. Flowers dark-red. Pedicels 8-28 mm. roneously localized as Philippine. Wimmer, I.e.,
Racemes 8-30 cm. Bracts narrow. Sepals saw Haenke's type specimen (Mus. Prague) and
recurved, glabrous, 10-15 mm. Corolla tube stated that it is conspecific with ''Lobelia laxiflora
15-20 mm, lobes 20-35 mm
long, posterior var. nelsonii McVaugh" from Guatemala, cor-
ones much narrower than the anteriors. roborating Merrill's statement about its native
Leaves linear-lanceolate, glabrous, 8-20 by country.
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 137

7. PHYLLOCHARIS
DiELS, Bot. Jahrb. 55 (1917) 122; Wimmer, Pfl. R. Heft 107 (1953) 724.—Fig. 22.
Delicate, erect or ascending annual herbs; stems producing roots in the basal
part. Leaves spirally arranged, petioled, often unequal at the base. Flowers solitary,
inserted on the midrib of the leaves. Calyx obliquely cup-shaped, 5-lobed. Corolla
5-lobed, dorsally split to the base, bilabiate, the 2 posterior lobes much prolonged,
narrow, almost free. Stamens 5, connate; anthers curved, unequal; filaments free
at the base. Ovary inferior, 2-celled; style filiform; stigma 2-lobed. Capsule obliquely
obovoid, opening with valves at the top, thin-walled with prominent nerves. Seeds
c\D, ellipsoid, verruculose or smooth.
Distr. In Malaysia: 4 spp., all endemic in New Guinea.
Ecol. Damp, shady, humous or rocky places in the undergrowth of rain-forests, 300-1650 m, ap-
parently rare.
Notes. The genus is only distinct tYom Lobelia in the epiphyllous flowers, with 2 much prolonged
upper lobes. A third difi'erential character mentioned by Diels, viz esetose anther tips, does not hold
for P. siibcordata. Diels assumed that the fruit is finally also loculicidally dehiscent at the base, but I
have seen no trace of such dehiscence.
I have seen only material of P. subcordata and P. saxicola. Of all species only one or very few
numbers have been collected and though the diff'erential characters appear satisfactory it remains to
be seen whether they will prove to be constant.
Sincere thanks are due to Dr Van Royen for putting his MS
revision of the genus at my disposal,
granting permission to publish the new species which he had fully described.

KEY TO THE SPECIES

1. Leaf-base cuneate.
2. Leaves oblong-elliptic, entire, with callose-fimbriate protruding veins. Flowers inserted at about
the middle of the midrib. Calyx glabrous, warty. Posterior corolla lobes at least twice as long
as the anterior ones. Seeds verruculose 1. P. oblongifolia

2. Leaves ovate to ovate-elliptic, dentate. Flowers inserted in the basal quarter of the midrib. Calyx
hairy. Posterior corolla lobes less than twice the anterior ones. Seeds smooth 2. P. schlechteri
.

1. Leaf-base rounded, truncate or subcordate. Seeds verruculose.

3. Leaves narrow-ovate to ovate, almost entirely glabrous, the base tending to be cordate; margin
crenate-serrate. Calyx glabrous 3. P. subcordata
3. Leaves ovate to elliptic, on both surfaces crispy-hairy, the base rounded to truncate; margin with

very small protruding callose vein-tips, almost entire to faintly dentate or shallowly crenate. Calyx
hairy 4. P. saxicola

1. Phyllocharis oblongifolia Diels, Bot. Jahrb. 55 (1917) 124, fig. 1 A-K; Wimmer, Pfl. R. Heft
(1917) 124, fig. 1 L-N; Wimmer, Pfl. R. Heft 107 107 (1953) 725.— Fig. 22h.
(1953) 724, fig. 108 c—
Fig. 22g. Stems 10-20 cm, glabrous below, the upper part
Almost glabrous herb, stems 15-30 cm. Leaves slightly puberulous.Leaves 2-5 by \V2-IV2 cm,
6-11 by 2'4-3 cm, oblong-elliptic, ± acuminate, ovate to ovate-elliptic, acute, base cuneate, very
base cuneate, entire but the veins protruding as sparsely hairy above, puberulous on the nerves
small callose-fimbriate teeth; nerves c. 8 pairs, beneath, margin dentate, ends of the veins cal-
arching; petioles 1-3 '/2 cm. Flowers inserted at lously protruding at the margin; nerves c. 6-7
about the middle of the midrib. Pedicels c. 5 mm. pairs; petioles glabrous, Vi-^Vi cm. Flowers
Calyx tube 1 '/2-2 by 2 mm, lobes lanceolate, inserted in the basal quarter of the midrib,
acute, 2-3 mm, both warty. Corolla tube V2-I Pedicels 5-6 mm, glabrous. Ci//v.v tube c. 1/2 1 mm
mm; posterior lobes 9 by V2 mm, at least twice as long, hairy, 10-ribbed; lobes 4 by lA mm, linear,
long as the anterior ones, narrow-spathulate, acute, with scattered hairs along the margin,
broadened at the base, acute; anterior lobes Co/-o//a tube 1mm; posterior lobes 7-8 by '/2 mm,
31/2-4 by 2-21/2 mm, ovate, acute, connate up to 1/, to Va longer than the anterior ones, linear,
the middle, at the inside near the posterior lobes acute, glabrous; anterior lobes 5-6 by 1 1/2-21/2
somewhat pilose, further glabrous. Androecium mm, oblong-ovate, acute, connate for about
21/2-31/2 mm
long; anthers 1-1 1/2 mm, posterior 2;, of their length, inside papillose. Androecium
ones dorsally ±
hairy. Seeds verruculose. 3 mm long; anthers mm, posterior ones dorsally
1

Distr. Malaysia: NE. New Guinea (Udu: hairy. Capsule 6-7 by 3 mm. Seeds smooth.
Schlechter 17445). Distr. Malaysia: NE. New Guinea (Bismarck
Ecol. Ram-forest, 300 m. Fl.fr. March. ^ts: Schlechter 18620).
2. Phyllocharis schlechteri Diels, Bot. Jahrb. 55 Ecol. In humous rain-forest, 1200 m. F/./r. Nov.
138 Flora Malesiana [ser. I, vol. 6^

Fig. 23. Laurentia longiflora (L.) Petermann. Java (Roepke).

Dec. 1960] Campanulaceae (Moeliono & Tuyn) 139

3. Phyllocharis subcordata Merr. & Perry, J.

Arn. Arb. 22 (1941) 387; Wimmer, Pfl. R. Heft
107 (1953) 725, fig. 108b.— P. lamiifolia Wimmer,
Ann. Naturh. Mus. Wien 56 (1948) 372; Pfl.
R. Heft 106 (1943) 11, fig. 8e.— Fig. 22a-c.
Plant almost glabrous, 5-20 cm. Leaves
1-3'/^ by 1-2 cm, ovate to ovate-oblong, acute,
truncate to cordate at the base, glabrous or
with some scattered hairs along the nerves
beneath, margin dentate; nerves c. 4 pairs;
petioles c. \-\V2 cm. Flowers inserted near the
base of the midrib, yellow (ex coll.). Pedicels
4-6 mm, glabrous. Calyx tube 1-2 mm, glabrous,
lobes 2-4 mm, linear, somewhat obtuse. Corolla
tube 0.5-1 mm, posterior lobes 6-8 mm, linear,
1-1 '/2 rnm wide at the base, twice as long as the
anterior ones, acute, glabrous, anterior lobes 3
by '/2 rnm, ovate, acute, connate at least up to
1

the middle, inside papillose. Androecium 2-3 mm;

anthers 1-1 V2 mm, posterior ones dorsally pilose,
anterior ones with a few setae on top. Capsule
4-4 V^ by 2-3 mm. Seeds about V2 rnm, verru-
culose, showing on the surface of the ripe fruit.
Distr. Malaysia: NE. New Guinea (Morobe
district:Yoangen: Clemens 3426, type; Mt Ako,
Malolo Mission: Clemens 4404A).
Ecol. Along a wet stream bank in mountain
forest, 750-1200 m. Fl. fr. June, Nov.-Dec.

4. Phyllocharis saxicola van Royen, no v. sp. —

Fig. 22d-f.
Caules, folia et calyx pilis crispis albis piibes-
centes. Folia ovata ad elliptica, ohtusiuscula, In-
tegra vel propter venulas ultra marginem in cal-
lositate breviter producta denticulata, basi rotun-
data ad fruncata. Flores in costae parte quarta
inferiore insert!. Semina verruculosa. Tvpus: Carr
14083, L.
Stems 5-12 cm, short crisped-hairy. Leaves
sparsely hairy on both sides, 1 by '/2-
ovate to elliptic, rounded, truncate to subcordate
at the base, margin subentire to very shallowly
sinuate-dentate with ^ remote vein-teeth;
nerves 3-6 pairs; petioles V5-I cm, hairy. Flowers
known to me in bud only, inserted near the base
of the midrib. Pedicels 5-7 mm, whitish hairy.
Calyx tube 1 mm long, whitish hairy, lobes
2-2 '/2 mm, linear, — acute, sparsely whitish
hairy. Posterior corolla lobes green with a crimson
Fig. 22. Phylloc/uiris siibcordata Merr. & Perry. median line, anterior lobes crimson, whitish pilose
a. Habit, x 2/3, b. flower, x 4, c. fruit, X 4. near the apex. Androecium c. 2 mm long; posterior
Phyllocharis saxicola van Royen. d. Habit, x 2/,, anthers with a few white hairs. Capsule 4-6 by
e. fruit, < 4,/. seed x 13. Phyllocharis obloiigi- 2-3 mm, very obliquely oblong-obovoid, 10-
folia DiELS. g. Flower, x 4. Phyllocharis schlech- nerved, pilose. Seeds verruculose, [A mm.
teri DiELS. /;. Flower, x 4 (a-c Clemens 4404, Distr. Malaysia: Papua (Lala River).
d-f Carr 14083, g-h after Diels). Ecol. On rocks in forest, 1650 m. Fl.fr. Dec.

8. LAURENTIA
MiCH. e.x Adans. Fam. 2 (1763) PI. 1 (1838) 511; DC. Prod.
134, 568; Endl. Gen.
7 (1839) 409; McVaugh, 67 (1940) 778; Wimmer, Pfl. R. Heft
Bull. Torr. Bot. CI.
107 (1953) 2S6.—Isotoma Lindl. Bot. Reg. 10 (1824) t. 964; Presl, Prod. Mon.
Lob. (1836) 42; DC. Prod. 7 (1839) 412; Benth. Fl. Austr. 4 (1869) \34.— Hip-
140 Flora Malesiana [ser. I, vol. 6^

pobroma G. Don, Gard. Diet. 3 (1834) 717; McVaugh, Bull. Torr. Bot. CI. 67
(1940) 782; N. Amer. Fl. 32 A, pt 1 (1943) 99.—Enchysia Presl, Prod. Mon. Lob.
(1836) 40.—Fig. 23.
Annual or perennial, laticiferous herbs. Leaves spirally arranged, rarely rosulate
and radical. Flowers in terminal racemes or solitary, axillary. Calyx lobes 5, sometimes
unequal. Corolla tube entire, limb 5-parted, almost regular to slightly zygomorphic
with unequal lobes. Stamens 5; anthers connate, sometimes bearing apical bristles;

filaments free at the base, and sometimes below the from the corolla,
anthers, free
or 2 adnate, or all inserted above the middle of the corolla tube. Ovary inferior,
2-celled; stigma 2-lobed. Capsule opening by 2 loculicidal valves at the top between
the persistent calyx lobes. Seeds co, ellipsoid, brown, up to 1 mm.
Distr. About 25 spp., in the Mediterranean (3), South Africa (18), Australia (10), and the Americas (4).
Notes. It is not unexpected that also in the affinity Lobelia, Laurentia, Isotoma, Palmerella, and
Solenopsis there is no unanimity of opinion about generic distinction, due to different evaluation of
characters. Lobelia is properly only distinguished from the Laurentia assemblage by a distinctly zy-
gomorphous corolla entirely or partially split on one side. McVaugh (Bull. Torr. Bot. CI. 67, 1940,
780) even suggested that this would not hold on the argument that it would not be present in L. sinaloae;
but this seems to be an error, cf. Wimmer, I.e. fig. 91 f. Whereas McVaugh is in favour of merging these
genera into Lobelia, it is curious that he maintained the single American species of Isotoma as representing
a monotypic genus, although it is manifestly allied to its Australian congeners. Gleason on the other
hand (Bull. Torr. Bot. CI. 52, 1925,93) believed that more attention should be paid to the characters
of corolla and anther appendages; this would lead to more than one genus in the assemblage, but would
also have the consequence that Lobelia, where a similar diversity is found, would have to be split up in
smaller segregates. At the present these characters have been used in Lobelia and other genera for the
distinction of infra-generic taxa. It seems satisfactory to follow this principle within the Laurentia
complex, as has been done by Endlicher, Wimmer, and others.
It should be admitted that another solution could be the subordination of Laurentia sens. lat. as a

subgenus under Lobelia, as there is, also in Laurentia, a tendency towards zygomorphism, but this is
characteristic for the entire tribe Lobeliae.
Adanson based his genus on the pre-Linnean Laurentia Mich. Gen. 18, t. 14 = Lobelia laurentia \^.
1753 = Laurentia gasparrinii (Tineo) Strobl.. As he reduced Lobelia L. to Laurentia a formal conser-
vation of Laurentia would seem necessary.

1. Section Isotoma

(R. Br.) PI. 1 (1838) 512; Wimmer, Ann. Naturh. Mus. Wien 56 (1948)
Endl. Gen.
335; R. Heft 107 (1953) 39S.—Lobelia sect. Isotoma R. Br. Prod. (1810) 564;
Pfl.

McVaugh, Bull. Torr. Bot. CI. 67 (1940) 194.— Isotoma Lindl. Bot. Reg. 10 (1824)
t. 964. Hippobroma G. Don, Gard. Diet. 3(1834)717. Laurentia subg. Isotoma
Petermann, Pflanzenreich (1845) 444, t. 118 fig. 665.
Leaves spirally arranged, coarsely sinuate-dentate or pinnatifid. Flowers solitary,
axillary, almost regular. Corolla tube long, slender, lobes spreading, almost equal.
Stamens inserted on the corolla tube above the middle.
Distr. Ten spp. endemic in Australia, one native in the West Indies, widely spread by culti-
vation and naturalized in the tropics.

1. Laurentia longiflora (L.) Petermann, Pflanzen- Isotoma longiflora Presl, Prod. Mon. Lob. (1836)
reich (1845) 444, t. 118 fig. 665; ed. 2 (1847) id.; 42; DC. Prod. 7 (1839) 413; Heyne, Nutt. PI.
Wimmer, in McBride Fl. Peru 6- (1937) 474; (1927) 1428; Back. Trop. Natuur 9 (1920) 129,
Ann. Naturh. Mus. Wien 56 (1948) 337, cum fig. 1-3; Fawc. &
Rendle, Fl. Jam. 7^ (1936)
var. runcinata (Hassk.) Wimmer; Pfl. R. Heft 138; Back. Bekn. Fl. Java (em. ed.) 8 (1949)
107 (1953) 405.— Lobelia longiflora Linne, Sp. fam. 184, p. 5; Steen. Fl. Sch. Indon. (1949)
PI. (1753) 930; Lindl. Bot. Reg. 14 (1828) t. 1200. 376; Quis. Med. PI. Philip. (1951) 955.— Isotoma
— Rapuntium longiflorum Mill. Diet. ed. 8 (1768) runcinata Hassk. Bonplandia 7 (1859) 189, ex
n. 7. Hippobroma longiflora G. Don, Gard. Diet. descr. — Fig. 23.
3 (1834) 717; McVaugh, Bull. Torr. Bot. CI. 67 Perennial (always?), erect or ascending, 10-30
(1940) 783; N. Am. Fl. 32A, pt 1 (1943) 99.— cm; stems up to 5 mm
diam., hairy to nearly
Dec. 1960] Campanulaceae (Moeliono & Tuyn) 141

glabrous, woody at the base, with ribs or wings Jamaica), in the lowlands from Florida and
running from the leaves downwards. Leaves Sonora to Brazil and Peru; introduced into Penn-
sessile, elongate-obovate, attenuated at the base, sylvania (U.S.A.), Hawaii, the Marianas, and the
acute or obtuse, mostly sparsely hairy, coarsely Old World tropics, almost throughout Malaysia.
sinuate-dentate, 3-16 by 1-3 cm. Flowers solitary, Ecol. In damp places, along ditches and
axillary, almost regular. Pedicels densely short- streams, against walls, from sea-level up to 300
hirsute, 3-15 mm. Bracteoies 2, at the base of (-1200) m. Fl. Jan. -Dec.
the pedicels, narrow, acute, up to 5 mm. Calyx This is the sole species which does not possess a
lobes inequal, narrow-lanceolate or linear, ir- trigger-hair on the two lower anthers, as described
regularly dentate, acute, hairy, 8-20 mm. Corolla in Isotoma by Hildebrand (Bot. Zeit. 27, 1869,
white, hairy, persistent, tube funnel-shaped, 476, f. 8-12) and Melville (Kew Bull. 14, 1960,
5-9 cm, lobes spreading, almost equal, lanceolate, 277-279, f. 1). In several American Laurent ias the
acute or obtuse, 10-25 mm. Filaments inserted two lower anthers have each 2 setae.
above the middle of the corolla tube, more or less Vern. Daun tolod, S, daun kendali, J, tambakis,
united, as long as or somewhat exceeding the N. Born., Philip.: estrella, Sp., Tag., lagrimao de
corolla tube, anthers curved forward, with a bearded San Diego, revienta caballos, Sp.
top, 4-6 mm, the 2 in front 2-3 mm. Ovary Note. The milky juice is said to be very
obconic to bell-shaped, (mostly 10-)ribbed, poisonous, in particular for horses. According to
densely hairy, especially along the ribs, 3-7 mm; BooRSMA (Heyne, Quis., //. cc.) its alcaloids
style equalling the anther tops or somewhat have a paralysing effect. The frequent occurrence
longer; stigma broad, flat, ±_ 2-lobed, bearing along streams seems to point to seed dispersal
long slender hairs beneath. Capsule nodding, by water; that against walls to seed dispersal by
ellipsoid. Seeds i^j, ellipsoid, foveolate-reticulate. ants, but there is no elaiosome.
Distr. Native in the West Indies (the type from

Cultivated species
Besides the already mentioned species of Lobelia, Backer mentioned in his Flora of Java (em. ed.)
8 (1949) fam. 183 the following species to be cultivated in the mountains of Java: Campanula medium
L., C. rapunculoides L., Legousia speculum-veneris (L.) FisCH., and Platycodon grandiflorum (J acq.) DC.
 JUGLANDACEAE (M. Jacobs, Leyden)

Juglancioceae represent a characteristic northern hemisphere family, in the New World

going south to Central America (Mexico, Costa Rica, Guatemala, Panama, Cuba, Hispaniola
and found S of the equator as fas as c. 30" S, absent from Africa, and overstepping the equator
also in the Malaysian region where Engelhardia extends to Java and New Guinea. This distri-
bution shows a remarkable resemblance with that of the Fagaceae-Castaneae which though absent
S of the equator in the Americas, occur in Africa in the Mediterranean part only, and though
rather well represented as far as New Guinea are also absent in Australia and the Pacific islands.
A noteworthy detail of this parallel is that although both are well represented in the Himalayan
region and the Indo-Chinese Peninsula no representative of either group is found in Ceylon
and the Deccan Peninsula!
Northwards the family extended much farther in Tertiary time and fossils are known from
Sakhalin, E. Siberia to 61 N (where at present Jiiglans occurs to 51 N), also Alaska (pollen
grains), Greenland, and Spitsbergen. Several genera which are now confined to East Asia or
North America occurred in Europe from the Upper Cretaceous until the Pliocene but became
gradually extinct there during the Pleistocene Ice Age. See also under Engelhardia.
Taxonomy. The family comprises 6 genera with c. 58 spp.. F/atycarya dLnd Pterocarya are
Asiatic, Alfaroa is American, while Engelhardia, Carya, and Jiiglans occur in both the Old
and the New World. As for the disputed monotypic genus Annamocarya Chev. (synonyms:
Caryojuglans Kirchh., Juglandicarya Reid & Chandler, Rhamphocarya Kuang), described
in 1941 from S. China and Tonkin, we refer to Leroy, Mem. Mus. Hist. Nat. n.s. Bot. 6
(1955) 66.
The family is a coherent one. Engelhardia and Alfaroa form a natural and rather primitive
group.
Thusfar three systems of subdivision for the Juglandaceae have been proposed. The first is
by Oersted, Vid. Medd. Nat. For. Kj0benhavn (1870) 159-174. The second is by
KoiDzuMi, who gives a complete subdivision with a taxonomic survey of the genera in Acta
Phytotax. & Geobot. 6 (1937) 1-16. The third is by Leroy, I.e. 1-246. The three systems have
in common, that they always keep Engelhardia together with Fterocarya, and Carya together
with Jiiglans.
The affinity of the Juglandaceae within the Amentiferae seems to be closest with the Myricaceae.
Palynology. According to Erdtman (Pollen Morph. & PI. Tax. 1952, 216) the pollen
grains have several characters in common with those in Betiilaceae, Casuarinaceae, Myricaceae,
and Rhoipteleaceae; those of Fagaceae etc. are less similar.
Phytochemistry. This family is known to accumulate different types of polyphenolic
compounds. The most characteristic constituent of Juglandaceae is naphtoquinon juglon. It
has been proved to occur in species of Jiiglans, Carya, and Fterocarya. In the tissues juglon
is present as a monoglucoside of the reduced form of the quinon (dihydrojuglon). Enzymatically
dihydrojuglonglucoside is split very rapidly and dihydrojuglon set free is concurrently oxydized
to juglon. Juglon is very toxic to fungi and seedlings of higher plants. It is also toxic to animals
and may well be the ichthyotoxic principle oi Juglandaceae; fresh barks and leaves of Jiiglans
rupestris Engelm., Carya illinoensis (Wang) K. Koch, Engelhardia spicata Bl., and Engel-
hardia polystachya Radlk. are reported in literature to be used as fish poison.
Besides juglon, flavonoid compounds and tannins are known to accumulate in considerable
amounts in Juglandaceae. The bark of Engelhardia formosana Hayata contains up to 7.5 °o
of flavonoid glycosides (engelitin, isoengelitin, afzelin, astilbin, quercitrin). Similar compounds
were isolated from leaves of Jiiglans spp. The tannins of Juglandaceae have been investigated
in recent years. They seem to belong to the hydrolyzable class and hydrolysis gives rise to
gallic acid, ellagic acid, and glucose. One tannin, juglanin, was proved to be isomeric with
corilagin. Ellagic acid and gallic acid occur also in the free state in the wood of Juglans siebol-
diana and Flatycarya strobilacea.

(143)
144 Flora Malesiana [ser. I, vol. 6^

Fig. Engelhardia rigida Bl. a. Fruiting twig, X '/2, d-f- variation in the ^ flowers, all x 6, g. ^ flower
1.

afterremoval of the stamens, with 2 perianth lobes and 3 bract lobes, /;. 3 flower from above, with
the bract lobes (b) and 4 perianth lobes.— £. serrata Bl. (?) b. Seedling, x V2.—E. spicata Lechen.
ex Bl. c. 5 Flower, X 6 (a Brass 11 061, 6 Burkill 9984, c Koorders 29792, d Endert 3447, e Endert
4407, f-h Clemens 494).
Dec. 1960] JuGLANDACEAE (Jacobs) 145

The high ascorbic acid contents (= vitamin-C) of different organs of Juglans regia reported
in literature are greatly exaggerated because juglon interferes strongly in the usual determination
of ascorbic acid.
The available data permit the characterization of the Juglandaceae phytochemically as a
family which accumulates many types of tannin-like polyphenolyc compounds. In this respect
it resembles many other woody families, e.g. Hamanwlidaceae, Fagaceae, etc. — R. Hegnauer.

1. ENGELHARDIA
Lechen. ex Blume, Bijdr. 10 (1825) 528; Fl. Jav. Jugl. (1829) 5, 'Engelhardtia';
CDC. Ann. Sc. Nat. IV, 18 (1862) 35; Prod. 16, 2 (1864) 140; Nagel, Bot. Jahrb.
50 (1914) 415.^Pten/ema Reinw. Syll. 2 (1826) 13.— Oreomwviea Oerst. Vid.
Medd. Nat. For. Kjobenh. (1856) 52.— Fig. 1-9.
Trees, mostly lepidote with golden yellow, glandular scales 0.1-0.2 diam. mm
Pith solid. Leaf buds naked, with a resemblance to hands. Young twigs densely
brownish tomentellous, young leaves mostly reddish. Stipules none. Leaves spirally
arranged (in Asiatic members), paripinnate; leaflets slightly asymmetrical, the
acroscopic side being mostly wider with higher inserted base than the basiscopic
side; midrib above flattish with a narrow, raised keel, nerves camptodromous.
Monoecious or dioecious. Flowers small, in catkins (each flower solitary) which
are often arranged to lateral or terminal panicles, the flower fused with a 3-lobed
bract; perianth 4-lobed. — d" Flowers: perianth often reduced and irregular; stamens
4-13 (in Asiatic members), (sub)sessile. —2 Flowers: perianth lobes in 2 whorls
partly connate with the ovary, the median pair exterior; ovary 2-carpellate, the
carpel sutures median, 1-locular with an incomplete, simple (in Asiatic members),
transverse septum, sometimes also with an abortive median septum; ovule 1,
erect, at the top of the main septum, conical with broad base, atropous, with 1
integument. Style in sect. Engelhardia well-developed with 2(-4) transversal elongate,
papillose, persistent stigmas, in sect. Psilocarpeae the 4 stigmas small and sessile.
In fruit the strongly accrescent bract forms a 3-lobed, obovate, scarious wing with
midribs and reticulate venation and partly adnate to the nut; the central (abaxial)
lobe is about twice as large as the lateral ones, sometimes a smaller adaxial simple
or irregularly dissected lobe is present. Nut indehiscent, pea-sized; pericarp carti-
laginous, mostly hispid with itching hairs. Seed fi\\\r\% the space between the septa;
testa forming an even covering of the seed; cotyledons strongly plicate and divided
into 4 lobes, later epigeal, radicle superior, endosperm none.
Distr. Five spp. in the Old World, from the W. Himalaya to China S of :i: 30 N, all over the Indo-
Chinese Peninsula, Malaysia, and Formosa; in America 2-3 spp. in Mexico, Costa Rica, and Guate-
mala. Fig. 2.
Ecol. Trees, mostly small or medium but sometimes growing very large, in evergreen, primary dryland
forest, apparently not on limestone. In Malaysia they prefer the mountains between c. 1000 and 2000 m.
although sometimes found in the lowlands.
Though Eiigelluirdia is a regular constituent of the fagolauraceous montane rain-forest in Java, none
of the species occurs gregarious or dominant, but in open grassy places they may occur in groups in
anthropogenous pseudosavannahs. Van Steenis observed on Mt Patuha (W. Java) abundant regenera-
tion of seedlings c. 10-200 cm high in a dense thicket of Eiiparoriiim iniilifoliiim on abandoned cinchona
plantations near Tjipadaruum, at c. 1800 m; the seeds were derived from three old fruiting trees which
happened to be in the adjacent forest border. See further under E. spicata.
Often (always?) shortly deciduous and then flowering. Pollination and seed dispersal are clearly
adapted to wind. However, the matter may be not so simple as that. Dr P. van Royen told me that
in New Guinea, where E. ligida is very common, he repeatedly found all the fruiting catkins, shed
as a whole, on the ground under the trees.
146 Flora Malesiana [ser. I, vol. 6^

Fig. 2. Distribution of Engelhardia Lechen. ex Bl. in the Old World, with number of species indicated
for each subarea.

Wood-ana t. Beekman, Meded. Proefstat. Boschw. 5 (1920) 96; den Berger, Meded. Proefstat.
Boschw. 13 (1926) 13, Determinatietabel van Malesie, Veenman, Wageningen (1949) 62 (hand lens.)
Heimsch, Lilloa 8 (1942) 164; Kribs, Trop. Woods 12 (1927) 16; Metcalfe & Chalk, Anat. Die. 2
(1950) 1287; Moll & Janssonius, Mikr. Holzes 6 (1936) 308; Tippo, Bot. Gaz. 100 (1938) 42. For
various opinions about the affinities of the family see Metcalfe & Chalk, I.e. —C.A.R.-G.
Morph. Not all authorities agree on the nature of the floral parts, called by Manning the floral
envelope, in his opinion consisting, when complete, of a 3-lobed bract, 2 bracteoles, and a 4-lobed
perianth. Other authors hold the view that the flower is surrounded by a 1-lobed bract with 2 bracteoles,
and a 4-lobed perianth, the lobes of which are some times called sepals. The nature of the bracteoles is
subject to interpretation. The smaller and facultative adaxial lobe of the fruit is, in my opinion, too
variable to represent anything but an outgrowth of the two adaxial margins of the bract lobes.
Hjelmqvist, basing his opinion on a comparison with the Myrieaceae, supposed that the flower was
originally unisexual and of a pseudanthic nature, i.e. it has arisen through union of a number of simply
built flowers. Nagel and Manning suppose that the flower was initially bisexual and has become
unisexual by reduction. This is supported by the fact that, very rarely, seemingly c? catkins are found
where part of the flowers have a prolonged bract. Such flowers often show both a reduced androecium
and a reduced gynoecium, i.e. a sterile stigmatiferous ovary surrounded by a subnormal number of
stamens.
The nut is also regarded as a sort of drupe with thin flesh which, however, is derived from the in-
volucre or perianth and not from carpel tissue.
The ovule seems only to be basal on a columella. Virtually it is not truly basal, but modified axile,
at the apex of one of the primary septa.
According to Hjelmqvist the integument is protracted upwards to a narrow tube in E. spicata and
E. parvifolia (the latter here reduced to E. serrata).
Several important studies have been published, all covering the whole family. We mention Nagel,
Bot. Jahrb. 50 (1914) 459-530; Manning, Amer. J. Bot. 25 (1938) 407-419 on the inflorescence, ibid.
27 (1940) 839-852 on the ? flowers, ibid. 35 (1948) 606-621 on the S flowers. Bull. Torrey Bot. Club
86 (1959) 190-198 on Engelhardia and Alfaroa in America; Hjelmqvist, Bot. Notis. Suppl. 2^ (1948)
5-171; Leroy, Mdm. Mus. Hist. Nat. n.s. Bot. 6 (1955) 1-246.
Fossils. Many records prove the wide occurrence of Engelhardia in the warmer parts of Europe
Dec. 1960] JUGLANDACEAE (JaCObs) 147

and also of N. America in the Tertiary. Dr W. H. Zagwijn kindly informed me, that Engelhardia pollen
forms an important part (up to 20 "„) of the anemophilous elements of a flora which existed in the SE.
part of the Netherlands and the adjacent region in Germany. This flora, of the Miocene and Pliocene
clay and browncoal formations, is remarkable for its strong E. Asiatic and American affinities, and
for the many tropical and subtropical elements it contains, especially in the Mid-Miocene, where these
make up about 50 ° „ of the flora. See also Zagwijn, Aspects of the Pliocene and early Pleistocene
Vegetation in the Netherlands, thesis Leiden I960. An endocarp of fE. nucifera (Lxjovjig) Madler is
depicted in Fortschr. Geol. Rheinl. u. Westf. 4 (1959) t. 1, by the same author.

Fig. 3. Fruiting Engelhardia spicata Lechen. ex Bl. above Sembalun, Mt Rindjani, Lombok, c. 2000 m
(de Voogd).

Variability. There is a significant difference between juvenile and adult leaves. The juvenile leaves,
on young trees and suckers, bear more pairs of leaflets, are thinner, longer acuminate, more serrate,
and denser hairy than those of mature trees. This holds for both the American species and at least for
part of the Malaysian ones.
The numerous field data on bark characters vary so much that it was found impossible to make reliable
descriptions.
In reproductive parts there is an astounding variation. The panicles of catkins are variously reduced,
often the single V catkin or the few ,-^ catkins remaining. In sect. Engelhardia the perianth is often liable
to much reduction. In the (^ flowers some stamens in a row may be lacking, or the indi\idual stamens
are reduced in size. In the nuts part of the perianth lobes may be wanting, or, on the other hand, may
have grown out excessively, up to nearly the length of the lateral wing lobes. The variation displayed
by the adaxial lobes is practically beyond description.
In the descriptions the + flowers are not mentioned because they are considered rather irrelevant for
diagnostic purpose. Their size varies much in the course of development and it is difficult to define
148 Flora Malesiana [ser. I, vol. 6^

in dry material the exactmoment when the stigmas are receptive; material with $ flowers is, moreover,
very rare in herbaria. The collecting of $ flowers and the making of observations on their anthesis is
strongly recommended to the attention of field workers.
Taxon. Several authors have, in the course of time, contributed to the present subdivision into 3
sections.
Sect. 1. Engelhardia. Sect. Pterilema (Reinw.) C. DC. Sect. Tric/wtocarpeae Nagel. Leaves spirally
arranged. Inflorescences lateral. Male flowers: subsessile; bract with 3 distinct narrow lobes; receptacle
elongate; perianth flat, irregular, reduced, often obscure; stamens in transversal rows of 2-3, hirsute.
Female flowers: stigmas commissural, hence transversal, linear, style far exceeding the perianth lobes.
Nut hispid, (sub)sessile, about halfway adnate to the wing, 2(-4)-celled at the base. Here belong E.
spicata Lechen. ex Bl. (the type sp. of the genus), E. rigidci Bl., E. serrata Bl., and E. apoensis Elmer
ex Nagel.
Nagel ew;e/?rf. Leroy. Leaves spirally arranged. Inflorescences terminal, sometimes
5ecr. 2. Psilocarpeae
also lateral. Male
flowers: stalked; bract very obscurely 3-lobed; receptacle orbicular; perianth regular,
4-lobed, lobes cucuUate, stamens (2-)3 at the base of each lobe, glabrous. Female flowers: stigmas split-
carinal, cushion-like, concealed by the perianth-lobes; no style. Nut scaly, not hairy, stalked, hardly
adnate to the wing, 2(-4)-celled at the base. Here belongs E. roxburghiana Wall.
Sect. 3. Oreomunnea (Oerst.) C. DC. (only American and not incorporated in the above generic
description). Leaves opposite or whorled. Inflorescence lateral. Male flowers: subsessile; bract obscurely
3-lobed; receptacle elongate, periant 2-4-lobed, flattish; stamens -^ 16-19(-23), irregularly arranged,
glabrous. Female flowers: septa nearly complete, ramified, one true septum transversal, a false septum
median; stigmas carinal, hence median, cushion-like on short styles just or almost exceeding the perianth
lobes. Nut (sub)sessile, not hairy but obscurely scaly, 8-celled at the base, 4-celled in the middle, 1 -celled
at the apex; the testa attached to all irregularities in the pericarp; wing for -^ 1/3 adnate to the nut,
often very large, with a large simple adaxial lobe completely concealing the nut. This section,
which comprises the 2-3 American spp., is considered to represent the most primitive group in
the genus.
Oreomunnea Oersted, often spelled Oreamiinoa, which is more correct from an etymological but not
from a nomenclatural point of view, is by several authors kept apart. Manning gave reasons for its
reduction to Engelhardia in Bull. Torrey Bot. Club 76 (1949) 196-209; in the same paper he commented
on characters in Alfaroa and Engelhardia.
Uses. The wood is locally applied for timber but considered of inferior quality. The bark is occasionally
used for fish-poison.
Notes. An attempt has been made to evaluate ail names given to Asiatic Engelhardias. I was not able
to place E. mollis Hu, Bull. Fan Mem. Inst. Biol. Peiping, Bot. 10 (1940) 161, from SW. Yunnan.
The author is greatly indebted to Professor Dr Wayne E. Manning, Lewisburg, Penns., U.S.A.,
who spent much time to go through the MS
and put the fruits of his lifelong work in the Juglandaceae
at our disposal in a most generous and liberal way.

KEY TO the species

1. Nut and stigmas. Bract of the $ flowers with 3 distinct narrow
hispid, (sub)sessile, with distinct style
lobes. Stamens hairy. Inflorescences lateral.
2. Leaflets underneath more or less densely set with yellow scales (hand lens!). Fruiting catkins on a
slender subterete stalk.
3.Nut subglobose, 3-5 mmlong, wing up to 4, rarely to 6 cm long. Leaflets widest about the middle
or above, the base acutish to obtuse. Stamens 3-7. Dioecious.
4. Leaflets entire 1. E. rigida

4. Leaflets (more or less distinctly) serrate or crenate 2. E. serrata

3. Nut elongate, ovoid, 6-8 mmlong, wing 5'/i —
71/2 cm long. Leaflets ovate, the base subcordate
to rounded. (Flowers unknown) 3. E. apoensis
2. Leaflets underneath sometimes with minute greyish (not yellow) scales, but mostly with hairs only
or glabrous. Stamens 8-13. Fruiting catkins on a vigorous, angular stalk. Monoecious.
4. E. spicata
1. Nut scaly, not hairy, stalked, stigmas concealed. Bract of the ^ flowers obscurely lobed, but 4 perianth
lobes distinct. Stamens glabrous. Inflorescence terminal, sometimes also lateral.
5. E. roxburghiana

1. Engelhardia rigida Bl. Bijdr. 10 (1825) 528; 155 p. l.—E. subsimplicifolia Merr. Govt Lab.
Fl. Jav. Jugl.(1829) 13, t. 3; Miq. FI. Ind. Bat. 1, Publ. Philip, n. 34 (1906) 6; Philip. J. Sc. 1 (1906)
1 (1856) 842; C. DC. Prod. 16, 2 (1864) 142; Suppl. 41; Nagel, Bot. Jahrb. 50 (1914) 477;
K. & V. Bijdr. 5 (1900) 175; Koord. Exk. Fl. Merr. En. Philip. 2 (1923) 24.— £. lepidota
Java 2 (1912) 53; Back. Bull. Jard. Bot. Btzg II, Schltr, Bot. Jahrb. 50 (1913) 66, f. 1; Nagel,
n. 12 (1913) 16; Nagel, Bot. Jahrb. 50 (1914) Bot. Jahrb. 50 (1914) 477; Rendle, J. Bot. 61
476; Koord. Atlas 4 (1916) t. 698; Bakh. /. in (1923) Suppl. 53.— £. zambalensis Elmer, Leafl.
Back. Bekn. Fl. Java (em. ed.) 7A (1948) fam. Philip. Bot. 9 (1934) 3195.— Fig. la, d-h, 4c-f.
Dec. 1960] JUGLANDACEAE (JaCObs) 149

Tree 15-30(-47)'m, sometimes with buttresses

up to 3 m high and 2 m out; the bark once
reported decorticating in small flakes. Leaf rachis
vigorous, blackish, (1 '/2-)3-6(-21) cm, mostly
glabrous or rarely puberulous, (l-)2-3
scaly,
(-5)-jugate; rather thick-coriaceous, ses-
leaflets
sile to 5 mm
stalked, blade 2'4-3|/2(-4) times as
long as wide, widest at the middle or rarely above,
5'/2-10'/2(-16i/2) by 2'/2-4(-7i/2) cm; base acutish,
top subacuminate, tip acute; margin entire (in
New Guinea exceptionally crenate towards the
top), surfaces more or less densely set with golden
yellow glandular-scales, especially underneath,
otherwise glabrous or rarely hairy on the nerves.
Dioecious. Catkins lateral on leafy and on slightly
older twigs, rarely serial in twos. Male catkins
solitary or 2-3 on a I/2-I cm long stalk, V2-5 1

cm long, rachis scaly and/or hairy. Bract 2-4 mm,

scaly, lobes irregular, narrow. Perianth 4-lobed,
the lobes resembling those of the bract, to com-
pletely reduced. Stamens (3-)4-6(-7), the anthers
equal or unequal, (Vi-)! mm, more or less hispid,
connective not or shortly protruding; if there is a
single proximal stamen, this has sometimes a
filament up to 1 mm, the others being sessile.
Fruiting catkins (6-)10-15(-22) cm including a
peduncle 2-3(-7) cm; peduncle subterete, in the
basal half with 2 scars; rachis rather slender,
more or less densely scaly and/or puberulous.
Nut (sub)sessile, 3 i mm
diam., hispid, wing
(2-)2'/2-3(-6) cm long with the nut, 3-8 mm
wide; perianth lobes sometimes very unequal,
one or both of the median lobes sometimes at-
taining half the length of the wing; style about as
long as the stigmas, the whole 2-4 long. mm
Distr.Malaysia: W. Java (rare), Borneo,
Philippines (W. Luzon, N. Mindanao), Central
and SW. Celebes (also Banggai Arch.), Moluccas
(Halmaheira, Morotai), New Guinea (also Japen Fig. 4. Fruits of Engelhardia. a. E. ruxburg/iiana
and Numfoor). In New Guinea rather common, Wall., ventrally, nat. size, b. the stigmas,
elsewhere rare. shown by removal of 2 perianth lobes, 4. E.
Ecol. Primary forest, only once reported from rigida Bl. c. Fruit dorsally, x 2, d-f, fruits
secondary forest, c. 1000-1 800(-2300) m. ventrally, showing variation in perianth and
Vern. Ki hiidjan, S, sibiri, Papua. adaxial lobe,x 2. E. serrata Bl. g. Longi-
Uses. The wood is used for canoes, native tudinal section of fruit, showing ovulum on
buildings, etc. columella (c) and partial septa (s), x 4 (a-b
Notes. Only in Borneo, the Philippines, and Clemens 27250, c-d Winckel 1540/3, e Brass
Celebes occur specimens with -jugate leaves.1
11051, / Kalkman 4363, g Jacobs 4677).
Closely related to E. serrata, and arguments
could be advanced to regard the two as subspecies.
Separation based on reproductive characters is leaflets were also found. Engelhardia zambalensis,
practically impossible. However, apart from the described on a specimen with entire leaflets but
different distribution, either species has its own much reduced towards the leaf base, is therefore
particular pattern of variability, as has been placed by me under £. rigida, but if one accepts
pointed out in the descriptions. For separation the narrowing leaf base as the key character, it
two characters have been found, notably the leaf would come under E. serrata.
margin, which is entire or serrate, and the decreas- Two specimens from Japen and Numfoor
sterile
ing in size of the leaflets towards the leaf base; are believed to belong here, notwithstanding they
this feature is generally far more pronounced in lack scales on the underside of the leaves.
£. serrata, where the basal leaflets are about half In Endert 3833 and Kostermans 7648, both
as long as the largest ones. These characters from E. Borneo, the fruit bears no long hairs,
mostly concur. In the cases where they do not, I only a soft pubescence and scales. From £. roxhurg-
am inclined to take the leaf margin character as hiana these specimens are distinguished by well-
decisive, since in a few E. rigida specimens from developed style and stigmas. In Robbins 569
New Guinea and elsewhere such small basal from New Guinea even the anthers bear scales.
150 Flora Malesiana [ser. I, vol. 6^

2. Engelhardia serrata Bl. F1. Jav. Jugl. (1829)

14, t. 4, 5c; MiQ. Fl. Ind. Bat.
1, (1856) 843;
1

C. DC. Prod. 16, 2 (1864) 141; Hook./. Fl. Br.

Ind. 5 (1888) 596; K. & V. Bijdr. 5 (1900) 172,
174; KooRD. Exk. Fl. Java 2 (1912) 53, f. 7;
Nagel, Bot. Jahrb. 50 (1914) 477; Koord.
Atlas 4 (1916) t. 699; Dode, Fl. Gen. I.-C. 5
(1930) 928; Quis. Philip. J. Sc. 76 (1944) 37;
Bakh. /. in Back. Bekn. Fl. Java (em. ed.) 7A
(1948) fam. 155 p. 3. E. palembanica MiQ. Sum.
(1861) 346, 139.— £. parvifolia C. DC. Ann. Sc.
Nat. IV, 18 (1862) 34; Prod. 16, 2 (1864) 141;
Nagel, Bot. Jahrb. 50 (1914) 477; Merr. En.
Philip. 2 (1923) 23.— £. nudiflora Hook. /. Ic.
PI. 18 (March 1888) t. 1747; Fl. Br. Ind. 5 (Dec.
1888) 597; Nagel, Bot. Jahrb. 50 (1914) 477;
Gamble, J. As. Soc. Beng. 75, ii (1915) 401,
incl. var. crenata Hook. /. ex Gamble; Ridl.
Fl. Mai. Pen. 3 (1924) 369. f. 158; Corner,
Ways. Trees (1940) 333, f. 1 17. E. permicrophylla
Elmer, Leafl. Philip. Bot. 9 (1934) 3194.— Fig.
lb, 4g, 5.
Tree (2-)20-25(^2) m, sometimes with but-
tresses up to 3-4 m high and 1 '/2 rn out. Leaf
rachis (3-)5-18(-21) cm, more or less hairy,
3-7(-9)-jugate; leaflets varying from small, co-
riaceous, and glabrous to large, herbaceous, and
hirsute (mainly on the nerves), (sub)sessile,
markedly decreasing in size towards the leaf-
base, the basal leaflets attaining about half the
length of the largest ones, 1 '/2-3 times as long as
wide, widest at the middle or above, (l-)2-13'/2
by ('/2-)I-4 cm; base narrowed, acutish to obtuse,
top acutish to subacuminate; margin crenate or
serrate sometimes from the base but at any
rate towards the top; surface dull, underneath
more or less densely set with golden yellow
glandular scales. Dioecious. Catkins lateral on
leafy and on slightly older twigs, the peduncles
solitary in the leaf axil or serial by twos or threes.
Male catkins in bundles of 2-3, sessile to 3 cm Fig. 5. Variation in of E. serrata Bl.,
leaflets
stalked, 3-10 cm long; perianth completely the acroscopic side on the a-d. x 2/3, e. leaf
left,

reduced or almost so: stamens (3-)5-7, sessile to base densely set with glandular scales (a Hout-
Vi mm anthers sometimes unequal,
stalked, soorten Gedeh 126, b Sulit 21595, c, e Holttum
V^-1 mm,
sparsely hirsute, connective not or 14857, d Clemens K
hardly protruding. Fruiting catkins 9-14 cm long
in all, 2-5 cm stalked, rachis as thick as that of Notes. In the Philippines several collections
the leaves, shallowly grooved. Nut (sub)sessile, possess very small leaflets, glabrous, viz 2-3 cm,
± 3 mm
diam., hispid, wing 1 '/2-3'/4(-334) cm the fruit wing being ~ 15-21 mm
long. In Celebes
long with the nut, 5-10 mm
wide; perianth lobes specimens occur in which the leaflets are slightly
rather equal, often not adnate to the style, style larger and 2-3-jugate, the fruit wing being
about as long as the stigmas, the whole 2-4 mm slightly smaller. In Malaya, Sumatra, Banka, and
long. Billiton the leaflets are generally 5-7 cm and
Distr. Upper Burma, N. Siam, Indo-China glabrous. In Java the leaflets are thin in texture,
(Laos, Cambodia), in Malaysia: Malay Peninsula, up to 9-jugate, 8-1 3 '/2 cm long, serrate from the
Sumatra (S of Lake Toba, also Bengkalis, Banka, base, and hirsute, especially on the nerves. Inter-
Billiton), W. Java. Borneo. Philippines (Luzon to grading specimens occur throughout the area.
N. Mindanao), Celebes, Moluccas (W. Ceram). Pierre 3301, from Cambodia, is a very bad
Ecol. Primary dryland forest (rarely secondary specimen with entire, acuminate leaves glandular-
forest)on sandy or clayey soil, from sea-level to scaly on both sides. Most probably it belongs here.
2200 m. In Malaya the leaves are shed at the end Jacobs 4677, from Mt Sago in W. Sumatra,
of a dry spell without fading into autumn tints in fruit, is the only specimen with entire leaflets.
(Corner). The leaf rachis is sparsely hairy, the leaflets are
Vern. Ki hudjan, {lang)kedi, {be)biri, M, ki 4 times as long as wide, the perianth lobes are
keper, S. not connate with the style. It tallies with a
Dec. 1960] JUGLANDACEAE (JaCObs) 151

specimen from the same locality that has serrate 4. Engelhardia spicata Lechen. ex Bl. Bijdr. 10
leaves. (1825) 528; Fl. Jav. Jugl. (1829) 8, t. 1, 5A; MiQ.
Most of the extra-Malaysian specimens, es- Fl. Ind. Bat. I, 1 (1856) 842; C. DC. Prod. 16,
pecially those from N. Siam, are very densely 2 (1864) 140; Kurz, For. Fl. Burma 2 (1877)
brown-pubescent on the twigs, the rachises of 491; ViDAL, Sin. Atlas (1883) t. 90A; Hook. /.
leaves and catkins, on the leaflets underneath and Fl. Br. Ind. 5 (1888) 595; K. &
V. Bijdr. 5 (1900)
the nerves above. The largest leaflet seen was 17'/2 165, 170, incl. var. a genuina et var. ft aceriflora,
by 5'/2 cm. The leaflets are sometimes nearly var. colebrookiana quoad specim. tantum;
y
entire, with their indument resembling E. spicato Koord. Exk. Fl. Java 2 (1912) 51, f. 6; Nagel,
var. colebrookeana, but have always yellow scales Bot. Jahrb. 50 (1914) 476, incl. var. aceriflora;
below. Leefm. Trop. Natuur 3 (1914) 46 with fig., 77;

3. Engelhardia apoensis Elmer ex Nagel, Bot.

Jahrb. 50 (1914) 477; Elmer, Leafl. Philip. Bot. 7
(1915) 2693. Engelhardtia sp. Corner, Ways.
Trees (1940) 333, f. 117.— Fig, 6.
Lfo/rachis 13-23 cm, more or less densely set
with scales and small hairs. Leaflets alternate,
7-10, thinly coriaceous, 2-4 mm
stalked; blade
± 2'/2-3 times as long as wide, ovate, (7-)9-13
by 2'/4-3(-5) cm; base rounded to subcordate, top
narrowed; margin comparatively finely serrate,
especially in the basal part; surface above glossy,
below glabrous or (in the type specimen) with

Fig. 6. Engelhardia apoensis Elm. ex Nagel,

leaflet and fruit, x 2/3 (CF 24424 Symington).

some minute tomentum on the nerves. Flowers

not known; inflorescences presumably lateral.
Fruiting catkins12-30 cm long in all, stalk 4-6
cm. Nut +
6-8 by 5-6 mm, the body ovoid,
elongate, wing (5'/2-)6-7'/2 cm long with the
nut, 11-18 mm
wide, adaxial lobe well developed,
4-6(-15)mm long, style 2-3 mm, stigmas 10 mm
long.
Distr. Malaysia: Malay Peninsula, Borneo
(N. Sarawak), Philippines (Mindanao: Mt Apo).
Notes. This taxon is remarkable for its ovate
leaflets and elongate nut.
The 7 examined specimens from Malaya match Fig. Engelhardia spicata Lechen. ex Bl. Large
7.
the type specimen from Mindanao, Elmer 1744, I tree, over 40 m tall, in primary forest near
very well, the Malayan plants having somewhat Tjikahuripan, Mt Gedeh, W. Java; stem base
longer catkins and larger fruits. In Richards 1521 8.84 m in girth, dbh 2.8 m (F. Kramer, who is
from N. Sarawak the leaves are entire. standing in front).
152 Flora Malesiana [ser. 1, vol. 6^

Gamble, J. As. Soc. Beng. 75, ii (1915) 400; C. DC. Ann. Sc. Nat. IV, 18 (1862) 34, t. 2 f.

KooRD. Atlas 4 (1916) t. 700; Merr. Sp. Blanc. 15; Prod. 16, 2 (1864) 141.— Fig. Ic, 3, 7-9.

(1918) 120; En. Philip. 2 (1923) 24; Koord. F1. Tree (5-)12-30(-36) m, sometimes with small
Tjibodas 7 (1923) 10, var. aceriflora (Reinw.) buttresses. Leaf rachis vigorous, blackish, (51/2-)
K. &v.; RiDL. Fl. Mai. Pen. 3 (1924) 368; 10-30(-40) cm, glabrous and/or scaly to hirsute
DocTERS VAN Leeuwen, Zoocecidia (1926) 103; and/or tomentose, (2-)4-5(-7)-jugate; leaflets
DODE, Fl. Gen. I.-C. 5 (1930) 930, f. 107, 3-7; firmly herbaceous to coriaceous, sessile to 5( 15) —
Chun, Sunyatsenia 4 (1940) 245; Corner, Ways. mm stalked, blade (1.7-)2.4-3.0(-3.4) times as
Trees (1940) 333, f. 117; Kanj. c.s. Fl. Assam 4 long as wide, widest about the middle or rarely
(1940) 229; de Voogd, Trop. Natuur 30 (1941) above, (6-)8i/2-16(-30) by (3-)5 •/2-6(-8) cm; base
103, f. 5-6, 125; Bakh. /. in Back. Bekn. Fl. very unequal, acute to subcordate, top shortly
Java (em. ed.) 7A (1948) fam. 155 p. l.—Pterilema acuminate, the tip obtuse to acute, rarely the top
aceriflorum Reinw. Sylloge 2 (1826) 13. E. rounded to acutish; margin entire, rarely serrate;
aceriflora Bl. Fl. Jav. Jugi. (1829) 11, t. 2, 5B; surfaces glabrous to hirsute, especially underneath
Mio. Fl. Ind. Bat. 1, 1 (1856) 842; C. DC. Prod. and on the nerves, sometimes underneath with
16, 2 (1864) 141; Hook./. Fl. Br. Ind. 5 (1888) domatia and/or thin Hat greyish (never yellow)
596, "acerifoiuC; Forbes &
Hemsl. J. Linn. Soc. scales. In juvenile specimens and suckers leaves
Bot. 26 (1899) 495; von Malm, Fedde Rep. 34 large, the leaflets serrate, hirsute. Inflorescences
(1934) 271. E. roxbiirghiami Lindl. e.x Wall. mostly bisexual, paniculate, lateral on leafy and
PI. As. Rar. 2 (1831) 85, t. 199 f. 7, quoad fnict.— on slightly older twigs, with i(-0) central $ catkin
Juglans pterococca Roxb. Fl. Ind. (ed. Carey) 3 and 2-5 basal rj catkins subtended by a caducous
(1832) 631, quoad fruct., 'plerococca'. Giro- — subulate bract 3-5 mm, rarely the basal catkins
carpus pendulos Blanco, Fl. Filip. ed. 2 (1845) also ?. Male catkins (5-) 10- 18 cm long in all,
55, ed. 3 (1877) 104, t. 387.— £. philippiiwiisis 0-1 cm stalked; bracts 2-4 mm, the apical lobe
sometimes mucro-like, the lateral lobes very
irregular, narrow, sometimes much reduced,
perianth whether or not reduced, very irregular,
the lobes up to 2 mm
long, a small proximal lobe
mostly present. Stamens 8-l2(-13), anthers (sub)-
sessile, equal or unequal, J; 1 mm
long, hirsute,
connective '/^-'^ rnm pointed. Fruiting catkins
(12-)21-40(-60) cm long in all, peduncle 2-IOI/2
cm, vigorous, angular, glabrous and/or greyish-
scaly. Nut (sub)sessile, 3-4 mm
diam., hispid,
wing (2-)2%-334(-6i/2) cm long with the nut,
(3-)7-9 mmwide, adaxial lobe undivided to 5-
lobed, often very irregular; perianth lobes fair-
ly equal, mostly small and connate with the style;
style about as long as the stigmas, the whole
^ 3 1/9-9 mm long.
India (in the montane Himalayas
Distr.
between about 75° and 90° E; Assam: Khasia,
Cachar, and Naga Hills, Manipur; E. Bengal:
Tripura, Chittagong), Tibet (Manning, //; litt.),

China (Yunnan, Kwangsi, /;.v.), Burma (S of

about 25° N), Siam, Indo-China (Laos, Cambodia,
Cochin-China), Hainan, in Malaysia: Sumatra
(Medan to Bencoolen), Malay Peninsula (Perak),
Java (W to E, very common). Lesser Sunda
Islands (Bali, Lombok, Sumbawa, Sumba,
Flores),Borneo (scattered), Philippines (Luzon,
Mindoro, Negros, Panay, Cebu, Mindanao;
rather common).
Ecol. Primary evergreen forest. Seems to prefer
the mountains up to 2000(-2500) m, especially
frequent in the Casuarina forests on the volcanoes
in Central and East Java. On the W. side of Mt
Jang in E. Java it is known to form locally pure
stands. Similar local dominance has been observed
by DE Voogd, I.e., on Mt Rindjani in Lombok;
he also observed it pioneering in mountain
savannahs. The dominance is due to succession
Fig. 8. Engelluirdia spicata Lechen. ex Bl. in and serai with Pittosporum, Homalanthus gigan-
;

clearing of coffee estate Kalisat, Mt Idjen, E. teus, Vernonia arborea, Dodonaea, Wendlandia,
Java (J. H. CoERT). etc. belonging to the pioneers of the rain-forest
Dec. 1960] JUGLANDACEAE (JaCobs) 153

Fig. 9. EngelharcHa spicata Lechen. ex Bl. in pyrogenous mountain savannah near Sembalun, Mt
Rindjani, Lombok, c. 2000 m (de Voogd).

which try to invade the pyrogenous Casiiarina small; twigs, leaf rachis. and leaflets underneath
jiinghiihuiana stands. densely hirsute, leaflets with rounded top, often
Often deciduous for a short time and then grey-greenish when dry. Fruiting catkins 16-26
flowering, not in definite periods. cm long in all, 21/2-8 cm stalked, the rachis
Vern. Ki hitdjcin. S, sowo, donghi, J, with various densely hirsute.
prefixes and variations. Distr. India (largely the same area as var.
Note. Easily distinguished from other species spicata, but not known from E. Bengal), S. China
by lacking the golden-yellow glandular scales (Kwangsi), Hainan, Burma (Kurz), N. and E.
on the lower surface of the leaves. If scales are Siam, Indo-China (Annam, Laos); in Malaysia:
present, these are flat, greyish and never golden- Philippines (Luzon, see below).
yellow. Notes. The material on which the Malaysian
record is based, Paraiso 25483 and Ramos &
colebrookeana (Lindl. ex Wall.) O. Kuntze,
vcir. Edano 37939. was identified by Dr Manning;
Rev. Gen. PI. (1891) 637, non K. & V. Bijdr.
1 I have not seen it.

5 (1900) 169, 172.— £. colebrookeana Lindl. ex E. spicata is a very variable species and the
Wall. PI. As. Rar. 3 (1832) 4. t. 208; Brandis, variation of its characters seems to occur more or
For. Fl. NW. & C. India (1874) 499; Hook./. less at random and not bound to a geographical
Fl. Br. Ind. 5 (1888) 596; Dode, Fl. Gen. I.-C. pattern. The reason that the combination of char-
5 (1930) 929; Merr. En. 23.— £.
Philip. 2 (1923) acters described above has received ta.xonomic
Yillosa KuRZ, For. Fl. Burma 2 (1877) 491, incl. recognition is solely because the specimens re-
var. Integra (according to Manning //; ////.). presenting this combination are conspicuous. All
E. esqidrolii Lev. in Fedde Rep. 12 (1913) 507, these characters, however, were also found to
cf. Rehder, J. Arn. Arb. 10 (1929) 118.— Fig. occur separately, and in other combinations as
4a-b. well. Therefore I adhere no more taxonomic value
Difl'ers from E. spicata var. spicata in the to this variety than as a random assemblage of
following characters: tree mostly comparatively marked paramorphs.
154 Flora Malesiana [ser. I, vol. 6^

In East Java, apart from the common form, a normal or slightly weaker lateral twigs, 2-4 cm
form dominates with densely hairy leaves, which peduncled, with central $ catkin (which may be
has incorrectly been referred to var. colebrookeana wanting) and 2-6 basal cJ catkins subtended by
by KooRDERS & Valeton. In my opinion this early caducuous bracts; rarely the $ catkin
form represents a local tendency in the population solitary. Flowers scaly. Male catkins 8-12 cm
that originated independently from the paramorphs long in all, V2-2 cm stalked; rachis slender, scaly,
on the Asiatic continent. Moreover, a complete not hairy; bract i 2 by 1 mm, obscurely 3-
series of intergrades exists between the glabrous lobed at the top, perianth lobes 4, not reduced,
and the hairy form in Java. strongly cucullate, 1-1 '/4 cm diam., with mem-
branous margin; stamens (8-) 12, inserted with
5. Engelhardia roxburghiana Wall. PI. As. Rar. (2-)3 at the base of each perianth lobe, filament
2 (1831) 85, t. 199, excl. fruct.; Brandis, For. and anther both Vi nim long, in some stamens
Fl. NW. & C. India (1874) 500.—Juglans reduced to 1/3 mm, glabrous. Fruiting catkins
pterococca RoxB. Hort. Beng. (1814) 68, nomen; 12-22 cm long in all, peduncle 2-5 '/2 cm, slender,
Fl. Ind. (ed. Carey) 3 (1832) 631, 'plerococca\ rachis tomentose, sometimes also scaly. Nut 4-8
descr., excl. fruct. — E. Hance, Ann.
chrysolepis mm stalked, transversely ellipsoid, 4-5 mm diam.,
Sc. Nat. IV, Nagel, Bot. Jahrb.
15 (1861) 227; scaly, not hairy; wing (3-)5-5i/2 cm long with the
50 (1914) 475; Corner, Ways. Trees (1940) 333, nut, 0.7-1.4(-l.8) cm wide, hardly connate with
f. 117; Metcalfe, Fl. Fukien 1 (1942) 45; P'ei, the nut, adaxial lobe almost none; perianth lobes
Bot. Bull. Acad. Sin. 1 (1947) 208.— £. wallichiana very equal and regular, 1-2 mm long, on top of
[LiNDL. ex Wall. Cat. (1831/2) no 4942] C. DC. the nut together enclosing the 4 sessile stigmas.
Prod. 16, 2 (1864) 141, incl. 3 chrysolepis; Hook. Distr. India (Assam), S. China (all provinces
/. Fl. Br. Ind. 5 (1888) 596; Forbes &
Hemsl. S of about 30° N lat.), Hainan, Formosa, Indo-
J. Linn. Soc. Bot. 26 (1899) 195; Nagel, Bot. China (Tonkin, Laos, Annam), in Malaysia:
Jahrb. 50 (1914) 475; Gamble, J. As. Soc. Bengal Sumatra (in the N. part especially, S to Bencoolen),
75, ii (1915) 402; Ridl. Fl. Mai. Pen. 3 (1924) Malay Peninsula (Kedah, Perak, Pahang, Penang),
370; Dode, Fl. Gen. I.-C. 5 (1930) 93\.—E. Borneo (Brunei and Kinabalu region). Everywhere
polystachya Radlk. Sitz. Ber. Bayer. Ak. Wiss. more or less rare.
Math.-Phys. Kl. 8 (1878) 385, ex descr.; Brandis, Ecol. Primary forests in hilly country between
Ind. Trees (1906) 620; Kanj. c.s. Fl. Assam 4 about 700 and 1500 m alt. Flowers and fruits
(1940) 301.— £•. pterococca O.K. Rev. Gen. PI. 2 not at definite times.
(1891) 637, quoad nomen et var. a roxburghiana Uses. In Sumatra said to be a fish-poison.
O.K. E. spicata var. formosana Hay. Fl. Mont.
Note. Wallich's treatment of this species com-
Form. (1908) 199.— £. formosana Hay. Ic. PI.
prises three elements: a two-line diagnosis from
Form. 6 (1916) 61; Kaneh. Form. Trees (1936)
Lindley's MS, an extensive analysis after Rox-
80, f. 35.— E.fenzelii Merr. Lingn. Sc. J. 7 (1931)
burgh's MS (the latter was to be published again,
300, ex descr.; Metcalfe, Fl. Fukien 1 (1942) 45.
under the name Juglans pterococca, in 1832), and
Tree ± 5-20 m, sometimes with low, rounded
a plate which is a copy of Roxburgh's drawing.
buttresses. Innovations bronze-coloured with
The diagnosis that Wallich took from Lindley
golden yellow glandular scales which become Hooker has already pointed out, to
refers, as
brown when older. Leaf rachis 5'/2-14 cm, as the E. spicata and the same is true for that part of
twig glabrous to sometimes densely brown
Roxburgh's plate and description where the fruit
puberulous, 2-4-jugate; leaflets coriaceous, on a
is given as hairy. The remaining part of Rox-
stalk 6-12 mm forming mostly a distinct joint burgh's plate and description clearly refer to
with the rachis; blade (2.4-)2.8-3.6(-4.0) times
E. roxburghiana in the present sense. Since only a
as long as wide, widest mostly below the middle,
minor part of Wallich's mixtum belongs to E.
sometimes halfway, (8-)10-16(-23) by (2-)3!/2-5 spicata,it is thought correct to accept Wallich's
(-71/2) cm; base very unequal, acute, often basis-
E. roxburghiana pro majore parte.
copically somewhat decurrent, top narrowed,
gradually bluntly acuminate; midrib narrowly
raised above; margin entire, surfaces in dry state Excluded
never even-coloured, but with dull shades of
greenish, greyish, brownish, or blackish, glabrous Engelhardia selanica Bl. Fl. Jav. Jugl. (1829)
but underneath yellow to brown glandular-scaly. 8 =Shorea selanica Bl. Mus. Bot. 2 (1852) 33
Probably sometimes dioecious, but mostly the {Dipterocarpaceae).
inflorescences bisexual, paniculate, terminal on
New York Bolanical Garde
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INDEX TO REVISED FAMILIES
Aceraceae . 4: 592 Dioscoreaceae
3, 4 293 Phytolaccaceae . . 229
Actinidiaceae s.str 37 Dipsacaceae ...
4: 4 290 Pittosporaceae 345
Aizoaceae 4: 267 Droseraceae ... 4 377 Plumbaginaceae 107
Alismataceae . 317 Elatinaceae
5: ... 4 203 Podostemaceae 65
Amaranthaceae . 4: 69, 593 Erythroxylaceae 5 543 Polemoniaceae . . 195
Ancistrocladaceae 4: 8 Ficoidaceae ... 4 267 Pontederiaceae 255
Aponogetonaceae 4: 11 Flacourtiaceae . . 5 1 Proteaceae 147
Basellaceae 5: 300 Flagellariaceae . . 4 245 Punicaceae 226
Batidaceae 5: 414 Gnetaceae. ... 4 336 Restionaceae . 416
Betulaceae 207 5: Gonystylaceae . . 4 349 Rhizophoraceae 429
Bixaceae . 4: 239 Goodeniaceae . . 5 335 Salicaceae 107
Burmanniaceae 4: 13, 592 Haemodoraceae . . 5 111 Salvadoraceae . 225
Burseraceae . 5: 209 Hamamelidaceae . . 5 363 Sarcospermaceae 32
Butomaceae . 5: 118 Hydrocaryaceae . . 4 43 Saururaceae 47
Callitrichaceae 4: 251 Hydrocharitaceae . 5 381 Scyphostegiaceae 297
Campanulaceae 6:, .107 Hydrophyllaceae . 4 207 Sonneratiaceae 4: 280,513
Cannabinaceae 4:' 223 Juglandaceae ... 6 143 Sparganiaceae . 4: 233
Capparidaceae 6: 61 Juncaceae ... 4 210 Sphenocleaceae . 4: 27
Caprifoliaceae 4: 175, 598 Juncaginaceae . . 4 '57 Stackhousiaceae 4: 35
.

Centrolepidaceae ^5: 421 . Malpighiaceae . . 5 125 Staphyleaceae 6: 49

.

CeratophyUaceae 4: 41 '
Martyniaceae . . 4 216 Styiidiaceae 4: 529
Chenopodiaceae 4: 99, 594 MoUuginaceae . . 4 '267 Styracaceae 4: 49
Cochlospermaceae 4:*^ 61 Moringaceae i^ 45 Thymelaeaceae 4: 349, 6: 1

.4
. . •
^

Combretaceae .
'
4:533 Myoporaceae . 265 Trigoniaceae . . 4: 59
Connaraceae . 5:495 Myricaceae . .4 . 277 Turneraceae . . 4: 235
Convolvulaceae 4: 338, 599 Nyssaceae ... 4 29 Typhaceae . 4: 243
Corynocarpaceae 4:'- 262 Papaveraceae . . \5 114 Umbelliferae . 4: 113,595
Crassulaceae . 4: 197 Pedaliaceae . . . ^4 >216 Valerianaceae . .4: 253
Datiscaceae 4: '382 Pentaphragmataceae . J j4 517 Xyridaceae 4: 366, 598
Dichapetalaceae . '5:'305 Pentaphylacaceae .'5 121 Zygophyllaceae .4: 64
Dilleniaceae 4: 141 Philydraceae ... 4 .:5
PRINTED IN THE NETHER LAND S BY
N.V. DIJKSTRA'S DRUKKERIJ V.H. BOEKDRUKKERIJ GEBR HOITSEMA GRONINGEN